Handbook of the Historiography of Biology (Historiographies of Science, 1) 3319901184, 9783319901183

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Historiographies of Science

Michael R. Dietrich Mark E. Borrello Oren Harman  Editors

Handbook of the Historiography of Biology

Historiographies of Science Series Editor Michael R. Dietrich Department of History and Philosophy of Science University of Pittsburgh Pittsburgh, PA, USA

The goal of this series is to provide definitive assessments of the historiography and the future of major fields and approaches within the history of science. Each volume will address the major trends in historical thought within a particular field, the major debates among historians of that field, and promising new directions that may shape future scholarship. Each volume is framed in terms of what a scholar should know about the history of work in that area, if they wanted to make a meaningful and original contribution to that field. Each volume will be written by experts in that field for graduate students and other scholars new to the history of that field. While other areas of history have extensive historiographic literatures, history of science has fewer resources from which to draw. The paucity of historiographical reflections by leading scholars in the history of science makes it more difficult for new scholars to join the field, as they try to assess the traditions of research on their own. These volumes will offer an informed introduction to major issues that will foster new, original research in the history of science. Editors will be asked to select topic areas/fields that they think have had a substantial and diverse body of scholarship. Each volume will be informed by different methods, theories, and perspectives that can be compared and contrasted in each volume. More information about this series at http://www.springer.com/series/15837

Michael R. Dietrich • Mark E. Borrello • Oren Harman Editors

Handbook of the Historiography of Biology With 5 Figures and 3 Tables

Editors Michael R. Dietrich Department of History and Philosophy of Science University of Pittsburgh Pittsburgh, PA, USA

Mark E. Borrello Program in the History of Science, Technology and Medicine Department of Ecology, Evolution and Behavior University of Minnesota Minneapolis, MN, USA

Oren Harman Graduate Program in Science Technology and Society Bar-Ilan University Tel Aviv, Israel

ISSN 2523-7748 ISSN 2523-7756 (electronic) ISBN 978-3-319-90118-3 ISBN 978-3-319-90119-0 (eBook) ISBN 978-3-319-90120-6 (print and electronic bundle) https://doi.org/10.1007/978-3-319-90119-0 © Springer Nature Switzerland AG 2021 This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. The publisher, the authors, and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, expressed or implied, with respect to the material contained herein or for any errors or omissions that may have been made. The publisher remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. This Springer imprint is published by the registered company Springer Nature Switzerland AG. The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland

Series Preface

While some areas of history have extensive historiographic literatures, history of science has fewer resources from which to draw than most. This scarcity of historiographical reflections by leading scholars makes it more challenging for newcomers who must try to assess traditions of historical research as they frame their own contribution to the history of science. As informed introductions to major themes in the writing of the history of science, we hope that this series will both help foster original research in the history of science and further discussion regarding historiographic trends. The goal of this series is to provide an assessment of the historiography and future of major approaches within the history of science. Each volume addresses the major trends in historical thought within a particular field, the major debates among historians of that field, and promising new directions that may shape future scholarship. Written for graduate students or scholars new to the history of science, each volume is framed in terms of what a scholar should know about the history of work in that area, if they wanted to make a meaningful and original contribution to that field. The volumes in the historiography of science series are not intended to provide comprehensive reviews of every topic discussed in the history of science. Editors of individual volumes select topic areas and fields that they think have had a substantial and diverse body of scholarship that have been informed by different methods, theories, and perspectives. Because we would like to foster more conversation about historiography, we see the idiosyncrasies of individual chapters, not as flawed and partial perspectives, but as opportunities to articulate diverse perspectives through an ongoing conversation. These volumes are open for revision through Springer’s Meteor publishing platform. Please engage with the authors and editors and help push this historiographic dialogue further. January 2021

Michael R. Dietrich

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Contents

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New Perspectives on the Historiography of Biology . . . . . . . . . . . . Michael R. Dietrich, Mark E. Borrello, and Oren Harman

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Charles Darwin and the Darwinian Tradition . . . . . . . . . . . . . . . . Janet Browne

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The Historiography of Modern Evolutionary Biology . . . . . . . . . . Mark E. Borrello

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The Historiography of Molecular Evolution . . . . . . . . . . . . . . . . . . Edna Suárez-Díaz

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The Historiography of Embryology and Developmental Biology Kate MacCord and Jane Maienschein

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Gregor Mendel and the History of Heredity Staffan Müller-Wille

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The History and Historiography of Eugenics . . . . . . . . . . . . . . . . . Paul Weindling

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The Historiography of Genetics . . . . . . . . . . . . . . . . . . . . . . . . . . . Michael R. Dietrich

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The Historiography of Molecular Biology Michel Morange

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Biomedicine and Its Historiography: A Systematic Review . . . . . . Nicolas Rasmussen

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The Historiography of Biotechnology . . . . . . . . . . . . . . . . . . . . . . . Nathan Crowe

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The Matter of Practice in the Historiography of the Experimental Life Sciences . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hannah Landecker

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Women in the Historiography of Biology . . . . . . . . . . . . . . . . . . . . Marsha L. Richmond

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The Historiography of Race and Physical Anthropology . . . . . . . . Tracy Teslow

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Local, Global, and Transnational Perspectives on the History of Biology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ana Barahona

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Organisms in Experimental Research Rachel A. Ankeny and Sabina Leonelli

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Scientific Biography Oren Harman

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Historiography and Immunology . . . . . . . . . . . . . . . . . . . . . . . . . . Warwick Anderson and Neeraja Sankaran

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The Historiography of the Sciences of the Brain and Nervous System . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. Chirimuuta

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Historiography of Marine Biology . . . . . . . . . . . . . . . . . . . . . . . . . Samantha Muka

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Historiography of Physiology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Andi Johnson

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Historiography of Plant Breeding and Agriculture Dominic J. Berry

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Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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About the Editors

Michael R. Dietrich is Professor and Chair of History and Philosophy of Science at the University of Pittsburgh. He studied Philosophy and Biology at Virginia Tech before earning a doctorate in Philosophy at the University of California, San Diego. As a historian and philosopher of twentieth-century biology, his primary interests are in the nature of scientific controversy. In numerous scholarly articles and chapters, he has explored controversies in evolutionary genetics and molecular evolution, as well as controversial figures, such as the émigré geneticist Richard Goldschmidt. He has coedited several books including Rebels, Mavericks, and Heretics in Biology with Oren Harman (2007), The Educated Eye: Visual Culture and Pedagogy in the Life Sciences with Nancy Anderson (2012), Biology Outside the Box: Boundary Crossers and Innovation in the Life Sciences with Oren Harman (2013), and Dreamers, Visionaries and Revolutionaries in the Life Sciences with Oren Harman (2018). He is currently writing a book on genetic drift with Roberta Millstein and Robert Skipper entitled Survival of the Luckiest: Perspectives on the History and Philosophy of Random Drift in Evolutionary Biology, as well as a biography of Richard Goldschmidt.

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About the Editors

Mark E. Borrello, Associate Professor of History of Science in the Department of Ecology, Evolution and Behavior, and Director of the Program in the History of Science and Technology, at the University of Minnesota, studied history and philosophy of science at Indiana University earning a doctorate in 2002. Before coming to the University of Minnesota, he was a visiting assistant professor at the Lyman Briggs School at Michigan State University. As a historian and philosopher of biology, his primary interests are in the development of evolutionary theory in the late-nineteenth and twentieth centuries. In numerous scholarly articles and chapters, he has explored the debate over the levels of selection idea from Darwin to the present. His 2010 book on this topic, Evolutionary Restraints: the Contentious History of Group Selection, was published by the University of Chicago Press. He is currently engaged in an investigation of the nature of individuality in developmental and evolutionary contexts. He has published on this topic with his colleagues Michael Travisano, William Ratcliff, and Ford Denison (PNAS 2012). His work has been supported by grants from the National Science Foundation. Oren Harman is the Chair of the Graduate Program in Science, Technology and Society at Bar Ilan University and Senior Research Fellow at the Van Leer Jerusalem Institute, where he hosts the public series “Talking About Science in the 21st Century” and the Science and Creativity Group. He was trained in history and biology at the Hebrew University, Oxford, and Harvard, and is a historian of science and a writer. He teaches evolutionary theory, history and philosophy of science, and writing. His books include The Man Who Invented the Chromosome (Harvard, 2004), Evolutions: Fifteen Myths That Explain Our World (Farrar, Straus, and Giroux, 2018), and the coedited trilogy, with Michael R. Dietrich, Rebels, Mavericks and Heretics in Biology (Yale, 2008), Outsider Scientists (Chicago, 2013), and Dreamers, Visionaries and Revolutionaries in the Life Sciences (Chicago, 2018). His book The Price of Altruism (W.W. Norton, 2010) (Bodley Head/Random

About the Editors

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House, 2010) won the 2010 Los Angeles Times Book Prize for Best Book of the Year in Science and Technology, was nominated for the Pulitzer Prize, and was a New York Times Notable Book of the Year. He is currently working on a book about metamorphosis.

Contributors

Warwick Anderson University of Sydney, Sydney, NSW, Australia Rachel A. Ankeny University of Adelaide, Adelaide, SA, Australia Ana Barahona Universidad Nacional Autonoma de Mexico, Mexico City, Mexico Dominic J. Berry London School of Economics, London, UK Mark E. Borrello Program in the History of Science, Technology and Medicine Department of Ecology, Evolution and Behavior, University of Minnesota, Minneapolis, MN, USA Janet Browne Harvard University, Cambridge, MA, USA M. Chirimuuta Department of Philosophy, University of Edinburgh, Edinburgh, UK Nathan Crowe History Department, University of North Carolina Wilmington, Wilmington, NC, USA Michael R. Dietrich Department of History and Philosophy of Science, University of Pittsburgh, Pittsburgh, PA, USA Oren Harman Graduate Program in Science Technology and Society, Bar-Ilan University, Tel Aviv, Israel Andi Johnson History and Sociology of Science, University of Pennsylvania, Philadelphia, PA, USA Hannah Landecker UCLA, Los Angeles, CA, USA Sabina Leonelli University of Exeter, Exeter, Devon, UK Kate MacCord Marine Biological Laboratory, Woods Hole, MA, USA Jane Maienschein Arizona State University, Tempe, AZ, USA Marine Biological Laboratory, Woods Hole, MA, USA Michel Morange IHPST, Université Paris I, Paris, France xiii

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Contributors

Samantha Muka Stevens Institute of Technology, Hackettstown, NJ, USA Staffan Müller-Wille University of Cambridge, Cambridge, UK Nicolas Rasmussen University of New South Wales, Sydney, NSW, Australia Marsha L. Richmond Wayne State University, Detroit, USA Neeraja Sankaran Independent Scholar, Bangalore, Karnataka, India Edna Suárez-Díaz Estudios de la Ciencia y la Tecnología, Facultad de Ciencias, Universidad Nacional Autónoma de México, Mexico City, Mexico Tracy Teslow University of Cincinnati, Cincinnati, OH, USA Paul Weindling Oxford University, Oxford, UK

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New Perspectives on the Historiography of Biology Michael R. Dietrich, Mark E. Borrello, and Oren Harman

Abstract

The Handbook of the Historiography of Biology is intended to foster a conversation about the historiographic traditions that have informed the history of biology. Explicit historiographical reflections by leading scholars in the history of biology will highlight important trends and innovations in the continuous stream of original research that has created this field. This will make it easier for new scholars to join the field and make their own original contributions. The German-born American historian Fritz Stern pronounced that “the historian must serve two masters – the past and the present,” and reminded us that “perhaps no one has changed the course of history as much as the historians” (Stern 1973). In his classic book, What Is History? the English historian E.H. Carr was even more blunt: “Study the historian before you study the facts” (Carr 1962), he admonished. Clearly, history never stands on its own. It is always constructed, filtered, placed within the context of what those who came before believed and wrote. With time, history becomes a palimpsest. To understand how successive generations have remembered the past, one must drill down, layer by layer. As a study of the science of history, historiography attempts to understand how historians work, how they frame their questions, how they use sources, and how historical scholarship reflects its different contexts. If history is a sign of its times, M. R. Dietrich (*) Department of History and Philosophy of Science, University of Pittsburgh, Pittsburgh, PA, USA e-mail: [email protected] M. E. Borrello Program in the History of Science, Technology and Medicine Department of Ecology, Evolution and Behavior, University of Minnesota, Minneapolis, MN, USA e-mail: [email protected] O. Harman Graduate Program in Science Technology and Society, Bar-Ilan University, Tel Aviv, Israel © Springer Nature Switzerland AG 2021 M. R. Dietrich et al. (eds.), Handbook of the Historiography of Biology, Historiographies of Science, https://doi.org/10.1007/978-3-319-90119-0_3

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historiography is like a geological expedition, allowing us to reconstruct the landscape and set of circumstances that helped produce its fossils. Historiographic analysis is valuable because as historians create their narratives describing the past, they are in dialog not only with their sources but with other historians, other historical narratives, wider social, cultural, and political changes, as well as shifting scholarly standards and methodologies (Christie 1990; Nyhart 2016). Every historian should be aware of the place of her narrative in its historiographic lineages, but those historiographic traditions are articulated and analyzed less frequently on their own terms. Still, some areas of history have extensive historiographic literatures. In 2020, the Blackwell Companion series, to provide just one example, has 72 volumes related to American history alone. While one of them is devoted to the history of American Science (Montgomery and Largent 2015) and Blackwell also has A Companion to the History of Science (Lightman 2016), by comparison to American history, the history of science, and especially the history of biology, have relatively few historiographic resources from which to draw (see, however, Abir-Am 2006, Creager 2017, de Chadarevian 2009, Finnegan 2008, Gilbert 1998, Greene 1971, Hopwood 2019, Jardine 2003, Kragh 1987, Löwy 2011, Paul 2016, Smocovitis 1996, Söderqvist and Stillwell 1999, Tanghe 2017). The Handbook of the Historiography of Biology is intended to foster a conversation about the historiographic traditions that have informed the history of biology. It is our belief that explicit historiographical reflections by leading scholars in the history of biology will highlight important trends and innovations in the continuous stream of original research that has created this field. This will make it easier for new scholars to join the field and make their own original contributions. History of biology is a relatively new field, but this doesn’t mean that it does not have rich historiographic traditions. While the Journal of the History of Biology was founded in 1968 (Dietrich 2017), histories of biology were being written earlier. The great span of biological inquiry was swept up in the synthetic histories of Charles Singer (1931) and Erik Nordenskiold (1946). As they do today, biologists frequently turned to history as they recorded narratives of their own fields, both in standard nonfiction and memoir form (Mayr 1982; Needham 1934; Sturtevant 1964; Watson 1968). The postsputnik expansion of science in the United States had a corresponding effect on the history of science. By the 1960s, a generation of historians of biology was coming of age, often trained at Cambridge University or at Harvard’s history of science program where Everett Mendelson founded the Journal of the History of Biology. The histories that these historians of biology produced were usually “internalist”: focusing on ideas, their origins, and their reception (see Shapin 1992). Reflecting on his own landmark textbook, Life Sciences in the Twentieth Century (1975), Garland Allen remembers that at the time, “There was virtually no institutional history, no sociological analysis . . ., virtually no economic analysis regarding the funding of biology, and no discussion of how social movements . . . influenced the development of biology.” (Allen 2016, 737). Although history of biology has never abandoned the history of ideas, from the 1970s, it raced to follow new developments in the history of science and an explosion of new biological research, often struggling to catch its breath.

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The concomitant rise of a generation of sociologists of science who often championed a constructivist view of science fed an older historiographic split between internalist and externalist approaches in the 1960s and beyond (Shapin 1992; Golinski 1998). Accounts of the discovery of nature were quickly supplanted by narratives that used social, political, and cultural contexts to explain the contingent development of scientists, their ideas, their institutions, and their reception (Nyhart 2016; Soler et al. 2014). At the same time, feminists were exposing another set of biases in the early history of science which often enshrined great men, reinforced an association of masculinity and science, and ignored the gendered context for scientific research (Rossiter 1982; Schiebinger 1989). The impact of this turn toward the social aspects of science-making was a tremendous diversification in the kinds of histories of biology that were written. Among the many social and political contexts associated with the rise of modern biology, historians began to draw significant connections between biology and business, whether in the form of agriculture or later biotechnology and genomics (Cook-Deegan 1995; Phillips and Kingsland 2015). Others focused on the politicization of biology in such infamous episodes as the rise of Lysenkoism in the Soviet Union (Graham 2016), or the vagaries of changing concepts of race (Barkan 1992). Still others traced the rich interconnections between biology and its social and cultural context, such as Richards account of biology and the German Romantic tradition (Richards 2002). As historians moved beyond the history of ideas, they began to broaden their view of science itself. Where earlier accounts focused heavily on scientific theories and how they changed, the growing recognition of experimental and material dimensions of biological research fostered new ways of describing what it was that biologists did. For example, while Alfred Sturtevant’s history of Drosophila genetics described life in the “fly room,” his account is largely of personalities and conceptual developments, whereas Robert Kohler’s account of the same history focuses deeply on how flies were created as technologies in the daily life in the fly room that was governed by a moral economy that regulated the course of research (Kohler 1994). Kohler’s account directly engages in the materiality of research and the social and institutional dynamics that informed the conditions for the group’s success. Of course, not all biology occurs in a laboratory and it is significant that Kohler’s next book followed biologists into the field to explore biological practice in that context (Kohler 2002). The result of this turn to practice has been a new appreciation of materiality, instrumentation, situatedness, and the work of creating biological operation, execution, and implementation (Kuklick and Kohler 1996). Focus on the practical work of biology inevitably becomes spatial. Kohler followed the transit of flies from lab to lab and practices from lab to field. This attention to travel and circulation of people and materials also draws on a growing appreciation of the global nature of science. Histories of Western science have been countered with histories of different regions, such as Lisa Onaga’s history of silkworm biology in Japan or Ana Barahona’s history of genetics in Mexico (Onaga 2010; Barahona 2019). Indeed, George Basala’s metaphors of a Western core of science and colonial peripheries have been replaced by much more sophisticated accounts of circulation and

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transformation of biological practices (Basalla 1967; Fan 2012; Raj 2013). And when biographers of biologists turn to their subjects, they have increasingly replaced “Great Men” accounts the likes of which George Sarton championed, with nuanced ancilla historiae, both of women and men, groups and even objects (Comfort 2003; Daston 2000; Harman 2010; Werskey 1979). The growth and diversification of history of biology is a sure sign of its vitality as a field. And yet the expansive literature now representing the history of biology is daunting. The Handbook of the Historiography of Biology is intended to provide an assessment of the historiography and the future of major subfields and approaches within the history of biology. Each chapter addresses major trends in historical thought within a particular subfield or approach, major debates among historians of that subfield, and promising new directions that may shape future scholarship. Each chapter is framed in terms of what a scholar should know about the history of work in that subfield, if they wanted to make a meaningful and original contribution to that subfield. Written for graduate students and other scholars new to the history of biology, The Handbook of the Historiography of Biology is not intended to provide a comprehensive review of every topic discussed in the history of biology. In developing our list of contributors, we matched topic areas to scholars with established expertise in that area. Of course, different authors will select different exemplars of scholarship and highlight different themes. Each author in this collection offers their particular perspective on the historiography of their subfield or topic area. Because we are seeking deep and broad but not necessarily exhaustive literature reviews, each chapter reflects the author’s judgment of the field and the trends that she or he has found most significant. Different authors undoubtedly would produce different perspectives on the same field. We do not view this as a shortcoming, but an opportunity to engage with each other about how the history of biology has been and will be written. This collection is intended to support a conversation among historians of biology regarding their work, its history, and its future. We have selected topic areas that we believe have a substantial and diverse body of scholarship. Because these chapters are not literature reviews, but historiographic essays, we selected fields informed by different methods, theories, and perspectives that can be compared and contrasted in each chapter. Many of the chapter topics that we have selected correspond to biological fields and traditions, such as biotechnology, immunology, marine biology, physiology, plant breeding and agriculture, or eugenics. Those fields with very extensive historical literatures are subdivided; for instance, the literature on Darwin is separate from the literature on the history of modern evolutionary biology. We have also included chapters organized by genre and historian’s topics, such as biography and women in biology. These chapters cut across biological fields but address a coherent body of scholarship devoted to their topic. We include these chapters because we think that the debates over the place of biography in history, for instance, are important, and that biography as a genre is developing in significant ways, and not about to disappear from the history of biology. In a similar fashion, scholarship on women and biology has undergone significant changes under the influence of feminist science studies that we thought were important to have presented for future scholars.

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This handbook is far from complete. There are fields that we would like to see discussed, such as the historiographies of natural history and botany or more in-depth treatments of the biological sciences in antiquity, the early modern era, and the eighteenth century. Unfortunately, we were not able to successfully solicit these contributions for this edition. We hope that this situation will be short-lived, however. Springer has generously agreed to allow this volume to expand in subsequent editions. So rather than see this as a fixed statement about the historiography of biology, we offer this volume as the beginning of a conversation. We hope that scholars will see the value in a collected historiographical resource and will contribute essays on different historiographies for future editions.

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Charles Darwin and the Darwinian Tradition Janet Browne

Contents Hero of Science, 1880–1959 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Archival Rapture . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Reconstructing Darwin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Common Contexts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Religion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Deconstructing the “Revolution” . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Social Darwinism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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Abstract

The chapter describes how historians have shifted from studying Darwin as an individual thinker to embrace a more panoramic cultural picture. Darwin’s rich archival record now reveals a great deal about nineteenth–century scientific practice. Fresh perspectives on evolutionary history and the so-called Darwinian revolution emerge by decentering Darwin. It is sometimes said in history of science circles that “Charles Darwin has been done to death” and that the field needs to move on. The criticism in a way is justified – although this chapter will propose that there are plenty of exciting and innovative projects still to explore. There was a time when the phrase “the Darwin industry” was also expressed as a criticism of a body of scholarly work that focused intensively on the manuscript archive (Ruse 1974, 1996). Frequently it was difficult for outsiders to see the larger historical purpose of many of these studies, and certainly some of the

J. Browne (*) Harvard University, Cambridge, MA, USA e-mail: [email protected] © Springer Nature Switzerland AG 2021 M. R. Dietrich et al. (eds.), Handbook of the Historiography of Biology, Historiographies of Science, https://doi.org/10.1007/978-3-319-90119-0_4

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close analyses of manuscript deletions and additions in Darwin’s private notebooks were not for the fainthearted. Yet the old Darwin is emphatically not the current Darwin. Times have changed. This chapter mostly concerns the scholarship that has emerged since the late 1980s to today, a period of about 40 years during which our ideas about history of science have themselves been transformed (Hodge and Radick 2009). The chapter does however reach back beyond that date to describe previous traditions because Darwin has always been a prominent focus for scholarly research. Essential foundations were laid down even as far back as the 1880s, according to the best practices of the day. In fact, an account of the historiography of Charles Darwin as a thinker and writer, and of the Darwinian tradition, reflects the emergence and changing structure of history of science in a manner that is well worth serious study in its own right (Churchill 1982; La Vergata 1985). For general access to everything one might wish to know about Darwin and the Darwinian tradition, an excellent summary can be found in Richard Milner’s Darwin’s Universe: Evolution from A to Z (Milner 2009) or in Bowler (1984) and Larson (2006). Two important websites have revolutionized the field: the Darwin Correspondence project (https://www. darwinproject.ac.uk/) and The Complete Work of Charles Darwin Online (http:// darwin-online.org.uk/). A great deal of scholarly research over these last four decades has established that Charles Robert Darwin (1809–1882) was not the only significant evolutionary thinker in the past and that the intellectual and social movement called “the Darwinian revolution” was not a revolution in the usual sense and hardly due to Darwin alone. The evolutionary movement was much broader-based than previously assumed and several simultaneous lines of thought and sociopolitical trends (especially industrialization and colonization) combined across the globe during the eighteenth and nineteenth centuries, creating conditions in which evolutionary ideas of many different hues could flourish, at first in Europe and then elsewhere. Darwin’s ideas therefore emerged within a diverse body of thought. They were not completely successful at the start, although On the Origin of Species, published in 1859, had a tremendous public effect. Moreover, as time went by, it was commonplace for contemporaries to combine Darwin’s ideas with other ideas of progress and change. So the route by which evolutionary ideas crystalized into a so-called “Darwinian” triumph is now one of the most useful questions that scholars in the history of biology can seek to answer (Ruse 1999; Smocovitis 2005). In general terms, modern evolutionary theory explains so much about the natural world, and the human condition, that historians need to be careful not to overemphasize those elements in the evolutionary story, such as natural selection, that hold central positions in today’s science. Because Darwinism is viewed as a success story, it is easy to fall into the trap of calling other evolutionary thinkers “precursors” or “successors” of Darwin when they were actually working in different frameworks. Indeed many modern biologists now consciously call their daily working theory “evolution” rather than Darwinism, primarily in order to recognize the dissimilarities in thought between then and now, but also perhaps to acknowledge features that would once have been thought non-Darwinian, such as epigenetics. In

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North America, it may also be a smart move to separate the modern theory from the historical figure of Darwin who is a central target for the creationist movement (Ruse 2006). Practicing biologists often contribute to the Darwinian literature with genuine understanding of the achievement, as discussed below. To think of the Galápagos Islands as a site of discovery or as a laboratory for evolution (Larson 2001), for example, is reflected in the important work on biological adaptation and divergence carried out there by Peter and Rosemary Grant (Weiner 1994). There is a satirical side to modern evolutionary biology, as is apparent in cartoons, blog texts, rap, songs, costume dramas, and imagery on the web. Lastly, whether to teach Darwinism has also been the subject of legal action in the courts and remains a hot topic in educational policy. Hence, evolution inspires public interest at a level scarcely seen in other areas of history of science. This adds special value and relevance to historical studies about its impact. New scholars in this area will therefore join a community of historians who give due weight to Darwin as a pivotal historical figure but are also keen to engage with issues relating to socioeconomic, religious, cultural, and political structures, not only in the nineteenth and twentieth centuries but also to participate in today’s concerns. One quick way of characterizing this approach is to say that historians of science are changing their focus from the individual to the collective, from treating Darwin as a single dominant figure in biology to embracing a more panoramic picture that takes account of wider historical trends and transformations. These wider historical trends might range from gender studies to literary analysis, visual representation, globalization, celebrity and reception studies, popular science, commemorations, the philosophy and metaphors of science, empire and race, social-network theory, scientific mythologizing, the consolidation of an acceptable canon of ideas, and so forth. The literature is sufficiently rich to provide opportunity to engage in the developing field of meta-history. At the same time, there are important continuing studies to be made in the relation of Darwinism to religious thought, in social Darwinism, eugenics, the history of anthropology, linguistics, and beyond.

Hero of Science, 1880–1959 Today’s historians enter a field that possesses several significant research legacies. The first studies of Darwin’s life and work were already begun by the time of his death in 1882. Writers and commentators at that time wished to show that science primarily advanced by intellectual breakthroughs. To praise Darwin seemed unproblematic to his contemporaries. Even before Darwin’s death, his friends were calling the evolutionary movement “Darwinism” (Wallace 1889). Early responses to Darwin as a historical figure were strongly connected to the emerging archival record. Darwin’s son Francis edited an unusually full threevolume Life and Letters (Darwin 1887), which included family recollections of Darwin’s daily life, a shortened (and expurgated) version of Darwin’s manuscript

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“Recollections of the development of my mind and character,” and a stirring propaganda piece written by Darwin’s close friend Thomas Henry Huxley about the early reception of the Origin of Species. Francis Darwin published more correspondence with the assistance of A. C. Seward (Darwin and Seward 1903) and transcripts of Darwin’s early drafts of the theory of natural selection (Darwin 1909). Henrietta Litchfield, Darwin’s eldest living daughter, prepared two volumes of her mother’s family letters (Litchfield 1915) that provide much social detail. Recollections by Darwin’s second son, George, and fourth son, Leonard, and a memoir by George’s daughter, Gwen Raverat (1952), added to the story. The family was thus active in producing and shaping Darwin’s public image in which he was cast as a gracious “gentleman” of science, anxious not to offend contemporary sensibilities, and considerate to those among his community who believed in a divine entity . The framing mostly took shape through the hands of Francis Darwin (as the editor of his father’s letters), who removed at his mother’s urging several sentences from Charles Darwin’s Recollections that referred to Christianity in critical terms (Barlow 1958, 11–12). But there was more than this. The overall thrust of the Life and Letters sought to show Darwin dedicated to observational science, struggling with ill health, and greatly loved by his friends and family. Although the Darwin family has been studied in its own right as a prominent intermarrying coterie in intellectual British society (Wedgwood and Wedgwood 1980; Berra 2013), and as part of the “intellectual aristocracy” (Annan 1955), it has not yet been studied as an interested party in the creation of a hero of science. The published documents at that time were more or less taken at face value by researchers. But in 1958, Nora Barlow (another Darwin granddaughter) published several editions of Darwin’s manuscripts that expressed her interest in the creative processes of science, including her grandfather’s first musings about evolution during the Beagle voyage and his reflections on his loss of religious belief. Her publication of the full text of Darwin’s “Recollections” (usually now called his autobiography) caused a minor sensation in revealing religious views that had previously been carefully manicured in Francis Darwin’s Life and Letters – Darwin even in one place expressed contempt for Christian doctrine (Barlow 1958). Barlow’s work needs to be given due prominence in the history of science for engaging with the postwar cultural shift toward exploring the private person. Much more, however, remains to be done on the role and impact of autobiographies in science and on issues relating to the construction of a scientific life (Soderqvist 2007; Shapin 2008). Darwin’s self-writing is an excellent place to start. The continuing family investment in curating Darwin’s legacy is worth noting. Barlow subsequently published Darwin’s Beagle diary and ornithological notes (Barlow 1963). Richard Darwin Keynes (a great-grandson) published Darwin’s zoological notes and specimen lists from the Beagle voyage (Keynes 2000). Randal Keynes (a great-great-grandson) published a study of Darwin’s family life called Annie’s Box (Keynes 2001) and advises a number of contemporary online digitization and educational projects relating to Darwin (Keynes 2016).

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Archival Rapture In the 1940s, an enormous collection emerged, including Darwin’s handwritten notes, letters, drafts of his writings, and his personal annotated research library of books and articles. The Darwin family passed this material to Cambridge University Library, UK, with financial assistance from the Pilgrim Trust, an American charitable body (Burkhardt et al. 1985a). A substantial collection was also established at the American Philosophical Society in Philadelphia, and there are notable smaller collections in other locations. The abundance, variety, and historical wealth of these records are stunning. Archival material in general is usually preserved in much smaller caches, sometimes even as single documents, with component parts often distributed across an institution according to accessioning practices or only to be found in separate institutions. Remarkably, the Darwin collection is nearly entire and all in one place: a point that goes a long way toward explaining the effect it had on scholarship. Not all of the papers were available immediately. The ball began rolling in the 1960s with a transcription by Gavin De Beer, director of the Natural History Museum in London, of Darwin’s transmutation notebooks. During his lifetime Darwin excised many of the most important notebook pages in order to use them for his continuing research, and De Beer had to work with incomplete materials. Some excised pages were located in the 1960s and the remainder emerged safe and sound in the 1980s, in a “Black Tin Box” that had remained in the family possession. The piecemeal publication of such significant documents, and the variable quality of De Beer’s transcriptions, provided strong rationale for a completely new transcription and integration of materials in one comprehensive volume in the 1980s (Barrett et al. 1987). This new transcription, accompanied by full scholarly annotations, is now the gold standard for understanding Darwin’s notebook reflections and theorizing. All of a sudden, it was possible to follow the development of Darwin’s reasoning, his excitement, fears, and steady work drawing on many different sources, and to acknowledge that he grasped the nettle of evolution some time before he developed the idea of natural selection. Scholars went wild with transcriptions, sometimes redoing what the family had previously done, other times locating fresh and unsuspected materials, such as Darwin’s early (and unsuccessful) attempt to explain heredity and variation (Olby 1963). Such restorations aimed to illuminate Darwin’s theory building, reflecting a developing historiographical concern to show science as a hard-won construction process rather than as a succession of inspired moments, although these could be discerned as well. Between 1970 and 1990, there was a steady flow of reevaluations of hot topics that transformed what scholars could hope to know about Darwin’s scientific creativity (Kohn 1989), as well as Darwin’s first encounters with evolutionary ideas, his education, literary tastes, fieldwork (Rudwick 1974), illnesses (Colp 1977), cultural circle (Manier 1978), and religious views (Gillespie 1979). The word “evolution” itself was examined (Bowler 1975). New studies emerged of Darwin’s botanical work and biogeographical thinking and of key concepts like

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his theory of divergence. Close examination of Darwin’s first few months in London after returning from the Beagle voyage led to a significant reevaluation of the role of the Galápagos finches in his thinking (Sulloway 1982). Important publications included the long manuscript from which Darwin abstracted the text of the Origin (Stauffer 1975) and a comprehensive checklist of Darwin’s working library with his annotations (Di Gregorio 1991). Fresh assessments of Darwin’s early engagement with human evolution were made (Herbert 1974–77), his ideas about the human mind (Gruber 1974), his early focus on reproduction (Hodge 1985), and understanding of European embryology (Ospovat 1981). A conference in 1982 explored all the scholarly results generated by this archival enterprise (Kohn 1985). A survey of these developments is given by David Oldroyd (Oldroyd 1984). A special aspect of this period in history of science circles was the dramatic upturn of scholarly interest in natural history and fieldwork. Darwin’s collecting on the Beagle voyage came under detailed investigation (Porter 1985). It was also realized that Darwin’s geological preoccupations were deeply significant to the development of his views (Rudwick 1974; Secord 1991; Herbert 2005; Sponsel 2018). And in 1974 a major project to publish all of Darwin’s surviving correspondence was initiated, taking physical shape in 1985 with a printed calendar of the known correspondence (Burkhardt et al. 1985a), followed by the first volume of the print edition of the correspondence (Burkhardt et al. 1985b). This is a scholarly edition of some anticipated 30 print volumes, with an associated digital site (https://www. darwinproject.ac.uk/). The project not only reveals Darwin as a deeply engaged scientific figure, with much previously unknown material becoming apparent, but it also marks a significant shift toward using the Darwin archive as a resource for understanding more than science itself in the nineteenth century. The impact of this scholarly edition can hardly be overstated. Among many other things, it shows that letters comprised a significant part of scientists’ research methodology and served for the circulation of ideas. A more recent project to place all Darwin’s manuscripts online with new transcriptions and bibliographical sources is under way at the American Museum of Natural History (Kohn 2015). With new archives to explore from the 1960s, a number of medically qualified scholars, who were frequently also practicing physicians or psychiatrists, offered their interpretation of Darwin’s illnesses and, as often as not, a retrospective diagnosis. There is hardly any parallel in other aspects of the history of science, and this is a fascinating area for any historian interested in the cultural framing of disease according to the professional expertise of the practitioner. Ralph Colp’s study (1977, updated 2008) is the definitive work here, providing all the known archival records and a thorough review of the diagnoses offered up to the date of publication. Some diagnoses focus on physical disorders such as Chagas disease, possibly contracted during the Beagle voyage (Adler 1959; Colp 2000). This has never been completely refuted as an option. Other diagnoses offer subclinical psychiatric conditions such as panic disorder or hyperventilation (Bowlby 1992). Darwin has been diagnosed as possibly suffering from multiple allergy syndrome, hypoglycemia, tinnitus, lactose intolerance, cyclic vomiting syndrome, infection with Helicobacter pylori, and chronic arsenic poisoning. This is an area that invites a careful meta-study. First, it

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could be asked why it might be useful to know what Darwin had and who wants to know? And second, it is a moot point how this illness (or illnesses) might bear on Darwin’s scientific productivity. Most authors appear to subscribe to the concept of “creative maladies” (Pickering 1974). This issue again reminds us that historical modes of investigation change over time and that the questions we ask of science’s protagonists often emerge from our own concerns. Only a handful of historians have looked at Darwin’s actual encounters with Victorian medicine (Browne 1990, 1998) or the influence of Darwinian ideas in medical practice during the last third of the nineteenth century (Bynum 1983). Today, there is significant interest in a Darwinian medicine that is concerned with explaining sickness and issues such as aging in evolutionary terms when being healthy would (to us) seem like an adaptive advantage (Nesse and Williams 1995).

Reconstructing Darwin The 1950s brought a second body of commentators to the field who had different questions to ask: questions primarily stimulated by the prominence of the new genetics and the prospect of upcoming commemorations of the sesquicentennial of Darwin’s birth combined with the centenary of publication of the Origin of Species. By this time, scholars were eager to integrate the emergence of Mendelian genetics around 1900 with the history of evolutionary theory. The history of science itself was also becoming professionalized, with the first of several departments or programs initiated in London (University College London), Indiana (Bloomington), and Massachusetts (Harvard). Two unusual books by an unusual scholar, Loren Eisley, bucked the existing archival trend. Eisley published Darwin’s Century (Eiseley 1958) in order to reveal the other people interested in evolutionary ideas, followed by Darwin and the Mysterious Mr. X, that described the rival ideas of the little-known naturalist Edward Blyth (Eiseley 1979). John Burrow and John C. Greene wrote accessible, wide-ranging accounts of the development of evolutionary ideas in general (Burrow 1966; Greene 1959). The neoconservative historian, Gertrude Himmelfarb, published a widely read account of the Darwinian revolution that mentioned some of the early difficulties of accepting natural selection theory (Himmelfarb 1959). She was criticized for disparaging science. Another contentious book, written more than 10 years earlier, and reissued in 1955, in time for the Darwin commemorations, was by Richard Hofstader, Social Darwinism in American Thought, that attacked the “dog-eat-dog” principles that lay at the heart of social Darwinism (Hofstadter 1944). Some indispensible tools were published at this time, too, including a fresh bibliography of all Darwin’s writings (Freeman 1977) and a variorum edition of the Origin of Species that lists all the changes made by Darwin through six editions, sentence by sentence (Peckham 1959). Both of these are now available electronically on darwin-online.org.uk. More specifically, a major American celebration of Darwin’s achievement was mounted in Chicago in 1959 (Smocovitis 1999), matched by other commemorative activities around the globe that have not yet been fully studied. The Chicago meeting

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aimed to make the fact of evolution clear to the high-school biology teachers who were the intended audience. It also showcased the synthesis of the science of genetics with the evolutionary ideas agreed by biologists during the previous 15 years or so. This “evolutionary synthesis” united genetic principles at the level of the cell, with the variability, selection of traits, adaptation of organisms, and gene flow in populations, demonstrating that Darwinian notions were a key unifying law of nature. In Chicago, it appeared to be an appropriate moment to display unity around Darwin’s ideas and to call on him as the founding father of the new biology, one in which human kind was emphatically included. A number of professionally trained academic biologists simultaneously began to contribute to the field. The evolutionary biologist Ernst Mayr epitomized the best (and occasionally the worst) of the biologist-historian tradition: he rewrote the history of biology to accommodate fundamental binary divisions in philosophical approach and praised Darwin as a unique figure standing between early theorists concentrating on ideal Platonic forms (whom Mayr thought were misguided) and those modern biologists who understood that evolution was all about populations, such as himself. Mayr’s definition of species rested on biological isolation, echoing the early drafts of Darwin’s theory. He held Darwin in high regard and was an accurate scholar (Mayr 1982). Though there are accounts of Mayr’s important role in modern biology (briefly in Barrow 1998), there is not yet a full examination of his impact as a historian on Darwin studies. Similarly, the paleontologist Stephen Jay Gould wrote with great effect in both history and biology and developed a substantial media presence. He disseminated evolutionary ideas through his columns in the American Museum of Natural History’s journal Natural History from 1974, in which he discussed points of view derived from Darwin’s life and writings, often opening his column with a historical anecdote (a number of volumes of collected essays were published, from Gould 1977 to Gould 1991). He called his column “This View of Life,” a quotation from the closing lines of Darwin’s Origin of Species: “There is grandeur in this view of life, with its several powers, having been originally breathed into a few forms or into one.” Gould described himself as a Darwinian, and yet he was a Darwinian of a very individual stripe (Sheldon 2014; Sepkoski 2008). He became an influential figure in the creationist disputes in North America in the 1980s, serving as a witness for science in the McLean v. Arkansas case of 1981 (Gould et al. 2006). Warm links to Darwin as a historical figure can be seen in the writings of other modern biologists who at times address the historical record, such as Richard Dawkins. Indeed this is a continuing aspect of the literature relating to Darwinism. There are many modern scientists who have an historical bent or are hybrid historianphilosopher-biological theorists. Some, such as Eva Jablonka, look back to Darwin to see where he stood on inheritance (Jablonka and Lamb 2005). Others regard themselves as hard-line neo-Darwinists (discussed elsewhere in this volume). And there are a number of attempts to bring Darwin up to date, several of which capitalize on a seemingly inexhaustible publishing phenomenon, while others attempt to show how Darwin as an author has shaped – and still shapes – modern evolutionary thinking (Jones 1999; Costa 2009; Reznick 2010).

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Common Contexts During the 1960s and 1970s, vigorous new trends in the professional discipline of history introduced fresh ways of approaching the history of science. Economic history, social history, reception studies, prosopography, and Marxist, Mertonian, and Weberian analyses came into play. In the history of science, these approaches were often expressed in terms of “internal” or “external” forms of analysis, where internalists emphasized the trajectory of ideas and externalists emphasized the social and cultural forces that produced and molded such ideas. To rising “externalist” scholars, the sociology of scientific knowledge (SSK) usually also included a commitment to the social construction of scientific facts. As far as Darwin studies were concerned, this was expressed most clearly in the influential work of the Marxist historian Robert Young, based in Cambridge UK. Young stimulated a fundamental change of direction by insisting that Darwin was best understood as a member of the elite classes of Victorian Britain and that his theories manifested, either more or less, the individualistic, capitalist economy of the industrializing nation. Scientific theories, he argued, were an integral part of wider intellectual, economic, religious, and political thought in Victorian Britain, a framework that he designated as a “common context” (Young 1969, 1985b). This was a far-reaching affirmation that nineteenth century thought cannot (and should not) be divided up into twentieth century disciplines. The very heart of Darwin’s theory, Young said, lay in the work of the political economist T.R. Malthus, from whom Darwin developed the idea of natural selection. Darwin’s direct and vivid engagement with Malthus has since served as an excellent case study of the intermeshing of science with society. “Darwinism is social” was a phrase often used by Young that became talismanic through the 1980s (Young 1985a). Young’s work on Darwin is not as much read as it once was. Nevertheless he fostered a group of remarkable historians of science from the 1970s, and his impact at an important moment in the rise of history of science as a field is highly rated (Bohlin 1991). His influence can be seen most explicitly in the social-constructionist biography of Darwin co-authored by Adrian Desmond and James R Moore (1991). One feature of this study is the proposal that Darwin delayed publishing his theory until the social and political circumstances of Victorian England were relatively placid, rather than publishing in the radical 1840s, when he first wrote it out in full. The question of whether such delay was deliberate, or even a delay at all, has been critiqued by John van Wyhe (2007). These decades of expanding historical vision fostered other trends in Darwinian studies from the 1980s, including the literary and visual. Scholars of Victorian literature began to explore science and culture as interrelated ways of knowing the world. Gillian Beer examined Darwin’s writing style and tracked the structural relationships between Darwin’s Origin of Species and literary works by novelists, primarily George Eliot (Beer 1983). Beer’s interpretations are not by any means the last words to be offered in this domain (Lightman 2007; Thurtle 2007; Gianquitto and Fisher 2014). In the United States, George Levine reflected on science as a deeply imaginative enterprise in Darwin and the Novelists (Levine 1988). These

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studies touched on the divergence between esthetics and science in the modern era. More recently Gowan Dawson has explored the problem of social respectability for Victorian readers of Darwin’s books (Dawson 2007). Darwin’s impact on the artistic and visual realm has also made waves in wider history of science scholarship. Jonathan Smith wrote comprehensively on Darwin and Victorian visual culture (Smith 2006), Julia Voss discussed the illustrative material that Darwin employed in his own writings (Voss 2010), evolutionary imagery and caricatures have been explored (Clark 2008; Browne 2001), Philip Prodger made a definitive study of the photographic archive that Darwin used for his work on the expression of the emotions (Prodger 2009), and two stunning exhibitions, “Endless Forms” (Donald and Munro 2009) and “Darwin: Art and the Search for Origins” (Kort and Hollein 2009) celebrated change and transformation in artworks of the period. Other notable trends in the literature show Darwin as engaging in detailed scientific work, often supported by exchanges with knowledgeable persons outside the modern category of “scientist.” An often cited example is Darwin’s exploration of the genealogy and breeds of pigeons that led him to discuss the topic with bird breeders and fanciers (Secord 1981). Darwin researched the systematics of barnacles (Ghiselin 1969; Stott 2003) and paid lifelong attention to plant fertilization, hybridity, and movement, discussing these with gardeners, colonial botanists, and many others, especially his close friend Joseph Dalton Hooker (Allen 1977; Ayres 2008; Endersby 2008; Bellon 2011). He valued female informants and used his family and friends as sounding boards for his ideas (Harvey 2009). This flourishing thread in Darwin studies originates in the archive, generating a much more realistic picture of Darwin at work and a stronger sense of the social construction of science (Browne 1995, 2005). Nowadays, it is frequently said that Darwin used his house and garden as his “laboratory” – a metaphor explored in De Chadarevian (1996).

Religion Usually regarded as the centerpiece of any inquiry into the Darwinian tradition, the question of the relationships, accommodations, and clashes between religion and evolutionary science, both then and now, is notably complex (Brooke 1991). Darwin’s personal beliefs have been widely documented and analyzed in a range of media outlets, including popular texts, journalism, religious studies, intellectual history, social history, the internet, and of course history of science. To some degree, this enterprise has gone as far as it can unless more documentary material turns up. A comprehensive account, including relevant quotations from Darwin’s papers, was given by Gillespie (1979). Some scholars center the onset of Darwin’s disbelief on the death of his daughter in 1851 (Moore 1989; Keynes 2001). The same view is adopted in the commercial cinema dramatization of Darwin’s life “Creation” in 2009. Moore provides an excellent social study of Darwin’s funeral in Westminster Abbey, focusing on the paradox of Darwin’s being lionized as a good man and yet the author of Origin of Species (Moore 1982).

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The role of natural theology in Darwin’s thoughts and the wide cultural influence of the theological author William Paley (Ospovat 1981; Fyfe 1997; Eddy and Knight 2005) are sometimes muddled in readers’ minds by Richard Dawkins’s best-selling The Blind Watchmaker, a book that addresses Paleyite design and modern evolutionary adaptationism (Dawkins 1986). One long-standing problem that deserves attention is how far did Darwin believe in progress (teleology), either as a God-given phenomenon or spontaneously arising from the natural selective process. A case has been made for seeing Darwin as the intellectual descendant of the romantic German nature-philosophers, inheriting their progressivist vision of unfolding biological form (Richards 2002, epilogue). It is also possible to view Darwin as following Adam Smith’s doctrine of market self-interest (Richards and Ruse 2016). More general accounts of the Victorian commitment to cultural progress are given by Bowler (1989) and Ruse (1996). It seems clear that Darwin’s remarks have sometimes been appropriated for diverse purposes – exactly what historians like to study. This phenomenon began soon after Darwin’s death with the publication in 1883 of a frank conversation about atheism that Darwin held with Edward Aveling and Ludwig Buchner (1883). The family took steps to publish their own sanitized account of Darwin’s religious views in the Life and Letters. A particularly telling myth has emerged of the purported deathbed conversion of Darwin to Christianity, as reported by Elizabeth Cotton (Lady Hope), an evangelizing speaker who visited Darwin’s home and village (Moore 1994). The creationist website “Answers in Genesis” uses quotations from Darwin to argue that some aspects of evolutionary theory were incredible to him, if not impossible. One particular quotation is often used: “The eye to this day gives me a cold shudder” taken from a letter to Asa Gray, dated by the Darwin Correspondence Project [8 or 9 February, 1860]. The purpose here is evidently to cast doubt on the edifice of naturalistic evolutionary thought: if the founder of the theory doubts it, where then do today’s practitioners stand? The richness of these topics for historical study lies in the fact that something does not have to be true in order to be historically important. The important thing is what people think has happened. The locus classicus for this remains the 1860 British Association for the Advancement of Science meeting in Oxford at which Samuel Wilberforce (Bishop of Oxford) challenged Thomas Henry Huxley over Darwinian theory. Very few written accounts were made of this event, and yet the view of Darwin’s scientific contemporaries was that “science” had roundly defeated the “church” (James 2005). Scholarship has moved beyond these events to explore the wider cultural embedding of religion in science and the causes and effects of their separation. Much that was formerly attributed to responses to Darwin’s Origin of Species can now be seen as being already under way. What is now called scientific naturalism was emerging in the Protestant world long before Darwin, although it evidently experienced a boost with publication of the Origin of Species (Turner 1974; Lightman 1987). It is also clear that although the Protestant (Anglican) faith provided the framework in which most British people operated through the nineteenth century, the grip of the church was loosening (Chadwick 1977; Hilton 1991). However, perhaps it was more the disruption of World War I that called traditional faith sharply into question than

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the Darwinian movement per se. In the nineteenth century, Anglicanism was the state religion, whereby the reigning monarch was the chief defender of the faith, the Anglican church historically owned a large proportion of the nation’s land, from which it claimed income, its bishops sat in parliament and voted on issues of the day, and the ancient universities operated as a powerful arm of the Anglican establishment. University professors at Oxford and Cambridge were required to be ordained as clergymen, although some opted not to take a parish or “living,” as it was called. However, education was nondenominational in Scotland, and in Edinburgh University several professors and teachers aligned themselves with dissent or even expressed non-belief of sorts (Desmond 1984). All of Darwin’s Cambridge University professors were clergymen, and at an early point in his education, he intended to become one too. In England, the first alternative to the Anglican university system was the foundation in 1826 of the nondenominational University College London. Later on, the technical colleges were established in South Kensington, London, including what was to become Imperial College, where Thomas Henry Huxley pioneered secular education and observational biology. Attacks on the Anglican church therefore represented more than just religious grievances: they were intrinsically social and political attacks too. Evolutionary ideas were often adopted by crusaders for reform beyond science. Darwin’s naturalistic approach – one that refused to call on a creator – was thought to support the notion that humankind could be self-determining, that the social hierarchy was not as fixed as the country’s leaders insisted, that a new horizon of intellectual and religious freedom might be glimpsed, and that the old patterns of elitism, wealth distribution, and conservatism might possibly be toppled (Desmond and Moore 1991; Desmond et al. 2007). Movements like these gained immeasurably by evoking the idea of a clash between science and religion. Now known as the “conflict thesis,” this has been a long-lived concept, especially in those nations where science is highly developed. John William Draper and Andrew Dickson White were early exponents of this point of view in the Anglophone world. Draper was a speaker in the British Association meeting of 1860. His book mostly criticized Catholic tradition, then a topical target, but also taking shots at Islam and Protestantism (Draper 1874). His characterization of the history of science as a battle between two contending powers was enormously influential, even finding expression in the positivist historians of the 1950s who believed that the development of science rested on breaking free from the supposed constraints of religious systems (Koyré 1957). In 1896, White published A History of the Warfare of Science with Theology in Christendom, condemning what he saw as restrictive forms of Christianity required by Papal authority. Neither of these books dwelt at any length on Darwinism, but the arguments they offered were attractive to many operating in that system (Moore 1979). The heart of the debates surrounding Darwin’s book lay most obviously in the origin of humankind and the role of the divine in creating the human soul, cognition, and morality. It proved extremely difficult for Victorians to imagine either physical or mental links between apes and humans, and Darwin himself acknowledged some of these difficulties in his book on mankind, The Descent of Man (1871), while yet

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arguing for the reality of the link. A wide variety of scholars have taken up these issues. Some address Victorian concerns about the origins of language (Alter 2002) or morality (Richards 1989). Others explore the controversies over evolutionary anthropology, racial science, or ape ancestry (Moser 1998), and in a variety of national contexts, such as Germany (Kelly 2012), Britain (Stepan 1982), France (Harvey 1983), or the development of primate science (Montgomery 2015) and paleoanthropology (Johanson 2009). Today there is an emphasis on dismantling the imagery of warfare (Lindberg and Numbers 1986; Brooke 1991; Bowler 2001). Biologists, as often as not, were religious believers in some form or another, and, equally, many church leaders came to feel that evolution was true to some degree (Ellegärd 1958; Livingstone 2014). There was in actuality little direct evidence of natural selection, rendering alternative mechanisms equally plausible, if not more so. Several authors after Darwin adopted various forms of evolution that kept a sense of direction in the natural world, often using Lamarck’s ideas of transformisme. In North America, Theodor Eimer proposed a neo-Lamarckian inner force that propelled their evolution in a linear progressive direction. He called this “orthogenesis.” Eimer’s scheme was directional but he believed that natural selection was ineffective as an explanation for change. Another non-Darwinian evolutionary scheme put forward by Edward Drinker Cope depended on embryological development pushing successive generations on to higher forms of organization (Bowler 1983). The English writer Samuel Butler popularized neo-Lamarckism by bringing back into view the inheritance of acquired characteristics through the work of three early evolutionists, Jean Baptiste Lamarck; Goethe; and, Charles Darwin’s grandfather, Erasmus Darwin (Paradis 2007). On the other hand, Asa Gray, Darwin’s main supporter in the United States, adopted the idea of variations arising and being directed in nature by a creative force (for him a Unitarian Godhead) while also accepting natural selection as an entirely mechanistic means for the modification of species (Dupree 1988). Others, such as R. W. Emerson and Moncure Daniel Conway, promoted a creed in which animal origins represented the world of sin that is left behind as religious transformation takes place (Bowler 2007). It was thus equally possible for there to be secular evolutionary thinkers who did not believe in natural selection and Darwinian thinkers who maintained a belief in the divine – and plenty of positions in between. The most famous conflict between science and religion in North America was the Scopes Trial held in Tennessee in 1925. Historical work indicates that this wellpublicized event should be regarded principally as a show trial for civil liberty (Larson 1997). Even though the court determined that John Scopes was guilty, it was not generally regarded as a “victory” for religion. The Broadway play of 1955, followed by the commercial movie “Inherit the Wind,” directed by Stanley Kramer and released in 1960, dramatized political maneuvering behind the scenes and also served as an allegory for the McCarthyism of the day. However, the trial deeply influenced Henry Morris who went on to develop and popularize scientific creationism (Numbers 2006). From the 1960s, new fundamentalist ideologies, principally young-earthism and intelligent design, emerged in North America. Perhaps the phenomenon at the start

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hinged on disillusion with rapidly changing moral values. Today, the rise of contemporary anti-evolutionary doctrines, often highly organized and media-savvy, expresses strongly felt ethnic, geographical, and socioeconomic issues, as well as theological ones (Ruse 2006).

Deconstructing the “Revolution” As already mentioned, major shifts have taken place in recent decades in understanding the Darwinian tradition. These comprise decentering the historical figure of Darwin and emphasizing the work of his contemporaries and other modes of evolutionary thought, particularly doctrines that rested on schemes of progression (Bowler 1988; Ruse 1996). Today we recognize that concepts of revolution should be treated cautiously, that natural philosophy did not emerge only in Europe, and that our vision of the scientific past has dramatically broadened to include all sorts of people, many different geographies, events, and historical trends (Smocovitis 2005). By the time Darwin published Origin of Species, the British nation was full of industrial diversification, commercial and professional specialization, religious tension, broad colonial expansion, and among the middle-classes much talk of “improvement” and “progress.” The self-congratulatory sense of the era was captured by the “Great Exhibition of the Works of Industry of all Nations” held in 1851 in central London, in the glass exhibition hall called the “Crystal Palace.” Few of the visitors to the Great Exhibition would have called themselves evolutionists but many believed in social, commercial, and industrial advance and in a colonial racial hierarchy in which white people were, to them, self-evidently at the top. One of the paradoxes still to be fully explored is the way that transmutationism and early evolutionary views could be discussed in these kinds of social bourgeoise circles and yet also represent a feared transgression from the status quo. Some views advocating social reform could be politically moderate: support for industrial and colonial advance, educational reform, and the extension of suffrage and representation in parliament were often topics of conversation in middle-class society. The anonymous transmutationary book authored by Robert Chambers in 1844 was just such a volume (Secord 2003). Yet, in other circles, to adopt transmutation or to promote points of view that advocated human self-determination, such as the doctrine of phrenology (Cooter 1984), would have been to identify oneself as a radical thinker who might favor atheism, materialism, or even political upheaval; someone who might plot to overthrow the elite establishment (Desmond 1989). Such views were subversive and usually circulated underground. They were probably derived from the transformisme advocated by French naturalists such as Jean Baptiste Lamarck and Etienne Geoffroy Saint Hilaire (Corsi 1989; Appel 1987), perhaps with a dash of German ideas about metamorphosis springing from naturphilosophie (Richards 1992), but also based in social reformist literature, such as the atheistical writings of Baron

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d’Holbach. It is easy to see how this swirl of radical ideas might simultaneously draw Darwin in and yet alarm his sense of social propriety and position (Desmond and Moore 1991). Ideas about transmutation in these various hands could take many shapes. It is therefore important for scholars to identify the social, religious, and educational settings of the people and movements they write about. Studies on the reception of Darwinism across the globe, and among different groups of people, take special care to reflect this caution (Glick 1974; Elshakry 2014). Decentering Darwin also encourages serious investigation of other thinkers. Herbert Spencer (1820–1903) was a remarkable polymath, avowed agnostic, and an evolutionary writer of note who began publishing on this topic some years before Darwin’s publication of Origin of Species (Paradis 2007). As his writings spread, he was crucial in the adoption of cultural evolutionary views and social Darwinism around the globe (Werth 2009; Lightman 2015). Yet Spencer is not given sufficient attention in the historical literature. Nor were his ideas Darwinian in any large degree: he based much of his philosophy on an inheritance of acquired characters. By contrast, Alfred Russel Wallace (1823–1913) has generated a scholarly industry comparable to that of Darwin, peopled by authors who are usually indignant on Wallace’s behalf for what is often characterized as a conspiracy to keep his name in the shadows. This, in itself, could make an excellent topic for analysis. Wallace’s autobiography (Wallace 1905), an early edition of his letters (Marchant 1916), and his many social reformist publications provide insight into his unusual mind. A number of authors have championed him (Mckinney 1972; Fichman 1981; Fichman 2003; Smith and Beccaloni 2008; Shermer 2002; Slotten 2004). His travels in South East Asia have been well documented (Berry 2003; Raby 2001), and his theories of evolution closely examined, although most of the time compared to those of Darwin, which undermines the case for thinking about him as an independent figure (Costa 2014). Conspiracy theories have been explored (Brooks 1984; Brackman 1980). It might be more revealing historically to position him in the movements typified by Chambers and Spencer rather than always to shoehorn him into the same story as Darwin. To decenter Darwin is the explicit focus of books by Peter Bowler, particularly The Eclipse of Darwinism (Bowler 1983) and Darwin Deleted (Bowler 2013). The latter opens with the question what if Darwin’s ship had sunk on its voyage? Would Wallace’s or Spencer’s version of evolution have been accepted to the same degree and what would biology look like today? Furthermore, decentering allows fuller recognition of the work of women in Darwin’s circle and more generally as critics and readers of his writings. The significant feminist points made in early publications (Richards 1983; Russett 1989) have continued to the present day. Among the areas researched are the ways in which Darwinian gender ideology shaped scientific participation for women (Hamlin 2014) and highlights the women who participated in Darwinian discourse (Kohlstedt and Jorgensen 1999; Gianquitto and Fisher 2014). Darwin’s concept of sexual selection has been explored historically (Milam 2010; Richards 2017) and also from the point of view of working theoretical biologists (Cronin 1991; Roughgarden 2004).

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Social Darwinism The leading theme of Social Darwinism, as usually understood, is a commitment to the notion that competition and selection are the natural (i.e., biological) foundation to all human affairs; therefore, human progress best emerges from a free market economy in which competition and selection are allowed full play. It is linked with laisser-faire economic practices, competitive commercial behaviors, the adoption of concepts of racial hierarchy, and bias in class and gender issues, as well as support for eugenics, imperialism, nationalism, and colonial appropriation (Hawkins 1997; Bannister 1989). As such, it was an ideology that had worldwide impact far beyond biology, especially in Europe and the United States from around 1870 to 1950. It is generally associated with the rise of Fascism in the 1930s, with Adolf Hitler’s crusade for racial purity, and with aggressive individualism in capitalist political systems and the business community (Weikart 2006). “Social Darwinism” is therefore a term that comes freighted with negative feeling, epitomized by Richard Hofstader’s Social Darwinism in American Thought, 1860–1915 (1944) that critiqued late nineteenthcentury American society and its economic values. The term Social Darwinism gained currency after Hofstader used it, although it seems to have appeared occasionally before then in the literature. It is almost always used pejoratively. Careful sociopolitical studies on the wide scope and problems of definition are presented by Moore (1986) and Leonard (2009). Because of its wide scope and multiple meanings drawing on biology, power, race, nation, and economics, Social Darwinism has been a fruitful research category for historians for a long time (Jones 1980). It is often said that the movement should really be characterized as “Social Spencerism” (Lightman 2015). Certainly, a great deal can be attributed to the popularity of Herbert Spencer’s writings (Spencer 1891). Spencer supported laissez-faire economics on the basis that struggle for survival would spur self-improvement and that this improvement could be inherited. Spencer, not Darwin, coined the phrase “survival of the fittest,” and Darwin, in later editions of Origin of Species, employed Spencer’s phrase. Spencer opposed any state activity that might be thought to maintain the unfit members of society although he believed in public causes such as altruism, suffrage, and land nationalization. The expression “survival of the fittest” became the leitmotif of what people commonly understood Darwinism and Social Darwinism to be about. Spender’s original definition of this phrase makes clear that “fittest” actually meant to him “most suitable for the conditions” not “strongest” as it subsequently came to mean. Spencer’s successful overseas lecture tours established the key ideas of competition and survival in the United States and elsewhere. By the end of the nineteenth century, these ideas were being put into action by the business entrepreneurs who masterminded the development of North American industry. In their view, the strongest and most efficient company would dominate the market and stimulate economic progress on the wider scale. Andrew Carnegie, the émigré Scots philanthropist and steel magnate, admired Spencer (Werth 2009). Such commitments were generally, if not exclusively, biased toward the political right. None of these thinkers

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believed in the emerging quasi-evolutionary socialist movement (Hale 2014) or in state support for the poor. It was assumed that any circumvention of Darwinism’s “natural laws” would permit “unfit” businesses to survive, thereby undercutting economic and national progress. An interesting sidelight on this field is that scholars have sometimes sought to maintain Darwin’s “purity” by separating him from the social consequences of his proposals (Shapin and Barnes 1979; Moore 1986; Richards 2013). Recent studies indicate that there is, nevertheless, much in Darwin’s Descent of Man published in 1871, on key issues such as eugenics and racial hierarchy, although he did not actually propose these as systems of thought. Darwin’s strong antipathy to slavery has been proposed as the motive force behind his evolutionary views (Desmond and Moore 2009), which seems a somewhat similar attempt to cleanse Darwin from the political effects of his theory. The survival of the fittest also supported contemporary notions of inbuilt racial difference and appeared to vindicate continuing fights for territory and political power on the international stage (Stepan 1982; Paul 2003). The evident success of white people in conquering and settling in foreign lands seemed to make “natural” the subjugation of indigenous peoples and near-extermination of populations such as Tasmanian Aboriginals. Conquest was deemed a necessary part of progress, at least by the conquerors. Eugenics was given its name and leading principles by Francis Galton in the 1880s, drawing on social and racial assumptions already well established but acquiring greater force when attached to evolutionary theory (Kevles 1985). Galton (who was Darwin’s cousin) feared that civilized societies tended generally to prevent natural selection working, in the sense that many of the “unfit” were preserved by medical intervention, charity, or religious doctrine, whereas in a state of nature, such people would die without reproducing. The so-called “worst” elements of society were the most fecund, he said. Galton promoted the idea of more births among the “worthy” middle classes. Eugenics became one of the most pervasive scientized movements of the early twentieth century, spreading widely through Europe and the Americas (Bashford and Levine 2010). It focused middleclass concerns about possible racial and national decline and projected them onto the “unfit” in society. Many eugenists were advocates for technological and scientific advance and in the early twentieth century often encouraged birth control. Many were committed socialists and supporters of women’s suffrage and yet also adopted nationalist, chauvinist, and elitist ideas in education and the public sphere. In the hands of the Fascist movements of the 1930s, these ideas were expressed through Hitler’s ideas of “lebensraum” and racial purity, resulting in state programs of sterilization and the genocide that followed. While Darwin’s Origin of Species or Descent of Man can hardly account for all the racial stereotyping, nationalist fervor, and bigotry to be found in the century after his death, there can be no denying the impact of his writings in providing an authoritative biological backing for a new ideology that combined science, eugenics, warfare, colonialism, notions of racial difference, and the superiority of western civilization.

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Conclusion What directions are Darwin studies taking now? Principally, the current trend is to expand our understanding that Darwinism was (and is) a global phenomenon. In fact, the spread of Darwinian evolutionary thought is an excellent area for exploring the interplay between the assumed universal nature of science and the tendency for regional differences to be expressed. There are an increasing number of significant studies that address the ways in which evolutionary ideas have circulated and are variously assimilated into specific national and cultural contexts. These studies have much to tell us about the movement of scientific ideas in general, about the transition in systems of thought from the local to the global, and the process by which concepts are taken up, adjusted, or rejected. There has been a dramatic upturn in studies carried out in contexts far from the Anglo-American context in which Darwinism first emerged and this – among other things – reflects a wish to decolonize scholarship. A corollary of this widening scope is that alternatives to Darwinian thought are increasingly on the historians’ agenda, such as the rise of epigenetics, and its relationship to Lamarckian ideas, as in the case of Trofim Lysenko in Russia (Graham 2016; Peterson 2016). The trend for historiography in this area is also more fully to reflect a fundamental shift in approach in which a single individual is now recognized as only one factor in the press of events that bring about major changes in science and society. This new focus does not aim merely to provide more “context” to Darwin’s achievement, or include more “precursors,” “supporters,” “rivals,” or “critics” (as in, e.g., Stott 2012), or to examine his actual role in propounding the ideas characterizing social Darwinism, or other cultural movements that drew on his writings. Now, we see that Darwinism was not nearly as securely established as Darwin’s modern followers would like to believe. There is scope to continue to explore the relationship between evolutionism and the history of early genetics (Gayon 1998), eugenics (Kevles 1985), religious reaction in the United States (Moore 1979; Numbers 2006), and the rise and fall of popular interest in Darwin as a person as shown in biographies (Lightman 2010; Browne 2010). Postmodernism encourages scholars to ask how did various individuals and ideas become high-profile and what these phenomena have to tell us about the changing times. The celebratory events that marked Darwin’s anniversaries are therefore ripe for much more study, especially beyond the AngloAmerican context, following on from the 1909 commemoration of Darwin’s birth (Richmond 2006), the 1959 anniversary of publication of On the Origin of Species (Smocovitis 1999), and the 150th anniversary celebrated in 2009 (Shapin 2010).

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Lightman B (ed) (2015) Global Spencerism: the communication and appropriation of a British evolutionist. Brill, Leiden Lindberg DC, Numbers RL (eds) (1986) God and nature: historical essays on the encounter between Christianity and science. University of California Press, Berkeley Litchfield H (1915) Emma Darwin: a century of family letters 1796–1896, 2 vols. John Murray, London Livingstone D (2014) Dealing with Darwin: place, politics and rhetoric in religious engagements with evolution. Johns Hopkins University Press, Baltimore Manier E (1978) The young Darwin and his cultural circle: a study of influences which helped shape the language and logic of the first drafts of the theory of natural selection. Reidel, Dordrecht. Reprinted Springer 2008 Marchant J (ed) (1916) Alfred Russel Wallace; letters and reminiscences, 2 vols. Cassell and Co., London Mayr E (1982) The growth of biological thought: diversity, evolution, and inheritance. Harvard University Press, Cambridge, MA McKinney HL (1972) Wallace and natural selection. Yale University Press, New Haven Milam EL (2010) Looking for a few good males: female choice in evolutionary biology. Johns Hopkins University Press, Baltimore Milner R (2009) Darwin’s universe: evolution from A to Z. University of California Press, Berkeley Montgomery G (2015) Primates in the real world: escaping primate folklore and creating primate science. University of Virginia Press, Charlottesville Moore JR (1979) The post-Darwinian controversies: a study of the Protestant struggle to come to terms with Darwin in Great Britain and America. Cambridge University Press, Cambridge Moore JR (1982) Charles Darwin lies in Westminster Abbey. In: Berry RJ (ed) Charles Darwin: a commemoration. Academic Press, London, pp 97–113 Moore JR (1986) Socializing Darwinism: historiography and the fortunes of a phrase. In: Levidow L (ed) Science as politics. Free Association Books, London, pp 38–80 Moore JR (1989) Of love and death: why Darwin gave up Christianity. In: Moore J (ed) History, humanity and evolution: essays for John C. Greene. Cambridge University Press, Cambridge, pp 195–229 Moore JR (1994) The Darwin legend. Baker Books, Grand Rapids Moser S (1998) Ancestral images: the iconography of human antiquity. Cornell University Press, Ithaca Nesse RM, Williams GC (1995) Why we get sick: the new science of Darwinian medicine. Random House, New York Numbers RL (2006) The creationists: from scientific creationism to intelligent design. Harvard University Press, Cambridge, MA Olby RC (1963) Charles Darwin’s manuscript of pangenesis. British J Hist of Sci 1:251–263 Oldroyd DR (1984) How did Darwin arrive at his theory? The secondary literature to 1982. Hist Sci 22:325–374 Ospovat D (1981) The development of Darwin’s theory: natural history, natural theology, and natural selection, 1838–1859. Cambridge University Press, Cambridge Paradis JG (2007) Samuel Butler, Victorian against the grain: a critical overview. University of Toronto Press, Toronto Paul DB (2003) Darwin, social Darwinism and eugenics. In: Hodge J, Radick G (eds) The Cambridge companion to Darwin. Cambridge University Press, Cambridge, pp 214–239 Peckham M (ed) (1959) The Origin of species by Charles Darwin: a variorum text. University of Pennsylvania Press, Philadelphia. Reprinted 2006 Peterson E (2016) The life organic: the Theoretical Biology Club and the birth of epigenetics. Univerity of Pittsburgh Press, Pittsburgh. Pickering GW (1974) Creative malady: illness in the lives and minds of Charles Darwin, Florence Nightingale, Mary Baker Eddy, Sigmund Freud, Marcel Proust, Elizabeth Barrett Browning. Oxford University Press, Oxford

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Smith J (2006) Charles Darwin and Victorian visual culture. Cambridge University Press, Cambridge Smith CH, Beccaloni G (eds) (2008) Natural selection and beyond: the intellectual legacy of Alfred Russel Wallace. Oxford University Press, Oxford Smocovitis VB (1999) The 1959 Darwin centennial celebration in America. Osiris 14:278–323 Smocovitis VB (2005) It ain’t over ‘til it’s over: rethinking the Darwinian revolution. J Hist Biol 38:33–49 Soderqvist T (2007) The history and poetics of scientific biography. Ashgate, Aldershot Spencer H (1891) Essays: scientific, political and speculative, 3 vols. Wiliams and Norgate, London Sponsel A (2018) Darwin’s evolving identity: adventure, ambition, and the sin of speculation. University of Chicago Press, Chicago Stauffer RC (ed) (1975) Charles Darwin’s natural selection, being the second part of his big species book written from 1856 to 1858. Cambridge University Press, Cambridge Stepan N (1982) The idea of race in science: Great Britain, 1800–1960. Macmillan, London Stott R (2003) Darwin and the barnacle: the story of one tiny creature and history’s most spectacular scientific breakthrough. Faber & Faber, London Stott R (2012) Darwin’s ghosts: the secret history of evolution. Random House, New York Sulloway F (1982) Darwin and his finches: the evolution of a legend. J Hist Biol 15:1–53 Thurtle P (2007) The emergence of genetic rationality: space, time, and information in American biological science, 1870–1920. University of Washington Press, Seattle Turner FM (1974) Between science and religion. Yale University Press, New Haven Van Wyhe J (2007) Mind the gap: did Darwin avoid publishing his theory for many years? Notes Rec R Soc 61:177–205 Voss J (2010) Darwin’s pictures: views of evolutionary theory, 1837–1874. Yale University Press, New Haven Wallace AR (1889) Darwinism: an exposition of the theory of natural selection with some of its applications. Macmillan, London Wallace AR (1905) My life: a record of events and opinions, 2 vols. Chapman & Hall, London Wedgwood B, Wedgwood H (1980) The Wedgwood circle, 1730–1897: four generations of a family and their friends. Studio Vista, London Weikart R (2006) From Darwin to Hitler: evolutionary ethics, eugenics and racism in Germany. Palgrave Macmillan, New York Weiner J (1994) The beak of the finch: a story of evolution in our time. Alfred A. Knopf, New York Werth B (2009) Banquet at Delmonico’s: great minds, the gilded age, and the triumph of evolution in America. Random House, New York Young RM (1969) Malthus and the evolutionists: the common context of biological and social theory. Past & Present 43(1):109–145. Reprinted in Young 1985b Young RM (1985a) Darwinism is social. In: Kohn D (ed) The Darwinian heritage. Princeton University Press, Princeton, pp 609–638 Young RM (1985b) Darwin’s metaphor: nature’s place in Victorian culture. Cambridge University Press, Cambridge

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The Historiography of Modern Evolutionary Biology Mark E. Borrello

Contents Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Biologists’ Histories . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . The Long Modern Synthesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Evolutionary Developmental Biology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Systematics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Histories of Selection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sexual Selection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ethology and Behavior . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sociobiology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Future Directions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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Abstract

Though Darwin and his copious work remain the focus of much of the history of biology even into the twenty-first century, the development of evolutionary biology throughout the twentieth century has been the subject of a great deal of significant historical analysis. As the scholarship on topics such as selection, speciation, or human evolution indicates the history of ideas is alive and well when it comes to evolutionary biology. Histories of evolution have not ignored wider social and cultural contexts either, as accounts of debates over sexual selection, group selection, and altruism, creationism, and sociobiology make clear.

M. E. Borrello (*) Program in the History of Science, Technology and Medicine Department of Ecology, Evolution and Behavior, University of Minnesota, Minneapolis, MN, USA e-mail: [email protected] © Springer Nature Switzerland AG 2021 M. R. Dietrich et al. (eds.), Handbook of the Historiography of Biology, Historiographies of Science, https://doi.org/10.1007/978-3-319-90119-0_5

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Introduction Though Darwin and his copious work remains the focus of much of the history of biology even into the twenty-first century, the development of evolutionary biology throughout the twentieth century has been the subject of a great deal of significant historical analysis. That said it is not at all obvious how to organize an analysis of the history of “modern” evolutionary biology in the twentieth century. As the scholarship on topics such as selection, speciation, or human evolution indicates, the history of ideas is alive and well when it comes to evolutionary biology. Histories of evolution have not ignored wider social and cultural contexts either, as accounts of debates over sexual selection, group selection, and altruism, creationism, and sociobiology make clear. For the purposes of this essay, I begin with how biologists have approached the history of their own field. I then contrast these accounts with those produced by historians. I organize the historian’s accounts topically and emphasize different historiographic approaches within each topic. I chose this topical approach because different topics have been subject to different levels of scrutiny by historians. This feeds into my final section where I consider future directions in the history of modern evolutionary biology.

Biologists’ Histories I begin with Ernst Mayr’s (1982) tome The Growth of Biological Thought and Stephen Jay Gould’s (2002) magnum opus, The Structure of Evolutionary Theory. Scientists’ histories often form the first pass on the history of a field. When written by participants, they can be a rich source of personal and collective memory. That said they can also be partial and biased, even self-serving. Nevertheless, Mayr, Gould, and other biologists have left a significant mark on the history of evolutionary biology. Mayr’s career spanned the twentieth century and by his own account was divided into three distinct stages. The first stage from the 1930s to the late 1960s was as an evolutionary biologist, ornithologist, and systematist. The second phase was dedicated to the history of biology and the final phase was a philosophical stage. The Growth of Biological Thought (1982) represents the culmination of Mayr’s historical work and, in some ways, served as the standard against which many historians of contemporary evolutionary biology measured their own work. Of course, Mayr’s historical vision was a very particular one. As historians Betty Smocovitis (1994, 1996) and Joe Cain (1993, 2009a) have argued, Mayr’s influence on our understanding of evolution is hard to overestimate. As Cain puts it, “Mayr’s strategy of inviting everyone into a big tent (chiefly of his own design), and engaging every conversation sincerely and seriously, has had lasting consequences. The seriousness with which scientists and historians have engaged each other on the topic of the great synthesis only adds to our collective sense of its importance” (Cain 2009a, p. 624). Reviewing Mayr’s last book, What Makes Biology Unique?, in 2005,

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the year he died, Smocovitis wrote, “What does come from Mayr’s monumental texts has shaped not one but at least a couple of academic generations in both the history and philosophy of biology (not to mention his influence on evolutionary biology). Mayr’s influence on the field has been deep, and I would argue defining” (Smocovitis 2005, p. 614). In an earlier paper that resonated with Cain’s assessment, Smocovitis pointed to the varied institutional and intellectual efforts of Mayr and others as “part of a process that led to the construction of a common language, a disciplinary discourse which would in turn lead to the emergence of a new ‘central’ science of evolutionary biology that would redefine the identities of the members. Critical to the process was the organizational role played by one chief ‘architect’ of the evolutionary synthesis, Ernst Mayr” (Smocovitis 1994, p. 242). Mayr’s legacy is complex and no doubt as the object and author of the standard story of the modern synthesis of evolutionary theory, his work demands careful attention. Perhaps not surprisingly, Mayr’s historical focus was the development of evolutionary biology in the twentieth century both as a set of theories and practices, and as an autonomous and fundamental field of biology. Mayr’s colleague at Harvard and generational successor, Stephen Jay Gould, has also made significant contributions to the history of biology and in particular, to the history of modern evolutionary theory. Indeed, Gould often presented his historical work as a counterweight to the received view presented by Mayr. As a historian, much as he did as a paleobiologist, Gould took it as his duty to highlight the contributions of the less orthodox views of evolutionary theory (i.e., the ideas of Richard Goldschmidt, the significance of contingency, multilevel selection, and his own theory of punctuated equilibria) and the roles they played (or did not) in the development of contemporary evolutionary theory. While Mayr’s history reflects the view of one of the architects of the modern synthesis, Gould’s work represents a challenge to that history. Indeed, Gould judged Mayr’s history of the synthesis, as presented in both Animal Species and Evolution in 1963 and The Growth of Biological Thought in 1982, as representing a very particular use of history to clarify and constrain the theory and practice of evolutionary biology. The differences between Mayr and Gould’s influential historical accounts is perhaps best illustrated in their assessment of the geneticist Richard Goldschmidt. Briefly, for Mayr, Goldschmidt’s role in the history of the modern synthesis was to serve as a target for perfecting the population genetic approach that was developed by R.A. Fisher, J.B.S. Haldane, and Sewall Wright. Goldschmidt, according to Mayr, was simply wrong in his assertions that “the decisive step in evolution, the first step toward macroevolution requires another method than the sheer accumulation of micromutations” (Goldschmidt in Mayr 1982, p. 562). Goldschmidt was wrong, too, about cytoplasmic inheritance (p. 788) and the nature of the gene (p. 737). Interestingly, in the same year that Growth was published, Gould published a forward to the reissue of Goldschmidt’s 1940 classic, The Material Basis of Evolution, titled “The Uses of Heresy,” where Gould uses Goldschmidt to demonstrate how contemporary evolutionary biology is coming around to some of Goldschmidt’s views. Gould as a biologist was sympathetic to Goldschmidt’s claims about the need for something more than “the sheer accumulation of

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micromutations.” One can imagine the atmosphere at the Museum of Comparative Zoology at Harvard where both Mayr and Gould made their academic homes. As Michael Dietrich has noted, when biologists such as Mayr and Gould invoke a figure such as Goldschmidt, they are using history as a kind of commemoration that can be put to use in contemporary biology (Dietrich 2011). Of course, historians also craft narratives with specific goals in mind, but those goals rarely include legitimating contemporary science. By contrast, as Pnina Abir-Am has argued, scientists’ uses of public commemoration are either aimed at articulating the public imagery of their discipline, at establishing organizational support within a field, and at creating a link to a prominent ancestor in order to legitimate the claims to authority of that ancestor’s descendants (Abir-Am 1999, p. 326, also see Creager 2010). In Gould’s case, Dietrich argues that he used Goldschmidt to demarcate orthodoxy and heresy in evolutionary biology. When Gould aligned himself with Goldschmidt, what he effectively communicated was his dissent from the orthodoxy still trenchantly defended by the previous generation of evolutionary biologists, as embodied in Ernst Mayr (see Mayr 1997). Biologists will continue to write histories of their field. Rather than take them as the first draft of history, however, examples such as these invite us to consider that biologists might have more complicated agendas.

The Long Modern Synthesis The modern synthesis, the evolutionary synthesis, or the neo-Darwinian synthesis has been taken to be a defining feature of evolutionary biology in the twentieth century. As such, questions regarding the nature, content, and meaning of the evolutionary synthesis have motivated historical inquiry since at least the late 1950s (Tax 1960, Vols. 1–3). Both biologists and historians have generated volumes, conducted symposia, and engaged in debates regarding the construction of the appropriate historical narrative and the identification of the key contributions (Mayr and Provine 1980). The modern synthesis has been described in a number of ways: the integration of theories from a wide range of fields (Darden 1986), the cooperative organizational solution to discipline building (Cain 1993), a conscientious effort on the part of working scientists to unify biology (Smocovitis 1994, 1996), and a hardening of biological investigation and explanation around the idea of individual-level adaptation (Gould 1983). Not surprisingly, these different accounts of the synthesis put different temporal bounds on it. For some it begins with early twentieth-century efforts to bring together Mendelism and Darwinism (Provine 1971). For others, Dobzhansky’s 1936 Genetics and the Origin of Species is a more natural starting place. Gould famously argued that the synthesis “hardened” in the postwar period, while more recent advocates of both sociobiology and evo-devo sought to “extend” it, albeit in different directions. For insight into the historical, biological, and philosophical issues associated with these developments, see Massimo Pigliucci and Gerd Muller, The Extended Evolutionary Synthesis, and Kevin Laland et al., “The extended evolutionary synthesis: its structure, assumptions and predictions.”

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William Provine’s The Origins of Theoretical Population Genetics (1971) could have been titled “The Story of the Modern Synthesis: Darwinism, Mendelism and Biometrics.” Perhaps this title would have eased the fears of some historians interested in reading this book without a background in genetics. This is important because Provine does not spend pages and pages delving into the intricate statistical and mathematical analysis that constituted much of this story. Rather, Provine provides an account of individuals, alliances, and debates which shaped, and sometimes stalled, but ultimately led to the synthesis of gradualistic Darwinian evolution, Mendelian genetics, and biometry, which is fundamental to present-day biological research. Provine’s Origins contributes to the history of biology in two distinct ways. First, it represents the first historical account of the development of population genetics and thus opened an exceedingly important area of biology to the historian. Second, Provine’s intellectual history is socially informed. Provine’s story of the evolutionary synthesis begins with an account of the varied reaction to Darwin’s theory of evolution by natural selection. He emphasizes two aspects of Darwin’s theory which concerned even his staunchest supporters, including Huxley and Galton, and led to what Peter Bowler has described as the “Eclipse of Darwinism” (Bowler 1992). These factors are Darwin’s insistence on the gradual nature of evolution and his lack of a coherent theory of heredity. Provine gives a lucid account of Galton’s statistical analysis of heredity and how this led to his saltationist assertion that evolution proceeded by “sports.” Galton’s work, especially his ideas about regression, sparked an interest in mathematical analyses of evolution, and hereditary phenomena in particular. Karl Pearson and W.F.R. Weldon sought to improve on the mathematics of Galton and to reestablish the gradual nature of Darwinian evolution through mathematical analyses presented in their new journal Biometrika, established in 1901. Simultaneously, William Bateson, who was not a mathematically inclined scientist, took from Galton’s work the saltationist view and, as a rediscoverer of Mendel’s work, melded these two approaches into a saltationist–Mendelian interpretation of evolution. The differing interpretations of Bateson, who advanced a discontinuous interpretation of evolution, and Weldon and Pearson, who supported a continuous evolution, form the backbone on which the rest of Provine’s story is supported. Throughout the body of this book Provine tracks the development of the debate between the Mendelians and the Darwinian biometricians. He examines various episodes which provide strong evidence for his position that science is often as socially contingent as it is data, or theory based. Provine repeatedly asserts (Provine 1971, pp. 25, 64, 69–70) that if the debate between the Mendelians and the Darwinians (read Bateson vs. Weldon and Pearson) had not been so vitriolic and personal, population genetics could have been established 15–20 years earlier. While Provine’s account laid out the broad contours of the Mendelianbiometrician debate, subsequent historians have revisited both the actors, the issues, and the dynamics of the controversy. Theodore Porter’s 2006 biography of Karl Pearson provides an exceptionally insightful background to the development of the statistical and rational approach to variation in populations that drove much of the later work population genetics. The

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possibility of an earlier resolution to this debate has also been explored in work such as James Tabery’s 2004 paper, “The ‘Evolutionary Synthesis’ of George Udny Yule” and in the 2014 Stoltzfus and Cable’s paper, “Mendelian-Mutationism: the forgotten evolutionary synthesis.” Recently, Gregory Raddick has offered an insightful new analysis in his forthcoming book Disputed Inheritance: The Battle of Mendel and the Future of Biology. The culmination of Provine’s account of the origins on the synthesis rests in the work of R. A. Fisher, J. B. S. Haldane, and Sewall Wright. Although Provine presents very brief biographical sketches of each of these men, his emphasis is on the conceptual reconciliation of emerging research on genetics and Darwinian evolutionary biology. Perhaps because this material is very technical, Provine’s account becomes more biologically descriptive. He does not explain how the differing views of Fisher, Haldane, and Wright are ultimately fused, yet understanding of their integration is essential to understanding what population genetics is. In the decades since, substantial new histories of population genetics have articulated both the origins and development of this field. Provine’s biography of Sewall Wright explains in detail how Wright, Fisher, and Haldane interacted with each other while developing different approaches to evolutionary genetics (Provine 1986). Hodge (1992) explores the philosophical views of Fisher and Wright as a way of offering a historiography of population genetics that explores three historical topics: Darwin as mechanist, the evolution revolution, and the probabilistic revolution in evolutionary genetics. In a pair of articles, Robert Skipper analyzes the persistent controversy between Fisher and Wright. Both extend the extant historical analysis of the Fisher–Wright controversy from the 1950s to the early 2000s by critically analyzing the debate in the late 1990s between biologists Jerry Coyne and Michael Wade over the scope and application of Fisher’s and Wright’s evolutionary theories. Skipper pays special attention to the debates over the scope of Fisher’s and Wright’s evolutionary theories, theories of dominance, and role of drift in evolution. Mark Borrello (2009) has also examined the work of Dobzhansky and Wright as it pertains to the question of the level of selection in the evolutionary process. Anya Plutynski (2006) revisits the Price-Ewens interpretation of Fisher’s fundamental theorem of natural selection paying special attention to its role in Fisher’s work on population genetics. Sahotra Sarkar (2007) discusses J. B. S. Haldane’s role at the origin of theoretical population genetics. Provine’s Anglo-American concentration skews the picture of the development of genetics. This has continued to be reflected in the literature. Though it is widely recognized that Russian geneticists were active in the 1920s and made various important contributions to population genetics – see the work of Mark Adams (1968) and Nikolas Krementsov (1996, 2005, 2010), for example – these contributions have not been integrated into the received history. Montgomery Slatkin and Michel Veuille (2002) offer an edited collection of essays written by biologists exploring the contributions to population genetics inspired by the French mathematician Gustav Malécot. But more work examining influences, schools of thought, and individuals outside the existing historiography is needed.

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Provine’s presentation of various episodes, including the acceptance of the pure line theory, and the rejection or ignorance of the work of Yule, Darbishire, and Hardy-Weinberg, clearly illustrate the role of personality and social relations in the process, and the progress of science. Indeed, Provine’s account laid the groundwork for a number of subsequent accounts of the modern synthesis. Immediately after his book appeared in 1971, Provine began working with Ernst Mayr to revisit the history of the evolutionary synthesis through the eyes of its biological architects. Based on a series of in-depth surveys given to selected biologists, this collection sought to use biologists’ own perspectives to reconstruct the evolutionary synthesis (Mayr and Provine 1980). While the volume made room for the role of hitherto unrecognized, or insufficiently recognized actors such as C.D. Darlington and Sergei Chetverikov, and provided a transnational perspective, significantly, Mayr cast himself in a central role (an essay by Gould, nonetheless, was included). In reinforcing a narrative of “architects” from different disciplines, this collection became a second major foil for subsequent work by historians. Provine’s own disaffection with the collaborative volume with Mayr was expressed in his 1988 essay rejecting the idea of a synthesis (Provine 1988). He argues that the search for a single synthetic theory of neo-Darwinian evolution is misguided and that the metaphor of synthesis should be replaced by that of constriction. According to this view, architects of the synthesis did not agree on a common theory, but agreed on what should be eliminated from modern evolution and so constricted evolutionary theory. Orthogenic and saltationist views, for instance, were systematically excluded from evolution during this time period. Later analysis of Richard Goldschmidt’s exclusion from evolutionary biology also support this model (Dietrich 1995; Davis et al. 2009). Adding further nuance from the biologist’s perspective, R. C. Lewontin (1981) critically discusses Dobzhansky’s contributions to evolutionary genetics, particularly the Genetics of Natural Populations series, in which speciation figures as a central theme. Speciation is the process or processes by which some lineage, or species, splits into two. Defining “species” and understanding the mechanisms of speciation are controversial topics in biology. Subsequent work has examined the vagaries of its conceptualizations and their trajectories. Wilkins (2011) offers a comprehensive historical survey of research on species concepts from antiquity. Jürgen Haffer’s (2007) biography of Mayr naturally addresses Mayr’s significant contributions to debates about species and the creation of the biological species concept. Leo Laporte’s (1994) work does much the same for G.G. Simpson’s contributions to issues regarding species and speciation in paleontology. In the context of the ongoing historiographical debates in the history of science that were prevalent in the 1990s, V. B. Smocovitis’ book on the role of the modern synthesis in the unification of the field of biology makes an important contribution. Smocovitis’ book Unifying Biology was an expansion of an article originally published in the Journal of the History of Biology in 1992. In the article, Smocovitis claimed that her central argument is simply that “the evolutionary synthesis signaled the unification of the biological sciences.” Indeed, she argued that by the completion of the synthesis, evolution had been purged of its metaphysical elements and biology

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was now bound together by the glue of evolution. This unification of biology was consistent with the goals of many scientists and philosophers of science of the era who were sympathetic to the creed of the logical positivists, the unification of the all the sciences. Although Smocovitis admitted that her characterization of the logical positivist program is oversimplified, she also argued that it is sufficient for her purposes. This was not consistent with the standard historical view, nor was it consistent with the memory of one of the most important participants, Ernst Mayr. In the standard account, many biologists resisted the attempts to include their science in the positivist project for fear that biology would cease to exist as an autonomous science and would be inexorably reduced to physics. Furthermore, as Smocovitis pointed out in her epilogue, Mayr disagreed with her interpretation. Mayr argued in a 1999 article in Trends in Ecology and Evolution that the unification of the field of biology was not an objective of the participants in the synthesis. Rather, according to Mayr, these individuals were responding to anti-Darwinians and straightening out the differences within their own fields. They could not be concerned with such far-reaching goals as the unity of biology. Challenges to the synthesis were also presented from within the scientific community. Niles Eldredge and S. J. Gould’s 1972 theory of punctuated equilibrium threatened the status of Darwinian theory and all that had been accomplished by the synthesis, not only intellectually but also institutionally and politically. These factors played an important role in previous interpretations of the modern synthesis and, as Smocovitis acknowledged in her epilogue, will continue to do so. The modern synthesis, on her account, is an ongoing process. Furthermore, the writing of the history of the modern synthesis is an ongoing process. As Gould and others had argued since the early 1980s, evolutionary theory had been expanding from the population genetic structure that had been codified in the modern synthesis, see Gould (1980, 1982) and Depew and Weber (1995). Historians of contemporary evolutionary theory have thus expanded their own analyses and begun to focus on the contributions to contemporary evolutionary theory from a broader range of biological perspectives including behavioral biology, ecology, and paleobiology among subfields of biology where the development of evolutionary biology was taking place. This is also a reflection of a trend in the history of science more generally that moved away from an exclusive focus on theory, and toward a broader approach that paid greater attention to the practice of biology in the twentieth century, both in the field and in the lab.

Evolutionary Developmental Biology Although a relatively recent field, evolutionary developmental biology has attracted the attention of a number of historians seeking historical precedents for the intersection of evolution and development. Amundson (2005) offers the most comprehensive single narrative of the effort to map the changing intersection of evolution and development. His work explores the history of the relationship between

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evolution and development through the lens of evo-devo; Amundson argues that extant histories were written in a way that vindicates neo-Darwinism and sets development aside. The essays in Laubichler and Maienschein (2007) provide more detailed analyses of relevant historical figures, such as Haeckel, Goldschmidt, and Bateson, as well as essays on important areas of research such as morphology and developmental genetics. The essays represent a broad range of approaches and analyses from scholars. What makes the volume most compelling is the authors and editors self-consciousness regarding the potential impact of these meetings and the resulting volume. This is a theme that is carried through each of the individual essays and most carefully and engagingly presented in the second chapter by one of the editors, Manfred Laubichler. He raises important questions specific to the history of evolutionary developmental biology and deftly connects them to the broader history of mid-nineteenth- and twentieth-century biology. His essay describes the challenges faced by historians whose object of study (in this case evolutionary developmental biology) is not well defined. Indeed, he argues, “we do not seem to be dealing with a single set of questions and methods; its distinctness from other areas of biology is still under debate; and we do not know how it will turn out. This makes the historian’s question of what is legitimately part of the story almost impossible to answer” (p. 15). Laubichler continues, pointing out that this is exactly what makes the field exciting for historians while simultaneously acknowledging the tentative nature of any of the historical conclusions derived in this volume. I think the best bit of this chapter is Laubichler’s conclusion where he describes the historian’s challenge to identify “the appropriate historical object – the specific research problems of ‘ontogeny and phylogeny’ – as the relevant epistemic object that allows us to connect the research of agendas of current evolutionary developmental biology with previous attempts” (p. 25). This approach to history foretells a deep connection between the work of historians, philosophers, and working biologists in developing an integrative history that will “reveal a lot about conceptual constraints, patterns of variation, and the problems of conceptual innovation within evolutionary developmental biology” (p. 28). This idea is clearly echoed in the organization and structure of the volume. The first part, “Ontogeny and Phylogeny in Early Twentieth Century Biology,” presents new perspectives on the history of embryology and developmental genetics from some of the most well-known voices in the field including Fred Churchill, Gar Allen, and Jane Maienschein. Fred Churchill’s chapter in particular stands out. His nuanced account of the changing meaning and significance of Haeckel’s biogenetic law demands careful attention and consideration not just from historians of evolutionary developmental biology but from all. In describing the particular lessons learned from the writing of this chapter, Churchill points out (consistent with Laubichler in the preceding chapter) that an integrated history is required here. He argues, “Until now, however, when we have presented our histories of the same biology, we have tended to frame our narratives in terms of either continuities or discontinuities or one succeeding the other. Our instincts should tell us that both are necessarily involved and intertwined” (p. 71). Indeed, in all of the chapters in this most historical section of the volume, the authors are concerned to evaluate the potential of the integrated

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perspective that is required for the proper historical understanding of the problems of developmental biology and their relation to evolutionary theory and the history of biology in general. The second section of the volume is the most diverse in terms of both subject matter and approach. One might characterize this section as the more distinctly philosophical, though in every case the approach is integrative. The contributors to this section represent philosophy, biology, and sociology in terms of disciplinary identity but all are concerned to incorporate and utilize the history of developmental biology in their analyses. In the second chapter of this section, the philosopher Alan Love’s chapter focuses on the concepts of innovation and novelty. He uses these philosophically rich and scientifically compelling concepts as a guide to “explore [the] morphological and paleontological perspectives for a history of Evo-Devo” (p. 269). This approach, according to Love, moves beyond the focus on developmental genetics and provides a richer historical account of the history of evo-devo that acknowledges the contributions of morphology and paleontology. His chapter also draws into focus the connection of evolutionary developmental biology to one of the other key areas of interest in the history and philosophy of biology, the importance of levels of organization and selection. He concludes that the recognition of this connection could have significant influence on the conduct of future research in evo-devo. “Arguably,” he writes, “higher levels of structural organization were neglected in the Modern Synthesis, and paying attention to this feature may be critical for the future unfolding of Evo-devo as a research discipline” (p. 291). This section also includes excellent chapters by the philosophers James Greisemer and William Wimsatt. Greisemer’s chapter focuses on the split between embryology and the Morgan school development of transmission genetics, which he uses to construct an argument about the importance of understanding scientific representations and explanations of complex processes. The result of this is a history of biology that avoids the “temptation to confuse the divergence of research styles in separating scientific social worlds with a historical progression of fields or lines of work that take the scientific limelight in turn” (p. 417). Griesemer’s message is both philosophical and historiographical; he reminds us that the divergence in research style and representational practice that occurred between genetics and embryology does not entail that nature is separated into heredity and development. We (historians, philosophers, and biologists) all benefit from the recognition of the entwined nature of these complex processes, as they manifest in the integrative approach of evo-devo. The last section of the volume, “Reflections,” consists of three papers by biologists working in evo-devo. Brian Hall’s chapter nicely integrates the message from Griesemer. He uses case studies to demonstrate the interdisciplinarity of evo-devo and to argue that rather than a trajectory from embryology to evo-devo we need to understand this new field as an integration of multiple fields around a common set of problems. The chapters by Gerd Muller and Gunter Wagner echo these interdisciplinary sentiments and provide some frameworks for future directions. The contributions are well written, wide ranging and engaging. For those who have been working in the history of biology for a long time it will come as a breath of fresh air

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(or at least as a new angle on some long standing questions). For newer scholars this volume is a rich resource for potential research. These should be complemented by further existing literature: Davis, Dietrich, and Jacobs (2009) explore the assimilation of developmental biology into evolutionary biology through the study of homeotic mutations. Scott Gilbert (2000) traces Waddington’s influence on the evolutionary synthesis, in particular his impact on the integration of evolution, development, and genetics at mid-century through the influential contribution of canalization and the epigenetic landscape. In his 2003 essay, Gilbert casts a wider net for precursors to contemporary evo-devo, tracing the historical impact of evolutionary morphology and genetics on the integration of evo-devo, but argues that ecology and medicine must also be appreciated as important influences on contemporary practice. Griesemer’s 2003 essay focuses on the integrated approach that has informed evo-devo research conducted by David Wake.

Systematics The Origin of Species famously contained only one image: an evolutionary tree. John Lindley, in 1830, was the first to define “systematics,” though his term was “systematic botany.” Generally, systematics is the comparative study and classification of groupings of organisms at the species level or higher over their evolutionary histories. In 1988 David Hull published Science as a Process where he presents the recent history of systematics as a case study. This case study serves as the primary evidence for a generalized model of how scientists interact and how these interactions lead to conceptual change. Ultimately, Hull incorporates this evolutionary model of science into a global theory of selection processes. This work provides an exceptional example of the ways in which history of science can be fruitfully used by philosophers of science. Later research on the history of systematics includes Joe Cain’s 2004 institutional history of the Society of Systematic Zoology and Josh Buhs’ (2004) history of ant systematics in the early twentieth century. The turn to consider scientific practices is well represented by Bruno Strasser’s 2010 examination of systematic practices, while Joel Hagen (2003) and Edna Suárez-Díaz (2013) consider the introduction and impact of statistical techniques and molecular data in systematics. Jann Sapp’s 2009 book, The New Foundations of Evolution: On the Tree of Life, traces the history of microbial evolution against the background of the tree of life. This book is consistent with Sapp’s earlier work on cytoplasmic inheritance and the evolution of symbiosis in that it concentrates on ideas in biology that are outside the mainstream. Indeed, one of Sapp’s goals as an historian is to constantly broaden our image of what is important in biology and why. Sapp is also unapologetic about his use of history to illuminate contemporary scientific debate. Historians are often hesitant to be explicit about this for fear of being labeled as “whiggish” but Sapp eschews such concerns. As one of the central characters of Sapp’s story, Carl Woese remarks in the foreword: “he finds his history on the unpaved trails of contemporary scientific exploration rather than safely recording his travels along the scientific

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super highways of the past” (p. vii). In New Foundations this strategy works exceedingly well. That is not to say that Sapp does not treat the early history of microbiology. Indeed the first ten chapters (of 22 total) are dedicated to a careful rendering of the developing interest in microbes beginning with Anton von Leeuwenhoek in the mid-seventeenth century and carrying through to the establishment of the genetic code in the middle of the twentieth century. What makes this history particularly interesting is the connection he draws with the contemporary theorizing in phylogenetics. He points out that previous histories of microbiology have concentrated on the relationship between microbes and agriculture or medicine. Here, his focus is squarely on the evolutionary history and diversity of microbes and the varied attempts to understand and explain these phenomena. Sapp deftly describes the challenges to a genealogical approach to bacterial classification and notes that for many researchers of Pasteur’s day, this approach was seen as neither necessary, or indeed, even possible. Debates ensued over whether classification was more properly based on morphological or physiological traits during this first of four distinct phases of the study of bacterial diversity that Sapp identifies. With respect to the idea of a universal ancestor, a standing question among researchers in microbial evolution, the early favorite was an autotrophic organism that arose approximately 1.5 billion years ago. The second phase was characterized by the identification of viruses as entities distinct from other microbes on the basis of electron microscopy and the prokaryote–eukaryote distinction was maintained as a fundamental division of the organization of life on the basis of the fact that prokaryotes “lacked a membrane bound nucleus, organelles, and sexuality comparable to other microbes” (p. 315). During this period, consensus on the universal ancestor shifted to a heterotrophic prokaryote that fed on the primordial soup and evolved over billions of years. The symbiotic origin of the mitochondria and chloroplasts as well as the idea of lateral gene transfer entered the discussion of the evolution of microbial forms. In the third phase of this history, essentially the rise of molecular phylogenetics, Sapp begins his focus on the work of the aforementioned Carl Woese and his collaborator George Fox. In 1977 Woese and Fox introduced archaebacteria to the world in a paper titled “Phylogenetic Structure of the Prokaryotic Domain: Three Primary Kingdoms.” In this paper they argued that the eukaryotes were not derived from the prokaryotes and that rather there were three distinct Ur-kingdoms in the history of life: the eubacteria, the eurkaryotes, and the archaebacteria, each having evolved distinctly from hypothetical entities called “progenotes.” The work of Woese and his collaborators and challengers make up the second half of this book, providing insights into the details of the restructuring of the tree of life that are hard fought and fascinating. The fourth and final phase of Sapp’s history examines the influence of genomics on microbial phylogenetics and the increasing evidence of extensive lateral gene transfer among bacteria. The significance of lateral gene transfer stirred controversy in 2009 when the The New Scientist ran a cover titled “Darwin Was Wrong: Cutting down the tree of life.” Though Sapp’s book was in press, at that point this idea that the new phylogenetics invalidates Darwinian theory is perhaps a point of overreach for Sapp. While there is no doubt that the research by Woese and others completely

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restructured the base of the tree of life it is not yet the time to throw out the Darwinian baby with the bathwater. Sapp has done the great service of bringing an area of biology that has been understudied into the light of historical examination. Andrew Hamilton’s (2013) collection of essays explores the origins and development of phylogenetic systematics. This builds on David Hull’s classic, Science As a Process. On Hull as both a commentator and an actor in these debates and developments, see Horder (2013). On the newly named sciences of the microbiome, which further complicate notions of individuality and therefore of systematics, see Ed Yong (2018) I Contain Multitudes: The Microbes Within Us and a Grander View of Life. Another important vein related to the history of systematics is the rise of the field of paleobiology exceptionally well told by David Sepkoski in Rereading the Fossil Record (2012).

Histories of Selection Selection – natural, artificial, and sexual – has been at the heart of evolutionary biology since Darwin. Major debates concerning the efficacy of natural selection to produce biological change were important features of late nineteenth- and early twentieth-century biology. After the Second World War, however, debates over the possibility of group selection and then multilevel selection fundamentally altered the meaning of this core concept. As Lindley Darden points out in her 1986 paper in Integrating Scientific Disciplines, Theodosius Dobzhansky was particularly attuned to the importance of these multiple levels and their role in the process of evolution (for work on Dobzhansky and the Russian genetics school see Adams). As Darden nicely illuminated, they consist of mutations and chromosomal changes at the level of the individual, selection, migration, and isolation occurring at the level of the population, and finally, fixation of the diversity attained at the preceding levels occurring at the species level (Darden 1986, p. 114). In the same volume, John Beatty pointed out that Dobzhansky achieved another kind of synthesis that was equally important. He argued that in addition to thinking of Dobzhansky’s contribution as a synthesis of theories operating at multiple levels, we must also understand his work as a synthesis of theory and observation. Beatty cited Lewontin to this effect writing, [T]he contribution of Dobzhansky’s experimental and field studies to the synthesis, [is] “The successful melding of the theory of gene frequency change with the known facts of genetic variation. . .marks the first real synthesis in biology of a complex mathematical theory with a large body of observation and experiment.” (Beatty 1986, p. 128)

Beatty went on to argue that evolutionary biologists did not go into the field or into their labs simply to test the synthetic theory. Rather, they first wanted to measure the values of the variables of the theory in particular cases. Second, they wanted to extrapolate from the values of the variables in particular cases to the overall relative importance of the various possible modes of evolutionary change (Beatty 1986,

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p. 128). According to Beatty, evolutionary biologists around the time of the synthesis had the goal of understanding the relative importance of the various possible modes of evolutionary change. A further, independent intervention in this debate should be credited to C.D. Darlington, a somewhat idiosyncratic member of the “architects” of the synthesis, who in his 1939 book The Evolution of Genetics Systems argued that selection works on genetic systems beyond the nuclear genes – a notion that foreshadowed both multilevel selection theory and the influential field of evolvability. On Darlington’s role in the historiography of evolutionary theory see Harman (2004). On Richard Goldschmidt’s use of the violin string, and other metaphors, to eschew the notion of the very existence of genes as biological entities implicated in development and amenable to selection, see Lamm (2008). On the conceptual distinctions relevant to current debates on levels of selection see Okasha (2006). Tracing the history of biological attempts to determine whether selection could lead to the evolution of fitter groups, Borrello (2010) takes as his focus the British naturalist Vero Copner Wynne-Edwards, who proposed that animals could regulate their own population levels and thereby avoid overexploitation of their food and other resources. By the mid-twentieth century, Wynne-Edwards became the primary advocate for group selection theory and precipitated a debate that engaged the most significant evolutionary biologists including Ernst Mayr, John Maynard Smith, George C. Williams, and Richard Dawkins. The resulting interpretations and arguments bled out into broader conversations about population regulation, environmental crises, and the evolution of human and animal social behavior. The debate over group selection was also closely tied to the development of inclusive fitness theory and the study of the evolution of altruism. Oren Harman’s (2010) biography of theoretician George Price, The Price of Altruism, provides another fascinating lens through which to view these developments in evolutionary theory in the twentieth century.

Sexual Selection With the publication of Robert Trivers’s account of natural selection, especially his 1972 paper, “Parental Investment and Sexual Selection,” it seemed for a time as though the “good genes” account of sexual selection on males, with its promiscuous males and coy females, would hold sway. However, later developments would broaden sexual selection considerably to include a variety of assumptions and models. Helena Cronin (1991) offers an account of the standard accounts of sexual selection and their historical origins. Sarah Hrdy (1981) reexamines and challenges that canon when she reviews the work of Bateman and Trivers, and argues that it has been inappropriately extrapolated to primates and showing how subsequent work in primatology demands new types of sexual selection explanations. In particular, she argues that sexual selection often operates on females, not just males. Joan Roughgarden expands the ambit of sexual selection even further, pointing out the substantial variability in sexual types upon which selection can act in Evolution’s

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Rainbow: Diversity, Gender and Sexuality in Nature and Humans (2013). The biologist Richard O. Prum argues for historical redress from a minority position in The Evolution of Beauty: How Darwin’s Forgotten Theory of Mate Choice Shapes the Animal World – and Us (2017). Erika Milam opens her (2011) book, Looking for a Few Good Males, with the provocative question: “How does a hen decide whether she is in the mood for sex?” (p. 1). This question sets the tone for her inquiry which ranges from the nineteenth century through the end of the twentieth, encompassing the thoughts, claims, experiments, and ideas of a wide range of biologists. As Milam deftly demonstrates, Darwin’s idea of sexual selection (which was briefly presented in the Origin then fully elaborated in 1871 in The Descent of Man) raised a plethora of contentious scientific, philosophical, political, and even experimental issues. Milam’s goal is to illuminate the constantly shifting topography of biological research on animal behavior and evolution, and to investigate and describe how the myriad applications of scientific knowledge of courtship behavior in animals served as a “biological basis of human courtship” (p. 6). One can immediately surmise that this research therefore bears on some of the most sensitive and controversial issues about the divide between humans and nonhuman animals, about the biology of sex and gender, about the boundaries between instinct and learning, and about the fuzzy line between biology and culture. Milam is also working on a historiographical project here. In the course of the book she demonstrates that the history previously written on the subject of sexual selection theory and, more specifically and importantly for her project, female choice has underrepresented the distinction between sexual selection research and research on mate choice, and has subsequently assumed that as interest in sexual selection theory waned in the twentieth century so too did interest inmate choice. She demonstrates that the “recovery of those forgotten elements of history of sexual selection merit inclusion and consideration,” while at the same time, “changing conceptions of animals’ minds affected not only the scientific reception of sexual selection and female choice as theories but also the ways in which the histories of these theories have been negotiated” (p. 3). The book is comprised of six chapters beginning with Darwin and Wallace and their divergent views on the topic of mate choice and concluding with a fantastic chapter “Selective History: Writing Female Choice into Organismal Biology.” This, the penultimate chapter, describes the impact of Robert Trivers’ work, especially his 1972 paper “Parental Investment and Sexual Selection,” that reinvigorated the debate and reconnected the issue of female choice with the broader theoretical issues in evolutionary theory that had been raised by the work of W.D. Hamilton, John Maynard Smith, and George Price. The influence of this paper is succinctly and convincingly presented by Milam and its impact explained as “convenient one-stop shopping; he synthesized the complex mathematics of John Maynard Smith and William D. Hamilton with the experimental evidence of Claudine Petit and Lee Ehrman, illustrating how they could be applied to field research, in a short, easy-toread form” (p. 141). In this chapter, the connection between mate choice and the most fundamental questions in evolutionary biology are made manifest. Trivers’

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work formed the backbone of Dawkins’ Selfish Gene and Wilson’s Sociobiology, and Milam describes these connections well. This chapter is also the site of Milam’s strongest and most interesting historiographical arguments. In the section “Constructing the History of Sexual Selection,” she places the recent work on female choice and sexual selection in the context of the ongoing struggle between molecular and organismal biology. She argues compellingly that the success of Trivers’ 1972 paper along with the growth of the field of animal behavior studies developed “a new historical authority, wiping out a more interdisciplinary picture of the history of female choice” (p. 151) which her own work recovers. Indeed, Milam suggests that “the eclipsed history of sexual selection is an exemplary case of the practicing biologists’ fascination with the history of their discipline and reflects the disciplinary affiliations of sexual selection in the 1980s” (p. 157). Milam’s work provides insight into the myriad modes biologists employ in the pursuit of their research including the construction of disciplinary histories. A persistent theme is the role of both anthropomorphism (imbuing animals with human qualities) and zoomorphism (interpreting human behavior as a complex form of animal behavior). Milam demonstrates the significance of these theoretical frameworks but more importantly, she emphasizes their changing significance as the scientific and social contexts shift.

Ethology and Behavior As the historian Richard Burkhardt wrote on the centennial of Darwin’s death in 1982, the development of evolutionary studies of behavior were a long time coming. “Considering the wealth of behavioral evidence Darwin cited in support of the general idea of evolution, and the variety of ways in which behavioral phenomena were related to key features of his evolutionary theorizing, and his strong claims about the continuity of between the mental faculties of the higher animals and man, one might expect that Darwin’s work would have immediately launched a vigorous new field devoted to the evolutionary interpretation of behavior. This is not however what happened” (Burkhardt 1983, p. 433). Nevertheless, ethology and behavioral biology have been the focus of many historians working on contemporary evolutionary theory, Burkhardt chief among them. In his (2005) account of the development of the field, Patterns of Behavior: Konrad Lorenz, Niko Tinbergen and the Founding of Ethology, Burkhardt explores the intricate relations between the study of animal behavior and the development of evolutionary theory in the twentieth century. The great strength of the book is that it shows concretely how a discipline gets constructed and achieves recognition and respectability. Burkhardt devotes considerable space to the summary and analysis of books and articles, but he is also careful to stress the “contingent and erratic” (Burkhardt 2005, p. 472) in ethology’s development. The history of the discipline of ethology is as complex and convoluted as any other. As Burkhardt observes, Lorenz and Tinbergen had predecessors, but not precursors. They borrowed eclectically modifying and rearranging ideas to suit their needs. Burkhardt follows his protagonists as they struggle with the vagaries and often unedifying politics of grant-seeking, academic

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appointments, conference organizing, and journal editing. Not surprisingly, only relatively late in their careers did the two men obtain institutional positions commensurate with their distinction as scientists, Lorenz at the Max Planck Institute and Tinbergen at Oxford. Lorenz and Tinbergen’s stature within the biological community, culminating in their joint award, with Karl von Frisch, of the Nobel Prize for medicine and physiology in 1973, translated into major public recognition and discussion of their ideas and their evolutionary perspective. Indeed, the history of behavioral biology provides fertile fields for the analysis of the expanded evolutionary theory in the twentieth century. Burkhardt’s history of ethology engaged the question that had motivated much of the philosophy of biology of the 1980s and 90s, namely the level at which Darwin’s mechanism of natural selection worked. As Burkhardt pointed out, the influential biologist Julian Huxley had a particular focus “on the benefit of the whole rather than the individual members of the species” (Burkhardt 1992, pp. 134–135). The combined influence of Huxley and animal ecologist Charles Elton created in their student, Wynne-Edwards, a field biologist with an integrated approach to the study of animal behavior ecology. But as Mitman has commented, “In the early 1920’s, the mainstream of ecological research centered on either autecology, with its focus on the physiological response of individual organisms, or on community analysis” (Mitman 1992, p. 72). Wynne-Edwards was educated and trained at Oxford in the midst of this division. Furthermore, “What we now call ethology – that biologically oriented, comparative, and naturalistic approach to the study of behavior that we associate with Konrad Lorenz and Niko Tinbergen – did not begin to become a coherent enterprise until the 1930’s and 40’s, and even then its status was highly problematic” (Burkhardt 1992). Wynne-Edwards’ staking out of social behavior as a mechanism for population homeostasis evolved through group selection creates an interesting comparison to the ethological approach to similar phenomena pursued by Lorenz and Tinbergen. Nevertheless, Wynne-Edwards’ interest in behavioral phenomena, such as the nonbreeding of sexually mature adults, that was not easily identified and ran counter to Darwinian interpretation, precluded his inclusion in the developing field of ethology. Through the 1930s and 1940s Wynne-Edwards pursued his ecological studies of behavior focusing on the social structure of breeding populations. He was encouraged in this work by the developments in the modern synthesis that identified populations as the unit of evolutionary significance. The emphasis on higher-level selection, which Dobzhansky called group selection, (following Wright), was of obvious importance to Wynne-Edwards. As Gregg Mitman argued, “the past historical focus on such luminaries as Mayr and Simpson has given a false sense of closure to a number of issues in evolutionary biology that continued to circulate in the literature, especially where evolutionary thought intersected with ecology and behavior. . . While most biologists by the 1940’s believed natural selection to be the causative agent behind evolutionary change, still the question of what level (s) selection operated on remained a highly contested and unresolved point” (Mitman 1992, p. 111).

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Mitman’s approach to the history of ecology and evolution concentrates on W.C. Allee and his colleagues and context at the University of Chicago. These men sought to “biologize human sociability.” Beginning with the assumption that nature is normative, these Chicago ecologists believed that what they saw in nature justified, and even demanded, particular social, political, and moral results. Not an unusual position at the time, the view took a special nonevolutionary twist at Chicago as the focus remained on physiology, development, heredity, and behavior of existing groups of organisms rather than on selection or adaptation of the traits of individual organisms. This led to an emphasis on cooperation rather than individual Darwinian competition. Focusing on the leading researchers and their research programs at Chicago, Mitman shows that the local setting and particular institutional conditions allowed considerable “drift.” Thus, ecology at Chicago reflected the local setting and “represented the borderland between the biological and the social sciences through the study of interrelationships between and among individual organisms and their environment” (Mitman 1992, p. 1). The Chicago school developed theories of animal ecology which postulated and then established the animal community as the basic unit of ecological action and evolution, gave animals agency to shape their environments, and made cooperation a major principle of evolution. Population ecology and the evolutionary synthesis presented the major challenges. The core of Mitman’s work is an insightful working out of Allee’s and Emerson’s development of animal ecology and their response to the revival of the Darwinist program and its synthesis with Mendelian genetics. This was not rationalization – biologists working through scientific paradigms shaped only through “internal” history of ideas, and then awkwardly fitting their science to social and political issues. Mitman shows clearly how external social and political commitments shaped the disciplinary and theoretical components of their scientific work. Whether the family or the “local group” generated animal society, whether heredity or social organization formed caste hierarchy in social animals, whether social conditioning or inherited predispositions shaped aggressive behavior, and whether a reproductive population was an aggregation of individuals or a superorganism are among the issues in community ecology studies of termites, birds, mice, and monkeys that Mitman analyzes. Mitman’s major themes, and the most socially significant interests of the scientists whose work and lives he chronicles, are conflict and cooperation, hierarchy and dominance, adaptation to or transformation of environments by organisms, and arguments over whether the basic unity of evolution is the community, the population, the individual, or the gene. In general, he argues, these animal ecologists, much like a later generation of sociobiologists, believed that the study of nature would yield direct social insights. Their animal ecology had several major themes: social organization and interaction with the environment, populations as superorganisms, group selection as an important evolutionary force, and cooperation and stability as ends of interactions.

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Sociobiology In a section of Patterns of Behavior, entitled “Whither Ethology?” Burkhardt briefly covers ethology’s position and the effect of E. O. Wilson’s (1975) Sociobiology: The New Synthesis. Wilson predicted ethology would shrink and merge with physiology and psychology, dwarfed by integrative neurobiology and sociobiology and behavioral ecology. Wilson’s Sociobiology represented an attempt to understand all of animal behavior (ultimately including human behavior) in terms of evolutionary adaptiveness. Sociobiology created a storm of contention almost immediately. The controversy was perhaps the most publicly debated episode in the field of biology since the Scopes trial in Dayton, Tennessee, in 1925 (Segrestrale 2000). Although Wilson had many supporters, he also suffered a great deal of criticism from within the scientific community and without. Biologists criticized Wilson’s methodology and adaptationist reasoning, philosophers of science challenged his attempt to “biologicize ethics,” and sociologists, educators, feminists, and others deplored his apparent ignorance of the social and political implications of his work. (See, for instance, Lewontin et al. 1984; Kitcher 1985). In his autobiography Wilson wrote that the reviews of Sociobiology “whipsawed it with alternating praise and condemnation” (Wilson 1994, p. 330). Along with recollections of the wide-ranging response to his theory, Wilson also recalled something about his methodology. He wrote: “In order to use models of population genetics as a more effective mode of elementary analysis, I conjectured that there might be single, still unidentified genes affecting aggression, altruism, and other behaviors” (Wilson 1994, p. 330). Wilson’s sympathy for the gene’s eye view is apparent in this passage. His commitment to genic (that is, gene-based individual) selection was such that he rejected most of Wynne-Edwards’s thinking, arguing “that one after another of Wynne-Edwards’ propositions about specific ‘conventions’ and epideictic displays were knocked down on evidential grounds or at least matched with competing hypotheses of equal plausibility drawn from models of individual selection [emphasis added]” (Wilson 1975, p. 110). In the past 30 years, sociobiology, by that name or some alternative – behavioral ecology is popular for the animal world and evolutionary psychology for the human – has proven to be one of the most exciting and fruitful areas of evolutionary biology. It is also firmly Darwinian, based through and through on selection arguments, generally stressing the virtues of social behavior for the individual. Fascinatingly, however, E. O. Wilson of all people has recently revised his position with respect to group selection. In a recent paper with his long-time collaborator Bert Hölldobler, Wilson now argues that “group selection is the strong binding force in eusocial evolution” (Wilson and Hölldobler 2005). Indeed, Wilson subsequently co-authored a paper with David Sloan Wilson, another enthusiast for group selection, going even further down this path. In “Rethinking the theoretical foundation of sociobiology,” the authors say flatly: “Current sociobiology is in disarray. . .Part of the problem,”

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they continue, “is a reluctance to revisit the pivotal events that took place during the 1960s, including the rejection of group selection and the development of alternative theoretical frameworks to explain the evolution of cooperative and altruistic behaviors” (Wilson and Wilson 2007). The history of sociobiological thought is an area of active research and is ready for further expansion. The connection of contemporary sociobiology to nineteenthcentury work on social Darwinism was made early by Kaye (1986). Other authors focus on more contemporary developments, such as the reception of sociobiology in China (Jianhui and Fan 2003) or Wilson’s recent change of heart about kin selection (Gibson 2013). For an example of a culturally sensitive contextualization of pre-sociobiology argument on human nature, see Milam (2019). There has been no biographical treatment of Wilson outside of his 1994 autobiography Naturalist, but Ullica Segrestrale has written a biography of William Hamilton who made important contributions to evolutionary approaches to altruism (Segrestrale 2013), and Oren Harman has done the same for George Price, another champion of altruism (Harman 2010). The political and ideological dimensions of the sociobiology debate have been described by Neil Jumonville (2002).

Future Directions As is made clear in the editor’s introduction to this volume, these essays are not meant to be comprehensive, and this is certainly true of this essay. Rather, I have pointed to what I think are some of the most important areas of historical interest in the study of modern evolutionary theory. In the coming years, the historical exploration and illumination of the integration of evolutionary theory into all of biology and much of medicine will provide rich material. Recent developments in microbiology, experimental evolution, microbiome research, and genetics will entice historians into new areas of research. The history of evolutionary approaches to behavior (both human and animal) invites further analysis which addresses the broader themes of race, gender, and class and the ways that these issues have influenced biology and the role that evolutionary biology has played in constructing and sometimes legitimizing these categories. History of modern evolutionary theory should continue to expand, engage, and embrace these challenging and important realities.

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The Historiography of Molecular Evolution Edna Suárez-Díaz

Contents Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Actors, Fields, Traditions. . . Disciplines? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Debates, Confrontations, and Negotiation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Technologies and Techniques . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trends and Topics to Explore . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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Abstract

This chapter revisits the main themes and questions addressed by the historiography of molecular evolution in the last three decades. Being a recently consolidated field in biology, historians have grappled with a sizable number of actors’ accounts, and with questions concerning the disciplinary status of the heterogeneous group of practices that in the early 1970s became known as molecular evolution. Historians of this field have paid special attention to scientific debates concerning the mechanisms and patterns of evolution at the molecular level, and the and to the adaptation and construction of novel molecular techniques, from serology and electrophoresis to automatic sequencing and bioinformatics, to produce molecular data and analyses. Finally, the chapter provides a roadmap of trends and possible future directions for historical research of the molecularization of evolutionary biology.

E. Suárez-Díaz (*) Estudios de la Ciencia y la Tecnología, Facultad de Ciencias, Universidad Nacional Autónoma de México, Mexico City, Mexico e-mail: [email protected] © Springer Nature Switzerland AG 2021 M. R. Dietrich et al. (eds.), Handbook of the Historiography of Biology, Historiographies of Science, https://doi.org/10.1007/978-3-319-90119-0_6

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Introduction The molecularization of evolutionary biology has been a rich, though not fully exploited, source of interesting research for historians of biology specializing in the second half of the twentieth century. A rich ecology of practices and theoretical debates, embedded in some of the political concerns and policies of the postwar period, is part of the emergence of the field of Molecular Evolution. Such a broad subject has been reconstructed from a variety of historical perspectives but is still open to new perspectives and interpretations waiting for the active engagement of historians of science. Today, the practice of evolutionary biology involves quite a bit of molecular biology. Establishing phylogenetic relations, testing hypothesis or models of evolutionary processes, and measuring the amount of genetic variation within a population are all activities requiring the use of molecular techniques and/or the use of sequence data banks and analytical tools. This is a relatively recent development, starting between the late 1950s and the early 1960s, when advances in protein chemistry provided compelling evidence of huge amounts of variation at the molecular level; unexpected phenomena, such as the presence of large fractions of repetitive DNA in eukaryotic cells; and processes, such as gene duplication that broadened, and eventually transformed, the theories, concepts, and practice of evolutionary biology. New technologies and experimental techniques, such as zone electrophoresis, protein finger printing, protein sequencing, and nucleic acid hybridization, were incorporated in the comparative analysis of biological molecules, provoking scientists to question long-held ideas, including the “inviolability” of the relation between protein structure and function, as biochemist Christian Anfinsen put it in 1959. Soon, protein chemists, biophysicists, immunologists, physical anthropologists, and the newly labeled molecular biologists understood the potentialities of the new technologies for understanding both evolutionary patterns and mechanisms and organized the first meetings that gathered most of those interested in molecular evolution, including some that would be its most furious critics in years to come.1 These developments climaxed a few years later with the publication of two versions of the Neutral Theory of Molecular Evolution, by Motoo Kimura’s in (1968, 1969) and Jack King and Thomas Jukes’ in (1969). The subsequent neutralist/selectionist controversy on the nature of evolutionary mechanisms, and the publication of the first international journal entirely devoted to developments in

1

These include the Conference on Evolving Genes and Proteins, at Rutgers University on September 17–18, 1964, and the Colloquium on the Evolution of Blood Proteins that took place in Bruges, Belgium, in the summer of the same year (Dietrich 1994). The same year, Emilé Zuckerkandl and Morris Goodman, representatives of what would eventually be called “molecular anthropology,” were invited to the Wenner-Gren Conference in Burg Wartenstein, Austria, to a meeting on the taxonomy of humans and primates (Zuckerkandl 1964, Goodman 1996). The proceedings of these conferences provide some of the major documentary sources of this period, concentrating a large number of early actors requiring further investigation, and of the first responses of the organismic evolutionists to the molecular newcomers.

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this field, the Journal of Molecular Evolution in 1971. The field of molecular evolution came to be known for some of its most controversial and transformative concepts and hypotheses: the molecular evolutionary clock, which marks a stochastic yet uniform tempo for evolutionary processes; the relatively recent divergence time between the major Primates (man, gorilla, and chimpanzees); the primacy of molecular over morphological markers in the reconstruction of phylogenetic relations; and the contention that, at the molecular level, natural selection plays a minor role in the fixation of variation, and genetic drift is the main evolutionary force. Not surprisingly, the history of the ensuing debates between the new molecular evolutionists and the “classic” or organismic practitioners of the field has occupied a large portion of the historians of science’s research and publications to date. At the same time, there is much more to the historian of Molecular Evolution than a focus on debates about the mechanisms of evolution: the transformative quality of molecular phylogenies and its impact on the systematics of numerous phyla, as well as the study of genetic lateral (or horizontal) transfer, have challenged the traditional representation of the tree of life, producing further ongoing debates. These developments are only beginning to receive historians’ attention, including large sections in Jan Sapp’s books (2003, 2009), historical accounts by Maureen O’Malley (O’Malley and Boucher 2005; O’Malley et al. 2010; O’Malley and Koonin 2011; O’Malley 2016), and participants’ accounts, such as those in Jan Sapp’s (2005) collection, Microbial Phylogeny and Evolution: Concepts and Controversies and Doolittle (1999, 2005). This chapter aims to provide a guide to the secondary literature published on the broad subject of the history of Molecular Evolution, organized according to major themes and subjects. It makes passing reference to molecular population genetics (see ▶ Chap. 6, “Gregor Mendel and the History of Heredity”), and other closely related areas of population biology (see ▶ Chap. 2, “Charles Darwin and the Darwinian Tradition”), and concentrates on how historians have dealt with the actors within the field of molecular evolution, the socio-professional boundaries of their field, the techniques and technologies associated with the study of evolution at the molecular level, and the most controversial debates involving molecular evolutionists. Each part includes reference to preceding and recent trends. Finally, this chapter provides a historian’s view of the impact that new historiographical perspectives and topics are having, or may have, on this important field in the history of the twentiethcentury biology.

Actors, Fields, Traditions. . . Disciplines? For students educated in the last two decades, it is hard to imagine a time when scientific theories, their production, testing, and eventual acceptance into respectable frameworks for research, ruled historical accounts of science. Not surprisingly, the very first detailed account of the rise of molecular evolutionary studies was authored by an interested practitioner, theoretical population geneticist Richard

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C. Lewontin in his 1974 book, The Genetic Basis of Evolutionary Change (Lewontin 1974). As a mathematically oriented geneticist, Lewontin was concerned with the recent advocacy by Motoo Kimura of the Neutral Theory of Molecular Evolution (NTME) and the subsequent elaborations co-authored with his colleague, Tomoko Ohta (Kimura 1968; Kimura and Ohta 1971). For Lewontin, the NTME was but the most recent continuation of the classical theory of population genetics first advanced by J.B.S. Haldane, James F. Crow, and Hermann Muller, around the concepts of the cost of selection and the mutational load (see Beatty 1987; Paul 1987; Dietrich 2006, also see ▶ Chap. 6, “Gregor Mendel and the History of Heredity” by Müller-Wille, this volume). According to Lewontin, the neutral theory originated in the debate between the classical and balance schools of population genetics (dubbed as such by Theodosius Dobzhansky in 1955) that concerned the amount of heterozygosity in natural populations (Lewontin 1974). This was a particularly relevant theoretical controversy, given widespread concerns regarding atomic fallout and radiation-induced mutations, which has received due attention from historians of biology.2 The use of gel electrophoresis in 1966, by Lewontin and Jack L. Hubby in Drosophila pseudoobscura, and by Harry Harris in human populations, to discover large amounts of genetic variation was problematic for prevailing theories based on natural selection.3 In Lewontin’s view, after the electrophoretic results were published, the balance-classical controversy was not solved but transformed into a new controversy between Kimura’s neutral theory and selectionist alternatives. Michael R. Dietrich, in a seminal historical paper, called this the Lewontin Historical Thesis (Dietrich 1994). Dietrich provided convincing historical and conceptual arguments linking the origins of the NTME to both debates in populations genetics and developments in molecular biology; thus, according to him, “[t]he problems and findings of molecular evolution [. . .] are essential features of the neutral theory that cannot be traced to the classical/balance controversy” (Dietrich 1994, p. 24). Historical research has shown that Lewontin’s claims did not recognize the revolutionary impact of new molecular techniques, problems, and approaches, in the understanding of evolutionary mechanisms and patterns that started at the end of the 1950s. Lewontin himself later agreed with the more detailed narratives produced by historians. Two historiographical points should be made in relation to Lewontin’s reconstruction of this history and historians’ responses that will help us introduce our subject.

2

There is a growing and relevant corpus of secondary literature on atomic fallout and mutation that includes besides Beatty (op cit, 1993), and Paul (op cit), the work of Susan Lindee (1992, 2005), Soraya De Chadarevian (2006), Karen Rader (2006), Jacob D. Hamblin (2007), Angela Creager (2009, 2013), and Mateos and Suárez-Díaz (2015) to name a few. See Dietrich (2006) for an overview of twentieth-century evolutionary genetic debates. On the Neutral Theory of Molecular Evolution, see also Provine (1990) and Crow (2008). 3 There is also a contested story about the third team at the University of Texas, who published on the same subject (Johnson et al. 1966). See Powell (1994) and Suárez and Barahona (1996) for diverging accounts.

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First, in writing their histories, scientists not only provide an interested interpretation of their own role in history but a legitimation (either conscious or unconscious) of what they consider to be the relevant factors and aspects of their scientific practice. As historian of molecular biology Pnina Abir-Am has noted, the problem arises when these histories provide, what she called, a second-order legitimation of science (Abir-Am 1985). Historians must not uncritically accept scientists’ narratives regarding their own history. In the case of molecular evolution, Lewontin, inadvertently as it happened, provided a history of the NTME that centered on mathematical population genetics, with experimental practices (like electrophoresis experiments) subordinated to their traditional role of providers of evidence for the testing of theories.4 However, as Dietrich further specified, “Lewontin’s Genetic Basis of Evolutionary Change has been jokingly called ‘101 Ways to Save the Classical and Balance Positions’. In many ways, Lewontin [was] trying to save this controversy, the question of the nature of genetic variation is the problem that has driven his career” (Dietrich 1994, p. 57). Dietrich’s critical account of Lewontin’s reconstruction was followed by my own account, which put a heavier weight on the diversity of experimental, comparative, and theoretical sources that gave way to the NTME (Suárez and Barahona 1996; Suárez-Díaz 2009). A second historiographical point follows from this. In the 1990s, in tune with contemporary developments in the history of science, and influenced by sociological and philosophical developments, theories were no longer the main object of research for historians of science. Their place was taken by models, the diversity of experimental practices and the material culture of the laboratory, and the collection and classificatory practices more in tune with expeditions, cabinets, and museums (see section “Debates, Confrontations, and Negotiation” below). The diversity of the sciences was captured in the radical idea of its heterogeneity, that is, the recognition that science was made up of incommensurable non-reducible practices. Peter Galison and David J. Stump’s volume on The Disunity of Science (1996), John Pickstone’s working practices (1993), and Ian Hacking’s styles of scientific reasoning (Hacking 1992) exemplify attempts to account for the newly recognized diversity of legitimate scientific practices, endeavors, and products (see ▶ Chap. 10, “Biomedicine and Its Historiography: A Systematic Review,” on practices). These ideas shaped the way historians of science have written about the actors, fields, and traditions involved in the creation and consolidation of molecular evolution. This is particularly notable because, in its very name, “Molecular Evolution” evokes the synthesis or at least the “coming together” of two supposedly antagonistic approaches to the study of living beings. This very fact has made the entire field particularly deserving of historical treatment and surely was the source of some authors’ interest in disciplinary history and socio-professional identities. Some of the 4

In May 2004, at the Dibner Seminar on Molecular Evolution at Woods Hole Marine Laboratory, Massachusetts, Lewontin generously accepted the interpretation Mike Dietrich and I had developed, in which experimentally oriented fields and traditions played a major role in explaining the origins of the NTME. Jim Crow also acknowledged our interpretation (November 16, 1996, personal communication).

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basic historical questions we may ask include how was it possible that one of the most reductionist fields of biological practice (molecular biology) met and even embraced the most comprehensive problems and approaches of biology (evolution)? The partial and provisional answers some of us have given to this question in the intervening years have usually relied on different analytical tools and different sorts of accounts. Starting with Soraya de Chadarevian, who called attention to the intersection of experimental and cabinet or collection practices in the work of hematologist Hermann Lehman (De Chadarevian 1998), Bruno Strasser has exploited Pickstone’s idea of working practices and explored how the conflicting moral economies of protein sequence collections and protein experimental work constrained the rise of the first sequence databases (Strasser 2010b, 2011). In other types of account, debates and paradoxes have played a positive role in the negotiation, construction, and delimitation of scientific identities (Dietrich 1998; Aronson 2002; Suárez-Díaz 2007; Sommer 2008). Finally, the flexible interpretation of concepts according to distinct scientific traditions may have contributed to the socio-professional dynamics of an incipient discipline in the early 1970s (SuárezDiaz 2009; Dietrich and Suárez-Diaz 2016). Indeed, a striking feature, as compared to other areas of research in biology, is that the men (with the salient exceptions of two notable women, Margaret Dayhoff and Tomoko Ohta) entering the field of molecular evolution came from a genuine variety of disciplines. Molecular biologists did not call themselves as such until late in the 1960s (Abir-Am 1992; De Chadarevian 1996, 2002), and during at least the 1960s and part of the 1970s, professional identities of students of evolution at the molecular level were not at all clear for its practitioners. Protein chemists, molecular anthropologists, medical geneticists, biophysicists, geneticists, and population geneticists interested in evolution had, however, a few important things in common in the 1960s: the familiarity with the new molecular techniques of protein analysis (DNA hybridization was used, but DNA sequencing was far away in the future) and the incorporation of knowledge on the molecular basis of heredity. For all these practitioners, informational molecules (or semantides) arguably provided the most direct evidence for evolutionary patterns and mechanisms. Other concepts, like the molecular evolutionary clock, and the idea of neutral mutations, also provided the “glue” for the initial collective endeavor. Complicating things for the socio-professional identity of practitioners, during the first decades (1960s and 1970s), they were constantly caught in the contrasting views of classical or traditional evolutionary biologists and the new – now called – molecular biologists. Though this problem has been indirectly taken up, it remains an interesting historical problem for institutional history. The Journal of Molecular Evolution, first published in 1971, and edited by Emilé Zuckerkandl, was instrumental in the construction of a socio-professional identity (Lenoir 1993) and in providing a venue for theoretical and experimental research in molecular evolution. As defined by its goals and editorial board, the field also included research associated with exobiology and the early evolution of life, besides the study of intra- and interspecific genetic (molecular) variation, as it is more usually defined. A decade later, in 1982, Masatoshi Nei and Walter Fitch launched a second journal, Molecular

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Biology and Evolution. Apparently, there was an inner conflict between Zuckerkandl and Nei and Fitch, who in 1992 founded the Society for Molecular Biology and Evolution (described in Suárez-Diaz 2009). In 1992, a third journal was created, Molecular Phylogenetics and Evolution, whose editor in chief was molecular anthropologist Morris Goodman (Hagen 2010). These internal professional struggles have yet to be accounted in the historiography of the field. Equally interesting may be the fact that we are witnessing the dissolution of the discipline of Molecular Evolution, into a myriad of statistical tools and mathematical models, supported by an enlarged experimental and computational apparatus (Suárez-Díaz and Anaya 2008), of the sort Hallam Stevens describes in his history of bioinformatics (Stevens 2013), and Sabina Leonelli has described in relation to big data and data-driven research (2012a, b, 2014). Though disciplines have been displaced a bit in a world of multidisciplinary approaches, the history of how Molecular Evolution came to be one of them, and how it has – in many ways – dissolved into a globally distributed network of database users, could provide clues on the dynamics and structure of contemporary biological research. Disciplines, practices, and debates in science happen, nevertheless, because of individual actors. The agents involved in the socio-professional dynamics of molecular evolution have so far included a few well-known names, who have deserved unequal amounts of published research. With the exception of Linus Pauling, few biographical accounts exist to date of the main actors, and historical accounts of these scientists’ work concentrate on their technical, conceptual, and theoretical contributions.5 Biographical profiles and interviews of the main actors in the origins of molecular evolution, as well as short narratives of the main topics, are to be found at the website of the History of Recent Science and Technology, supported by the Dibner Institute for the History of Science and Technology at Caltech (with John Beatty, Michael Dietrich, and Jay Aronson among the participants), and the Dictionary of Scientific Biography (for instance, Dietrich and Crow 2007 on Motoo Kimura).6 Concerning their participation in the origins of molecular evolution, Pauling and Emilé Zuckerkandl have attracted a lot of historical research, with prominent roles in works by Michael Dietrich (1998), Gregory Morgan (1998), Edna Suárez-Díaz (2007, 2009), and Marianne Sommer (2008); available also is an interview of Zuckerkandl by Morgan.7 Other important actors and their contributions to molecular evolution also have deserved the historians’ attention. Morris Goodman’s 5

Linus Pauling’s life and contributions has been the object of several biographies, including Eaton (2003), Goertzel and Goertzel (1995), Eaton (2003) and Hager (2011). Lily Kay (1993) explores his multiple roles role in the consolidation of molecular biology; Hamblin (2007) makes reference on his participation in the atomic fallout debate. 6 http://authors.library.caltech.edu/5456/1/hrst.mit.edu/hrs/evolution/public/index.html (accessed April 14th, 2016). 7 http://authors.library.caltech.edu/5456/1/hrst.mit.edu/hrs/evolution/public/clock/zuckerkandl.html (accessed April 14th, 2016).

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immunodiffusion experiments have been the focus of research by Joel Hagen (2010), who also interviewed Goodman (an autobiographical account is Goodman 1996).8 Alan Boyden’s serological taxonomy has been reconstructed in its search for objective methods and in relation to earlier serology by Bruno Strasser (2010a); Roy J. Britten and Eric Bolton’s development of nucleic acid hybridization as a tool to measure “genetic affinity” among species has been studied by Suárez-Díaz (2001, 2013, 2014a). In one of the few gendered studies in the field, Tomoko Ohta has been studied in her participation in the neutralist-selectionist controversy by Tomoko Yamashita Steen (1996); also, Margaret Dayhoff’s work at the interphase of comparative (or collecting) and experimental practices, and her involvement in the creation of GenBank, has been the subject of Bruno Strasser’s research (Strasser 2010b, 2011). In Strasser’s view, gender and the moral economies of biomedicine played an important and constraining role in Dayhoff’s career. Vincent Sarich and Allan Wilson’s controversial participation at the debates concerning the molecular evolutionary clock have been the focus of Dietrich (1998). Jay Aronson (2002) and Jonatan Marks, himself a participant and historian, have dealt with the work of early molecular anthropologists, including John Buettner-Janusch, William Curtis, and Morris Goodman (Marks 1996, 2002, 2008). Paradoxically, the work of Motoo Kimura is waiting for a thorough understanding of his mathematical achievements in the context of Japanese-US relations but also in its contributions to the construction of stochastic modeling in mathematics. So far, Kimura has been the focus of works by Tomoko Yamashita Steen (1996), William Provine 1990, Michael Dietrich (1994, 2006), and geneticist and colleague James F. Crow (1995, 2008; see also Dietrich and Crow 2007). Other relevant actors remain to be explored in their different contributions to the field. These include biochemists Ernest Baldwin (a disciple of Frederick G. Hopkins’ “general biochemistry”), Marcel Florkin (though see Ross 2008), and Christian Anfinsen (though see Creager 2006, and Suárez-Díaz 2017); population geneticists Richard C. Lewontin, Masatoshi Nei, and Jack L. King (though see Dietrich 1994); bioinformatics pioneer Walter Fitch (though see Suárez-Díaz and Anaya 2008); biochemists Richard Dickerson, Emmanuel Margoliash, and Emil L. Smith (though see Aronson 2002 for the last two); and biochemist and exobiologist Thomas Jukes, to name a few. Some, like molecular anthropologist John Buettner-Janusch (1924–1992), are waiting for a proper account of their exceptional lives. See, for instance, Buettner-Janusch (1962). Finally, a word for those excluded by socio-professional boundaries must be said. They saliently include the small but prestigious community of comparative biochemists, who studied the evolution of metabolic pathways among biological groups, largely marginalized by the molecular biology bandwagon (see Anfinsen 1959, for a contemporary “explanation” on the priority given to molecular genetics

8

As part of the Oral History Project at the Dibner Institute, see the interview of Goodman, by Hagen: http://authors.library.caltech.edu/5456/1/hrst.mit.edu/hrs/evolution/public/goodman.html (accessed April 14th, 2016).

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and protein chemistry, against “classical” biochemistry). Florkin and Baldwin, to whom I referred above, were the main representatives of the interest in comparative biochemistry (Ross 2008), an approach later incorporated in Thomas Jukes’s book (1966) and in the development of exobiology and early evolution of life, a neighbor field associated with mainstream molecular evolution since the 1970s. This is, clearly, an area deserving further historical investigation, particularly with regard to the early work of Catalonian biochemist Joan Oró and Sri Lankan scientist Cyril A. Ponnamperuma;9 so far, these developments have received the attention of James Strick (2004), whereas the Cold War context of the NASA space program has been addressed by Audra J. Wolfe (2002).

Debates, Confrontations, and Negotiation For historians of science, the late 1980s and 1990s was a period of increasing interest in the role of debates in the construction of scientific consensus. No field in postwar biology generated more debates and contrasting views than evolutionary biology, probably because of the combination of its central place in biology, but also because of the relevance of population genetics in high-stake political issues, like the effect of radiation fallout on human populations, and public health as related to genetic risks. The arrival of the molecular vision of life in the 1960s made matters worse. Historian Pnina Abir-Am goes so far as to say that molecular biology challenged the centrality of evolution in biology (Abir-Am 1985). For historians of molecular evolution, a common theme has been to situate the ensuing debates within the broader distinction and conflict between traditional organismal biology and the new molecular biology of the 1960s (Dietrich 1998; Hagen 1999; Smocovitis 1992). But Abir-Am’s observations go beyond a clash of perspectives. John Beatty originally situated the contrast between the “classic” and the “new” biology in its proper political and economic context (Beatty 1990, 1994). This context, involving harsh personalities but also professional stakes and funding, has been described by naturalist Edward O. Wilson, in a biographical chapter properly called “Molecular Wars” (Wilson 1994). Participants were forced to articulate the distinctions and the territory of each approach, in order to accommodate the newcomers (Simpson 1964). Ernst Mayr, George G. Simpson, and Theodosius Dobzhansky, the so-called architects of the Evolutionary Synthesis, participated in what Dietrich calls “an unprecedented counterattack” (Dietrich 1998, p. 85). In this contested territory, historians of biology have paid attention, first and foremost, to the neutralist/selectionist controversy. The controversy lasted at least two decades, and its participants – particularly in the early stages – involved notable organismic evolutionists, like George G. Simpson, and molecular evolutionists like 9

There is solely a Catalan biography of Joan Oró (1923–2004) written by Miquel Pairolí in 1996, to which I have not had access, despite his relevant contribution to the field of the origins of life and the RNA world. An entire volume was dedicated to the memory of Ponnamperuma (1923–1995) by the Journal of the International Society for the Study of the Origin of Life, 28(2), in April 1995.

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Emilé Zuckerkandl, Vincent Sarich, and Allan Wilson (Dietrich 1998, 2006). It quickly evolved into a complex mathematical contest between Motoo Kimura, Tomoko Ohta, Jack King, and others on the neutralist side, while Lewontin, Ledyard Stebbins, Francisco Ayala, Walter Fitch, Charles Langley, Morris Goodman, John Gillespie, and others took the selectionist side, and still others caught in-between, like Crow (Dietrich 2006). In his historical account of the controversy, Michael Dietrich (1998) has pointed to the main issue at stake: the causal distinctiveness of organismal and molecular evolution, a theme taken up also by Provine (1990) and Crow (2008). Molecular evolutionists maintained that while natural selection was the prevailing force acting at the organismal level, genetic drift and the fixation of neutral mutants at constant rate (the molecular clock) were the main evolutionary cause at the molecular level. Classical evolutionists opposed this distinction, which ran against the integration of evolutionary biology under the principles of the modern evolutionary synthesis. According to Dietrich, molecular evolutionists invoked the “molecular/morphological paradox” to articulate their position within the field of evolutionary studies. By claiming that different mechanisms took place at different levels, they differed and negotiated the discussion with organismal evolutionists (1998, p. 87). Increasingly, the constancy of the rate of evolution, and not only the amount of variation in natural populations (as we saw in section “Introduction”), became the most potent powerful evidence for the neutralist band (but see section “Debates, Confrontations, and Negotiation,” on the “electrophoretic revolution”).10 Thus, the molecular clock became a crucial element in the origins of the controversy, and its development, with typical experimental and theoretical regress as depicted in the sociology of science literature. A thorough account of the origins of the molecular evolutionary clock has been written by Gregory Morgan (1998). But the clock itself was the result of previous and contemporary work by protein chemists, a topic that deserves thorough historical treatment. The idea that each protein may change at a specific rate had been explored by Christian Anfinsen (1959), who concluded that each individual protein, depending on its particular combination of restricted sites, its three-dimensional requirements, and the nature of its active site, varied to a given rate during evolutionary time. Historian of molecular biology and biochemistry Angela Creager has called attention to this particular aspect of Anfinsen’s work (Creager 2007, 2008). The idea was developed independently by other biochemists and taken to its ultimate consequences by Zuckerkandl and Pauling (1962, 1965b). The neutralist/selectionist controversy involved a number of different but connected issues, which have been the subject of several historical accounts. One of them concerned the systematics and evolution of humans and apes. Indeed, the It should be noted, as historians have argued (see previous section), that in his first version of the NTME, Kimura (1968) ascribed a higher rate of substitution at the molecular level as evidence of the role of genetic drift over natural selection at the molecular level. In King and Jukes’ version (1969) and in subsequent versions of Kimura’s theory, however, it was the constancy of the substitution rate, or the molecular clock, which was also at the center of the mathematical evolutionary model debate (Kimura 1969).

10

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level of confrontation between organismal and molecular evolutionists (and between neutralists and selectionists, not always the same) grew up from the entanglement of questions concerning the adequacy of molecular markers, with questions regarding the evolutionary relation between humans and apes, as shown by the secondary literature on this issue (Dietrich 1998; Morgan 1998; Aronson 2002; Suárez-Díaz 2007; Sommer 2008). Between the 1960s and the 1980s, the subject of human evolution seems to have invited the most contested exchanges and the most excessive rhetoric on the superiority of molecular markers in the study of evolution (Zuckerkandl 1964, 1965a; Suárez-Diaz 2007; on molecular biology’s rhetoric, see also Abir-Am 1992). Eventually, the contrasting positions evolved into a principled methodological dispute as to the superiority and causal primacy of informational molecules over organismal explanations, embedded in a rhetoric of promises that championed the view that one sole molecule would be enough to reconstruct the history of life (Aronson 2002; Suárez-Diaz 2007). Within this broad scenario, particular attention has been paid to the controversies concerning the use of the molecular clock by molecular evolutionists and anthropologists. The famous confrontation between Zuckerkandl, Goodman, and Simpson at the Wenner-Gren Conference in Austria, 1964, has been recounted by Dietrich (1998), while the rhetoric involved has been analyzed by Suárez-Diaz (2007) and Sommer (2008). Other participants of the debate in those early years included Emanuel Margoliash, Curtis Williams, and Ann Hafleigh, who confronted their molecular reductionism against G. G. Simpson’s argument in favor of a variety of traits, markers, and judgment in the taxonomy of the primates (Aronson 2002). As Aronson has shown, Simpson was not alone, and some early molecular evolutionists like John Buetner-Janusch and Robert Hill also argued that natural selection caused molecular evolution (Aronson 2002, p. 457). A couple of years later, starting in 1965, biochemist Allan Wilson started to collaborate with graduate student and anthropologist Vincent Sarich to establish the time of divergence between humans and apes. They first used the so-called micro-fixation technique. Later, like Morris Goodman, Sarich and Wilson also used immunological techniques to measure the amount of reactivity between the albumins of more than 20 species; they obtained similar results to Goodman’s, but they gave a different interpretation, one according to the molecular clock that drastically shortened the divergence time between chimpanzees, gorillas, and humans (Wilson and Sarich 1969). The experiments and the confrontation that followed have been narrated in detail by Dietrich (1998, pp. 105–109). Last, but not least, the writing of histories of molecular evolution would benefit from accounts where experimental techniques have taken center stage in a scientific controversy. One of those debates concerned the use of nucleic acid hybridization to solve the so-called Primate trichotomy (Marks 2002, 2008; Marks et al. 1988; Suárez-Díaz 2014a). Though this debate also shares a focus on human evolution, the core of it revolved around the technique, the nature of data produced, and the experiments’ calibration and its statistical treatment. While hybridization had proved to be a transformative technique in the classification of birds (Cracraft 1987; Sibley and Ahlquist 1990), its adoption in matters of primate evolution led to one of the

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most contentious debates in molecular anthropology at the end of the 1980s (Sibley and Ahlquist 1984, 1987; Sibley et al. 1990; Marks et al. 1988; Dickerman 1991; Suárez-Díaz 2014a). If we include bioinformatics tools, the amount of debates regarding the methods and techniques of today’s molecular evolutionary studies becomes explosive and a rich mine of sources and subjects for historians of science (Suárez-Díaz and Anaya-Muñoz 2008).

Technologies and Techniques How molecular technologies and techniques were introduced in, and produced for the study of evolutionary patterns and mechanisms, has been another important topic for historians of this field. At the beginning of the 1980s, professional historians had become interested in the history of molecular biology (see ▶ Chap. 7, “The History and Historiography of Eugenics,” by Weindling, this volume). Later, Soraya de Chadarevian and Harmke Kamminga referred to a broader process, the molecularization of biology and medicine (De Chadarevian and Kamminga 1998), the forging of new alliances between researchers, industry, and the state after World War II. In this context, the molecularization of evolutionary biology can be seen as part of the revolutionary transformation of postwar life sciences. In the case of evolution, the process of molecularization started in the late 1950s and early 1960s (this does not mean that previous developments cannot be seen as part of a longer history, as I argue below). The tools and results of protein chemistry research, and what eventually became molecular biology, provided unexpected data and phenomena that challenged deep-held beliefs and changed the practice of evolutionary biology. New experimental methods and instruments literally opened an epistemic space that in the 1970s and 1980s threatened to severe the molecular from the organismic understanding of evolution.11 Historical accounts of Molecular Evolution have emphasized the work of protein chemists in the discovery of structural and functional constraints in proteins and the role of technologies in the establishment of huge amounts of heterozygosity in natural populations (see section “Introduction”) and in the formulation of the idea that proteins could stand a large amount of variation without affecting their function (see section “Actors, Fields, Traditions. . . Disciplines?”). Those developments shaped the understanding and the evidence in favor of different evolutionary mechanisms at the molecular level. As we will see in this section, however, there is also a growing secondary literature on the early uses of molecular techniques and experimental practices in the construction of molecular taxonomies, that is, the knowledge of patterns of evolutionary change. Some of these tools, like one-century-old serology, but also zone gel electrophoresis, protein finger printing, or protein and DNA sequencing, were developed in neighboring fields like immunology and protein chemistry. The history of these techniques has been a subject for historians of molecular biology and molecular On the notion of epistemic space in the field of heredity, see Müller-Wille and Rheinberger (2012).

11

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evolution in their association with relevant actors (see section “Introduction”); most of them, however, still deserve a careful attention in their applications in different disciplinary and geographical contexts. A biologist’s extensive account of the myriad techniques used by molecular evolutionists is included in Powell (1994) and Avise (2004). Historical accounts of the technologies of the field usually make reference to the first attempts to use serological data at the beginning of the twentieth century. In 1904 George Henry Falkiner Nuttall (1862–1937), a well-known American-British bacteriologist and immunologist working at Cambridge University, published Blood Immunity and Blood Relationship, a book that became a common reference for those seeking a quantitative measure of the relationships between species. Bruno Strasser has given a brief account of his work as a crucial antecedent for Alan Boyden’s serological taxonomy (Strasser 2010a). Though Boyden’s methods influenced later molecular anthropologists, like Morris Goodman and Curtis Williams, Nuttall’s early attempts deserve a thick contextualization, since it is too easy to see him as a precursor of later work. Other isolated antecedents in need of proper historical contextualization include the comparison of crystallographic features of large molecules by E. T. Reichart in the early twentieth century (see Dietrich 1998). The 1930s and 1940s saw the radical transformation of much of physiology into biochemistry. In 1937 Ernest Baldwin published An introduction to comparative biochemistry, and in 1949 the Belgian biochemist and historian of science Marcel Florkin published a book on the biochemical evolution of metabolism, L’evolution biochimique (translated to English in 1949). In contrast to Nuttall’s isolated efforts, theirs was part of a collective endeavor within general biochemistry, which has been the subject of study of a chapter in the unpublished doctoral thesis by Sage Ross (2008). Comparative biochemistry, however, was left out of the limelight in the 1960s, when molecular genetics and informational molecules took professional primacy, a recurrent theme in the historiography of molecular biology, as studied by Abir-Am (1992) and De Chadarevian (1996). Moreover, Florkin’s attempt was organized around the notion of the “orthogenetic evolution of biochemical systems” (1949, p. 33). Nevertheless, the method of comparing the metabolic pathways among different groups of organisms was valuable and continued in Thomas Jukes’ monograph, Molecules and Evolution (1966). The methods of biochemistry became part of molecular evolution in the 1960s, no less because of the interest at the NASA in the development of exobiology, and the study of the origins of life at that time (by Oró and Ponnamperuma, among others, who participated, for instance, in the Viking program). Again, this is an entire field in need of historical exploration, and one in which there is an incipient but excellent historical literature on the contexts where these programs took place, particularly by James Strick and Audra Wolfe (for a broader context, see Krige 2010; Krige et al. 2013 to name a few). As we arrive to the revolutionary molecular techniques of the 1960s, the first one that comes to mind when talking of evolution is gel electrophoresis. Electrophoresis has been so important to the study of population genetic variation that Dietrich unequivocally calls it “the Electrophoretic Revolution” (1994, 2006). Gottlieb

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(1971), Avise (1974), Vesterberg (1989, 1993), and Powell (1994), for instance, are practitioners who have written useful historical accounts of the development of electrophoretic methods. As such, they provide celebratory technical accounts of the many variations and innovations belonging to this family of techniques. These works are full of suggestive paths of action for historians of science and technology, though they typically lack the skeptical historian’s eye. Lewontin (1991) and Oliver Smithies (1995) are also interested actors who have written anecdotally rich accounts of the early days of zone electrophoresis. These sources make clear the enormous diversity of material arrangements included under the rubric of “electrophoresis” and the particularities of those variations in relation to specific biological problems. For their part, historians of science Lily Kay (1988) and Howard Hsue-Hao Chiang (2009) have provided detailed historical research on the development of moving boundary electrophoresis (Kay) and zone or molecular sieving electrophoresis (Chiang). In Kay’s analysis, Arno Tiselius and the apparatus he first developed and built in Caltech are seen as an important piece in the advancement of the molecular vision of life. In Chiang’s, zone electrophoresis is a tool that opened the analysis of (mostly blood) proteins to individual characterization. Both Chiang and the scientific participants agree in one significant aspect: zone electrophoresis, in its myriad variations (paper, starch gels, polyacrylamide gel, two dimensional, and whatever variations), was a technology available to an unheard number of users and applications. Electrophoresis became a cheap, small (as compared with the long, roomy demanding Tiselius apparatus), mobile, and easy tool to learn and perform in order to measure intrapopulation genetic variation. Electrophoresis (or better, the family of methods falling under that name) effectively revolutionized the study of genetic variation in more than one way. Combined with other methods for chemical analysis, electrophoresis evolved into a set of useful techniques applied also in the study species relations. One such example is protein finger printing, which combined chromatography and electrophoresis and developed by Vernon Ingram in 1956 in the context of Pauling’s studies of sickle cell anemia and molecular disease. It was used by Zuckerkandl as a first comparative molecular method and immediately provided conflicting results, as has been accounted in the secondary literature on debates (section “Actors, Fields, Traditions. . . Disciplines?”). Another such example was immunodiffusion, a method developed by Morris Goodman, which has been given enough detail in Joel Hagen’s study of Morris Goodman (Hagen 2010). A different group of techniques includes the nucleic acid hybridization method developed by Eric Bolton and Roy J. Britten (Giacomoni 1993; Suárez-Díaz 2001, 2013, 2014a), a technique that illustrated the interaction between biophysics and evolution, whose aim was to give a measure of genetic similarity between species. Indeed, historical accounts of the role of molecular techniques in systematics share a common underlying theme in the historiography of molecular evolution: the continued effort to improve and develop methods that provide quantifiable, objective estimations of species affinity (similarity) by molecular evolutionists, a theme taken

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up in several works by Joel Hagen (1999, 2001, 2003), but present in most secondary literature of the field (see also Porter 1992). A word apart deserves the historical research on protein and nucleic acid sequencing. Moreover, though sequencing has been studied for its impact on the construction of phylogenies, historians have not attended the transition from electrophoresis to the use of DNA in studies of intrapopulation variation. At the end of the 1970s, John Avise introduced restriction enzymes to analyze DNA variation, creating a new source of data for the study of evolutionary mechanisms (see Powell 1994; Avise 2004). While this area of historical research clearly overlaps with the historiography of molecular biology (see ▶ Chap. 7, “The History and Historiography of Eugenics,” by Weindling, this volume), and genomics, there are some basic readings that a student interested in molecular evolution may want to revise. Cook-Degan’s (1994) and Keating et al. (1999) remain as standard and detailed narratives of automated sequencing and other technological aspects of the Human Genome Project. But the early history of sequencing has been addressed by Soraya de Chadarevian (1996, 1999), who has treated it as an overlapping endeavor of biochemists and molecular biologists. Stephen Hilgartner (2004) and De Chadarevian (2004) deal with the specific social arrangements where large sequencing projects are undertaken. Miguel García-Sancho also has written extensively on the long history of sequencing and its insertion on larger computing facilities (2010, 2011, 2012). There is a lot of ground left, however, and interesting historiographic and historical questions remain to be explored with regard to the role of DNA and genomic technologies in the study of evolution. For starters in the field of bioinformatics and evolution, the reference books on the history of computers in this context are Ceruzzi (1998) and Edwards (1997), while the introduction of computers into the biological sciences is the subject of De Chadarevian (2002 Chap. 11) and November (2012). But more specifically, the introduction of computers into systematics and comparative sequence analysis has been addressed by Hagen (2003), Strasser (2011), Stevens (2013, Chap. 5), and tangentially by Suárez-Díaz (2010). Also, the establishment of the first protein and nucleic acid databases linking experimental and comparative work has been addressed, mainly by Bruno Strasser in his accounts of the work of Margaret Dayhoff (2010b). The consequences of the genomic revolution for the biological sciences and its changing political and social context are thoroughly analyzed by STS literature on biomedicine, and excellent starting points are Beatty (2000), Adam Bostanci (2004), and Barnes and Dupré (2008). A participant’s account of the early days of databases is given by Temple Smith (1990). As with technologies coming from protein chemistry in the 1960s, it is hard to establish the borderline between molecular biology (or genomics) tools and those of molecular evolution, as more and more disciplines have come to rely on what Stevens characterizes as biological research “out of sequence.” The distinctiveness between genomics or bioinformatics in general, and molecular evolutionary tools, seems to lie in the particular algorithms and tools, as embedded in common software

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and individual statistical methods used to establish phylogenetic or similarity relations or to test evolutionary hypothesis. This is relatively unexplored terrain for historians of science and a thrilling window to what contemporary biological research means. So far, Michael Dietrich (1996) has addressed the use of Monte Carlo experiments in molecular population genetics and an unpublished reconstruction of multiple-hit models to account for point mutations in DNA (Dietrich 2008). Suárez-Díaz and Anaya (2008) offer an account of the methodological decisions incorporated in the construction of molecular phylogenies. In the case of modern databases, some of the first mining algorithms derived from practices of protein comparison (Suárez-Díaz 2010). Clearly, this is a field of historical inquiry that requires notable skills in computational sciences and bioinformatics.

Trends and Topics to Explore In this essay, I have presented specific areas and topics deserving attention from historians of science. Few characters and controversies have concentrated the historians’ interest since the 1990s, when molecular evolution first became approachable from the perspective of the history of recent science. Molecular evolution, however, is not so recent anymore. With time, perspective comes, and the chance to explore people, tools, and places left out from the preferred topics of decades past. New or older topics, however, also can be seen in the new light of changing perspectives in the historiography of the sciences. The study of molecular evolution would greatly benefit by studies exploring its deep ties with broader social and political processes taking place in the decades after World War II. By this, I mean that historians need to address the molecularization of evolutionary biology not only at the level of the introduction of molecular techniques into the study of evolution. A thicker account of the development of this field would come from an understanding of the forging of alliances between political and public health concerns and the study of the molecular basis of evolution (Anfinsen’s book title). Interestingly enough, the attention to this broader context also redirects the historical investigation to new geographical areas. Recent historical research has shown that during the early and mid-1960s, electrophoresis was being used to investigate human population variation in relation to medical genetics around the globe (Souza Sebastiao and Ventura Santos 2014; Lipphardt 2014; and Suárez-Díaz 2014b; see the introduction to the issue by Bangham and De Chadarevian and comments by Susan Lindee in that issue). This means that well before Hubby, Lewontin, and Harris had published their results within the frame of theoretical population genetics, measurements of genetic variation of human populations, besides the common serological studies of Mourant (1954) and Boyd (1963),were well entrenched as part of medical genetics and medical anthropology. Such developments belong to early applications of biomedicine to public health concerns in the United States and many other countries, particularly in Africa, Asia, and Latin America (Suárez-Díaz 2017). The new zone

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electrophoresis of the 1960s was the right tool for the job: mobile, cheap, and easy to use, not requiring large laboratory investments, or particularly sophisticated skills. Moreover, these studies took place with the help of national and international agencies, such as the World Health Organization. A very different perspective arises with the epistemological problems associated with the new practices of molecular evolution. Evolutionary biologists are nowadays educated in the skills needed for bioinformatics, in a broad sense. This is true for scientists working on phylogenetics and also for those who focus on population genetics. The concepts of traditional philosophy of science do not seem fit to cope well with the practices of current biology. Popular concepts such as “data-driven research” still need much philosophical reflection and historical depth (with Sabina Leonelli and Hallam Stevens’ work leading the way). And new problems related to the organization of (networks of) science need to be critically addressed by historians and other students (sociologists, ethnographers) of science. Data are produced, or manufactured, by subrogate laboratories in different parts of the world (like China and South Korea) with huge automated capabilities. Nevertheless, the need for curators and naturalists with a broad biological culture is much needed to give sense to the data deluge. In this frontier of biological research, historians of science may give proper context to the promises and pitfalls of the most recent technologies.

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5

The Historiography of Embryology and Developmental Biology Kate MacCord and Jane Maienschein

Contents Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82 Embryos and the Enlightenment of the Eighteenth and Early Nineteenth Centuries . . . . . . . . . . 83 Embryos and Evolution in the Late Nineteenth Century . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86 Experimental Embryology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88 Early Twentieth-Century Understanding of Embryos and Development . . . . . . . . . . . . . . . . . . . . . . 90 From Embryology to Developmental Biology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94 Nonmolecular Narratives in the History of Developmental Biology . . . . . . . . . . . . . . . . . . . . . . . . . . . 95 Evolutionary Developmental Biology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97 Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100

Abstract

Embryology is the science of studying how embryos undergo change over time as they grow and differentiate. The unit of study is the unfolding organism, and the timeline upon which embryology is focused is brief compared to the life cycle of the organism. Developmental biology is the science of studying development, which includes all of the processes that are required go from a single celled embryo to an adult. While embryos undergo development, so to do later stages of organisms. Thus, development is broader than embryology, and it focuses on the processes more than on the entities being developed. The fields overlap, and in some senses embryology gave way to developmental biology as new techniques

K. MacCord (*) Marine Biological Laboratory, Woods Hole, MA, USA e-mail: [email protected] J. Maienschein Arizona State University, Tempe, AZ, USA Marine Biological Laboratory, Woods Hole, MA, USA e-mail: [email protected] © Springer Nature Switzerland AG 2021 M. R. Dietrich et al. (eds.), Handbook of the Historiography of Biology, Historiographies of Science, https://doi.org/10.1007/978-3-319-90119-0_7

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and new questions, in particular genetic analyses and methods, allowed researchers to “see” more inside of organisms and manipulate the processes that are required for their unfolding. This article examines the ways in which developmental biology manifested from embryology, while also retaining aspects of the scientific goals and approaches of the earlier field of embryology. It also looks at the ways in which the study of both embryos and their processes of development have intersected with evolution, both in the nineteenth century and throughout the late-20th century emergence of the field of evolutionary developmental biology.

Introduction “Developmental biology” became a field by that name only in the 1960s, and at the same time, it became a complex cluster of often uncoordinated questions and methods. Before that time, researchers who studied the development of individual organisms tended to call themselves embryologists and to focus on the ways embryos arise and change over time. Historians have largely muddled the two together, though embryology involved different questions, methods, and interpretations focused especially on morphological change rather than the later more molecular and genetics-driven developmental biology. Some historians and biologists have assumed that the field only emerged when it was called “developmental biology” and that no serious research took place before. We show that this is not the case and that it is important to explore how researchers have for millennia sought to understand how individual organisms come into existence, grow, and become the appropriate kind of living thing. As a result of the complexity of the topic of developmental biology, in this essay we look at selected episodes to explore the history of understanding how organisms develop, and we set aside plants to focus on animal development. We largely skip the early modern period when curious people began looking through microscopes and applying a scientific method to their studies and focus mainly on the nineteenth and twentieth centuries. For the late eighteenth into the nineteenth century, much of the historical work was carried out by embryologists seeking to present and interpret their own predecessors (e.g., Needham 1934; Russell 1916, and others we discuss below). Their histories reflect their own predispositions about what was most important for embryology. More recently, historians have provided broader overviews to help illuminate some of the themes, questions, and methods involved in this earlier work. For the late nineteenth-century period, more historians have taken up the study of embryology as it relates to evolution in particular. Those historical interpretations reflect a recent resurgence in interest in connections between “evo” and “devo,” as evolution and development have been represented in the “evo-devo” movement. Study of development in the twentieth century has had much more attention, both recently and in previous decades. We explore the major themes that emerge from that

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study, including the importance of experimental methods and analytical approaches. As “embryology” became known as developmental biology in the second half of the twentieth century, molecular and genetics approaches took on a dominant role and brought new questions and methods that gave developmental studies a new focus. To some extent embryos largely disappeared from the story. They reappear at the end of the twentieth century, with in vitro fertilization, cloning, stem cell research, and most recently gene editing. We end this essay with some twenty-first-century topics to show how diffuse the study of development has become.

Embryos and the Enlightenment of the Eighteenth and Early Nineteenth Centuries General histories of science largely ignored embryology and development, but several review articles provide a useful overview of the themes. This section points to some of the leading thinkers about the history of embryology. Because these historians focus on different themes, and do so in different ways, we point to the most significant contributions as a way to get at how the topic has been interpreted. Historians Tim Horder and Nick Hopwood offer recent valuable overviews, while embryologists looked at their own history in many of the earlier studies of the period. A summary article by Tim Horder on “History of Developmental Biology” reviews the early science and provides references that we need not repeat here (Horder 2010). He noted the contributions of Aristotle, Leonardo da Vinci, William Harvey, Marcello Malpighi, and other later luminaries as well. Horder succinctly pointed out that “Of progress in the eighteenth century, it is easy to conclude simply that theory ran ahead of data.” (p. 3) In contrast, later work became more based on observation, description, comparison, and eventually experimentation. While theory played a leading role in investigations of embryos, the early period through the eighteenth century was not devoid of empirical contributions to understanding organismal development. One line of research involved examining how the contributions of male and female parents come together to make one individual or how sexual reproduction occurs. Aristotle had considered this problem but was only able to conclude that the fluids from male and female combine – through the action of the four kinds of causes (material, formal, efficient, and final) that he thought operated on all phenomena. The question of just how fertilization and reproduction work only received close attention starting in the late eighteenth and nineteenth centuries. Another insightful review article, “Embryology” by Nick Hopwood (2009), shows the rich diversity of research into development that falls under the broad umbrella of that title. Medical study of reproduction and development, comparative embryology, and evolutionary questions all brought different sets of questions and methods. There was, Hopwood showed, no one clear discipline or established institutional home for embryology or development. This complexity has affected the historical research as well, as historians have concentrated on particular areas and

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ignored others (Hopwood 2009). Looking back to the earlier twentieth century, the history of reproductive research gained the attention of zoologist Francis Joseph Cole. He became fascinated by the history of protozoology and of sexual reproduction, and his 1930 volume Early Theories of Sexual Generation provided a starting point for generations of biologists and those interested in history of biology. In his own research as well as his historical studies, Cole showed that the process of development of an individual animal has a starting point, namely, with the coming together of the egg and sperm, and that there is a regular and predictable process of generation that occurs as a result. His focus on the early history of understanding that early development revealed the importance of microscopic observations and materialistic thinking in order to see the processes involved. By the mid-nineteenth century, the early ideas had led to an understanding of the process by which a sperm cell fertilizes an egg cell to produce a fertilized egg and thus an embryo. Historian John Farley (1982) picked up on that theme and carried it forward into the early twentieth century with his Gametes and Spores: Ideas about Sexual Reproduction 1750–1914 that introduced the variety of different ways of thinking about germ cells and reproduction during that period. At about the same time that Cole published his historical study, embryologist Joseph Needham published his own detailed three-volume study, Chemical Embryology. His work on that massive project led him to reflect also on the historical study of embryos, and he included chapters on the topic. At the same time, he delivered a series of historical lectures at the University of London. A few years later, in 1934, he published those lectures as A History of Embryology, which he revised for a 1959 edition (Needham 1934). There he looked at themes in embryological research up to the end of the eighteenth century, which provided a background for the way researchers in the early twentieth century were approaching problems of development. Needham showed that the study of embryos had a rich history and had long raised curiosity among leading and less well-known researchers. Careful observation and experimentation in order to create new conditions, which expanded the range of what could be seen, added to the theoretical background that Horder noted. After giving us rich descriptions of a sequence of observations, in the last section of the book, Needham pointed to themes that continued into the nineteenth century. In particular, competing theoretical frameworks for the research started from competing assumptions about whether development of an individual from its earliest starting point occurs gradually, with form and function emerging over time out of unformed matter in motion (the epigenetic view) or whether the earliest stages were already laid out in some way and simply grew larger (the preformationist view) (Maienschein 2005; Roe 1981; Bowler 1971 on preformation and epigenesis). Needham noted that he had initially titled his lecture series “Speculation, Observation, and Experiment,” but he realized that experimentation did not really come to embryology until the end of the nineteenth century with Wilhelm Roux. He suggested that he would hold a discussion of that period for a second volume to appear later, though he apparently did not publish such a book (Needham 1934, p. 230). Around the same time that Needham was working out his ideas about the history of embryology, Edward Stuart Russell was also thinking about how life develops.

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Trained as a classicist and zoologist, Russell turned his attention to philosophy of biology and especially to questions about what organisms are and how they develop. His Form and Function appeared in 1916 and his The Interpretation of Development and Heredity in 1930 (Russell 1916, 1930). This second book asked how an individual develops from an unformed material into a complex and organized organism. And to what extent, Russell asked, is the result an organized whole that is more than or different from the sum of the parts? His questions and his approach involved reviewing ideas of the past and examining current discussions, including explorations of the idea of an “organism as a whole” that had become popular with Jacques Loeb, William Ritter, Charles Manning Child, and others of the time. Another contribution from the 1930s that also focused on earlier periods of embryology is embryologist Arthur William Meyer’s 1939 volume The Rise of Embryology, which focuses especially on the contributions of early embryologists up to the work of the epigenesist Karl Ernst von Baer (Meyer 1939). As Meyer noted, the 1930s had attracted considerable attention to embryology, but the historical growth of the ideas remained largely unexamined and unfamiliar. Perhaps in part because of the increased interest in developmental questions and research, a few scientists such as Meyer and Needham turned their attention to reviewing and interpreting the historical study of embryos and development as well. Though Meyer continued his study of von Baer and von Baer’s contemporaries, which led to the 1956 volume Human Generation: Conclusions of Burdach, Döllinger and von Baer, it was not until the 1960s that we again see such concerted attention to the history of studies of development (Meyer 1956). At that point, embryologist Jane Oppenheimer led the way to serious scholarly study of the history of embryology. While Cornell zoologist and historian Howard Adelmann spent much of his career focusing on the publications and correspondence of Italian embryologist Marcello Malpighi (Adelmann 1966), which resulted in the definitive study of this outstanding Italian anatomist’s meticulous anatomical studies, Oppenheimer looked at embryology more broadly. Some of the essays that Oppenheimer collected into an important volume signaling the new field of history of embryology include essays on the classic early figures: William Harvey, William Gilbert, Sir Thomas Browne, John hunter and his brother William Hunter, and Karl Ernst von Baer. One essay focuses on Ross Granville Harrison, who headed the department at Yale where Oppenheimer had studied and whom she shows as having established an experimental program of research that provided an exemplar for what embryology had become by the twentieth century. Other chapters look at crosscutting themes, assessing what questions embryologists were asking and with what methods. These works offer descriptions of central ideas and contributions in understanding development and also provide historical interpretations that begin to place the biological research in the context in which it developed. As Oppenheimer noted: “a number of us who are working embryologists feel that our life in the laboratories is made more meaningful to us when we know something about our intellectual forebears” (Oppenheimer 1966, p. v).

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While Meyer and then Oppenheimer did an excellent job of first interpreting and then making von Baer’s work available to twentieth-century English-reading audiences, later generations of professional historians of science looked at von Baer and the early nineteenth-century period in which he worked in different ways (Oppenheimer 1986). Timothy Lenoir’s study of different forms of vitalism and teleological thinking, including the work of Friedrich Blumenbach, Karl Friedrich Kielmeyer, and von Baer, emphasized their search for laws and explanations about fundamental issues concerning life (Lenoir 1989). While not everyone accepted his interpretation, Lenoir certainly opened a discussion about central questions related to biology. More recently, Sabine Brauckmann has been working through von Baer’s published work and his archives with meticulous care to explore his ideas about development and also about evolution (e.g., Brauckmann 2008). In collaboration with embryologist and historian Scott Gilbert, Brauckmann has looked at von Baer’s descriptions and understanding of gastrulation (Brauckmann and Gilbert 2004). And others, such as Brian Hall and Fred Churchill, have looked more closely at the emergence and understanding of germ layers in development (Churchill 1991; Hall 1998). As MacCord explains (MacCord 2013), embryologists began to understand the importance of germ layers in defining the appearance of form from what starts initially as unformed material. For most animals, the process of gastrulation, which initiates the formation of germ layers, marks a change that leads to visible differentiation and begins a period of growth. It also, developmental geneticists realized later, comes along with increased gene expression, but that is another story.

Embryos and Evolution in the Late Nineteenth Century Von Baer remained focused on embryos and did not accept the idea of evolution by natural selection. Yet he and other embryologists recognized that the development of individual organisms is shaped by history in some way, if only because a past creator set out the way individuals develop. That simplistic account did not provide sufficient explanation for most researchers interested in development through the middle and second half of the nineteenth century, and they had at least to think about what evolution might mean for development. Biologists who studied development after Darwin at least placed evolution in the background of their study of embryos, and some tried in various ways to bring embryology and evolution more closely together. Those attempts tended to focus on morphological change, and they ran up against limitations in the methods available. It was only in the later twentieth century that researchers sought explicitly to bring development and evolution into the same explanatory framework through “evo-devo” or “devo-evo,” as we discuss later. Darwin’s theory of evolution changed the thinking about biological organisms in general, and he also gave embryology a special role in providing information about evolutionary relationships. Yet he had little to say about the developmental process itself. In Chapter XIII of his On the Origin of Species, Darwin pointed to embryos as

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especially valuable. He had observed a variety of embryos in different species and was aware of some of the contemporary work such as von Baer’s. Pointing to what seemed to him clear similarities among embryonic forms of different species, Darwin concluded that “the leading facts in embryology, which are second in importance to none in natural history, are explained on the principle of slight modifications not appearing, in the many descendants from some one ancient progenitor, at a very early period in the life of each, though perhaps caused at the earliest, and being inherited at a corresponding not early period. Embryology rises greatly in interest, when we thus look at the embryo as a picture, more or less obscured, of the common parent-form of each great class of mammals” (Darwin 1859, pp. 432–433). To reiterate, embryology was thought to be important for understanding evolution and in particular the relationships among organisms. By implication, knowing those evolutionary relationships might help illuminate why embryos were similar, but Darwin’s evolution did not provide any explanation for why embryological development occurred as it did nor information about the processes involved. Ernst Haeckel provided the strongest claims that the results of evolutionary history actually define and direct development. For this outspoken German zoologist, “ontogeny recapitulates phylogeny.” In other words, the developmental process of an individual organism will repeat or “recapitulate” the developmental process of the phylogenetic group to which that organism belongs. Evolutionary history started to provide an explanation for embryology, according to Haeckel. Yet his evidence remained largely theoretical and based on the apparent similarities that Darwin had also observed. Only recently have professional historians of science turned their attention to Haeckel, and Robert Richards and Nick Hopwood in particular give us very vibrant views about this larger-than-life zoologist, his work, images, and impact (Richards 2008; Hopwood 2015). In his day, Haeckel was reviled or revered, largely depending on the reviewer’s own theoretical leanings. Those inclined toward the social implications of Haeckel’s world view, grounded as it was in materialism and a sense of progress for individuals as well as species, praised his work and its popular impact. Researchers immersed in the details of meticulous study of cells and processes of development, such as Wilhelm His, were appalled by Haeckel’s tendencies toward bold and expansive conclusions. Much of the historical study of Haeckel and his recapitulation ideas are concerned with the social impacts of his theorizing. One result of Haeckel’s popularity and accompanying controversy was a shift to marine studies. Haeckel himself studied a variety of marine organisms, seeking to get at the “Urform” that Haeckel believed to be the earliest form of animal life and from which all subsequent forms arose. He was confident that these would be found in the sea and therefore observed, drew, and compared a variety of marine invertebrates. As a result, he gathered his collecting buckets, waders, and other equipment and headed off to the seaside to collect organisms in jars and study them at the tables of the rooms he rented. His critics began to do the same, sometimes also looking for evolutionary relationships and sometimes looking instead at fundamental biological descriptions of form and function.

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Especially important for marine studies was Anton Dohrn, a graduate from the University at Jena where Haeckel taught. Dohrn heard about evolution and became an enthusiastic convert. He also became convinced that studying marine organisms would help reveal both fundamental living phenomena including embryonic development as well as evolutionary relationships. Thus, marine studies would illuminate both ontogeny and phylogeny. And so Dohrn used his tremendous energy to attract the funding and moral support that led to founding the Stazione Zoologica in Naples in 1872. Longtime Stazione archivist Christiane Groeben has given us many historical resources, cited in her 2013 summary, to help understand the importance of that special place and the kinds of work done there (Groeben 2013). The earliest work started with natural historical descriptions of development of different organisms and comparison across them. Haeckel inspired and was inspired by another German zoologist, August Weismann. As a passionate naturalist, Weismann studied a wide diversity of living organisms in his efforts to understand how they work. In his masterful book, August Weismann: Development, Heredity, and Evolution, historian Frederick Churchill discusses of Weismann’s interactions, friendship, and frustrations with Haeckel and most other leading figures in zoology of his long life. Churchill spent 50 years in careful study of Weismann’s research and shows us that leading nineteenthcentury biologists did not think of either genetics or embryology or evolution as separate. Rather, the phenomenon of development and life in general intertwines all those factors, and the goal for the science of life should be to interpret the interconnections. Churchill also showed that Weismann kept one foot in the conceptual traditions of the nineteenth century and another in the experimental and analytical traditions that came to dominate the twentieth century (Churchill 2015).

Experimental Embryology It is not the case that the period around 1900 brought experimentation to biology for the first time. Yet the push for “Entwicklungsmechanik,” or developmental mechanics, in particular made experimental approaches and mechanistic interpretations the norm for developmental studies. In 1964 with a first edition, and then in 1974 with a second and expanded edition of Foundations of Experimental Embryology, Jane Oppenheimer and fellow embryologist Benjamin Willier provided a set of classic papers that they felt helped define experimental work in embryology. The first of these came from Wilhelm Roux and was his 1888 manifesto for experimentation (Willier and Oppenheimer 1964). In introducing the Roux article, the editors explained that when comparative and descriptive methods had become inadequate for explaining the development of individual organisms, Wilhelm Roux led the way when he “founded a new discipline, causal analytical embryology, which he called developmental mechanics” (Willier and Oppenheimer 1964, p. 3). Roux also founded a new journal for the field and expected leading embryologists to publish their work there. Most did, but Roux proved himself such a heavy-handed editor that other journals arose as

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alternatives, including the Journal of Experimental Zoology with Harrison as the managing editor. Willier and Oppenheimer explained the way Roux had punctured one of two cells in a developing frog embryo to determine whether each cell could develop on its own or only as a part of the whole. Roux had concluded that the embryo acts as a mosaic of differentiated cells, each with its own “fate.” Though it turned out that Roux’s theory overreached his empirical data, this experiment nonetheless has remained an important introduction for students of embryology. In part, this may be because of the way that Willier and Oppenheimer introduced the work to a wider readership. As embryologist turned historian, Oppenheimer especially looked closely at the ideas and methods of the embryologists she studied. The esteem in which she was held by professional embryologists gave her an authority that other historians might not have had. It took a few more decades for professional historians of biology to emerge and become accepted, at least among historians of science even if not always among the embryologists themselves. The professionalization of the history of biology as a new academic field within the history of science brought additional perspectives. In particular, Garland Allen and William Coleman, two graduate students working with Harvard historian of science Everett Mendelsohn, suggested that the late nineteenth century might be viewed in terms of a “revolt from morphology.” Coleman introduced the idea at the end of his 1971 book, Biology in the Nineteenth Century: Problems of Form, Function, and Transformation. Intended as an introductory volume in a series for students and nonspecialists, the volume became a starting point for students in the history of biology. Coleman emphasized what professional historians were coming to see as a distinct break at the end of the nineteenth century, namely, as a move away from the reliance on historical methods and toward experimentation as the new ideal for biology. “To the embryologist, the bacteriologist, or the student of heredity and variation,” Coleman wrote, “the model was truly the new ideal. In its name – experiment – was set in motion a campaign to revolutionize the goals and methods of biology” (Coleman 1971, p. 166). Following Coleman, Garland Allen, took the turn to experimentation theme further in his contribution to the same introductory series in his 1975 book, Life Science in the Twentieth Century (Allen 1975). For Allen, the turn of the twentieth century saw a “revolt from morphology” that was truly revolutionary and that cast out the earlier morphological, descriptive, and historical approaches. According to Allen’s interpretation, a younger cohort of embryological researchers, including Thomas Hunt Morgan, Edwin Grant Conklin, and Edmund Beecher Wilson, rejected the natural historical traditions of the past and pushed for experimentation. Experimental embryology provided Allen’s central case for this revolt. Although other historians questioned aspects of Allen’s interpretation, especially the notion of a revolt or clean break between historical or morphological life sciences and experimental sciences (Benson 1981; Maienschein 1981; Rainger 1981), it was clear that the study of development had begun to change in important ways around 1900 (Maienschein et al. 1981).

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Early Twentieth-Century Understanding of Embryos and Development The 1980s brought increased attention to the history of embryology from embryologists and historians alike. Timothy Horder, Jan Witkowski, and C. C. Wylie edited A History of Embryology (1986), a volume for the British Society for Developmental Biology that both reviewed older themes and introduced many new ones. Though the title includes “embryology,” the set of essays included analysis of biochemical, molecular, morphological, physiological, and other perspectives on the understanding of development. The focus remained largely on early twentieth-century themes, and the editors brought together biological, historical, philosophical, and sociological points of view. In his review of the volume, Gilbert noted that, “There is a great need for a major new history of embryology, as developmental biology, the anagenic descendant of embryology, is becoming pivotal to all areas of biology. It is in a remarkable period of growth, expanding in one direction into the molecular basis of gene regulation and in another direction into the developmental basis of evolutionary change” (Gilbert 1987). The edited volume and other work of the time pointed to a number of themes emerging among historians by the 1980s that deserve a closer look, including among many others: the role of research institutions like the Marine Biological Laboratory (MBL) and Stazione Zoologica and University of Chicago, the importance of cells, regeneration, the organizer, metaphors and interpretation, and social views of embryos. While the earliest historiographic approaches, often by biologists reflecting on their own work, had emphasized the role of individuals and the ideas and interpretations, professional historians increasingly adopted a wider range of questions and methods. Individual scientists and their ideas remain important, of course, but often as part of larger institutions, changing scientific practices, and social influences. Indeed, historians have come to recognize the importance of particular people, places, methods, organisms, concepts, and other choices in the history of embryological work. Social factors, metaphors, ethical worries, and other considerations outside the parameters of laboratory work, some historians have argued, have helped to shape the study of development. We can see the diversity of historical approaches with some examples. In efforts to identify institutions important for the emergence of professional biology, historians have looked especially at the Stazione Zoologica in Naples (Groeben 2013) and the MBL in Woods Hole, Massachusetts (Maienschein 1985, 1989). As historian of science, Philip Pauly put it, “By the middle of the 1890s, an important group of American scientists and academic administrators, centered around such leading universities and colleges as Johns Hopkins, the University of Chicago, Columbia, Bryn Mawr, and Harvard, believed in the existence of a science of biology” (Pauly 1988, p. 121). Increasingly, these scientists went to the seashore to explore life and to carry out their biological research, with embryology as one of the core fields of study (Lillie 1944; Maienschein 1989; Benson 2001). They also found new research-oriented universities like the University of Chicago especially

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welcoming, and Chicago has had close ties with the MBL since its beginning (Maienschein 1988). These institutions provided a place to train graduate students by letting them jump in and carry out research themselves, embracing the importance of “learning by doing” and engaging in “hands-on discovery.” They also allowed biologists from different institutions and different specialties to come together in the summers and learn from each other, often developing long-term friendships and research collaborations. Marine biology lent itself to studies of developing individuals, through embryology, and also to study of cells as a way of studying, step-by-step, the processes of individual development. Embryologists at the end of the nineteenth century observed and compared many different types of embryos to observe the emergence of form. They saw patterns repeating regularly in different organisms, and they described in detail what they came to call the developmental stages (Hopwood 2007). In addition to the marine invertebrates and more accessible species of amphibians and such favored by zoologists, medical researchers had a fascination with human embryos. The Carnegie Institution of Washington Embryology Department gathered human embryos and fetuses in a collection that was later housed at the Walter Reed Army Medical Hospital and then also digitized and made available virtually through several venues (Maienschein et al. 2004). Such stages and collections remained a standard for those studying development until the advent of photography, films, and other technologies became the standard for displaying what counts as normal or pathological in development. Around 1900 cell lineage studies brought together study of cells with study of development to trace how cells change and give shape to embryos and organisms. Comparing these changes in cells across different organisms provided a source of information about development and possibly also about evolution, though it soon became obvious that the cells vary widely and the approach had limitations. The close examination of cells by Oscar Hertwig and Theodor Boveri in Germany, and Edmund Beecher Wilson and his colleagues such as Edwin Grant Conklin and Charles Otis Whitman in the USA, provided excellent observations and interpretations of what was happening as one cell divided into two, into four, and so on. Only recently have historians begun to look more closely at the history of cell biology in its own right, and the ways that study of cells and study of development were connected remains a largely untapped field. Almost a century since cell lineage research connected the study of individual cells to organismal development, human embryonic stem cell research opened new doors and new questions for biologists and for historians and led to regenerative biology as a hot topic, starting in 1998, and the first human stem cell research. Regeneration was not a new topic, but it gained new attention. In fact, Thomas Hunt Morgan had carried out a number of studies on regeneration in the 1890s, and in 1901 he published a volume that summarized research to date that showed a strong fascination among biologists for the powers of regeneration. The questions for Morgan and his predecessors were: Which organisms could regenerate, under what conditions, and how did regeneration work? (Morgan 1901). Almost a century later, biologist Charles Dinsmore enticed a set of biologists and historians to explore

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the history of studies of regeneration, with much of the discussion focused on the periods leading up to Morgan’s work (Dinsmore 1991). Stem cell research has since added to this discussion by directing thinking about regeneration into the context of often-exaggerated hopes for regenerative medicine. We are likely to see more historical work on the topic in the future as the science develops further. As those studying regeneration raised questions about how organisms could respond to changing conditions to regenerate missing parts, others used experimental methods to create new conditions in order to discover the effects of disturbing development. Perhaps the most important of these methods for the first half of the twentieth century was transplantation. Hans Spemann in Germany and Ross Harrison in the USA (after he received a medical degree in Germany as well as a Ph.D. in the USA) each used amphibians for their transplantations. Spemann cut out parts that would normally give rise to eyes or ears or limbs in one individual; then he transplanted them to another individual to see what would happen. Would they develop as normal eyes or ears or limbs, but in the new place? Or would they adapt to their new surroundings and blend in with the local tissue? The pieces had a strong tendency to retain their initial capacity, and yet they also adapted significantly – or else they would not have developed at all. This suggested a combination of intrinsic capacity to become a particular part, along with a strong ability to adapt to changed conditions. Development, clearly, is a complex process. Spemann’s work was well-known, and in 1935 he received a Nobel Prize for the work the he had begun with his student Hilde Proescholdt (later Mangold); unfortunately she died before the Prize was awarded (Hamburger 1984; Horder 2001; Sander and Faessler 2001). Meanwhile, Harrison was also transplanting cells and tissues. In 1907, he transplanted nerve cells (neuroblast cells) completely out of a frog and into a dish with frog lymph. The cells kept differentiating as they would have done normally inside the organism, with the fibers reaching out to make neural connections. With this work, Harrison showed the power of tissue culture and also of apparent selfdetermination of cells under some conditions. The work of Spemann and Harrison led Spemann to hypothesize the existence of a process of induction, through which an “organizer” induces surrounding cells to begin differentiation at the appropriate times and under the appropriate conditions. The search for the organizer and resulting experiments led to what Harrison referred to as a “gold rush” for embryological research (Hamburger 1988; Maienschein 2010). Such work dominated embryology through much of the 1930s, despite the fact that Johannes Holtfreter challenged the concept of a specific organizer and showed that many different kinds of tissue, including inert material, could stimulate the same result. Holtfreter’s questions strengthened calls by others for more sophisticated biochemical studies. By the 1950s, the gold rush of morphological experimentation gave way in part to molecular approaches and developmental genetics (Hamburger 1996). Despite the great enthusiasm for experimental embryology, and despite the prominence of Spemann and Harrison in the field, little historical analysis of the period appeared until Jane Oppenheimer identified it as an important topic.

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Then Horder, Allen, Maienschein, Gilbert, and others undertook historical explorations from different perspectives (Oppenheimer 1967; Horder 2001; Allen 1975; Maienschein 1991; Gilbert 1991). Yet the most important historical study of Spemann and his approach is surely by distinguished neuroembryologist Viktor Hamburger. Late in his career, Hamburger turned his considerable talents to history, in particular examining the lab and director under whom he had received his Ph.D. Few scientists have so successfully carried out such a challenging professional project, including discussions of the personal side while also developing a critical distance for the historical analysis. Hamburger’s use of two different type fonts throughout his volume allows him to maintain two different voices (Hamburger 1988). While Hamburger’s approach is grounded in close study of the science itself and the people involved in carrying out research, other historians have adopted alternative views. Donna Haraway, better known for her later, more socially oriented studies, wrote her dissertation and first book about three efforts to interpret developmental processes. Her Crystals, Fabrics, and Fields looked at the use of three different metaphors by, respectively, Harrison, Needham, and Paul Weiss. Inspired by Thomas Kuhn’s ideas of paradigm shifts in science, Haraway shows how each of the chosen metaphors both reflected and helped shape the research approach of the proponent (Haraway 1976). Embryologist Scott Gilbert received his Ph.D. from the Johns Hopkins University in biology, and he also received a master’s degree in History of Science while working with Haraway. His own historical and social studies of metaphor usage in biology complement his scientific use of metaphor in his own work. Gilbert’s Developmental Biology textbook, first published in 1985 and now in its tenth edition, has introduced generations to the field, including its theoretical and metaphoric side (Gilbert 1985, 1991). A number of insightful social scientists have pointed to other ways that social factors impact science. For example, Emily Martin has shown that the apparently straightforward understanding of fertilization is highly gendered. After examining a wide range of descriptions of the process by which sperm and egg combine, she noted that most describe a very traditional “boy meets girl” story. Both scientists and those writing about the science invoke a kind of triumph of the male in penetrating the female myth. She pointed to these stereotypical male-female roles in 1991 and challenged writers to be more aware and more accurate in their descriptions (Martin 1991). More recently, Lynn Morgan pointed to another social side of studying development, namely, censorship. At her own Mt. Holyoke, she discovered that a once wellused collection of human embryos and fetuses had been removed. Institutions became concerned that antiabortion politics had created such a heated social environment that these embryos were no longer seen as neutral scientific objects but as something ethically tainted. In her Icons of Life: A Cultural History of Human Embryos, Morgan lays out a number of examples of the cultural role and meaning of human embryos (Morgan 2009). Evelyn Fox Keller has contributed extensively to our understanding of the history of biology, including especially the history of genetics and development. Her 2002

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text, Making Sense of Life: Explaining Biological Development with Models, Metaphors and Machines looked at another side of developmental studies. And her more recent look at the persistent disputes about whether life results from “nature” or “nurture” in The Mirage of a Space Between Nature and Nurture shows ways in which cultural and social values shape biological thinking about life (Fox Keller 2010). Science and its intersections with society and cultural values play out especially in evolutionary work. Even researchers focused on developmental biology bring in their own convictions about the relative importance of historical evolutionary factors and local embryological factors in shaping the processes and patterns through which the individual organism becomes organized and differentiated.

From Embryology to Developmental Biology Decades before historians started to explore the social aspects and impacts of embryos (and embryological research), embryology witnessed its own social upheaval. In the wake of World War II, the field of embryology was transformed into a new field called developmental biology. This transition was replete with journals and society name changes. For example, the first issue of the US journal Developmental Biology came out in 1959, along with statements of unity from leading scientists like Weiss (Hopwood 2009; Oppenheimer 1966; Pradeu 2016). Despite appeals of unity within the field, historians have pointed to this as an era of “fragmentation,” where the old recourse of studying embryos as whole organisms was pushed to the boundaries as the field experienced increasing diversification in terms of research questions, fields of influence, and methods (Crowe et al. 2015; Hopwood 2009; Horder 2010). Historians who have dealt with this shift and its subsequent history have tended to privilege the biochemical and molecular aspects of the emerging field, focusing on narratives that centralize the molecularization of developmental biology (Burian and Thieffry 2000; Crowe et al. 2015; de Chadarevian 2000; Gilbert 1996; Hopwood 2009; Morange 2000; Oppenheimer 1966). Molecularization in this context designates the “substitution of molecular level analyses of biological problems for traditional cellular, tissue, or organismlevel analyses” (Burian and Thieffry 2000, p. 316). Bridging her career as an embryologist and as a thoughtful historian, Oppenheimer (1966) contributed the introductory chapter to the 25th Symposium of the Society for Developmental Biology (previously the Society for the Study of Development and Growth). This chapter, which was praised by the conveners of the symposium, deals with “The Growth and Development of Developmental Biology” and traces the historical unfolding of this field starting with the first symposium on development and growth held in 1939 (Oppenheimer 1966). At that time, the study of genetics and heredity remained largely separate from the study of development. Embryology included the work of Harrison on tissue development, Spemann on the organizer and induction, Holtfreter on the developmental potential of isolated pieces, and so on. All this work ignored genetics, largely because researchers did not see

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how to connect the study of heredity and development. Oppenheimer concluded that recently the two had converged and transformed biology. Following along the lines that Oppenheimer laid, Hopwood has further highlighted the importance of molecularization for the field of developmental biology. “Developmental biology was a joint initiative of self-consciously ‘modern’ embryologists and geneticists, biochemists, cell biologists, and molecular biologists who saw a field ripe for their skills. . .the new field’s key generalization was development as differential gene expression” (Hopwood 2009, p. 309). In addition to the overviews provided by Oppenheimer and Hopwood, historians interested in the molecular aspects of developmental biology have focused on the works or research programs of individuals and, particularly, on molecular biologists who crossed the boundary into developmental biology. Historians of genetics Michel Morange and Soraya de Chadarevian have given us glimpses into the shifting research programs during the 1960s of Francois Jacob and Sydney Brenner, respectively, each of whom left a bacterial model in order to apply gene regulatory questions to eukaryotes (mice, in the case of Jacob, and C. elegans, in the case of Brenner) (de Chadarevian 2000; Morange 2000). While Morange’s work explores in detail the T-complex model and the metaphor of “program” that surrounded Jacob’s work, de Chadarevian’s focuses on the tools that molecular biologists (e.g., Sydney Brenner) brought with them as they moved to developmental questions. The topic of genetics is covered elsewhere in this volume (see ▶ Chap. 6, “Gregor Mendel and the History of Heredity”), and so we do not address the intersection of genetics and development in further detail here.

Nonmolecular Narratives in the History of Developmental Biology While genetics has had an undeniable impact on the field, a great deal of research within developmental biology has not utilized molecular techniques (Sunderland 2011). Following from a computational study of the General Embryological Information Service, Crowe et al. (2015) have recently pointed out that genetics cannot account for the diversification of topics that came along with the advent of developmental biology. If it is the case that genetics cannot account for the diversification of topics within developmental biology post-1945, what can? That is, what does a focus on the molecularization of developmental biology leave out? We can look to a dizzying array of research problems, such as morphogenesis or regeneration, for example, where molecular methods were not co-opted for the study of developmental processes along the same timeline that can be applied to the phenomena of differentiation. This is a research area poorly populated by historians of science but immensely rich in subject matter and different stories. Here we focus on just one topic, morphogenesis, and a few scientists whose work has proven hugely influential. Those few who have focused their historical lenses outside of the molecular (or chemical/biochemical) trend in developmental biology have tended to focus on individuals and localized research programs, preferring to trace the conceptual and

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practical work embedded in the science rather than broader networks of individuals or the social settings of the research. Looking back in 1996, developmental biologist Viktor Hamburger provided an overview of the work of his close friend and colleague, Johannes Holtfreter, whose work never included molecular or even biochemical methods (Hamburger 1996). Holtfreter started out in Spemann’s laboratory, receiving his Ph.D. in 1925. His work over the next 15 years produced impressive insights into the determination of different parts of the embryo during different stages of development, answered the question of whether induction was mediated by a chemical or physical force, and even investigated the dynamic morphogenetic movements and properties (e.g., cell motility and cell adhesion) that shape the embryo. Holtfreter’s research was guided by a search for the mechanisms of development, to the extent that when, at a conference in Utrecht in 1931, he heard Spemann speak about the “metaphysical concept” of the organizer and its “vitalistic agency,” he set up a series of experiments to test the organizer’s capacity for neural induction himself (Holtfreter 1991, p. 117). His results dealt a blow to Spemann’s vitalistic concept of the organizer, showing that the organizer was “merely the source of certain chemical substance(s) that initiated neural differentiation in the responding ectoderm” rather than some metaphysical entity (Holtfreter 1991, p. 117). With regard to Holtfreter’s research on morphogenetic movements, Hamburger tells us that it was his “seminal contribution to envision these processes as the manifestations of inherent properties of cells and cells assemblies, and, furthermore, that these dynamic properties are inseparably connected with the histological and organological fate of the different regions” (Hamburger 1996, pp. 215–216). Holtfreter’s approach to morphogenesis was a radical departure from the tradition of the time because it extended analytical embryology to the cellular level and one that greatly impacted future morphogenetic research (Hamburger 1996; Dietrich 2007; also see Gilbert 1991 and elsewhere for more extensive discussion of Holtfreter’s work). In his autobiography, Embryologist, developmental biologist John P. Trinkaus traces his own research program in morphogenesis from the 1930s through the 1990s. Embryologist is a very personal history of Trinkaus’s research, combining anecdotes and stories about his life with detailed conceptual and practical aspects of his research (as well as the research of those colleagues that he admired). Trinkaus describes being so inspired by Holtfreter’s work on cell motility and adhesion that it directed his career, noting, “Here was a subject of outstanding developmental importance whose mechanism was poorly understood but which was definitely attackable, as Holtfreter had so brilliantly demonstrated” (Trinkaus 2003, p. 89). Trinkaus’ career was spent understanding the mechanisms of cell motility and adhesion during epiboly in Fundulus – work for which he received the first ever Edwin Grant Conklin Medal from the Society for Developmental Biology in 1995. Also notable about Trinkaus for our purposes was his aversion to genetics, “The morphological beauty of embryology won out over the logical beauty of genetics” (Trinkaus 2003, p. 56), and his reflection on a general trend in developmental biology that, “. . .the bulk of research in developmental biology in recent

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decades has concentrated on cytodifferentiation, leaving somewhat to the side the equally important problems of how the cells got there in the first place, how they change form, and how they arrange themselves into different tissues and organs” (Trinkaus 1984, p. 3). Trinkaus’s friend and fellow graduate student at Johns Hopkins University, John Saunders, similarly embarked on a nonmolecular career track with a focus on tissue and cellular interactions. Trinkaus discusses briefly in his autobiography Saunders’ discovery during his graduate studies of the apical ectodermal ridge (AER), the thickening of ectoderm on the developing vertebrate limb bud that directs limb outgrowth (Trinkaus 2003). Saunders continued work on the AER but also looked at other problems of vertebrate limb development, including cell death and the formation of digits. It was through his investigations of cell death that he discovered, along with his colleague, the zone of polarizing activity (ZPA), a region on the outgrowing limb that is central to limb patterning (Tickle 2002). Developmental biologist, Cheryll Tickle, whose own research on avian limb patterning led to the discovery that retinoic acid could mimic the effects of the ZPA, has shown that Saunders’ work on the ZPA was the result of grafting experiments and that, while his work to uncover the role of the ZPA was molecule-free, his discovery spurred a long debate over competing models of limb patterning and a search for chemical and molecular mechanisms (Tickle 2002). John Tyler Bonner, often referred to as one of the “grand old men” of the Dictyostelium community, similarly pursued questions of morphogenesis that focused on organisms rather than genes or molecules. Historian of science Mary Sunderland (2011) has shown both that Bonner’s investigative pathway fell outside of the traditional molecular narrative, due both to his emphasis on applying an array of experimental strategies to a range of organisms and the intractability of Dictyostelium as a genetic model at the time and that he conceptually laid the groundwork for evolutionary developmental biology. In his 1952 text, Morphogenesis: An Essay on Development, which was written in the Woods Hole, MA, laboratory of famed cell biologist and embryologist Edwin Grant Conklin, Bonner laid out an argument for uncovering shared developmental principles by studying organismal diversity in a theoretical context (Bonner 1952; Sunderland 2011). Unfortunately, not much heed was taken of Bonner’s arguments for studying development across the animal kingdom at the time of publication.

Evolutionary Developmental Biology In the field of evolutionary developmental biology (or “evo-devo”), Bonner’s vision of scouring organismal diversity for mechanisms of developmental change has been achieved. The field of evo-devo is broad, encompassing many organisms, techniques, goals, and theoretical and epistemic frameworks. It is a field bound together by the tenet that understanding development gives the best insights into understanding organismal evolution and one in which no unifying theoretical framework has emerged.

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The twentieth-century concrescence of evolutionary and developmental biology into evo-devo has been traced back to several key events: Stephen Jay Gould’s publication of Ontogeny and Phylogeny (Gould 1977), the 1981 Dahlem Conference on “Evolution and Development” organized by John Tyler Bonner (1982), and the discovery of the universality of Hox genes (See Carroll 2005). Each of these moments spurred the broader scientific community to reclaim an evolutionary perspective for developmental biology – a field that had largely been left out of the Modern Synthesis. The history of evo-devo has been told along two separate, yet complementary, narrative arcs. In the first arc, which has been called “the standard narrative” (Laubichler and Maienschein 2013), a direct conceptual lineage is traced from Darwin, through the Modern Synthesis, to an “expanded evolutionary synthesis” in which development is incorporated into the evolutionary theoretical framework that the Modern Synthesis provided (Carroll 2005, 2008; Raff 2000). The evolutionary framework of the Modern Synthesis is one tied to genes, particularly population genetics, and the role of natural selection in guiding evolutionary processes. Studies of development are seen in this evo-devo narrative as providing insights into the limitations of natural selection, i.e., the developmental constraints that affect the direction of evolution. As evolutionary developmental biologist Rudy Raff put it, in modern evolutionary theory, “Micro-evolutionary processes are considered sufficient to explain macro-evolutionary history. However, developmental processes are emergent, and not predictable from the properties of genes or cells; therefore, starting with a particular ontogeny, some phenotypes might be readily achieved and others impossible. Developmental mechanisms are crucial, both to large-scale evolutionary changes, and also to small scale evolutionary processes” (Raff 2000, p. 78). The standard narrative is neat and tidy, but as Laubichler and Maienschein (2013) point out, there is “an implicit progression of ideas, with inclusion of new empirical facts and methodological approaches within the general framework of Darwinism leading to an increasingly more complete understanding of the evolutionary process” (2013, p. 375). The problem with the standard narrative is that it leaves out several traditions that have played a major role in conceptually shaping evo-devo and, in doing so, gives the impression that it is a unified, theoretically coherent field. Unfortunately, this is not the case at the present time. A precursory look at current publications in the field or attendance at an evo-devo conference shows just how theoretically and epistemically diverse the field is. In fact, at the inaugural meeting of the Pan-American Society for Evolutionary Developmental Biology in August 2015, one of the main talking points for the “Future of EvoDevo” session was how to form a more (theoretically) unified field so that funding bodies would continue to recognize the research (MacCord was in attendance). Laubichler and Maienschein (2007, 2013) offer an alternative narrative arc that complements the standard narrative by filling in the history with an understanding of traditions that focused on the origins of variation, rather than the inheritance of them. In an edited volume that arose from the 2001 Dibner Seminar at the MBL “From Embryology to Evo-Devo (Evolutionary Developmental Biology)” and a follow-up

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workshop the next year, Laubichler and Maienschein (2007) brought together historians, philosophers, and scientists to explore the historical trajectory of research that joined together evolution and development. While contributors like Scott Gilbert, Brian Hall, and Alan Love had previously made contributions to this problem space (see, for instance, Gilbert 2003; Hall 2000; Love 2003), it was a singular accomplishment to unite a group that could both broadly survey evo-devo history and use these historical insights to analyze the future of the field. Topics in this volume range from understanding the modern significance of research programs of historical figures like William Bateson (Newman) and Richard Goldschmidt (Richmond) to tracking the contributions of different fields outside of evolutionary biology and developmental biology (Love). As broad a perspective as Laubichler and Maienschein offer, there remains a great deal of territory for historians to cover. In light of the current theoretical and epistemic disunity of the field, historical and philosophical investigations can only serve to benefit any future synthesis of evo-devo. Thus, this is an area where young historians of science can both carve out their own niche and have a tremendous impact on shaping the future of a scientific field.

Conclusion Developmental biology (formerly and sometimes still known as embryology) has a long and rich history, and scientific research in this area is unfolding at a growing rate. Despite the centuries of theoretical, empirical, and, more recently, experimental investigations of the phenomena and processes that shape life, this is an area of inquiry that is only sparsely populated with historical analyses. The future of historical work (which sounds a bit like an oxymoron) in this area is wide open for fresh, young historians of biology to dig in and offer new perspectives. In closing, we point to some forward-looking techniques and areas where contextualized understandings of development offer an excellent place for an intersection between academic and public knowledge. Instead of focusing on particular topics that could benefit from more historical attention, we note that there are so many theories, practices, and discussions ripe for study. Throughout, we have noted a number of them, and there are many, many others. There is a huge potential in the historical study of developmental biology for new digital and computational approaches and techniques that bring new questions and new discoveries of patterns worth exploration. These techniques also enable historians to reach a broad, public audience like never before. Digital publications, such as the online, open access Embryo Project Encyclopedia (EPE), edited at Arizona State University, have enabled publically oriented scholarship that is approaching one million page views a year. The project welcomes submissions of shorter and longer articles and can be accessed through http://www.embryo.asu.edu. The EPE, along with the MBL History Project (a digital project that preserves and communicates the history of science at the Marine Biological Laboratory in Woods Hole, MA, at http://history.archives.mbl.edu), is also starting to integrate

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computational approaches, which will allow both scholars and the public to query data harvested from digitized content of the MBL archives. As these projects expand, computational methods like citation analyses, topic modeling, and semantic network analyses will become available, allowing all users to explore the connections between researchers, institutions, and concepts in new ways. Software for such analyses is currently being developed in places like Manfred Laubichler’s Digital Innovation Group at Arizona State University through http://diging.asu.edu. Such new approaches, new studies of existing topics, and also studies of emerging areas within the rapidly growing field of developmental biology make this an especially dynamic area for historians. In fact, we hope that we have missed other intriguing areas for research, because that just means the field is even richer.

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Gregor Mendel and the History of Heredity Staffan Müller-Wille

Ihr Lettern, meines Forschens Sprossen ... Gregor Mendel, 1830s

Contents Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mendel Ahead of His Time . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mendel Stuck in Time . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . The Many Times of Mendel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Outlook . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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Abstract

Gregor Mendel’s paper “Experiments on Plant Hybrids” (1866) has become a paradigmatic case in the historiography of the life sciences because production and reception of a “discovery” sharply fell apart, thus raising fundamental questions about the relationship between scientific achievement and “its” time. In this chapter, I am providing an overview of answers that have been given to these questions by various historians. In a first section, I cover commentators who have claimed that Mendel was “ahead” of his time, and that contemporaries failed to recognize his achievement. I then move on to scholars and scientists who argued against this position, claiming that Mendel was not anticipating twentiethcentury genetics, but was in fact representative of an older research tradition. In a First line from a poem that Gregor Mendel sketched in the late 1830s during his time at the gymnasium of Opava (Troppau). It can be translated as “Oh letters, rungs of my research ...”; the poem celebrates Johannes Gutenberg’s invention of movable print. Quoted from Iltis (1924, 14) S. Müller-Wille (*) University of Cambridge, Cambridge, UK e-mail: [email protected]; [email protected] # Springer Nature Switzerland AG 2021 M. R. Dietrich et al. (eds.), Handbook of the Historiography of Biology, Historiographies of Science, https://doi.org/10.1007/978-3-319-90119-0_8

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last step, I turn to the more recent cultural history of heredity according to which Mendel was embedded in a local culture that combined a variety of advanced and traditional strands of nineteenth-century life-sciences. Overall, I am arguing that one should not overestimate the coherence and dominance of presumed “paradigms,” “epistemes” or “styles” in biology.

Introduction There is no doubt that Gregor Mendel was a child of the nineteenth century. In 1822, he was born into a family of ethnically German small holders in Hynčice (German: Heinzingen) in northeastern Moravia (today Czech Republic), joined the Augustinian Abbey St. Thomas in Brno (Brünn) in 1843, received a thorough education in natural sciences at Vienna University, and finally rose to the position of abbot, which he held until his death in 1884. Like countless contemporaries, he was a keen naturalist, joined local associations for natural history and the promotion of agriculture and industry, and published occasional notes and essays on agricultural pests, plant hybridization, and meteorology. Nothing in Mendel’s biography – including perusal of the Emperor Ferdinand Northern Railway to visit the libraries of Vienna and participation in a group tour to see the Great Exhibition in London in 1862 – points beyond the intellectual horizon of a polyglot citizen of the Austro-Hungarian Empire and active participant in the vibrant culture of industrializing Central Europe.1 And yet, the historiography of the life sciences has usually portrayed Mendel as being out of step with his time, either by making him out as the “founding father” of genetics, a quintessentially twentieth-century discipline, or by portraying him as a late representative of “hybridism,” a research tradition in the plant sciences deeply rooted in eighteenth-century species fixism. These divergent interpretations are due to what is often called “the long neglect” of Mendel’s main literary achievement – a 47-page long report on hybridization experiments with pea and other plant varieties that was published in 1866. The aim of this publication, as Mendel himself put it, was to establish “a generally valid law for the formation and development of hybrids.”2 Yet, with some exceptions, this publication went unnoticed until 1900. In that year, three botanists – Hugo de Vries (1848–1935), Carl Correns (1864–1933), and Erich von Tschermak (1871–1962) – claimed to have independently “rediscovered” the laws that Mendel had presumably laid down already. Within a few years, his paper, as well as the eponymous laws, became widely known among biologists as providing the foundations for the new discipline of genetics, a discipline that was perceived to hold revolutionary potential for 1

For Mendel’s biography, see Iltis (1924, English translation 1966), Orel (1996), and Klein and Klein (2013). I will rely on Orel for biographical details throughout this chapter. 2 Mendel (1866, p. 3); Mendel (2016a, p. 3, s. 6). There exist several English translations of Mendel’s essay (Mendel 1901, 1902, 1966, 2016b). I am quoting from the latest critical edition and translation, citing page and sentence number. The phrase “long neglect” can be traced back to Glass (1953, p. 148).

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controlling, manipulating, and understanding life. Since then, the 34 years of silence around Mendel’s paper have either been made out as a tragic case of oversight that hampered scientific progress or else as a phantasm resulting from anachronistic projections of later developments in science onto earlier episodes in its history.3 Given the curious fate of Mendel’s paper, it comes as no surprise that it is one of the most heavily scrutinized texts in the historiography of biology – almost certainly second only to Darwin’s On the Origin of Species (1859) – and that the resultant body of scholarly work is ruptured by passionate debates about what it was, exactly, that Mendel discovered and what place this discovery occupies in the longue durée history of the life sciences in general and the history of heredity more specifically. No less than three substantive review articles are available, which chart the evolving landscape of these historiographical debates throughout the twentieth century. Their authors – Robert C. Olby, Jan Sapp, and Vítezslav Orel (1926–2015) – fall themselves on different sides of the debates and yet agree in one fundamental point: They all portray the debates as arising from a divorce between scientist historians looking back at the origins of their discipline and professional historians of science deconstructing the resultant “origin myths.”4 This chapter proposes that this picture of a divorce between scientist and professional historians involves elements of mythmaking itself. Whether or not, and if yes, in what sense Mendel can be seen as having brought about a breakthrough in understanding heredity has also been debated among scientists, for the simple reason that the science of heredity, its aims, methods, and core assumptions, was understood and evaluated very differently by different schools and generations of scientists. Mendel’s paper became a paradigmatic case in the history of science not simply because its historical meaning was contested, but more specifically because production and reception of a “discovery” sharply fell apart in this case, thus raising fundamental questions about the relationship between scientific achievement and “its” time. In the following, I am going to provide an overview of answers to these questions. I will first cover commentators who have claimed that Mendel was “ahead” of his time and that contemporaries failed to recognize his achievement. I will then move on to scholars and scientists who argued against this position, claiming that Mendel was not anticipating twentieth-century genetics, but was in fact representative of an older research tradition. In a last step, I will turn to those who have held that both of these positions rest on a false dichotomy and that Mendel

On references to Mendel’s paper before 1900, see Olby and Gautrey (1968). On its “rediscovery” in 1900, see Jahn (1958), Olby (1985, ch. 6), Rheinberger (1995), Stamhuis et al. (1999), Harwood (2000), and Simunek et al. (2011). Olby (1985, 219–234) provides English translations of some sources mentioning Mendel before 1900. On the history of Mendelian genetics, see Dietrich ▶ Chap. 8, “The Historiography of Genetics,” this volume. 4 Sapp (1990, p. 146), Orel and Hartl (1994, p. 445), and Olby (1997, section “Scientific Disciplines”). All three review articles are accessible online at MendelWeb (www.mendelweb.org), an internet resource created by Roger B. Blumberg in 1995, but not updated since 1997. It offers a wealth of other useful material, including the German original and Bateson’s 1902 translation of Mendel’s paper for download. 3

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was embedded in a local culture that combined a variety of advanced and traditional strands of nineteenth-century life sciences. Overall, I am going to argue that one should not overestimate the coherence and dominance of presumed “paradigms,” “epistemes,” or “styles” in biology. Life, as well as the scientific disciplines that study it, is just too indeterminate and inchoate to develop in such an orderly fashion.

Mendel Ahead of His Time The historiography of Mendel begins with his three rediscoverers. All of them acknowledged his priority and added that his paper had “fallen into oblivion” (de Vries), had “hardly attracted any interest” (Correns), or, as Tschermak put it, only had come to light after its “simultaneous ‘discovery’ by Correns, de Vries and myself.”5 Yet, already in this epochal moment in the history of heredity, disagreements about how to read Mendel became apparent. De Vries was the first to announce the “discovery” of Mendel in a “preliminary note” that he had submitted for publication on March 14, 1900, to the German journal Berichte der deutschen Botanischen Gesellschaft. Without much ado, he brought Mendel into the fold of his own mutation theory, claiming that Mendel had only shown for a “special case (peas)” what he, de Vries, had been able to establish with “general validity,” namely, (1) that hybrids show one of two “antagonistic characters” only (what is sometimes called the law of dominance) and (2) that these characters “separate in the formation of pollen and egg cells” (the law of segregation).6 Correns followed suit, with a paper submitted to the same journal on April 24. With thinly veiled sarcasm, he stated that he had not “considered it necessary to secure priority for a ‘post-discovery’ [‘Nach-Entdeckung’].” Against de Vries, Correns argued on the basis of actual quotations from Mendel’s paper that the latter had not formulated two laws, but only one, which stated “that the hybrids produce germ- and pollen cells that correspond in equal quantities to all the constant forms that emerge from the combination of the traits that were united by fertilisation.” What de Vries’ called the “law of segregation” (Spaltungsgesetz) was actually included by this law, whereas de Vries’s law of dominance was simply fictitious and had never been endorsed by Mendel.7 Tschermak went even further in dissociating Mendel from any particular “laws” of inheritance. His contribution to the Berichte, submitted on June 2, did not embark on an interpretation of Mendel’s original paper, but simply took his empirical results for granted.8 A year later, however, Tschermak published the first scholarly edition of 5 Vries (1900, 85), Correns (1900, 159), and Tschermak (1900, 239). English translations of de Vries’s and Correns’ papers can be found in Stern and Sherwood (1966, 107–138), but I am here providing my own. 6 Vries (1900, 84–85). 7 Correns (1900, 166–68). Correns quotes Mendel’s own formulation of the alleged law; cf. Mendel (2016a, p. 29, s. 4). 8 Tschermak (1900, 235).

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Mendel’s paper in a renowned book series of classical contributions to natural science. According to him, all that Mendel had shown was that traits of organisms differ regularly in their “valency” (Werthigkeit) and that they do so in three respects: heritability, number of individual carriers, and degree of expression of traits.9 His account of how Mendel had been forgotten now showed a slight change in wording. Mendel’s paper had not been “discovered” but rather “‘rediscovered’” (“wiederentdeckt”; quotation marks in the original).10 It is likely that he took over this curious locution from the British botanist William Bateson, who mentioned “de Vries’s papers announcing the ‘rediscovery’ and confirmation of Mendel’s law” in the published version of a paper presented to the Royal Horticultural Society on May 8, 1900. Bateson also used the phrase “simultaneous rediscovery” – now without quotation marks – about half a year later in an introductory note that preceded the first English translation of Mendel’s paper.11 It is worth contemplating these details for two reasons. First, the various uses of the term “discovery,” and especially its derivatives, and the frequent use of quotation marks indicate that it is a very slippery term, even, and perhaps especially, to scientists. Not only does it seem that scientific discoveries, just like films, can experience “remakes”; the meaning of the term also shifts back and forth between uncovering a timeless fact of nature and lifting a past effort to describe such a fact from the vast archives of modern science. Sociologist of science Augustine Brannigan has argued for an “attributional model of scientific discovery,” according to which priority disputes, but more importantly scientific debates about how best to interpret phenomena, provide the social context in which “certain happenings” are declared “discoveries.” The “simultaneous rediscovery” of Mendel by de Vries, Correns, and Tschermak served him as a paradigmatic case.12 My point here is a different one, though, namely, that such processes of attribution generate among scientists what I suggest to call a “spontaneous” history of science. Even for scientists, facts of nature are inextricably bound up with texts that try to communicate them, and these texts never simply speak for themselves, but always need historical uncovering and interpretation to reveal their proper meaning.13 Given the central role genetics played in twentieth-century life sciences, it therefore does not come as a surprise that Mendelian scholarship, to this day, has heavily depended on contributions from scientists. Virtually all the archival sources we possess from Mendel have been unearthed and edited, often with an accompanying scholarly apparatus, not by historians of science but by scientists: Tschermak

9 Tschermak (1901, 54). In the second edition (1911), Tschermak changed his mind and presented Mendel much more in line with what we today would consider as conventional Mendelism. 10 Ibid., 55. 11 Bateson (1900–1901, 60) and Mendel (1901, 2). 12 Brannigan (1979, 450). 13 The French Marxist philosopher Louis Althusser suggested that the “spontaneous philosophy of scientists” could provide an antidote to the tendency of philosophy “to speak about nothing but itself” (Macherey 2009, 19).

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and Bateson, as already mentioned, edited and translated his original paper, and Correns soon followed with an edition of the letters Mendel wrote to prominent Swiss botanist Carl Nägeli (1817–1891).14 The centennial of Mendel’s paper saw another two critical editions, with accompanying materials and commentary by geneticists R. A. Fisher (1890–1962) and Curt Stern (1902–1981), respectively, and in the same year the Czech poultry geneticist Jaroslav Kříženecký (1896–1964) produced a transcription of Mendel’s original fair copy.15 Even the few handwritten annotations that have come down to us from Mendel’s readings and experiments were brought to light by scientists like the embryologist Sir Gavin de Beer (1899–1972), Dutch botanist and conservationist Jacobus Heimans (1889–1978), or German viticulturalist Franz Weiling (1909–1999) – not to speak of the countless, and often obscure, primary sources that support the three biographies of Mendel, all of them composed by scientists.16 These efforts did not simply aim to celebrate Mendel as the founding father of genetics. What was also at stake, as we have seen with the rediscoverers, were fundamental questions relating to the nature of Mendel’s discovery. In a classic essay, which appeared in the first issue of the journal Annals of Science (1936), the statistician R. A. Fisher formulated these questions succinctly: “What did Mendel discover? How did he discover it? And what did he think he had discovered?”17 And Fisher was all but naïve about the prospect of deciding these questions on an entirely objective basis. His essay was in fact the first historiographical contribution to Mendelian scholarship and came to the oft-quoted conclusion that: Each generation, perhaps, found in Mendel’s paper only what it expected to find; in the first period a repetition of the hybridization results commonly reported, in the second a discovery in inheritance supposedly difficult to reconcile with continuous evolution. Each generation, therefore, ignored what did not confirm its own expectations. Only a succession of publications, the progressive building up of a corpus of scientific work, and the continuous iteration of all new opinions seem sufficient to bring a new discovery into general recognition.18

The target of this conclusion, and of the article as a whole, was a complex one. It included those (the “first generation”) who allegedly had only been interested in the empirical verification of Mendel’s experimental results, but also those (the “second generation”) who had offered a particular theoretical reading of these results. Fisher’s argument was therefore twofold. On the one hand, he rejected the idea – prominently defended by Bateson in the polemical context of what came to be known as the “Mendelism-Biometry debate”19 – that Mendel had been opposed to Darwin, that Mendel’s main achievement had been to disprove continuous variation, and that the

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Correns (1905); Nägeli’s responses to Mendel are lost. Bennett (1965); Stern and Sherwood (1966); Kříženecký (1965, 57–92). 16 Beer (1964); Heimans (1968, 1970); Weiling (1992). On Mendel biographies, see note 2. 17 Fisher (1936, 137). 18 Fisher (1936, 137). 19 Bateson (1902). This was the first book-length critical study of Mendel’s paper, and a second revised edition appeared in 1909. 15

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dominance of Darwinian ideas had been responsible for his neglect. On the other hand – and now following up on earlier clues from Bateson – Fisher proposed that Mendel did not at all proceed empirically, as a superficial reading of his paper might suggest. Instead, he had constructed his experiments, as well as the account he gave of them (including some manipulation of data to fit expectations), in order to produce “a carefully planned demonstration of his conclusions.”20 Both lines of argument chimed well with Fisher’s own contributions to the so-called Modern Synthesis of Mendelian genetics with the Darwinian theory of natural selection on the one hand and to the foundations of mathematical population genetics on the other.21 Fisher’s essay extended the “long neglect of Mendel” by another full 30 years.22 If nineteenth-century researchers had overlooked his results, and what these revealed about the mechanism of inheritance, early twentieth-century Mendelians had misunderstood his unique approach, which combined experiment with mathematical treatment and had failed to see that it was compatible with Darwinian natural selection. Mendel thus turned into the paradigm case of a scientist who fell out of the discursive boundaries drawn by scientific communities. In 1953, American geneticist H. Bentley Glass produced a map-like diagram for his contribution to a Festschrift honoring Arthur O. Lovejoy, the “father” of the history of ideas. Bearing the title “The Chain of Ideas from Goethe and Oken, Who Represent the Naturphilosophie, to Modern Genetics,” it did not even include Mendel who, as Glass supposed, lay “out to the side of all these streams of thought.”23 For the history of heredity more generally, this portrayal of Mendel as a lone and mis- or not-at-all understood researcher had a curious consequence. Books dedicated to this subject, up until the mid-1960s, were of two kinds. They were either histories of genetics, taking their starting point in 1900, the year of rediscovery;24 or they were histories of “precursors” to Mendelism, choosing the same year as their vanishing point.25 Alongside this, there existed a literature that dealt separately with a related subject: the history of generation.26 All of these studies retain their value, both for their sophistication and the wealth of material they contain. The three strands would only be brought together, however, after the Long Neglect story had been seriously challenged in the late 1970s.

Fisher (1936, 124). Whether Mendel, consciously or not, falsified “his data to produce results that were, too good to be true’ has been the subject of a fierce debate among statisticians, geneticists, and historians of science that continues to this day; see Franklin et al. (2008) for a collection of important contributions to this debate. The allegation of data manipulation was not new when Fisher wrote his article. Bateson’s main opponent, the biometrician Raphael Weldon (1860–1906), had actually raised it in 1902 already; see Radick (2015). 21 On the Modern Synthesis, see Borrello ▶ Chap. 3, “The Historiography of Modern Evolutionary Biology,” this volume. 22 Beer (1964). 23 Glass (1953, 158). 24 Carlson (1966); Sturtevant (1966). 25 Roberts (1929); Stubbe (1963); Robinson (1979). 26 Lesky (1951); Roger (1963/1998); Gasking (1967). 20

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Mendel Stuck in Time While not themselves contributing to historical scholarship, sociologists and philosophers of science in the first six decades of the twentieth century were quick to pick up the story of Mendel’s neglect and subsequent parallel rediscovery. Cultural anthropologist Alfred L. Kroeber (1876–1960) used it in his article “The Superorganic” (1917) – in which he argued for the independence of cultural evolution from the biological and psychological evolution of individuals – to point to “the inexorable fate in store for the discoverer who anticipates his time,” and such references became routine with later sociologists and anthropologists intrigued by the phenomenon of multiple and roughly simultaneous discoveries and inventions.27 In the French tradition of historical epistemology, references to Mendel and his rediscovery were made just as routinely. Thus, Michel Foucault (1926–1984) proclaimed in his inaugural lecture at the Collége de France, held on 2 December 1970, that Mendel had spoken “of objects, employed methods and placed himself within a theoretical perspective totally alien to the biology of his time” and that it would require a “whole change in scale, the deployment of a totally new range of objects in biology ... before Mendel could enter into the true [dans le vrai] and his propositions appear, for the most part, exact.”28 In the same year, molecular biologist François Jacob published his magisterial Logic of Life: A History of Heredity, in which Mendel, once more, but now in greater detail, was credited as having introduced an entirely new approach to biology – as well as constituting a new object, the gene – by focusing on discrete character pairs in his experiments and analyzing their results with the help of combinatorial analysis and statistics.29 All of these accounts do not divorce scientific achievements from contemporary contexts; quite on the contrary, the point of telling Mendel’s story was, and often still is today, to demonstrate how powerful prevailing discursive contexts are in advancing or else obstructing innovation. The only way to counter this story, without falling back into sheer positivism, was to say that the story itself was simply wrong; that Mendel had, in fact, not discovered what he was supposed to have discovered 16 years after his death; that he had not been a “lone genius” but a well-connected naturalist; and that he had not been neglected at his time but received due attention from his peers. In other words, what was called for was a new reading of relevant historical sources for hints that would root Mendel firmly within traditions of nineteenth-century biology. Helpfully, Mendel himself was quite explicit in placing himself within such traditions. Not only did he list predecessors in the first paragraph of his paper, but

27 Kroeber (1917, 198); Ogburn and Thomas (1922); White (1949). Robert K. Merton (1910–2003), in his classic article on multiple discoveries (1961), considered the “case of Mendel” as “too well known” to even expand on it any further; see Merton (1996, 308). 28 Foucault (1970/1971, 16). The quoted passages rely heavily on earlier work by Foucault’s mentor Georges Canguilhem (1904–1995); see, e.g., Delaporte (1994, 37, 51). 29 Jacob (1970/1996, 202–9).

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he also concluded it with a critical review of their experiments (again, an exercise in “spontaneous history”).30 In a book that appeared in the centennial year 1966, alongside the two new English editions of Mendel’s paper mentioned above, Olby followed up these references in a two-pronged attack on the received view of Mendel. Not only did he show that Mendel had been working in a long-established and well-defined older research tradition which dealt with hybridization, rather than heredity as such, and dated back to a hybridization experiment carried out in 1759 by the Swedish naturalist Carl Linnaeus (1707–1778); it also turned out that heredity as a research subject of its own was not so much defined by Mendel, nor by Charles Darwin who remained a “lifelong generation theorist,” but by the likes of Francis Galton (1822–1911) and August Weismann (1834–1914).31 Olby was the first professional historian of science to write a monographic account of Mendel and the history of inheritance. As he recalls in the second edition of his book, Origins of Mendelism had grown out of a PhD thesis project he had undertaken with population geneticist and historian of science Alistair Crombie, who later became known for transferring the concept of “style” from the study of art to the study of science and its history. Aware of his own “‘Whiggish’, ‘presentist’ ... tendencies” in the first edition – that is, tendencies to select and evaluate protagonists and events in the history of science in view of their contributions to the current state of art32 – Olby aimed to shift emphasis in his amended second edition “away from a positivist and empiricist interpretation of the history of genetics and toward a constructivist view in which knowledge is seen as under-determined by the facts, their meaning being dependent to an important extent upon the theoretical presuppositions of the observer.”33 This shift in emphasis was already evident in a highly influential journal article Olby had published in 1979. It bore the provocative title “Mendel no Mendelian?” and turned far more explicitly against “inflated whiggish interpretations” of Mendel that were prevalent among scientists and in the popular press. Relying on Paul Forman’s classical study of mythmaking in the historiography of X-ray crystallography, Olby surmised that the “orthodox view” of Mendel as the founding father of genetics had originated and continued to prevail because “geneticists and plant breeders have introduced mythical elements into their reconstructions of the history of genetics which historians have failed to identify and reject.”34 Less polemical in substance than this may sound, and flanked by the sociological analysis Brannigan provided in the same year, “Mendel no Mendelian?” provided ample historical evidence for a new assessment of Mendel’s achievements that 30

Mendel (2016a, 38–47). This last section was also included in what became the most popular English edition of Mendel’s paper. It originally appeared as an appendix in Castle (1916, 281–321) and was then reprinted as an inconspicuous brochure by Harvard University Press until 1965. It is all the more mysterious why Zirkle (1951, 99), otherwise a very attentive scientist historian, claimed that “Mendel himself described none of the earlier research.” 31 Olby (1966, 10). On Darwin as a “lifelong generation theorist,” see Hodge (1985). 32 See Jardine (2003) for critical reflections on the concept of “Whig history.” 33 Olby (1985, xi–xiv). 34 Olby (1979, 53–54).

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inspired a whole new generation of Mendelian scholarship. A consensus emerged over the long run that Mendel had certainly not endorsed the particulate view of genetic factors that was promoted by mid twentieth-century population and molecular genetics and also had not formulated his “laws” in the same way as modern biology textbooks do to this day.35 But controversy persisted regarding the question how far back in history Mendel should eventually be pushed. Some, including Olby, argued that Mendel was fully up to date with current problems in contemporary cell and evolutionary biology, but failed to convince his peers that he had come up with a plausible solution.36 Others, among them evolutionary biologist Ernst Mayr (1904–2005) in a review of Olby’s book, identified an element of outdated essentialism in Mendel and sometimes even went as far as claiming that he had written his essay specifically to disprove Darwin’s theory of natural selection.37 Olby captured the two sides in this controversy nicely, by speaking of the “Darwinian” and the “Linnaean” Mendel.38 For the history of heredity more generally, Olby’s intervention signaled the end of accounts that assumed that heredity was an obvious problem only waiting for its eventual solution by Mendel and the Mendelians. Jacob had already made the forceful point in 1970 that heredity was a relatively late conceptual product of modern biology and that its history formed a complex landscape of evolving and competing disciplinary formations. Each “stage” in the history of heredity was characterized by a whole series of conceptual, technological, and methodological breakthroughs that made new domains of enquiry accessible to researchers, but also limited them to a “range of possibilities defined not only by current theories and beliefs, but also by the very nature of the objects accessible to investigation, the equipment available for studying them and the way of observing and discussing them.”39 While disagreeing with Jacob’s very general, transdisciplinary periodization, Mayr produced a similar account in his The Growth of Biological Thought: Diversity, Evolution, and Inheritance (1982). Like Jacob, he saw biological disciplines going through “periods of stagnation and periods of greatly accelerated advance.”40 Placing emphasis much more exclusively on conceptual shifts than Jacob, Mayr told the history of heredity through shifts along conceptual dichotomies – from essentialism to population thinking, from preformation to epigenesis, from soft (or Lamarckian) to hard inheritance, from blending to non-blending inheritance – that have profoundly influenced subsequent discussions among historians and philosophers of biology.41

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Kampourakis (2015). Campbell (1982), Olby (1985, ch. 5), and Sandler and Sandler (1985). 37 Mayr (1973); Callender (1988); Bishop (1996). 38 Olby (1997), section IV, “Mendel and the Darwinians.” 39 Jacob (1970/1996, 11). 40 Mayr (1982, 127). 41 Burkhardt (1994); Müller-Wille (2011). 36

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Despite this influence, or rather just because of it, Mayr attracted substantial criticism from professional historians of science for what they perceived as his “Whiggish” tendency to focus selectively on the history of those ideas that would eventually shape twentieth-century theories of evolution.42 A good example is Peter J. Bowler’s book on the history of hereditarianism, entitled The Mendelian Revolution: The Emergence of Hereditarian Concepts in Modern Science and Society (1989). Bowler agreed that the “advent of Mendelism represented ... a conceptual revolution of major proportions.”43 But drawing inspiration from Thomas S. Kuhn’s theory of paradigm change and the Edinburgh School’s “strong program” in sociology of knowledge, he emphasized the social and ideological dimensions of this revolution. Especially eugenics, which had hardly played any role in Jacob’s and Mayr’s histories of heredity, was thus foregrounded as a major force in promoting hereditarian ideas, while Mendel’s original contribution receded into the background of the countless efforts in late nineteenth-century biology to conceptualize heredity from the point of view of a “developmentalist” or “pre-Mendelian” paradigm that did not yet separate clearly between transmission and development. “[N]ew laws or theories are not simply ‘discovered’,” Bowler insisted against scientist historians, but “invented to satisfy the cultural values of the scientists and of the public with whom they interact.” Mendel’s paper had simply failed to conform to these values at the time when it was published, but happened to do so 34 years later.44

The Many Times of Mendel The idea that Mendel’s “discovery” was an invention, or fabrication even, met stubborn resistance above all from one Mendelian scholar: Vítězslav Orel (1926–2015). Orel was a student of Kříženecký’s and played an important role in preparing the 1965 Gregor Mendel Memorial Symposium in Brno, which was organized by the Czech Academy of Sciences with sponsorship from international organizations such as UNESCO, the International Union of Biological Scientists, and the International Atomic Energy Agency. The Symposium attracted an international audience of scientists, including Conrad H. Waddington (1905–1975), Nikolay Timofeev-Ressovsky (1900–1981), as well as Jacob, Stern, and Stubbe as speakers.45 Subsequently, Orel became the director of the Mendelianum – a museum dedicated to Mendel’s life and work on the premises of the monastery in Brno – head 42

Bowler (1988). For a defense against these attacks, see Mayr (1990). Bowler (1989, 7). 44 Bowler (1989, 7, 44, and 94–95). Ironically, Bowler relied on an earlier explanation of the “long neglect” that had been proposed by two geneticists, Iris and Laurence Sandler; see Bowler (1989, 108). 45 Sosna (1966, vii–xi). The conference proceedings are available from the Wellcome Library’s Digital Collections (URL https://wellcomelibrary.org/item/b18019900). 43

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of a research unit on Mendel at the Agricultural University in Brno, and editor of a journal entirely devoted to Mendelian scholarship, the Folia Mendeliana. He held these positions until his retirement in 1990, but continued to publish on Mendel until his death in 2015, often in co-authorship with anglophone scientists and historians.46 It is tempting to dismiss Orel’s lifelong insistence that Mendel was the “first geneticist” as a sign of the hero worship that scientists easily fall prey to and that Olby and Bowler criticized so incisively. But the Gregor Mendel Memorial Symposium was not just any commemorative event. It had been organized to achieve the symbolic “rehabilitation” of Mendelism in the Communist Bloc after the official downfall of Lysenkoism (and Nikita Khrushchev) in 1964.47 In order to make this happen, it was essential to secure a substantial connection between Mendel as a historical figure of local significance in Brno, then part of the Communist Bloc, and twentieth-century genetics as an international science. And something else needed to happen. Just like the Soviet theoretical physicist Boris Hessen (1893–1936) had exposed the social and ideological “roots” of Isaac Newton’s Principia in 1931 in order to fend off accusations in his home country that the pursuit of “pure” science was economically and ideologically useless, it had to be shown that Mendel’s work as well was firmly grounded in progressive technological and political developments of his time.48 Orel, as well as the many contributors he recruited for Folia Mendeliana, did this to great effect. Their collective efforts revealed that Mendel was not only firmly grounded in hybridism, but operated at the intersection of several additional strands of nineteenth-century scientific culture: Through his training at the University of Vienna under the physicist and mathematician Christian Doppler (1803–1853), Mendel became familiar with methods of analytic experimentation involving the application of combinatorial analysis and probability theory. His lifelong interest in meteorology, documented in several publications, additionally deepened his understanding of methods in descriptive and predictive statistics. At the University of Vienna, he was exposed to the teachings of botanist Franz Unger (1800–1870) who promoted a “physics of the plant organism” that involved experimentation and the search for mathematical laws governing the distribution of organisms in space and time, and throughout his life, Mendel moved in intellectual circles that discussed evolution also for its political implications (tensions between German and Czech nationalists were running high in the Year of Revolution 1848). Unger was also

46

Paleček (2016). Due to complex local developments in post-communist Brno, there are two Mendel museums now in Brno. The Mendelianum was moved to the Moravian Museum, while a new Mendel Museum sponsored by the Masaryk University was established in the monastery. Folia Mendeliana continues to appear (see http://www.mzm.cz/en/folia-mendeliana/), but its contents are unfortunately not yet available online. 47 Orel (2005). “Rehabilitating” Mendel was not just an academic question: Kříženecký lost his university position in 1948 and even spent 18 months in jail in 1958 (Orel 1992). At around the same time, Orel lost his job as head of the poultry research unit at the Agricultural University in Brno (Paleček 2016). 48 On Hessen, see Freudenthal and McLaughlin (2009).

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teaching cell theory, and a visit by Jan Purkyňe (1787–1869) to Brno in 1850 contributed further to Mendel’s very advanced understanding of fertilization and plant development. Finally, membership in the Brno Sheep Breeders’ Society immersed Mendel in long-standing debates among local naturalists, farmers, and industrialists about breeding methods and inheritance. These dated back to the early nineteenth century, and Mendel’s predecessor as abbot, Cyril Napp (1792–1867), played a leading role in them. The monastery was, in fact, one of the biggest players in the thriving economy of Brno – the “Manchester of Central Europe,” as contemporaries called it – and it was Napp who actively encouraged Mendel to pursue his hybridization experiments.49 What is crucial about these different strands is the fact that they were independent of each other and that each had its own time of birth, flourishing, and decline. While biologists in the late nineteenth century did not clearly separate between hereditary transmission and development, breeders already engaged in a well-articulated discourse of heredity in its first few decades.50 Doppler’s experimental methods were developed in the 1830s, but such methods only began to infiltrate biology, especially developmental mechanics and biochemistry, in the very late nineteenth century.51 “Austro-Ungerian” botany had seen its heyday in the 1840s and 1850s, but continued to be influential until it waned with increasing acceptance of Darwinian evolution, just as hybridism did.52 Questions regarding the cytological basis of sexual reproduction and inheritance, in contrast, were only beginning to be asked at Mendel’s time, and the field would only see its “watershed moment” in the 1880s.53 These strands were largely independent of each other and fail to form a coherent paradigm, and depending on which one accepts as a benchmark, Mendel appears as “ahead” or “backward” in time. Their conjunction in Mendel’s work, by contrast, was certainly unprecedented and remained unique. On occasion of his retirement, Orel described his scholarly accomplishment as having “pointed out the achievements of Mendel in the special cultural milieu of this country [i.e., Czechoslovakia].”54 And indeed, particularly his work on Mendel’s background in breeding resonated very well with a whole wave of projects on the cultural history of heredity that were initiated in the early 2000s. Various factors motivated these projects. There was first of all the general shift within the discipline of history of science toward a focus on local contexts of knowledge production and the processes of communication and translation that connect these.55 Second, and

49

Orel’s biography (1996) continues to be the best guide to the large body of literature on these different strands. The first volume of Klein’s biography (2013) adds much interesting detail about the German-Czech context of Mendel. 50 Wood and Orel (2001, 2005). 51 Allen (2002). 52 Gliboff (1999) and Radick (2011). 53 Farley (1982); Churchill (1987); Dröscher (2015). 54 Matalová (1992, 118). 55 Shapin and Ophir (1991); Dear (1995); Secord (2004).

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concomitantly, eugenics and agricultural biology had become subjects of serious historical scrutiny in the 1980s and 1990s, suggesting, as Bowler had most prominently, that any history of heredity not attending to relevant developments in the political, medical, and agro-industrial sphere would be essentially incomplete.56 Third, and perhaps most importantly, inheritance as a concept in both the social and the life sciences entered a crisis in the early 2000s, raising the prospect that non-genetic, “alternative” inheritance systems might again move center stage in the sciences.57 Writing the history of heredity was a way to escape the conceptual grip that the history of genetics had had on historiography. And it is no coincidence that historians embarked on this project precisely in the moment when biologists as well began to explore inheritance beyond genetics.58 A whole range of essay collections and book-length studies, which paint a varied and complex picture of the history of pre-Mendelian heredity, has resulted from this reorientation.59 In terms of methods and approaches, the cultural history of heredity is quite diverse, if not to say, amorphous. But there are a few shared features, such as a focus on the metaphors used to describe reproductive processes. “Heredity” itself is a concept that was imported from legal discourses into the life sciences, and it does not appear to have come into general use in its biological sense before the nineteenth century.60 Historians of law, as well as historians of literature who have looked at expert and non-expert discourses of succession and transmission of land, privileges, titles, and money, have therefore provided important input to the cultural history of heredity, especially by drawing attention to the multiplicity and complexity of ways in which property (and properties) are transmitted in different historical and cultural contexts.61 An emphasis on heterogeneity also characterizes the literature on the cultural history of heredity that has more narrowly focused on conceptions of inheritance within the life sciences. When biologists like Darwin and Galton first began to look into heredity in the mid nineteenth century, they relied on empirical knowledge from very different sources: breeders’ tracts on the uses (and dangers) of in- and outbreeding domestic animals; medical treatises like the Traité philosophique et physiologique de l’hérédité naturelle (1847–1851), in which the French “alienist” Prosper Lucas (1814–1899) discussed the inheritance of mental disease in two hefty volumes; or the burgeoning literature in physical anthropology, anthropometry, and human statistics that had emerged out of the eighteenth-century tradition of a “natural history of mankind.”62 Bowler was right, then, to place emphasis on what

56 Russell (1986); Allen (1991); Olby (1993); López Beltrán (1994). On the history of eugenics, see Weindling ▶ Chap. 7, “The History and Historiography of Eugenics,” this volume. 57 Keller (2000). 58 See, e.g., Jablonka and Lamb (2005). 59 For an attempt at a synthetic overview, see Müller-Wille and Rheinberger (2012). 60 López Beltrán (2004a); cf. Radick (2012). 61 Parnes et al. (2008); Weigel et al. (2013); see also Gayon and Wunenberger (1995). 62 Müller-Wille and Rheinberger (2007).

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he called the “Mendelian” revolution; it was indeed the late nineteenth century in which heredity was consolidated as a widely recognized subject of research, speculation, and explicit debate. Unlike Bowler, however, more recent studies have highlighted theoretical pluralism and coexistence of a diversity of research traditions, rather than discursive closure as a key characteristic of this period.63 Finally, despite their focus on local contexts of knowledge production and transmission, cultural studies of heredity have also promoted a renewed attention to the longue durée of heredity in its various incarnations. Thus, for some very specific contexts – such as medical theories explaining familial diseases, the breeding of domestic animals, and political and legal debates about the status of nobility – it has been possible to trace hereditarian ideas as far back as the early fourteenth century.64 Continuities proliferate and extend back even further to the ancient world, inside and outside of Europe, if one casts the net wider and shifts the focus from the concept of heredity to the broader category of reproduction, hence including disciplines like gynecology and demography.65 It is medical practice and theory, in particular, with its inherent political dimension of governing life for the sake of health and its consequent exposure to the contingencies and accidents of “real” life, that challenges the historian’s imagination with a bewildering array of coexisting conceptions of heredity, both “ancient” and “modern,” at any point of its history.66 Seen against this background, modern hereditarian ideas appear less specifically tied to the name of Mendel, but rather associated with the long-drawn processes of nation state formation, the rise of capital and industrialization, as well as European colonialism and imperialism.67 The story of Mendel’s “long neglect” thus appears less and less as a focal historiographical riddle. Nothing about the sciences of heredity before 1900 seems to exclude Mendel as “abnormal” nor predetermines his “rediscovery” as founding father of a new science at the beginning of the new century. It has therefore become conceivable to think of Mendel’s innovation as firmly grounded in the hybridist tradition and yet taking it a crucial step further by creating an experimental system that later Mendelians could exploit productively.68 “Mendel’s discovery,” in all its different senses, was the result of thoroughly contingent circumstances that fail to fit with a clear taxonomy of “paradigms,” “epistemes,” or “styles.” In the light of current epigenetics and theories of extended inheritance, it therefore seems perfectly possible that other inheritance mechanisms than genetic transmission could have been foregrounded in the life sciences at the beginning of the last century, and to raise the counterfactual question how twentiethcentury life sciences would have developed if Mendelian views of inheritance had

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Rheinberger (2008) and Kampourakis (2010). Lugt and Miramon (2008). 65 Hopwood et al. (2018). 66 Gaudillière and Löwy (2001); Gausemeier, Müller-Wille and Ramsden (2013); Gausemeier (2014), and Porter (2018). 67 López Beltrán (2004b); Müller-Wille (2007); Waller (2012); Müller-Wille and Brandt (2016). 68 Müller-Wille and Orel (2007); Rheinberger and Müller-Wille (2017). 64

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not prevailed.69 In other words, the central historiographical riddle for the historian of heredity today with regard to “Mendel’s discovery” is not anymore why it was neglected but how it could have happened at all.

Outlook “History is ... never history, but history-for” according to anthropologist Claude Lévi-Strauss.70 We do not tell stories to the dead, but for our contemporaries and in reaction to present situations that have been shaped by past events but do not represent them.71 Writing history is therefore an inherently anachronistic practice and conditioned by assumptions about the historicity of the subject of our choice.72 Ever since Hessen’s exacting demand to understand Newton “in his time,” it has become commonplace for historians of science, and eventually part of their professional identity, to accuse “scientist” historians, and each other, of anachronisms. But what defines the time scale against which anachronism is to be determined remains a wide open question that will be answered differently by different historians. From this perspective, it can be as anachronistic to call Mendel a “Linnaean” as it may be to call him “the first geneticist.” This does not mean that any historical account is legitimate. Misrepresenting sources, ignoring their contexts, and falling back on facile teleological explanations are bad practice, in history as much as in politics. What it means, though, is that we should not be surprised that developments in the sciences themselves, among many other factors, can precipitate major changes in perspective in the history of science, as I have argued for the “cultural history of heredity” in the last section of this chapter. The opening up of biology for a plurality of inheritance systems in the last two decades has drawn attention to a whole range of potential areas of historical inquiry into heredity that we still know far too little about and that I want to outline in conclusion. One of the ironic results of the long-lasting fixation with Mendel is that we know little about the wide variety of hereditarian ideas around 1900 and especially their legacy in the twentieth century. The little we do know has emerged from the study of debates between Mendelians and other schools of thought and practice like the biometricians or proponents of cytoplasmic inheritance.73 But studies of continuing traditions within embryology, cytology, or biochemistry in their own right are relatively rare, as are studies that trace back contributions of prominent Mendelians to their formative years before the annus mirabilis 1900. A promising start has been made with attempts to write the history of evolutionary developmental biology 69

Radick (2016). Lévi-Strauss (1962/1966, 257). 71 Collingwood (1994). 72 Canguilhem (1966/2005). 73 Sapp (1987); Olby (1989). 70

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(evo-devo), epigenetics, and stem cell research, but many more lacunae – such as the continuing influence of Nägeli on early twentieth-century colloid research – need to be filled before we can even get a clearer picture of how pervasive the dominance of Mendelian genetics actually was in the biological and biomedical sciences of the twentieth century.74 A further aspect waiting to be elucidated is the entanglement of heredity with politics. Again, this may come as a surprise, since the many studies that exist on the history of eugenics and scientific racism seem to evince a very clear connection between heredity and politics. These tend, however, to understand “biopolitics” – against the intentions of Michel Foucault who introduced the term – as a politics of the bureaucratic nation state and the scientific elites it coopted and thus conceal how concepts of inheritance also operate on the more mundane level of families and property transmission. Heredity itself is a very male idea and its history inseparable from the history of patriarchy.75 Yet it is rare that perspectives from gender history and historical anthropology are employed in the study of heredity, and the history of hereditarian ideas in relevant disciplines, such as gynecology and pediatrics, has up until recently hardly figured in the historiography of heredity.76 The recent expansion of the cultural history of heredity to a more inclusive cultural history of reproduction – no doubt motivated by the rapid development of new reproductive technologies – is bound to open up new avenues of historical inquiry in this respect, and historical anthropologists and social anthropologists have a wealth of conceptual tools to offer for such inquiries.77 A final area in the cultural history of heredity of which we know too little concerns ideas and practices in non-European and premodern contexts. To wit, various attempts to explain similarities between parents and their children are well covered for some of the more prominent early modern natural philosophers.78 However, the vast domains of medical and agricultural literature, which premodern Europe and other civilizations such as the Arab-Islamic world or China produced, have hardly been touched upon by scholars with a specific view on heredity.79 One may argue as some have done, including the author of this chapter, that heredity is a concept that simply did not exist outside of modern Europe. But that may itself just reflect a historical bias that was brought about by Mendel’s discovery.

74 Laubichler and Maienschein (2007); Barahona et al. (2010); Dröscher (2014). On Nägeli and colloid chemistry, see Liu (2016, ch. 4). 75 Jordanova (1995). 76 See Arni (2015) for an interesting exception. 77 Franklin (2013); Sabean et al. (2013). 78 Smith (2006). 79 For notable exceptions, see Lugt (2004) and López-Beltrán (2007).

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The History and Historiography of Eugenics Paul Weindling

Contents Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Evolution and Eugenics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Conceptual Shifts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Positive and Negative Eugenics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gender . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Imperial Roots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Internationalism/Transnationalism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Critics of Eugenics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Victim Narratives . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Newgenics and Resources . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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Abstract

The history of eugenics has been one of the most dynamic and challenging areas within the history of biology offering critical new perspectives in modern social history. Eugenics offered modern, scientifically based “solutions” to crime, poverty, and social deviancy. The eugenics discourse became complex with diverse positions on the questions of race, birth control entitlement, and welfare benefits. From a methodological point of view, discourse risks ignoring how context and social processes shape meaning, overlooks issues of impact, and can amount to little more than an old-fashioned narrative of intellectual progress. One challenge is to look beyond discourse to such social processes as professionalization and the construction of welfare institutions and practices. Eugenics provides insight into wider social transformations showing how science represented differing social agendas.

P. Weindling (*) Oxford University, Oxford, UK e-mail: [email protected] # Springer Nature Switzerland AG 2021 M. R. Dietrich et al. (eds.), Handbook of the Historiography of Biology, Historiographies of Science, https://doi.org/10.1007/978-3-319-90119-0_9

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Introduction Historical research on eugenics has sustained itself as innovative and constantly raising new social, gender, epidemiological, and cultural perspectives for investigation since the 1970s. Over the past 50 years, the history of eugenics has been one of the most dynamic and challenging areas within the history of biology and history of modern science. As history of medicine moved from marginality to entering the historical mainstream, the history of eugenics provided critical new perspectives in modern social history. Once it was realized that eugenics was not a prerogative of the right but part of the process of transition to welfare in a modern society, multiple avenues for historical investigation opened. There was a wide spectrum in terms of political and religious affiliation, and accordingly eugenic ideas showed considerable variation concerning reproductive choice. This was apparent as feminists and sexual reformers developed more libertarian forms of eugenics and clashed with authoritarian advocates of family policy and reproductive controls (Grossman 1995). Scientific positions varied and innovated in line with ongoing research in a plethora of disciplines from anatomy to zoology. A range of medical specializations such as endocrinology, gynecology, psychiatry, and venereology as well as statistics and physical anthropology have been of relevance to achieving the health and fitness of future generations. Eugenics offered modern, scientifically based “solutions” to crime, poverty, and social deviancy. The eugenics discourse becomes complex with diverse positions on the questions of race, birth control entitlement, and welfare benefits. From a methodological point of view, discourse risks ignoring how context and social processes shape meaning, overlooks issues of impact, and can amount to little more than an old fashioned narrative of intellectual progress. One challenge is to look beyond discourse to such social processes as professionalization and the construction of welfare institutions and practices. Eugenics provides insight into wider social transformations showing how science represented differing social agendas.

Evolution and Eugenics Eugenics (literally “good birth”) was a term invented by the Victorian statistician Francis Galton in 1883 to define the science of breeding a better race. At the heart of eugenics has been the sustaining by scientifically based measures of physical and psychological fitness over generations. Galton was concerned about how industrialization resulted in profligate breeding among “degenerates” (defined by deviant social traits and mental capacity), whereas the fit and healthy were restricting their fertility. This was the result of industrialization distorting the natural balance of reproduction: Galton defined population structure by a bell curve to depict the distribution of mental and physical capacity. Eugenics was designed to correct the distortion by curbing the fertility of persons deemed “unfit” and offering inducements for persons of higher reproductive value to have at least three children.

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The science of heredity rapidly developed from 1850 to 1900 in terms of observable cellular processes of reproduction and experimental plant and animal breeding. Eugenics involved analyzing the statistics of biological variation and the transmission of characters from one generation to the next. Evidence-based measures could be imposed to sustain fitness over generations. This new science – or more accurately the combination of sciences – focused on problems of hereditary health. Galton prescribed new social practices such as a certificate of good health prior to marriage (so screening for sexually transmitted diseases), and his analysis of changing disease patterns was innovative. The cultivating of good heredity qualities for the collective entity of the nation or “race” provided rationales for welfare measures, albeit on a selective basis. By 1900 demands for more radically interventive measures arose to “solve” social problems. State medical officers saw a sense of duty to maintain the health of the population as a collectivity, so introducing a new ethic in medicine that duty to society and future generations overrode the medical ethic of care for the individual. The eugenically minded conceived an agenda to segregate and sterilize persons deemed “unfit.” It could mean detention in custodial institutions and the controlling and segregating of ethnic groups (such as Roma/“gypsies,” aboriginals, and native populations) so that the hereditary health of future generations should not be “contaminated.” Among those vulnerable to negative intervention were offspring of racially mixed marriages, adolescents labelled “mental defectives,” and mothers deemed “incompetent.” Ideas of differential qualities – whether of “health,” “intelligence,” or of a supposed racial entity – were used to define the differing “value” of persons. Eugenics was therefore elitist in the sense that it was critical of mass democracy and in giving an expert elite powers over human reproductive capacity. “Positive” measures ranged from improved diet, housing (Rosental 2016), and the better management of pregnancy and birth, whereas “negative” practices involved segregation, sterilization, and – at their most extreme – eradication of those deemed to be of “lesser value.” Eugenics marked a break with Victorian ideas of mass improvement by education and the inculcation of moral codes: power should pass to a meritocratically selected expert elite (Paul 1984). In this sense eugenics reflected modernist social structures of policy and planning by technocracy (Weindling 1989). Eugenics marked a change in the secularization of attitudes to sexuality and reproduction. Although a statistical necessity, Galton saw eugenics as a natural religion, realizing that popular belief in eugenic measures would be essential to ensure compliance. Ideas of sin and divine predestination were replaced by notions of hereditarily determined disposition to disease and intellectual attainment. Galton devised statistical methods (such as the correlation coefficient) to address speculative points in his cousin Charles Darwin’s theory of Pangenesis (1868; Holterhoff 2014). Darwin speculated on circulating gemmules which transmit characters and traits to the next generation. Although some historians have alleged that “social Darwinism” led to racist policies, this is highly implausible. Darwin was opposed to slavery and accepted the unity of “mankind” albeit that there were subspecies or “races.” It is an ahistorical distortion to attribute racial thinking to Darwin, whose concept of a species was not fixed but in continuous variation.

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Darwin’s Descent of Man (1871) showed variations on the issues of race and the plausibility of Galton’s social theories. Eugenicists set their fledgling science within issues of changing patterns of morbidity and mortality and of population and socioeconomic structures. They offered pioneering analysis of the fundamental social and biological implications of changing patterns of the “transition” from infectious to chronic degenerative diseases, the reduction of infant mortality, and the change to smaller, “nuclear” families (Soloway 1990). Having diagnosed the transformation of social structures and values, eugenicists prescribed rescue measures from (in their view) a horror scenario of the profligate reproduction of the physically and psychologically unfit with a surfeit of low-quality births. Eugenicists adopted a critical stance toward what they decried as the new modern norm of the “the two child system” and the pathological effects of industrialization in terms of physique, poor health, and depraved psychology. One prescription was that families of high eugenic value should have three children. The reproduction of others deemed unfit should be discouraged: children with unmarried mothers were believed to be prone to degeneracy. Within the wider context of Victorian politics, Galton’s ideas represented a fundamental break from liberal notions of moral self-improvement by education and a shift to meritocracy and a society governed by experts. They modified utopian models of society and ideas of sexual equality, as advocated by the French philosopher Auguste Comte and the Victorian John Stuart Mill, by regulating sexuality and reproduction. The issue of reproductive controls and their intention regarding the perfection of mankind prior to Galton raises issues regarding whether there was a “pre-Galtonian eugenics” among Enlightenment and liberal philosophers seeking perfectibility and the strengthening of state power through population increase; indeed earlier utopian models of the qualities of an ideal society reach back to ancient philosophy. Some looked to Plato’s notion of state regulated reproduction and classical ideas of an ideal physique (Carol 1995; Chapoutot 2016). Eugenicists found inspiration in utopian and especially rural colonies. The German physician and pioneer of “racial hygiene” Alfred Ploetz (1895) found a rift between concept and practice in the Indiana community, New Harmony, or in the Oneida commune, and saw how biological principles could provide a model for social action. Whereas Darwin opposed slavery and placed faith in reformist political agendas, Galton’s 1865 article on “Hereditary Talent and Character” in Macmillan’s Magazine and his 1869 study of Hereditary Genius stood for a shift to meritocracy and professional elitism. The historian Ruth Schwartz Cowan (1985) noted the far-reaching political implications: Galton’s scientized social model represented a critique both of hereditary aristocratic privilege and of its converse, the commercialism of mass industrial society as physically and psychologically degenerate. Galton’s model was of a stable society with physical and psychological attributes distributed as a Gaussian curve. Galton correlated attributes like height and intelligence but considered there to be a “natural” tendency over generations to revert toward a mean as regards to the distribution of characters. This meant higher numbers at the midpoint of the distribution. At the other end of the spectrum were

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those with “low grade” and “high grade” characters. Each generation had to identify talent through regular inspections and examinations. This “natural” model was under threat because of the pathological distortions of industrialization. As Donald Mackenzie (1981) noted in a fundamental and pioneering study of mathematical statistics as socially constructed, here was an ideology of a sector of the professional middle class claiming social leadership. Eugenics thus diagnosed the “social pathology” of modernity but also defined the therapeutic role for the professional elite as the scientific arbiter of social affairs. This meritocratic social model of the structural and ideological effects of industrialization and urbanization was to be transferred to other “developing” societies. Eugenicists called for surveys of the physiological and mental qualities of populations and preventive sifting of migrants. Eugenicists were pioneering in diagnosing issues of the transition to modernity in terms of the biological implications of social structures and values, but they condemned modern individualistic culture in mass urban settings as degenerative. Eugenicists felt entitled to privileged positions of power in a modern technocratic society. They claimed power because they could provide “solutions” to the menace of the mentally and physically disabled. They categorized as inferior “races,” the native and nomadic peoples such as Roma, Sami (laps), the travelling Yenish, bushmen, and aboriginals. Eugenicists looked at the package of traits that they variously defined as a “race” and devised scientistic measures for how mankind could direct its own evolution. Eugenicists were prescient in the consideration of the onset of social changes of modernity and as a critique of curative medicine, philanthropy, and state welfare as exacerbating social problems. Eugenicists in tackling the problem of the reproduction of the “unfit” have taken a lead in providing rationales and measures not just of selective welfare but for regimes of prejudice and persecution. This is sinister science to impose regimes of control, segregation, and destruction. Eugenics was not an autonomous science but a set of sciences applied to “solve” issues ranging from mental health to migration. The forms of projecting eugenic ideas and values have developed from public discourse to state imposed propaganda. In the “liberal era” from the 1880s to the 1900s, eugenics was a scientifically informed public discourse with articles in Victorian reviews, public lectures at events like the British Association for the Advancement of Science, and exhibitions (such as the International Health Exhibition, South Kensington, of 1884) when the public underwent psychological tests. In the decade before WW1, eugenics became organized with new specialist journals, dedicated societies to further eugenic aims, and a first International Congress held in London 1912. The shift from discourse to organizations occurred at a time when new welfare structures were being organized. New corporate foundations such as those of the Rockefeller Foundation and Carnegie Trusts looked for scientific solutions to intractable and costly welfare problems of poverty, crime, and mental illness. National research institutes had variously private and state support. The Galton Laboratory in London derived from Galton’s bequest. In the USA Mrs. Mary Harriman used funds from a railroad fortune to establish the Eugenics Record Office in 1910 under C.B. Davenport close to his Cold Spring Harbor experimental station and in 1918 handed ownership of the Office over

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to the Carnegie Institution of Washington with additional finance. John D. Rockefeller Jr provided operating costs for 4 years. But the Carnegie ceased to fund by December 1939, once the Rockefeller Foundation (RF) took a negative view of Cold Spring Harbor since the early 1930s (Allen 1986). John D. Rockefeller Jr and the photography entrepreneur George Eastman made personal donations to the American Eugenics Society after it was founded in 1923. State support was crucial for the Swedish State Racial Biological Institute in 1922 under Herman Lundborg and in Germany for the Kaiser Wilhelm Institute for Anthropology founded in Berlin in 1927 (Schmuhl 2008; Weindling 1985). Eugenic permeation of public health gave the nascent movement considerable power which can be seen in highly original structures. During the 1890s medical officers conceived ideas of regular medical inspections and “health passports” for preventive surveillance of total populations. From around 1900 eugenicists were active in major organizations to combat sexually transmitted diseases, alcoholism, and other chronic degenerative diseases (Porter 1999). During WW1 eugenic ideas of healthy reproduction of at least two children for the nation and race became mainstream. State and local authorities began to build up hereditary data banks (Erbkarteien). Interwar eugenicists had a major role in the new welfare states in Europe and in isolationist and New Deal, USA. From the 1920s a series of notable films publicized eugenic issues, the first being The Black Stork of 1917 (Pernick 1996). Progressive minded geneticists such as JBS Haldane in his 1923 Daedalus, or, Science and the Future on in vitro fertilization and Hermann Muller’s Out of the Night (1935) on the possibility of freezing germ cells outlined elaborate biologically based utopias. Warning of degeneration was a central theme in naturalist authors such as Gerhart Hauptmann, Thomas Hardy, and Emile Zola, and the pioneer author of science fiction H.G Wells saw the relevance of eugenics. Conversely, the author Aldous Huxley part satirized and part prophesied about eugenic hatcheries in his dystopian Brave New World, and Sinclair Lewis’ novel (1925) Arrowsmith comically depicted a fake eugenic family in the public health paradise of “Nautilus.” Scientific critics of eugenics realized the dangers of coercion and medical abuse. Rather than an undisputed inexorable rise, historians have neglected the extent to which eugenicists were confronted with forthright criticisms of how they posed a threat to civil liberties. Figures like the social scientist Friedrich Hertz engaged in a sustained critique of eugenics. The historically neglected critique of eugenics circumscribed influence. While criticisms culminated in the UNESCO Statement on Race of 1950 (1969), there was a scientistic reaction resulting in successive modifications. The debate on eugenics peaked at the time of the Nazi hegemony. During the 1930s eugenicists divided over whether the Nazis should be applauded for eugenic measures such as coerced sterilization or denounced for misapplication of hereditary science. The international community of eugenicists split on these issues: a leading eugenicist like Julian Huxley (1935) was forthright in his criticisms of the falsehoods of Nazi race theory and of anti-Semitic racial policies. Indeed, “pseudo-science” was a term introduced to undermine eugenic and racial biological schemes. In Nazi

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Germany some scientists clashed with racial ideologues, whereas others, notably Eugen Fischer and Otmar von Verschuer, integrated science with the ideology of race purity. The eugenicist Fritz Lenz initially advised the SS on marriage procedures, but as the Holocaust got underway, he disengaged from racial policy. Interwar and Vichy France (Drouard 1992) and Fascist Italy stressed positive pro-natalism or puericulture, exemplified by the new model community Fertilia in Sardinia. The eugenically minded demographer Corrado Gini regularly briefed Il Duce on fluctuations in the birth rate (Cassata 2006; Ipsen 1996; Quine 2012). Disentangling the complex elements of eugenic science from other strands of Nazi ideology concerning “living space” and anti-Bolshevism and anti-Semitism remains a complex historical challenge which is still only very partially achieved. At times racial hygienists were rebuffed by Nazi ideologues, but at times reproductive and psychiatric research indicates a far-reaching grasp on the mechanisms of mass destruction. The physician and anthropologist Josef Mengele exemplifies how much there is still to disentangle. Mengele had dual qualifications in anthropology and in human genetics and while at Auschwitz from May 1943 took a key role in selecting Jews and Roma for forced labor or poison gas; from May 1944 he retained several hundred Jewish twins and dwarves for racial and hereditary research (Müller-Hill 1984). Issues arise as to the role of eugenics in Nazi allies such as Hungary and Romania, and other authoritarian states, as well as Nazi sympathies in the wider international eugenics community. By the end of the war, eugenicists saw the way forward as denouncing Nazi atrocities as having nothing to do with eugenics (Blacker 1952). Post-WW2 eugenic policies permeated international measures to combat the “population explosion” on a global basis (Connelly 2008). How to write this history varies between the idea of an expert conspiracy or enlightened benefactors. The US birth control lobbyist and feminist Margaret Sanger promoted women’s understanding of the benefits of contraception. Fundamental issues arose regarding access to birth control, especially the contraceptive pill. Eugenicists sought alliances among major religions, while they clashed with feminists over freedom in sexuality, birth control, and prevention and interruption of unwanted pregnancies. Genetics was pioneered by the Moravian monk Gregor Mendel (1866) and then rapidly developed after “rediscovery” of the Mendel’s hereditary principles of the transmission of characteristics in 1900. Genes were located on the chromosomes, and genetics as a science rapidly advanced in terms of gene function, composition, and structure. The German physician and biologist Alfred Ploetz (1895) outlined a new science of “racial hygiene” based on what he called chromosomal engineering. The focus on the chromosome as the carrier of hereditary substance opened the way to appreciation of the Mendelian ratios. Between 1900 and 1940, Mendelism was developed by the German pioneers of “racial hygiene” Erwin Baur, Eugen Fischer and Fritz Lenz (1921), and Ernst Rüdin. Relations between genetics and psychology (e.g., developed by Rüdin’s research on the inheritance of “dementia praecox” or “schizophrenia”) shaped policies of institutionalization, sterilization, and (in Nazi Germany) systematic killing of the mentally ill and disabled. As the dreams of eugenicists have come to be realized in modern reproductive technologies, and

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genomic research has advanced, there have been successive historical investigations of the changing science of human heredity and of social aspects of the organization and dissemination of eugenics. Molecular biology with the post-WW2 discovery of the double helix structure of DNA and new reproductive technologies such as the contraceptive pill opened the way to a modern understanding of the processes of heredity. But whether this occurred in ways linked to eugenic thinking on population quality or as a fundamental break marking the demise of eugenics has been historically controversial (Kay 1993; Paul 1995). Questions remain as to the demise or transformation of eugenics as human genetics. The “chromosomal engineering” envisaged by Ploetz but not realizable in the 1890s has become possible, at least to some extent with respect to assisted reproduction, gene line editing, animal cloning, and gene therapy. This presents researchers and the public with new bioethical challenges.

Conceptual Shifts Until the 1970s the chronicling of eugenics was positivistic and written by insiders to legitimate and publicize “pioneers.” The noted British eugenicist Carlos Paton Blacker (1952) published on Eugenics Galton and After. The Eugenics Review (1909–1968), the Annals of Eugenics (1925–1954), and the German journal Archiv für Rassen- und Gesellschaftsbiologie (1904–1944) contained numerous eulogies of eugenic pioneers. Samuel Jackson Holmes (1924) published a notable (and still usable) bibliography documenting the international range of periodical papers on eugenic topics. Marie-Thérèse Nisot (1927–1929) produced an early comparative survey, taking a positive view of eugenics. Modernizers favored welfare state and international agencies, shifting away from imperialism to a global population-based perspective. The mercurial biologist Julian Huxley (1935) opposed the older imperialist racism and then decisively Nazi race theory, contributing to the anti-Nazi We Europeans (1936). The tone of eugenic histories was futuristic and oriented to rational planning (Wolstonholme 1963). The interpretation was of a non-imperialist idea of “reform eugenics” oriented to state welfare. Huxley (1946) as the first Director-General of UNESCO controversially saw eugenics as the basis for international education and science. The US civil rights movement and wider movements for civil liberties, abortion law, and homosexual law reform of the 1960s generated not only a complete reappraisal of attitudes to race and gender but also a critical approach to studies of science. The situation arose when a new critical history and the traditional positivism ran parallel. In 1963 Mark Haller (1963) published his landmark Eugenics: Hereditarian Attitudes in American Thought. The old guard of biologists, notably Julian Huxley (1946), the cytologist J.R. Baker (1974) writing on race, the botanist C.D. Darlington (1969) on sociobiology, and Frederick Osborn in the USA as founder of Eugenics Quarterly (1954–1970, then continued as Social Biology) continued to propagate racialized and scientized versions of history (Stern 2005). But the situation began to change by 1970 as the social critique of science began to take off. Such new

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thinking impacted on science itself. This marked change of social views can be seen in Watson (1971). There was a transition to critical views of scientifically based experts in the late 1960s. For the claim was that experts should be accorded state resources without public accountability. The result was a new politics of scientific knowledge. This could be seen in 1970 with the establishing of the Society for the Social History of Medicine in UK, as well as feminism and gender studies. Geoffrey Searle (1976) pursued a wider sociopolitical significance in terms of Edwardian welfare politics. Robert M. Young’s critical studies of Darwin and social Darwinism (1985) provided a background to social studies of eugenics. The historical debates with Young’s radical science group gave rise to publications on the roots of sociobiology (Webster 1981). Donald Mackenzie (1981)’s powerful analysis of British eugenics represented a landmark study. Coming out of the sociologically informed Edinburgh science studies group, key issues were how social and economic status translate themselves into scientific innovations and applications. Mackenzie considered eugenics as offering a social handle on mathematics by showing that statistical methods were socially constructed. Mackenzie focused on the elite – or active nucleus – within British eugenics, discerning how different types of statistics were related to socioeconomic position. The focus of most studies of eugenics in the 1980s was strongly on the genetic component of eugenics, albeit socially constructed and ideologically driven. Essentially the concern was the misapplication of genetic knowledge, sometimes because the science was flawed and sometimes because the application was racist and in clear conflict with the libertarian ideals of civil rights activism and the takeoff of feminism in the USA. These were the concern of pioneering generation of historians of US genetics, notably Garland Allen (1986), Barry Mehler (1988), Daniel Kevles (1985), and Diane Paul (1995). German eugenics attracted pioneering attention of the human geneticist Adela Baer had uncomfortable encounters with a resistant German generation in the late 1970s. The geneticist Benno Müller-Hill (1984) conducted pioneering research and interviews indicating the falsehoods in narratives current by the 1980s. Jonathan Harwood (1994) produced a finely researched analysis of contrasting scientific styles among of German geneticists. Harwood demarcated between animal and human genetics although it could be argued that animals were often seen as models for humans. A slim but influential work, Bernhard Schreiber’s Men Behind Hitler (1975) received serious but critical attention from the Oxford historian, Hugh TrevorRoper (1975) and has appeared in multiple formats since the 1970s. Schreiber alleged international linkages between Nazi eugenics and population and mental health organizations, by analyzing membership lists. The post-war German genetics community divided between apologetics and disclosure: the geneticist Widukind Lenz supported critical historical investigations of his father’s (i.e., Fritz Lenz’s) generation. Müller-Hill (1984, 1998) broke new ground by locating hitherto concealed documents on the German Research Fund’s support of Mengele and programs of hereditarian research. He documented links with Nazism, exposing

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how aged scientists told him falsehoods contradicted by documents from the period of National Socialism. Case studies of hereditary biology indicate the rising funding of coercive and racial research (Cottebrune 2008). Germany and Austria still need to make good incomplete evidence on issues like the victims of medical killing and the subsequent use of body parts and brain tissue for research on into the post-WW2 era. Mitchell Ash (2002) has opened up the problem of the construction of “resources” for science: who the victims of Nazis like Mengele remains only very partially answered (Weindling 2014). The neglect of the identities and narratives of research subjects is a gaping flaw in studies of East Central Europe. Identifying impacts on research subjects represents an exercise in historical accountability by major funders of research such as the Max Planck Society and German Research Fund, as well as by individual German and Austrian universities. Swiss eugenics showed how the sterilization schemes of Rüdin could be applied in cantonal democracies (albeit with women excluded from voting), as well as throughout Nazi Germany. The social history of biology offers a powerful resource in terms of concepts and analytical approaches to the social dimensions and impacts of science and as markers of social mentalities and structures. Social historical approaches to science meant a ready response to Mackenzie’s landmark analysis (1981): prosopography remains a powerful research tool, as do analyses of institutional structures and funding. Biographical and institutional studies extended understanding of how eugenics permeated British regions, for example, Greta Jones (1992) on the Belfast Eugenics Society, and internationally. Pauline Mazumdar (1992) extended the range of coverage of British eugenics, by considering a wider range of sciences, looking beyond genetics to immunology and epidemiology. Kenneth Ludmerer (1972), Barry Mehler (1988) and Garland Allen (1986) tackled US eugenics with a primary focus on applied animal and human genetics. Successive research has examined the interaction of US foundations, scientific lobbyists, and legislation throughout the USA as regards issues like migration controls and the role taken by eugenicists in sustaining WASPish dominance of the isolationist interwar USA. US analytical models were a powerful inspiration for studies of eugenics in other contexts. During the 1980s Robert Proctor (1988), Sheila Weiss (1987), Dikoetter (1998), Otsubo (1998) and Paul Weindling (1989) tackled German “Rassenhygiene” from different analytical perspectives: Proctor examining the role of science in Nazi ideologies of health and reproduction and Weindling indicating impact on public health organizations and administrative structures. A science policy approach was taken by the group working with Peter Weingart (Weingart et al. 1988). William Schneider (1990) opened up the issue of French eugenics as distinct from pro-natalism. The dynamic of studies of eugenics in state contexts spread to Scandinavia (Broberg and Roll-Hansen 1996), where there was a powerful impact on welfare measures with imposition of “negative” sterilization. Other distinctive forms of eugenics were discerned by Mark Adams (1990) for Russia/the Soviet Union (Krementsov 2006, 2011). Latin America was opened up by Nancy Stepan (1991), although her model based on limited sources for only three countries has been substantially revised (e.g., by Eraso (2008)) There followed studies of

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South Africa, Australia, and New Zealand (Klausen 2004; Garton 1994, 2010; Wanhalla 2007, 2013). A global history of eugenics emerged in studies of the transition from imperialism to globalism. Wolfgang Eckart (2012) looked at the German colonies in shaping ideas of racial hygiene. The African basis of the human genetics of Eugen Fischer on the Herero showed how imperialism fuelled racial research. Fogarty (2010) examined eugenic linkages to the French colonies. The interwar period showed a powerful take up of eugenic issues at a time of “national self-determination” embedded in the Treaty of Versailles. The successor states in Eastern Europe and the Baltics all showed a strong eugenic impact in areas like public health and anthropology (Turda and Weindling 2006; Felder and Weindling 2013). Countries like Greece (Trubeta 2013) and Romania (Bucur 2002) show a strong role of expert elites within distinctive ideological frameworks. Other historical studies have examined Asian cultures, notably India, China, and Japan (Dikoetter 1998; Otsubo 1998). Historians of science saw how scientific and medical structures had a distinctive autonomy and that scientific rationales needed uncovering. The studies of German eugenics in the 1980s challenged a simplistic account of a shift from volkisch racism (an undifferentiated assortment of ideologies including social Darwinism) linked to anti-Semitism which mutated into Nazism. George Mosse (1964) successfully linked a range of disparate themes – including sport and sex reform – but paid scant in regard to social structures of professionalization and administrative and civic structures. Mosse’s positioning of eugenics has thus been problematic in ignoring how linkages between racial hygiene and eugenics were forged. Placing the ideology, lobbies, and institutions of health before those of race opened up a hitherto historically unexplored terrain, offering a corrective to Nazism as an outgrowth of volkisch anti-Semitism. The sociologist Zygmunt Bauman (1989) offered the intriguing concept of a “gardening state” (drawing on T.H. Huxley) but insufficiently developed issues of scientific expertise. Whereas earlier work empirically analyzed institutions, biographical profiles, and rationalizing ideologies, studies using the concept of biopower/biopolitics risk being empirically lax as the concept becomes a substitute for substantive explanation. Although enriching national historical narratives, the danger is one of legitimating genocidal nationalism in that being a dedicated nationalist does not excuse exterminatory agendas (Weindling 2016). The problem with the concept of biopolitics is that the conceptual label becomes by default an explanation, while the specificities of what happened to whom are glossed over. Wider issues of how a diversity of scientific and social agendas were constructed, implemented, and impacted on people’s lives become redundant. This simplifies eugenics as a reference point in general historical narratives, whereas an authentic analysis would reveal diverse actors, policies, opponents, and victims. The eugenic components of “race” and “hygiene” have been subject to a wide range of conceptual variation. Eugenics has therefore been subject to constant variation, shaped by ideological and social parameters. Some historians have seen a single trajectory, such as from Darwin to genocide, whereas others see a history of

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discontinuities and variations. It has been necessary to engage with how “race” was scientifically constructed as an underpinning of policies, ranging from welfare and psychological evaluation to persecution and physical destruction (Stone 2002). The theorist of genocide, Raphael Lemkin (1944), recognized how German “professional genocidists” imposed coerced population and sterilization policies in occupied Poland. Historicizing eugenics allows theorized discourse to be integrated with the biographical collectivities of the involved persons, policies, programs, and wider social transformations. The admittedly arduous reconstructions provide historical authenticity for motivating ideals and programs. Wider interpretative frameworks such as “social Darwinism,” “biopower,” and “modernity” are otherwise meaningless, vapid, lax, and lazy without analysis of social forces like professionalization, policies and persons, and the entanglements of conflicting interests, discontinuities, and oppositional groupings.

Positive and Negative Eugenics Galton (1904) distinguished between desirable and undesirable. Ever inventive, he devised health examinations prior to marriage and – on the positive side – maternity insurance. The British obstetrician Caleb Williams Saleeby (1909) formalized the distinction between “positive” and “negative” eugenics. This posed the challenge of how concerns with reproduction impacted on different branches of medical practice and public health administration. This in turn raised issues of gender and population policy, over which there have been a series of clashes over reproductive rights. New systems of public health insurance and state welfare resulted in an expansion of institutions of containment – tuberculosis sanatoria and psychiatric hospitals segregated the sick and deviant between the 1880s and WW1. The Eugenics Education Society achieved noted success in 1913 by establishing a system of “colonies” for “mental defectives.” For many juveniles it meant detention on a lifelong basis (Thomson 1998). The historian of eugenics has been faced with a long march through the history of “welfare” institutions, when issues of segregation took priority over cure. Indeed, eugenicists accused curative medicine of allowing persons with pathogenic genes to freely circulate and reproduce. Tubal sterilization or vasectomy was a surgical technique pioneered from 1890 and then used by Albert Ochsner for the so-called degenerates. Another technique for sterilization that was increasingly considered was that of the newly discovered X-rays by Conrad Röntgen in 1895 in Würzburg and by the time of the WW1 was proven to have sterilizing effects. Soon after 1900 came calls for sterilization of chronic alcoholics and other bearers of hereditary degenerative traits. Eugenicists rapidly saw how sterilization could be used to prevent the proliferation of unwanted progeny. They put themselves in the position of medical guardians of the nation and race and assumed powers over the capacity to father and bear children. Sterilization was hailed as a way of cutting institutional costs. Different target groups were proposed. In 1903 the German psychiatrist Ernst Rüdin called for sterilization for alcoholics. Psychiatric targets included “mental defectives,” schizophrenics, and “idiots” – all highly contentious diagnostic

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categories. Calls for sterilization were extended to racial groups, e.g., Sinti and Roma, Yenish, Jews, aboriginals, Sami, mixed race African-Germans and AsiaticGermans, and vagrants more generally. The first US state to pass sterilization legislation was Indiana in 1907 and then followed, notably, by California where the highest numbers of sterilizations took place (Hansen 2014, King and Hansen 2013, Kline 2001). How these policies were imposed, the target groups and individual narratives of experience, and finally issues relating to disclosure, apology, and compensation have all involved immense historical effort in unravelling medical and psychiatric routines. In 1928 the Protestant Swiss canton of Vaud imposed sterilization (Jeanmonod and Heller 2000). Sterilization became routine in the Nordic welfare states. In 1929 Denmark imposed sterilization on the basis of “voluntary” consent (which could mean professionals using psychologically coercive arguments). The UK with its entrenched system of colonies for mental defectives failed in 1930/1931 to legislate for a proposed law. As in other contexts (notably Finland), sterilizations were carried out outside the law; in the UK these were at a localized and low level (six sterilizations of blind and presumed mentally subnormal children are known to have occurred in Leicester). Much has been made of the US influence on the German sterilization law of 14 July 1933, although the files indicate that the German legislators considered a range of other countries and cases (such as the Danish precedent). Sterilization legislation was imposed in 1934 in Norway, in 1935 in Finland and Sweden, in 1937 in Estonia and Latvia (Felder and Weindling 2013), and in 1938 in Iceland. Municipal sterilization occurred in Zurich and in Basel from 1937 sterilization guidelines of the Medical Society and in Bern and St. Gallen with Christian conservative support. By way of contrast “Latin” (or Southern European), societies such as Italy, Spain, Portugal (Cleminson 1994, 2014), and Romania (Bucur 2002) preferred positive eugenic measures such as financial incentives of a “bachelor tax” or targeted child support. However, one must caution against any gross oversimplification of “Latin eugenics” as dominated by positive eugenic “puericulture.” “Latin” contexts such as Romania and Argentina (Eraso 2008) saw strong advocacy of negative measures such as sterilization. “Biotypology” flourished in non-Latin contexts such as interwar Czechoslovakia. Polarities between “negative” Nordic and “positive” Latin eugenics are gross oversimplifications. Nazi German sterilization had far-reaching institutional/state implications and was unparalleled in its scale. Alfons Labisch and Florian Tennstedt (1985) have examined the restructuring of the state public health services on a hereditary biological basis. Researchers notably Ernst Rüdin and state medical officers such as Arthur Gütt of Reich Ministry of the Interior gained considerable power. New offices for race and hereditary health were established throughout Germany winding up the innovative Weimar German system of social medicine. Between 1934 and 1945, roughly 350,000 persons were forcibly sterilized in Germany and Naziannexed Austria. In addition there were ca. 1000 sterilizations in the so-called Reichsgau Sudetenland (annexed from Czechoslovakia). There were also sterilizations in the annexed former free city of Danzig and the Eastern Baltic Memel

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peninsular. Questions arise regarding sterilization of unfit in the German-annexed Warthegau area of western Poland and for the repatriated ethnic Germans. These operations show a far-reaching transformation of German medicine to issues of population and racial engineering. One issue opened up by Gabriele Czarnowski has been reproductive research involving forced abortions and the use of fetal tissue as a research “resource.” How a scientifically minded profession exploited such policies for opportunistic research has to be reconstructed on a case-by-case basis to have an evidence basis for its extent. Comparative studies showed the prominence of eugenic elites in a range of social systems. At first the comparisons were poorly researched: examples are Loren Graham (1977)’s conceptually innovative but empirically defective study of Russian/Soviet eugenics with Germany. A comparative volume edited by Mark Adams (1990) laid out the American, British, German, and Russian cases. Kevles (1985) produced an account of USA and British eugenics, highlighting the religious idiosyncrasies of the major scientific actors.

Gender Foucault’s ideas of biopower and repression took longer to obtain impact: their influence can be seen in the work of Anne Carol (1995) on France and in studies of Swiss eugenics. The primary agency is the state (albeit weak and localized in Switzerland), and eugenically minded, and generally male professionals. Feminist studies of gender and patriarchy preceded the English-language translating of Foucault: a notable example is that of Jane Lewis’ Politics of Motherhood (1980) showing how the eugenically minded medical official Charles Newman at the British Ministry of Health blamed mothers for the persistent high rates of infant mortality. Historical writing on professional power and gender politics reinforced the conceptually suggestive analyses by the French philosopher Michel Foucault (1963, 1976). Issues of “biopolitics” (1911) and associated ideas of coercive power have historiographically complex roots in terms of the politics of gender and health. Not only did eugenicists blame and manipulate women as part of the politics of paternalism but also women had a leading role in the organizing of a militant, organized eugenics. This shows the plurality of forms that eugenics could take. The League of Protection of German Mothers espoused radical Nietzschean ideas of feminist empowerment. Sybil Gotto was a leading organizer of the (British) Eugenics Education Society concerned with prostitution and sexually transmitted diseases. Women were prominent in the German Racial Hygiene Society, which at one level defined itself as a breeding community. Eugenics deepened feminism showing a concern beyond the politics of voting rights to maternity, reproduction, and health. Richard Evans interpreted biological concerns as illiberal and proto-Nazi rather than emancipatory and reformist. Ann Taylor Allen (1988) has argued resolutely for the deficiencies in Evans’ interpretation based on a narrow and mistaken conceptualization of an illiberal social Darwinism and eugenics, instead of seeing diverse biologically framed positions.

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For Germany, Gisela Bock (1986) – drawing on a theoretical model by Allan Chase (1977) on the supposed racist legacy of Malthus – analyzed gender issues in German reproductive science. Karin Hausen (1984) analyzed how the public health official Hans Harmsen along with the Pro Familia organization pursued an antifeminist agenda in his support for the “German Mothers’ Day.” From the 1970s German history was reconceptualized from below by activists – whether in terms of the radical left, notably Karl-Heinz Roth (1984), feminists, and disability rights campaigners. The “Gesundheitstag” (Health Rally) organized in Berlin in 1980 and at Bremen in 1983 offered opportunities for developing radical reinterpretations of German history – including reproductive rights, an issue marginalized by political historians. Vienna eugenics had – as Ploetz moved rightward – too many socialists, Jews, and women for it to qualify as a constituent of the German Racial Hygiene. Doris Byer (1988) and Maria Wolf (2008) contributed pioneering studies of the Austrian discourse on the “human economy,” and Claudia Spring (2009) has analyzed sterilization for Austria. Interwar Austria and especially “Red Vienna” saw new fields open for women in medicine, nursing, and social work. Vienna saw a pioneering birth control clinic established in 1921, while female empowerment and National Socialism gave rise to a classic debate between Claudia Koonz (1993) and Bock about women and Nazi values. Leading female protagonists for birth control such as Marie Stopes in the UK and Margaret Sanger in the USA show strong linkages between birth control advocacy, eugenics, and feminism (Carey 2012; Hodges 2008). Whether birth control should be under medical control or a women’s right was a key issue as eugenicists often clashed with libertarian minded advocates of reproductive rights: radical advocates of Sexpol skirmished with authoritarian racial hygienists intent on professional sanctioning of fitness to reproduce. Other case studies of feminism and eugenics have teased out the spectrum of feminist opinion on eugenic measures. Authors, activists, and social workers have engaged on a protracted debate over feminism and fitness to reproduce in social settings ranging from Alberta (McLaren 1990) to Zagreb.

Imperial Roots The world’s first eugenics societies were as much internationalist as nationalist. The history of German eugenics is far broader than the history of Germany as a political entity, as the founders wished to promote the health of the “white races.” For the intention was to reach out to a wider world of German colonies and German ethnic groups beyond the frontiers of the united German imperial state. Austria-Hungary (and then after 1918 Austria), German-Swiss cantons, and German settlements in Eastern Europe all had eugenic advocates and groupings. “Loss” of territories and of German colonies after WW1 made the problem of the survival of the Germanic races all the more acute, in the context of starvation and the penalties of the post-war settlement. German eugenicists strode to take a lead internationally, influencing the development of eugenics in the wider world, notably in Nordic Scandinavia and the USA with its large German immigrant population.

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The eugenics movement addressed key themes like modernization and associated changes in family structure and sexuality and epidemiology with the shift from infectious to chronic degenerative diseases. In 1905 the Racial Hygiene Society was founded in Berlin, providing eugenics an organizational basis, and one of a number of societies to improve national health. The new Society’s message was that fitness was a duty to the race. Alcohol, tobacco, sexually transmitted diseases, and tuberculosis – with their own public prevention organizations – were “racial poisons.” Germany – and Austria – had rapidly expanding cities, a declining birth rate, high rates of children born outside marriage, and a range of infectious and chronic degenerative diseases. There were strong German linkages to Scandinavia and to Eastern and Southeast Europe, so that eugenics spread among scattered ethnic German groups such as the “Saxon Germans” in Romania and the Volga Germans. Eugenics impacted too on a multiplicity of medical disciplines as well as the social sciences. Textbooks and handbooks represented a significant means of dissemination of racial ideal as opposed to pathogenic types (Baur et al. 1921; Fangerau 2003). The Eugenics Education Society was founded in London in 1907 on the initiative of Sybil Gotto, later Sybil Neville-Rolfe (1949), and other radical thinkers on sexually transmitted disease, maternity, and the birth rate. Much effort has gone into analyzing the Society’s activities, membership, and structure. Its early members were less concerned with technical issues in demographic statistics (the statistician Pearson (1938) kept a distance) than chronic degenerative diseases, mental deficiency, and reproductive matters. The structure of the EES had local groups such as at Cambridge, Oxford, and Belfast. It also had overseas groups, such as in Australia, New Zealand, India, and South Africa. Further groups crystallised as in colonial Kenya in 1933 (Campbell 2007). This indicates how imperial centers had links throughout the world. Post-WW1 eugenics flourished in an international political context of national self-determination, as well as concerns that the falling birth rate could not replace war losses. Ethnic minorities sought to defend their vulnerability in the new nation states concerned about the ethnic hegemony of the majority groups. Jews divided between Zionists, who were keen to promote a muscular Judaism and those who looked toward eugenics in host societies. Jewish organizations concerned with genealogy and health proliferated (Efron 1994). Ethnic Germans sought to defend their position in contexts like Czechoslovakia, Yugoslavia, the Volga region and Ukraine, and the Siebenbürgen region of Romania (Georgescu 2010). These eugenic groups were beyond the state, as the scattered Germans looked toward a “Greater Germany” for financial and ideological support.

Internationalism/Transnationalism International agencies espoused eugenics as a scientific and rational resolution of global problems. The first societies were international – without a national descriptor as “German” or “British” – albeit in imperial contexts. Competitive rivalry for international leadership arose first at the International Hygiene Exhibition at Dresden

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in 1911 and then at the International Congress at London in 1912 made uneasy by rising Great Power conflicts. There followed congresses in New York in 1921 and 1932, also under strained circumstances. An International Federation of Eugenics Organisations succumbed to Nazi power politics (Kühl 2013). US foundations, notably the Carnegie, Rockefeller, and Milbank Memorial Fund, provided the model for expert-driven interventions, backed by immense wealth. The Milbank, very much an expert-driven think tank, shifted from medical to population issues and to the modern model of “development” and “transition theory” (Weindling 2003). Eugenics fitted well these foundations’ concerns with population and global development. Interwar international agencies were more cautious as regards eugenics. The League of Nations Health Office had its statistical service developed by Edgar Sydenstricker seconded from Milbank. Yet the Health Organization’s charismatic Director Ludwik Rajchman otherwise steered clear of eugenics and birth control. By way of contrast, the International Labour Office under Albert Thomas was more open to overtures by the feminist campaigner Margaret Sanger. The post-WW2 era of the Cold War saw major interventions. John D. Rockefeller III was concerned about rapid growth of Asiatic populations. The Population Council from 1952 acted as intermediary lobbying the United Nations to develop programs for population control. The Council interacted with the Population Office at the UN: eugenics was rebranded as “population ecology.” The Ford and Carnegie Foundations supported the World Population Congress in Rome in 1954 and subsequent initiatives in population control. International and national aid agencies supported sterilization policies in “third world.” From the 1960s the contraceptive pill provided new possibilities. China’s one-child model provided a radical “solution” at a time of improving health within a rural economy. Rising standards of education and prosperity have made the policy redundant.

Critics of Eugenics The Victorian Darwinist T.H. Huxley presciently warned against state selection on the basis of what he first termed as “pseudo-science.” A range of scientific critics of selectionist theories of heredity expressed caution about selection and its theoretical basis (Soloway 1982). Liberal politicians, notably Josiah Wedgwood in Britain, consistently opposed eugenic schemes from the mental deficiency act of 1913 to sterilization Jones 1986. Central Europe saw the emergence of both a eugenic and a critical discourse. The Austrian social scientist Friedrich Hertz published a pioneering critique of “modern racial theories” (Hertz 1904). Hertz’s critique of “modern race theories” took place at the point when eugenics and racial hygiene were emerging as organized movements seeking control by expert elites over areas of social policy with medical and scientific claims and when anti-Semitic organizations began to take an interest in the biology of race. Advocates of eugenics took Hertz’s views seriously. Adopting a legal and sociological approach to the problem of race, Hertz advocated the view

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that: “Race theories are little else but the ideological disguises of the dominators’ and exploiters’ interests” (Hertz 1928). He pointed out that there was no significant linkage between physical and mental characteristics and that race theories lacked a convincing empirical basis. The rise of Nazism prompted an international coalition to resist Nazi race theory. Franz Boas, the eminent anthropologist based in New York, gave significant support. Ignaz Zollschan devoted himself tirelessly in organizing collaborations between scientists, politicians, and religious leaders, not least the Papacy. French activists established an anti-racist populist organization Races et Racisme to combat the spread of “racist” ideas. Elazar Barkan (1992) traces a shift from biological concepts of race to “culture,” becoming fully evident in the UNESCO Declaration on Race (1950).

Victim Narratives Whereas the international health exhibition of 1884 recruited novelty hungry test subjects, accounts of eugenics generally fail to incorporate the experiences and narratives of research subjects. Tony Kushner (2004) has made innovative use of the Mass Observation Archive to critique British public attitudes on race. Being a Nazi racial research subject was a matter of defying the racial stereotypes promoted in a racist exhibition against Jews at the Vienna Natural History Museum. When the face masks taken during a Nazi round up of “stateless Jews” were discovered in 1998, this was an opportunity to gather survivor recollections of the Vienna Jews herded into the Prater sports stadium and then used for anthropological research (Berner and Spring 2004). Research on East European eugenics in Hungary and Romania has failed to include the key dimension of the responses of target populations. This weakens any account that stretches into the era of the Holocaust in East Central Europe. Foucault analyzed how the “medical gaze” was dehumanizing and within closed institutions had an exterminatory potential. Under Nazism there were synergies between clinics and a concentration camp as a device to secure the racial health of the German race. An example is how Eva Justin studied a group of 41 Sinti adolescents at the children’s home at Mulfingen, before they were transferred to the Auschwitz Zigeunerlager (“Gypsy Camp”) under SS doctor Josef Mengele. What is necessary is a full-scale reconstruction of all such instances of racial atrocities. Resistance to Nazi racial research involved evasion, deceiving the researcher, and sabotage. Narratives by Mengele twins and the Ovitz family of (mainly) dwarves show how some twins were not really twins and how they cheeked and challenged their deadly tormentor. The journalist Hans-Joachim Lang (2011) has analyzed survivor memories and biographies. Much myth surrounds Nazi eugenics. These myths arose from a situation when documents were destroyed and the lack of reliable information for victims. A persistent myth was that the SS had a special breeding program to produce blond and blue-eyed children based at the Lebensborn (Lilienthal 2003). Another

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misconception was that the Germans researched to increase the numbers of multiple births. Myths have been particularly attached to Mengele and the sterilization experiments in Auschwitz. The Germans also systematically kidnapped children, particularly in Poland in order to be germanised, and some 20,000 single mothers gave birth at Lebensborn homes. Nazi eugenics was until the 1970s obscured by the history of anti-Semitism. This overlooked distinctive professional and scientific issues not only under Nazism but also stretching from the late nineteenth century on into the post-WW2 periods. One myth which has been brushed aside is that Nazism was antithetical to the scientific community. Research on Nazi eugenics has taken a variety of forms. The dissemination of racial ideas involves examining the racialization of administrative structures and policies and the extent to which Nazi racial agencies drew on eugenics. This in turn raises longer-term issues of the biologization of social policy, expert power, and evolving policies on birth control, family welfare, and genetic counselling. German scientific, political, and ideological transformations provide points of comparison with other countries, as well as wider issues of regulatory structures. The issues of compensation and rehabilitation are one of the most dismal in the history of eugenics. Victims have received scant redress and in the case of Nazi atrocities were subject to neglect and a total lack of rehabilitation. Apart from Germany, compensation has been rarely offered. But far more valuable are healthcare entitlements and, in the case of sterilization, the reversing of the operation. At the end of the war, any comprehensive program to purge German medicine of its racial element was only very partially realized (Weindling 1989; Schmuhl 2008). Although the demand for an operation to reverse sterilization was articulated in post-war Germany, this was not provided by German medical officials. The reversal of sterilization would in fact have had good chances of success in cases of male vasectomy. It would have provided the most effective form of redress. Compensation for victims of sterilization can be characterized as late (from 1980 in the Federal Republic of Germany) and limited. Compensation in terms of a single 5000 DM payment was only granted from 1980, and a monthly pension supplement of 300 DM (now 1200 euro) was approved. A full apology to the victims by the German state has yet to be made, although there have been a series of partial gestures. Compensation is an issue that few historians have engaged with, although revealing much about experiences of eugenic victims. During the 1950s and 1960s Federal Republic of Germany, the 1933 sterilization law was not viewed as a Nazi law but as comparable to USA, Canadian, and Scandinavian laws. It therefore remained on the statute book, but not actively in operation. The League of Persons Damaged by “Euthanasia” and Compulsory Sterilization (Bund der “Euthanasie”-Geschädigten und Zwangssterilisierten) was founded in 1987. It has campaigned for a full repeal of the law and a full apology: both aims have only partially been realized. In September 2014 only 364 surviving victims were claiming this pension, a tiny fraction of the ca. 450,000 sterilized. In contrast Austria has not had a specific scheme but has provided compensation under its generic Nazi victims law (Opferfürsorgegesetz) rather than specifically for sterilization victims. Switzerland decided not to compensate, despite public lobbying for this.

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Coerced sterilization continued in Sweden until the mid-1960s, but the law was finally repealed only in 2012, and the Canadian province of Alberta repealed its 1928 Sexual Sterilization Act in 1972. Coerced sterilization found an echo in population control programs fuelled by the ideology of a global population explosion in the Cold War period. From 1976 certain Indian states have been the targets of programs that were nominally voluntary but in practice involved high levels of coercion (Sharma 2014). China’s one-child policy (recently relaxed) has been the most notorious biopolitical project (Dikoetter 1998). Socially discriminated groups and ethnic minorities such as Sinti and Roma, the Yenish, Native Americans, and the Nordic Sami (Laplanders) have been vulnerable to sterilization. Global population thinking on birth control has become less interventive as birth rates diminish with the participation of educated women in the labor market. From the 1960s vasectomy has become an accepted form of voluntary contraception in the West. Disability rights campaigners have extended the positive acceptance of the variety of physical and mental states. The biological and genetic notion of schizophrenia was questioned by radical psychiatrists, and from the 1970s medication has been found to be more effective in the management in mental disorders. Few other countries provided compensation apart from Sweden. In the USA North Carolina offered $20,000 for persons still alive to claim by June 2014, and Virginia compensated in 2015. Of Alberta and British Columbia as the two Canadian provinces where sterilizations took place, compensation claims have only succeeded in the Alberta courts. There were doctors who simply went ahead to sterilize irrespective of legislation. In Finland there were a high number of sterilizations for which authorization under the then law was not obtained. In Switzerland there were cantons where sterilizations were carried out without any legislative framework. Sterilizations for transgender operations in Sweden have now ceased, and victims will be compensated. One can see how professionals have been interventive in bodies of persons deemed “unfit” and of lesser value in economic and reproductive terms. Reproductive health is an area of major medical advance, but there can be sinister abuses. It is a history that is open-ended and unfinished (Dowbiggin 2008).

Newgenics and Resources Whether eugenics has ended has been much debated. The name “eugenics” was eventually dropped by societies in the UK and USA. Kevles (1985) argued eugenics ended discredited by the defeat of Nazism, whereas Diane Paul (1995) sees continuities to the screening for genetic diseases and in wholesale disease prevention programs. Genetic counselling has been criticized as to the extent that it is professionally directed. Molecular biology has been controversial, as regards implications of discoveries in the biochemistry of the gene on to the human genome project and genetic engineering schemes. Wider definitions of “newgenics” include prenatal implantation and elimination of genetic defects. The acceptability of the technique varies, with a marked contrast between positive acceptance in the UK and higher rates of rejection in Germany.

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Historical perspectives inform the public on ethical issues. The Oxford Handbook of the History of Eugenics (Bashford and Levine 2010) is a tour de force in terms of global coverage. It indicates how eugenics touched most countries of the world and variety of forms from imperialism to decolonization. The handbook covers the Near East with interwar Turkey and Iraq and pulls together significant coverage on the Far East, especially China and Japan. The Human Genome Project in the USA built in a historical component as part of its ethics dimension (Kevles 1992). In the USA Cold Spring Harbor has pioneered a website on the history of eugenics with select but significant documents. The Galton Institute in the UK has made available online sources on Galton and the Eugenics Review. The Wellcome Trust has a “Codebreakers” hereditary science documentary program http://wellcomelibrary. org/collections/digital-collections/makers-of-modern-genetics/ making available significant sources. The program was sadly not proactive in seeking to locate lost and missing archives of émigré eugenicists. Sterilization in Alberta was the focus of the Living Archives of Eugenics program which approaches issues from the perspectives of victims and informs a wider public: http://eugenicsarchive.ca/database/documents/ 5233c4395c2ec5000000008a. How invasive research impacted remains under-researched. A database on victims of Nazi coerced experiments covers victims of genetic and reproductive research (Weindling 2014). Germany has been slow to have online resources. Lutz Kaelber’s excellent site (based at University of Virginia) on the history of euthanasia is a significant resource. The Leopoldina German Academy of Sciences sponsors the digitization of the immense Ernst Haeckel papers indicating public penetration of evolutionary ideas. The balanced and useful sites were a response to the Catholic activist Eugenics Watch. This site makes available the membership listings of the US and UK eugenics societies. Scientologists have focused on Nazi psychiatry. Holocaust deniers have intervened in an attempt to discredit key historical memoirs, for example, that by the prisoner pathologist Miklós Nyiszli (2012) on Josef Mengele, ignoring the veracity of his first-hand observations. Eugenics has entered the historical mainstream. Its historiography shows a rich variety of methods and perspectives but uncritical oversimplifications. At first the historical mainstream marginalized eugenics, seeing it as no more than populist race prejudice and anti-Semitism; then interpretations swung the other way, and (equally uncritically) eugenics was everywhere from the welfare state to immigration controls. Gratifying as this for those who spotted the significance of eugenics when it appeared to be little more than scientized eccentrics and extremists, there remains the need for greater definition. One way forward is to figure in opponents of eugenics and race theory (Chesterton 1922; Hertz 1904; Iltis 2017). Galton diagnosed fundamental issues as regards the role of science-based expertise and the power of science to address changes in fertility and morbidity. The innovative power of science has sustained eugenics as a historical force. It is not just that the history of eugenics has been one of the most vigorous branches of the history of science, but immense opportunities for historical research remain.

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The Historiography of Genetics Michael R. Dietrich

Contents Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . History and Memory . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Creating Classical Genetics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . The Long History of the Gene Concept . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genes in Action: Development and Genetics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Better than Darwin: Evolutionary Genetics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Human Genetics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . National and Transnational Narratives . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Future Directions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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Abstract

Research on the history of genetics has followed trends in the history of biology more generally as it has moved from histories focused on ideas to more socially contextualized histories, including more geographically diverse histories and histories that center around gender and race, and then to histories that focus on scientific practice and material culture. As new historiographic foci have emerged, older approaches have not been replaced. There remains a strong tradition of considering research in genetics primarily in terms of its scientific development. Genetics is a rich and fast-moving field and there are still many stories to be told. Branching out from more familiar topics presents an opportunity for original research in the history of genetics.

M. R. Dietrich (*) Department of History and Philosophy of Science, University of Pittsburgh, Pittsburgh, PA, USA e-mail: [email protected] © Springer Nature Switzerland AG 2021 M. R. Dietrich et al. (eds.), Handbook of the Historiography of Biology, Historiographies of Science, https://doi.org/10.1007/978-3-319-90119-0_10

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Introduction While heredity has been studied scientifically for hundreds of years, genetics became established as a science in the twentieth century. If we take the so-called rediscovery of Mendel in 1900 as an origin point for this new science of genetics, then the field’s beginning is marked by the dispute between advocates of Mendelism with its emphasis on discrete traits and particulate inheritance and advocates of a biometrical approach using statistical methods to measure gradual change in continuous traits. Resolution of this dispute required attention to the role of statistics, the limits of generalization, and the phenomena of inheritance. At the same time, genetics was becoming an icon of experimental biology and applied biology. Early genetics rested on elaborate practices of controlled matings that promised precise understanding of hereditary patterns and ultimately control of those processes. Hereditary control was of special interest to plant and animal breeders and to those who would see similar systems of breeding extended to humans (see Paul Weindling’s ▶ Chap. 7, “The History and Historiography of Eugenics,” in this volume). These elements of quantification, experimentation, and practical application became defining features of genetics as it developed as a field. As historians have grappled with these different dimensions of genetics, they have produced histories that began with its ideas and methods, but quickly moved to the social, cultural, political, and economic implications of its applications. Although it is a relatively new science, the history of genetics is well established as an area of scholarship. Historiographically, research on the history of genetics has followed trends in the history of biology more generally as it has moved from histories focused on ideas to more socially contextualized histories, including more geographically diverse histories and histories that center around gender and race, and then to histories that focus on scientific practice and material culture. As new historiographic foci have emerged, older approaches have not been replaced. There remains a strong tradition of considering research in genetics primarily in terms of its scientific development. Genetics is a rich and fast-moving field and there are still many stories to be told. Branching out from more familiar topics presents an opportunity for original research in the history of genetics. The topics that I focus on here do not provide a comprehensive account of scholarly research in the history genetics. Instead, I have selected topics that highlight recent historical scholarship as well as topics that illustrate trends from early genetics to contemporary genomics. (“Mendel and Mendelism” are covered in Staffan Mueller-Wille’s ▶ Chap. 6, “Gregor Mendel and the History of Heredity,” in the volume and “Molecular Genetics” is discussed in Michel Morange’s ▶ Chap. 9, “The Historiography of Molecular Biology”). I begin with scientists’ own histories before moving into scholarship produced by historians.

History and Memory The first pass on the history of a science is often written by its participants. Genetics is no different. Centennial celebrations of Mendel’s paper on pea hybrids in the mid-1960s provided the occasion for systematic historical reflections on the field.

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Alfred H. Sturtevant, a leading geneticist, provided one of the first histories that included the famous rise of Drosophila as a research organism in his book, A History of Genetics (1964). Leslie C. Dunn’s A Short History of Genetics (1965) offered a more balanced survey of topics but did not extend beyond the first half of the twentieth century. Mendel was more directly commemorated in Royal Brink’s collection from the Mendel Centennial Symposium sponsored by the Genetics Society of America, Heritage from Mendel (1967). Curiously, a commemoration of the centennial of Mendel’s birth by the American Society of Naturalists in 1922 also included Francis Galton. Essays written by E. M. East, Arthur Harris, T. H. Morgan, and George Shull traced twin origins for their new field in an attempt to bridge the recent dispute between those who identified as Mendelians and those who advocated a biometrical approach to heredity (East 1923; Harris 1923; Morgan 1923; Shull 1923). As memorial reflections, these historical narratives sought to create a usable past: they offered historical narratives that were useful to contemporary biology. In the case of the 1922 centennial, they acknowledged a rift that threatened the founding of their new field and signaled its resolution by commemorating both Galton and Mendel. By the mid-1960s, the same need was not seen and Galton was allowed to fade as Mendel was given center stage. Such uses of science are common enough that historian Pnina Abir-Am has documented scientists’ uses of history to create and reinforce an image of their field, to rally organizational efforts within a field, and to legitimate contemporary science by invoking a historical lineage to some founding figure (Abir-Am 1999). In the case of Mendel’s centennial, Audra Wolfe has made an intriguing case for contextualizing this celebration in terms of Cold War geopolitics (Wolfe 2012). Scientists’ histories have not been limited to major milestones such as the Mendel Centennial. The pages of Genetics have seen hundreds of articles on the history of genetics. Early on many of these articles were biographical and often functioned as extended obituaries. In the 1980s, however, geneticist James F. Crow began editing these articles under the rubric of “perspectives on genetics” and contributed many of his own reflections on the field there. Under Crow, topical or thematic narratives became more common and historians began to contribute histories written for practicing geneticists. (See the collection of perspectives articles in Crow and Dove 2000.) None of this is to say that scientists’ histories are necessarily poor history. Scientists’ histories are often rich in technical detail and reflect personal experiences that may not be recorded in any other form. However, the goals of scientists’ histories may not be shared by historians who rarely write to legitimize contemporary science or to place themselves in a scientific lineage. Of course, every historian has an axe to grind, and it is to those agendas that we will now turn.

Creating Classical Genetics When the historians Garland Allen and William Provine began to write about the history of genetics in the early 1970s, they focused on the science that they took to define the basis for early or classical genetics (Allen 1978; Provine 1971). Over time

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subsequent scholars have revisited this early period to make a case for more sociologically oriented history, histories that engage with scientific practice, histories that include broader social and institutional elements, and just more expansive histories of genetics in the first decades of the twentieth century. Provine’s book, The Origins of Theoretical Population Genetics, placed the origins of genetics in the context of evolutionary biology. Charles Darwin’s theory of evolution by natural selection famously lacked a theory of heredity that was essential for explaining how traits persisted across generations. Followers of Darwin understood that heredity should allow for slow gradual change over generations that would not allow new variations to get “swamped” by the prevailing trait. Darwin’s own theory of pangenesis postulated particles, pangenes, that determined the fate of different body parts and were collected and transmitted in the gametes. Modifying pangenes during an organism’s lifetime became a way of ensuring that variants were introduced frequently enough to accumulate. Darwin’s cousin, Francis Galton, famously refuted the existence of pangenes but supported continuous inheritance by developing statistical tools for precisely describing the similarities between characters. Using correlation and regression, Galton reconsidered heredity from a statistical point of view. Because characters were continuous, Galton believed that their distribution in a population was best described by a statistical distribution. The effects of selection were reconsidered in terms of effects on that distribution’s mean and variance. Selection could move the mean of a population over a number of generations to create a new characteristic population mean. The relationship between parent and offspring was presented in terms of a “law of ancestral heredity” where a particular character of an offspring can be determined from the diminishing physical contribution of its ancestors. Galton’s Natural Inheritance (1889) inspired Karl Pearson and W. F. R. Weldon to develop a statistical approach to biology and evolution that they called biometrics. Within the biometrical tradition, Weldon and others applied statistical methods in order to support gradual Darwinian evolution by natural selection. Weldon and others collected statistical evidence from various traits which were thought to demonstrate the effect of selection in reducing population variability. These efforts convinced the biometricians that statistical methods were essential for understanding heredity and evolution. William Bateson had also been impressed with Galton’s work, but was not convinced that statistical methods were the best tools or that either evolution or heredity should be understood as continuous or blending. In 1894, Bateson argued in his book, Materials for the Study of variation with Special Regard to Discontinuity in the Origin of Species, that discontinuous variations were common and saltational evolution of new species was probably the norm. The dispute between Bateson and the Biometricians began with Weldon’s hostile review of his book. It was transformed into the Mendelian–Biometrician controversy when Bateson read Mendel’s paper on a train ride in 1900. Bateson translated Mendel’s paper into English and immediately began championing it as the key to heredity and evolution. As a result, Weldon and Pearson would debate the significance of Mendel’s paper vociferously over the next 10 years.

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Bateson’s Mendelism emphasized discrete pairs of traits that left no room for statistical distributions. These pairs of traits were associated with pairs of cellular particles that obeyed what Carl Corren’s had labelled as Mendel’s laws. Abrupt changes from one form to another ran counter to Darwinian gradualism and seemed to support its opposite, a saltational view of evolution driven by what Hugo de Vires called mutations. In Provine’s narrative, the controversy that spun out from these two positions thoroughly entangled genetics with the study of evolution and the creation of new statistical methods and modes of quantification. Most importantly, this founding rift in genetics was healed by the creation of population genetics and its new mathematical theories of how populations vary and change over time. For Provine, the Mendelian-biometrician controversy was resolved when R. A. Fisher, J. B. S. Haldane, and Sewall Wright demonstrated how Mendelian genetics and Darwinian evolution could be reconciled (Provine 1971). Where Provine found the controversy’s resolution in population genetics, Nils Roll-Hansen located it in Wilhelm Johannsen’s distinction between the phenotype and genotype (Roll-Hansen 1980; see also Falk 2008). Such emphasis on the scientific grounds for disagreement between the major players in the MendelianBiometrician controversy was not shared by more sociologically oriented scholars of the dispute. Donald MacKenzie and Barry Barnes argued that framing the controversy in terms of an intellectual disagreement that was rationally resolved would not do. Instead, they invoked symmetry to argue that consideration of a full range of sociological differences should be considered as an irrationalist counter-explanation (MacKenzie and Barnes 1975; Shapin 1982). The ensuing debate became somewhat methodological as the sociologically inclined resisted accusations of social determinism in favor of a standard of association, not causation, between social structures and knowledge or scientific position in the controversy. Bateson’s wariness of eugenics and Pearson’s advocacy of eugenics became important points of difference as a result, casting the dispute in wider terms. As Robert Olby notes, one important consequence of these different approaches to the controversy was to recognize the importance of social history that went beyond intellectual factors in the origins, persistence, and resolution of this foundational dispute (Olby 1989). Because the Biometricians and Mendelians drew on extended networks of biologists during the course of this dispute, historian Kyung-Man Kim argues that the controversy was resolved by all of the members of this extended network, not by just the principle antagonists who remained strongly polarized (Kim 1994). A. D. Darbishire, for instance, set out to refute Mendelism with a set of experiments on albino and waltzing mice. Bateson wrote devastating critiques reinterpreting Darbishire’s results in Mendelian terms. Darbishire himself was convinced when he tested his hybrids and realized that statistical analysis of external appearance was not a reliable guide to genetic constitution. Darbishire’s defection infuriated Pearson, but this was one of several shifts that altered the course of the debate (Kim 1994; Ankeny 2000). Pearson, Bateson, and Weldon have all been subject to renewed historical inquiry as founding figures of British genetics (Cock and Forsdyke 2008; Magnello 2004; Radick 2015). Theodore Porter has written a far reaching biography of Pearson

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(Porter 2004), while Gregory Radick and Marsha Richmond have been investigating Bateson’s research group (Radick 2013; Richmond 2001, 2006). Charles Pence (2011) has reconsidered the roots of Pearson’s and Weldon’s commitments to statistical methods, and Radick has been reimagining the foundations of this dispute and their consequences for a gene-centric science of heredity (Radick 2005). James Schwartz contributed a well-written popular account of the pursuit of the gene from Darwin to DNA (Schwartz, 2008). More scholarship in this area is in the wings at the time of this writing. Garland Allen took a very different approach to the first decades of genetics. By approaching genetics through a biography of Thomas Hunt Morgan, Allen tied genetics to on-going debates in developmental biology as well as debates over the place of heredity in evolution. Morgan’s winning of the Nobel Prize in 1933 for his advocacy of the chromosomal theory of inheritance and development of Drosophila as an unparalleled genetic tool makes him a natural choice for a founding figure. Allen’s biography traces Morgan from his training and early work in embryology through his early interest in chromosome cytology to the creation of a major American research group which demonstrated that genes were pieces of chromosomes that could be mapped, traced across generations, and mutated to create new traits. Although Morgan’s “Fly Room” was filled with some of the most influential geneticists of the twentieth century, curiously only Herman J. Muller has been subject to a full biography (Carlson 1981). Alfred Sturtevant and Calvin Bridges certainly deserve more consideration. The gendered division of labor in Drosophila genetics has been documented (Dietrich and Tambasco 2007), but important and influential centers of Drosophila research beyond Morgan’s remain unconsidered. When the turn to practice in the history of genetics hit its stride in the 1990s, Robert Kohler revisited the Fly Room in his influential book, Lords of the Fly (1994). Kohler reimagines the rise of Drosophila genetics as the creation of a new biotechnological system where the Fly Room becomes a “breeder reactor” that produces the mutations that make fly genetics possible, while Morgan institutes a moral economy that regulates the group’s work ethos. Importantly, Kohler also follows the trajectory of Drosophila genetics beyond its beginnings in the chromosomal theory of inheritance to trace its members’ involvement in evolutionary genetics and developmental genetics, which we will discuss below. Kohler’s organismal focus set an agenda for further work such as Karen Rader’s important history of the development of mouse genetics at the Jackson Laboratories (Rader 1998, 2004) and Rachel Ankeny’s history of the rise of C. elegans as an experimental system (Ankeny 2001). For an extended analysis, see Ankeny and Leonelli’s ▶ Chap. 13, “Organisms in Experimental Research,” in this volume. Social, institutional, and economic histories present a different take on the early decades of genetics. Rather than trace the refinement of ideas or methods, many historians have questioned the conditions that influenced the reception of Mendelism, especially the close ties of genetics, first to agriculture and then to biotechnology (Palladino 1994; Kimmelman 2006, see Berry’s ▶ Chap. 22, “Historiography of Plant Breeding and Agriculture,” in this volume and Crowe’s ▶ Chap. 11, “The Historiography of Biotechnology,” in this volume). Diane Paul and Barbara

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Kimmelman’s important essay on this question frames the positive reception of Mendelism in the United States in terms of the nature of federal support for agriculture and the success of hybrid corn both in scientific and economic terms (Paul and Kimmelman 1988; see also Fitzgerald 1990). Garland Allen expands on these themes to link the reception of Mendelism in the United States to both a commitment to a kind of mechanistic materialism and to scientific, industrialized agriculture (Allen 2000). In contrast to these social histories, Roll-Hansen and others have claimed that Mendelian genetics offered methodological and technological innovations for breeders that contributed to its success (Roll-Hansen 2000). More recently, this challenge of explaining the usefulness of Mendelism to breeders has been addressed by Berris Charnley and Gregory Radick (2013), while the success of Mendelism, its relation to agriculture, and to the move toward more scientific agriculture have been addressed by Jon Harwood in a chapter in Denise Philips and Sharon Kingsland’s far-reaching collection on New Perspectives on the History of Life Sciences and Agriculture (Harwood 2015; Phillips and Kingsland 2015). Given how interrelated genetics and agriculture have been, it is striking that plants in general have not been given more attention. This is not to say that notable plant geneticists have been overlooked. Nobel laureates such as Barbara McClintock have been subjects of several historians beginning with Evelyn Fox Keller’s biography in 1983 which cast her as an enigmatic feminist savant (Keller 1983) continuing through Nathaniel Comfort’s biography which sought to unravel the mythmaking around McClintock (Comfort 2001). Lee Kass continues to carefully explicate McClintock’s science (Kass 2013), but in general maize genetics has not received the treatment that fruit fly genetics, mouse genetics, or C. elegans has received (see Ankeny and Leonelli’s ▶ Chap. 13, “Organisms in Experimental Research,” in this volume). Betty Smocovitis’s research on the development of Crepis as an experimental system and Sabina Leonelli’s work on the history of Arabidopsis stand out as exceptional studies of plant systems (Smocovitis 2009; Leonelli 2007). Indeed, Smocovitis’ work on Ledyard Stebbins and Masou Kodani stand as models of what histories of individual plant geneticists have to offer (Smocovitis 2000, 2011). While there has been some outstanding work on individual figures and certain developments, plant genetics deserves more synthetic history such as that undertaken in Helen Curry’s Evolution Made to Order (Curry 2016). Curry effectively extends the history of plant breeding beyond its early efforts by examining how technological tools such as x-ray radiation and the mutagen colchicine allowed unprecedented genetic manipulation and control. Curry’s combination of a wider chronological scope and integration of history of genetics and history of technology make it a valuable new contribution.

The Long History of the Gene Concept The gene championed by Thomas Hunt Morgan and his colleagues in the early twentieth century offered a powerful unifying object for genetics as a new discipline. Morgan’s own writing on The Theory of the Gene endowed it with explanatory

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power (Morgan 1926). Biologist Elof A. Carlson recognized the central role of the gene in classical genetics and in 1966 published a critical history of the gene just in time for Mendel’s centennial celebrations (Carlson 1966). Subsequently, the gene has become its own historical object with historians and especially philosophers of biology elaborating its conceptual history. Carlson’s history traces the articulation of Morgan’s theory of the gene and its material basis on the chromosome, but also documents the fragmentation and demise of what would later be called the classical or particulate gene. Focusing on ideas and experiments, Carlson describes how the discovery of position effects undercut the idea that the gene was an autonomous entity, like a bead on a string. Instead, Sturtevant’s discovery that a gene’s position relative to its neighbors meant that it was not a neatly isolated unit of structure and function. Doubts about the gene were amplified by German geneticist Richard Goldschmidt, a critic of Morgan’s style of genetics, who declared that “the gene was as dead as a dodo!” (Allen 1974). In its place Goldschmidt proposed that all mutations were in fact chromosomal rearrangements and that those rearrangements altered a hierarchy of genetic structures with accompanying consequences for function. This attention to chromosomal rearrangement around the time of the Second World War inspired careful mapping of rearrangements that led to the left-right test and then to ideas of genetic complementation that allowed fine mapping of the limits of units of structure and function on the chromosome. Issues raised in Carlson’s narrative were pursued by historians such as Garland Allan who wrote about the tension between Morgan’s group and Goldschmidt (Allen 1974). Later, Dietrich reconstructed Goldschmidt’s alternative vision of genetic units (Dietrich 2000a, 2008) and Holmes and Summers offered a detailed history of how fine mapping was used by Seymour Benzer to rethink the nature of the gene and mutation (Holmes and Summers 2006). Philosophers began to grapple with the gene as it appeared in arguments concerning the relationship between the classical gene and the molecular gene and the possibility that Mendelian genetics could be reduced to molecular genetics (Schaffner 1976; Hull 1979). These accounts did not tend to rely on careful historical reconstructions of gene concepts as much as philosophical distinctions regarding reduction. That said, Peter Beurton, Raphael Falk, and Hans-Jörg Rheinberger’s 2000 collection The Concept of the Gene in Development and Evolution: Historical and Epistemological Perspectives sought to bridge the gap between historical and philosophical approaches. In a similar vein, Evelyn Fox Keller’s The Century of the Gene, also published in 2000, followed the gene concept’s history in order to critique contemporary genetics and its role furthering ideas of determinism and evading complex problems of regulation, organization, and development in biology. Recently, a new appraisal of his discoveries and ideas – including “nuclein” as constituting a kind of alphabet producing hereditary variation based on differential stereochemistry – have positioned the nineteenth century Swiss physician and biologist Friedrich Miescher as an important precursor to twentieth century work on DNA (Veigl et al. 2020). Post Watson and Crick and “central dogma” developments in molecular genetics have driven historical reappraisals of the gene as

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scholars trace how biologists’ gene concepts have tracked new discoveries (Portin 1993; Burian and Zallen 2009; Griffiths and Stotz 2014; Kampourakis 2017). Discoveries that eukaryotic genes were not continuous, but were divided into introns and exons, led to discoveries of alternative splicing schemes that allow several proteins to be produced from a single DNA transcript (Morange 2000b). Genomic gene concepts have the challenge of trying to encompass these new kinds of phenomena which radically diverge from earlier structure-function accounts of the gene (Griffiths and Stotz 2014).

Genes in Action: Development and Genetics Thomas Hunt Morgan famously decided not to address the question of how genes actually produce traits. As someone trained in embryology, Morgan thought that researching how genes acted in development was not going to be as productive as researching patterns of gene transmission from generation to generation (Morgan 1934). Many geneticists outside of the United States did not agree, and traditions of physiological and developmental genetics grew, especially in Germany (Harwood 1993). Although Morgan’s views were historically contextualized by Garland Allen in 1978 (Allen 1978), they remain a topic of conversation in the historical literature (see Maienschein 2016; Allen 2016). In 1987, Jan Sapp’s Beyond the Gene seriously considered traditions of genetics beyond Morgan’s. Building on an analytic framework offered by Pierre Bourdieu, Sapp casts debates over cytoplasmic inheritance as a struggle for authority to define the central questions of genetics. Morgan’s gene centric approach was in this view a means of delimiting legitimate topics for genetic research, and cytoplasmic elements or developmental processes that complicated the connection between gene and trait were set aside. Robert Kohler has argued pointedly that Sapp’s interpretation, as well as Garland Allan’s interpretation (Allen 1986), portray Morgan’s Drosophila network as “a ‘hegemonic’ establishment, from which genetics had to be liberated by people with a broader biological outlook” (Kohler 1994). Kohler sees Morgan’s choice to focus on transmission problems as constrained by the success of his experimental system. The challenge of creating an equally successful system addressing problems in developmental genetics was a legitimate barrier. Nevertheless, Morgan’s decision did not even fully constrain his own group as members such as Jack Schultz and George Beadle pursued questions bearing on gene action (Kohler 1994; Berg and Singer 2005) Setting aside Morgan, there is a rich history of geneticists who have grappled with gene action, developmental genetics and genetic regulation (Burian et al. 1994; Davis et al. 2009; Galperin 1998; Gilbert 1988, 1998; Keller 2014; Laubichler and Rheinberger 2004; Maas 2001; Rheinberger 2000; Sapp 1983, 1987). Scott Gilbert’s many articles on development and genetics represent the most consistent voice on this topic. As both a developmental biologist and historian, he was first to place genetics in an embryological context (Gilbert 1978) and has consistently argued for the historical recognition of research on physiological and developmental genetics

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(Gilbert 1988, 1991, 1998). Unfortunately, some biologists’ histories of developmental genetics tend to emphasize stagnation and speculation until the insights of molecular genetics makes progress possible in the 1970s and 1980s (Gilbert et al. 1996; Lewis 1994; Love and Raff 2003). Such a view discounts the work of notable geneticists, such as Sewall Wright and L. C. Dunn in the United States and Alfred Kühn and Richard Goldschmidt in Germany (Deichmann 2011; Dietrich 2000a, b; Rheinberger 2000; Richmond 2007). Indeed, Nobel Prize winning work by George Beadle and Edward Tatum, Francois Jacob and Jacques Monod, and Edward Lewis all spoke to significant issues of gene action and regulation in adult and developing organisms before the molecularization of genetics (Berg and Singer 2005; Morange 2000a, b, 2020).

Better than Darwin: Evolutionary Genetics Evolution and genetics have been entangled from the start. As a result, histories of modern evolutionary biology, especially histories of the evolutionary synthesis, have addressed the development of genetics (Provine 1988; Smocovitis 1996; Cain 2002). Given his important role in twentieth century evolutionary biology, many historians have focused on the work of Theodosius Dobzhansky. Although no biography of Dobzhansky has been written, Mark Adams’ 1994 collection, The Evolution of Theodosius Dobzhansky : Essays on His Life and Thought in Russia and America, offers multiple historical perspectives that describe the arc of Dobzhansky’s life, the Russian tradition of population biology in which he was trained, his genetics training in Morgan’s Fly Room, and his incredibly influential research program. Earlier reprintings of Dobzhansky’s series of papers on the genetics of natural populations included important historical assessments by William Provine and Richard Lewontin (Dobzhansky 1981). Betty Smocovitis’ account of the history of evolutionary synthesis still offers one of the best accounts of Dobzhansky’s role in setting a program for evolutionary biology (Smocovitis 1996). Dobzhansky’s views on drift, selection, and genetic variability are expertly contextualized by John Beatty in papers following Dobzhansky’s genetic research in the postwar period (Beatty 1987a, b). Dobzhansky’s interests in the biology of race have also been carefully considered by Lisa Ganett (Gannett 2013). Finally, Dobzhansky appreciated the importance of population size for different evolutionary processes. Because he thought the Brazilian rain forests would harbor a very large population of Drosophila, he conducted field work in Brazil where he had a tremendous impact on the development of genetics (Araújo 2001; Pavan and Brito da Cunha 2003; De Carvalho 2020). Following Provine, a consistent thread in the history of evolutionary genetics has been an emphasis on controversy (Provine 1989). Dietrich provides a brief overview of the history of evolutionary genetics in terms of controversies starting with the Biometricians and Mendelians, the founders of population genetics, and then moving to the classical-balance controversy and the neutralist-selectionist controversies (Dietrich 2006). As mentioned above, Provine rooted the resolution of the

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Biometrician-Mendelian controversy in the rise of population genetics as formulated by its principle founders, R. A. Fisher, J. B. S. Haldane and Sewall Wright (Provine 1971). Fisher, Wright, and Haldane approached evolution and population genetics from different training backgrounds, different mathematical perspectives, and different assumptions about evolution (Sarkar 1992; Gayon 1998, on Fisher see Box 1978; Dietrich and Skipper 2013, on Wright see Provine 1986, on Haldane see Subramanian 2020; Dronamraju 2017). Fisher and Wright were engaged in a series of disputes from 1929 until 1962 when Fisher died (Provine 1986). While they debated many things, the core of their difference lay in their general theories of evolution: Wright’s shifting balance theory appealed to a range of evolutionary processes and emphasized population subdivision while Fisher’s large population approach placed its emphasis on natural selection (Provine 1992; Skipper 2002; Grodwohl 2017a, b). The relative importance of genetic drift was a flash point in this dispute and remained so in evolutionary genetics despite efforts to find some empirical resolution (Millstein 2008). While histories of these disputes in population genetics have been soundly intellectual in their orientation, histories of the classical-balance dispute have grounded it firmly in its Cold War context (Beatty 1987a, b). The classical and balance positions were at once stances on the nature of genetic variation, the way the selection acted on that variation, the ways atomic radiation produced mutation, and eugenic ideals for human variability. On the balance side, Dobzhansky advocated for high levels of genetic variability and a new form of selection that maintained it. On the classical side, Herman Muller argued for low genetic variability, a more traditional role of selection (Dietrich 1994). As the person who received a Nobel Prize for demonstrating that radiation caused mutation, Muller was deeply concerned that radiation was contributing to an accumulation of harmful mutations that he dubbed “our mutational load” (Carlson 1981; Paul 1987). In addition to carefully describing the dynamics of this controversy, John Beatty contextualizes it in terms of both research on radiation genetics and increasing social and political concern regarding radiation exposure and atomic energy policies (Beatty 1991, 1993). Susan Lindee’s history of the Atomic Bomb Casualty Commission delves even deeper into this sociopolitical context describing the ABCC efforts as a colonialist form of science imposed by occupying US forces in Japan (Lindee 1994). More recently, Luis Campos has offered a longer view on the relationship between radiation and life providing a trajectory from nineteenth century radium research to Muller’s own transformation of the gene (Campos 2015).

Human Genetics One of the major challenges facing historians of human genetics is the effort to disentangle it from eugenics origins. Virtually every biologist who identified as a geneticist in the early twentieth century had an interest in eugenics. Too often, however, historians treat eugenics in isolation from genetics or they see genetics as a source of legitimacy for eugenics. Just as agriculture helped bolster genetics as a

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science, eugenics also legitimated genetics to the public (see Weindling’s ▶ Chap. 7, “The History and Historiography of Eugenics,” in this volume). The interplay between eugenics and human genetics has been addressed by a number of historians (Fortun and Mendelsohn 1999; Weiss 2006). Nathaniel Comfort’s The Science of Human Perfection (2012) provides a compelling narrative of how geneticists have been driven by a desire for human improvement from Galton to the human genome project. Attempts to claim that human genetics somehow was legitimated around the time of WWII do capture changes in the human genetics as it professionalized, but the eugenic impulse was not abandoned even if it was expressed in more coded terms in the postwar period (Duster 1990). In the postwar period, there was an attempt to distance human genetics from its deeply racist origins (Barkan 1992; Gormley 2009). However, as histories of race and race mixing research show, explaining why human geneticists moved away from some racist views is not as simple as appealing to revulsion with Nazi racial science (Provine 1973; Paul 1998; Caballero and Aspinall 2018). Moreover, whatever changes occurred as geneticists adopted new attitudes toward race and endorsed new statements through their professional societies, racist science re-emerged in terms of the race and IQ debates of the 1970s and continues to have its advocates in the genomic era (Panofsky 2014; Reardon 2005). These later histories are much more socially contextualized than some earlier approaches. Panofsky, for instance, diagnoses the persistent controversy in behavioral genetics not merely as a feature of their involvement in debates over race and IQ, but in terms of struggles for authority, legitimacy, and autonomy that ultimately produced a fractured field with an archipelagic structure. The medicalization of human genetics around the time of WWII involved a complex shift toward creating a category of genetic disease. Diane Paul and Jeffrey Brosco’s research on PKU dives deeply into the PKU paradox, why a relatively rare disease became something that was tested on a mass scale (Paul and Brosco 2013). Mary Mitchell takes a more institutional perspective when she describes the work of the Social Issues Committee of the American Society of Human Genetics as they address PKU testing in the context of American abortion politics (Mitchell 2017). Troy Duster’s classic Backdoor to Eugenics (1990) also addresses the history of genetic testing and especially the social stigma and racial politics that has surrounded such efforts. Alexandra Stern (2012) reconstructs the rise of genetic counseling as a discipline that accompanied these tests. Susan Lindee’s Moments of Truth in Genetic Medicine (2008) offers a broader range as it discusses early human twin research, Viktor McKusick’s research with Amish communities, and research and treatment of familial dysautonomia. While the history of any genetic disease could be told from the perspective of the researchers, Lindee offers an account that gives voice to both subjects and scientists. In doing so, she argues that scientific authority was distributed, and the recognition of scientific reality was not the purview of just geneticists but was shared by researchers, subjects, families, and communities. Counting chromosomes has been a task for geneticists since 1902. Counting human chromosomes, however, was notoriously difficult until the 1950s. The challenges of early human cytogenetics have been recounted by a number of historians (Kottler 1974; Martin 2004; Chadarevian 2015, 2018, 2020; Lima-de-

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Faria 2003; Santesmases 2017). While Maria Stantesmases contextualizes human chromosome counts in terms of cancer biology, Soraya de Chadarevian pushes back against narratives about the preeminence of molecular biology, arguing that advances in human cytogenetics in the 1960s and 1970s were as important as molecularization for fostering projects such as the human genome project of the 1990s. It is important to recognize that cytogenetics extends well beyond humans and was a crucially important field of research. Oren Harman’s biography of Cyril Darlington captures a significant portion of this larger view on chromosome research, even if this account is naturally focused on Darlington and his circle (Harman 2004). While Darlington’s importance and influence should not be underappreciated, there are many developments in cytogenetics that deserve further research (Smocovitis 2011).

National and Transnational Narratives Genetics has never been limited to the English-speaking world even if its history in those nations has been told in greater detail. Because of its connection to centuries old breeding practices as well as the rise of scientific agriculture and eugenics, genetics spread around the world fairly quickly. Historians have addressed this geographic dimension of genetics with histories of distinct national styles of research, histories of the institutionalization of genetics in different countries, and in histories that take a comparative and transnational perspective. To mark the 100th anniversary of the American Society of Zoologists in 1989, Ron Rainger, Keith Benson, and Jane Maienschein brought together two collections of essays on the rise of American biology, The American Development of Biology (1988) and The Expansion of American Biology (1991). Drawing together portraits of American biologists, these volumes introduced a cast of characters and placed them within the institutional and social context of developing twentieth century American science. Jane Maienschein’s 1991 book, Transforming Traditions in American Biology, 1880–1915, expanded on this theme of nationally distinct approaches to the history of biology by focused on a founding generation of American biologists including Edmund Beecher Wilson, Edward Grant Conklin, Thomas Hunt Morgan, and Ross Harrison. Maienschein describes the roots of a kind of American distinctiveness in biology as they charted a path away from earlier German traditions of morphology within a nation eagerly expanding its commitment to education and scientific expertise. Morgan’s genetics was part of this national trend. The idea of national styles has been most fully articulated in Jon Hardwood’s 1993 book on German genetics, Styles of Scientific Thought: The German Genetics Community, 1900–1933 (Harwood 1993). Based on an analysis of every academic geneticist in pre-Nazi Germany, Harwood proposes two major styles of thought: the comprehensive and the pragmatic. As a result of the emphases of its dominant educational tradition, Harwood argued that most German biologists were comprehensives, while most Americans held the more practically oriented pragmatic style.

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That said, Harwood’s analysis focused on the German community leaving an as yet unfilled invitation to analyze the American genetics community in similar terms. Histories of genetics in different national contexts are becoming much more common with specific accounts of genetics in France, Mexico, Japan, the Soviet Union, and elsewhere (Araujo 2001; Pinar 2002). In a series of articles, Richard Burian, Jean Gayon, and Doris Zallen undertook a systemic examination of the development of genetics in France (Burian et al. 1988; Burian and Gayon 1999; Gayon and Burian 2000; Gayon and Burian 2004; Gayon and Zallen 1998; Zallen and Burian 1992). From its roots in French seed companies through the development of regulatory ideas of genetics, these articles develop a portrait of a national tradition of genetics that doesn’t make claims regarding styles of thought per se but does reflect the influence of French institutions and culture (Bonneuil 2006). Ana Barahona’s writing on the origins of genetics in Mexico develops similar connections to agriculture, as does Kaori Iida’s history of genetics in Japan (Barahona 2013; Barahona and Ayala 2005; Iida 2009, 2015b). Barahona’s and Iida’s histories, as well as Nurit Kirsch’s work on early genetics in Palestine/Israel (Kirsh 2004), also feature influential founders as essential for creating the institutional space and support that allowed genetics to flourish in these different countries. The history of Soviet genetics, by contrast, has been less about developing traditions of research (although see Gaissinovitch and Adams 1980; Graham 1990; Cohen 1991) and more about the impact of the controversial agronomist Trofim Lysenko. Casting his views in opposition to the Mendel-Morganism of the West, Lysenko promoted the vernalization of wheat as a means to increase yields. Several historians have discussed how acceptance of Lamarckian inheritance of acquired characteristics and opposition to Western genetics fed into the Cold War dynamics (Joravsky 1986; Roll-Hansen 2006; Graham 2016). Because of the extensive response by non-Soviet geneticists, the Lysenko affair became one of the first instances of comparative history of genetics in the literature (DeJong-Lambert and Krementsov 2017; Iida 2015a). Of course, many national histories involve comparisons to other national practices of genetics, but the move to transnational history involves focusing more on the circulation of people, texts, organisms, and tools. For example, Lisa Onaga’s research on silkworm genetics and Dietrich’s research on balance theories of sex both focus on systems of exchange and circulation across national boundaries where genetics interpretations are contingent and contested (Onaga 2010; Dietrich 2016, see Barahona’s ▶ Chap. 17, “Local, Global, and Transnational Perspectives on the History of Biology,” in this volume).

Future Directions Genetics owes part of its authority to popular and scientific beliefs in genetic determinism and biological essentialism (Nelkin and Lindee 1995; Lewontin 1991). Many scholars have engaged in critiques of genetic determinism, but it is time for careful histories of the debates over the different dimensions of genetic determinism, especially in light of its implications for essentialist views that

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supported discrimination in a range of forms. Aaron Panofsky’s insightful analysis of the history and organization of behavioral genetics offers one model of how this topic can be approached via analysis of the characteristics of a field where such issues were actively and publicly debated (Panofsky 2014). James Tabery’s 2014 book on the nature-nuture divide offers another (Tabery 2014; Griffiths and Tabery 2008). Focused histories of individual actors, such as biologist Richard Lewontin, would complement field-based approaches, on the one hand. On the other, because genetic determinism has been such a multidisciplinary topic, grappling with perspectives from geneticists, philosophers, pundits, and educators will be essential to contextualize this history.

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The Historiography of Molecular Biology Michel Morange

Contents Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A Renewed Historiography of the Classic Period of Molecular Biology . . . . . . . . . . . . . . . . . . . . . . The Exploration of New Fields . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . The Long 1970s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . The 1980s: An Explosion of Discoveries . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . The 1990s: The Rise of Genomics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Entering the Post-genomic World . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . The Need for longue durée Studies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

175 177 180 180 183 184 186 187 189 190

Abstract

This chapter reviews the historiography of molecular biology from its early “classical period” through 1970s and 1980s into the genomic era beginning in the 1990s. I emphasize the need to find connections between the “classical period” and more contemporary genomic and post-genomic histories. I argue that synthestic and longue durree histories are essential for capturing the trends embodied in molecular biology research.

Introduction In 1980, Horace F. Judson published an article entitled Reflections on the historiography of molecular biology (Judson 1980), just 1 year after the publication of his history of molecular biology, The Eighth Day of Creation (Judson 1979). He critiqued the rare historical studies then available, in particular the descriptions M. Morange (*) IHPST, Université Paris I, Paris, France e-mail: [email protected] # Springer Nature Switzerland AG 2021 M. R. Dietrich et al. (eds.), Handbook of the Historiography of Biology, Historiographies of Science, https://doi.org/10.1007/978-3-319-90119-0_11

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by René Dubos of Oswald Avery’s personality and work (Dubos 1976) and Robert Olby’s The Path to the Double Helix (Olby 1974). Judson’s view was that all previous authors had been unable to acknowledge the major transformation wrought by molecular biology, i.e., the transition of a three-dimensional conception of specificity to a one-dimensional informational one. He reiterated the same idea 13 years later, in an article published in Gene (Judson 1993). Long gone is the time when the history of molecular biology could be reduced to one conceptual transformation! Anyone seeking to write the historiography of molecular biology faces a twofold problem. First, what should be included? Quite early, historians noticed that molecular biology did not have the acknowledged characteristics of a scientific discipline (Olby 1990). And the situation rapidly worsened: the rare signs of institutionalization of molecular biology disappeared and it was progressively diluted; it “evaporated” into the different biological disciplines that adopted its techniques and concepts (Powell et al. 2007; Rheinberger 2009). From the middle of the 1980s, a series of new disciplines emerged – genomics, systems biology, and synthetic biology – whose relations to molecular biology are unclear. The American historian Lily Kay preferred in 1993 to avoid the term discipline and to describe the emergence of a “molecular vision of life” (Kay 1993). More recently, the death of molecular biology was announced and was considered by some as good news after too many years of harsh reductionism (Morange 2008). This first problem – of what to include in a historiography – is probably not unique to molecular biology but is more obvious than for other branches of science. It is compounded by a second problem, which is shared by all historical studies of scientific developments: the increasing wedding of historical studies with philosophy on the one side and social studies on the other. The focus has shifted and more attention is paid to the description of experimental systems, models (and in particular animal models), and representations, but also to the social organization of the work and the wider context in which it is pursued. This diversification of studies is certainly a plus for historians but also has its flaws. Many studies are skewed toward a goal that differs from their own goal. For instance, a study may be designed to show the importance of one type of representation in science. Examples from molecular biology will support the hypothesis favored by the authors but will not be directly related to the history of molecular biology: the historical description is used for a purpose other than that of history. One example will illustrate this point. Exploratory experiments, besides hypothesis-driven experiments, are deemed to be characteristic of the scientific method, and their expansion through the new postgenomic technologies has been amply discussed by philosophers (Waters 2007). Richard Burian convincingly demonstrates the importance of these exploratory experiments for the characterization of small regulatory RNAs (Burian 2007). But will this study enable the historian to understand the significance of the search for regulatory RNAs in present-day biology? In addition, this can generate bias in the choice of subjects. Genome sequencing (in particular human genome sequencing) and post-genomics have been extensively studied because they represent a new organization of scientific research and a new way of funding it. The impact on the

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whole discipline was seen as obvious but was not really examined. This transformation of historical studies has not created bias where it did not exist before; rather, it has simply replaced some types of bias by others. Such biases create a rich source of study for future historians: some areas have yet to be examined, whereas others have only been explored by the scientists involved and await a historical critique. Given the difficulty of defining molecular biology plus the spread of historical information through various studies, I will distinguish different periods in the historiography of molecular biology. The first period is that of “classic” molecular biology, the topic of Judson’s historiographical account. This historiography has been considerably enriched since Judson’s time by the description of work that had not attracted the attention of historians before and by new light cast on some wellknown episodes. Authors refrain from examining the effects of this work on the rise of molecular biology and instead look at the context in which it was done. It is a virtue of historical studies that ground already covered can be reexamined by shifting the focus! The second period I shall consider covers the investigations conducted by molecular biologists between the end of the 1960s and today. I will organize it in four sections: the “long 1970s,” the 1980s and the rapid accumulation of data generated by the tools of genetic engineering, the rise of genomics, and, lastly, the post-genomics era. I shall then address issues only briefly considered in the preceding parts and which can only be elucidated by historical studies of longer periods. Examples are the slowly progressing field of gene therapy and changing conceptions regarding the mechanisms of aging, which are rich subjects for future historical studies. In conclusion, I will justify my conviction that current post-genomic studies can be placed under the banner of molecular biology and that long-term studies are needed.

A Renewed Historiography of the Classic Period of Molecular Biology Historians have at their disposal new sources, such as the autobiographies of important, but hitherto silent scientists – that of “the third man of the double helix” is an emblematic case (Maurice Wilkins: Wilkins 2003) – and letters that had disappeared and miraculously reemerged, such as those of Francis Crick (Gann and Witkowski 2010). These documents have added details, but one must admit that they have not revolutionized our vision of the race to the double helix. Even Robert Olby’s richly documented biography of Francis Crick did not open up new avenues to historians, although it is a source of reliable information for further studies (Olby 2009). The same is true of the remarkably precise historical studies of Frederic Larry Holmes (2001; Holmes 2006). Nevertheless, the use of quantitative methods allows to revisit the “impact” of some of these famous discoveries, for instance, the discovery of the DNA structure (Gingras 2010). The frantic search for novelty and scoops may irritate when authors claim to reappraise the contributions of scientists but in fact simply shift the emphasis from

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one interpretation to another, without adding new material for future historians (see, for instance, Stegenga 2011). But these new studies are not always so disappointing. A serious biography of Rosalind Franklin was welcome (Maddox 2003). And it is useful to have a precise and annotated English translation of the first study of the gene by Max Delbrück (Sloan and Fogel 2011), in part because of the attention drawn to it by Erwin Schrödinger’s What is Life? (Schrödinger 1944). By shedding new light on important episodes, authors can escape the burden of previous interpretations and provide a vivid and renewed description of well-known events (Cobb 2015). Biographies and autobiographies can also reveal additional interactions between the protagonists – the exchanges between William Astbury, the first scientist to produce X-ray diffraction patterns of DNA, and Oswald Avery – that show how the circulation of information between the founders of molecular biology was much more extensive than is generally admitted (Hall 2014). It is always possible to reinterpret well-known historical descriptions. Michel Foucault used to say that the best material for the historian is what resists his interpretation of events. Time favors the historian: if historical facts are, for a long period, hard to fit into the interpretative framework imposed by historians, then a reinterpretation is probably needed. One example will help to illustrate how it is possible to put new interpretative wine into old bottles! One of the first results of molecular biology was to establish a relation between genes and enzymes, the famous one gene-one enzyme (protein) relation, a favorable niche for the elaboration of the Central Dogma, and the decipherment of the genetic code. This simple presentation raises at least three difficult historical issues. The first is that the existence of this relation was already widely admitted, and the importance of the discovery appears disputable. The second difficulty arises from the two-step establishment of this relation. In a first step, George Beadle and Boris Ephrussi tried to characterize the role of genes in Drosophila development by studying two mutations affecting eye color. The second step was accomplished by Beadle and Edward Tatum on Neurospora by studying the genetic control of well-described metabolic pathways. The development of a new experimental system may play a decisive role in the transformation of scientific knowledge. But what is puzzling in this case is that the project itself changed: whereas Beadle and Ephrussi tried to discover the mechanisms of gene action in development, Beadle and Tatum proposed a general mechanism valid for all genes. Finally, what was also surprising in the early work of Beadle and Ephrussi was that they both mentioned hormones as intermediates of gene action, a reference that seemed to have no place in their project. The way to overcome these contradictions is simply to admit that there was an implicit model of gene action that existed before the one gene-one enzyme relation, in which hormones had a central place and which the new model replaced. The solution emerged when Hans-Jörg Rheinberger studied the parallel and so far neglected work of Alfred Kühn on Ephestia (the flour moth) in Germany and brought to light the explicit reference of this researcher to hormones as intermediates of gene action (Rheinberger 2000; Morange 2015b). Even more interesting is when historians (or scientists who become historians) explore new areas such as studies of DNA repair (Friedberg 1997). This field of

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research, overlooked in previous histories of molecular biology, is closely linked to work on phages and is also important in work on cancer, on how different types of radiation affect the human genome, and on the mutagenic power of chemical compounds in the environment. Conceptually, these studies were also important in showing that DNA stability is not a property of the DNA molecule itself but of the whole cell and that organisms are able to control their rate of mutations (Keller 2000). In a similar way, Angela Creager explored the use of radioactive elements in biological research, pursuing the work initiated by Hans-Jörg Rheinberger (Creager 2013). This study, like the previous one, shows that the history of molecular biology is illuminated when viewed in a wider landscape. More generally, the development of the new technologies characteristic of molecular biology has received a lot of attention from historians (for a recent contribution, see Chiang 2009). Many studies have been devoted to electron microscopy. There are two reasons for this particular interest. The first reason is that long process leading to the recording of electron microscopy images has been scrutinized as an emblematic example of how scientific representations are constructed. In particular, the criteria used to discriminate between “true images” and artifacts have been hotly debated by historians and philosophers (Rasmussen 1993, 2001; Hudson 2003; Cambrosio et al. 2008). The second reason is that an electron microscope is a complex and expensive machine whose acquisition and use requires political and financial support. It represents a point of reference, a “totem” for the members of the laboratory (Strasser 2006). It is also important to acknowledge that the Rockefeller Foundation program, considered by most historians to have been highly important in the development of techniques to study macromolecules and in the rise of molecular biology, was shared and even anticipated by other institutions in the world. In 1927, in Paris, the Edmond de Rothschild Foundation created the Institute for Physical-Chemical Biology (IBPC in French) where physicists, chemists, and biologists worked on and facilitated the development of new technologies, notably chromatography and ultracentrifugation (Morange 2002). This points to the need to abandon historiography centered on the scientists and institutions involved and to devote more attention to a cultural history in which the layperson’s vision is often as important for the development of science. Important attempts have also been made to place early contributions to molecular biology in their scientific context. Two examples will illustrate this. Oswald Avery’s demonstration that the transforming principle of Pneumococcus was made of DNA was an important step toward the chemical characterization of the genetic material. But the discovery of the phenomenon of transformation is often seen as an anomaly in the work of its discoverer, the microbiologist Frederick Griffith. Pierre-Olivier Méthot recently showed that the discovery of transformation had its full place in Griffith’s plan for epidemiological control of infectious diseases (Méthot 2016). Similarly, there is a contrast between the important contribution of French biologists to the rise of molecular biology and the delayed creation in France of the first university chairs in genetics after the Second World War (40 years after the United States!) and the attachment of most French biologists to a Lamarckian

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tradition. Recent work has shed light on this paradox through a better description of the French neo-Lamarckian tradition and its close relation to the physiology of Claude Bernard. The favored vision of heredity that emerged was a dynamic one, which explains the attention paid to strange phenomena such as lysogeny – the existence of a dormant state of bacteriophages within bacteria. Temperate bacteriophages, responsible for lysogeny, represented one of the two experimental systems that gave rise to the first description of genetic regulatory mechanisms, the other being enzymatic adaptation (Loison 2015). Viruses and bacteria have been model organisms for molecular biologists, and the long history of these experimental systems, in particular the tobacco mosaic virus and the bacteriophage, has been scrutinized by historians (Creager 2002; Summers 1999). The role of bacteriophages in the rise of molecular biology is now well described, but it would be helpful to have a precise historical description of their decades-long therapeutic use in the USSR, in particular during the Second World War. Such a study would be useful at a time when the use of bacteriophages appears to offer a way to overcome antibiotic resistance. For years, the story of molecular biology has been reduced to the history of three schools of research: the American phage group headed by Max Delbrück and Salvador Luria; the Cambridge laboratories of Francis Crick (and Jim Watson), Max Perutz and John Kendrew, and Sydney Brenner and Seymour Benzer; and the French Pasteurian group of André Lwoff, Jacques Monod, and François Jacob. This small circle has progressively been widened to include other scientists – William Astbury in England and Boris Ephrussi in France – but also research groups in Belgium (the Rouge-Cloître group in Brussels), the Geneva group, and contributors from Germany, Italy, and Spain. There are also studies on work done in Mexico and South America. The panorama is far from complete, and we lack well-documented studies of what happened in Japan, China, Russia (USSR), and other Eastern European countries. The obstacles created by the domination of Lysenkoism and the role of physics departments in the initial development of molecular biology in the USSR are well known. But the historian of science would like to know whether the molecular biology that developed in these conditions adopted specific characteristics that distinguished it from other forms of molecular biology.

The Exploration of New Fields The Long 1970s Between the most visible success of molecular biologists, the decipherment of the genetic code at the beginning of the 1960s, and the rapid accumulation of puzzling observations and discoveries permitted by genetic engineering techniques at the end of the 1970s and the beginning of the 1980s, was a period of more than 10 years poor in discoveries, and for this reason called the “long 1970s” or “the crossing of the desert,” which did not immediately attract the attention of historians (Morange 1997). The time when this period could be neglected is clearly behind us. Indeed,

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it appears increasingly crucial for a redefinition of molecular biology and of its limits, models, and tools. Instead of being a vacuum, these years were a time of active research that exploited sophisticated technologies such as differential molecular hybridization (Suarez 2001; Suarez-Diaz 2013; Morange 2014). These years were also rich in the production of models: models of genetic regulation (Suarez-Diaz and GarciaDeister 2015), which became a priority after the operon model was proposed by Monod and Jacob; but also models of development and evolution, since in these years many molecular biologists moved toward the study of higher organisms and their development: the Britten-Davidson model of 1969 or the T-complex developmental model proposed by Dorothea Bennett in 1975. There were new observations on which some of these models were based: discovery of the abundance of short repeated sequences in the genome of eukaryotes and of the existence of large nuclear RNAs undetectable in the cytoplasm and the description of some of the steps (capping, addition of a polyA tail) leading to the formation of mature mRNAs in eukaryotes. Apart from the few exceptions mentioned before, this period has not been fully explored by historians. We have precise descriptions given by scientists themselves (Darnell 2011), and further studies will certainly reveal new facets of this rich time. These years were a period of confrontation but also of cross-disciplinary exchanges. The challenges that molecular data posed to evolutionary theory have been amply studied by historians (see ▶ Chap. 4, “The Historiography of Molecular Evolution,” in this book). The accumulation of protein sequences led to the production of the first “atlases” and databases (Strasser 2010). But other aspects of this expansion of molecular biology, and of a definition of its limits, have not received the same attention from historians. For instance, the huge debate that surrounded the characterization of the first memory molecules, the most famous of which was scotophobin, has not received adequate historical treatment: the material is there, with excellent historical descriptions provided by some of the protagonists (Irwin 2007). These years corresponded also to a rapid development of cell biology. Cells were no longer reduced to bags of proteins but were shown to be organized in different compartments with specific structures and functions. A new model was proposed (and rapidly accepted) for the cell membrane, and intracellular signaling pathways were described, as were the processes of protein maturation, secretion, and transfer to different cell compartments. A new powerful technology (immunofluorescence) allowed direct access to these processes. In the same years, Peter Mitchell proposed a model for the production of oxidative energy within cells that accorded mitochondrial membranes a major role, which was progressively confirmed. Molecular biology was replaced by molecular cell biology or molecular and cellular biology. These developments were boosted by genetic engineering techniques and led to an explosion of observations in the late 1970s and early 1980s. Once again, we have eyewitness accounts, but we lack full historical studies extending the description given by William Bechtel in Discovering Cell Mechanisms: The Creation of Modern Cell Biology (Bechtel 2006). The legacy of cell biology in the 1970s led to the more

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recent construction of chimaeras between a luminescent protein (such as GFP) and the proteins under study. It permitted observation of the movement of proteins in living, unfixed cells. The long history of the scientific observations that allowed the development of these powerful new tools has been wonderfully described (Pieribone and Gruber 2006). Some interesting contributions have also focused on the changes in representations that accompanied this transition to molecular cell biology (Serpente 2011, 2013, 2015). These years were also crucial for the emergence of new animal models. The nature of the “right model” for the study of development was amply debated. Sydney Brenner’s choice of the nematode C. elegans has been well described (De Chadarevian 1998; Ankeny 2001; Brown 2003). The molecularization of Drosophila has also been studied (Weber 2007), as has the more recent choice of Arabidopsis as a model plant (Somerville and Koornneef 2002; Leonelli 2007). But the importance of other models, such as the aggregating amoeba (Dictyostelium discoideum) and protozoa (Tetrahymena and Paramecia) and the increasing role of the mouse in biomedical research remain to be fully explored. The development of these new models led to the abandonment of the unique bacterial and bacteriophage model that had dominated the first steps of molecular biology (see ▶ Chap. 13, “Organisms in Experimental Research,” this volume). Two major transformations of molecular biology also occurred at this time: the development of genetic engineering tools and slow but constant progress in structural biology. The first has not escaped the attention of historians, but most studies have focused on the rise of biotechnologies, the establishment of new relations between universities and companies, and the consequences for intellectual property issues, which have been in the limelight for the last 40 years (Yi 2015). The best of these studies (see, for instance, Rasmussen 2014) do not overlook the numerous difficulties encountered by molecular biologists in obtaining the correct expression, in bacteria or yeast, of an active hormone or enzyme. These issues are addressed in another chapter. But despite this abundance of studies, some problems have yet to be explored. How were these technological developments progressively organized in a network, permitting the isolation and directed modification of genetic fragments and their transfer between different organisms? There were also alternatives, such as different approaches to the sequencing of nucleic acids, DNA and RNA: not all of them have been fully studied (Okada et al. 1966). The replacement of the notion of episome by that of plasmid bears witness to these changes, which have yet to be explored fully (Grote 2008). The ambitions underpinning these first attempts at genetic modification are worthy of study and should be positioned in the wider perspective of modifying human and animal species. The structural face of molecular biology has been neglected by historians, in comparison with its informational side, a point already made by Robert Olby more than 35 years ago (Olby 1979)! For instance, the “hubris” that typified the characterization of macromolecular machines at the end of the 1990s has been wholly ignored by historians. Doubts have also been voiced concerning the place of structural biology within molecular biology. Structural biology has always been a part of biochemistry, and its progressive transformation is not punctuated by

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dramatic episodes comparable to the discovery of messenger RNA or the decipherment of the genetic code! But what would molecular biology be without the structure of DNA? This structure is emblematic of molecular biology’s success, because of its beauty, but mostly because it showed how a function – carrying genetic information – could be explained by the structure of the molecule that served this function. This credo remains that of most biologists: complex functions cannot be reduced to the parts of a system, but a precise structural description of these parts will, in one way or another, help to understand the complex functions. This is also the conviction underpinning the search for targeted drugs. When molecular biology triumphed in the mid-1960s with the characterization of the genetic code, the structure of only two proteins, myoglobin and hemoglobin, had been described at low resolution, which was insufficient to explain their functions. Decisive progress was made in the following years. A precise description of the structure of hemoglobin was provided and used to explain the mechanisms by which oxygen is taken up by the protein in the lungs and released to the tissues. Simultaneously, the first enzyme was crystallized (lysozyme) and its catalytic power explained. This was the beginning of a long series of technical improvements that facilitated the preparation of crystals, reduced the time needed for the analysis of diffraction data, and opened the door to the crystallization of larger protein complexes or classes of proteins (membrane proteins) until then resistant to crystallization. These transformations were the extension of previous efforts (described, for instance, by Soraya de Chadarevian in her account of the Cambridge lab (De Chadarevuan 2002)), boosted by the new techniques of genetic engineering that facilitated the preparation of pure (possibly modified) proteins that are easier to crystallize and to analyze. A new representation of proteins emerged in parallel with the accumulation of structural data. While the increasing role of computers in protein structure determination and representation has been well documented by historians, the dominant representation today has been done by hand and has only one creator, Jane S. Richardson. This is a wonderful and maybe unique example of a new representation, elaborated by one person and adopted by the whole scientific community. It deserves more attention from historians to add to Richardson’s own accounts. This new structural vision and representation pushed aside other types of representation that emphasize the importance of lower levels of organization, the quantum level, and the fluidity of protein structures. It would be a useful historical contribution to study these alternative models at a time when fluidity, plasticity, and quantum phenomena are again highly valued in biology.

The 1980s: An Explosion of Discoveries A short and incomplete list of these discoveries will be sufficient to give an idea of the way biological knowledge was rapidly renewed: the description of archaea, the third branch of the tree of life (at the end of the 1970s); the progressive

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characterization of the complex mechanisms of transcription in eukaryotes (and of its regulation with the discovery of “enhancers” (Darnell 2011)); the distinction within the DNA molecule between introns and exons and the description of the splicing process; the discovery of the different mechanisms generating the complexity of antibodies and receptors in the immune system; the isolation and characterization of the first oncogenes and tumor suppressor genes (Weinberg 1996); the isolation of the first developmental genes (the homeobox-containing genes) and the discovery of their conservation in evolution; the characterization of mutations modifying the duration of life in yeast (Guarente 2003) and in the nematode C. elegans but also in humans; and the progressive rise of protein engineering. To this list, one must also add a discovery made in the 1990s but, which has often been considered as “postmature,” the discovery of small regulatory RNAs. In this long list, what are the potential topics for historical study? Some issues, like the complex mechanisms of transcriptional regulation, seem to concern the specialists, and a historian could barely add anything to the precise accounts already written by some of the protagonists (Darnell 2011). Other discoveries are better candidates, when biologists consider that they did not occur at the right time but were delayed by “obstacles” (with the meaning given to this term by Gaston Bachelard) that prevented acceptance of the solution finally considered as the right one. Such is the case of the splicing process and of small regulatory RNAs. In these cases, the participation of historians is also required by scientists because of disagreements concerning the relative contributions of the various protagonists. Characterization of the obstacles that prevented earlier acknowledgement opens the door to evidence for a shift in the explanatory framework. Other discoveries are of interest, not in the way they were made but through the transformations that they effected in other disciplines. This is the case of the characterization of the first developmental genes, which led to the emergence of Evo-Devo, a new scientific field at the boundary between evolutionary biology and developmental biology (see ▶ Chap. 3, “The Historiography of Modern Evolutionary Biology,” this volume). As I will discuss later, it is obvious that the significance of these discoveries cannot be fully appreciated independently of the long history of experiments, models, and theories previously developed to account for these phenomena. The discovery of oncogenes and tumor suppressor genes cannot be understood without reference to the models proposed for the origin of cancer throughout the twentieth century. And the structural and functional characterization of mutations affecting the duration of life is only significant when viewed within the succession of models that were proposed for aging. Some descriptions and historical work have already been accomplished in this sense, but much remains to be done.

The 1990s: The Rise of Genomics The origin of the human genome sequencing program and the debates that punctuated its development have been well described by the scientists themselves (Davies

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2001; Sulston and Ferry 2002) and by historians (Bostanci 2004): the need to sequence the whole genome or only its expressed parts; the value (or not) of sequencing in parallel the genomes of other organisms; the strategy of establishing a genetic and physical map of the genome with the contribution of the “French DNA” (Rabinow 1999; Kaufmann 2004) before sequencing its fragments, as opposed to a “shotgun” approach in which fragments are obtained by random cuts in the chromosomes, sequenced, and reassembled from sequence overlapping on computers; the attempt by Celera Genomics headed by Craig Venter to patent gene sequences and the free-access policy of the International Consortium; and the accuracy of sequencing methods as opposed to the need for a technological jump. The term “genomics” itself was abruptly introduced by Victor McKusick and Frank Ruddle as the title of a new journal (McKusick and Ruddle 1987). The new organization of the work and the increasing role of computers (Francoeur and Segal 2004; November 2012) and of computational biologists and databases have also been explored by historians (Garcia-Sancho 2011), as have the expectations (and disillusions) attendant upon the program. However, the human genome sequencing program has attracted too much attention, and a precise description of other genome sequencing program, in particular those that yielded results before the human genome program, is still lacking. The most interesting studies are those in which these developments are repositioned in a long trajectory: the long history of sequencing (Garcia-Sancho 2012b) or of the increasing role of bioinformatics (Stevens 2013), with the transition from “big machines” to personal computers (Garcia-Sancho 2012a). Particularly interesting from this point of view is the progressive construction of databases, from the first atlas of protein sequences (Strasser 2010, 2011) to the multilevel databases of sequences, but also structural and physiological data (Richardson and Stevens 2015). The development of these databases has its roots in the “natural history” tradition, which was relevant even in the early days of molecular biology (Strasser and De Chadarevian 2011). The creation of these databases also generated a moral framework for the research and for the behavior of biologists (Leonelli and Ankeny 2012). The functional classification of sequences generated the production by computational biologists of “ontologies,” which, once created, orient the data interpretation (Leonelli 2010). This is a wonderful example illustrating the need to integrate historical, philosophical, and sociological studies. Edna Suarez-Diaz has emphasized the role of evolutionary questioning, both in the development of databases and in the progress of bioinformatics (Suarez-Diaz 2010). Reciprocally, the comparison of sequences has become one of the main heuristic tools in genomics and has had various consequences. The first is that organisms to be sequenced in priority were increasingly selected because of the evolutionary questions that deciphering their genomic sequences would solve. In addition, if model organisms were supported by the rapid sequencing of their genomes, newly sequenced organisms would more easily be converted into new models. The exchange of information created by the development of databases facilitated the circulation between models, and expansion of their number, a revolution in comparison to what happened in the first years of molecular biology.

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Research is now clearly oriented toward a description of the diversity of the living world and not of the principles of structure and function shared by all organisms.

Entering the Post-genomic World I will designate by the term post-genomics the period that started with the completion of the human genome sequencing program. What characterizes this new period of biological research is unclear (Richardson and Stevens 2015), to the point that it has been suggested that the new name was proposed to avoid a decrease in funding, once the human genome had been fully sequenced. Some of its characteristics had emerged in the previous period: the importance of databases, the time spent by biologists in front of computer screens, the increasing role of data-driven research, the emergence of new types of representations, and the increasing part played by huge scientific consortia; for instance, the ENCODE consortium seeks to attribute a functional role to the noncoding sequences of the genome. I have selected three areas of research that seem emblematic of post-genomics: systems biology, epigenetics, and metagenomics. Systems biology and epigenetics preceded the emergence of post-genomics, but both have characteristics commonly attributed to the current state of biological research. The father of systems biology was Ludwig von Bertalanffy in the 1950s, but this new discipline became fashionable only at the end of the 1980s (see Powell et al 2007 for a brief historical record). It is possible to distinguish two quite different forms of systems biology. One, which flourished in the 1990s, had huge ambitions: to find in the structure of the networks formed by the different molecular components a new “logic of life” that had so far escaped the attention of molecular biologists, whose studies were focused on individual molecular components. After some disappointments encountered by this strong program, the present dominant vision is more Cartesian: it does not break with the molecular studies but simply considers that precise knowledge of all the molecular components and their interactions will yield a hitherto unattainable degree of understanding of organisms. An important issue that has been addressed by historians and philosophers, but not yet definitively resolved, is whether (and eventually how) this new systemic vision differs from the mechanistic vision that has dominated “classic” molecular biology (Braillard 2010; MacLeod and Nersessian 2015). Epigenetics also predated post-genomics. Today, epigenetic work mainly consists of the study of DNA and histone (chromatin) modifications that alter gene expression and are transmitted through mitosis and, in some (rare) cases, through meiosis. It is a very rich, but also very complex field of research, at the boundary between developmental biology, evolutionary biology, ecology, and even ethology. It is a pity that there is no complete history of epigenetics, of the origin and multiple uses of the term, a history not written by those who push the development of this discipline. The claimed close association of epigenetics with post-genomics suggests that the latter leaves more room for the effects of the environment and less to genetic determinism than traditional molecular biology.

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The third, highly visible field of research in post-genomics is metagenomics – the possibility of simultaneous access to an ensemble of microorganisms (and their genomes) without the bias represented by the in vitro cultivation of the different microorganisms. One area of predilection is the study of the microorganisms present in the digestive tract – the microbiota. The relations between this ensemble of microorganisms and the functions of the human body, as well as the anomalies in the composition of the microbiota that may be related to human diseases, are very active fields of research. Diseases that are concerned are not only those affecting the digestive tract, such as Crohn’s disease, but even psychiatric diseases. Metagenomics is an extension of genomics but also belongs to systems biology. Are there discoveries that would be characteristic of metagenomics? One case, studied by Maureen O’Malley, is the discovery of proteorhodopsin genes (rhodopsin being the protein involved in light capture) in many microorganisms not previously considered to capture the sun’s energy (O’Malley 2007). Although it is a beautiful example of exploratory research, it is not evident that the systemic nature of metagenomics is involved in this discovery: the fact that the different microorganisms form a system has no role in this story. While the initial observation was not hypothesis-driven, O’Malley shows that the development of the work was based on previously obtained information on the structures and functions of rhodopsins.

The Need for longue durée Studies I have already mentioned that some issues cannot be fruitfully addressed by focusing on short periods of time: the meanings and expectations invested in epigenetics, the transformations in the models of the origin of cancer, and the relations between molecular biology and evolutionary biology require, to be fully understood, a description of the theories, models, and experimental systems that were used during previous, often long periods of time. This does not mean that these epistemic tools were the only players in, for instance, the complex history of cancer research, but rather that if they are ignored, or more often given a nonhistorical description, a full understanding of the transformations that occurred will not be possible. I will provide six examples in which a well-informed historical study would be highly useful: six potential research projects! The first concerns synthetic biology (Porcar and Pereto 2014). The authors of a recent book exploring the design project of synthetic biologists uncritically accept the radical novelty of this discipline claimed by some of its most famous exponents (Ginsberg et al. 2014). In contrast, Evelyn Fox Keller has resurrected the work and writings of Stéphane Leduc, who in the 1910s claimed to have created synthetic forms of life and who introduced the expression “biologie synthétique” (Keller 2002). Others have alluded to Mary Shelley’s novel Frankenstein or to the myth of the Golem. But what has not been carefully studied, and is far more significant, are the relations between genetic engineering and synthetic biology. One particularly interesting facet of this missing story is the history of protein engineering, from the first rational approaches to protein design in the 1980s to the currently extensive use

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of directed evolution – the selection among random mutations induced by the experimenter. The development of enzymes from antibodies – the famous abzymes – and the failure to obtain enzymes with a catalytic power comparable to that of “natural” enzymes is a history waiting to be written. In addition, this discipline was progressively transformed by the introduction of evolutionary models and concepts, such as the notion of a trade-off between protein stability and, for instance, catalytic efficiency. With this historical framework in mind, the comparison often made between the tinkering action of evolution and the “engineering spirit” of synthetic biologists appears simplistic or more exactly an illusory strategy to sell the new discipline. A second, closely related but independent issue is the process of protein folding. The problem stems from the sequence hypothesis proposed by Francis Crick simultaneously with the Central Dogma, according to which a protein spontaneously folds to reach the lowest accessible state of free energy. The present state of research is well known, with the use of powerful computers to predict the three-dimensional structure of a protein from its sequence. Between the two, a rich and complex history has yet to be recounted: the debates on the mechanisms of conformational changes of protein in the 1960s (for or against the allosteric theory); the discovery of prions in the 1960s and the models progressively proposed to explain their “replication” and their pathogenic power; the dramatic change in protein folding models in the 1990s (with the elaboration of the funnel model); the discovery of protein chaperones, assisting protein folding, and the interpretation of their activities (integrated in the so-called heat-shock response); and the present emphasis on the dynamics of protein structures following the fast development of nuclear magnetic resonance technologies. Some minor parts of this history have been scrutinized by historians, such as the discovery of prions and of their role in neurodegenerative diseases. But the studies have often been biased toward a specific issue – does the existence of prions violate the central dogma of molecular biology? – masking the relation between the question of the nature of prions and the research on the mechanisms of protein folding. An additional interest of this field of research is its discrete but real interdisciplinarity. I also mentioned the history of the characterization of regulatory RNAs – siRNAs (small interfering RNAs), microRNAs, and, more recently, the long noncoding regulatory RNAs. They have an important place in current research. Their delayed discovery, the debates on the Nobel Prizes attributed for the discovery of siRNAs, demonstrates that there is a need for serious historical work. Once initiated, many long-forgotten studies and debates will reemerge: the demonstration in the 1930s of the acquired immunity of plants after viral infections; the early models of regulation by RNAs in the 1960s; the use of antisense RNAs in the 1980s to control gene expression. Even the debate on junk DNA and the notion of biological function finds its place in this wide historical perspective. Expanding the historical framework helps to interpret recent historical events through older ones. This historical background is even more necessary when ethical and societal issues are directly associated with particular fields of research. This is the case of the experiments designed to edit the genome (and to modify the germline) with the CRISPR-cas 9 system. The historian can provide a precise history of the way these

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new tools were elaborated and replace the biased historical record produced by the scientists themselves (Morange 2015a). But history can provide more. How is it possible to discuss these new developments without describing the first difficult steps of gene therapy in the 1980s, the “dreams” of molecular biologists in the 1960s, the ambitions of evolutionary biologists in the 1930s, and the projects of eugenicists in the first half of the twentieth century? The same is true for regenerative medicine. The history of research on stem cells is rich, yet only some of its episodes have been described so far. The early observations on plants, the progressive elaboration of the notion of hematopoietic stem cells, and the three decades separating the early observations on the stem cells of teratocarcinomas, tumors of the gonads, from the preparation of mouse embryonic stem cells, are still poorly described. In a sense, stem cells were of more interest to molecular biologists than to embryologists, because they were a material well adapted to molecular studies. A last puzzling issue is the question of aging and its mechanisms. The history of theories of aging concerns more than the history of molecular biology! But soon after theories of aging emerged, molecular biologists proposed, in the 1960s and 1970s, molecular models of aging (Jiang 2014) and even suggested the existence of a “program of aging.” Interest in the evolutionary models of aging soon rendered these first molecular models obsolete. But, in parallel and independently, the question of aging was taken up again through the characterization of mutations affecting the duration of life, in yeast, C. elegans, and in humans. The description of the mechanisms of chromosome replication and the discovery of telomeres and telomerase shed new light on the process of aging (Brady 2007). These molecular observations do not provide a coherent picture of the aging process, but they underpin some of the plans for germline genome editing designed to prolong human life.

Conclusion Was it reasonable to include the developments in genomics and post-genomics in a historiography of molecular biology? There are so many differences between the practices of molecular biologists in the 1950s and present-day biologists. High-throughput technologies have replaced the traditional tedious techniques, to the point that research is considered as no longer hypothesis-driven but data-driven. The nature of work has also changed: the long hours spent at the bench have been replaced by equally long hours in front of the computer screen, searching for information in the increasingly rich databases, a transition from wet to dry experiments. The research is done by interdisciplinary teams, as shown by the long lists of authors contributing to publications in the post-genomic era. The feeling of radical novelty is shared by the researchers involved but also by social scientists who act as heralds of the new disciplines. This vision is too common not to deserve closer examination. Consider, for instance, epigenetics. The importance that it has acquired is considered as emblematic of the new post-genomic biology. Paradoxically, epigenetics is said to have been founded by Conrad

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Waddington in 1940, before the rise of molecular biology! It is easy to show that epigenetics as it is practiced today has nothing in common with what Conrad Waddington called “epigenetics.” This obviously shows that we cannot trust the official historical discourses to assess the novelty of presently flourishing disciplines. It is probably high time to rehabilitate the notion of longue durée, a concept introduced by Fernand Braudel (Braudel 1958) and adapted to the history of science by Frederic Larry Holmes (Holmes 2003). In an illuminating contribution, Mathias Grote has recently tried to extract from this vague notion what could be significant for current historians of science (Grote 2015). He first distinguishes longue durée studies from big picture histories, the latter implying a longue durée in a limited number of cases. He sees an obvious place for longue durée in the repetition of practices, what he calls “sustainability in scientific practices.” Good examples are found in classification. The existence of natural history practices in early molecular biology as well as in the current use of databases is another example of these sustainable practices (Strasser and De Chadarevian 2011). This helps us gauge whether molecular biology is dead and has been replaced by new disciplines. The main practice of molecular biology consisted in isolating macromolecular components and characterizing their structure with a view to explaining their functions. The best evidence for the existence and role of these components came from the isolation of mutations affecting their function. Has this practice disappeared or been dramatically transformed? The answer is obviously no. The purification of macromolecular components and the characterization of their structures still have a major part to play in biological research. The search for mutations has adopted new forms – through Genome-Wide Association Studies (GWAS) – but the link between DNA modifications (mutations) and alterations of function remains. Most of the work done in epigenetics consists of the characterization of DNA and protein modifications and the functional consequences of these modifications. The rise of systems biology would have threatened this practice if its early supporters had been right, if the description of the form of the network (system) had been the only information needed to understand how the system works. In this case, the practice of biologists would have been dramatically changed. But this has not occurred: a precise description of the structure of the components is still favored. The properties of a system are more than the properties of the individual parts. But it would be a blow to the intelligence of most molecular biologists of the 1950s and 1960s to suggest that they ignored that! In all the new disciplines and research fields that we have described, in synthetic biology as in work on genome editing, there is a continuity of practices with the “classic” era of molecular biology, beyond the obvious differences in the tools used by biologists (Morange 2020).

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Biomedicine and Its Historiography: A Systematic Review

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Contents Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . What Is Biomedicine? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Biomedicine’s Postwar Development . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . The Distribution of Activity in Biomedicine and in Its Historiography . . . . . . . . . . . . . . . . . . . . . . . Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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Abstract

In this essay I conduct a quantitative systematic review of the scholarly literature in history of life sciences, assessing how well the distribution of the activity of historians aligns with the distribution of activities of scientists across fields of biomedical research as defined by expenditures by the cognate institutes of the United States NIH. I also ask how well the distribution of resources to the various research fields of biomedicine in the second half of the 20th Century has aligned with morbidity and mortality in the United States associated with the cognate disease categories. The two exercises point to underexplored areas for historical work, and open new historical questions about research policy in the US.

Introduction I have taken an unusual approach in this essay to ask a question that, to my knowledge, has not been addressed before: how well, in terms of subject matter, does our historiography describe the domain of biomedicine? I shall define the subject matter by treating it as an actor’s category; that is, I define biomedicine according to the way the concept was N. Rasmussen (*) University of New South Wales, Sydney, NSW, Australia e-mail: [email protected] # Springer Nature Switzerland AG 2021 M. R. Dietrich et al. (eds.), Handbook of the Historiography of Biology, Historiographies of Science, https://doi.org/10.1007/978-3-319-90119-0_12

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used by scientists and policy-makers in the institutions they created at mid-century. I shall briefly describe the early history of the institutionalization and growth of this kind of research activity after the Second World War. Then I shall bring quantitative evidence to bear in a way that is somewhat novel to map the distribution of resources, hence research activity, among the major fields within the biomedicine of the United States during the five decades that followed the War. (I focus on the United States because it was the polity supporting and shaping biomedical science far more than any other and because it exerted additional influence as the West’s Cold War leader.) The first major purpose is to ask in quantitative terms how well the distribution of research effort among biomedical fields matches the importance of the different fields of research to the health of the US population – as political leaders and technocrats of the day acknowledged that they should – or whether, on the other hand, there are interesting historical questions about why some fields attracted disproportionately little or much cultural and political favor. This leads me to propose a number of topic areas for further research by historians of biomedical research policy and politics. An additional purpose for charting the distribution of health research resources is that it enables one to address the novel question noted above and an issue central for this collection: assessing how well the biomedical historiography aligns with the historical trajectory of biomedicine itself. That is, I ask whether historians are distributing their efforts to the various fields of the biomedical enterprise in a way roughly matching the past distribution of efforts by biomedical researchers and policy-makers generating the activity itself. Here, too, we will be able to identify apparent mismatches between events and attention and offer some hypotheses about why certain fields appear relatively underor overstudied by historians. To identify mismatches I apply a quantitative method to the historiography – systematic literature review – rare for the history of science and medicine but standard in certain biomedical fields. In a stand-alone review article, I would follow these three quantitative sections – dealing in turn with the biomedical research enterprise over time, its fit with population health needs, and its fit with the historiography purporting to describe it – by drilling down into several particularly active areas of biomedical historiography, looking qualitatively at the prevalent questions being addressed and fashionable approaches to answering them. However, given that this review is part of a collection in which many of the other historiographic articles deal with areas of life science that overlap what I (following the US National Institutes of Health, NIH) have broadly defined as biomedicine, I will leave such qualitative analyses to the other authors. The main aim of my largely quantitative survey is to point out historiographic lacunae and raise new questions for historians; it is also intended to demonstrate that quantitative systematic review methodology can be productive, but not to rule out other more qualitative approaches.

What Is Biomedicine? The first task of any literature review must be to define the subject matter. Bibliometric tools (such as Google Ngram for book contents or ProQuest Historical Newspapers for American journalism) show that as a term in English usage, and

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arguably too as a generally accepted concept, “biomedicine” appears around 1960 and becomes ubiquitous by 1970. So, while both clinical research and experimental biology with deliberate medical application certainly dates to the nineteenth century, for example, in Koch’s and Pasteur’s famous vaccine trials and Helmholtz’s and Bernard’s work on nerve action, and while furthermore medicine has increasingly borrowed concepts, practices, and authority from laboratory sciences since the late nineteenth century (Vogel and Rosenberg 1979; Starr 1982; Ludmerer 1985; Rothstein 1987; Porter 1998), the postwar period manifests this melding of science and clinical technique on a new scale. After the Second World War, government funding for research dramatically expanded in the United States, quickly overshadowing private philanthropies such as the Rockefeller Foundation and Institute – which before the War had been the preeminent patrons of both biological and medical research, and not only in the United States. In the postwar United States, the position of dominant patron was rapidly assumed by the National Institutes of Health. In the mid-1970s, for example, NIH funding accounted for 40% of US health R&D, as compared with about a 25% share from all other government funders, about 30% from the private sector, and less than 5% nonprofits and philanthropies. The overwhelming dominance of the NIH among all US funders of life science is also attested by a comparison of the agency’s 1970 budget of more than $1 billion with the $49 million research budget of the National Science Foundations’ (NSF) division of Biological and Medical Sciences (Table 1). Indeed, in terms of resources, the NSF as a patron of life science was roughly equivalent to one of the more modest NIH Institutes (Strickland 1972; Brown 1979; Shyrock 1980; Bloom and Randolph 1990; Kohler 1991; Appel 2003; Hamowy 2008). As implied by the mushrooming NIH budget and proliferation of NIH Institutes from the late 1940s through the 1960s, briefly discussed below, health-oriented science became a social priority in the United States like never before. That commitment to biomedical research, by the world’s richest nation amidst a global scientific community shattered by war, made the United States the dominant force in health research globally. Indeed, it was not until the mid-2000s that the US share of world health research and development spending fell beneath 50%.1 Beyond mere spending power, we may also point to literature indicating that in biomedicine as in science generally, during the Cold War many Western countries tended to emulate the “Free World’s” leader in order to demonstrate solidarity and furthermore that the East Bloc often felt compelled to compete with US scientific strengths (Gaudillière 2002; Krementsov 2002; Krige 2006). This cultural leadership in the postwar period offers another reason why the American biomedical research enterprise shaped others throughout the world. Thus, putting aside qualms about provincialism, for

1

US public and private combined biomedical research spending still accounted for half of world spending in the mid-2000s (Moses et al. 2015). I am not aware of reliable figures earlier than this, but given the postwar state of European and Asian economies until the 1980s, it would be safe to suppose that before the 1990s the United States would have accounted for far more than half.

Founded 1937 1948 1948 1948 1949 1950 1950 1962 1962 1968 1969 339,380 158,600c (88,350 res) 24,870d

1960 budget 91,257 62,237 34,054 10,019 67,470 46,862 41,487

1970 budget 181,454 160,634 97,342 28,754 NA 131,761 97,315 76,095 148,294 22,828 17,423 1,061,007 462,490 (316,410 res) 49, 450c

1980 budget 999,869 527,488 215,364 68,303 212,000a 341,206 241,966 208,953 312,468 112,989 83,893 3,428,935 975,130 (836,830 res) 170,000e

1990 budget 1,634,332 1,072,354 832,977 135,749 727,000b 581,477 490,409 442,914 681,782 236,533 229,234 7,576,352 2,100,000 300,000f

BMS, BBS Directorate of Biological and Medical Sciences/Biological Behavioral and Social Sciences Although total Institute budgets are given here, approximately 90% represents research expenditure. For NIMH, incorporated in the late 1960s as a research division within another non-NIH agency dedicated mainly to service delivery (ADAMHA), research expenditures are given for 1980 and 1990. In 1992 NIMH, together with NIDA and NIAAA, was reconstituted as an Institute when that agency was reorganized. (Sources: NIH figures from http://officeofbudget.od.nih.gov/approp_hist.html) a ADAMHA research division budget from http://www.ncbi.nlm.nih.gov/books/NBK235741/ b ADAMHA research division budget from http://www.ncbi.nlm.nih.gov/books/NBK235734 c Budget obligations (http://dellweb.bfa.nsf.gov/NSFFundingbyAccount.pdf), and d New grants, both from Appel (2003) e Total divisional appropriation (http://library.cqpress.com/cqalmanac/document.php?id=cqal79-1183790); research obligation not available f Approximate estimate from http://www.ncbi.nlm.nih.gov/books/NBK235736/

NIH Institute NCI NHI (NHLBI) NMI (NIAID) NIDCR NIMH NIAMD (NIDDK) NINDB (NINDS) NICHD NIGMS NEI NIEHS NIH total NSF total NSF BMS/BBS (res)

Table 1 Budgets of NIH Institutes founded before 1970 and NSF research budgets (in current $1000s) over time

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present purposes I will define “biomedicine” as the areas of research supported and conducted by NIH. This essay will emphasize the half-century from 1945, which is to say the period in which biomedicine as we know it first appeared, up to the recent horizon of what can be considered history. But I will take it as given that many of the particular research disciplines and fields that flourished under the “biomedicine” aegis existed in some form since the early twentieth century. In accepting this postwar actor’s category, I am aware that I am bypassing a significant body of sociological literature – and controversy – seeking to define the essence of the biomedical, and of associated “biomedicalization,” and when exactly the postmodern form of “biopower” emerged (in the 1980s for many authors; Rabinow and Rose 2006; Sunder Rajan 2006; Clarke et al. 2009). However, for purposes of the present review, we do not need to theorize biomedicine deeply; we only need to recognize biomedical research and historical work describing it. For this limited scope, the actor’s category works well and reduces the risk of anachronism. So, for present purposes “biomedicine” is defined operationally as what NIH did (and does). As the names of the Institutes indicate – as do the introductory sections of their present web sites – the research supported by NIH is defined not so much by the disciplines of academic life science (cell biology, developmental biology, molecular biology, neuroscience) as by the disciplines and specialties of academic medicine, which roughly correspond to human organ systems and their diseases. Thus the National Cancer Institute (NCI) sought (and seeks) to expand the knowledge base and techniques of the conjoined specialties of hematology and oncology; the National Institute of Mental Health (NIMH) psychiatry; the National Heart Institute (NHI, later, Heart, Lung and Blood Institute, NHLBI), cardiology; the National Institute of Allergy and Infectious Disease (NIAID) the gradually diverging specialties of immunology and infectious disease; and so forth. Biomedicine’s autodefinition according to medical specialties, by its central agency the NIH, presents an immediate challenge to the historian (and historiographer) of biology. Historians of life science typically define their own subject matter according to the fields one finds in a university faculty of arts and sciences, corresponding to the way that the key figures we have mainly focussed on, biology professors, define themselves. Scientists with very closely related research programs, when employed in medical schools, often find themselves and their departments defined at least indirectly by reference to clinical practice. Indeed we may already sense a shortcoming of the professional historiography, because the list of Nobel Laureates in Physiology or Medicine – a Prize mapping well onto “biomedicine” in the postwar period – shows that many of the century’s leading biomedical scientists held physician’s degrees and have defined their own careers in terms of medical speciality and problem, in NIH fashion. Sampling the Prize every 5 years or so, consider Max Theiler (1951, virology, on yellow fever), Dickinson Richards (1956, cardiology, on catheterization), Frank Macfarlane Burnet (1960, immunology, on transplant compatibility), CB Huggins (1966, oncology, on prostate cancer), or Earl Sutherland Jr (1971, endocrinology, on hormone

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signalling).2 Such medically oriented figures (with a few exceptions) have received but little attention from professional historians, in that historians of medicine typically attend more to patient care while historians of biology attend more to biologists oriented toward basic science. Thus, a great deal of past biomedical science has fallen between the stools, so to speak, of the scholarly fields of history of biology and history of medicine. In the next section, I will explore the validity of this impression through a systematic review of the historiography concerning the many areas of biomedical research encompassed by NIH activity. All this is not to say that historians of science have failed to notice scientists funded by NIH and therefore biomedicine as viewed from Bethesda. Of course they have studied NIH-funded scientists, as one expects given that the NIH was the dominant funder of life science in the United States, both basic and applied (to use the commonplace postwar distinction). World-leading departments in fields we think of as completely removed from the clinic were built on NIH money, for example, the molecular biology hothouse of Stanford’s Biochemistry Department (in the medical school), as ably discussed by Doogab Yi (2015), where luminaries like Paul Berg and Arthur Kornberg solved the riddles of gene expression in E. coli bacteria. But historians typically act as if NIH medical agendas had no influence over what they did. The biology, historians suppose, was the same no matter who funded it (so long, at least, as it was scientist-initiated and scientist-evaluated research). This conceit meshes with the internalist premises of Mertonian and Kuhnian history and philosophy of science scholarship from the Cold War era, which in turn reflects the self-conception of many scientists of the time – a construction of good science as independent of whatever practical or political implications it might have that was encouraged by the American National Security State in its early Cold War heyday (Hollinger 1995; Fuller 2000; Shapin, 2008, Chap. 3). This purist conceit among biologists played a key role in the cultural Cold War, I have argued (Rasmussen 2014, Chap. 1), in the same way that the modernist conceit that good art seeks only to advance the potential of its medium and discipline (e.g., in abstract expressionist painting) found favor among Guggenheim Foundation and State Department decisionmakers promoting the image of a Free World where artists and intellectuals were not enslaved to political agendas (Saunders 2000; Osgood 2006; Wilford 2009). This same conceit allowed university physicists developing the maser in classified Navy-sponsored research projects to imagine that they were simply pursuing paradigm-driven science for its own sake (Forman 1992) – and similarly, biologists pursuing the genetics of cancer with tobacco industry funding (Bero 2005; Proctor 2011, Chap. 16; Brandt 2012). It is time, I would urge, for historians of life science to shake off the Cold War hangover and broach the question of whether the science really was the same regardless of who funded it, even when it

2

http://www.nobelprize.org/nobel_prizes/medicine/laureates/, accessed 2 July 2015. I have not counted how many Physiology or Medicine Laureates have higher medical, dental, or veterinary degrees, but the proportion is very substantial [why not count and give stat to us?]. Another interesting question is whether that proportion has changed over time.

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was peer-reviewed. But here I stray from my central task of comparing the biomedical territory’s distribution to its historiographic map. After we have a more reliable measure of key divergences between the attention of professional historians of biomedicine and the historical terrain they aim to describe, I will venture some speculations to explain these divergences.

Biomedicine’s Postwar Development Before we compare the recent historiography to the past biomedical research enterprise it describes, a review of that enterprise’s emergence is in order. For the United States, the rise to prominence of biomedical research on the political scene has been comparatively well studied. Biological and medical research contracts of the wartime Committee on Medical Research (a branch of Vannevar Bush’s science agency, OSRD) were in 1946–1947 adopted wholesale by the NIH’s director Rolla Dyer under Surgeon General Thomas Parran, the expansionist chief of the late New Deal Public Health Service (PHS). While the National Science Foundation proposed by Bush was delayed in Congress by disputes over its exact form and purpose (until 1950), by the end of 1947, the NIH had transformed itself from a research sponsor mainly of in-house research at its Cancer Institute, as well as the home to smaller PHS laboratories with infectious disease and other public health functions, to a patron of academic life science research on a scale an order of magnitude larger. Riding a wave of popular enthusiasm for the idea that the main threats to health and longevity could be conquered through science, a throng of new National Institutes of Health was conjured by lobbyists such as Mary Lasker and their Congressional allies – the National Institute of Mental Health in 1946 (but funded and formally established in 1949), the Heart, Microbiology, and Dental Institutes in 1948, and the Diabetes and Stroke Institutes in 1950 (Kevles 1977; Fox 1987; Strickland 1989; Kleinman 1994; Hamowy 2008). All boasted a campus laboratory in Bethesda but sponsored much more research extramurally through grants. As shown in classic work by Stephen Strickland (1972), based largely on Congressional and other political discourse, the rapid growth of the NIH within PHS during this period emerged as an unintended quasi-compromise from the clash over Harry Truman’s 1948–1950 effort to institute national health insurance. This initially popular plan was notoriously defeated by an all-out lobbying and advertising campaign against “socialized medicine” by the American Medical Association and its business allies (Poen 1996; Mayes 2004, Chap. 3; Boychuk 2008, Chap. 3). Public funding of research to improve medical care (for paying patients) became a palatable alternative to publicly funded medical care, not just for physicians and drug companies but also for health advocates and for politicians concerned to demonstrate both to constituents and to the polarized Cold War world that the United States cared about its citizen’s health. Given an impoverished postwar Europe encumbered by a greater dedication to health care and other social welfare, this concerted effort by the world’s richest nation quickly made it the world leader in life science research and likewise clinical technique. This is not to suggest extreme divergence between the United States and Western Europe, however. On the one hand, other Western nations

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seeking to retain great power status, like France, also emulated the American hightechnology biomedical turn in life science (Gaudillière 2002). On the other, the American Federal government found indirect ways to subsidize health care, such as through the Veterans Administration and hospital construction, long before Medicare was instituted in 1966 (Hamowy 2008). Under the NIH aegis in the 1950s and 1960s, the life sciences as a whole participated in what Roger Geiger (2008) has called the “hypertrophy of disinterested research” at universities and other scientific institutions in the early Cold War era. NIH funding grew exponentially from the early 1950s, reaching a peak in 1966 at just over $1bn constant 1960 dollars. But the growth of the research enterprise outstripped Federal funding; already in 1966, at peak funding, grant success rates dipped below 1/2 and dropped to 1/3 over the next 5 years as competition intensified (Fig. 1). Therefore in the same period that the Mansfield amendment reduced military funding for physical sciences at American universities, Federal funding for life sciences declined in real terms – and was perceived to move in a more narrowly pragmatic direction too, so that (for instance) at Stanford Paul Berg worried in 1970 that NIH might no longer support any work on bacterial viruses because it was too remote from medical application (Yi 2015, 55). NIH funding did not return to its 1966 level in real terms until 1972, with an infusion of money from the 1971 National Cancer Act (“Nixon’s war on cancer”), and despite this infusion remained at an approximate plateau of a little more than $1 billion 1960 dollars for more than a decade. Real NIH research spending did not start to expand 0.7

2,500,000

0.6

2,000,000

0.5 1,500,000

0.4 0.3

1,000,000

0.2 500,000

0.1

0

o NIH Budget ($1960)

0

YEAR

R01 success rate

Fig. 1 NIH grant success rates and budget (constant 1960 $1000s), 1960–1995. (Sources: For NIH appropriations, http://officeofbudget.od.nih.gov/approp_hist.html; for NIH grant success, http:// www.nih.gov/UploadDocs/Estimated_success_rates_1962-2008.xls (both downloaded February 7, 2013); and for historical (urban CPI) inflation values from the US Bureau of Labor Statistics, http://www.usinflationcalculator.com/inflation/consumer-price-index-and-annual-percent-changesfrom-1913-to-2008 (downloaded June 9, 2013))

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well beyond the 1966 level until after 1984. This late 1960s funding stall and long plateau, combined with a biomedical enterprise still expanding after two decades of consistent growth, produced some very hard years for US researchers in the mid-1970s – as indicated by unprecedented low grant success rates. The distinct existence of this interruption in funding growth for life science research is not well appreciated by historians of biomedicine and still less by other historians of science (e.g., Agar 2012, p. 435); it needs to be incorporated into our understanding of everything that happened in the life sciences around the 1970s, such as the emergence of biotech (Rasmussen 2014). Moreover, it intriguingly suggests that US biomedical research spending tracks the intensity of the Cold War conflict, escalating from around 1950 to the late 1960s and then again from the middle 1980s to the early 1990s – the periods labelled by some historians as the First and Second Cold Wars (Halliday 1983; Painter 2002). The American biomedical funding plateau corresponds to, but is 3–4 years longer on both ends than, the 1970s era of economically rational détente between these wars as usually defined. This parallel, and the possible role it suggests for biomedical research as another weapon in the Cold War(s), deserves further investigation by historians of science, medicine, and policy. As for describing the changing distribution of biomedical research effort in the United States over time, this is easily done (to a reasonable approximation) by tracking the budgetary history of the various NIH Institutes. A simple glance at Table 1 reveals, for instance, that cancer research was an especially favored area in 1960 and so too was heart disease. Both research fields remained generously supported through the subsequent two decades, while mental health research appears to have lost favor between 1960 (when its share of resources exceeded heart disease research) and 1980 (when its share was less than half). These observations are not novel, although they still do bear further investigation by historians of US science policy. However, new historical questions could be opened if we could identify in some unbiased way those cases where research areas were especially over- or underfunded. This in turn would require, for comparison, some measure of unproblematic or politically rational funding distribution (not that rational political decisions require no explanation!). To enable this, in Table 2, I compare the relative contributions to the health burden of the United States, as reflected by national mortality and morbidity statistics, of the various disease groups, with the budget allocations of the several NIH Institutes dedicated to research on cognate biomedical problems. That the burden of disease should guide biomedical research expenditure proportionately is not only almost axiomatic for health policy today but has been generally accepted for the whole of the postwar period (e.g., among the Senators in the 1948 appropriation hearings to fund the freshly established National Heart Institute, as well as the Surgeon General Leonard Scheele).3 The task of correlating causes of death and disability to NIH Institutes is not

3

Second Deficiency Appropriations Bill, Hearings Before the Subcommittee of the Committee on Appropriations, US Senate (80th Cong., 2nd sess.), on H. R. 6935 (Washington: GPO, 1948), 17 June 1948, p. 141 (testimony of Oscar Ewing and Leonard Scheele), pp. 141–143 (statement of Senator Claude Pepper).

46,862 [13.8%] 41,487 [12.2%] 34,054 [10.0%] 67,470 [19.9%]

100%a

NIAMD/ NIDDK NINDB/ NINDS NIAID/ NIAID NIMH/ ADAMHA (research) NICHD/ NICHD

Total

80.1%

208,953 [6.1%]

76,095 [7.2%]

87.8%b

341,206 [9.9%] 241,966 [7.1%] 215,364 [6.3%] 212,000 [6.2%]

D 1980 NIH Institute budget, current $1000s [% share] 999,869 [29.2%] 527,488 [15.4%]

131,761 [12.4%] 97,315 [9.2%] 97,342 [9.2%] NA

C 1970 NIH Institute budget, current $1000s [% share] 181,454 [17.1%] 160,634 [15.1%]

76.3%

442,914 [5.8%]

581,477 [7.7%] 490,409 [6.5%] 832,977 [11.0%] 727,000 [9.6%]

E 1990 NIH Institute budget, current $1000s [% share] 1,634,332 [21.6%] 1,072,354 [14.1%]

5.2%//1.9% [750–76// 740–79] 84.0%//82.1%

15.6%//20.9% [140–205] 41.5%//40.9% [400–02, 442–47, 450] 3.2%//3.7% [260, 592–94, 540–41] 11.3%//8.6% [330–34] 3.9%//3.2% [480–93] 3.3%//2.9% [581, suicide]

F % total mortality [cognate illness ICD codes], United States 1960//1980

1742 [11.5%] 4878 [32.1%]

3.2% 7.4%

58.8%

15,194

1707 [11.2%]

2643 [17.4%]

2.1%

17.2%

894 [5.9%]

1404 [9.2%]

H No. [% of included] cognate historical literature, 1990–2015, all journal 1926 [12.7%]

1.4%

13.0% (heart disease only)

14.5%

G % total morbidity, cognate illness group, United States 1981–1982, YPLL65

312

22 [7.1%]

128 [41.0%]

74 [23.7%]

19 [6.1%]

13 [4.2%]

6 [1.9%]

I No. [% of included] cognate historical literature, 1990–2015, elite journal 50 [16.0%]

a

In this calculation a value of $190 m is imputed to ADAMHA research, midway between the final NIMH appropriation of about $227 m for 1966 and the earliest ADAMHA research budget figure available to me of $153 m for 1977 (http://www.ncbi.nlm.nih.gov/books/NBK235734/table/ttt00015) b Institute budgets may not sum exactly to 100% total NIH Budget, because the latter reflects only current appropriations and also includes central administration funds, while the Institute budgets reflect commitments and may include fund transfers. Furthermore, the 1960 total budget includes NIDCR, while the 1980 and 1990 budgets also include many other Institutes. It appears that NIH expenditures exceeded appropriations slightly in 1960

NHI/ NHLBI

NCI/NCI

B 1960 NIH Institute budget, current $1000s [% share] 91,257 [26.9%] 62,237 [18.4%]

A NIH Institute 1960 name/ 1990 name

Table 2 Share of NIH budget, US disease burden, and historiography. (By Institute domain)

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especially difficult, as the names of the Institutes correlate nicely with the coding of the disease groups used by the Centers for Disease Control (for long another branch of the Public Health Service) in describing the national vital statistics. For example, of the 15 leading causes of death for 1980, the second, malignant neoplasms (ICD classification codes 140–208 on death certificates) correspond neatly to the brief of the National Cancer Institute; the third, cerebrovascular events (ICD codes 430–438, and no other neurological disease classes are listed among the top 15 causes) correspond neatly to the brief of the National Institute for Neurological Disorders and Stroke, while the first, heart diseases, combined with the fifth, chronic obstructive pulmonary disease, and the ninth, atherosclerosis (ICD codes 380–398, 402, 404–429; 490–496; and 440, respectively), correspond neatly to the brief of the National Heart, Lung and Blood Institute. As crude indicators of the mortality burden of the (psychiatric) conditions lying within the National Institute of Mental Health’s domain, I have, following others, combined suicide and liver disease, since alcoholism and other substance abuse are the latter’s major cause (morbidity is discussed separately below). Column F records the contribution to national mortality of the causes of death corresponding to NIH Institutes, and to offer some sense of how the causes of mortality are shifting through the postwar period, these are given both for 1960 and 1980. Some Institutes, such as General Medical Sciences and Dental and Craniofacial Research, do not correspond to major causes of death not otherwise recorded, but these are minor components of the NIH portfolio; indeed six of the Institutes (Cancer, Heart, Infectious Disease, Diabetes and Kidney, Neurology and Stroke, and Mental Health), account for approximately 100% of the NIH budget in 1960 and, with Childhood Development, still account for more than 75% of the budget in 1990, despite the addition of many of new Institute level budgetary units by that time and the splitting off of disease domains to new Institutes, such as the loss of Blindness from NINDS in 1968 and Arthritis from NIDDK in 1986. This analysis focuses mainly on these dominant branches of the biomedical enterprise (see Table 2, columns A–E). While mortality statistics, available through the causes of death recorded on death certificates, are a relatively accessible and politically potent measure of the relative social burdens of various diseases, a nation governed by the rationality of public health (as currently framed) would instead apportion its health research resources according to morbidity – that is, suffering and lost productivity due to illness, independent of death. Unfortunately, however, there is no single generally accepted measure of health burden. For example, the main measure used today in the United States, the DALY (disability-adjusted life year), differs from that used in much of the former British Commonwealth, the QALY (quality-adjusted life year). Even the standardized DALY measure specified by the WHO for international comparisons in recent Global Burden of Disease studies produces highly variable results (Polinder et al. 2012). Such measures also require detailed preference questionnaires, surveys, or other sociological data on particular health conditions that are not retrospectively obtainable. Indeed, morbidity measures were first explored only in the late 1930s, largely by public health advocates for increased attention to chronic diseases, and remained contentious and experimental through the 1960s (Weisz 2014). The first National Health Examination Survey, the earliest baseline against which subsequent

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US health status trends can be judged, was only conducted in 1959–1962.4 National morbidity figures have only been routinely published since 1981 and are based on a simple measure calculable from the same death certificate data as mortality, the YPLL65 (years of productive life lost before age 65; age 75 from 1996). Although it also fails to assess impaired health not associated with shortened life expectancy, YPLL does improve on raw mortality as a health burden measure by accounting for the age of victims. Because nothing better is available, and also because they represent the health policy decision base circa 1980 (the approximate midpoint of the postwar twentieth century), I present YPLL65 US morbidity figures for 1981 in column G of Table 2 as a second rough measure by which the rationality of the distribution of health research funds may be assessed. Columns H and I refer to the distribution of historical research efforts and will not be discussed until the next section, after we have concluded our analysis of the match between biomedical research effort and the burden of cognate illness categories. Taking each described budget year as a rough measure of funding distribution for the decade in which it stands as midpoint, these figures cover the period from 1955 (before which funding patterns had not stabilized) to 1995 – essentially the entire postwar twentieth century. Therefore we may draw some broad conclusions about the postwar distribution of biomedical research efforts based on the budget shares of the seven NIH Institutes described in columns A–E in Table 2, as compared with the shares in national mortality and morbidity of the corresponding illnesses in columns F–G. For example, as many historians have noted, cancer research claimed a large and fairly stable share of research resources throughout the postwar period (the dip in 1970 is more apparent than real, as it immediately preceded a major boost from the National Cancer Act of 1971). However this 27% (1960) or 29% budget (1980) share was not dramatically far out of step with the illness’s impact on health (about 16% of national mortality in 1960 and 21% of mortality in 1980 but 15% of morbidity). Heart disease also attracted a consistently large share of research resources, and likewise this large share is easily explained by the condition’s visibly large health burden. Indeed, one would be justified in asking why in 1960 (before morbidity was formally considered, and mortality dominated the epidemiological visibility of disease), heart disease research attracted only 60% of the funding won by cancer research, despite killing more than twice as many Americans. But asking why heart disease was relatively underfunded is nearly the same question as asking why cancer research attracted particular favor, because these two research areas together accounted for nearly half of the biomedical research enterprise at the time. On the other hand, research into metabolic disorders such as diabetes and kidney disease (whose chief cause is diabetes) seems to have enjoyed even greater relative favor in 1960, when the research share of the fields under NIAMD aegis was fourfold higher than their cognate mortality share, and 1980, when what had become NIDDK enjoyed a budget share two and a half times its mortality share (and seven times morbidity share, although YPLL does not capture arthritis). This analysis suggests that some historical explanation is needed to account for the flourishing 4

Data available at https://www.icpsr.umich.edu/icpsrweb/ICPSR/series/00197

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of this field, which is dedicated to the main chronic diseases of aging other than cancer and heart disease, but to my knowledge no historians have interrogated it. As I argue below, this area of biomedicine is in in fact radically understudied by historians. We might also observe that infectious disease appears to have commanded a disproportionately high share of research funding as compared with mortality and also morbidity (in 1980), but that this share was declining until the 1980s – a reversal due to an influx of new AIDS research funding, one presumes as for neurology and neurological disorders, this biomedical field seems to have attracted a share of research funding in both 1960 and 1980 that is very close to the mortality burden attributed to cerebrovascular disease at both times, which suggests both that mortality was a key determinant of health research priorities in both 1960 and 1980, as expected, and also that there is no very interesting problem to study in why this field’s research funding share declined 42% over the 20-year interval (because cognate mortality declined proportionately). Still, that morbidity was starting to play some role in attracting health research funding by 1980 is implied by the National Institute of Child Health and Development budget share of about 6%, three times the mortality contribution of congenital and early childhood illness – but still only one-third the contribution to morbidity at the time. The emerging role of formal morbidity assessment in the establishment and funding of this Institute in the early 1960s, together with the other arguments offered for funding research in perinatal medicine and childhood development, is another worthwhile but largely unexplored question for historians of biomedicine. Mental health research funding presents special problems of assessment. At face value it would seem that the National Institute of Mental Health was grossly overfunded in 1960, when it accounted for one-fifth of the total NIH budget, double the budget of the National Institute of Allergy and Infectious Diseases (whose cognate illnesses had a similar contribution to mortality) and even somewhat more than the National Heart Institute (whose disease domain claimed more than ten times the mortality share attributable to mental illness, even excluding the cardiopulmonary causes of death that did not officially fall within its portfolio before its 1969 reorganization as the NHLBI). By contrast in 1980 NIMH had seen a 70% decline in budget share or more precisely in the research budget of the mental health agency to which NIMH then belonged (and which would again spin off NIMH in 1992, along with the National Institute of Drug Abuse and National Institute on Alcohol Abuse and Alcoholism, formerly programs within NIMH), from six times the national mortality share of suicide and substance abuse to about two and a half times that mortality share. It would seem that this branch of research had lost the political favor it had once enjoyed in the 1950s and early 1960s and by 1980 was attracting research effort more proportional to mental health’s contribution to the national health burden. Advocates of mental health research (and psychiatry in general) have long argued, of course, that the health burden of mental illness far exceeds what is visible in mortality-based statistics; indeed, at the height of early postwar enthusiasm for all things psychiatric, Surgeon General Parran urged Congress to establish the NIMH on the grounds that “mental diseases equal all physical diseases in subtracting from the total vigor, the total fitness of our population” (Grob 1996). Yet as noted earlier, no national morbidity data were available at the time to support such a claim, although it was a widely recognized problem that around half of the national

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hospital beds were occupied by long-term mental health patients. Even today the measures of health burden that elevate the morbidity profile of mental health, and have been criticized for exaggerating its contribution to total burden such that antidepressant drugs may be displacing more cost-effective public health interventions, still tend to place anxiety and depression in fourth place, far behind heart disease and at about the same level as metabolic disorders (in developed countries) (Brhlikova et al. 2011; Polinder et al. 2012; Bemme and D’Souza 2014). In sum, historians have good reason to investigate why mental health attracted so much cultural attention and political favor in early postwar United States. They also have good reason to ask how and why the field lost favor in the late 1960s and 1970s, a question much less thoroughly addressed. To sum up, in this section we have examined the shifting distribution of biomedical research funding, and therefore (approximately) research effort, among the major fields of biomedicine, and also the relationship of research effort to national burden of disease. This comparative exercise has been fruitful in raising new hypotheses and problems for investigation by historians of biomedical science as well as US research policy. We have confirmed the observation of many historians that cancer appears to have attracted disproportional support. We have raised the more novel question of why heart disease, responsible for three times as many deaths, never achieved the same national attention and resources. We have pointed out that the cluster of diseases associated with the other main chronic diseases of aging (diabetes, kidney disease, and arthritis) appear to have, on the other hand, garnered a disproportionately great share of resources and suggested this as an area for historical investigation. We have also suggested that a similar, large overinvestment in mental health research occurred in the early postwar period, as well dramatic decline by the end of the 1960s, both meriting further historical investigation. We have noted that funding for infectious disease research was disproportionately large in the postwar period and that its share was in decline until the 1980s, suggesting that historians might investigate the mobilization of resources in reaction to AIDS to explain this reversal. We have also suggested that the emerging role of morbidity burden, as opposed to mortality, as a metric used in research policy decision-making from the 1970s, is another topic deserving historical investigation.

The Distribution of Activity in Biomedicine and in Its Historiography Much as we can compare the budget shares of various NIH research areas to the shares of national morbidity and mortality claimed by cognate disease groups, in order to identify important issues for the history of health research policy, we can compare NIH research area budget shares to the share of the historical literature devoted to biomedical activity in the same cognate disease groups. That is, we may ask about the distribution of the efforts of historians of biomedicine – historiographic “mindshare” (if I may) – as compared with the distribution of past activity within the biomedical enterprise itself. Taking the share of the budget claimed by each NIH Institute as a tangible (if still approximate) measure of postwar American biomedical research activity in that scientific domain is not hard to defend, given (1) the way NIH dwarfed by an order of magnitude other sponsors in the

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main biomedical research fields it funded and (2) the indisputable premise that research activity within modern biomedicine correlates (as a first approximation anyway) with the quantity of money in a field available to pay for salaries, facilities, supplies, and graduate fellowships. This measure can be extrapolated internationally to some extent – albeit one not assessable on the basis of current historical literature – given that US biomedical research spending throughout the postwar twentieth century represented the bulk of world spending (as noted above). The other side of this comparison, measuring the distribution of historians’ efforts among biomedical research fields, is a more complex matter and one without much precedent in the historiography. However, systematic literature reviews have become a commonplace method in biomedical research in recent years. I have adopted such an approach to study the historiography of biomedicine, inspired not just by the attractions of reflexivity but also by the impressive body of “evidence-based medicine” studies that assess questions of bias in the clinical trial literature (e.g., the degree to which, ceteris paribus, published clinical trials sponsored by the manufacturer of a novel drug more often report significant benefits for the new drug as compared with independent trials) (Lundh et al. 2012). Mine can be thought of as an analogous project to measure bias in historical literature. The foundation of any systematic review is an extensive and representative database of the relevant field of literature, together with a sampling method that is unbiased with respect to the question at hand. To identify recent historical publications on biomedical research fields, in June 2015 I searched the EBSCO History of Science, Technology and Medicine database established by the History of Science Society and Wellcome Trust, limiting searches to journal articles published since January 1, 1990 (mainly because before this date journal coverage in the database is very incomplete and also because little of the postwar period was remote enough to be treated as history before the 1990s). The search terms, allowed anywhere in the bibliographic entry, were “Research” and any of five other terms distinguishing the NIH research area in question. These five search terms were derived iteratively from the mission statement on each relevant NIH Institute web page, exploring close cognates and variants to achieve the maximum number of total hits for the compound search. British and American spelling variants were both used, together counting as only one of the five search terms; for example, historiography concerning the National Cancer Institute’s research domain was identified with search terms Research AND Cancer OR Neoplasms OR Tumor OR Tumour OR Oncology OR Hematology OR Haematology, while that of the National Heart (Lung and Blood) Institute was identified with terms Research AND Cardiology OR Cardiovascular OR Cardiac OR Cardiopulmonary OR Heart.5 Boolean mode was employed, no language was specified (although the database is mainly Anglophone),

5

The other search term clusters were as follows: Mental Health, Research AND Psychiatry OR Behaviour OR Behaviour OR Mental Health OR Psychopharmacology OR Insanity; Kidney and Metabolic Disease, Research AND Diabetes OR Kidney OR Metabolic OR Gastrointestinal OR Arthritis; Infectious Disease, Research AND Immunology OR Bacteriology OR Virology OR Parasitology OR Allergy; Neurology, Research AND Neurology OR Brain OR Stroke OR Spinal OR Cerebral; Childhood Disease, Research AND Childhood OR Neonatal OR Pediatric OR Pediatric OR Congenital OR Embryology.

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and the automatic related term search function was deactivated. The number of hits for each Institute-cognate biomedical research domain is given in column H of Table 2, together with percent of total hits in searched domains – although the latter should not be regarded as a reliable measure of the total literature because the database contains many duplicate entries, and there was no effort to restrict hits to sources actually dealing with biology or medicine. It must also be considered that at least as many of the historical items retrieved in these searches were short retrospective items published in biomedical research journals, typically written by practitioners in these fields, as were scholarly articles by professional historians. While such auto-historiography of biomedicine merits investigation in its own right, it lies beyond the scope of the present review. To assess biomedical research area mindshare in the professional research literature for history of science and medicine, I searched within the total hits retrieved above by limiting results to a subset of elite scholarly journals. This elite subset, intended only as an unbiased sample of the professional historiography, was derived from the journals ranked in the top half (of 59) categorized in the field history and philosophy of science, according to the 2013 Thomson-Reuters Web of Science Journal Citation Index (which was the latest available at the time). Only regularly publishing journals with a scope including all of biology or medicine were included (thus excluding Journal of the History of Neurosciences, as well as Annals of the History of Computing) and also specifying history as their central purpose (thus excluding the American Journal of Bioethics and, perhaps regrettably, Social Studies of Science). While somewhat arbitrary, the eight included journals do represent a plausible sample of the professional historiography of biomedicine: Bulletin of the History of Medicine, British Journal for History of Science, Isis, Journal of the History of Biology, Journal of the History of Medicine and Allied Sciences, Medical History, Social History of Medicine, and Studies in History & Philosophy of Science (Part C). This elite subset happened, conveniently, to be picked out by limiting the much larger overall search results in each domain to items with the term “history” in the publication title/subtitle; these results then were manually searched. Articles specified as book reviews or introductions (with no further designation of historical subject matter) were excluded and also any item shorter than five pages. All duplicate records were eliminated. No article was included unless it specified “20th century” in its keywords or clearly referred to events in the twentieth century in its title or keywords (excluding birth and death dates). In the very few cases that an article was listed in multiple research domain searches, it was allowed to count for each. The number of hits for each Institute-cognate biomedical research domain in the elite journals is given in column I of Table 2, together with the percentage share of total elite items identified in all domain searches. A number of conclusions emerge concerning the distribution of effort in the twentieth-century biomedical historiography as compared with distribution of research effort within postwar biomedicine itself. Restricting myself to cases of pronounced mismatch, where the share of historical attention is more than double or less than half the share of a biomedical field’s research activity in the past, these include the following. First, the historiography of biomedicine pays attention to

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psychiatry and related fields to a degree far exceeding their importance in past biomedicine. Even compared with the early postwar period, when the biomedical fields associated with the National Institute of Mental Health exhibited special dynamism thanks to an excessive (I have suggested) 20% share of health research funding, the historiography redoubles this favor with a surprising 42% of the elite journal literature. That historians pay special attention to mental health research of the past may owe something to the relative accessibility of the field’s content to scholars trained in the humanities or to the closeness of the field’s content to themes subsequently fashionable in the humanities (e.g., social control, deviance, race, gender construction). Probably both. Explaining such disproportionate attention among historians is an intriguing topic for future reflective research, although not as important in my view as correcting it with new empirical work elsewhere. Second, and similarly, the historiography of biomedicine pays exaggerated attention to fields associated with infectious disease. Microbiology at the end of nineteenth century could not be more worthy of historical attention. As historians have emphasized time and again, the successes of this field of lab research not only revolutionized thinking and practice in medicine but also catalyzed the wider convergence of the two as biomedicine during the middle third of the twentieth century. However, the bacteriological revolution was already victorious by the first decades of the twentieth century; for example, between 1900 and 1938, the US national death rate from tuberculosis declined by three quarters and that from pneumonia two-thirds, before any impact attributable to antibiotics.6 That infectious disease research, including virology and allergy as well as bacteriology, still accounted for a much greater share of the postwar US biomedical research budget than can be justified by its contribution to national mortality can be plausibly explained by a number of mutually reinforcing explanations. For example, the penicillin-driven retreat of bacterial infection as a feared cause of death after the Second World War inspired ambitions that all remaining infectious diseases, such as those due to viruses, could similarly be cured (and not just prevented). Also polio, that dreaded scourge of children, was almost eliminated from American experience, practically overnight. At the same time, the simplicity of bacteria and viruses made them attractive experimental objects for more basic disciplines like biochemistry and the emerging, glamorous field of molecular biology, which achieved some spectacular scientific successes. Further, immune disorders became an intellectually fashionable field, even if not responsible for a large burden of disease – partly due to metaphoric relevance to Cold War problems of political control, according to some historians (Rasmussen 1993; Anderson and Mackay 2014). But a plausible hypothesis about why historians should redouble past popular and political enthusiasms for microbiology-related research is less

6

Forrest Linder and Robert Grove, Vital Statistics Rates in the United States, 1900–1940 (Federal Security Agency; GPO 1947), available at http://www.cdc.gov/nchs/data/vsus/vsrates1900_40.pdf; accessed 2 July 2015); Mortality Tables: Table 12.

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obvious.7 Perhaps some of this interest is driven by the recognition of historians in the late twentieth century that the period of freedom from fear of infectious disease was transitory, since it would soon return as a prominent problem in the developed world (e.g., with AIDS and mounting bacterial drug resistance), while less developed countries never achieved as much benefit from postwar advances. The roots of pressing social and medical problems of the present ought quite properly to attract extra attention from historians. Another, somewhat less charitable explanation is historiographic momentum: topics already attracting attention from historians tend to exert a continuing attraction to historians, because training and previous work by established historians instructs budding historians and shapes their consciousness of the past. Taken too far, this process can be likened to mapmakers relying only on old maps to draw new ones, as in Scholastic cartography. Third, the biomedical research fields centered on heart disease, corresponding to the NHLBI’s remit, are drastically understudied relative to their importance as activities within postwar biomedical research. At first thought, it seems hard to explain the relative neglect of heart disease by historians since it was an area of research that not only expanded so much in the postwar era that it soon rivalled cancer research in terms of resource allocation but also attracted enormous glamor, for example, in spectacular cardiac surgery techniques and blockbuster pharmaceuticals modifying the heart’s action (e.g., beta blockers) and blood properties (e.g., pressure, clotting, fat profile). Here perhaps the likeliest explanation is a type of historical accident that we may call “inertia,” the flip side of what I called “momentum” above: historians often take up subjects close to their doctoral supervisors’ expertise, but those few who have studied heart disease have for whatever reason trained fewer students who went on in the field. And why were there few senior historians of medicine in the heart disease field to begin with? Here it may be that the relative neglect of heart disease within medical research before the Second World War has had a follow on effect upon historians of medicine. Thus, in the 1960s and 1970s, there was as yet not much heart disease research old enough to qualify as history – certainly not much compared with microbiology, still in its heyday with the afterglow from penicillin, the polio vaccine, and earlier conquests of infectious disease (and long established as historical genre of the Microbe Hunters ilk). The 1970s were when many of the senior generation of historians of science and medicine, now in or entering retirement, wrote their dissertations – and the period when the field was expanding and many academic positions were filled at research universities. Thus I would suggest in the realm of our disciplinary ecology what population biologists call a “founder effect,” perhaps coupled with incidental causes 7

It might be supposed that historians have mostly attended to the early twentieth-century golden age of bacteriology, when it achieved its most dramatic successes. However, it seems this is not the case: the majority (38/74) of elite journal articles returned by the searches in the review dealt substantially with events after 1945. My takeaway impression is that the center of gravity of the retrieved literature in this area lies in the 1930s and is motivated by questions about how postwar biomedicine took the shape it did.

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that propagated rather than corrected the historiographic neglect of the heart over time. Fourth, the fields of metabolic disorders, kidney disease, and diabetes, corresponding to the NDDK’s remit, are also drastically understudied relative to their importance as activities within postwar biomedical research. In the case of kidney disease and metabolic disorders (a cluster of conditions associated with chronic diseases of aging), we might venture that, in addition to the founder effect and inertia among historians just discussed, there may operate another effect based in professional formation. Just as historians may seek to share in the glory of especially glamorous past events by becoming authorities on them, they may fear obscurity by studying areas of science and medicine forgotten by the public and neglected by fellow historians (whose opinion, after all, determines the bulk of professional reward). This last hypothesis, attraction to glamorous and still memorialized past events, may of course apply to any marked case of selective historical attention and historical neglect, not just infectious disease and kidney disease. Why the introduction of the polio vaccine drew so much more acclaim in the past than renal dialysis, and left a more lasting aura of glamor, is another intriguing question worthy of historical investigation. Here we may begin to consider the role of professional historians in managing society’s official memories of science and medicine, sometimes unthinkingly accepting a script written by past journalists as a starting point. But we can only ask such questions, which arise from comparing public and professional commemoration (both past and present) with some independent measure of importance, if we stray from the well-worn paths left by the existing historiography and look afresh at the landscape we purport to be describing.

Conclusion There is nothing inherently wrong with historians researching the topics most frequently studied by other historians. To some extent this is inevitable; as just noted, young historians often adopt the study areas of their mentors as part of their apprenticeship, giving rise to momentum (and, as a side effect, inertia). Furthermore it is vital to the spirit of historical scholarship that the stories arising from existing historical writings, popular and professional, be subject to scrutiny and evidencebased reinterpretation – “revisionism,” to those uncomfortable with scrutiny of convenient myths. However, there is something wrong if we do nothing more, treading ever-deeper historiographic ruts fixed on our crude inherited maps, while oblivious to vast, largely unexplored parts of past biomedical terrain lying nearby. It is not my intention to suggest that too much history of microbiology and history of psychiatry has already been done, and sufficient history of cancer research. The amount of biomedical activity in the recent past so vastly dwarfs the activity of its historians – think of the ratio of living biomedical researchers to historians of biomedicine! – that even on many relatively well-studied topics we have only scratched the surface. Even so, I do hope that the large lacunae on our maps of the

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past, highlighted in this review, will tempt some adventurous historians of biomedicine into less explored territory.

References Agar J (2012) Science in the 20th century and beyond. Polity, London Anderson W, Mackay IR (2014) Intolerant bodies: a short history of autoimmunity. Johns Hopkins University Press., Baltimore Appel T (2003) Shaping biology: the National Science Foundation and American biological research, 1945–1975. Johns Hopkins University Press, Baltimore Bemme D, D’Souza NA (2014) Global mental health and its discontents: an inquiry into the making of global and local scale. Transcult Psychiatry 51:850–887 Bero LA (2005) Tobacco industry manipulation of research. Public Health Rep 120:200–208 Bloom FE, Randolph MA (eds) (1990) Funding health sciences research: a strategy to restore balance. National Academies Press, Washington, DC. Available at http://www.ncbi.nlm.nih. gov/books/NBK235742/ Boychuk G (2008) National health insurance in the United States and Canada: race, territory, and the roots of difference. Georgetown University Press, Washington, DC Brandt AM (2012) Inventing conflicts of interest: a history of tobacco industry tactics. Am J Public Health 102:63–71 Brhlikova P, Pollock AM, Manners R (2011) Global burden of disease estimates of depression–how reliable is the epidemiological evidence? J R Soc Med 104:25–34 Brown R (1979) Rockefeller medicine men: medicine and capitalism in America. University of California Press, Berkeley Clarke AE, Mamo L, Fosket JR, Fishman JR, Shim JK (2009) Biomedicalization: technoscience, health, and illness in the U. S. Duke University Press, Durham Forman P (1992) Inventing the maser in postwar America. Osiris 7:105–134 Fox DM (1987) The politics of the NIH extramural program, 1937–1950. J Hist Med Allied Sci 42:447–466 Fuller S (2000) Thomas Kuhn: a philosophical history for our times. University of Chicago Press, Chicago Gaudillière J-P (2002) Inventer la biomedicine: La France, l’Amérique et la production des savoirs du vivant (1945–1965). Éditions la Découverte, Paris Geiger R (2008) Research and relevant knowledge: American research universities since World War II. Transaction, New Brunswick Grob G (1996) Creation of the National Institute of Mental Health. Public Health Rep 111:378–381 Halliday F (1983) The making of the Second Cold War. Verso, London Hamowy R (2008) Government and public health in America. Edward Elgar, Northampton Hollinger DA (1995) Science as a weapon in Kulturkampfe in the United States during and after World War II. Isis 86:440–454 Kevles DJ (1977) The National Science Foundation and the debate over postwar research policy, 1942–1945: a political interpretation of science – the endless frontier. Isis 68:5–26 Kleinman D (1994) Layers of interests, layers of influence: business and the genesis of the national science foundation. Sci Technol Hum Values 19:259–282 Kohler R (1991) Partners in science: foundations and natural scientists, 1900–1945. University of Chicago Press, Chicago Krementsov N (2002) The cure: a story of cancer and politics from the annals of the Cold War. University of Chicago Press, Chicago Krige J (2006) American hegemony and the postwar reconstruction of science in Europe. MIT Press, Cambridge, MA

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Ludmerer K (1985) Learning to heal: the development of American medical education. Johns Hopkins University Press, Baltimore Lundh A, Sismondo S, Lexchin J, Busuioc O, Bero L (2012) Industry sponsorship and research outcome. Cochrane Database Syst Rev (12):Art. No.: MR000033. https://doi.org/10.1002/ 14651858.MR000033.pub2 Mayes R (2004) Universal coverage: the elusive quest for national health insurance. University of Michigan Press, Ann Arbor Moses H 3rd, Matheson D, Cairns-Smith S, George B, Palisch C, Ray Dorsey E (2015) The anatomy of medical research: US and international comparisons. JAMA 313:174–189 Osgood K (2006) Total Cold War: Eisenhower’s secret propaganda battle at home and abroad. University of Kansas Press, Lawrence Painter D (2002) The Cold War: an international history. Routledge, London Poen MM (1996) Harry S. Truman versus the medical lobby: the genesis of medicare. University of Missouri Press, Columbia Polinder S, Haagsma JA, Stein C, Havelaar AH (2012) Systematic review of general burden of disease studies using disability-adjusted life years. Popul Health Metrics 10:21. https://doi.org/ 10.1186/1478-7954-10-21 Porter R (1998) The greatest benefit to mankind: a medical history of humanity. Norton, New York Proctor R (2011) Golden holocaust: origins of the cigarette catastrophe and the case for abolition. University of California Press, Berkeley Rabinow P, Rose N (2006) Biopower today. BioSocieties 1:195–217 Rasmussen N (1993) Freund’s adjuvant and the realization of questions in postwar immunology. Hist Stud Phys Biol Sci 23:337–366 Rasmussen N (2014) Gene jockeys: life science and the rise of biotech enterprise. Johns Hopkins University Press, Baltimore Rothstein W (1987) American medical schools and the practice of medicine : a history. Oxford University Press, New York Saunders FS (2000) Who paid the piper?: the CIA and the cultural cold war. Granta, London Shapin S (2008) The scientific life: a moral history of a late modern vocation. University of Chicago Press, Chicago Shyrock R (1980) American medical research past and present. Arno, New York Starr P (1982) The social transformation of American medicine. Basic, New York Strickland S (1972) Politics, science, and dread disease: a short history of United States medical research policy. Harvard University Press, Cambridge, MA Strickland S (1989) The story of the NIH grants programs. University Press of America, Lanham Sunder Rajan K (2006) Biocapital: the constitution of postgenomic life. Duke University Press, Durham Vogel MJ, Rosenberg C (eds) (1979) The therapeutic revolution: essays in the social history of American medicine. University of Pennsylvania Press, Philadelphia Wiesz G (2014) Chronic disease in the twentieth century a history. Johns Hopkins University Press, Baltimore Wilford H (2009) The mighty Wurlitzer: how the CIA played America. Harvard University Press, Cambridge, MA Yi D (2015) The recombinant university: genetic engineering and the emergence of Stanford biotechnology. University of Chicago Press, Chicago

The Historiography of Biotechnology

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Contents Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Popular Conceptions of Biotechnology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . First-Generation Scholarship: Understanding the Biotech Revolution . . . . . . . . . . . . . . . . . . . . . . . . . Second-Generation Histories: Broadening Scope and New Definitions . . . . . . . . . . . . . . . . . . . . . . . Third-Generation Histories: Revisionist Histories . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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Abstract

Compared to the vast literature on the history of biology as a whole, scholarship on what we might term the “history of biotechnology” has only recently arrived in the past 30 years as historians have become interested in the field. Although scholars have studied the history of biotechnology for only a short length of time when contrasted with subjects such as Charles Darwin or genetics, histories of biotechnology have changed and diversified both in approaches and topics since biotechnology became a choice of focused study for historians in the early 1980s. Since that time, it has become a robust field of scholarly activity and promises to be an important part of the history of science, technology, and medicine in the future. In this chapter, I provide a structure for understanding the progression of these histories from the beginnings of the scholarly engagement in the early 1980s through the present.

N. Crowe (*) History Department, University of North Carolina Wilmington, Wilmington, NC, USA e-mail: [email protected] # Springer Nature Switzerland AG 2021 M. R. Dietrich et al. (eds.), Handbook of the Historiography of Biology, Historiographies of Science, https://doi.org/10.1007/978-3-319-90119-0_13

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Introduction Compared to the vast literature on the history of biology as a whole, scholarship on what we might term the “history of biotechnology” has only recently arrived in the past 30 years as historians have become interested in the field. Although scholars have studied the history of biotechnology for only a short length of time when contrasted with subjects such as Charles Darwin or genetics, histories of biotechnology have changed and diversified both in approaches and topics since biotechnology became a choice of focused study for historians in the early 1980s. Since that time, it has become a robust field of scholarly activity and promises to be an important part of the history of science, technology, and medicine in the future. In this essay, I hope to provide the reader with a broad overview of the literature by articulating the topics and ideas to which scholars have paid most attention and the intellectual debates that have emerged from their works. Normally, in an essay about biotechnology, one might expect that a clear definition of biotechnology would appear in the introduction. However, because of the variety of ways that historians and other scholars have defined the subject, providing a definition of biotechnology in a historiographical essay would be less than helpful. In fact, I would argue that how authors have defined biotechnology, either explicitly or implicitly, is one of the more distinctive features of the literature. A primary example, for instance, is how historians have given primacy to recombinant DNA technologies, which dominated definitions of biotechnology since its rise in the 1970s or whether the history of biotechnology should instead deal with much longer periods and include a broader array of materials and ideas. Because of how varied the use of term “biotechnology” has been, one of the central themes of this essay will concern how historians’ definitions of biotechnology have been contested, reimagined, and enlarged over the past 30 years. Researchers have been drawn to various biotechnologies and emphasized different aspects of them in order to explain how biotechnologies have impacted our modern world. They have shown how biotechnologies have redefined conceptions of life; restructured the practice, organization, and business of science; and provided a window into politics, society, and pertinent institutions. While some have delved deeply into the scientific and technological details, others have been drawn to the various cultural, legal, and ethical definitions and debates that biotechnologies have engendered. Not surprisingly, what scholars have been attracted to over the past 30 years of scholarship has often been a reflection of their contemporary perspective. I have organized this essay by grouping the historiography of biotechnology into three successive generations, which, in part, was a choice motivated by the larger motifs that I believe bind the scholarship of each generation. The first generation of scholars, which emerged in the mid-1980s, was directly responding to a growing fascination with biotechnologies, particularly recombinant DNA, in both science and the public at large. Some of these early scholars created narratives that saw the development of recombinant DNA as a significant break from what had come before, producing a whole host of distinct changes to the practices and institutions of science. Others sought to place recombinant DNA and its effects into larger

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narratives, seeing recombinant DNA as an apex of a story that has long, unbroken roots as far back as the nineteenth century, if not earlier. A second generation of scholarship on the history of biotechnology arose in the late 1990s and was characterized, in part, by skepticism toward the exceptionality of the biotechnology revolution in the late 1970s and early 1980s. Unlike some firstgeneration scholars who sought to argue that the advances of the 1970s were simply recent nodes along a specific long-standing narrative, second-generation scholars tended to argue that what observers in the 1980s had originally described as novel outcomes from recombinant DNA’s discovery could be found in multiple places within the history of science and technology. In practice, this meant that secondgeneration scholars articulated many episodes throughout the twentieth century that looked similar to what supposedly made post-1970s biotechnology unique or exceptional, effectively undermining arguments as to the necessity for treating the history of biotechnology as something different from the history of science and technology generally. Additionally, second-generation scholars, who were influenced by neighboring fields, such as sociology and anthropology, and were skeptical about equating biotechnology with recombinant DNA, appended several new topics to the biotechnology rubric. Consequently, by the middle of the 2000s, historians had applied to the term biotechnology to a wide array of times, places, and topics. By the end of the first decade of the twenty-first century, a third generation of scholars emerged with a renewed focus on biotechnology in the 1970s and 1980s. The historical accounts created by the second-generation scholars and a quarter-century of perspective motivated this third generation of historians to revisit the development and impact of recombinant DNA. These revisionist histories focused on the larger contexts of the period, which had become more visible given the improved perspective. Seen as neither revolutionary nor ordinary, revisionist historians treated the 1970s and 1980s as a distinct historical period in a way that previous historians had never done, though they did disagree on what they thought characterized the era. Greater perspective on the historical trends in biotechnology also inspired historians to engage in a number of new topics, which were sometimes motivated by new sources that helped articulate previously underdeveloped subjects and sometimes motivated by efforts to historicize the biotechnologies that were (and still are) emerging. This periodization of the historiography of biotechnology, of course, is not seamless. The dividing lines between generations are not well-defined; some historical accounts might feel pioneering and belong better in later generations, whereas others may seem connected to earlier historiographic periods. Additionally, there are issues that have motivated scholars throughout each generation rather than being indicative of a particular era. For instance, questions surrounding biotechnologies and the application and impacts of intellectual property rights have inspired scholars since the 1980s and still continue to be a significant topic for historians.1 In this essay I will primarily deal with historical treatments of biotechnology, that is, those

For instance, the 2015 History of Science Society meeting featured “Roundtable: The New Historiography of Science, Technology, and Intellectual Property Law,” History of Science Society Annual Meeting, November 19–22, 2015, San Francisco, California

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narratives that explicitly seek to understand issues of change over time. However, this leaves out a substantial literature by scholars interested in understanding issues surrounding biotechnology that are not primarily driven by historical questions. Nevertheless, these works, and particularly their themes, have become important to historians, and I will spend some time discussing them when appropriate. Note, however, that this will be far from an exhaustive discussion of the literature but rather a recognition of their importance. Before I begin a tour of the first-generation biotechnology histories, it is important to recognize that non-historians have written about biotechnologies for a long time, and still do. Much of this writing, which includes everything from science fiction to popular science writing, has been for a general audience. These writings are certainly important in a broad sense to the historiography of biotechnology as they have helped shape larger cultural and social attitudes about the value of biotechnologies. Undoubtedly, many of these views have helped inspire scholarly interest in the history of biotechnology, and it is worth articulating some of the larger trends in these genres before we get started with discussions of academic treatments of biotechnology.

Popular Conceptions of Biotechnology At the end of the 1970s, there was a dramatic increase in the usage of the words “biotechnology” and “biotechnology revolution,” which carried forward throughout the 1980s and 1990s. One way to interpret this explosion in the use of these words is to presume that the development of recombinant DNA created a new fascination with the products, processes, and directions of the biological sciences.2 That, however, is untrue. Though often not labeled as “biotechnologies” before the late 1970s, writers from a multitude of backgrounds had been exploring issues in and around what would later be labeled “biotechnology,” including genetic engineering and cloning, for decades before recombinant DNA entered the scene. Often intended to reach large public audiences, these writings have inspired many of the reactions to emerging biotechnologies over the years, and they can be broken down into two general categories. The first is that which resides in the realm of literature and science fiction. The second involves what might broadly be labeled as science writing, which became increasingly focused on the biosciences after World War II.

The easiest way to see the growth of the terms “biotechnology” and “biotechnology revolution” is to track their usage within the English language using Google’s “ngram” viewer, which inspects millions of digitized materials. For further reading about google ngram data, see Jean-Baptiste Michel et al., “Quantitative Analysis of Culture Using Millions of Digitized Books,” Science 331, no. 6014 (January 14, 2011): 176–82 and Yuri Lin et al., “Syntactic Annotations for the Google Books Ngram Corpus,” in Proceedings of the ACL 2012 System Demonstrations (Jeju Island, Korea: Association for Computational Linguistics, 2012), 169–74, http://www.aclweb.org/ anthology/P12-3029

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The number of novels (and later movies) employing various aspects of what might be called biotechnologies is too numerous to discuss here, but three, in particular, are important because of how often they are invoked by scholars and non-scholars alike when talking about biotechnology. The themes and ideas presented in Mary Shelley’s Frankenstein; or the Modern Prometheus (1818), H.G. Wells’ The Island of Dr. Moreau (1896), and Aldous Huxley’s Brave New World (1932) consistently appear in discussions about the implications of biotechnology. These fictional stories warn of hubris, unintended consequences, and the startling effects of “playing God,” as it has been often put. With the creation and use of the atomic bomb during World War II, the fears of what scientists could create, the intentions behind the work, and the consequences of their actions only reinforced many of the fictional tropes articulated by Shelley, Wells, and Huxley. And since these books were widely understood in the Western world to be important literary works, they were often assigned reading during the course of one’s education, making them part of a shared literary canon that many could invoke. When people wanted to talk about the implications of cutting-edge biology in the popular press, including recombinant DNA debates of the 1970s, the names of Victor Frankenstein and Dr. Moreau and the visions of Huxley’s dystopian future were consistently applied to help describe potential dangers of these biological advances.3 The other category of biotechnology writing worth discussing is what might best be labeled as science writing. As biology emerged in the 1950s as the next great science (physics having dominated the first half of the twentieth century), many biologists and physicians became important public figures, playing the roles of popularizers of current biomedical research and advertiser for what the biosciences would soon be able to do. For example, Hermann J. Muller, Joshua Lederberg, and Peter Medawar, three Nobel Prize winners, spoke passionately about what biology and medicine could already do and what these fields would soon accomplish. In 1959, for instance, Medawar gave a series of radio lectures for the British Broadcasting Company on the “Future of Man” that became compiled into a widely read book soon afterward. Similarly, Muller and Lederberg both argued that the new biology developed after World War II would provide many solutions to the world’s problems, though they did acknowledge that these solutions could radically transform society.4

You can also see these fictional tropes invoked in other controversies, like those associated with organ transplantation in the 1960s and cloning and in vitro fertilization the 1970s, among many other controversies throughout the twentieth century. As just one pertinent example, in his article about recombinant DNA patents, Daniel Kevles included a 1980 political cartoon that depicted Dr. Frankenstein with his creature standing outside a U.S. Patent Office, see Daniel J. Kevles, “Ananda Chakrabarty Wins a Patent: Biotechnology, Law, and Society, 1972–1980,” Historical Studies in the Physical and Biological Sciences 25 (January 1, 1994): 134) 4 Examples of these types of writings can be found in the conference proceedings of the 1962 conference “Man and his Future” – a name inspired by Medawar’s work – sponsored by the Ciba Institute, which featured multiple Noble Prize winners and famous public intellectuals. Gordon Wolstenholme, Ed., Man and His Future (Churchill; Ciba Foundation, 1963) 3

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In the mid-1960s, the new biology that Lederberg, Muller, and Medawar espoused as transformative attracted attention of journalists, public intellectuals, and scholars interested in discussing the technological possibilities emerging during this period. Many of these authors reinforced the idea that science and society were in the midst of a revolution due to what was being done in the biology laboratory. Authors like Gordon Rattray Taylor in The Biological Time Bomb (1968) and Albert Rosenfield in Second Genesis: The Coming Control of Life (1969) played the role of heralds of the future, writing for a public audience about the possibilities of genetic engineering, in vitro fertilization, and organ transplantations. Lederberg himself wrote a weekly syndicated newspaper column from 1966 to 1970 entitled “Science and Man” in which he discussed for a broad audience the impact on society of new and impending scientific advances. These authors generally wrote with awe about what the biosciences seemed capable of accomplishing, often raising ethical questions in the process but rarely articulating comprehensive arguments about whether the new technologies should be embraced or embargoed. Some of these discussions about what biology would be able to do in the future inspired theologians and philosophers to articulate ethical platforms regarding advances in the life sciences. In the 1970s, for instance, theologian Paul Ramsey laid out several ethical arguments against many new biotechnical interventions which were being discussed at the time, but had yet to become a reality, such as genetic engineering, reproductive technologies, and cloning (Ramsey 1970). In contrast, Joseph Fletcher, who like Ramsey was an active member in the newly formed bioethics community, defended the application of genetic engineering and cloning on humans (Fletcher 1974).5 The number of books aimed at the public about biotechnology proliferated over the course of the 1980s, 1990s, and early 2000s. Beyond the advent of recombinant DNA, the Human Genome Project, the development of genetically modified organisms (GMOs), the creation of a cloned sheep named Dolly, and the rise of stem cell technologies, just to name a few major topics, inspired scores of books that looked at the promises and perils of these new technologies.6 New critics arose, such as Jeremy Rifkin, who made his name by advocating for the complete abandonment of research programs focused on biotechnologies like recombinant DNA (Howard and Rifkin 1977; Rifkin 1984), and Vandana Shiva (1991; 1997; 2001), who saw modern agricultural practices and their products as problematic for both the planet and a variety of non-Western societies. The counterparts to these critics also flourished since the 1970s, writing many works that, like Lederberg and Muller in the 1950s and 1960s, shared with the public a real excitement about the

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A good discussion of the various intellectual debates surrounding genetic engineering and its related sciences from this period through the 1990s can be found in John Evans’ Playing God?: Human Genetic Engineering and the Rationalization of Public Bioethical Debate (Chicago: University of Chicago Press, 2002) 6 For a small sample, see Cook-Deegan 1995; Kolata 1998; Bowring 2003; Druker 2015; Knoepfler 2015

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advances in the biosciences and optimism about how they could solve many of the world’s problems (given the proper support, of course).7 One thing that helps distinguish many of these works compared to the scholarship that I will discuss in the next section is that their intentions are to largely act as heralds, informing the broader public of the effects and implications, good or evil, of biotechnologies. In general, they are mostly ahistorical, providing little to no historical analysis despite the fact that they often include histories to help readers understand how the contemporary work came to be. Overall, these books fill a role that is much closer to commentary than histories as they generally aim to inform the larger public about contemporary issues rather than elucidate the past. Consequently, for historians today the significant number of books about biotechnology that have appeared over the decades should be seen much more as primary sources than anything else, as they can be read as voices in the debates and definitions that have surrounded biotechnologies throughout the twentieth century.

First-Generation Scholarship: Understanding the Biotech Revolution Beginning in the early 1980s, a first generation of scholarship appeared that examined the historical development of biotechnology, and what effects these biotechnologies had wrought on society. At this time, however, for much of the Western world, and particularly the United States, biotechnology had become synonymous with recombinant DNA techniques and its products. The mid- and late 1970s had seen a storm of controversy surrounding the use and application of recombinant DNA. There were condemnations of its continued development from not only distrustful groups within society but scientists themselves also debated the relative safety of recombinant DNA technologies. Scientists held closed-door meetings to examine the issue, the most famous being the 1975 Asilomar Conference, congress considered legislation to regulate the work, and cities considered banning recombinant DNA research altogether. On top of these ethical remonstrations, recombinant DNA also seemed to spark a new sector of the American economy, with the first company, Genentech, receiving one of the most astounding initial public offerings in US history when it went public in October 1980, as its first-day increase in share price set a record for Wall Street at the time (Hughes 2011, p. 158). Overall, by the early 1980s, recombinant DNA was being marketed as a revolutionary development for everyone: scientists, university administrators, entrepreneurs, drug developers, and patients to name only the ones most directly involved. Not surprisingly, the earliest histories of biotechnology, which began to emerge in the early 1980s, were a reflection of the zeitgeist of the period, when 7

The number of books that could be cited here is immense, but good examples since 2010 include Collins (2010), Wohlsen (2011), Church and Regis (2012), and Venter (2014)

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society had become infatuated with the idea of a “biotechnology revolution” driven by recombinant DNA. Due to the perceived political, economic, and social importance surrounding biotechnology, many scholars turned their attention to understanding the origins of the new revolution and whether it was decidedly different than what had come before it. Historians were not actually the first to write histories of biotechnologies but rather sociologists and philosophers who were interested in how biotechnologies, and, particularly, recombinant DNA, had effected many traditional institutions and ideas. One of the earliest scholars who focused on writing a historical account of biotechnology was Sheldon Krimsky, a philosopher who was part of the newly constructed bioethics community that had arisen in the 1970s. As a member of the government’s task force to review recombinant DNA, Krimsky had had a front-row seat for many of the debates over the topic and wrote Genetic Alchemy: The Social History of the Recombinant DNA Controversy (1982), which discussed the ethical controversies raised with the development of the technology, how those debates changed over the decade, and the subsequent policies that developed in response. As one of the first books to trace a historical path through the recombinant DNA debates, it served as an initial road map for scholars interested in understanding the controversies surrounding recombinant DNA. Recombinant DNA, however, seemed to involve more than ethical debates, and sociologists became particularly interested in how these new technologies had transformed the institutions in and around science. Martin Kenney was one of the first scholars to do so and his Biotechnology: The University-Industrial Complex (1986) focused on the effect of recombinant DNA by looking at the new relationships that had formed between commercial entities and public universities since its arrival. Though Kenney acknowledged that biotechnology could be defined very broadly – humans had, he noted, used and exploited organisms for millennia – he explicitly employed a narrow definition of the term, writing that he was “restricting the word biotechnology to the new biological techniques that found commercial applications during the 1970s and 1980s” (Kenney 1986, p. 2). Kenney articulated how this definition of biotechnology had developed since the 1950s, drawing upon a narrative that began with public and philanthropic funding of molecular biology in the 1930s and moving to a contemporary analysis of the complex relationships between private biotechnology corporations and university biology professors and departments in the 1980s. Much like the fears that came with Eisenhower’s 1961 warning about the new military-industrial complex, so, too, did Kenney illustrate the potential dangers of the new university-industrial complex that had arisen by the mid-1980s. Kenney’s focus was also on the institutional level, analyzing how companies and universities had changed in response to the development of recombinant DNA. Of course, Kenney noted, these larger social and institutional changes had a significant impact on the activities of individuals, namely, professors, university administrations (from department politics to institutional priorities), and corporate leaders, reshaping priorities and restructuring relationships between all of these actors. Moreover, Kenney described the connections between private businesses and publicly funded university

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professors as a novel product of the biotechnology revolution, one that had permanently reshaped the landscape and not necessarily for the betterment of society. Two years later, Jack Ralph Kloppenburg, Jr., also a sociologist, published First the Seed: The Political Economy of Plant Biotechnology (1988), which depicted a much longer history of biotechnology than the one that Kenney had put forward. Kloppenburg focused particularly on the technologies involved in the production of plants and plant varieties and placed recombinant DNA techniques within a longer narrative of the ways in which humans have tinkered and transformed seeds to maximize their benefit for humans. Methodologically, Kloppenburg relied on a political economy framework, tracing how technologies affected the commodification of seeds and the flow of capital investment, how seeds were produced and who produced them, and how both commodification and production issues affected institutions in the United States. Kloppenburg was explicit about his desire to connect the “old” biotechnologies of plant breeding with the “new” biotechnologies that multinational corporations like Monsanto were using to create novel plant varieties. In doing so, he showed how many of the changes that seemed to be occurring in the biotechnology revolution of the 1980s had historical antecedents. For instance, the Bayh-Dole Act of 1980 was widely seen to be innovative federal legislation that changed the rules concerning intellectual property rights on inventions made using federal funding, allowing universities and corporations to own the patents gained from government-funded research. As Kloppenburg illustrated, however, the Bayh-Dole Act was just the most recent node along a string of government policy interventions over the course of American history used to encourage innovation in agriculture. Like Kenney, Kloppenburg was critical of the new biotechnologies, denouncing the policies that led to a significant transfer of wealth and power from the public domain into the private sector. Perhaps not surprising due to their training as sociologists, what Kenney and Kloppenburg had in common was how they both articulated the ways in which biotechnologies shaped and reshaped the sociological institutions in the production of science, particularly in regard to commercial activities. Neither of the authors critically examined the techno-science itself, instead focusing on the effects of these developments. The social transformations related to recombinant DNA biotechnologies also attracted the attention of historians of science. Susan Wright, one of the earliest historians who examined the implications of recombinant DNA technologies, argued that the state had created powerful commercial motivations for scientists to help build the new biotechnology industry that emerged in the late 1970s and early 1980s (Wright 1986a, b, 1994). In her 1994 book, Molecular Politics: Developing American and British Regulatory Policy for Genetic Engineering, 1972–1982 impressive in its heft alone, Wright analyzed the social and political regimes in the 1960s and 1970s that produced the environment in which recombinant DNA moved from a controversial endeavor to one seen as less of a threat than originally anticipated. Wright also analyzed the regulatory environments that emerged in response to recombinant DNA techniques, showing how the differences between British and American regulations affected the development of the biotech sector in the early 1980s.

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In the early 1990s, historian Robert Bud crafted a narrative that reframed the discussions surrounding biotechnologies from one focused on the impact of biotechnologies to one focused on characterization of what makes biotechnology distinctive (Bud 1991, 1992, 1993). In The Uses of Life (1993), Bud placed biotechnology’s beginnings in the emergence of zymotechnology in the mid-nineteenth century. For him, zymotechnology, or the science of brewing and fermentation, contained the kernels of consistency that he thought still resided in the contemporary forms of biotechnology. The core of biotechnology, Bud claimed, was the not just the way that people had sought to control natural processes but more importantly how people have exploited these processes for profit. The emphasis on the industrialization of nature allowed Bud to include many industries that had been overlooked or forgotten in contemporary discussions of biotechnology that focused almost exclusively on recombinant DNA.8 It is important to note, however, that Bud privileged only the scientific and technological advances that became commercialized and left other advances related to the increasing control over nature to be explored in different histories of science and technology.9 To trace the evolution of biotechnology industries, Bud focused on how the various forms of the word “biotechnology” (biotechnics, bioengineering, biological engineering, etc.) had been employed. Bud did not set any particular geographic boundaries to his history and traced how the definition of biotechnology and its cognates had changed over time as Hungarian, Czechoslovakian, German, French, English, and American usages interacted to create a nebulous concept that could be used in a variety of settings for a variety of purposes.10 Bud’s etymological story showed what types of technologies had been defined as biotechnologies throughout history and also how these biotechnologies became associated with a variety of ethical, cultural, and political concepts, all of which, he claimed, underpinned the multifaceted definitions of, and reactions to, biotechnology in the 1980s. Bud built a history of biotechnology that did two important things. Firstly, Bud argued that the essential aspect of biotechnology was its connection to industry. The “uses of life” were meant not to articulate how humans had used organisms to their advantage but rather how we had industrialized and profited from them. Secondly, and perhaps most importantly, was that Bud’s history undermined the “revolutionary” narrative of the 1980s. Bud showed that there had been a long history of the industrial use of biological processes, and recombinant DNA was simply the most recent biological mechanism to be exploited.

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Bud is not the only historian who was attracted to the industrialization of biological products. See Neushul (1993) 9 For example, the types of histories that Philip Pauly illuminated in Controlling Life: Jacques Loeb & the Engineering Ideal in Biology (1987) which historians have found to be particularly useful in understanding the changes in twentieth-century biosciences that became relevant to the development of biotechnologies 10 Bud was tapping into the emerging literature during that time in the history of science and technology on “boundary objects,” which he implied was what biotechnology was for many actors in his story. He explored the concept of biotechnology as a boundary object more specifically in Bud (1991)

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The creation of a long history of biotechnology was both lauded and criticized by fellow historians. On the one hand, Bud illuminated a history that had previously been understudied, bringing together a wide variety of sources from various countries and contexts. On the other hand, to some historians Bud’s focus on the connections between industry and biotechnology felt disconnected from modern conceptions of biotechnology that seemed to be associated with far more than business-related issues. Bud, for instance, was not interested in understanding how the relationships between various public and private scientists and entrepreneurs had changed over time but rather was more focused on establishing that there had always been a connection. Bud himself even seemed to be aware of this problem when he began his final chapter by writing, “Practically the entire period covered by this book so far could be considered as the ‘prehistory’ of biotechnology, for only in the 1980s did biotechnology acquire internationally recognized significance” (Bud 1993, p. 189). Where Kenney had begun with the definition and concept of biotechnology as recombinant DNA, Bud ended his account with this idea and in doing so undermined the self-styled biotechnology revolution of the 1980s as a revolution in name only. Beyond the discussions of biotechnology’s origins and effects, historians identified an additional area related to biotechnology worthy of study: patents. Due to the importance of the 1980 US Supreme Court decision, Diamond v. Chakrabarty, which allowed companies like Genentech to capitalize financially on the techniques of recombinant DNA, several scholars turned their attention to issues surrounding intellectual property and biotechnology. Kloppenburg’s work certainly fits into this trend as he looked at the government’s role in the commodification of seeds, but other scholars made notable contributions as well (Weiner 1986, 1987; Lesser 1989; Krimsky 1991; Kevles 1994). These scholars specifically highlighted the impact of the new changes in patent laws and began to use intellectual property discussions as a window into larger cultural and social forces. Daniel Kevles (1994), for instance, showed how the judges’ in the Diamond v. Chakrabarty decision had to go beyond simply a legal analysis of patent law to consider many of the ethical, social, and cultural criticisms that biotechnologies had become associated with by that point. These histories of patent law demonstrated that it was not simply the techniques of recombinant DNA with which people became particularly uncomfortable but rather that it was the ability to patent these techniques and their products that drove many of the biotechnology-related controversies in the 1980s. Ultimately, all of these first-generation works were reactions to the growing interest in and importance of biotechnology during this period. The specter of recombinant DNA can be found in all of them. Kenney, Kloppenburg, and Wright had pointed to specific social, political, and economic factors that had contributed to the rise of a recombinant-DNA-fueled biotechnology revolution in the 1980s and highlighted how these biotechnologies reshaped society. Bud’s work, on the other hand, created a history that highlighted the long-standing history of biotechnology’s industrial roots, undermining the idea that the biotechnology of the 1980s was drastically different – i.e., “revolutionary” – from what had come before it. In all cases, recombinant DNA acted as a powerful driving force, either explicitly or implicitly, for historians of biotechnology during this period.

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Overall, the histories written between the mid-1980s and the mid-1990s laid the foundation for rapid growth in scholarship in the field. These initial forays into the history of biotechnology, however, were increasingly scrutinized by other scholars as time passed. How these early historical accounts defined what was and was not biotechnology, and particularly the distinctiveness of recombinant DNA-fueled biotechnology, was undermined by the next generation of scholarship. This scrutiny developed not only from the heightened awareness of historians to these issues but also from the increased attention that biotechnology received from other scholars.

Second-Generation Histories: Broadening Scope and New Definitions During the late 1990s and early 2000s, a new group of historians began to look closely at biotechnology. These historians were inspired by a couple of major forces. Firstly, biotechnology continued to be a prominent topic in the biosciences. Unlike the 1980s, however, the concept of biotechnology was connected to more than just recombinant DNA. The Human Genome Project, breakthroughs in cloning, and advances in stem cell science, just to name a few headlines, generated immense coverage in both the scientific and popular presses. Secondly, biotechnology became the focus for many scholars outside the discipline of history, and though these works often did not privilege histories in their work, they did broaden both the scope of what types of things should be considered biotechnology and what made biotechnologies particularly compelling research subjects. Because of the importance of this scholarship for historians, it is worth briefly discussing some of the major areas of focus. The work by scholars associated with the social studies of science and science and technology studies (STS) in the 1990s and 2000s transformed many of the definitions and concepts surrounding biotechnology. These scholars included anthropologists, sociologists, and interdisciplinary academics who were interested in the interaction and development of science and society. Emerging in the 1970s, but becoming a powerful voice within critical studies during the 1990s, STS and social studies of science researchers began using biotechnology to explore a variety of theoretical concepts and analytical frameworks for understanding science. In contrast to the work of historians, who by the 1990s had become focused on issues surrounding recombinant DNA and the industrial uses of life, these scholars applied the word “biotechnology” more broadly. Many of these studies focused on the social construction of knowledge. In the 1990s, anthropologists Paul Rabinow (1996) and Joan Fujimura (1996) found biotechnologies to be fertile ground to explore these issues. Though the topical foci of each of these works – Rabinow on the development of polymerase chain reaction (PCR) in the 1980s and Fujimura on cancer research in the 1970s and early 1980s – required each author to illuminate historical aspects of their techno-sciences, creating these histories took a back seat to the broader analysis of how science was made. Their focus on technical objects as biotechnologies, however, is an example of how

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these scholars did not adhere to the concepts that the first generation of biotechnology scholars had created but rather utilized a bigger definition that connected “bio” and “technology.” The ways in which biotechnologies controlled, changed, and reconceptualized the body became particularly important to scholars in the social studies of science and STS fields. Recognizing the ways that humans had come to manipulate and control the body, particularly in light of advances in cloning and stem cells, scholars broadened the traditional definitions of biotechnology that focused on recombinant DNA and industrial applications to include the technologies and science related to reproduction, organ transplantation, and medical technologies (Clarke 1998; Brodwin 2000; Rapp 2000; Mamo and Fishman 2001). By categorizing these technologies as biotechnologies, the 1950s and 1960s quickly became important decades in the history of the field, when technologies such as birth control pills and heart transplantation techniques were developed and commercialized. Notably, in these narratives the end of the 1970s became more known for the emergence of in vitro fertilization rather than recombinant DNA; cutting and pasting DNA by itself had little impact on human bodies unless paired with the reproductive technologies that these scholars highlighted. The concepts of increased control over the body that scholars highlighted dovetailed with historical narratives that had not been originally tied to biotechnology but became particularly important to this area of scholarship, namely Philip Pauly’s 1987 work, Controlling Life, which articulated an engineering ideal that emerged at the turn of the twentieth century and flourished in the postWorld War II work of biologists like Gregory Pincus, pioneer of the pill. These larger conceptions of biotechnology and the body allowed scholars to see how many of these body-related biotechnologies became intimately connected with the evolving neoliberal state. Scholars focused on these issues, which often has been labeled biocapitalism, have argued that the development of biotechnology has been an outcome (or taken advantage) of neo-capitalism regimes and thus look to at how the physical products of biotechnologies – i.e., the blood, cells, tissues, embryos, etc. – have become economically valuable (Waldby and Mitchell 2006; Cooper 2008). Through this critical lens, scholars interested in biocapital have explored many issues concerning concepts of individuality, gender, power, and race, showing how biotechnologies have changed conceptions of life in new and unintended ways.11 With broadened concepts and definitions, STS and social studies of science scholars found biotechnology to be a rich topic that could be used for critical studies. For instance, biotechnology has been used as a powerful cross-cultural example for analyzing issues related to technology and gender (Haraway 1997; Creager et al. 2001), science and democracy (Jasanoff 2005; Hilgartner et al. 2015), science and empire (Jasanoff 2006; Briggs 2002), and postcolonial studies (Kowal et al. 2013;

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Discussing biotechnology in the context of biocapitalism is a particularly rich area of scholarship. For further reading see Fortun (2008), Helmreich (2008), Loeppky (2005), Parry (2004), Rajan (2003, 2006, 2012), and Thacker (2005)

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Kowal 2013; Radin 2013). In these works, the focus on biotechnology allowed for rich discussions on the co-production of knowledge, the influence of the state, and the comparisons across multiple cultures, spaces, and temporalities. Ultimately, the focus by STS scholars and social scientists on biotechnology produced many new insights into the function and impact of biotechnologies within society and vice versa. One of their most enduring impacts to the historiography of biotechnology was how these scholars applied the term biotechnology to many technologies, sciences, and techno-sciences that had originally been overlooked in the first-generation histories. Because their questions were often motivated by different methodologies compared to historians, however, they did not seek to explicitly intervene into the historical literature about biotechnology the way that historians did during this period. Historians during this second generation of scholars often undermined the preconceptions built into the first-generation historical narratives. Much of this historical work was done through focused studies in articles and chapters in edited volumes rather than via monographs. These narrower works allowed historians to examine, for instance, the interactions between biologists and industry in new ways, calling into question whether the policies and relationships that formed around biotechnologies during the “biotech revolution” in the early 1980s were changes in degree rather than in kind compared to earlier episodes. By calling into question the momentousness of the biotech revolution, however, these historians were not simply supporting a long history view. Rather, they tended to complicate the history of biotechnology, turning it from a discussion about tracing a single historical thread into a debate about a nebulous set of practices more indicative of twentieth century life sciences as a whole. A number of historians published work that challenged the notion that conceptions of biotechnology should be deeply connected to development of recombinant DNA and its aftermath. Excellent examples of this can be found in the work of Nicolas Rasmussen (1999b, 2001, 2002), Jean-Paul Gaudillière (2001; 2005a; Gaudillière and Rheinberger 2004), Angela Creager (1998), and Lilly Kay (1998). Both Rasmussen and Gaudillière incorporated the development of vitamins, hormones, and pharmaceuticals in the first half of the twentieth century into narratives of biotechnology. Rasmussen (1999b) showed, for instance, how the relationships between academic life scientists, pharmaceutical companies, and university bureaucracies that formed in the 1930s and 1940s had many similarities to the types of relationships that flourished around recombinant DNA technologies. Similarly, by writing about research on blood in the 1940s, Angela Creager (1998) illustrated how some of the entrepreneurial pursuits of the post-recombinant DNA academic world were not as novel as historians like Kenney had portrayed. Creager’s work also showed that biotechnology did not need to be defined simply through technoscientific objects but rather by the types of relationships formed between entities – i.e. biotechnology is not best understood as a history of a particular technique or technology but rather a commercial relationship that emerges between researchers and companies. Lilly Kay (1998), on the other hand, argued that recombinant DNA was simply an outcome of a much larger goal that had been at the heart of molecular

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biology’s existence since the 1930s and thus should not be seen as new nor specifically connected to DNA even. These narratives of the pre-1980s showed that many of the same elements that scholars labeled “revolutionary” about the so-called biotechnology revolution actually had foundations that stretched well back into the early twentieth century. These narratives differed from Bud’s work from the early 1990s for a number of reasons. Primarily, they highlighted biotechnologies that were outside the etymological boundaries of Bud’s definition. Additionally, they were focused on the relationships between public and private entities in the production of knowledge, an aspect that Bud was not particularly interested in articulating. Nevertheless, these secondgeneration histories did support a “long history” view of biotechnology and made historians reevaluate whether their emphasis on the birth of biotechnology being connected to recombinant DNA was indeed the best way to understand the topic. They did, however, continue to highlight the commercial aspects of biotechnology despite extending these narratives to earlier periods. The essays by Kay (1998) and Creager (1998) that helped historians rethink biotechnology’s origins were published in a landmark volume edited by Arnold Thackray, entitled Private Science: Biotechnology and the Rise of the Molecular Sciences (1998). Notably, Private Science included many of the original scholars who were prominent in the first-generation histories, featuring chapters by Bud, Wright, Kenney, and Kevles. These chapters contained many of the same ideas present in the authors’ trailblazing work, though often the arguments they forwarded in Thackray’s volume had matured and become more nuanced. Bud’s (1998) chapter, for instance, drew more explicit connections between post-1970s recombinant DNA biotechnology and the deeper history that he articulated in The Uses of Life, connections that reviewers had criticized him for not making particularly well in his earlier work. Private Science also provided opportunities for new voices to weigh in on the history of biotechnology, broadening the scope of what was considered biotechnology and undermining some of the existing historiography in the process. Alberto Cambrosio and Peter Keating (1998) examined the history of monoclonal antibodies, more fully articulating the relationship between biotechnology and biomedicine and highlighting an important innovation from the 1970s that had been overlooked in earlier biotechnology histories. Cambrosio and Keating expanded these topics later in Biomedical Platforms (2003). Michael Fortun (1998) and Stephen Hilgartner (1998) separately discussed issues surrounding the Human Genome Project (HGP), a big science project that had been initiated in the late 1980s and consequently absent in the first generation of biotechnology histories. Biotechnology and biomedicine became more explicitly connected during the second generation of scholarship. For example, Soraya de Chadarevian and Harmke Kamminga’s edited volume, Molecularizing Biology and Medicine: New Practices and Alliances, 1910s–1970s (1998), showed how many aspects of the history of biomedicine and pharmaceuticals could be construed as a part of the history of biotechnology. In the 2000s, Jean-Paul Gaudillière (2001, 2005a, b, 2007, 2008a, b) focused on the development and commercialization of pharmaceutical drugs,

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highlighting how the physical objects themselves were windows into what Gaudillière called “the complex networks that bind together the various professionals involved in their invention” (Gaudillière 2005a). In doing so, Gaudillière showed how pharmaceutical drugs fit within the historiography of biotechnology, as the development of drugs also dealt with many of the points that the first generation of scholarship articulated as making biotechnologies distinctive, including patenting issues, changing university-industry relationships, and the relationship to the history of molecular biology. Gaudillière also helped turn a focus to the scientists themselves who worked at these pharmaceutical companies, a group that had often been overlooked in the past and would gain further attention in the next generation of scholarship. By the end of the first decade of the twenty-first century, historians and scholars had created a robust literature that expanded both the types of things that should be understood as biotechnologies and the historical antecedents of the field. Instead of understanding recombinant DNA in the late 1970s as the primary catalyst for a social, institutional, and cultural revolution in biology, historians and biotechnology scholars had undermined the importance of recombinant DNA and demonstrated that many of the transformations that took place in and between private and public institutions during the biotech revolution had been modeled off of much older relationships. Moreover, by the end of the second generation of scholarship on biotechnology, a fertile exchange had occurred between STS scholars and historians, giving way to a great deal of crossover among these scholars between various fields by the end of the 2000s.12

Third-Generation Histories: Revisionist Histories In the late 2000s, historians returned to the dramatic rise in biotechnology activity in the 1970s and early 1980s. By this point in time, however, it had become clear that many of the promises of the biotechnology revolution had failed to live up to the hype that had instigated the first generation of scholars. Armed with a deeper understanding of the subject, more sources, and greater perspective, historians reinterpreted the biotechnology revolution that supposedly emerged during this period, focusing not on the impact and novelty of biotechnologies as many first-generation historians did but rather on understanding why biotechnologies flourished in specific times and places and the contexts that drove those changes. 12

There are great many examples that could be cited here, but perhaps most indicative of the intersection between historians of biotechnology, STS scholars, and social scientists can be seen in special issue journal volumes focused on biotechnologies such as the Studies in History and Philosophy of Biological and Biomedical Sciences special issue “Between the Farm and the Clinic: Agriculture and Reproductive Technology in the Twentieth Century,” vol. 38, June 2007, and the Social Studies of Science volume on post-colonial studies and biotechnologies, vol. 43, August 2013

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One of the new ways that historians approached this period was by concentrating on the larger social context of the 1960s and 1970s, which they argued allowed for the rapid growth of the biotechnology industry. In doing so, however, historians created several competing arguments about which contexts were the most important. Eric Vettel (2008) argued for the importance of the 1960s counterculture movement in San Francisco as a driving force in the creation of the subsequent biotech industry. Social pressure, Vettel claimed, pushed the big bioscience departments in the San Francisco Bay Area to transition from a focus on “fundamental” research to “applied” research. For Vettel, the new orientation of the research universities in the Bay Area led to the creation of the Cetus Corporation, which he dubs the first modern biotechnology company. Notably, the Cetus Corporation was not built to exploit the new recombinant DNA technologies but rather microbial screening machines. However, Vettel believed the combination of applied research and venture capitalism that fueled the rise of the Cetus Corporation to be the same driving force of the biotechnology revolution of the late 1970s and early 1980s. In 2011, Sally Smith Hughes and Soraya de Chadarevian separately argued for the importance of an entrepreneurial culture in the 1970s for the rapid growth of the biotechnology industry. In contrast to Vettel, Hughes profiled the rise of Genentech, arguing that the history of this company, the first company to go public using recombinant DNA technology, embodied the driving culture and ideals that not only fueled the growth of biotechnology in the early 1980s but was often the blueprint that typified the industry as a whole. Hughes’ account stressed the entrepreneurial spirit of the founders and the success that scientists found within the scientific and business worlds. The outcome of these arguments and narratives, however, is that the book reads largely as a hagiography of Genentech.13 Focusing on the British context, de Chadarevian (2011) argued that there was no “patent culture” within the United Kingdom during the 1970s like that of the United States. Despite a number of scientific breakthroughs in the twentieth century, such as penicillin and monoclonal antibodies, British scientists and institutions were not mindful of the entrepreneurial possibilities of their work and thus missed out on the surge in biotechnology activity that took place in the United States during the 1980s. Rather, de Chadarevian claimed, a patent culture had to be deliberately developed. While Vettel and Hughes concentrated on the contexts that drove the development of specific companies, Doogab Yi (2007, 2008, 2009, 2011, 2015) resituated the development of recombinant DNA techniques and its transition from a basic laboratory tool to the driver of modern biotechnology. To do so, Yi analyzed both the work of scientists and the institutional, moral, and economic changes occurring at universities and within biomedical laboratories during the 1960s and 1970s that led 13

Though carefully curated, one of the most important aspects of Hughes’ work relates to the series of oral histories that she helped conduct with the pioneers of these early biotech companies. Hughes used these oral histories as the basis of her book, and they are available through the University of California Berkeley’s Regional Oral History Office, which can be found at the website for the Regional Oral History Office, Bancroft Library, University of California, Berkeley. See http:// vm136.lib.berkeley.edu/BANC/ROHO/projects/biosci/oh_list.html, accessed February 16, 2016

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to the rise of the biotechnology sector as a whole. In Yi’s accounts, previous recombinant DNA heroes Cohen and Boyer faded into the background compared to the scientific community in San Francisco led by Paul Berg and Stanford’s biochemistry department. Yi’s work undercut the traditional story of recombinant DNA and the subsequent biotechnology revolution, which had been “mythologized” through not only the retellings of scientists themselves but also through the several of the histories outlined in this essay. Rather, Yi showed how recombinant DNA was a part of a larger story of biological investigations during this period. New sources and nearly 30 more years of perspective allowed Nicolas Rasmussen (2014) to return to some of the earliest questions posed by the first generation of scholars. Like Kenney, Rasmussen was interested in how the new influx of commercially driven investments affected the practice and institution of science during the late 1970s and 1980s and particularly how these affected scientists working within commercial enterprises. To get at these questions, Rasmussen relied on newly available legal documents and court testimonies to explore how academic traditions of science and scientists became transplanted, integrated, and eventually transformed over time as the biotech industry grew. He also investigated how long-standing reward structures of getting credit for scientific work through peer-reviewed publications became upended as press conferences and patent applications became more important. Rasmussen focused on biologists themselves as active participants in all of these changes, undermining other historical interpretations that emphasized larger macro-trends and structural transformations as the agents of change or a technological determinism created by recombinant DNA’s discovery. Similarly to Yi, Vettel, and Hughes, Rasmussen also strove to create a larger context for the radical growth of the biotech industry in the late 1970s, weaving together narratives from the history of molecular biology and biophysics with the history of medicine, Cold War politics, science policy, patent law, and emerging ethical debates – topics that Rasmussen had explored throughout much of his career (Rasmussen 1999a, b, 2001, 2002).14 Two larger themes emerged from these revisionist accounts. First, Vettel, Hughes, de Chadarevian, Yi, and Rasmussen did not support the narrative that the technologies developed during this period were responsible for the social, economic, and institutional changes that took place. Rather, each author focused, in their own way, on the changes that occurred within particular institutions or groups that made it possible for the dramatic growth of what became commonly referred to as biotechnology in the 1980s. In the process, the role of recombinant DNA, though certainly important, became decentralized in favor of focusing on larger contexts. Secondly, though each of these authors told a different story about the rise of biotechnology industry, their work does, in effect, support the idea that the biotechnology of the late 1970s and 1980s should be seen as the beginning of a distinct historical time period. For these historians, questions concerning whether biotechnology had a long history or was a product of a revolutionary development were not relevant. This latter theme

14

Much of this paragraph was taken from my review of Rasmussen’s book (Journal of the History of Biology 2015)

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sits as a correction to the scholarship that took place in the second-generation histories, which tended to deemphasize the importance of the growth of biotechnology in the 1970s and 1980s. The reassessment of the biotechnology revolution coincided with a growing interest in the historiography of biotechnology. Jean-Paul Gaudillière was one of the first historians to write historiographical articles on the subject. As someone who had already written several histories of biotechnology by this point, Gaudillière examined the “biotechnology problem” (2009b) in which he argued that biotechnology should not be understood as a discipline nor an industry nor even a particular set of technologies but rather as a larger explanatory framework by which to interpret modern biosciences. The life sciences, he argued, had entered into their biotech phase, which was characterized by the construction of neoliberal market relationships between the academic world and agricultural and biomedical industries. These relationships pervaded all of modern biosciences, meaning that specific biosciences like molecular biology and recombinant DNA could not be disentangled from the larger pattern. Ultimately, Gaudillière argued that historians should employ a new framework when thinking about twentieth-century life sciences, what he called “ways of regulating,” which effectively privilege and expand the importance of histories of biotechnologies for historians of the life sciences.15 Beyond revisionist histories and historiographies, during the 2000s and 2010s, historians continued to examine long-standing issues in the field while also investigating new areas of biotechnology that had previously been overlooked. Historians had been discussing patents and intellectual property rights since some of the earliest biotechnology histories of the 1980s, and they continued to explore this issue by incorporating more stories that took place outside the United States. Historians explored how many Western countries regulated, through government and nongovernment actions, the proliferation and successes of biotechnologies (Kevles 2002, 2007). They also articulated the interplay between patents and biomedicine, a category that historians had made virtually synonymous with biotechnology, by focusing on European pharmaceutical companies (Gaudillière 2008a, b; Slinn 2008). In 2008, the Max-Planck-Institut für Wissenschaftsgeschichte hosted an international conference on patent rights and biotechnology, which became an edited volume in 2009 (Gaudillière 2009a). The volume had a particularly strong focus on patents and plants, a topic that historians showed had a much longer and internationally robust history than simply the 1980s patent issues that arose in the United States due to the Diamond v. Chakrabarty decision. Historians had long defined biotechnology by the manipulation of organisms, and most histories had focused more on the issues of manipulation than on the organisms themselves. How these created organisms could be understood as biotechnologies 15

Gaudillière also notes, much like I argue in this essay, that one of the main disagreements in the literature concerns whether molecular biology, particularly recombinant DNA technologies, should be seen as revolutionary or a part of a longer history. He labels this to fight the discontinuity/ continuity argument and believes that it ultimately is a problematic way to understand the importance of biotechnologies during the twentieth and twenty-first century

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became an important area of research in the 2000s. Many of the studies that began to consider animals as biotechnology have their roots in historians’ interest in model organisms. Kohler (1994), for instance, came to understand the domestication and development of the fruit fly for early genetics research as the creation of a laboratory technology. This laboratory-animal-as-technology motif has been pursued in many other historical studies on model and standardized organisms since then (e.g., Creager 2001; Rader 2004; Kirk 2012) and has also been an important topic for philosophers of science (Leonelli and Ankeny 2011). Yet none of these scholars engaged explicitly with the literature surrounding the history of biotechnology. Seeing the changing role of laboratory organisms as a part of the history of biotechnology became more prominent in the first decade of the twenty-first century. As genetically modified organisms became important to agricultural, which include both crops and livestock, historians became more interested in understanding their impacts (Munns 2015; Garcia-Sancho 2015; Berry 2014; Saha 2013). Science studies scholars also wrote histories related to the products of biotechnology. Hannah Landecker (2007) and Sarah Franklin (2007), for example, used the analytical tools of social studies of science and STS scholars to upend traditional narratives of biotechnology. Landecker treated the development of cell culturing techniques, which allowed biologists to keep individual cells alive outside the body, as the creation of a new form of biotechnology that reshaped the way that biologists understood life. Rather than associating her narrative with the prominent histories of biotechnology related to molecular biology and recombinant DNA, Landecker instead connected her definition of biotechnology to histories of twentieth-century developmental biology and biomedical research. Her history of cell culturing, she claimed, was not reflected in etymology of the word biotechnology but rather fit into a larger history of “technologies of living substances” (Landecker 2007, pp. 1–2). Sarah Franklin (2007) also upended conventional interpretations of the significance of the cloned sheep, Dolly, by inserting the creation of sheep into a broader narrative of animal husbandry, agricultural research, and reconfigured genealogical relationships. Ultimately, the third generation of biotechnology, which I would argue we are still currently in, can be characterized by a maturation of the field as a whole. At the time of this publication, the history of biotechnology encompasses the traditional definitions of recombinant DNA as well as a multitude of other concepts including human reproduction technologies, agricultural products, and the intellectual property rights issues. Moreover, the revisionist accounts by Rasmussen and Yi, for instance, which reinterpreted the biotechnology revolution of the 1970s, suggest an ability to not only reflect a particular time period but on the historical literature itself. Similarly, the historiographical discussions like that of Gaudillière (and of the presence of this chapter) indicate that there has been enough literature published on the history of biotechnology to begin to pick out patterns and trends. Generally, the future of the history of biotechnology will map closely to new developments in the biosciences, just as they have always been closely related to the current work being done when they were written. The first generation of biotechnology histories, for instance, focused intently on the recombinant DNA revolution, either in its origins and effects or questioning whether it was a revolution at all. As

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1990s drew to a close and the new biotechnologies of the 1980s had become more fully integrated, both in how they affected the practice of science and its institutions, historians began to see more parallels to the past. Similarly, as cloning and stem cells became prominent issues in the late 1990s and 2000s, histories emerged that reflected these anxieties. Third-generation histories of biotechnology also follow this trend. In the 2010s, historians and related scholars have focused on the impact of computers to the biomedical process (Strasser 2011; García-Sancho 2012; Stevens 2013; Onaga 2014), and synthetic biology (Campos 2012; Calvert 2012; Mackenzie 2013; Boudry and Pigliucci 2013), continually expanding conceptions of biotechnology in the process but also taking part in a long-standing tradition in the history of biotechnology. Given the range of topics and concepts I have placed within this third and extant generation of scholarship, it may appear that the history of biotechnology has become indistinct from the history of biology in general. This is partly an outcome of the way that the label of biotechnology has been more liberally, and perhaps problematically, applied to describe a multitude of topics within the biosciences. This could both be a justifiable evolution in the concept and/or an outcome of scholars attempting to keep their work connected to the relevance of biotechnology today. Yet it is also important to realize that the essential aspect of commerciality, and all the issues surrounding it, that drove the earliest scholars to the history of biotechnology, and has fueled the interests of scholars since, has saturated the discourses and practices of modern biosciences. In other words, perhaps what once made biotechnology distinct enough to validate its own histories has now permeated the biosciences in such a way that it has become simply a matter of discretion when something should be labeled biotechnology. One major consequence of this is that the history of biotechnology will continue to be an important field of historical research going forward. Perhaps the other major consequence will be that this messiness in both the historiography and in the subject itself will provide the fodder for subsequent generations of historians of biotechnology.

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The Matter of Practice in the Historiography of the Experimental Life Sciences

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Hannah Landecker

Contents Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . The Practice Turn as an Experimental Turn . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . How to Do Things with Practice . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Experimental Systems and Epistemic Things . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Temporality and Experimental Systems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Model Systems and Biomedical Platforms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Studies of Visualization . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Conclusion: Questions and New Directions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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Abstract

This chapter reviews the trajectory of the practice turn in histories of experimental biology. With a focus on “how to do things with practice,” the methodological implications of a focus on material practice are discussed. A map of the overlapping territories of experimental systems, epistemic things, biomedical platforms, visualization practices, and experimental bodies is traced out together with the source materials that are central to these approaches, such as gray literature, protocol manuals, and laboratory notebooks. The argument is presented that studies of literature, rhetoric, narrative and concept are not opposed to studies of material practice, and indeed present opportunities going forward for new syntheses and integration of approaches.

H. Landecker (*) UCLA, Los Angeles, CA, USA e-mail: [email protected] # Springer Nature Switzerland AG 2021 M. R. Dietrich et al. (eds.), Handbook of the Historiography of Biology, Historiographies of Science, https://doi.org/10.1007/978-3-319-90119-0_14

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Introduction What is a practice? According to the Oxford English Dictionary, it is the habitual doing or carrying on of something; usual, customary, or constant action or performance. It is conduct. To the contemporary reader, it might seem strange to think about doing histories of experimental science without attending to practices, to those habits and carryings-on that occur in laboratories, fields, and academies: to not account for how things were customarily done, with what materials and instruments, particularly biology. How would one do biology the science without biology the stuff – without living matter and without the means to observe, cultivate, prod, and test within living things and ecologies? Indeed, the use of biological matter to make other practical tools for research and industry, from enzymes to antibiotics, points to a manifestly practical character to this endeavor (Bud 1994; Podolsky 2014). It seems to follow quite obviously that to do history of biology the science, one should think about the histories of matter and enactment. Nonetheless, like other areas of history of science, this subfield quite explicitly and self-consciously discovered practice not so long ago. Historians of biology participated in and indeed shaped what has been called the “practice turn,” a broad trend sweeping the larger terrain of scholarship in the last decades of the twentieth century (Schatski et al. 2001; Pickering 1992).1 A turn implies leaving a given route or direction and choosing another, which in its most general terms was characterized by its proponents as a departure from approaches focused on theory, language, representation, values, ideas, individuals, or structures and a turn toward practical action, toward how things are done rather than what things mean (Trentmann 2009, p. 297). Notable in this literature is the rather unconscious use of the word “actual,” with the repeated assertion that turning to practice enables the researcher to access how science is or was actually done. This phrasing implies that previous approaches did not capture actual science but an impoverished version of science leftover once it was abstracted from the settings in which it was done. The function of such language was to differentiate experiment-focused accounts of science from earlier intellectual history approaches to the history of science and, after Kuhn, from accounts of theory-driven change in science. Like all generalizations, this one is useful for the big picture but might be better understood as a productive source of historiographical renewal than a turn. It opened older material up to fresh readings; suggested new framings for historical inquiry; challenged some rickety binaries such as external/internal, constructed/real, and theory/practice; and prompted new empirical approaches to different kinds of sources. Representation was not left behind, but a rich vein of studies into the practices of visualization, inscription, and model-building opened, from microscopy to wax models (Schickore 2007; de Chadarevian and Hopwood 2004). Language

1

See Rouse (2007) for a review of Practice Theory in the Philosophy of Anthropology and Sociology. Law (2009) provides a useful overview of “Actor Network Theory and Material Semiotics” in the Blackwell Companion to Social Theory.

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and rhetoric were not abandoned but were resituated within the larger scope of discursive practices tethered to particular technologies and contexts, from models to data display (Kay 2000, p. 18). It occurred to scholars that they should be looking at instruction manuals, protocols, preparations, scientific newsletters, and instruments as well as scientific treatises, at places where procedures are explained and debated as well as places where theories are argued. Even values could be reinvigorated as empirical objects of study when recast as enacted value practices (Dussauge et al. 2015). Arguably, we are still turning. Today, practices and the materiality of scenes of practical action in life science are taken as central for history of biology, to the extent that contemporary authors are less likely to mark out this orientation as requiring an explicit methodological or historiographical discussion; they are more likely to just go ahead and do it or mix it in with biographical and institutional information. It has become unsurprising to students, and I have been asked, “but who doesn’t address practice now?” Yet this is unwarranted complacency. The general orientation toward practical matters does not in itself dictate which practices and what empirical sources should or could be attended to nor what micro- or macroscale narratives should be pursued. It is one thing to herald attention to materiality; it is quite another to decide which matter in a complex experimental setting to attend to. Thus we continue to work through the historiographical implications of writing histories of science that are or include histories of practice, whether or not those implications are explicitly addressed by any given author. Below, I revisit some of the central texts of the “practice turn” that are particularly relevant to the history of experimental biology and that should continue to be included on any reading list of the historian of biology. Necessarily partial, the texts chosen for discussion are those that have framed ensuing studies into experimental biology in interesting ways and may continue to be valuable provocations for new projects going forward.

The Practice Turn as an Experimental Turn It can be quite confusing to try and navigate what was meant by practice in the changing winds of the debates around realism and social construction of science in the latter half of the twentieth century. Therefore I begin here with a few key aspects of the shift in attention to practice in the history of biology and how, conversely, the examination of biological experimentation has been productive for theorizing science as practice. One the one hand, a turn to practice seemed a way to challenge the image of science as objective truth produced by geniuses unaffected by politics or culture; the process of production of concepts and truth claims was thereby under scrutiny. Thus the now classic Leviathan and the Air-Pump was titled that way and not Leviathan and the Concept of a Vacuum; it was not about which account of nature was right but about the ways in which experimental practice produced knowledge and certainty with particular technologies and techniques in the specific historical setting of seventeenth-century English society, where other modes of knowing such as natural philosophy were current (Shapin and Schaffer 1985).

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Practice therefore was an important term in the examination of how knowledge was produced in context and was often enrolled in the project of showing knowledge to be socially constructed. On the other hand (and this is where it can get confusing), practice was reclaimed after a while as a way to navigate away from social constructionism in its strong form and toward a better grasp on the role of the material world in constraining, shaking up, or shaping knowledge production. Experimental life sciences provided an excellent venue for exploring these ideas on new empirical ground, departing from a literature that was heavy on physics.2 An important event in the development of this literature about biology and medicine is the 1979 translation into English of Ludwik Fleck’s Genesis and Development of a Scientific Fact, originally published in German in 1935 (Fleck 1979). Examining the history of syphilis, Fleck’s narrative centers on the relationship between the concept of the disease, the community of researchers and doctors, and the development of the Wassermann reaction, in which blood drawn from patients could be tested for the presence of antibodies to the spirochete bacteria recently identified as the causal agent of the disease. Giving the name “thought collective” to the group that any scientific practitioner belonged to, and “thought style” to the kind of thinking shared among them, Fleck used the example of the development of the Wasserman test to give empirical historical detail to the interaction of thought, perception, experiment, and instrumentation that over time coalesced as an accepted reality and a set of facts about syphilis. The title itself points to the central argument: facts are made in and through historically and socially specific settings. Fleck was writing explicitly counter to reigning depictions of science as a disembodied objective truth. His emphasis on science as a collective cognitive process that was fundamentally social in nature is often noted as an influence on Thomas Kuhn and The Structure of Scientific Revolutions (1962). Yet the empirical core of the book is about the development of a diagnostic technique and pays close attention to the negotiations between scientists and matter, as in this passage that gives room both to the work of thought and the ways in which the material world pushes back on the experimenter: The work of the research scientist means that in the complex confusion and chaos which he faces, he must distinguish that which obeys his will from that which arises spontaneously and opposes it. This is the firm ground that he, as representative of the thought collective, continuously seeks. (1979, p. 95)

The translation of Fleck’s text, with its interest in “how knowledge making relates to the recalcitrance of. . .the material world,” was therefore significant both for elaboration of what it meant for scientific knowledge to be socially constructed and for later attempts to bring the materiality of experimental work more to the forefront of explanation (Creager 2002, p. 320). Historians began to look at other 2

This is not to say that one couldn’t take the same approach with physics. Andrew Pickering’s book Constructing Quarks: A Sociological History of Particle Physics (1984) and Peter Galison’s Image and Logic: A Material Culture of Microphysics (1997) were both decisive in showing the way.

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instances in the history of experimental practice from this perspective, and simultaneously, broader trends in scholarly attention enabled the broad rediscovery of Fleck’s text as newly relevant. For example, Andrew Pickering reads Fleck’s account as allowing for the agency of the material world, as well as the effects of practice in shaping the social group of expert practitioners competent to carry out the test. In his reading, scientific practice as a “mutual tuning of social and material agency” is at the center of Fleck’s study and fits nicely with his own formulation of the “mangle of practice” to describe the dynamics of “accommodation and recalcitrance” in scientific activity (Pickering 1992, 2005, p. 163). The same year that Fleck’s book was translated saw the publication of Laboratory Life: Construction of a Scientific Fact, by Bruno Latour and Steve Woolgar (1979). This was an ethnographic study of a laboratory at the Salk Institute engaged in a highly competitive race to isolate and identify a peptide hormone. Here, too, the process of consolidating a discovery that becomes accepted fact was observed as it unfolded, with the authors calling attention to processes by which rats and reagents are transformed into results and papers. They noted that practices of establishing truth claims about nature disappear by the time the textual account of the experiment is published and called for the abandonment of cognitive accounts of change in science. The concern with the productivity of practice and the methodological centrality of following it in time was what Latour would come to call “science in action,” which again reinforced the theme of empirical attention to the complexity of action among scientists, technicians, citations, institutions, funders, reagents, machines, and so on, rather than any account of what was going on in scientists’ minds (Latour 1987). Followed by a series of works including the Pasteurization of France, which took on a figure most often treated hagiographically as the prepared mind at the very origins of experimental biology (Louis Pasteur) and reset him into a landscape of other agential “actants” including microbes and cows, Bruno Latour’s work continues to be an important point of reference for the history and sociology of science (Latour 1993, 2005). Not only has it made historians of biology rethink the role and place of the matter and practice of experimental work, but the cross-disciplinary influence of this work across the social sciences and humanities has meant that a much larger constituency of scholars has learned to care about the history of biology than might have otherwise been the case. Equally, Donna Haraway’s formulation of the material-semiotic nature of things in the life sciences has been and continues to be influential (1991). Protesting the strong version of social construction of science as incommensurable with a feminist political commitment to the project of “crafting reliable knowledge about the ‘natural’ world” in the name of being able to live in it better, Haraway was concerned to beat a path of “situated knowledge” that neither gave everything up to social construction nor practiced a naïve realism (1991, p. 184).3 The material-semiotic 3

In that spirit, this review should not be read as a view from nowhere. I went through graduate school at the MIT Program in Science, Technology, and Society between 1994 and 1999, perhaps the height of this literature explicitly focused on the historiography of practice, and went on to a postdoctoral appointment at the Max Planck Institute for the History of Science, and these experiences no doubt profoundly shape my understanding of this literature.

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actor was a way to argue for objects of knowledge as generative and active agents in a politically and historically constituted field of language and technical practices. This twinned attention to material (bodily) production, and signifying practices was meant to capture the dynamic and generative nature of the biological things and to not assume that they existed as objects prior to or outside politics, society, and culture. Taking issue with a narrow focus on experimenters and machines in the work of Latour and other social studies of science focused on laboratories, Haraway’s accounts of immune systems, primates, genetically modified mice, and other objects of knowledge and economic value emphasized their simultaneously semiotic and concrete nature, connecting to a broader web of gender and power than was present in other accounts (1997). In retrospect, what seemed like an argument with oppositions at the time now seems more like differentiated facets of a fractiously unified movement: an intense flourishing of scholarship seeking a way out of the either-or of theory or practice, language or action. Having gained much from the perspective taken by the social construction of science, many authors were nonetheless dissatisfied and sought to move on to a mode of analysis with some form of allowance both for ideas and matter, for persons and things. Thus one sees an ensuing proliferation of language in the 1990s that abandoned Fleck’s “making” of scientific facts as well as the strong “construction” language and explored ideas of enactment or performativity (Mol 2002). Objects such as the material-semiotic actor discussed above contained duality within themselves, meant to capture intersections and interactions as a way out of uncomfortable polarizations of constructed or real, theory or experiment, and language or practice: the mangle of practice (Pickering 1995), knowledge machineries (Knorr-Cetina 1999), epistemic things (Rheinberger 1997), looping of human kinds (Hacking 1998), agential realism (Barad 1998), and a general import of the language of material culture in the history of science (Galison 1997). It is not inconsequential that much of this thinking was done in and through experimental life science; i.e., it is not that the “turn” occurred elsewhere and came to the history of biology, but it is the history of biology itself that has provided much of the empirical ground through which this theory has arisen. Perhaps exactly because biology laboratories are profuse collections of humans, animals, cells, molecules, reagents, machines, and instruments, and the objects of inquiry are frequently manifestly in possession of “lives of their own,” they are very good places to think about concatenations and negotiations of material and social agency (Hacking 1983). Moreover, as Haraway insisted, increasingly more things to do with life in the world seemed at stake in biological laboratories for agriculture, medicine, and health, with genome projects occupying the economic and cultural centrality that once was accorded to moon shots and particle accelerators. Laboratory ethnographies, such as Karin Knorr-Cetina’s Epistemic Cultures: How the Sciences Make Knowledge, compared molecular biology and high-energy physics, further underscoring the growing power of biology as a distinctive endeavor (1999). Not coincidentally, this turn to practice of experimental life sciences occurred in a larger social and economic scene of growth of life and biomedical science both in scale and importance. In 1995, for example, an article reviewing the historiography

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of the experimental life sciences opened with self-assured certainty that its topic was important because “experimental biology has come to be seen as the most powerful force in the modern reconception of the nature of life and in the radical transformation of medical practice” (Kevles and Geison 1995, p. 97). Lawyers and legislators were turning to practice, too (although they wouldn’t have put it in those terms), deciding on which forms of biological practice counted as patentable arts, such as recombinant DNA techniques (Hughes 2001); the patenting of various cell lines, microbes, plants, mice, and eventually gene sequences brought the question of the objects of life science and their productivity and technicity very much to the fore (Kevles 1994; Haraway 2001; Murray 2011). Indeed, they would not have been patentable matter if one could not spell out the role of productive “nonobvious” human invention in distinguishing them from the realm of nature; the practices such as recombinant DNA techniques or cell fusion and the instruments such as PCR machines also became economically valuable both as material things and as part of intellectual property portfolios (Rabinow 1996). Thus the genome, genetically modified organisms, cancer model systems, peptide hormones useful to pharmaceutical development, and pretty much anything to do with genes and molecular biology emerged as interesting and even urgent topics for historical investigation. That is probably enough generalization about a topic that is supposed to be about actually doing versus later abstraction from the scene of action. One of the great ironies of the literature discussed above is that one often finds these works discussed only in terms of the concepts they propose – the epistemic things, the actants, and the theoretical claims about how science is actually done – rather than the empirical ground used to anchor those claims, from peptide hormones to complement fixation tests. Of course one wants to be able to generalize in relation to the particular instance, but it seems paradoxical that science as practice should be so easily abstracted from the empirical practices of the social scientists and historians themselves. What falls away if one leaves the empirical ground behind is not so much the particularities of this or that case study but the important practical questions of how an orientation to practice directs one to do research differently, to pay attention differently in the archive or the interview and also to think carefully about what gets left out by such an approach. This kind of attention to the historian’s practice is essential to the ability to be reflexive and critical about the principles of selection for empirical sources for historical inquiry.4 From a student’s point of view, however, the question is not just what idea of scientific practice is proposed by a text but how to design and do a research project after decades of discussion of these topics. Therefore, the rest of this essay focuses on how to do things with practice.

4

The exception here would be Latour’s Science in Action: How to Follow Scientists and Engineers Through Society (1987). This and later works on the actor-network approach put emphasis on its role as a descriptive technique and a methodological toolkit for grasping the relations between things rather than its existence as a theory (Law 2009).

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How to Do Things with Practice What should a person read if one wants to know how to do things with practice? Historians do not have recourse to ethnographic immersion in the moment in question, so how are they to follow the “performativity. . .of heterogenous assemblages” of persons and things? (Introna 2013, p. 337). Tacit and artisanal knowledge, particularly in the work of unrecognized laborers such as technicians and craftsmen, leave fewer traces for historians. At the same time, there is clearly a role for innovative thinking about how to look for sources and a flourishing of arguments about moving away from “elite” centers from which theoretical change was seen to flow, with the assumption that practice somehow was then changed in a downstream manner after people began to think differently. How have scholars gone about framing their accounts of the history of experimental biology as a history of practical action and interaction with material agency, with what sources and methods? Further, it is important to identify gaps and questions remaining unaddressed in this literature. A number of thematic streams will help organize this necessarily partial overview: experimental or model systems, infrastructures and platforms, and visualization practices and technologies. Throughout, I will emphasize what these themes imply for the empirical work of doing histories of practice. Finally, I will turn to some interesting outstanding questions for the field.

Experimental Systems and Epistemic Things The experimental system is a good place to start. This refers to the collection of organisms, biological matter, machines, and techniques that are the “material functional units of modern knowledge production” (Rheinberger 2010a, p. 244). An actor’s category, in that many experimental biologists shape their work and identity around particular experimental systems, this has also become an important unit of historical inquiry through the work of philosopher, historian, and molecular biologist Hans-Jörg Rheinberger. His important book, Towards a History of Epistemic Things, is focused on the development of an experimental system of in vitro protein synthesis through which transfer RNA was established as an intermediary and step in the process of gene transcription and translation (1997). It deals with a particular laboratory at the Massachusetts General Hospital in the period 1947–1962 and a set of scientists working intensively with an identifiable experimental system: a distinct set of materials such as cell extracts, measurement techniques for assaying protein synthesis such as radioisotope counting, and instruments such as ultracentrifuges. It is, as Rheinberger writes, a history of objecticity, rather than a history of objectivity, of the material culture of the sciences (1997, p. 4). Transfer RNA is the epistemic thing, the unknown answer to questions that are themselves discovered in the process of manipulation of the system. It emerges out of the experimental arrangement as a simultaneously conceptual and a phenomenal thing, which comes to be known through how it is instrumentally registered. Epistemic things can in turn become part of experimental systems, once they are stabilized enough.

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How exactly does one trace the emergence of an epistemic thing and then its subsidence into a stabilized element of an experimental system? In Rheinberger’s study, sources include “classic” archival sources such as letters and scientists’ papers but make extensive use of research notebooks, grant applications, meeting minutes, images and drawings, scientific papers, and other historical documents directly relevant to experimental design and manipulation of instruments. Also important is the repeated return to François Jacob’s observations about experimentation in biology, letting the detail of the case study interact with a practitioner’s voiceover, even if the particular practitioner of molecular biology (Jacob) was not the one under study. The chapter structure of the book oscillates between the details of the history and philosophical and historiographical concepts worked out through that history, chief among which is this idea of the epistemic thing: things embodying concepts that emerge out of experimental practice.

Temporality and Experimental Systems Orientation of experimentation toward the future is a particular finding of Rheinberger’s analysis, and close attention to the empirical detail of the temporality of the experimental process is one of its methodological hallmarks. This version of experimental temporality departs from earlier ideas of science developing by theory changing first with experiments later confirming or disproving theories: this entails every experiment looking backward to an established theory for its direction. Attunement to the language, iconography, and practice of temporal action is therefore key to seeing how practice gropes its way into the as of yet unknown. Of course, time is part of experimentation in manifold other ways, as Henning Schmidgen shows in The Helmholtz Curves (2014). Time was not just an object of research for Helmholtz’s nerve physiology (as in reaction time) but was embedded in the “acceleration technologies” of the steam engine and the electrical telegraph taken to physiological experimentation. Attention to instruments in which time was a part of the functioning of the instrument, and the necessity of temporal alignment of various instruments, is one layer of excavating the temporality of experiment and not just its spatiality as an assemblage or network of things and people; Schmidgen further highlights Helmholtz’s desire for speed in demonstrating physiological facts, the temporality of publication and precedence, and the chronically short-of-time scientist. Temporal practice and temporal orientation are thus a part of the study of experimental practice and the effort of the historian to render “the peculiar density of scientific time more palpable” (Schmidgen 2014). Practically speaking, examination of apparatuses, images, letters, equations, and instructions sits alongside reading the published accounts of the work and its theoretical elaboration. Of note, too, is the parallel reading of Marcel Proust, indicating that an orientation to practice does not restrict a historian to staying in the laboratory but instead opens out axes of connection to the larger cultural, social, aesthetic, industrial, and economic world. On this issue of the temporal orientation of scientific work, an entire subfield of literature sits at the intersection of the history and the philosophy of biology around

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the question of “exploratory experimentation” versus theory-driven experimentation (see Schickore 2016 for an overview). Key to doing histories of exploratory experimentation is finding the documentary and material traces of experiments or techniques that feel around in the world without being “theory-driven” and shaped by theoretical expectation terms of their direction and outcome. An example would be Maureen O’Malley’s analysis of metagenomic techniques in the discovery of proteorhodopsin in marine bacteria, whose particularities she uses to explore the differences and commonalities of exploratory and theory-driven experimentation (2007). For historians, the “bottom-up” history of practice rather than the story of elites disseminating theory that then changes practice is proven by empirically following practices, as in Christophe Bonneuil’s study of plant breeding practices in early twentieth-century French agriculture (2006). He argues that rather than asking whether Mendelian theory reshaped breeding practice (or failed to do so), one should examine what horticulturists, agricultural scientists, and breeders did and see that “in contrast to the classical Mendelian experimental system and strategy, plant breeders designed an experimental system that was populated by millions of individuals and hundreds of traits,” aiming not to account for ratios of traits but to “harness a vast genetic lottery and then sort it out, thereby assessing hundreds of potentially interesting new combinations” (2006, p. 299). The “microtechniques” of the breeder’s attention to the plants and the inscription and description techniques of this experimental system are at the center in this account. The focus on practice acts as a productive zone of contact between historians and philosophers of biology. In general, philosophers are more likely to be focused on generalizable accounts of how science works to make knowledge and less likely to take interest in some of the questions central to historians. For example, an empirical focus on practices opens out connections to the social, political, and cultural world in which laboratories operate and how instruments, techniques, reagents and instruments are embedded in the larger social histories of communication, war, industrial chemistry, and business. From radioisotopes to vacuum tubes to silk to wheat, scientific experimental practice both makes and is made by the larger world, and historians are particularly interested in this aspect of practice and in charting the links between “pure” and “applied” science and “academic” and “industrial” domains – rather than further entrenching differences between them by assuming that science only happens in the academic realm and everything else is derivative (Creager 2002; Garson 2015; Onaga 2015; Bonneuil 2006). Many historians take the position that history of experimental science is impoverished without empirical work on the locality and specificity of laboratory or institutional practice. While “there was a time when historians regarded universality as the natural condition of science and local specificity as a problem of adjustment. . .the scholarship of science studies has reversed field in the last generation,” showing that looked at culturally, “we find that tools and concepts alike, mathematical skills as much as laboratory results are anchored at local institutions and practices do not travel readily or seamlessly” (Porter 2012, p. 219). Indeed, close attention to practice for some historians is synonymous with practice-in-context. For example, Soraya de Chadarevian has argued that experimental practices central to

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the production of twentieth-century molecular biology cannot be understood except in the institutional and local laboratory context in which they took place, as in her study the X-ray analysis of protein structure in the Laboratory of Molecular Biology in Cambridge (2002). Yet practices and findings are constantly moving out of the local context and being implemented and interpreted elsewhere in other contexts, creating a question of micro-macro-relations in accounts of practice, as well as some tensions with philosophical accounts of practice, even those that are historically grounded and details (Rheinberger 2009). Such deep detail about context, particularly the larger scene of institutional and social context, is not generally the aim of philosophers’ work, even when interested in the “context of discovery” and not the “context of justification” of experimental work (Schickore and Steinle 2006). Despite these differences, there is a considerable productive overlap between philosophers’ and historians’ concerns about practice in experimental biology, and they are more productively read together than apart. A number of other works give further insight into methods for doing histories of experimental systems. The collected volume of essays Reworking the Bench: Research Notebooks in the History of Science lends insight into the use of research notebooks as rich sources for seeing the role of exploratory experimentation, failure, and mistakes in the laboratory, as well as the gap between the self-presentation of scientists in publication versus the research record not written for broad consumption (Holmes et al. 2003). The historian Larry Holmes was particularly fond of the laboratory notebook as a source for reconstructing the “investigative pathway” followed by experimental scientists; Holmes’ particular expertise was the history of experimental biochemistry, and to him we owe extremely detailed accounts of the work of Hans Krebs and many other figures in the history of biochemistry (1991, 2004). Holmes is an important guide for any historian of science seeking to use such sources, as he also discusses the challenges of using and interpreting these sources, even with the scientists who generated the notes alive and right there as interlocutors, as detailed in Meselson, Stahl and the Replication of DNA (2001). There are also dangers in reconstructing the detail of experimental work in narrative, the chief of these being that one will lose the attention of potential readers who may not want to reexperience every road not taken. Rheinberger’s collected essays in the volume Epistemology of the Concrete (2010a) offer further insight and reflection, particularly into how to think about instruments in the laboratory. In particular, the set of short essays about the intersections between instrument and object found in biological preparations, physiological apparatuses, microscopy, radiography, electrophoresis, and others is a set of concise examples of using historical data about instrument function and operation to understand the materiality of knowledge production. A number of useful examples show the creative empirical possibilities opened up by thinking about the things in the experimental scene. Even though historians cannot do the kind of ethnographic observation of science in progress that anthropologists do in the present, there are still many nontraditional sources that can help one assess scientists’ feel for their materials and the iterative nature of using and making as tinkering proceeds. For example, the so-called gray literature was explicitly regarded by its authors as a different form of communication for a different

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subject matter than publication – the Drosophila community had a newsletter printed with “This is Not a Publication” on its cover that went into enormous detail about the kinds of procedures for fly husbandry that would never make it into formal publication (Kelty 2012). Gray literature such as model organism newsletters is a rich source for the ins and outs of food composition and bedding materials and simultaneously plays an important social role in the community of practitioners who recognize each other not just as the producers of findings but as the makers and doers of a shared field. Discussions about technique, such as what Jutta Schickore has called the “reflexive” conversations about microscopy in manuals and protocols, open out an untapped source for historians interested in arguments about how to do things that are at the same time fundamental arguments about what is seen or sensed in experimental procedures (2007). The very form in which records are kept, tabulated, organized, and represented says a great deal about the organization of perceptual processes within the experimental setting, an insight which has been underlined by the growing importance of data practices in contemporary life sciences but is as true of the hand-drawn tables, kymographs, and breeding notebooks of earlier periods as it is of data ontologies in bioinformatics (Stevens 2013; Brain 2015; Bonnueil 2006). As Robert Brain reminds us in the setting of social science research, there is an ontology to the instruments of measurement, such as the questionnaire (2001). Other nonconventional historical sources such as biological supply company catalogs, technical instrument manuals, grant proposals, and biological sample collections become important points of reference not just for how experiments were done but how experimental matter was kept, systematized, put in suspended animation, rendered, represented, and organized (Strasser 2011; Radin 2013). A curious subgenre of this literature might be called bodily practices of experimentation. In the refusal of a disembodied, only-intellectual, theory-driven, and pure science, historians have turned to the bodily practices of scientists as one way to ground scientific activity and thought in the visceral, quite literally. Steven Shapin’s book Never Pure is subtitled (in part), “Historical Studies of Science as if It was Produced by People with Bodies. . .,” and to do science as if there were bodies in the laboratory along with all the other materials, one has to find historical sources about this presence (2010). Indeed, self-experimentation – because it finds its way into documentation in ways that some other bodily data might not – has been put to good use in histories of X-rays and exercise physiology (Herzig 2005; Johnson 2015). Historian Robert Kirk reminds us that experimenters were not unaware of the material effects that their own bodies and practices had on their animals, for example, in the physiological differences in animals on weekends versus weekdays. He shows that the history of animal care practices includes a kind of experimental science, attempting to measure “how researchers, animal caretakers, and animal technicians formed part of the social environment for laboratory animals” and thereby affected their reproductive and growth rates or their response to drugs (2014, p. 249). In cases where emotion and affect are themselves objects of inquiry, all the bodily affects in the laboratory are implicated, including those of the investigators (Dror 1999). In this literature, histories of experimental biology intersect in

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productive and interesting ways with cultural histories of gender and the body, a welcome corrective to the sometime rather antiseptic accounts of biomedical platforms and epistemic objects. Historian of technology Adelheid Voskuhl reminds us that just because one has a text or object related the how things were done in the past, it is not obvious how one should use that source in historical work. Working with reenactment as a way to explore the history of experimental practice, Voskuhl argues against taking historical documents about practice as historical authorities or transparent windows onto the past. One has, she argues, to read “any text on ‘action’ as the remnant of action rather than an account of action” or drawing on the historian of medicine Adrian Wilson’s ideas of a “hermeneutic stance” on the part of the historian toward the historical source. While these reflections were written in the context of thinking about reenactment of an experiment as a mode of historical inquiry, they are applicable to any text on action: a historical source is regarded as the product, or effect, of a procedure that lies in the past rather than a witness, an authority, or a window to former centuries: the existence of a record of the past is not linked to its function as a document for a researching historian, but only defined through the past period that gave rise to its being. The appropriate question to pose when studying an item of evidence, for example, would thus be: how did this document come into being? Or: how can one infer from its existence to proceedings in the past and what kind of historical explanation does this document allow? (Voskuhl 1997, p. 339)

In short, the varied kinds of sources that a historian might use to access the nature, process, and nonverbal aspects of a past experimental procedure do not speak for themselves. One cannot just turn to practice in some transparent manner; there are still a great many choices about how to find, organize, select, and interpret the potential historical materials that are remnants of past action.

Model Systems and Biomedical Platforms Experimental systems are not the same thing as model organisms, though the two are easily confused because of the centrality of model organisms in twentieth-century experimental life sciences and the fact that model organisms are essential elements of many experimental systems (see ▶ Chap. 13, “Organisms in Experimental Research”, by Ankeny and Leonelli, this volume). This is further muddied by the use of the term model systems by historians to designate experimental systems organized around model organisms: “A model system in biology refers to an organism, object, or process selected for intensive research as an exemplar of a widely observed feature of life (or disease)” (Creager et al. 2007, p. 5). Model systems are attractive as organizing rubrics for histories of science, because they are so central to the conduct of experimental science. Many people and things are entrained to maintaining and stabilizing the model system, and so it provides a narrative organization for addressing the wide variety of things and processes involved in scientific activity. Mice and monkeys, flatworms and corn have charisma

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as organisms with lives and worlds of their own, and in many ways make for tractable narrative devices with a “point of view” on the experimental process that is differently embodied from the experimenter’s, from Darwin’s Dogs to the Guinea Pig’s History of Biology (Townshend 2014; Endersby 2007). Two classic works of history of biological experimentation centered around model systems are Robert Kohler’s Lords of the Fly: Drosophila Genetics and the Experimental Life (1994) and Angela Creager’s The Life of a Virus: Tobacco Mosaic Virus as an Experimental Model, 1930–1965 (2002). Kohler’s emphasis on the way that the social relations between Drosophila geneticists were mediated through the material practices of fly cultivation and exchange, on the technical nature of the living thing, and the laboratory as an environmental niche for the organism changed the way subsequent historians thought about writing histories of biology. Creager’s Life of a Virus emphasized how close attention to the visualization, isolation, and representation of a central model system was a way to address change in twentiethcentury science without requiring recourse to sweeping theoretical change. The detailed case study of tobacco mosaic virus shows how “the experimentalist in biology strives for the typical in his organism or setup, but leaves behind any dreams of a final theory” (2002, p. 3). This is because “What scientists take from exemplars such as TMV is not necessarily a way to see the world differently as a way to handle or materially configure the world differently” (2002, p. 7, emphasis added). Both Kohler and Creager thus show how scientists’ work with model systems led them to handle the material world differently, which Creager argues allows the historian to see experimental practices as not mere responses to contextual factors but achievements that “set in motion trends ranging far outside the laboratory walls” (2002, pp. 2–3). Model systems and model organisms are important narrative devices for both scientists and for historians of science. As noted above, a model system or animal is a part of but is not the same as an experimental system. An experimental system encompasses a broader set of instruments, reagents, and biologicals. Yet other cuts through the material culture of the experimental scene are possible, such as approaches that look to infrastructure, all those things which make research possible yet remain quite unnoticed because of their ubiquity, their origins in the everyday world of consumer goods, or their apparently mundane role in things. Such infrastructural elements can appear in multiple experimental systems and be constitutive of the form of life of model organisms and thus are differently distributed in experimental science than these other kinds of units of practice. These can be living or nonliving elements of infrastructure, from radioisotopes to cell cultures (Creager and Landecker 2009). One can say with some certainty, for example, that experimental research in twentieth-century life science would be entirely different without the presence of fridges and freezers (or for that matter, electricity), yet once the historian trains her eye on the fridge and its contents, different histories come into view (Parry 2004; Radin 2013). Even in the case of animals, not all of the animals that scientists use are model organisms. Experimental animals and their availability as a kind of stabilized, standardized experimental reagent for use in testing, assays, and production of yet other research tools such

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as antibodies have played a vital role in twentieth-century experimental life science (Kirk 2010). Often, the establishment and contestation of standards for these infrastructural elements of science can provide rich insight into infrastructural practice, as demonstrated by Robert Kirk’s immaculately detailed studies of the records of the British Laboratory Animals Bureau and the standardization of germ-free animals (2010, 2012). The history of recent biological sciences has also necessitated attention to the building, maintenance, and use of database infrastructures and the changing relationships between the biology and computing (Stevens 2013; Ribes and Polk 2015). Bruno Strasser has argued that practices of collecting, naming, comparing, displaying, and organizing natural objects can be identified in the making of the molecular sequence database GenBank; through analyzing databases as tools for producing knowledge, he argues that a hybrid of experimental and natural history styles of practice emerged in the late twentieth century (2011). One cut taken across experimental practice with a particular eye to the articulation of medical and biological research space in biomedicine after World War II is Peter Keating and Alberto Cambrosio’s analytic of the “biomedical platform” (2003). As they explain, the biomedical platform is less about the production of “local and unprecedented ‘epistemic things’...” and more about “the constitution and circulation of protocols, instruments and substances between laboratories and the establishment of conventions that allow them to be used in the generation of biomedical facts” (2003, p. 3). They use the example of immunophenotyping, which originated as an experimental system in immunology and oncology, and become a biomedical platform for the diagnosis of leukemia via the “interlaboratory constitution” of cell surface markers and the equipment and techniques used to stabilize them as biological entities that are also pathological signs. The platform is what coheres across laboratory, clinical, and public spaces, enabling the bodies of patients, samples, test results, and clinical entities to function coherently across diverse disciplinary, technical, and architectural spaces; biomedical platforms are the “material and discursive arrangements that act as the bench upon which conventions concerning the biological or normal are connected with conventions concerning the medical or pathological” (2003, p. 4). Practically speaking, the biomedical platform approach draws ones attention to “specific combinations of techniques, instruments, reagents, skills, constituent entities. . .spaces of representation, diagnostic, prognostic, and therapeutic indications, and related etiologic accounts” (2003, p. 4). It also necessitates historical attention to the way in which newer platforms often do not replace older ones but are technically and conceptually articulated with them. While this might seem proper to analyzing the practice of medicine and medical standardization more than the practice of experimental life science per se, one should not be too quick to draw a line between the clinical and the experimental or the clinical and the infrastructural that becomes the experimental and vice versa. While the story of recombinant DNA as an experimental technique is well known, for example, one has to think only of the lesser-known detail that the first restriction enzymes came from a patient in a San Francisco hospital suffering from an antibiotic resistant ailment (Creager 2007). It is exactly the articulation of the clinical and the experimental, and the protocols and

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practices, that keep things and people flowing between these domains that Keating and Cambrosio point us toward.

Studies of Visualization To return for a moment to the scene sketched above of the 1980s and 1990s, in which I characterized the period as a kind of proliferation of hybrid thought-matter, theoryaction, and language-thing forms, visualization practices occupy a special place in the development of scholarship on experimental practices. A convenient origin for this thread lies in the important chapter on microscopes in Representing and Intervening by philosopher Ian Hacking (1983). In it, he argues that we “do not see, in any ordinary sense of the word, with microscopes” (1983, p. 187). Rather, we have various modes of intervening in and working on the observed phenomenon. We do not have a “picture of a gene” in any straightforward sense, because, “biological microscopy without practical biochemistry is as blind as Kant’s intuitions in the absence of concepts” (1983, p. 205). Only when there is good a “map of interactions” between the specimen and the imaging radiation, do we see with a microscope (1983, p. 207). This philosophical inquiry into realism turned attention onto the work of making things visible, discernable, and believable, which was a necessary mixture of visual, verbal, physical, and experiential elements. As with other elements of the late twentieth-century turn to practice elaborated above, the study of visualization has attracted much effort because of its mixed status as simultaneously conceptual, phenomenological, and technical. Norton Wise has argued that the work of making things visible is an important but also underexplored element of the historical examination of experimental practice. We need, he argues, to develop a “materialized epistemology” that unites sensual with ideational knowing (Wise 2006, p. 75). Images and strategies of depiction are an important locus of the material culture that links science to art, popular culture, and pedagogy (Anderson and Dietrich 2012); drawing and photography, for example, are central to how scientists and publics come to see objects such as chromosomes (de Chadarevian 2015). Scientific images and their making are sometimes sites of intense political contest, as in the history of embryology (Morgan 2009; Hopwood 2015; Maienschein 2016). Also interesting is the difficult line between the artifice and technical skill necessary to make things apparent, such as in specimen preparation, and the artifact, produced by the instrument and tricking the observer into counting it as part of the observed reality (Strick 1996). In one way or another, all of the different cuts taken through experimental practice, from experimental systems to infrastructure, model organisms, to instruments, involve the visualization and representation of biological things, temporalities, and processes. Whether one is feeling one’s way into the unknown contours of an epistemic object or making determinations of normal and pathological in oncology samples, practices of representation, observation, visual analysis, and visual communication, pedagogy and explanation are involved. For some scholars, therefore, visualization is the cut to take in examining histories of experimental practice,

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centering analysis on instruments central to visualization, such as kymographs, microscopes, camera lucidas, cinematographs, autoradiographs, or on practices of drawing, sculpting, diagramming, and fixing specimens (de Chadarevian 1993; Rasmussen 1999; Schmidgen 2004; Wellmann 2011; Landecker 2011; Maienschein 1991, 2016; Hopwood 2015). This topic of course could occupy its own chapter in this book and bleeds into other areas of topical concern for historians of the life sciences, such as the role of simulation and modeling (Keller 2000; Creager et al. 2007). Here, too, the history of biology finds zones of productive contact with other disciplines; historians of film and media have much to teach historians of science about working with visual materials, and there is a flourishing of books by cinema scholars on the history of scientific and medical film (Gaycken 2015; Curtis 2015; Ostherr 2013). Visualization thus represents a special subcategory of the turn to practice, and although many of the studies cited in the sections above are also to greater and lesser extent about visualization, it still merits a few words in its own right. Observation is fundamentally mediated by the sense of sight, though smell and sound and touch are sometimes brought in to play. Pictorial representation comes in many forms, as drawing, tracing, imaging, or recording. Indeed, even if a scholar sets out to do a historical project on some aspect of life science without meaning to include visualization, the archive itself may well push back with the manifest presence of many nonverbal traces of past scientific activity. In addition, following the cut of visualization cannot be constrained to scenes of experimentation, as it is exactly the “immutable” mobility of images and their multiple roles in science and culture that make them such interesting historical objects (Latour 1990; Dietrich 2007).

Conclusion: Questions and New Directions The problem with turns is that the work of distinction and departure often leaves many useful things behind. The abandonment of cognitive accounts of science might have seemed necessary when accounts of science were nothing but stories of genius. The rhetorical work necessary to elevate sources that had not even been noticed before, or were dismissed as ephemeral or banal, required diminishing more mainstream empirical sources such as publications or focusing on materials and methods sections of papers and turning away from narrative and rhetoric. Even though many of the concepts detailed above attempt to embody both intellectual and material elements in one package, they were articulated in overt rejection of older approaches that were only about intellectual or theoretical developments. This meant that arguments for the new hybrid semiotic-material or phenomeno-technical unit of historical entity tended to overemphasize the materiality and technicity aspect of the hybrid in distinguishing between new approaches and old, thus somewhat undermining the wish to overcome such binaries about whether something was conceptual or material. At this point it seems a good juncture to historically situate these distinctions in the context of their making and recognize that through this body of work, we can see that “intellectual history is fully compatible with an interest in

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objects and practices and should be reckoned not as a rival or competitor to cultural history, but as its best ally” (Porter 2012, p. 216). Is it possible to move toward an intellectual history of practice? (Landecker 2007). Beyond questions of shopworn distinctions between intellectual and material culture approaches to history, there is also the question of the different scales of inquiry that have emerged in the literature in the last 30 years. The slightly idiosyncratic review of examples touched on above gives some idea as to the problem: we have studies of single laboratories; single controversies; model organisms used by many laboratories and industries; infrastructures that cross not just laboratories but regions and nations, institutions, and groups of practitioners such as horticulturalists or oncologists; and to a certain extent disciplines such as genetics or nerve physiology. There is in addition the question of scales of time. Examples of studies of practice attentive to time might examine the question in the context of a discovery that spans a few years, might unfold the many scales of time present in a single moment or event, or pay attention only to the temporal dynamics of the local context under study. If the turn to practice “has quite naturally privileged micro-histories” and used case studies to move away from sweeping histories of ideas, then is it possible to use fine-grained analysis of practices and materials for doing larger or longer scale histories (Rheinberger 2009; de Chadarevian 2009)? Finally, there is the question of what might have been left behind in the turn to practice. Many of the texts that established this as an empirical and theoretical stance in the history of science made claims about accessing science as it is “actually” done. Indeed, the offhand way in which the word “actually” is used suggests that there are some unexamined assumptions in this founding literature about where science is actually done and where, by implication, it is not actually done. The need to legitimate attention to matters of practice is now past; it is an accepted and central part of contemporary ways of approaching the history of experimental biology. Can we therefore recuperate attention to how science is also actually done in writing and reading and pedagogy and communication, a direction already suggested by engaging with visualization practices? How is science actually done in and through cognition and emotion in relation to visual and tactile experience? There is room for rethinking “a feeling for” the organism, for surprise in relation to the material world, and for the role of sound, smell, and taste in experimental settings (Keller 1983; Fortun 2015). Is there space to reengage older literatures on rhetoric of science or renewed interest in narrative and affect, in relation to all that we have learned from turning to practice?

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Organisms in Experimental Research

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Rachel A. Ankeny and Sabina Leonelli

Contents Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tracing Organisms Through Biographies, Research Fields, and National Trends . . . . . . . . . . . . Organisms as Units of Study . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Organisms in and as Research Practices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Current Directions: Comparative, Quantitative, and Integrative Work Beyond the Western Lab Environment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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Abstract

Research on non-human organisms has been a major focus in the scholarship of historians of biology, especially over the past 25 years. This chapter identifies four overarching trends concerning historical scholarship on the use of nonhuman organisms for experimental purposes, paying attention both to its style and epistemic goals, and to the species and research locations that have been studied and documented. The first trend (1970s–1980s) focused on organisms as one of the many other components of epistemic cultures, the second (1990s) on organisms themselves as units of historical study, the third (late 1990s–2000s) on the organisms in relation to their experimental and institutional context, and the fourth (ongoing) on the diversification of methods and types of research under examination, including multispecies work and the study of practices in a wider range of biological subfields and across geographic locations.

R. A. Ankeny (*) University of Adelaide, Adelaide, SA, Australia e-mail: [email protected] S. Leonelli University of Exeter, Exeter, Devon, UK e-mail: [email protected] # Springer Nature Switzerland AG 2021 M. R. Dietrich et al. (eds.), Handbook of the Historiography of Biology, Historiographies of Science, https://doi.org/10.1007/978-3-319-90119-0_15

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Introduction Non-human organisms are central to much of biological practice and play crucial roles in informing researchers’ theorizing and intellectual trajectories. Biologists’ perceptions of what defines life develop hand in hand with the observational and interventionist strategies used to study the characteristics, life cycles, and behavior of organisms, particularly when organisms are brought into controlled experimental environments. Thus unsurprisingly, non-human organisms have been a main focus in the scholarship of historians of biology, especially over the past 25 years. This chapter explores the existing literature and outlines overarching trends concerning historical scholarship on the use of non-human organisms for experimental purposes. A temptation in approaching a review of this scholarship is to focus solely on well-known historical examples, particularly since several famous biologists have come to be strongly associated with the particular organisms on which they worked. Thomas Hunt Morgan has become synonymous with the fruit fly Drosophila melanogaster, for instance, while Barbara McClintock has come to exemplify research on maize, Max Delbruck on phage, Sydney Brenner on the nematode Caenorhabditis elegans, Eric Kandel on Aplysia, and so on. However, far from being associated solely with one research group, some of these organisms have become so popular so as to function as anchors for entire scientific communities, with journals, infrastructures, funding streams, and discussion venues dedicated explicitly to them and thousands of researchers around the world adopting them as their main materials for experimental work. Particularly since the advent of the largescale genomic sequencing projects associated with the Human Genome projects, these widely used, highly tractable organisms have been ubiquitously referred to in biological discourse as “model organisms,” that is, non-human species that are easy to breed and maintain in large numbers under laboratory conditions and which are extensively studied in order to understand a range of biological phenomena, with the hope that data and theories generated through the use of the model will be applicable to other organisms (Ankeny and Leonelli 2011). The most widely acknowledged inventory of these organisms includes those that have been officially recognized by the US National Institutes of Health as model organisms for biomedical research, such as mouse, rat, zebrafish, fruit fly, nematode worm, and thale cress. The merits of other organisms as potential model organisms are under regular debate (Behringer et al. 2009). Given its defining role for twentieth-century biological science, it is of course important for historians to study the emergence and development of model organism research. This approach to inquiry aligns with other “big science” initiatives emerging in the same period in other disciplines (Agar 2012) and constitutes an excellent platform to examine the role of scale and infrastructures in knowledge production, as well as the importance of translational discourse and attempts to apply biological results to questions relating to human health and disease, as well as food security concerns (Leonelli and Ankeny 2012; Leonelli 2016). At the same time, model

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organisms constitute a tiny fraction of the enormous variety of species used by life scientists to conduct research (Dietrich et al. 2014) and are quite distinct in various senses from the highly diverse set of experimental organisms with which researchers have investigated and interacted over the course of the last three centuries. As Adele Clarke and Joan Fujimura (1992) aptly put it, the choice of an organism for research often amounts to determining what is the “right tool for the job,” and there is a staggering diversity of jobs for which experimental organisms have been used within biological research that historians have labored to document at least to some extent. In what follows, we provide a taxonomy of this historiographical work, paying attention both to its style and epistemic goals and to the species and research locations that have been studied and documented. As is the case with all broad taxonomies, this one acknowledges the presence of exceptions and outliers, yet we believe it is broadly correct and will be useful for others in the field, particularly when attempting to uncover themes, areas, periods, and organisms that are yet unexplored. The first trend that we identify, running from the 1970s until the end of the 1980s, is the treatment of non-human organisms as one among many components within local research cultures, with no specific prominence attributed to them within the narrative and few questions asked around their status, epistemic roles, or practical significance. We have identified at least two reasons for this tendency. The first is associated with the particular historic episodes that have been considered: the specific cases of experimental work under examination often utilized more than one type of organism, and hence within these histories, there was not a focus on particular species, which in turn influenced the overall intellectual approach used by historians in this period. This scholarship typically hybridized the methods and traditions of intellectual, biographical, and/or institutional history with more detailed attention to scientific research and thus placed more emphasis on other aspects of the research process rather than on the materials and technologies which it involves, including organisms. The second trend, which became particularly notable in the early 1990s, was a turn to using organisms themselves as units of historical study. Hence rather than focusing on theories, problems, researchers, institutions, or disciplines, organisms (and usually individual species) became the main characters in these narratives. Together with this change came increased attention to scientific work and practices and greater emphasis on the provenance, characteristics, and behaviors of individual species. Many of these histories stressed the agency and specificity of biological materials used in experimental research and the link between attributes that organisms have and the type of research approach and focus being pursued. The third trend can be viewed as a hybrid of the first two outlined above and came to prominence in the late 1990s. This type of historical scholarship tends to combine the analysis of institutions, scientists, and fields within biology with close attention to the organisms themselves within the context of the experimental work being performed. There is particular attention to the research practices, methods, and technologies adopted in biological laboratories, to the tensions and opportunities

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created when importing and using organisms in those environments, and to Rheinberger’s idea of using biological materials of various types as “things to know with” (1997). This literature focuses on a range of themes, including the ways how certain organisms came to be foci, particularly those that are considered “model organisms,” how they are integrated into specific projects and bring together disciplines or fields, and the details of particular setups within experimental settings and their implications particularly for theoretical and conceptual work. Finally, the fourth trend that we investigate is currently underway, and we wish to encourage the continued expansion of the field in these directions. It involves the use of more diverse methods for studying the history of experimental organisms as well as attention to a wider range of subfields in biology and related research areas. In terms of methods, there have been some attempts to pursue more quantitative findings to supplement the almost exclusively qualitative scholarship that documented above that has dominated the field to date. An additional part of this trend is to utilize methods, literature, and concepts not only from the history of biology but also from philosophy, sociology, and anthropology, including continental approaches. As the majority of literature on the history of experimental organisms has focused on their uses in genetics and molecular biology, other biological subdisciplines have begun to be mined for insights into similarities and differences in these scientific practices. Finally, there has been greater emphasis on multispecies research and on groups of organisms as units of analysis, as well as on nonAnglo-American settings which have been largely neglected in the Anglophone literature. It is significant to note that these historiographic trends developed in parallel with, and arguably in reaction to, trends within twentieth-century biology itself. In the first half of the century, biologists predominantly worked on problem-driven research using a variety of organisms, an approach that was reflected in the outlook and choice of case studies by historians. As biological research became increasingly focused on genetic and then molecular approaches, some individual species, and especially the model organisms discussed above, commanded overwhelming attention, and historians also came to pay greatly increased attention to research on particular species in the latter half of the century. The turn of the millennium brought a renewed interest in integrative and comparative research across species, biological fields, and geographical locations, with an increasingly global research culture emerging in parallel to the Open Science movement and the internationalization of networks and funding sources. At the same time, historians also enlarged their vision beyond the local, to embrace more complex comparative and international narratives, sometimes through the adoption of new historical methods. In what follows, we devote a section to each of the trends that we have identified, with a concluding section outlining the reasons why investigating organisms constitutes a useful lens for historians of biology, though of course by no means the only useful one. Before delving into the material, we should note that our analysis will focus on organisms as conventionally defined, so we do not examine other types of experimental systems which arguably come to have the status or role that organisms do as research materials, such as cell cultures and probes (which are examined in

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detail by Landecker (2009), Creager and Landecker (2009), Landecker (▶ Chap. 12, “The Matter of Practice in the Historiography of the Experimental Life Sciences,” this volume), and Crowe (▶ Chap. 11, “The Historiography of Biotechnology,” this volume)). Furthermore, we consider solely non-human organisms, since an analysis of human experimental subjects would lead us beyond the life sciences and into medicine; more generally we tend to focus on literature from history of science and do not explore the voluminous literature from the history of medicine on animal experimentation, rights, and vivisection. Hence we limit our analysis to laboratory settings, since the growing literature on the use of organisms in other research contexts (such as observational fieldwork, zoos, museums, and clinics) is highly multidisciplinary and evidences a variety of different concepts and trends in comparison to that which focuses on experimental organisms. Finally, while drawing attention to some of the key contributions from continental Europe and elsewhere that are likely to be of interest to historians of science, we primarily analyze trends in Anglophone scholarship, and we focus largely on scholarship from the 1980s onward, since many of the seminal works in the history of biology that appeared before this time tended to be large-scale narratives without any detailed explorations of experimental organisms (e.g., Coleman 1971; Allen 1978, to name just a few).

Tracing Organisms Through Biographies, Research Fields, and National Trends Although much contemporary literature in the history of biology has a strong emphasis on organisms as the organizing trope around which accounts of scientific practice are constructed, this focus is relatively new. Relevant literature in the 1970s and 1980s tended toward broader narratives examining particular scientists or institutions or the emergence of certain research fields or national styles of doing biology. This type of historical work often explored research with non-human organisms as part of studies with much wider scope, for instance, noting the different species chosen and handled by various researchers. Much of this literature generated detailed investigations of the findings, models, or theories that resulted from the use of organisms in the lab and placed strong emphasis on how the adoption of specific organisms shaped existing or emerging individual careers, biological fields, or institutions and related conceptual and organizational trends. Thus organisms were part of a larger story: this literature did not tend to probe or conceptualize the use of organisms as a main focus nor to emphasize their materiality and its constraints as a key theme. The biographical genre details the rise in prominence of particular biologists and in so doing also devoted some attention to the organisms on which they worked. A key example here is Garland Allen’s book (1979) on Thomas Hunt Morgan and those with whom he collaborated, in which the focus is the group’s scientific work, while Drosophila itself is discussed but remains a relatively small part of the story (note that Allen’s (1975) article, discussed below, takes a different approach, as does Carlson’s (1981) biography of H. J. Muller). Both Evelyn Fox Keller (1983) and

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later Nathaniel Comfort (2001) comment extensively on the significance of Barbara McClintock’s handling of maize within their biographies of the scientist, although the organism itself retains a secondary role in their narratives. For instance, Keller emphasizes McClintock’s “feeling for the organism” but more as a means of exploring McClintock’s own intellectual and career development. A more journalistic take can be found in Horace Freeland Judson’s (1979) Eighth Day of Creation which follows the trajectories of key scientists involved in the development of the field of molecular biology, with passing reference to some of the research organisms used, particularly in the later period; again here, the scientists are the main actors with the organisms presented merely as instrumental to the scientific practices examined. A second theme within this literature can be found in the numerous discussions of emergent research traditions or fields in biology, which again touched on experimental organisms but did not explore their specific roles, characteristics, or epistemological status in any great detail. An early example of this approach is Nicholas Mullins’s (1968) sociologically focused Kuhnian-influenced exploration of the origins of the field of molecular biology via research with bacteriophage by the “Phage Group,” which draws considerably on the volume edited by John Cairns et al. (1966) tracing the origins and accomplishments of the Phage Group through milestones as seen by its participants; this study is a classic in the history of biological community formation, but the organism is not central to this discussion. William Coleman’s examination of Claude Bernard’s views on the discipline of psychology (1985) does address the epistemological importance of experimenting on living organisms and yet does not devote much attention to the type of organisms used by Bernard in his research. Bernardino Fantini’s (1985) investigation of organismal choice in embryological and genetic studies in the early twentieth century specifically compares work on sea urchins with work on fruit flies as a means of examining distinct research traditions which each of the organismal types is argued to “symbolize.” Jan Sapp’s (1987) history of cytoplasmic inheritance discusses the use of marine invertebrates and protozoa within this subfield of genetics without documenting and analyzing precisely how organisms were handled or the underlying conceptual or epistemological frameworks associated with the choice and use of these organisms. Finally, considerable scholarship has been done on the use of non-human organisms in the context of the development of biological research within a particular locale or institutional or national context. This literature seamlessly blends analysis of macro-trends at the national and international levels with investigations of practices within specific institutions and labs. A key example is Jonathan Harwood’s (1987) work analyzing and comparing the rise and professionalization of genetics in postwar Germany and the United States, which mentions the importance of work on Drosophila while also critiquing historians’ tendency to focus exclusively on T. H. Morgan’s work. Timothy Lenoir (1982) and Lynn Nyhart (1987, 1995) also focus on biological trends in nineteenth-century Germany and particularly the development and eventual decline of morphology as a prominent field of research. Richard Burian, Jean Gayon, and Doris Zallen (1988) reconstruct the distinct trajectory

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taken in France in terms of the reception of Mendelian genetics, including passing discussions of research with mice and Drosophila. Considerable attention also was devoted to documenting American trends in a variety of biological subdisciplines and periods. Among these works, Gerald Geison’s (1987) edited collection on American physiology, which spanned from the 1850s to the 1950s and featured chapters by Adele Clarke and Louise Marshall specifically on research materials including organisms, is of particular note for our purposes, as it documents themes that come to be prominent in subsequent stages of the history of biology to be reviewed below. In addition, two volumes on twentieth-century American biology commissioned by the American Society of Zoologists and edited by Ronald Rainger et al. (1988) and Keith R. Benson et al. (1991) and Maienschein’s (1991) own book on trends in American biology at the turn of the twentieth century touch on key themes relating to the use of organisms within larger narratives about institutional and national styles, as well as providing close attention to prominent biologists and their practices. In summary, the literature in the history of biology that was published in the 1970s and 1980s did explore the use of non-human organisms, sometimes in detail but as one among many components and typically as another form of instrument or technology along with others. Much of the science explored in this period utilized more than one type of organism (with some exceptions such as McClintock on maize), and hence it is not surprising that no one organism is central to any particular research program or the narratives about it. Perhaps most importantly, this scholarship typically hybridized the methods and traditions of intellectual, biographical, or institutional history with more detailed attention to scientific research and to show less influence of philosophy of biology or science and technology studies. Hence there tended to be less emphasis on themes from these literatures such as the materiality of organisms and how this impacts on research practices, or on epistemological considerations, as compared to the subsequent stages of research on these topics.

Organisms as Units of Study We contend that a turning point in the historiography of biology occurred in the early 1990s and involved focusing on organisms – rather than scientists, theories, problems, institutions, or disciplines – as the unit of, and narrative thread for, historical study. This widespread change in perspective resulted in increased attention to scientific work and practices together with focus on the contingencies, characteristics, and behaviors of individual species particularly those imposed by their materiality. Thus scholarship in this period also placed considerably more emphasis on the agency and specificity of biological materials used in experimental research, especially organisms. This turn in the field was due in part to the growth of interdisciplinary scholarship in studies of biology but also to influences of themes from fields outside of history, notably science and technology studies and particularly sociology and anthropology and philosophy of biology. There are a few early precedents for

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this type of work, such as Allen’s (1975) article which rejects the typical “discovery” account of Morgan and Drosophila (on the historiography of discovery accounts, see Woolgar (1976) and Löwy (1990)) and outlines previous research work with the organism including the social and collective efforts as well as the qualities of the organism itself that led to Drosophila’s adoption. Crucial to the emergence of this historiographical trend was the interdisciplinary volume edited by Adele Clarke and Joan Fujimura (1992) entitled The Right Tools for the Job: At Work in the Twentieth-Century Life Sciences, which grew out of an organized session at held at the biennial meeting of the International Society for the History, Philosophy and Social Studies of Biology (ISHPSSB) in 1989. As the title indicates, the collection’s main focus is on identifying and discussing tools deployed in order to do specific types of work in the life sciences, and several of the contributions focus on particular organisms as “tools.” Drawing on scholarship from science and technology studies and sociology, anthropology, and philosophy of science as well as history of science, the volume’s approach is constructivist and ecological, showing that the conditions of scientific practice are highly specific and situated, as outlined in the introduction (Clarke and Fujimura 1992). Among the articles focused on organisms, of note is Gregg Mitman and Anne Fausto-Sterling’s (1992) exploration of the rise and fall of the flatworm Planaria particularly in C. M. Child’s work and how it became embedded with conceptual, social, and personal assumptions that contributed to explaining its lack of success as an experimental organism (except for pedagogical purposes). A chapter on R. A. Emerson’s work with maize in agricultural genetics by Barbara Kimmelman (1992) illustrates how this organism was “right” for not only scientific and technical reasons but also for various social reasons. In both cases, the authors explicitly use their cases to challenge to scientific and historical representations of Drosophila as an organism particularly well-suited for genetic research. Another piece of scholarship that made key contributions to this trend was the special issue edited by Muriel Lederberg and Richard Burian (1993) in the Journal for the History of Biology. Their mandate was to explore organismal choice, particularly what characteristics make specific organisms suitable for particular kinds of research and how do those qualities evolve and adapt to shift in techniques, questions, and research environments (Lederman and Burian 1993) and which again came out of a special symposium held at ISHPSSB in 1991. In this work, the relationship between the choice of a species and the kind of research produced was conceptualized in at least three different ways. In some cases, the “job” to which an organism is assigned is primary, and the organism is secondary in the sense of being sought and even constructed to fill that particular role. This narrative underlies the contributions by Bonnie Clause (1993) on rat, Robert Kohler (1991, 1993) on Drosophila, and Doris Zallen (1993) on the use of algae for photosynthesis research. In other cases, the “job” is created partially or completely by the features and behavior displayed by the organism in the lab: part of Kohler’s story also fits this picture, as does the contribution by Muriel Lederman and Sue Tolin (1993) on viruses. Still other contributions, notably F. Larry Holmes’s (1993) article on the frog, reject the teleological analysis of organismal use implied by the use of the

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terminology of “jobs” and “rightness” and instead focus on the ongoing processes characteristic of scientific work. He provides an overview of experimental uses of frogs, which enables him to highlight their ability to withstand pain as a major motivation for their adoption as biological materials in physiology. A milestone in this genre was Kohler’s (1994) book Lords of the Fly: Drosophila genetics and the experimental life, whose appearance is widely recognized to have marked an important moment in the evolution of scholarship on research organisms (his approach was foreshadowed in several articles including his 1991 and 1993 described above). Ironically enough, this book returned to the classic organism Drosophila but took an atypical approach, explicitly exploring the material culture and way of life of experimentalists who worked on the fruit fly, which he conceptualizes as their “co-worker.” His simultaneous attention to the technological, biological, and moral aspects of both Drosophilists’ work practices and of the organism itself set a precedent for much history of biology that was to follow particularly due to its ecological vision and especially scholarship exploring experimental organisms. The book emphasizes the commensal relationship between organisms and scientists who use them, including the idea that laboratory organisms undergo a form of “domestication,” hence drawing on diverse historiographic trends in the more general literature on human-animal relationships (e.g., Serpell 1986; Ritvo 1987) as well as more ecological approaches to history (e.g., Cronon 1991; Worster 1990). Kohler also saw his work as a call for scholars to avoid the technicalities and specificities which he viewed as endemic within histories of special sciences at that time, including the biological sciences, thus providing a model for development of more “general” histories of science through shared focus on experimental practices. This literature grew in dialogue with the more general trend during this period toward attending in more detail to material cultures in scientific practice and the dynamics of experimentation, particularly in history and philosophy of science (e.g., Hacking 1983; Shapin and Schaffer 1985; Gooding 1990). As noted in Andrew Mendelsohn’s (2003) dialogical paper “Lives of the Cell,” this scholarship raised a range of innovative questions, created creative tensions, and had an overall revolutionary effect on historical and philosophical discussions around the choice and use of organisms in research. Within scholarship specifically focused in the life sciences, this trend arguably culminated in Hans-Jörg Rheinberger’s (2010) proposal to use whole experimental systems as units of analysis, particularly notable for our purposes because several of the systems he discusses in detail center around the choice and handling of specific organisms (such as Ephestia, Pisum, Eudorina, and tobacco mosaic virus). Other important influences included the debates on the epistemic role of standardization (usefully reviewed in Timmermans and Epstein (2010)) and expanded discussions and problematizing of organisms as “boundary objects” (Star and Griesemer (1989); for an application of this concept, see Keller (1996) on Drosophila embryos’ transformation from transitional objects to boundary objects). It also is clear that many historians of biology integrated consideration of the Latourian emphasis on the active role of non-human actants within social networks (what

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Latour (1993) calls “heterogeneous engineering”) into their accounts, as well as actively considering literature from the history and sociology of technology that encouraged viewing experimental organisms as technologies which undergo construction in a similar way to any form of scientific instrumentation (e.g., Bijker et al. 1987). Another relevant concept was Steven Shapin’s (1988) application of E.P. Thompson’s term “moral economy” to scientific workplaces, which encouraged other historians to relate the use of research materials to the social, ethical, and institutional norms and conditions of laboratory work. An essay on research materials in the reproductive sciences by Clarke (1995, revised from her contribution to Geison ed. 1987) similarly stresses the need to develop a richer concept of “ecology of knowledge” relating to the organization of research materials (including organisms) and the development of techniques to study them, hence forcing more attention to the social and material conditions associated with knowledge production in science. While particularly influential within history of biology, the tendency to focus on organisms as a thread for historical narratives also came to be popular in cultural and intellectual approaches to history in this period. This extensive literature typically forgoes in-depth discussions of the role of particular organisms in research, focusing instead on their importance in trade and food cultures (see, e.g., the many monographs devoted to the potato, none of which explores its use in scientific research, such as Salaman et al. (1985), Zuckerman (1999), Reader (2011), Smith (2011), and Gentilcore (2012)). A notable exception is the Reaktion series on “biographies” of animals, which includes brief discussions on organisms as experimental subjects in the case of the rat (Burt 2006), chicken (Potts 2012), octopus (Schweid 2013), leech (Kirk and Pemberton 2013), and rabbit (Dickenson 2013). Jim Endersby’s (2009) A Guinea Pig’s History of Biology exemplifies the fruitfulness of combining approaches from history of science and cultural studies to create a narrative about organisms, as demonstrated by the wide appeal that the book generated well beyond traditional academic audiences. The use of organisms as a main thread also was common among historically oriented narratives by scientists themselves, sometimes in collaboration with historians (e.g., Gurdon and Hopwood 2000 on Xenopus; Laubichler and Davidson 2008), although these accounts tend to be much more internalistic and focused on pragmatic issues within the lab. To name just a few, these include S. G. Ernst (1997) on sea urchins, Francois Jacob’s (1998) more popular book on part of his work with mice and flies, and John T. Bonner (1999) on slime molds. Chris Somerville and Maarten Koornneef (2002) on Arabidopsis is an interesting exception, as they draw particular attention to social dynamics and community building on a global scale for Arabidopsis, reflecting the conscious effort done within the community to advertise and expand the range of research uses for the model plant. In summary, a key stage in the historiography of biology for those studying non-human organisms was the turn to using organisms themselves as the central units of historical study. This trend was accompanied by increased attention to scientific work and practices and greater emphasis on the provenance, characteristics, and behaviors of individual species and opened up

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conceptual spaces and new questions that have come to characterize scholarship in this area.

Organisms in and as Research Practices Historical scholarship in the life sciences since the mid-1990s has tended to explore the diverse practices of scientists who work with various organisms with focus on a range of themes, including the ways in which these organisms are used as resources for longer-term collaborative projects, how they are integrated into specific projects, the details of particular setups within experimental settings and their implications, and the broader cultural and institutional contexts in which these organisms are employed. In a sense, this trend can be viewed as a hybrid of the first two outlined above, as the scholarship tends to be attentive to the research practices, methods, and technologies adopted in biological laboratories and to the tensions and opportunities created when importing and using organisms in those environments. Rheinberger’s (1997) idea of using biological materials of various types as “things to know with” was particularly influential for scholars working on these topics. Thus this scholarship tends to combine the analysis of institutions, scientists, and fields within biology in some cases national trends with close attention to the organisms themselves within the context of the experimental work being performed. In addition, the emergence of this trend can be argued to have run in parallel to the developments in the life sciences themselves, particularly within molecular biology where the concentration of resources around few model species was increasingly overtaking research emphasizing variation and biodiversity (for historiographic work on this issue, see Churchill 1997; Laubichler 2000; Geison and Laubichler 2001; Ankeny 2010). As outlined in the introduction to this paper, this period saw the rise of the term “model organism” within the biological and biomedical sciences (particularly due to the Human Genome projects and their associated large-scale genomic sequencing efforts) to refer to species used as gateways to understanding fundamental processes in ways that can then be generalized to other organisms and fostered numerous analyses and critiques of these concepts (Gest 1995; Bolker 1995; Ankeny 2000; Gilbert 2009; MacLeod and Nersessian 2013). Key individual species were the primary focus of scholarship in this period, with a tendency to examine the rise and use of canonical model organisms within their institutional and community contexts. For example, Karen Rader (1998, 2004) traces how standardized mice came to have the prominence which they now have in contemporary biomedicine, including the methods for balancing their natural attributes with laboratory-induced features, with particular focus on the Jackson Laboratories; this work is particularly important due to its stress on processes of standardization and their effects. Soraya de Chadarevian (1998) and Rachel A. Ankeny (2001, 2000) explore the development and use of the nematode Caenorhabditis elegans as an experimental organism in genetics, developmental biology, and neurobiology in the context of the Cambridge Laboratory of Molecular Biology, with special attention to various aspects of community formation (see also

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de Chadarevian (2002)), a theme that continues in Sabina Leonelli’s (2007) investigation of the use of the mustard cress Arabidopsis thaliana, which traces how the material features of the organism in particular together with the growth in and efforts of the international community associated with this work together made Arabidopsis the most well-researched plant model organism. Marcel Weber (2004) examines the material characteristics and experimental culture (particularly the technique of chromosomal walking) that made Drosophila into the most successful model organism in the 1980s for positional cloning. A second theme in this period is how organisms can be used in projects which utilize diverse disciplinary perspectives or bring together several fields. For example, Rheinberger (2000) examines Alfred Kühn and colleagues’ work in Germany on the flour moth Ephestia, particularly his use of this organism in projects derived from a range of disciplinary perspectives or research traditions including embryology, physiology, genetics, and biochemistry and the hybridization of these fields within their experimental system. Angela Creager (2002) examines Wendell Stanley’s laboratory’s use of tobacco mosaic virus and how the experimental techniques and instruments that they developed came to be used by others studying TMV and beyond in other fields of research. Christopher W. P. Lyons and Karen-Beth G. Scholfthof (2015) follow the evolution of the wild grass Brachypodium distachyon to its current status as a model organism, drawing together distinct trajectories which ground contemporary research on it, including studies of taxonomy, host-pathogen interaction, and biofuels. Some scholarship stresses interactions between organismal use, experimental practices, and the resulting theories. For instance, Judy Johns Schloegel (1999) explores Tracy Sonneborn’s research with the protozoan Paramecium aurelia and how his detailed knowledge about this organism helped to shape his defense of it as a research organism as well as his advocacy of a more unifying definition of species. Schloegel and Henning Schmidgen (2002) examine the use of unicellular organisms in late nineteenth- and early twentieth-century psychophysiological research and how these organisms forced researchers to change their views on the ontological status of these organisms which in turn had major impacts on fundamental concepts in psychology. Scott Gilbert (2009) analyzes the demands that research with different species imposes on conceptualizations of evolutionary developmental biology and its relation to the rest of the life sciences, and V. Betty Smocovitis (2009) shows how the adoption of the weed genus Crepis as biological material at Berkeley in the 1920s and 1930s grounded a radical rethink of systematics in the context of the modern synthesis. John T. Bonner’s work with the slime mold, Dictyostelium discoideum, and how it contributed to his views on developmental theory and practice, is explored in detail in Mary Sunderland (2011). Robert Meunier (2012) traces the use of zebrafish in developmental biology from the 1970s on and its use as a platform for mechanistic models. In general, this literature emphasizes the practical, biological, and epistemic implications of importing organisms into a new ecosystem: the laboratory. The advantages, disadvantages, and peculiarities of these types of moves have been well-discussed in the scientific and philosophical literature and in historical

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scholarship with reference to specific case studies. Scholars have worried in particular about the extent to which organismal features and behavior adapt to experimental environments, which often results in radical changes in the characteristics of organisms, thus potentially compromising any attempts to draw easy inferences from the study of life in the laboratory to knowledge about life in the wild (e.g., Griesemer and Wade 1988; Griesemer and Gerson 2006). A related, though perhaps less prominent, focus is the extent to which specific features of organisms affect the construction and long-term development of laboratory facilities. This concern, which was already voiced in Bruno Latour’s (1993) account of Louis Pasteur’s isolation of germs, has been further developed by Edmund Ramsden and Robert Kirk, who have documented the sophisticated interplay between experimenter’s objectives, organismal behaviors, and the design and modification of the space where animals are kept in the case of rats in behavioral psychology (Ramsden 2011a, b, 2012) and sheep in the investigation of the psychology and physiology of stress (Kirk and Ramsden 2018) and the production of standardized and germ-free animal strains (Kirk 2008, 2012, 2013). The importance of organism choice in the context of discipline building continues to be subject of debate but with special emphasis on the ways in which particular species fit (or not) and shape the demands and affect (or fail to affect) the intellectual and institutional directions of given communities and areas of research. Daniel P. Todes (1997, see also his book 2001) looks at the systems of production in Pavlov’s laboratories, including the “dog technologies,” and traces their effects on the resulting experimental practices, relations within the laboratory, physical structures, and products. Cheryl Logan (2001, 2002) traces the use of rats in a variety of experimental settings related to psychology, with particular attention to the interplay between the practices associated with the use of these organisms and the underlying conceptual and experimental assumptions that accompanied such research; a key theme in her work also are the tradeoffs between the benefits of standardization and the potential limits on making claims that are more generalizable. A growing trend particularly in the 2010s has been attention to other areas of human activities in which organisms are actively co-opted but which have relevance for experimental science. Aside from the abundant work on Darwin and his pigeons (e.g., Secord 2011), documentation of the intersections between animal fancying and experimental scientific research remains relatively limited, exceptions being Christian Reiß’s (Reiß 2012) work on axolotls where he investigates the relation between the popularity of aquaria and nineteenth-century zoology in Europe and Endersby’s (2013) contribution on the public appeal of primroses, Hugo de Vries’s organism of choice, in relation to the establishment of early twentieth-century mutation theory. Sheep breeding and its contributions to knowledge of heredity are examined in Roger Wood and Vítêzlav Orel (2001), with particular focus on the activities of nonscientist sheep breeders. The increasing amount of research on dogs, which includes work by Kirk and Ramsden discussed above (as well as Kirk (2014)), is particularly interesting insofar as it documents the research implications of these animals’ roles as companions as well as “workers” engaged in the provision of specific services (such as assistance for the blind, in hunting, and as sniffer dogs for

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smelling dangerous or illegal substances in law enforcement). A key early example which fruitfully exploited the intersection of history of biology and institutional and agricultural history is Deborah Fitzgerald’s (1990) work on hybrid corn. Harriet Ritvo’s work (e.g., 2010) also is of note, as an example of broader historical explorations in the growing field of animal studies, in this case focused on the importance of non-human animals to human culture. In short, organisms became an extremely popular focus of historical studies in the 1990s onward, in parallel with trends in the biological sciences that emphasized using individual species as cornerstones for research programs. Key themes included how and why certain organisms come to be utilized, particularly as model organisms, and the importance of processes of standardization and community building, how organisms can be used to bring together several disciplinary perspectives or fields, and how work with particular organisms helps to shape underlying concepts or theories in the life sciences.

Current Directions: Comparative, Quantitative, and Integrative Work Beyond the Western Lab Environment There are a number of ways in which historians are building on the existing sophisticated work carried out on individual species over recent decades. To begin, there has been a recent push toward broader methodological approaches, such as the integration of quantitative analysis and related large data collection with the almost exclusively qualitative scholarship based on in-depth interpretative study of predominantly textual sources that has dominated the field to date. Quantitative methods can include citation and network analysis, as well as data-intensive forms of research such as text and data mining from digital archives and statistical records, which can be supported by computational analysis and typically require the commitment of larger research groups (including individuals with skills from other disciplines, such as statistics, computer science, and information systems). Examples focused on experimental organisms include early work by Churchill (1997) with regard to quantitative tracking of institutions and training trends and Michael Dietrich and colleagues’ (2014; Crowe et al. 2015) use of extensive data mining to document long-term, broad trends in the choice and use of and funding support for specific organisms within particular fields. Katherine McCain (1991) pioneered the use of citation analysis in correlation with detailed accounts of the origins and development of research within specific labs, an approach we feel could be usefully applied to other areas and periods, as well as application of more “computational” approaches as recently advocated by historians of science (e.g., Laubichler et al. 2013). Another approach to promoting methodological innovation with regard to historical studies of organisms and the research associated with them is to integrate insights from the history and philosophy of biology more explicitly with literature and concepts from cultural and intellectual history, sociology, and anthropology, including continental approaches (which, as we noted in our introduction, have

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largely proceeded without intersecting with scholarship in the history and philosophy of biology). A key early example which blended historical and sociological approaches, albeit using work with organisms largely in a nonexperimental context, was the paper by Susan Leigh Star and James Griesemer (1989) discussed above in which the terminology of “boundary objects” was coined. More recent examples include Nick Hopwood’s (2015) article on public views on amphioxus (Branchiostoma); Sarah Franklin’s (2007) book on Dolly which contains useful historical material on the economic, social, and scientific significance of the creation of this clone, presented within a broader anthropological frame that seeks to contextualize the significance of this animal’s creation; and Kirk’s (2008) work on guinea pigs, which blends history of science with economic and social history to analyze how specific laboratory organisms were sourced and what the resulting consequences were in terms of their experimental handling and the knowledge thus produced. More emphasis on the range of financial and economic situations in which organisms are selected, sourced, and disseminated for experimental work is also crucial to comparing the handling of specific species and identifying patterns attached to specific cases. This gap is particularly evident in the case of research on mouse, where the commercial value attached to transgenic mice over the last two decades has fundamentally altered the directions and dynamics of molecular biology, particularly in its medicine-facing incarnations, as documented, for instance, by Gail Davies (2013). Aside from methodology, what has become evident of late is the need to diversify the historiographical foci employed in historical research on experimental organisms. For a start, the majority of literature to date has focused on genetics and molecular biology, thus leaving aside the numerous roles and uses of non-human organisms in other biological subdisciplines. It is particularly critical to pay more detailed attention to the use of experimental organisms in fields such as immunology and psychology, as well as in emerging fields such as synthetic biology, which have so far mostly been analyzed by philosophers (see, e.g., Fagan’s (2013) work on the use of organisms in stem cell research). Luis Campos’ (2015) book on radium, for instance, mines a treasure trove of new material on hitherto unacknowledged contributions to plant development and evolutionary engineering by researchers working on various organisms. Furthermore, some of the most philosophically inspired approaches to experimentation on non-humans are starting to look beyond research on individual species and thus increase the body of scholarship that compares the handling of different organisms in various types of experimental contexts, as well as work across species and on groups as units of analysis, including not only research on population science but also on entomology, microbiology, zoology, ecology, and other fields concerned with dynamics outside, rather than only within, individuals. Examples include Ankeny and Leonelli’s (2011) comparison of the history and use of model species, Rasmus Winther et al. (2015) on modeling populations, Griesemer (2015) on the role of model taxa, and Maureen O’Malley (2013) and Alan Love and Mark Travisano (2013) on microbial cultures. All of these topics are in need of more extensive and detailed historical research.

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Such historical approaches can sometimes require conceptualizing research environments as going beyond the lab and as including fields, zoos, clinics, hospitals, museums, and other places of relevance to the study of non-human organisms for scientific purposes. This type of focus is already well-established in environmental and agricultural history; a prominent example is Kathy Cooke’s research on early twentieth-century breeding research on chicken and its intersections with farming practice (1997). However, these approaches have yet to be fully exploited in historical studies of the role of experimental organisms in research. Some prominent scholars have started to make inroads into exploring the use of organisms in these environments, although for the most part their focus remains on broad cultural, scientific, and institutional trends rather than on the organisms themselves. For instance, Kohler (2002) mentions several organisms in his exploration of research at the border between lab and field and yet does not specifically discuss their contributions to shaping these boundaries. Similarly, Harwood (2005, 2012), Berris Charnley (2011), Giuditta Parolini (2015), and Dominic Berry (2015), among others, have probed research cultures and practices at the intersections of farming techniques and knowledge, agricultural policy and governance, and biological research (particularly Mendelian genetics), yet this strand of research does not tend to place emphasis on the role that specific species of plants, and particularly wheat and barley, played in British and German agriculture-focused research. Looking instead at biomedicine, Ilana Löwy and Jean-Paul Gaudillière have documented the use of animal models in hospitals and clinics. Löwy (1992) in particular devotes considerable attention to the conditions under which specific organisms such as rabbits, guinea pigs, and bacterial strains were handled in order to yield experimental results that could inform theories and practices regarding vaccination on humans. In contrast, Gaudillière mentions experimental organisms in his discussion of different types of biomedical research (e.g., 2008) but does not explore more detailed questions concerning the role of non-human organisms in medical environments. More historical work also is being carried out on the role played by zoos and botanical gardens in promoting experimental and observational research on non-human organisms. Research focused on contemporary history includes Carrie Friese (2013) on the cloning of endangered animals in zoos for conservation purposes and Lene Koch and Mette V. Svendsen (2014) on capuchin monkeys initially used for psychiatric research and then brought into a private zoo, where they became subjects of an altogether different type of experiment concerning conditions of life in captivity. Finally, another area in need of expansion concerns the handling of organisms in natural history and other types of museums, a strand of research which is exemplified by Erika Milam (2009), Sunderland (2013), and of course Star and Griesemer (1989). Finally and perhaps most importantly, Anglophone histories of experimental organisms are broadening beyond Europe and North America to include more research on scientific practices with experimental organisms in the former Soviet Union and Russia, Asia, South America, and Africa, as well as continuing work in other languages that document national episodes and trends (of course there has been some previous excellent scholarship on these locales, some of which has been

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discussed above). The globalization of historical outlook is becoming increasingly prominent across a range of subfields and presents fascinating methodological questions concerning the meaning and possible methods for making cross-cultural comparisons, the difficulties of mastering the relevant languages and primary sources, and the complex work of contextualization involved in investigating multinational research programs (see ▶ Chap. 15, “Women in the Historiography of Biology”). Examples of recent relevant work focused on non-Western settings include emerging research on nineteenth- and twentieth-century Russian biology, particularly botany and its relation to agriculture (Loskutova and Fedotova 2015), and Lisa Onaga’s (2010) analysis of the use of silk worms in early twentiethcentury Japan. Illustrating the power of cross-national comparisons, the recent volume New Perspectives on the History of Life Sciences and Agriculture (2015), edited by Denise Phillips and Sharon Kingsland, brings together contributions spanning over two centuries of research in Germany, France, Italy, Russia, the United States, Japan, Austria, Java, and China.

Conclusion Our journey through the Anglophone historiography related to the experimental use of non-human organisms, though limited in scope and timescale, emphasizes the extensive interest paid by historians of biology to this topic and the variety of approaches and styles used to pursue it. In closing, we would like to address a question that lies at the core of such work and its extension into the future: whether and how such literature is still relevant and for what? In other words, what is the point of this work, and why is it still an active research area, given all the scholarship that has already been published which elucidates the roles and impacts of non-human organisms in biological experimentation? We contend that documenting the history of biology while paying attention to the organisms used as experimental tools, and the reasons and circumstances for their use, is important for at least five reasons. First, it helps to unravel the material basis of theoretical developments and the extent to which practical and concrete concerns shape and guide experimentation and its outputs. Second, it forces focus on questions concerning where, be it geographically, institutionally, culturally, and otherwise, research is being conducted, since the choice and use of non-human organisms vary dramatically depending on social norms and availability (e.g., whether researchers can source the organism in question locally or need to procure it through trade or exchange). Third, it brings to the fore questions about the limits and opportunities related to standardization practices, attempts to enact experimental control, and the status of technology and instrumentation in scientific practice, all of which are at their most complex when dealing with living entities. Fourth (and related), it encourages a reflexive outlook on biological practices and the paradoxes of studying life by interfering with it and isolating individual organisms or groups from their wider environments.

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Finally, normative questions about whether and in what sense research on a particular experimental organism is appropriate, and for which purposes, have clear social, political, and economic implications for how science is conducted and how knowledge is constructed. Historical work exploring the roots of organismal choice and its impact on research can inform the conduct of contemporary science and how we understand the underlying epistemic structures and scientific practices relating to this field of research. For example, many criticisms have been raised by scientists as much as those who study science about the overly limited selection of reference species exemplified by the pursuit of model organisms for biological experimentation (Bolker 1995; Davies 2007). Historical research has corroborated these critiques, by showing how the model organism concept has been prominent in biological research agendas over the last three decades, making it more difficult to pursue biological research on organisms not considered to be model organisms and thus potentially limiting research on questions that do not fit these particular models (but see Dietrich et al. 2014; Crowe et al. 2015). At the same time, attention to historical sources can help to counter some of the accusations of reductionism and intellectual myopia leveled to the founders of model organism communities, since it reveals that these scientists’ willingness to temporarily sacrifice attention to biodiversity was in most cases a strategic choice to allow the building resources and knowledge toward a goal of a truly integrative biology and was accompanied by the intention to expand the number of species under investigation as soon as practically possible. In this respect, it is also important to note that a large body of work in philosophy and social studies of science has also addressed the role of organisms in biological research, focusing on questions such as how knowledge is created using non-human organisms, what such organisms represent, whether and in which sense they should be regarded as models, how processes of idealization and abstraction contribute to and warrant their use, when and why arguments about projectability of data and other results are well-founded, what the relationship is between such organisms and the experimental contexts within which they are utilized, and how the epistemic structures and shared scientific practices within the communities of scientists focused on these organisms influence the ways in which the research is conducted and how these organisms are understood. As we noted in the previous section, some of this work has made a significant impact on historiography, pushing historians to pay more attention to the framing and objects of their scholarship. Accordingly, we encourage more productive dialogue between historians and philosophers of biology on this topic, as well as better awareness among historians in this field of related useful discussions in the social sciences. A recent debate in anthropology, for instance, concerns the idea of “multispecies ethnographies” focusing on hitherto undocumented encounters among “non-charismatic” species (Kirksey and Helmreich 2010), which are those types of organisms defined in opposition to the “charismatic” species such as whales and tigers that are used by environmental activists as flag bearers for conservation concerns. Beyond this single example, there is a vast scholarship on human-animal relations and intersections between species in anthropology and cultural studies which could be explored to see whether some of

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these ideas can be co-opted to expand the quantity and variety of species and research settings under investigation and also the impacts of human-non-human organismal relations on biological research. In turn, new historical work can provide critical starting points for conceptual discussions of interest to all students of the life sciences, whether they are philosophers, sociologists, geographers, or anthropologists. A case in point is the potential expansion of historical work on the use and handling of microbial communities within and beyond the laboratory, which was discussed in the previous section. Such work could open up questions regarding what counts as an “organism” in the first place, how organisms are grouped and standardized, and what impact these practices have on their representational power vis-à-vis other organisms or phenomena, and indeed how organisms are described, classified, counted, and conceptualized across periods, disciplines, and experimental cultures. Acknowledgments Funding via the Australian Research Council Discovery Project (DP160102989) “Organisms and Us: How Living Things Help Us to Understand Our World” (2016–20)

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Leonelli S (2016) The disruptive potential of data publication. Notes Rec: Royal Soc J Hist Sci 70:20160036. https://doi.org/10.1098/rsnr.2016.0036 Leonelli S, Ankeny RA (2012) Re-thinking organisms: the impact of databases on model organism biology. Stud Hist Phil Biol Biomed Sci 43(1):29–36. https://doi.org/10.1016/j. shpsc.2011.10.003 Logan CA (2001) ‘Are Norway rats ... things?’: diversity versus generality in the use of albino rats in experiments on development and sexuality. J Hist Biol 34(2):287–314 Logan CA (2002) Before there were standards: the role of test animals in the production of scientific generality in physiology. J Hist Biol 35:329–363 Loskutova MV, Fedotova AA (2015) The rise of applied entomology in the Russian Empire: governmental, public, and academic responses to insect pest outbreaks from 1840 to 1894. In: Phillips D, Kingsland S (eds) New perspectives on the history of life sciences and agriculture, vol 40. Springer international Publishing, Switzerland, pp 139–162. https://doi.org/ 10.1007/978-3-319-12185-7_8 Love AC, Travisano M (2013) Microbes modeling ontogeny. Biol Philos 28(2):161–188. https:// doi.org/10.1007/s10539-013-9363-5 Löwy I (1990) Variances in meaning in discovery accounts: the case of contemporary biology. Hist Stud Phys Biol Sci 21(1):87–121. https://doi.org/10.2307/27757656 Löwy I (1992) From guinea pigs to man: the development of Haffkine’s anti-cholera vaccine. J Hist Med Allied Sci 47:270–309. https://doi.org/10.1093/jhmas/47.3.270 Lyons CWP, Scholfthof K-BG (2015) Watching grass grow: the emergence of Brachypodium distachyon as a model for the Poaceae. In: Phillips D, Kingsland S (eds) New perspectives on the history of the life sciences and agriculture. Springer, Dordrecht, pp 479–501 MacLeod M, Nersessian NJ (2013) Building simulations from the ground up: modeling and theory in systems biology. Philos Sci 80(4):533–556. https://doi.org/10.1086/673209 Maienschein J (1991) Transforming traditions in American biology, 1880–1915. Johns Hopkins University Press, Baltimore McCain KW (1991) Communication, competition, and secrecy: the production and dissemination of research-related information in genetics. Sci Technol Hum Values 16(4):491–516. https://doi. org/10.1177/016224399101600404 Mendelsohn JA (2003) Lives of the cell. J Hist Biol 36:1–37 Meunier R (2012) Stages in the development of a model organism as a platform for mechanistic models in developmental biology: zebrafish, 1970–2000. Stud Hist Philos Sci Part C: Stud Hist Philos Biol and Biomed Sci 43(2):522–531. https://doi.org/10.1016/j. shpsc.2011.11.013 Milam EL (2009) The experimental animal from the naturalist’s point of view: behavior and evolution at the American Museum of Natural History, 1928–1954. Trans Am Philos Soc 99 (1):157–178 Mitman G, Fausto-Sterling A (1992) Whatever happened to Planaria? C.M. Child and the physiology of inheritance. In: Clarke AE, Fujimura JH (eds) The right tools for the job: at work in the twentieth-century life sciences. Princeton University Press, Princeton, pp 172–197 Mullins NC (1968) The development of a scientific specialty: the phage group and the origins of molecular biology. Minerva 6:828–843. https://doi.org/10.1007/BF01881390 Nyhart LK (1987) The disciplinary breakdown of German morphology, 1870–1900. Isis 78 (3):365–389. https://doi.org/10.1086/354473 Nyhart LK (1995) Biology takes form: animal morphology and the German universities, 1800–1900. University of Chicago Press, Chicago O’Malley MA (2013) Philosophy and the microbe: a balancing act. Biol Philos 28(2):153–159. https://doi.org/10.1007/s10539-013-9360-8 Onaga L (2010) Toyama Kametaro and Vernon Kellogg: silkworm inheritance experiments in Japan, Siam, and the United States, 1900–1912. J Hist Biol 43(2):215–264. https://doi.org/ 10.1007/s10739-010-9222-z

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Parolini G (2015) The emergence of modern statistics in agricultural science: analysis of variance, experimental design and the reshaping of research at Rothamsted Experimental Station, 1919–1933. J Hist Biol 48(2):301–335. https://doi.org/10.1007/s10739-014-9394-z Phillips D, Kingsland SE (eds) (2015) New perspectives on the history of life sciences and agriculture. Springer, Switzerland Potts A (2012) Chicken. Reaktion Books, London Rader KA (1998) The ‘mouse people’: murine genetics work at the Bussey Institution, 1909–1936. J Hist Biol 31:327–354 Rader KA (2004) Making mice: standardizing animals for American biomedical research, 1900–1955. Princeton University Press, Princeton Rainger R, Benson KR, Maienschein J (eds) (1988) The American development of biology. Rutgers University Press, New Brunswick Ramsden E (2011a) From rodent utopia to urban hell: population, pathology, and the crowded rats of NIMH. Isis 102:659–688. https://doi.org/10.1086/663598 Ramsden E (2011b) Travelling facts about crowded rats: rodent experimentation and the human sciences. In: Howlett P, Morgan MS (eds) How well do facts travel? The dissemination of reliable knowledge. Cambridge University Press, Cambridge, pp 223–251 Ramsden E (2012) Rats, stress and the built environment. Hist Hum Sci 25:123–147. https://doi. org/10.1177/0952695112471005 Reader J (2011) Potato: A history of the propitious esculent. Yale University Press, New Haven Reiß C (2012) Gateway, instrument, environment: the aquarium as a hybrid space between animal fancying and experimental zoology. NTM Z Gesch Wiss Tech Med 20(4):309–336. https://doi. org/10.1007/s00048-012-0079-4 Rheinberger H-J (1997) Toward a history of epistemic things: synthesizing proteins in the test tube. Stanford University Press, Stanford Rheinberger H-J (2000) Ephestia: the experimental design of Alfred Kühn’s physiological developmental genetics. J Hist Biol 33:535–576. https://doi.org/10.1023/A:1004858314375 Rheinberger H-J (2010) An epistemology of the concrete: twentieth-century histories of life. Duke University Press, Durham Ritvo H (1987) The animal estate: the English and other creatures in the Victorian age. Harvard University Press, Cambridge Ritvo H (2010) Noble cows and hybrid zebras: essays on animals and history. University of Virginia Press, Charlottesville Salaman RN, Burton WG, Hawkes JG (1985) The history and social influence of the potato. Cambridge University Press, Cambridge Sapp J (1987) Beyond the gene: cytoplasmic inheritance and the struggle for authority in genetics. Oxford University Press, Oxford Schloegel JJ (1999) From anomaly to unification: Tracy Sonneborn and the species problem in Protozoa, 1954–1957. J Hist Biol 32(1):93–132. https://doi.org/10.1023/A:1004464509024 Schloegel JJ, Schmidgen H (2002) General physiology, experimental psychology, and evolutionism: unicellular organisms as objects of psychophysiological research, 1877–1918. Isis 93:614–645 Schweid R (2013) Octopus. Reaktion Books, London Secord JA (2011) Nature’s fancy: Charles Darwin and the breeding of pigeons. Isis 72(2):162–186 Serpell JA (1986) In the company of animals. Blackwell, Oxford Shapin S (1988) The house of experiment in seventeenth-century England. Isis 79(3):373–404. https://doi.org/10.1086/354773 Shapin S, Schaffer S (1985) Leviathan and the air-pump: Hobbes, Boyle, and the experimental life: including a translation of Thomas Hobbes, Dialogus Physicus de Natura Aeris by Simon Schaffer. Princeton University Press, Princeton Smith AF (2011) Potato: a global history. Reaktion Books, London

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Smocovitis VB (2009) The ‘plant Drosophila’: E. B. Babcock, the genus Crepis, and the evolution of a genetics research program at Berkeley, 1915–1947. Hist Stud Nat Sci 39:300–355. https:// doi.org/10.1525/hsns.2009.39.3.300 Somerville C, Koornneef M (2002) A fortunate choice: the history of Arabidopsis as a model plant. Nat Rev Genet 3(11):883–889. https://doi.org/10.1038/nrg927 Star SL, Griesemer JR (1989) Institutional ecology, ‘translations’ and boundary objects: amateurs and professionals in Berkeley’s Museum of Vertebrate Zoology, 1907–39. Soc Stud Sci 19:387–420 Sunderland ME (2011) Morphogenesis, Dictyostelium, and the search for shared developmental processes. Stud Hist Phil Biol Biomed Sci 42(4):508–517. https://doi.org/10.1016/j. shpsc.2011.07.002 Sunderland ME (2013) Teaching natural history at the Museum of Vertebrate Zoology. Br J Hist Sci 46(1):97–121. https://doi.org/10.1017/S0007087411000872 Timmermans S, Epstein S (2010) A world of standards but not a standard world: toward a sociology of standards and standardization. Annu Rev Sociol 36(1):69–89. https://doi.org/10.1146/ annurev.soc.012809.102629 Todes DP (1997) Pavlov’s physiology factory. Isis 88(2):205–246 Todes DP (2001) Pavlov’s physiology factory: experiment, interpretation, laboratory enterprise. Johns Hopkins University Press, Baltimore Weber M (2004) Philosophy of experimental biology. Cambridge University Press, Cambridge Winther RG, Giordano R, Edge MD, Nielsen R (2015) The mind, the lab, and the field: three kinds of populations in scientific practice. Stud Hist Phil Biol Biomed Sci 52:12–21. https://doi.org/ 10.1016/j.shpsc.2015.01.009 Wood RJ, Orel V (2001) Genetic prehistory in selective breeding: a prelude to Mendel. Oxford University Press, Oxford Woolgar SW (1976) Writing an intellectual history of scientific development: the use of discovery accounts. Soc Stud Sci 6:395–422. https://doi.org/10.1177/030631277600600306 Worster D (1990) Transformations of the earth: toward an agroecological perspective in history. J Am Hist 76(4):1087–1106. https://doi.org/10.2307/2936586 Zallen DT (1993) The ‘light’ organism for the job: green algae and photosynthesis research. J Hist Biol 26(2):269–279 Zuckerman L (1999) The potato: how the humble spud rescued the western world. North Point Press, New York

Scientific Biography

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Oren Harman

Contents About Biography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . About Biographers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . The Uses of Biography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Art or Craft? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . The Future of Biography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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“Biographies are but the clothes and buttons of the man,” wrote Twain, “the biography of the man himself cannot be written.” In this chapter, with due deference to Twain, I examine how scientific biographies – of both men and women – have been written historically, ever since the Greeks, and how they have evolved over time, in and out of step with the history of science. Notions of truth, objectivity, and the role of science in society are reflected in biography, as are literary fashions, rendering biography a valuable prism through which to evaluate our ever-changing cultural mores and norms. So, too, are biographers here examined, to consider the alchemy between subject and portrayer. To what uses have biographies been made? Is biography writing an art or a craft? And what will be its future? These and other questions are addressed in this historiography of “scientific biography.” “The trouble with life,” according to Martin Amis, “is its amorphousness, its ridiculous fluidity. Look at it: thinly plotted, largely themeless, sentimental and ineluctably trite. The dialogue is poor, or at least violently uneven. The twists are either predictable or sensationalist. And it’s always the same beginning, and the O. Harman (*) Graduate Program in Science Technology and Society, Bar-Ilan University, Tel Aviv, Israel e-mail: [email protected] # Springer Nature Switzerland AG 2021 M. R. Dietrich et al. (eds.), Handbook of the Historiography of Biology, Historiographies of Science, https://doi.org/10.1007/978-3-319-90119-0_16

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same ending” (Amis 2000, 7). Mark Twain put his finger on a more fundamental conundrum, beyond the concerns of the novelist: “What a wee little part of a person’s life are his acts and his words!” he wrote. “His real life is led in his head, and is known to none but himself. All day long, the mill of his brain is grinding, and his thoughts, not those of other things, are his history. These are his life, and they are not written. Every day would make a whole book of 80,000 words – 365 books a year. Biographies are but the clothes and buttons of the man – the biography of the man himself cannot be written” (Twain 2010, 221). So it is both impossible to write them, and even if it were possible, lives would make little sense. Charming. Ah, but how we love biography!1 Biographies are courted by editors and perennially adorn bestseller lists. They are, for more than a few of us, some of our best-remembered reads, trusted windows into the soul – what we call the “human condition.” Depending on their hero, they are also a window into art, history, poetry, politics and world affairs, military strategy, music, literature, sport, comedy, and philosophy. And if the nearly 5000 lives of scientists, engineers, and medical men and women printed over the past four hundred years in Latin, Spanish, French, Italian, German, Dutch, Swedish, Russian, and English are any indication, then they are a preferred prism into science, too. As Thomas Söderqvist, a student of the genre, has shown, the first vitae of the new men of science – Copernicus, Tycho, Peiresc, Galileo – appeared in the seventeenth century, and we have never looked back. It was ten a year by the end of the nineteenth century, and is ten times that today. Add the innumerable obituaries, entries in biographical dictionaries, éloges in academy publications, and memorials, and you have a metascientific genre that exceeds almost all others, giving philosophy of science and history of science a run for their money (Söderqvist 2007b, 1–3).2 A funny thing, then, that scientific biography, as a genre, has attracted relatively little comment, in relation to biographical writing in other fields to be sure.3 It has paled in relation to the preoccupation with most thematic topics in the history, sociology, and philosophy of science, technology, and medicine, too – disciplines, societies and academies, museums, expeditions, laboratories, clinics, “the field,” 1

A 2011 Harris poll found that 76% of Americans who read at least one book a year report reading both fiction and nonfiction, with 29% of the nonfiction accounted for by biography. 2 Söderqvist goes so far as to state: “It is not too far fetched to suggest that scientific biography as a whole may in fact have had a stronger cultural and political impact than any other genre of metascientific writing in the last four hundred years.” 3 More attention has been afforded to literary and art biography, even biographies of economists, as genres. See, for example, Backschneider (1999), France and St. Clair (2002), Weintraub and Forget (2007), and Cline and Angier (2010); also, as examples of reflection coming from practitioners of the genre, Holroyd (2002) and Hamilton (2007). There are academic groups advancing biography writing and research, such as the Zentrum für Biographik at Humboldt University, the dedicated Journal of Historical Biography based in Cadana and Journal: Life Writing in Australia, and, for the last thirty years, the interdisciplinary quarterly, Biography, from the University of Hawaii Press, featuring articles exploring the “theoretical, generic, historical, and cultural dimensions of lifewriting; and the integration of literature, history, the arts, and the social sciences as they relate to biography.”

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science and cinema, science and literature, diaries, archives, paradigms, women – you name it. More prevalent have been bibliographical essays on the quickly growing biographical literature, descriptive rather than analytical, and almost entirely lacking in reference to biography as an historical category in its own right.4 But this is beginning to change.5 The “presence/absence” paradox, as it has been called, is slowly being redressed.6 Why do we love biographies so much? After all, while D’Israeli may have admonished to read “no history: nothing but biography, for that is life without theory” (D’Israeli 1832), Kundera was more tough minded (and probably closer to the truth): Biography, he wrote, is “an artificial logic imposed on an incoherent succession of images” (Kundera 2011). It is a question that may have as many answers as there could be biographies, and perhaps best left to such hearts and minds as described by Twain. After all, it’s the rare description of the reason for loving something, or someone, that captures it better than the actual love itself. But there are words to spend on scientific biography, nevertheless. It’s history, and changing uses over time, principally, before the questions raised by its craft, not least the challenges faced by its craftsmen and women, and its horizons. Biography is not only a sign of the biographer but a sign of the times, a weather vein by which to measure the changing intellectual temperature – in fact, a fascinating window into the history and philosophy of science as evolving disciplines. And so we turn first to biography as genre, seeking to lay foundations for our discussion.

About Biography Like the United States, or ice cream, biography has a history, and that history can teach us a lot about the development of views surrounding science. In particular, looking closely at the evolution of the genre can shed light on how different cultures at different times answered the question: What is science? And, What is not science? So, too, can it trace an oscillating trajectory concerning whether scientific thought is considered to be independent of politics, and of culture, of time, and place, whether such as thing as objective “truth” exists, and what the role of the scientist is or should be in society. Alongside such considerations, the genre of scientific biography invites reflection on how we write about science and scientists, artistically, and why that matters. And so, from the outset, the biographical genre addresses itself to two kinds of concerns: the first contextual, having to do with science and scientists as agents and 4

A good example is Hessenbruch 2000. One exception is Fullmer 1967; another is Hankins 1979, further discussed below. 5 Besides the above exceptions (Söderqvist 2007a, b; Govoni and Franceschi 2014), examples include La Vergata 1995; Shortland and Yeo 1996; Richards 2006; Harman 2011a. 6 Thomas Söderqvist 2007a, b deserves credit for breathing life into the subject. Biography, however, continues to be virtually ignored as a historiographical category in science studies and STS. An exception is Kragh 1987, and more recently, Kragh 2015. Further exceptions are discussed below.

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subjects of history, the second aesthetic, having to do with how lives and the ideas entwined in them are expressed and communicated. These have been called the cultural-historical poetics of the genre, on the one hand, and its literary poetics, on the other hand (Söderqvist 2007). An appreciation of biography flutters like an intoxicated butterfly between these two linked considerations, for as Adam Gopnik has remarked, “Whatever our official pieties, deep down we all believe in lives. . .. if you ask a cultural-studies maven who believes in nothing but collective forces and class determinisms how she came to believe in this doctrine, she will begin to tell you, eagerly, the story of her life” (Gopnik 2004, 90). How then have scientific lives been told throughout history? The question beckons, too, in light of the fact that the term “scientist” was only invented – by the English polymath William Whewell – in 1840. Let’s begin in ancient times. Romans and Greeks very much enjoyed writing the lives, or Bioi, of fellow countrymen (very seldom women) of note, but these represented a set of “overlapping traditions, embracing works of varying form, style, length, and truthfulness” (Pelling 1996, 241). Usually bioi were a celebration of someone’s exemplary life, oftentimes including a section on his opinions, or doxai, oftentimes detailing the “succession” of teachers and influences. In many cases, as in Plutarch’s Parallel Lives of Greek and Roman heroes, they were a form of “aretology,” paeans to virtue (arête) to be used for moral instruction – a display of the rewards of duty performed, the traps of ambition, the fall of arrogance (see Duff 1999; Hadas and Smith 1965). But eulogies, encomia, bioi, and “doxographical” writings bled into each other with little demarcation; beyond the common themes of celebration, moral edification, and intellectual lineages, there is no well-defined biographical genre of which to speak. In the event, returning to Whewhell, a distinction between a “philosopher” and what we today call a “scientist” is difficult to make in Greco-Roman antiquity (Lloyd 1991, 301). Still, there were certainly ancient men who addressed their inquiries to the natural and physical world, to the mysteries of the heavenly bodies, and the joys of mathematics. Lisa Taub has taken a close look at one of them, Pythagoras, according to Alfred North Whitehead the founder of European philosophy and mathematics. Comparing three ancient bioi of her subject – written some seven centuries after his death in the fifth century BCE by Diogenes Laertius, Porphyry, and Iamblichus, respectively, Taub shows nicely how the writing of ancient lives often served as a way for a particular school of thought to construct its own intellectual history. In the case of Pythagoras, this included both a scientific/mathematical and religious/ethical philosophy, aimed at defining and strengthening the school by advocating a Pythagorean lifestyle, especially in Iamblichus. Characteristically for ancient bioi, the past is harnessed for the present, and the future. And so the intellectual and aretological aspects are not always easy to disentangle, nor is historical record from myth and lore: Alongside the claim that it was Pythagoras who first introduced weights and measures into Greece, Laertius tells us that once he disrobed, Pythagoras’ thigh appeared to be gold, and that “after two hundred and seven years in Hades he has returned to the land of the living.” Porphyry and Iamblichus’ accounts are often interpreted in the light of the rise of the Christian gospels, and the Neophythagorean attempt to counter their

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hero with a hero of their own. No wonder Iamblichus starts with a lineage tying his protagonist to Zeus (Taub 2007, 25). If many ancient renderings are protreptic rather than Rankean, Medieval lives of natural philosophers, such as accounts of the learned men of the Golden Age of Islam, or of Latin saints, lean heavier on hagiography, to be sure, than on contextualization. As the Renaissance ushered in the Hellenic tradition of writing secular lives, hagiography became wedded to the new humanistic sensibility, including writing on women.7 But it was a tradition of orationes funebres, in the manner of eulogies of university professors, and memorials in biographical dictionaries, that dominated the telling of sixteenth- and seventeenth-century scientific lives (Hizlerus 1566; Adam 1605). It is perhaps with Descartes’ Discours de la Méthode, published in 1637, that we most strongly confront, immediately and strikingly, the chasm between our own conception of identity and those of earlier ages.8 The tones of the author’s selfawareness, the timber of his moral integrity, his approach to responsibility, his notion of personhood, differ starkly from our modern sensibilities in this autobiography. Descartes is not interested in presenting an account of his actual comings and goings, thoughts and doubts, warts and virtues, secret and plain-to-view histories. Rather than capturing his own state of mind, he is at pains to describe what the state of mind of a man of his calling should be, indeed to paint the contours of the appropriate mentality of a natural philosopher of his age. Likewise, before him, Bacon in New Atlantis and the Advancement of Learning sought to rebel against an ideal of the philosopher inherited from classical antiquity and the Christian Middle Ages, in which otium, or contemplation, was valorized over negotium, or the active participation in the affairs of state. “The techniques of self-examination and self-investigation opened up by the wholesale attempt to transfer monastic religious values to the population at large,” writes Stephen Gaukroger, “and by the sense that one was responsible for the minute details of one’s daily life in the form of new norms of appropriate behaviour gave way to the possibility of a new understanding of one’s psychology, motivation and sense of responsibility, and of shaping one’s personal, moral and intellectual bearing”(Gaukroger 2007, 47). Both Bacon and Descartes could sense it: the times called for a radical new kind of thinker, not a limp Scholastic cleric but rather a public hero in whom the moral sage and the natural philosopher combine in equal measure to create new works for the betterment of mankind. It was a type, not an individual in the biographical sense we hold today, which is why Descartes begins the Discours by saying: “I intend this text only as a history, or if you prefer, a fable.” Studying this type isn’t exactly engaging in biographical work, but it is closer to biography than the history of philosophy or of scientific discovery (Gaukroger 1995, 2001).

7

On the uses and perils of Medieval biography writing, see Chance 2005. It was Marcel Mauss 1938 who first argued that the very notion of personhood was a Roman invention, and has been developing since.

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Söderqvist narrates what happened next (Söderqvist 2007a, 245–258).9 Gradually, as with Menckenius’ 230-page account of the life of Girolamo Fracastoro from 1731, and later Stoever’s two-volume account of Linneaus from 1792, the short éloges of the seventeenth century were supplemented by true vitae, book-length tomes resembling, at least superficially, what we would consider biographies. These were invariably written by scientists, rather than historians, as were histories of the scientific disciplines – mathematics, chemistry, physics, technology, biology, zoology – now increasingly instituted in the universities and professional societies, such as the British Association for the Advancement of Science, established 1831. Indeed, there was no great separation or tension between historiography of science and biography, other than that those scientists who tended to concentrate on one neglected the other.10 What united them, in the historian of science Arnold Thackray’s estimation, was that, in the wake of the Industrial Revolution, both represented materials “steadily accumulated to explain and justify the role of science in the national life” (Thackray 1980, 10). Science was coming into its own, elbowing for cultural room and challenging the humanities. As the genre grew in scope, so did it diversify, with writers and journalists diminishing the proportion of biographies contributed by scientists (and family members). Increasingly, the somewhat unsmiling format of life-and-letters gave way to colorful portraits aimed at showing the human face of the (sometimes frightening) scientific enterprise. Scientific biography was losing its character of high litterateur just as the history of science was coming into its own as an academic discipline. As the first professional practitioners convened in Paris at the turn of the century to discuss the boundaries of their field, and later created its first journal, Isis, in 1913 in Belgium, they were at pains to promote the history of science, rather than science necessarily, and the influence of scientist writers gradually waned. But could scientific biography account for serious history of science? George Sarton was optimistic. Like August Comte, he believed that science drives history forward; the history of science would chronicle the advance. Except that progress in science was less a function of collective action, as in politics, it was rather “largely the history of a few individuals” (Sarton 1948, 61–63). Genius matters. Indeed, “above all,” Sarton wrote, “we must celebrate heroism whenever we come across it. The heroic scientist adds to the grandeur and beauty of every man’s existence” (Sarton 1936, 45). Biography was important because every detail of the scientist’s life – as long as it was respectably germane to his science – was relevant to “the basic tragedy of mankind – the struggle for knowledge.” “Every human aspect must be considered because this is not simply a scientific matter but a human one. . . ‘Le style, c’est l’homme’” (Sarton 1957, 43). This sentiment was echoed by Douglas McKie, founder in 1936 of the Annals of Science, the British counterpart to Isis which had since migrated to the United States. His pages, he

9

Many of the sources in the following section come from this account. The historiography of science didn’t come into its own until the turn of the nineteenth century (see Lauden 1993).

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determined, would employ biography generously to “recall the story – nay, the romance, of love for natural knowledge and devotion to its pursuit. . . to help evoke a widened sympathy and interest for the subject” (McKie et al. 1936, 103). Thus the history of science recognized biography as a distinct genre, and, to begin with, embraced it. It didn’t last long. Biography was “a defective kind of antiquarianism,” pronounced Herbert Butterfield in the first issue of the Bulletin of the British Society for the History of Science in 1950 (Butterfield 1950). Serious history of science dealt with the synthetic study of historical changes in scientific thought, not with “anecdote,” or, as the American Medievalist Charles Homer Haskins had called it, the distribution of “medals for modernity” (quoted in White 1947, 422). It culled its sources from more than mere personal papers, and considered the notion of the scientist “hero” the legacy of a quaint nineteenth century ideal. Indeed, this was the tone concerning historical biography more generally. When Arnaldo Momigliano published his classic study of biography in ancient Greece in 1971, he told his readers: “When I was young, scholars wrote history, and gentlemen wrote biographies,” and he was right (Momigliano 1993, 1). Now times were changing. And so a cold war generation of historians, born in and around World War I and including I.B. Cohen, Alistair Crombie, Marshall Clagget, A. Rupert Hall, Charles Gillispie, and Thomas Kuhn, became strongly influenced by the sensibilities of the émigré Frenchman Alexander Koyré whose Études Galilèennes exemplified biography that eschewed psyche and style and disposition and temperament in favor of disembodied intellectual history. Scientific ideas were what counted, not personalities (Koyré 1939). It may perhaps not be surprising that many of this generation of historians of science had trained to begin with in the sciences; their insistence on “pure” intellectual history and rejection of “romance and antiquarian curiosity” (Guerlac 1963, 799) reflected a desire to justify their new calling. Prominent historians of science without a science background, at any rate, like Edward Lurie in his Louis Agassiz: A Life in Science (1960), or L. Pearce Williams, in his Michael Faraday: A Biography (1965), were less at pains to make this distinction. Neither were a new breed, the “science writer,” such as the former war correspondent Ronald W. Clarck, who produced popular, and excellent, biographies of Julian Huxley, J.B.S. Haldane, Albert Einstein, Bertrand Russell, Thomas Alva Edison, Sigmund Freud, Benjamin Franklin, Charles Darwin, and Ernst Chain. Biography was gaining serious commercial appeal just as it was being shunned within the profession. When Crombie founded the journal History of Science in 1962, contra McKie, he didn’t invite biographical studies, and the editorial board of the Dictionary of Scientific Biography cautioned contributors in 1970 that personal biography be “intentionally kept to a minimum consistent with explaining the subject’s place in the development of science” (Gillespie et al. 1970, x). The skepticism of biography inherited by the Cold War generation of historians of science was more or less imbibed wholesale by the generation that took its cue from Kuhn’s The Structure of Scientific Revolutions. This had much to do with the newly espoused intellectual aesthetic that cultural values and political ideology undermine individual contribution because individual cognition is derivative of collective

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arrangements. Science studies had gone social. What was the use of delving into the mind of a Clausius if thermodynamics was an invention of an age? And so those who sought to defend the genre, like Thomas Hankins, or Robert Young, did so by fashioning biography as ancilla historiae: lives made sense because they helped contextualize social history.11 The stratagem worked. Adrian Desmond called his magisterial biography of Huxley a “story of Class, Power and Propaganda.” D’Israeli’s quip about biography constituting history minus theory was “good enough, perhaps, for the littérrateur,” he explained, “but it would be a rash biographer today who ignored the new contextual and sociological approaches to science in “Darwin’s Century” (Desmond 1997, 235). It was this approach to biography as a kind of crutch to social history that ushered the genre back into the fold of the history of science. But a further development, more radical, challenged the notion of “disembodied selves,” and hence the Koyréan ideal, most dramatically. Already in the twenties, Virginia Woolf shocked contemporaries with A Room of One’s Own, sparking a debate in which the use of narrative process to tie between autobiography and biography undermined the Victorian epitomes upon which her generation had been reared (Gualtieri 2000). But it would take a different age of feminists and gender studies scholars, in the 1970s, to mount a challenge to social history – and literature – and place biography center stage again. How, after all, can a life be written, when the personal and the social are so inextricably linked? When cognition itself is so radically gendered? (Govoni 2014). It was against this backdrop that science historian Evelyn Fox Keller’s biography of the American cytogeneticist Barbara McClintock, A Feeling for the Organism, became a minor classic in the history of science and in gender studies (Keller 1983). Fox Keller’s protestations that she was not advocating a feminine, but rather a “gender-neutral,” way of scientific thinking in describing McClintock’s challenge to the genetical theory of her day did little to quell the excitement (Keller 2014). Science may be perceived by many of its practitioners and followers as a detached, supranational community of rational men churning out objective knowledge, but there was more to it than that. There were women, and “feeling,” and “intuition,” and, of course, a more modern corpuscularized “social.” Drawing on a new generation’s mastication of Kuhn, historians of science now began to conceive of science as belonging to a place, as dis-unified, in possession of no unique set of values, dependent on tacit knowledge, and “performed” – rather than revealed – by practitioners with diverse moral and cognitive styles. Truth, with a capital T, gave way to credibility, and consensus.12 Whatever else, scientists were not disembodied generators of immaculately insolated ideas (Browne 1998). Lives matter. Gender matters. Practice matters. Relationships matter. Indeed, even things like love affairs and diets matter, “food for thought” in Steve Shapin’s words (Shapin 2010, 5). And so

11

In effect, they suggested that biography become to social history what Aquinas had suggested philosophy is to theology, a handmaiden, or ancilla. See Hankins 1979 and Young 1987. 12 On the ways in which this development reflected, and came on the heels of, changes in science itself and its practitioner’s own perceptions, see Shapin 2010.

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alongside its reinvention as a crutch to social history, the feminist challenge, too, and the evolution of modern sociology of science, saw the genre of biography resurge in the latter part of the twentieth century. We’ll return to McClintock a bit later. Today, biographers of scientists may wonder, together with general historians, what sets biography writing apart from microhistory, or what role biography can play in “lowering the tone” (Shapin 2010; Lepore 2001). But promotion committees in history of science departments no longer look down upon the genre. Gone are the days of Claude Bernard’s “Art is I, Science is We,” or T.H. Huxley’s exhortation that “What we care about in a light is that it shows the way, not whether it is a lamp or a candle, tallow or wax” (Bernard 1957, 42–43; Huxley 1900, 2; quoted in Shapin 2010, 9). To the contrary, owing to its appeal to large audiences, historians of science now lament the fact that professional writers perennially steal the very best stories of lives from under their noses.13 Biography can serve many ends, not just one or two, as we shall see. Before we turn to these, let’s take a closer look at biographers themselves. For as we have begun to appreciate in this potted history of the genre, biographies reveal just as much, if not more, about their writers and their times as they do about their subjects.

About Biographers Who is the ideal biographer? Is it someone who knew her subject or not at all? The question seems useless unless to consider works we have all loved. St. Augustine’s confessions, Pepys’ diary, Rousseau and Franklin’s and Twain’s and Zweig’s and Ramon y Cajal’s and Anne Frank’s and Patti Smith’s self-renderings, all have been read with relish. Being autobiographies, they exhibit the highest form of intimacy, but also tendentiousness, vanity, selective memory, and, in some cases, plain lying. Biography is a different category, and one in which those who have had close personal knowledge of their subjects have produced enduring accounts, from Tacitus on Agricula, through Boswell’s Johnson, all the way to Doris Kearns Goodwin on LBJ. But achieving intimate knowledge of someone else doesn’t necessarily require having known them first hand, as Manuel’s Newton, or Berg’s Lindbergh, or Morris’ Teddy Roosevelt, or John Eliot Gardiner’s Bach prove abundantly (and feel free to choose your own favorites). The wonder of the human brain, as many primatologists and philosophers have remarked, is its ability to slip into another. If firsthand knowledge of their subject is special but not essential for biographers, learning about the world of the biographer himself is often essential for the reader’s understanding of the subject. “Before you study the historian,” E.H. Carr famously tutored, “study his historical and social environment” (quoted in Thackray 1970, 12). This has been termed the “Sobel effect” by Miller (2002) in reference to Dana Sobel’s international bestseller, Longitude 1995. One could add Isaacson 2007, 2011, 2017 to the list, Gleick 1992, Bird and Sherwin 2006, and many others. 13

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My favorite examples of this truism are Bertolt Brecht’s three versions of his play Galileo. Galileo I, or the “Danish” version, was penned in 1938 where the Marxist playwright had been living since 1933 in exile from Nazi Germany. This Galileo is a persecuted intellectual fighting surreptitiously if unsuccessfully against the Inquisition, and the play a liberal defense of freedom against tyranny. “Watch out as you pass through Germany,” Brecht gives his broken hero the line as he passes the heretical Dialogue Concerning the Two Chief World Systems to be smuggled out of Italy, “keep the truth under your coat” (Brecht 1966). But in 1940 Brecht had to flee the advancing German army, settling in the United Stated a year later. It was there that he wrote the second, “Laughton” version of his play (named after the lead actor), completed in 1946, and heavily reworked. For having now acquired a very different conception of science due to its (mis)use and abuse during the war, Brecht’s Galileo morphs into a Marxist defense of a social conception of science against the liberal view that the truth is an end in itself. “The atomic age made its debut in Hiroshima in the middle of our work,” Brecht reflected. “Overnight the biography of the founder of the new physics read differently.” Scene 14, in which Galileo analyses his own recantatory behavior, illuminated the metamorphosis most vividly: Rather than the broken paragon of truth of the “Danish” version, Galileo now becomes a collaborator of the Church, a criminal, someone who has misused his prodigious intellect by subordinating it to religious authority. “I sold out,” the Tuscan astronomer admits to his former pupil Andrea Sarti, “I surrendered my knowledge to the powers that be” (Schroeer 1980). But this was not the end of it. Shortly after the play opened in Hollywood and New York to roaring crowds, Brecht was summoned before the Congressional Committee on Un-America Activities on account of his Communist leanings. His convictions thereafter hardened, the playwright left America, first for Switzerland, and finally to East Berlin, where until his death in 1956, he worked on the third, and most popular, “Berlin” version of the play. It is here that Galileo is most strongly condemned for robbing science of its social significance: “Out of the new astronomy,” Brecht explains, “which initially deeply interested the new class of bourgeoisie since it gave impetus to the revolutionary social currents of the time, he (Galileo) made a sharply defined scientific specialty, which of course, precisely because of its purity – i.e. its indifference to the productive process – could develop with relatively little interference” (Brecht 1957, 205). Galileo I was a broken herald of truth, Galileo II a sell-out to power, and in Galileo III, the playwright finally comes full circle, replacing the starry messenger with his true hero, the people. Brecht was a dramatist, not an historian, but whomever thinks his Galileos have no relevance to scholars is fooling himself.14 “The historian projects into history the interests and the scale of values of his own time,” (Koyré 1958; translation in Schroeer 1980) Koyré counseled, and a brief survey of 25-year, then jubilee, then centennial appreciations of either Relativity or the first World War, or jubilee, centennial, and 150 year accounts of either Darwinism or the Emancipation

14

For a wonderful historiography of Galileo and his trial, see Finocchiaro 2007.

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Proclamation reinforce the point. So much is obvious.15 But with biography, especially, the personality of the biographer becomes all the more important, because writing a life, as many have remarked, is akin to conducting “a love affair, even a marriage” (a life was once panned in the New York Times Book Review for being closer to a “one-night stand”) (Maddox 1999). This isn’t always the case. As Jill Lapore reminds us, biographers can be more similar to crazed stalkers than loyal husbands (Lepore 2001, 134). Indeed, some biographers think of themselves as burglars, opening hidden locks to discover forgotten treasures, or as hunters, or detectives, carefully pursuing bashful facts. Others imagine themselves physicians of memory, or moralists, setting the record straight (Harman 2011b). What is perhaps most crucial, however they fancy themselves, is the fit between biographers’ own proclivities and those of their subjects, a matter of mysterious alchemy. I remember the Pulitzer prize-winning American historian Joseph Ellis being accosted in the press for having lied to his students about his military service (Ellis claimed to have paratrooped with the 101st into Vietnam, when in reality he had passed his service quietly teaching history at West Point). What didn’t the hyenas heckle about: that this was a travesty, that the one-year suspension from Mount Holyoke was a joke, that Ellis’ scholarship was now suspect and needed to be gone through with a steel comb. My thoughts traveled elsewhere: might Ellis have been such a brilliant biographer of Thomas Jefferson precisely because his own character flaws and gnawing self-doubt allowed him to penetrate the maddening contradictions of the American Sphinx?16 The same spirit baffled me when I read Ray Monk’s virtuoso biography of Wittgenstein, and was then disappointed with his competent but rather less sparkling life of Bertrand Russell, in two volumes. The solution had to be that Monk had no sense of humor, I determined, recounting Wittgenstein’s dour aspect and the twinkle in Russell’s eye, proud of my psychological acuity. Except that some years later I met Monk and had to throw my brilliant thesis out the window. The discrepancy between the two biographies remains, at least to my mind, a mystery (Monk 1990, 1996, 2001). Perhaps not surprisingly, the biographer’s own life history often plays a role in his or her choice of subject. Paola Govoni suggests that, despite Fox Keller’s denial, she chose McClintock among other reasons due to the fact that she herself experienced discrimination as a female PhD student in theoretical physics at Harvard in the late 1950s and early 1960s, just as she claimed her subject had earlier, thus providing the impetus to theorize science, in cahoots with McClintock, as “a place where gender could disappear.” Nathaniel Comfort, whose biography of McClintock appeared nearly twenty years after Keller’s, took Keller to task for perpetuating a myth, arguing powerfully among other things based on notebooks and 15

On the factors that tend to stabilize biographical accounts, see Kragh 2007. See the thoughtful essay by the graduate student Bonnie Goodman, “Has Scandal Taken its Toll on Joseph Ellis?,” History News Network, 11/21/2004, http://historynewsnetwork.org/article/8656, in which she suggests that Ellis’s biography of George Washington, which he began writing while on suspension leave from Mount Holyoke, may too owe its success to Ellis being the right person to puncture the myth of Washington as “a man who could tell no lie.”

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correspondence that had not been available to Keller – as well as with the distance from a now deceased McClintock which Keller lacked – that contrary to the claim of an “intuitive” style of science-making, if anything McClintock with her so-called “controlling elements” could just as easily be cast as advancing “masculine” modes of thinking, and that, in any case, she had never been neglected and marginalized, but was rather celebrated, if sometimes misunderstood, and justly challenged. Comfort, Govoni suggests, however self-reflective (Comfort 2001, 2011), may himself have been attracted to the corn geneticist precisely because he was married at the time to Carol Greider, who together with her PhD supervisor Elizabeth Blackburn had helped create in their University of California lab the precise “gender-free” environment which led to the discovery of telomerase and, ultimately, a shared Nobel Prize (Govoni 2014, 20–22). I am not sure whether any of this is true. My lack of certainty, I believe, stems from the fact that I once told myself that I chose the cytologist Cyril Dean Darlington as a subject for a biography because of a shared passion for extrapolating from the very tiny to the very large, and later the troubled polymath George Price, because of a burning quest to discover the origins of altruism (Harman 2004, 2010). Today I think Darlington caught my attention due to his arrogance, and Price due to his vulnerability; extrapolation and altruism seem almost beside the point. I expect to revise my current theory before too long. Why certain minds are drawn to famous figures, of whom many biographies have been written, whereas others seek out unknowns, is also a question for which answers can be offered, and re-offered, ad infinitum. Clearly, different biographers are endowed with different talents. But successful choice of subject remains in some fundamental sense a mystery. Perhaps this is because self-understanding and identity, by their very nature, are shifting sands: like collective memory, they are based on an evolving picture of the past and of the present (Yerushalmi 1996; Luria 1968).

The Uses of Biography Having glimpsed the history and practitioners of scientific biography, let’s consider for a moment the different uses to which the genre lends it name, a consideration that highlights the plurality of the enterprise. In a hair splitting exercise worthy of the great Talmudists, literary scholars have enumerated no less than sixty genres of “lifewriting,” a term that became current in the 1980s (Smith and Watson 2010). Scientific biographies have not sported such a plethora, perhaps, but they too have exhibited impressive variety.17 We’ve already considered two biographies of the cytogeneticist Barbara McClintock, that of Evelyn Fox Keller attempting a history based to a large extent on McClintock’s own perceptions of her career, and that of Comfort addressing itself 17

Nye (2015) suggests that there are three principle forms of biography: the life of the scientist, the scientific life, and lives of scientific collaboration. We consider alternative groupings.

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explicitly to the supposed myth that was born from these perceptions. So biography can deal as much with a life as it can with a reputation, whether gained or selfconstructed or both. The recognition of the importance of self-styling, in fact, has been singled out as an important antidote to what David Aubin and Charlotte Bigg have called the “genius versus context” dichotomy. In their respective biographical work on the French astrophysicist Jules Janssen and his British counterpart Norman Lockeyer, they attempt to rescue what they call “the premature obliteration of individual agency” brought about by this polarization. Victorian London and Third Republic Paris were homes to different conceptions of the utility of science, and the two pioneers of the “new astronomy” had different kinds of casts of mind, to be sure. But self-styling, too, had much to do with the Frenchman’s successful bid for an independent, government sponsored observatory, as opposed to the Englishman’s dithering at the margins of the scientific establishment (Aubin and Bigg 2007, 54). So did Galileo’s fortunes, Brecht’s exegeses notwithstanding, depend on his selffashioning as a courtier, as Mario Biagioli has nicely shown (Biagioli 1994). “A strange and generally speaking rather tiresome farrago of hearsay, imperfect observation, wishful thinking and credulity amounting to downright gullibility” was how Peter Medawar, the 1960 Nobel Laureate in Physiology or Medicine, scored Aristotle’s contributions to biology (Medawar and Medawar 1985). As a recent intellectual biography of the man who “invented science” and laid the foundations for modern biology nicely shows, this, too, was a carefully constructed myth, indeed the original myth of modern science, in which Aristotle was “the giant who had to be slain so that we could pass through the straights of philosophy to reach the open sea of scientific truth that lay beyond” (Leroi 2014, 353). Francis Bacon had played a major part in this, fashioning Democritus as a new philosophical champion of antiquity. So did Descartes, whose bête machine reduced the complex of Aristotelian changes to local motion alone. Medawar was re-enacting this myth for a new generation, but, as an earlier biographer grasped, in order to glimpse the merits of the ancient sage that others have missed, every generation must take pains to read Aristotle anew (Jaeger 1948). This historiographical truism has been at the center of Michael Dietrich’s approach to the twentieth century geneticist Richard Goldschmidt: the ways in which Goldschmidt’s ideas gained longevity and salience for scientists and historians of science through their self-serving representation by the American paleontologist Stephen J. Gould as evolutionary iconoclasm, he once told me, is of greater interest to him than Goldschmidt’s life itself (Dietrich 2011).18 Biographies, as Bernadette Bensaude-Vincent has argued, can be mediators between memory and history in science. “The key notion,” she writes, “which I find useful for biographers and historians, is that texts are framed by other texts and by the writer’s and reader’s cultural contexts. Intertextuality emancipates science biographers from the dull and vain task of fighting against myths and distorted memories. It is an invitation to a more positive attitude of a

18

Another example is Rupke 2005.

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hermeneutical reconstruction of historical realities by displaying the wide variety of their potential meanings. (Bensaude-Vincent 2007, 182)

Legends, in other words, are telling. Rather than combated against, they should be embraced to further understanding. So biographies train their headlights at lives, reputations, myths, or the working out of the relationship between all three. More than anything, however, they have become prisms through which to view an age. A Google books Ngram viewer shows dramatically how biographies that include the words “Life and Times” in their title have skyrocketed ever since the 1940s, whereas those including the words “Life and Letters” have plummeted. It is instructive, too, to see the peak in the latter between the wars, a reflection of the “great man” approach prevalent in those days, now almost entirely neglected.19

This is a victory for social history, hands down, with biography playing the role of ancilla historiae. The physiologist J.S. Haldane, the conservationist and adventurer Carl Akeley, and the geneticist Craig Venter are three (of many) examples of largerthan-life figures who have been cast by their biographers as emblems of their industrial, colonial, and capitalist-entrepreneurial age, respectively (Goodman 2007) (Kirk 2010) (Shreeve 2004), but a true dramatis persona is not always necessary for the task. As Jim Endersby explains in “A Life More Ordinary: The Dull Life but interesting Times of Joseph Dalton Hooker,” the very wearisomeness of a life, at least on one occasion, has been marshaled to celebrate the fireworks of an era (Harman 2011a, 611–631). Embracing its role as prism (Tuchman 1985), scientific biography has been used to probe everything from ideological politics to national and international politics to culture and gender politics to the politics of science itself.20 Lives in the service of

19

https://books.google.com/ngrams/graph?content=life+and+letters%2Clife+and+times&year_ start=1800&year_end=2000&corpus=15&smoothing=3&share=&direct_url=t1%3B%2Clife% 20and%20letters%3B%2Cc0%3B.t1%3B%2Clife%20and%20times%3B%2Cc0 20 There are innumerous examples. For an interesting sample, compare Medvedev 1969, Joravsky 1970, and Nils Roll-Hansen 2004; also, Koerner 2001, Anker 2002, Canales 2015, Ferry 1998, Kevles 1998, as well Chilvers 2007, Selya 2007, Schiebinger 2014, and Abir-Am 2014.

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understanding the intellectual and political sociology of disciplines, too, have proven valuable, both within those disciplines and between them. The categories of “rebels,” “outsiders,” and “dreamers,” for example, have been used to create new historiographies of the modern life sciences, focusing in the first case on the dynamics of dissent of individuals who went against the grain, whether successful or not, in the second on the historical importance of the infusion into biology of thinkers who were not trained as biologists, but who nonetheless used their skills – as linguists, philosophers, computer scientists, engineers, physicists, and mathematicians – to probe important biological questions, and finally on those who followed their visions beyond “normal science” to new horizons (Harman and Dietrich 2008, 2013, 2018). In all such cases, the notion that biography should concern itself merely with the job of telling important, or instructive, or unusual lives is challenged by the imperative of contextualization. Biography can and is written to explicate science, however, not just culture, or the culture of science. One aspect of this, as in Richard Burkhart’s classic biography of Lamarck, is to rescue the totality of a scientist’s intellectual system, when only a biased fraction of it has survived to posterity (Burkhardt 1977). Another, different approach has been to look carefully at the daily routines and mental processes of scientists, at their intellectual and technical talents and vulnerabilities. As with Frederic L. Holmes’ painstaking study of the day-by-day laboratory work that led to the biochemist Hans Kreb’s description of the citric acid cycle (later named for him), the idea is to demonstrate how scientific knowledge is achieved by a particular brain in action, and to tie closely between experimental work and theory-making. Scientists’ influences may transcend the laboratory, however, rendering their social and political worldviews relevant to their science. In Darwin’s Sacred Cause, Adrian Desmond and James Moore unpack the distinct logic of The Origin of Species and The Descent of Man by considering Darwin’s motivations: the very theory of a common origin, they claim, was borne of a burning desire to fight the injustice of slavery (Holmes 1991; 1993; Desmond and Moore 2011). Still, there is a deeper sense in which science can be explicated by biography, one in which the science itself, rather than the path leading to it, is rendered more transparent to the reader. A well-known example is Sylvia Nasar’s A Beautiful Mind in which the palpable psychiatric troubles of the American mathematician John Nash are used to help the reader grasp some of the abstract rational weirdness of Game Theory (Nasar 1998). Something similar is achieved in Graham Farmelo’s biography of Paul Dirac, The Strangest Man, in which Dirac’s almost counterintuitive personality is summoned to scrutinize the head-bending realities of quantum physics (Farmelo 2009). Janet Browne’s magisterial two-volume biography of Charles Darwin beautifully succeeds to get across the gradual, trudging, and local nature of the mechanism of natural selection by minutely detailing Darwin’s own meticulous, methodical, and largely home-based scientific enterprise (Browne 1996; 2003). Indeed, science and biography often construct a two-way street: Thomas Söderqvist’s study of Niels Jerne plays with the parallels between the protagonist’s revolutionary theory of immunity, in which antibodies exist premade in the body and

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are selected by external antigens rather than created by them, and Jerne’s own selfunderstanding as a person who always had “a set of viewpoints in stock, which can be put to use on different occasions.” According to the biographer, it was the projection of the latter that opened the way to the discovery of the former, but whatever the direction of the causal arrow, both the life and the science are illuminated by the juxtaposition (Söderqvist 2003). I, too, sought to penetrate the illusive psychology of George Price, as well as the paradox of altruism, by pitting one against the other. Constructing a “double helix”-like architecture that weaved chapters on the evolutionary mystery with those describing the troubled life of Price, I hoped to show how George’s inability to find a scientific litmus test for selfless versus selfish giving illuminates the scientific question, as much as the scientific question illuminates his life (Harman 2010). But there are ways to practice biography other than as ancialla historiae, or for the purpose of the explication of knowledge, or the paths to knowledge, or for unraveling myths, or for rescuing forgotten theories, or as fodder for intertextual exegesis (Söderqvist 2011). One explicit purpose has been and continues to be to encompass the universal in the particular, since as Robert Frost said, the artist only needs a sample, and a life mercifully narrows the scope to the manageable, including in scientific affairs (Tuchman 1985, p. 134). Yet another is to advance the public understanding of science: Matt Ridley’s biography of Francis Crick in the “Eminent Lives” series is an example, introducing lay readers to the beauty of molecular biology and the mysteries of consciousness; Martin Nowak’s intellectual (auto) biography on the science of cooperation and altruism is another (Ridley 2006; Nowak and Highfield 2011). A further purpose is a kind of bait, meant to get readers interested in science by introducing remarkable, if anecdotal, scientific lives (James 2009). Biographies are also written as a form of eulogy, both by colleagues in publications like The Biographical Memoires of Fellows of the Royal Society, or by family members (Box 1978), or indeed by those who have accepted the charge as a labor of love, or for the purpose of historical redress (Sayre 1975). Two thousand years after Plutarch, biography, and especially autobiography, continues to exist as a form of aretology, too.21 In what is sometimes referred to today as “Aristotelian virtue ethics,” lives of scientists are not examined solely for their intellectual achievements per se, but rather as exemplars of “the good life,” or as achievements in and of themselves.22 Truth be told, many biographies are a combination of some, or even all, of these subgenres, which hardly ever exist as forms entirely distinct one from the other. More than a few fine biographers find themselves considering historical redress, the public understanding of science, exegesis, ancilla historiae, myth making and exploding, the construction of scientific knowledge, “the good life,” even eulogy, in their biographical writing (Todes 2014) (Wulf 2016). But there is a further

21

And genealogy. See Nye 2009. Kandel 2006 can be read as a companion to Kandel 2012. Another autobiographical example is Bishop 2003. A complex biographical example is Stoltzenberg 2004. 22

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motivation, one I take to be central to many writers. For biography is pursued as a form of belles-lettres, too, the marshalling of a legitimate unit of analysis to elicit feeling, paint in words, sculpt in structure, indeed to create a work of beauty. It is this possibility that leads to our next consideration.

Art or Craft? We turn once again, in her essay “The Art of Biography,” to Virginia Woolf. “What do we mean by calling a book a work of art?” she writes: At any rate, here is a distinction between biography and fiction – a proof that they differ in the very stuff of which they are made. One is made with the help of friends, of facts; the other is created without any restrictions save those that the artist, for reasons that seem good to him, chooses to obey. (Woolf 1939)

The facts of fiction are verified by one person alone – the artist – and their authenticity lies in “the truth of his own vision.” This world, wrote Woolf, is “rarer, intenser, and more wholly of a piece than the world of authentic information,” which, by comparison, represents “a life lived at a lower degree of tension.” The artist’s imagination at its most intense fires out what is perishable in fact; he builds with what is durable; but the biographer must accept the perishable, build with it, imbed it in the very fabric of his work. Much will perish; little will live. And thus we come to the conclusion, that he is a craftsman, not an artist; and his work is not a work of art, but something betwixt and between.

Not that, like ceramic plates on walls, nonfiction is useless. To the contrary, since humans are incapable of living entirely in the intense world of the imagination, “on that lower level of work . . . a biographer is invaluable” since “for a tired imagination the proper food is not inferior poetry or minor fiction . . . but sober fact.” This was a backhand compliment if ever one was given, a posthumous (and therefore unassailable) rub against her deceased Bloomsbury Circle colleague, Lytton Strachey. In Eminent Victorians, and his studies of queens Elizabeth and Victoria, influenced by Freud, Dostoyevsky, and probably Shaw, Strachey had invented a new form of biography wedding psychological empathy with irreverence and whit. At his most successful, his former friend now dispensed, he could do more to stimulate the imagination than any poet or novelist “save the very greatest.” But Lytton, Woolf was making perfectly clear, had been fooling himself if he thought such work could ever approach the level of her creative genius. The record must be set straight: It was, she admitted, one imagines with a self-satisfied smile, “a very cruel distinction.” But is it true? After all, whether they choose to paint exclusively in blue during the month of November, abide by the rules of the sonata form, or sculpt in wood, or in granite, artists are always bound by constraints, fiction writers included. “Technique is the scaffold upon which the highest spirit climbs,” Rachmaninoff is reported

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to have determined (in a quote I am not sure isn’t pure invention). The joys of artistic freedom notwithstanding, one can argue that the majority of constraints placed upon artists are of no making of their own. Instead, they are imposed by the hard “facts” of language, logic, dexterity, and perspective, by the arrow of time and the direction of causality, the timber of the ear and the glint of the iris, and the contours of the sulci. Kant would have considered Woolf’s distinction to be more a matter of degree than of kind, and perhaps Kant would have been right. In that case, the “curse” of mere craftsmanship may be lifted from above biography. It is a good question to debate, but one destined, because in the eye of the beholder, never to be settled. Whether art or craft, there are challenges that most biographies face, still. One relates to using the subject as source, a problem shared with journalists and historians, but uniquely palpable for biographers, especially those writing about the living (“History will be kind to me,” Churchill once promised, “for I intend to write it”) (Popkin 2005; Smocovitis 1999; Comfort 2011). Another is whether to write thematically or chronologically, sometimes referred to as “Seutonian” versus “Plutarchian” approaches (Leo 1901). A third is how to write about people who have left little behind, or about those for whom many accounts already exist (Greenblatt 2005; Powers 2007). Yet another is how to deal with the complication, as Woolf’s own biographer explained, that “A self that goes on changing is a self that goes on living: so too with the biography of that self” (Lee 1999, 11). With biographies of scientists, the trade-off between deep explication of the science and readability is always an issue.23 And, of course, there remains the business of selecting, among the myriad details of a life, those moments and words and relationships and thoughts that count, and are telling. As Barbara Tuchman reminds us, however democratic we wish to be (if indeed we do), “a portraitist does not achieve a likeness by giving sleeve-buttons and shoelaces equal value to mouth and eyes” (Tuchman 1985, 145). Biographers have found different solutions to such challenges.24 The art or craft of making a life “speak,” of putting a reader in the subject’s shoes while maintaining for him or her the requisite critical distance, is an art, or craft, not easily accomplished. Whether or not biography “lends to death a new terror,” as Oscar Wilde feared, the conscientious biographer shoulders the weight of responsibility. It is a daunting task. One can never step into the same life twice, but herein lays the beauty.

The Future of Biography Sometimes it’s impossible to describe the achievements of an individual without understanding their network: lives in science may stand alone, but often they depend on collaboration. That is what Silvan “Sam” Schweber found in his QED and the As one editor once said to me, “every further equation cuts the readership in half.” The use of portraits, or alternatively original art, is just two of the many tools biographers have marshaled. See Fara 2007 and Redniss 2010.

23 24

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Men Who Made It, weaving Tomanaga, Feynman, Dyson, and Schwinger into one intellectual fabric (Schweber 1994); Michael Lewis, too, rightly, could not disentangle Amos Twersky from Daniel Kahneman (Lewis 2017). As “Big Science” continues to grow, with authorship of many papers slipping into the hundreds, it will be an increasing trend; the collaborative “labography” has already been christened as a new biographical genre (Otis 2007; Nye 2009). Gone are the days, in theory at least, of the Renaissance genius or Victorian gentleman-scientist, unlocking the mysteries of a coy Universe.25 This may have the effect of softening the sting of hagiography, dulling the distortion visited upon science history by the overemphasis on individuals. Since readers develop their own affections, however, and collectives can be just as inspiring as singletons, I am not certain to what degree. In the end, like the gene or the atom or the twenty-four hour day, we seem to relate best to the fundamental entities, even as we are doing our darnedest to depart from them. Gopnik was right, we do believe in lives, and I don’t see that changing. But collective biographies can be justified not only on intellectual-scientific grounds. Even when the science itself is more or less beside the point, they may thrive as forms of ancilla historiae. The Visible College describes five wildly differing British scientists: Needham the biochemist, Bernal the crystallographer, Haldane the geneticist, Levy the mathematician, and Hogben the experimental zoologist. But casting them together allowed Gary Werskey to explain how Science and the Left went hand in hand in Britain between the wars, and what this meant both for the left and for science (Werskey 1978).26 The same holds true for lives corralled in the service of explicating whatever form of “otherness.” Collective biographies are one departure from the classic form known to moderns, and hardly the most radical. Today there are “biographies” of objects (Daston 2000), organisms (Moyal 2004; Nicholls 2006), diseases (Mukherjee 2010),27 molecules (Ball 2000), genomes (Ridley 1999), even equations (Bodanis 2001). Purists may cringe at the idiomatic misuse, but any one who has taught a history of science graduate course in scientific biography will know just how useful the expanded term can be for organizing a student’s mind: “Think of mitochondria like you would a person,” “Narrate the birth and apotheosis and denouement of chicken pox,” “Construct a biographical arc of the science of the evolution of consciousness” – the possibilities are endless, for students and scholars alike. “Know the grave doth gape for thee thrice wider than for other men,” the King warns Falstaff. Personification, the bard knew, is a tonic of the imagination.28 As biography grows beyond the person and the group to nonhuman beings, and things and ideas; as it continues to evolve from hagiography to existential biography

25

A curious exception is Lovelock 2000. Two nonscientific collective biographies that stand out as effective vehicles for explicating the birth of American pragmatism, on the one hand, and European utopianism, on the other hand, are Menand 2001 and Manuel 1962. 27 Also, Oxford University Press’s series, Biographies of Disease, on Asthma, Diabetes, etc. 28 William Shakespeare, Henry IV, Part 2, Act 5 Scene 5. 26

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to metabiography, and from historical reconstruction to subtle historiography, it will be attended by new challenges. Some of these will be due to our brave new world, in which digital clouds and tweets are replacing letters and notebooks. Oral history projects are on the rise, I’d argue, because it remains unclear to somewhat angst-full historians precisely how they are going to collect their data in this day and age.29 A related challenge will have to do with the charms and pitfalls of “Big Data”: sifting through the reams of materials to paint the eyes at the expense of the shoelaces will be more of an issue in the future than it has been in the past, and whether they wish for it or not, the rising field of Digital Humanities will increasingly become central to biographers. Yet another challenge will be our changing notions of identity: when human-cyborgs start walking among us, or even just when enhancement begins to blur the distinctions we have come to view as natural, how will we write lives? What will become of agency? Of feeling? Nobody knows. Still, many of the age-old challenges faced by Iamblichus and Augustine and Stoever, whatever they thought they were doing, will probably continue to pertain; to a meaningful degree, a life is a life is a life, the talons of time’s changing perspective notwithstanding. The same ambivalence that visited Boswell when he considered which adjective to use in describing Johnson’s rage, or happiness, or Doris Kearns Goodwin when she deliberated over an embarrassing personal detail in the life of LBJ, or Lincoln, will keep the future biographer awake at night, still. For when all is said and done, biography is not a two-way but a three-way affair between subject, author, and reader. It is meaningless without an audience. And in seeking to distill what we care most about, a kind of Promethean theft of self-knowledge in miniature, it captures us, or at least has the potential to. Like ice cream, it is a guilty pleasure, and when it’s good, you don’t want it to end. Why do we love biographies so much? There are many answers. Having come to the end of this disquisition, we leave the last word to the playwright: “My advice to you is not to inquire why or wither, but just enjoy your ice cream while it’s on your plate.”30

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29

Examples include projects launched by the Chemical Heritage Foundation, the Royal Society, the American Institute of Physics, the Max Planck Institut für Wissenschaftsgeschichte, the British Library, and many more. 30 Attributed to Thornton Wilder.

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Women in the Historiography of Biology

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Marsha L. Richmond

Contents Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Historical Background . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . The Women’s Movement and the History of Biology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scholarship on Women . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Efforts to Recover the Work of Women Scientists . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Biographical Studies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Women in Different Areas of Biology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Women Nature Writers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Women and Biology Education and Teaching . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Feminist Studies of Biology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Areas for Future Work in the History of Women in Biology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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Abstract

Women no less than men have been drawn to the study of nature, but only after they were given access to higher education were they able to participate in science in greater numbers. The historian’s gaze has accordingly been transfixed on the accomplishments of men and a few extraordinarily talented women. This began to change in the 1970s when a new wave of women scholars and feminists turned to recovering the accomplishments of individuals, exploring opportunities and hindrances to women’s participation in science, and identifying sexist ideologies and gendered assumptions within biology that influence not only women’s ability to do science, but also broader social, economic, and political rights of women. The result is a rich body of literature on women’s contributions to, and a feminist critique of, biology. M. L. Richmond (*) Wayne State University, Detroit, USA e-mail: [email protected] # Springer Nature Switzerland AG 2021 M. R. Dietrich et al. (eds.), Handbook of the Historiography of Biology, Historiographies of Science, https://doi.org/10.1007/978-3-319-90119-0_17

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The amazing thing is that there are so many of them. In almost every journal in every month since the mid-nineteenth century, the names of women biologists abound. In almost every area within the discipline women have pursued and publish their research. In every class of every year (since its inception in 1888), women have represented no less than 40–60% of the total number of students at the Marine Biological Laboratory (MBL) at Woods Hole. And yet, when we think of the history of biology, when we tried to recount the concepts and techniques that have moved the science forward, we end up finally by shrugging our shoulders and asking, “But where were the women? What did they do?” Is it possible that for all their work and all their efforts most of these women, whose careers often spent many decades, and in fact contributed so little to their science that for all intents and purposes they might never have existed? Or, is it more probable, as I would like to argue, that the work of the scientists has succumbed to the same fate as has the work of women artists and composers; that is, although their work has frequently been woven into the fabric of the discipline, it has only rarely been associated with the names of the women responsible for it. Gabriele Kass-Simon (1990, 215)

Introduction Since Gabriele Kass-Simon first asked “But where were the women? What did they do?,” there has been an outpouring of literature that attempts to answer these critical questions. In the past 25 years, a considerable body of scholarship has appeared that examines women’s participation in science, and especially biology, from a number of perspectives. Once women were given access to higher education in the latter half of the nineteenth century, many more individuals were able to contribute to biological investigation. Current historiography of women in biology includes numerous biographical studies of women, and we are also beginning to recognize how the discipline has developed as a result of women’s contributions, both individually and collectively. The picture is not all glowing, however. Science, including biology, has historically been a male-dominated human activity. Women’s participation in scientific work has been hampered by dominant patriarchal social and cultural systems that systematically excluded, marginalized, or overlooked their efforts. The historiography of biology thus reflects these operational hindrances: not only have fewer historical studies been devoted to women than to men in biology, proportionally speaking, but there has traditionally been an implicit androcentric bias within the history of biology that tends to disregard women’s contributions. There is, in short, a propensity for historians to privilege men’s contributions and disregard those of women. Such a scenario can be better understood by surveying the disciplinary development of the history of biology as a subset of the broader discipline of the history of science. Within the history of science, there exists a long-standing tradition of documenting the accomplishments of the “great men” (and, to be sure, occasionally the “great women”) or the “knowledge-makers” in science. Such a predisposition naturally tends to neglect those who engaged in normal science, whether women or

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men, whose scientific activities did not attract widespread recognition. Thus, the scientific work of women, people of color, and other junior figures has not figured prominently in the historical record. In the case of women, only with the rise of second-wave feminism in the 1970s did such exclusionary practices begin to be called into question (Evans 2003). The rise of feminist scholarship, coupled with greater numbers of women entering the history of science, encouraged the publication of a vigorous body of literature on women in science. Within this corpus, works have documented women’s contributions to biology and critically examined the role of gender in the life sciences. This expansive scholarship demonstrates that not only have women been long engaged in biological studies but that broadening the focus of investigations to include women as well as men in historical studies provides both a more accurate and dynamic picture of the development of the life sciences. This essay presents an overview of general trends in historical writing about women in biology from a number of vantage points. First, it surveys the historical development of the history of science in the early twentieth century and the rise of the history of biology in the 1960s. Next, it describes the different periods and genres of writing about women in biology, highlighting contrasting analytical and theoretical perspectives as well as themes that have been explored. It concludes with practical suggestions about how to conduct research on women in biology as well as future directions for scholarship.

Historical Background Literature on women is a subset of the general history of biology, which is itself one of the fields encompassed by the history of science. To provide a framework for the historiography of women in biology, I will first present an overview of the discipline’s history and then chart the rise of interest in women’s studies. Professional interest in the historical development of science emerged as an academic field in the early twentieth century, largely through the efforts of the Belgian chemist George Sarton (1884–1956). Sarton founded the journal Isis in 1912, and when he moved to the USA in 1915, he brought the journal with him. Sarton became a lecturer at Harvard University in 1920 and professor in 1940, retiring in 1951. In 1924, he and other like-minded scientists founded the History of Science Society (HSS) to serve as a subscription society to support the journal. The predominant focus of much of the early scholarship in the history of science was on the physical sciences and especially the “great men” who contributed to the Scientific Revolution of the sixteenth and seventeenth centuries. Given that most early practitioners had been trained in the sciences (and that a scientific background was generally regarded as a qualification for doing history of science), it is not surprising that males dominated the field until after the Second World War. Among the 62 participants at the 1957 symposium “Critical Problems in the History of Science,” intended to chart new directions in the fledgling discipline, three were women, two of whom (Dorothy Stimson and Marie Boas) contributed to the

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subsequent published volume (Clagett 1959; Stimson 1959) (Interestingly, Stimson, writing about the value of the history of science in liberal arts programs, specifically focused on women’s colleges.). As Arnold Thackray commented retrospectively, “The roughly two dozen individuals who acted as speakers and commentators at the Madison meeting came close to constituting the totality of professional historians of science in the English-speaking world of their day” (Thackray 1995, vii). Gradually, new academic programs in the history of science were established in the 1950s and 1960s, training in historical methodology rose in importance, and more women were attracted to the new discipline.1 It is noteworthy that Stimson (1890–1988), a Copernicus scholar and professor of history at Goucher College, was elected president of HSS in 1953. It was another 35 years, however, before the second woman, Mary Jo Nye, became president in 1988. Of course, women historians did not necessarily focus on women in science. Indeed, given the struggle for acceptance in the discipline that women faced at the time, doing so would have impeded their career advancement. Nonetheless, the entry of more women into the profession in the 1960s was a precondition for an awakening of interest in women’s science studies in the 1970s. By the late 1960s, the history of the life sciences joined the physical sciences as a mainstream field of study. Its emergence as a subdiscipline dates from the founding of the Journal of the History of Biology in 1968 under the editorship of the Harvard historian of biology Everett Mendelsohn. In the Editorial Forward to the first issue, Mendelsohn pointed to current disciplinary trends: “While the physical sciences have long served as the paradigm for work in the history of science, and several specialized journals have published articles in this field, this imbalance is now being redressed. Many historians of science are now turning their attention to the complex and often challenging problems of the history of biology, and a new generation of scholars has taken biology as the focus for their historical analyses” (Mendelsohn 1968). Given Harvard’s continuing prominence in the discipline, most of the authors of the first articles were either Harvard graduates or faculty. As a sign of the times, a woman, Judith Pound Swazey, was among this group (Swazey 1968), and Shirley A. Roe, a Harvard Ph.D., served as Book Review Editor (becoming Associate Editor in 1981 and Co-editor in 1989). Mendelsohn drew attention to new directions in the history of science. As he noted, the discipline was beginning to expand beyond its early focus on tracing the history of scientific ideas: Contemporary scholarship in the history of science makes changed demands upon the author; these are demands for methodological awareness and realization that other fields of historical study have brought new sophistication to the writing of history. While hard data will always serve as the basis of history, the simple narrative is no longer acceptable,

1

Harvard University remained the leading program for training historians of science, but new departments were created at the University of Wisconsin (1940s), the University of Oklahoma (1950s), and Indiana University (1960), and a program in the history of science formed at Cornell University in the mid-1960s.

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particularly when dealing with the emergence of ideas. The best history will be characterized by penetrating and critical analysis of changing concepts and altered methods of experiment and observation. Biology, in particular, must be studied in terms of its relationships with the other sciences and with the intellectual currents of its day. It may be examined as well for its interaction with the institutions of the society which spawns it.

In this way, Mendelsohn specifically tied the history of biology to studies that emphasized social (“external”) as well as intellectual (“internal”) influences.

The Women’s Movement and the History of Biology Women’s studies likewise emerged along with other social movements of the late 1960s. Yet a focus on the history of women in science developed later; it was not until 1976 that JHB published the first article explicitly dealing with the “woman question” in biology. Maryanne Cline Horowitz’s essay entitled “Aristotle and Woman” well exemplifies the change occurring in scholarship as a consequence of the rise of the women’s movement (Horowitz 1976). Horowitz, a Renaissance historian who was soon appointed a Research Associate in Women’s Studies in Religion at the Harvard Divinity School, began by noting that Aristotle’s view of “female human nature” required critical analysis, not simply because this topic had been overlooked by leading Aristotelian scholars, but more importantly because his writings had inspired “many of the standard Western arguments for the inferiority of womankind and for the political subordination of women to men in home and in society” (Horowitz 1976, 183). Her text thus illustrates a central position of feminist historians of the time. It also stimulated a group of young biologists to begin thinking about androcentric bias in biology (Biology and Gender Study Group 1976). Horowitz’s article provoked a response by the political philosopher Johannes Morsink. Considering the question “Is Aristotle’s Biology Sexist?,” Morsink distinguished between biological sexism and political sexism, which allowed him to conclude that Aristotle’s biological view on women, and his form-matter hypothesis (that in reproduction males provide the form while females provide the matter), could have been based on sound scientific principles rather than simply reflecting sexism. If so, he argued, “Aristotle’s biology, though not now sexist in conception and origin, was still sexist because of the consequences it had.” In this way Morsink concurred with Horowitz’s central thesis, while he nonetheless defended Aristotle’s science. “When seen in its historical and scientific context,” he concluded, “the theory testifies to Aristotle’s scientific acumen, and it would seem that in constructing it Aristotle stayed within the boundaries of accepted scientific practice” (Morsink 1979, 84, 112). These two articles well reflect the impact that the rise of the women’s movement in the late 1960s and 1970s had on scholarship. Not only do they illustrate a new focus on “the woman question” in science, but they also indicate contemporary concern about the possible sexist views of scientists and in science itself. From the

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mid-1970s, increasing attention was given to bringing women into the history of science and new impetus and highlighting the work of women within the history of biology as well as feminist critiques of science. With an expanding cohort of women entering the history of science in the late 1960s and 1970s, the discipline itself began to change. Within HSS, for example, an ad hoc Committee on Women was created in 1972 to examine the status of women in the discipline.2 The following year the committee recommended the establishment of a Standing Committee on Women whose purpose was “to evaluate the ongoing status of women, to maintain a roster, and to hear complaints,” which particularly involved inequities in hiring practices. In 1982, the Committee offered a detailed report that assessed progress that had been made. This committee determined that “the number of women in the HSS jumped 43% between 1971 and 1980, although (because the Society was also growing rapidly) the women’s percentage of total membership rose only from 14.7% to 17.1%. However, . . . women’s participation at HSS meetings between 1974 and 1980 varied from year-to-year and location to location, but averaged 20.8%, largely because of ‘Works in Progress’ sessions.” One area where the Committee pressed for change was an increase in the number of women publishing in and serving on editorial boards of history of science journals. In 1972, the committee found that Isis, the journal of the Society, was not involving women in editorial activities proportionate to their Society membership. “Women had written 11.5% of the articles in Isis in 1968–72, were asked to write 7.6% of the book reviews, and held only one position on the editorial board.” A decade later progress could be reported: whereas in 1979, when Arnold Thackray assumed the editorship, there were no women serving as advisory editors, by 1981 there were six (26%) (“Women in the History of Science, 1973 to 1981” 1982). The push for equal employment opportunities for women initiated in the early 1970s also began to bear fruit. While the Committee on Women promoted several initiatives, it recognized its limitations: “So much has happened in the elaboration of the jurisdiction of the Equal Employment Opportunities Commission and the Department of Labor’s Office of the Federal Contract Compliance Program since 1973 that employment and advancement issues are now more clearly in the realm of government than a professional society.” The HSS Women’s Committee lobbied for the open posting of positions and began to keep track of job openings and those seeking positions, and it prepared annual surveys of the number of women hired. With the coming of affirmative action and an expanding job market, a greater number of women were hired by universities or other institutions. As reported in 1983, the job surveys between 1974 and 1981 showed an increased number of openings in the history of science. “Women have applied for most of these jobs but only in the last year or so have their proportion of offers paralleled their numbers of applications” (“Women in the History of Science, 1973 to 1981” 1982, 5).

2

(In establishing a women’s committee, HSS was following the lead of the American Historical Association, which established a Coordinating Committee on Women in the History Profession in December 1969 (Smith 1994)).

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The rise in the number of women faculty members served as role models to incoming women graduate students. This development was significant. In 1963, for example, Marie Boas Hall (1919–2009) and Alfred Rupert Hall (1920–2009) were hired at Imperial College of Science and Technology in London, and over the years they cultivated a generation of graduate students in the history of science, many of whom were women who eventually assumed leadership roles in the profession (James 2012). In the USA, the establishment of the HSS Women’s Caucus served to provide effective role models for junior scholars. Again, although the number of women in the profession does not directly map onto interest in the study of women in science, the two are correlated.

Scholarship on Women The initiatives encouraged by the women’s movement were important in effecting change in the discipline. Not only did women seek to gain increasing recognition within the profession, they also focused on expanding scholarship on women in science. Before 1970, there was almost no literature on women in science, with the notable exception of Rev. John A. Zahm’s classic book Woman in science (1913), written under the pseudonym of H. J. Mozans (1991). Indeed, the only woman scientist generally identified was Nobel Laureate Marie Curie (Ogilvie 1996). Within a decade, however, the situation had changed significantly. Within the field of general history, by the late 1970s, the field of women’s history was “the most rapidly growing new specialization in the profession” (Smith 1994, 16). The history of science soon followed suit. In fact, Signs: A Journal of Women and Culture devoted its fall issue in 1978 (vol. 4, no. 1) to the topic of women and science. By the 1980s, the history of women in science was acceptable as a dissertation project. In 1984, Londa Schiebinger submitted her thesis on “Women and the Origins of Modern Science” to the Harvard History Department. In the introduction Schiebinger mentioned a central topic in the women’s movement in recent years: the exclusion of women from various aspects of social life based on the assumption of their “natural” incapacity. “A question then immediately arises: what have been the social or cultural origins of the exclusion of women from public spheres of life? If we reject nativist explanations for the division of labor in society, we are forced to look to history and the confluence of forces – intellectual, social, and economic – which have brought about the exclusion of women from science.” Identifying a gap in the history of the sciences, Schiebinger noted that “those few exceptional women who did contribute to science have not been thoroughly investigated; nor have the mechanisms which have blocked women’s contributions and fully documented or analyzed” (Schiebinger 1984, 6–7). Her dissertation examined both the reasons for the exclusion of women from science in the sixteenth and seventeenth centuries and a number of individuals who contributed to science. Marilyn Ogilvie soon documented the new focus on women’s studies by tabulating the number of sessions devoted to women’s history at HSS annual meetings

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between 1972 and 1994. Whereas in 1972 “there was neither a session nor an individual paper devoted to gender issues or women in science,” the next year there were two. Throughout the 1980s, meetings regularly included at least two sessions devoted solely to women in science, and there were papers on women as part of other sessions. By 1993, the number of presentations swelled; “there were numerous sessions with papers concerned with women in science, feminism, and gender in science, with five sessions devoted exclusively to the topic.” This greater interest in women’s studies led Ogilvie to conclude that “interest in feminism, science, and gender issues, and specifically women and science has increased exponentially” (Ogilvie 1996, ix–x). Contemporary scholarship supported her claim.

Efforts to Recover the Work of Women Scientists Historians approached women’s participation in the sciences from several different vantage points (see especially Kohlstedt 1995). Given the paucity of information about women scientists, it was natural that some of the earliest works were those of “recovery,” works that sought to identify women in science from all eras, but with a particular focus on the nineteenth and twentieth centuries. This scholarship included general surveys, biographical dictionaries, and individual biographies. A central theme of this literature was the different “barriers” women faced in trying to do science. The groundbreaking work of this period was the publication in 1982 of Margaret Rossiter’s Women Scientists in America: Struggles and Strategies to 1940, which eventually was expanded by three additional volumes. In justifying the work’s focus on late nineteenth and early twentieth centuries, Rossiter cited the opening of higher education to women as crucial in allowing them to participate in science. Identifying three general periods – before 1880, 1880–1910, and after 1910 – she noted that: “Of these the second is both the most interesting and the most important, for it was then that acceptable conditions for women’s presence in science were worked out” (Rossiter 1982, xvi). The opening chapter, appropriately enough, discussed women’s colleges, given that most women scientists in America before 1900, because they were barred from Ivy League faculties, held positions in women’s colleges and a few coeducational universities. In subsequent chapters, Rossiter highlighted “women’s work” in science; the “manly profession” of science; academic, government, and industrial employment; double standards and under-recognition; and the prizes and honors women created to compensate for this. Although thematically organized, Rossiter covers the work of women in the life sciences as well as other fields. The statistical analysis she prepared of women who received doctorates from American universities indicates, for example, that counter to nineteenth-century trends, that zoology rather than botany “was the most widely taught and therefore most accessible of all the sciences to women” in the first half of the twentieth century (Rossiter 1982, 151). Rossiter’s book not only advanced women’s scholarship but inspired others to begin working on the history of women in science. This rich and well-documented work – founded on truly

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herculean archival research – was unparalleled for the time and indeed, along with three subsequent volumes, remains a prime reference work to this day (Rossiter 1995, 2012). In addition to this broad survey, another significant area within works of recovery was biography. Most notable in this genre was the publication of a series of biographical dictionaries aiming to provide basic biographical information about women in various fields of science, across different time frames and from different countries. Marilyn Ogilvie’s Women in Science: Antiquity through the Nineteenth Century Ogilvie (1986) was foundational. She eventually published two more volumes, including one co-edited by Joy Harvey (Ogilvie 1996; Ogilvie and Harvey 2000). Another scholar who has prepared useful biographical source books is Mary R. S. Creese. A research chemist, in 1998, she published Ladies in the Laboratory? American and British Women in Science, 1800–1900: A Survey of their Contributions to Research (Creese 1998). This work is organized differently than a dictionary. “The field-by-field examination brings out patterns and concentrations in women’s research (in both countries) and allows a systematic comparison of the two national groups. Through this comparison, new insights are provided into how the national patterns developed and what they meant, in terms of both the process of women’s entry into research and the contributions they made there” (Creese 1998, back cover). For those working in the life sciences, this book is of special interest: Part I is solely devoted to the life sciences, covering botanists “from early explorers to plant geneticists,” lepidopterists, zoologists (from museum taxonomists to morphologists and embryologists), naturalists and general biologists, as well as women who worked in the medical sciences. In 2004 Creese extended her coverage to Western Europe, listing the work of women in Scandinavia, Ireland, France, Belgium, the Netherlands, Germany, Austria-Hungary, Switzerland, and Italy (Creese 2004). This volume likewise highlights women in the life sciences, although it is not as comprehensive as the first volume. Two additional volumes have also appeared, one on women scientists from South Africa, Australia, New Zealand, and Canada and another on women working in Imperial Russia. These works fill a significant void by highlighting the participation of women around the world in scientific work, especially in the life sciences (Creese 2010, 2015). These and similar works have proved to be excellent source books for those working on the history of women in biology (see also Herzenberg 1986). In addition to general and national biographical dictionaries, scholars have also examined women who were associated with particular disciplines and institutions. For example, Helga Satzinger and Ute Deichmann have explored women in German genetics (Satzinger 2004, 2009; Deichmann 1997). Annette Vogt has published on German women who worked in universities as well as a very useful biographical dictionary of women associated with the Kaiser Wilhelm Institutes in Germany (Vogt 1996, 2004, 2008). These studies highlight the work of women who may not have been leaders, but who nonetheless contributed to the normal work of important research institutions. A similar effort by scientists themselves focuses on women’s contributions to Japanese science (Kozai et al. 2001).

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Biographical Studies In addition to biographical dictionaries, scholars have enriched the historiography of women in science by studying the lives of individuals, adding both depth and nuance to our understanding of the experiences of women who devoted themselves to scientific study. Since the 1970s, a large body of biographical literature on women in science has been amassed, focused on those who were exceptional and widely recognized as well as on those who simply contributed to normal science. Indeed, it was historians of women in science who were largely responsible for reaffirming the value of biography in the history of science. As Paolo Govoni noted, the status of biography “was at a low ebb in the 1960s,” but beginning in the 1970s, “specialist books and articles of scholars in various fields began to examine and evaluate biography as a genre, and the relations between the self doing the research and the self researched began to be explored” (Govoni 2014, 10). The genre of biography, as fueled by feminist concerns of rewriting “master narratives,” has not only enriched women’s studies but also science studies in general.3 To date, there have been a profusion of books, edited volumes, and articles on the lives and work of women in biology. Among book-length biographical studies, most explore the lives of exceptional women who gained wide recognition for their scientific work. While it is impossible to list all of these works, some notable books for the recent history of biology include biographies of the Nobel Prizewinning geneticist Barbara McClintock by Evelyn Fox Keller and Nathaniel Comfort and of the marine biologist and ecologist Rachel Carson (Keller 1983; Comfort 2001; Lear 1997; Lytle 2007; Musnil 2014). Other notable biographies are those of the American zoologist Alice Middleton Boring (Ogilvie and Choquette 1999), the German geneticist Paula Hertwig (Gerstengarbe 2012), and the French Darwinist Clémence Royer (Harvey 1997). Given their study of biologically interesting chemical structures or medically interesting molecules, biochemists’ biographies are notable, including Georgina Ferry’s biography of the biochemist Dorothy Hodgkin, two biographies of the X-ray crystallographer Rosalind Franklin, and the recent life history of the biochemist and microbiologist Marjorie Stephenson which are noteworthy (Ferry 1998; Maddox 2002; Sayre 1975; Ŝtrbáňová 2016). Although biographies of women prior to the nineteenth century are rare, the study of the extraordinary entomologist Maria Sibylla Merian (1647–1717) is exceptional (Todd 2007). Given the difficulty of finding archival materials on women extensive enough to support full-length studies, most of the biographical literature on women scientists has appeared in the form of journal articles or chapters in collected volumes. A number of works stand out by addressing particular themes important to women in the life sciences. A notable early work is the 1987 Uneasy Careers and Intimate Lives: Women in Science, 1789–1979, edited by Pnina Abir-Am and Dorinda Outram (Abir-Am and Outram 1987). The volume includes articles on women in botany, ornithology,

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(On feminist methodology for narrative research, see Bloom (1998)).

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medicine, as well as mathematical biology, in addition to those in chemistry and astronomy (Shteir 1987; Ainley 1987; Slack 1987; Harvey 1987; Abir-Am 1987). Moreover, the pieces do not simply provide biographical details but explore women’s strategies to combine their scientific proclivities with marriage and family. A companion volume, Creative Couples in Science, appeared in 1996, edited by Helena Pycior, Nancy Slack, and Pnina Abir-Am, and describes the lives of women whose scientific work was carried out within the context of marriage (Pycior et al. 1996). Like the earlier volume, it includes articles on several women who worked in botany, zoology, and medicine (Ainley 1996; Cohn 1996; Garber 1996; Henson 1996; McGrath 1996; Tucker and Groeben 1996; Slack 1996). Similarly, a collection of Isis articles on women in science edited by Sally Gregory Kohlstedt includes six articles on women in biology, some biographical and others analytical (Kohlstedt 1999). More recently, another collected work on scientific couples has appeared, For Better or For Worse: Collaborative Couples in the Sciences, edited by Annette Lykknes, Donald Opitz, and Brigitte Van Tiggelen, which explores how women (and a few men) collaborated with their spouses or partners (Lykknes et al. 2012). Again, such collective volumes offer both individual stories and a comparative perspective, which helps establish general tendencies about women’s experiences in science.

Women in Different Areas of Biology One way of grouping the large number of individual biographical studies of women is to focus on those contributing to particular biological fields. This approach is more than simply expedient. Women were given access to higher education in the sciences in the latter decades of the nineteenth century, and by 1900 those graduating with university degrees in the life sciences were seeking employment in science. This coincided with the emergence of a number of new subdisciplines in biology, including genetics and cytogenetics, ecology, and biochemistry, all of which needed workers to carry out investigations and to staff the new research laboratories that were being created. Because these areas were not yet established, and hence not attractive to men seeking career security, they offered women unprecedented opportunities to work in science. Thus women were proportionally more visible in these fields. Women were prominent in the new field of genetics, for example, and a number of studies have documented their work through biography and prosopography.4 Less attention has been focused on biochemistry and ecology, both of which present rich

4

(For Britain, see Richmond (Richmond 2001, 2006, 2007b) and Love 1979. For the Netherlands, Norway, and Germany, see Stamhuis (1995), Stamhuis and Monsen (2007), and Stamhuis and Richmond (2014). For the USA, see Ogilvie (2007), Richmond (2012), and Dietrich and Tambasco (2007). On cytogenetics, see Brush (1978), Ogilvie and Choquette (1981), Cross and Steward (1993), and Echeverría (2000) (all of which focus on the work of Nettie Marie Stevens); see also Richmond (2010), and Williams (2016). For Germany, see Satzinger (Satzinger 2004, 2008, 2009a, b) and Deichmann (Deichmann 1997, 2014))

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opportunities for future scholarship.5 Women were also present in more established fields as well, including marine biology, protozoology, developmental biology, molecular biology, primatology, and anthropology.6 Although not exhaustive, this literature demonstrates the energetic nature of studies of women in the life sciences.

Women Nature Writers A particularly vibrant subset of studies of women in biology is the focus on “nature writers.” This genre has primarily been advanced by literary scholars who explore the natural history writings of women who were not formally schooled in the sciences – women “on the edge of science” (Gates 1998). Most authors were autodidacts who learned about nature through experience as well as through reading books. Women’s botanical writing particularly flourished in the eighteenth and nineteenth centuries and aimed to enlighten women, children, and general readers about various aspects of plant life; it also encompassed the herbal tradition as well as botanical art and design. Some women engaged in collecting and describing new plant species, not simply to advance knowledge but also as a source of income, often supported and encouraged by male patrons and collectors. A series of exceptional books exploring women nature writers have been published since the 1980s. These include works by Ann (Rusty) Shteir, Barbara T. Gates, and Tina Gianquitto (Shteir 1996; Gates and Shteir 1998; Gates 1998; Gianquitto 2007a). A number of articles have also appeared in collected volumes (Shteir 1987, 1997a, b; Benjamin 1999; Shuttleworth et al. 2001). Linked to this tradition are those women who, despite being autodidacts, considered themselves scientists rather than nature writers. Several of the women who either assisted Darwin in his botanical studies or whose own studies were inspired by his evolutionary theory have been studied (Harvey 2009; Gianquitto 2007b, 2013; Gianquitto and Fisher 2014; Love 1983). Relatedly, Kimberly Hamlin has explored Darwin’s impact on the early women’s movement in the USA (Hamlin 2014).

Women and Biology Education and Teaching At a time when women were marginalized from participating in mainstream professional biology, they were nonetheless able to pursue their interest in the life sciences through teaching. Teaching was indeed one of the earliest career options for women interested in science. Many taught natural history as part of the primary school

5 (For biochemistry, see Richmond (2007a), Needham (1982), and Ŝtrbáňová (2004). For ecology, see Norwood (1993), Marcil (2015), and Slack (1996)). 6 (Marine biology, Sloan (1978) and Zottoli and Seyfarth (2015); protozoology, Warner and Ewing (2002); developmental biology, Keller (1996) and Gilbert and Rader (2001); molecular biology, Spanier (1995); primatology, Fedigan (1994, 2001); anthropology, Wylie (1997, 2001)).

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curriculum, while others found places in the secondary schools (and particularly the new high schools for girls that opened in the nineteenth century), which offered instruction in natural history disciplines. A few were fortunate to gain appointments at one of the women’s colleges and or even at a coeducational university. The impact of women who taught biology was enormous although often unacknowledged. Sewell Wright, for example, decided to major in biology after taking a course at Lombard College from Wilhelmine Entemann Key (1872–1955), one of the first women to receive a Ph.D. from the University of Chicago, studying under C. B. Davenport and C. O. Whitman. Two notable contributions specifically explore women’s education and teaching in the life sciences: Ellen Burstyn’s study of the Anderson School of Natural History (Burstyn 1977) and Sally Gregory Kohlstedt’s book-length examination of the American nature study tradition (Kohlstedt 2012). Also of note are two studies of biology teaching at Mount Holyoke (especially Cornelia Maria Clapp and her successor Ann Haven Morgan), one of the leading women’s colleges in the USA for those interested in biology (Levin 2005; Warner and Ewing 2002). The biography of Dr. Alice Middleton Boring by Marilyn Ogilvie and Clifford Choquette highlights the interesting career of someone who chose to teach biology in China rather than having to deal with some of the prejudices against women in the USA (Ogilvie and Choquette 1999) and who inspired a number of women students to embark on careers in medicine or science.

Feminist Studies of Biology As we have seen, significant progress has been made over the past few decades in trying to answer the question, Where are the women? Biographical studies of individual women have provided a better sense of who they were and what they did. Another aspect of this inquiry, however, involves a questioning of social, economic, and political systems that disadvantaged women who wished to pursue the study of nature. While historical studies can reveal the nature of the frameworks in which women’s work in science was embedded, philosophical and theoretical analyses provide critical insights into how these systems were constructed. The feminist critique of science emerged in tandem with historical and literary studies of women in biology, likewise building on foundational scholarship that emerged in the 1970s. As scholars began to explain why women were absent from historical accounts, a more radical reading within women’s studies began to emerge. As Jacqueline Zita noted, “In the early 1970s, articles, conference papers, and discussions began to appear, sparked by the discovery of women’s absence in the sciences, both as practitioners and as accurately understood subjects of study. Gradually, ‘the woman question’ in science (or why aren’t there more women scientists?) was transformed into the [sic] ‘the science question’ in feminism (or what is wrong with science?). In this transition, the initial reformist criticism of science’s exclusionary practice as regards women was altered into a new radical interrogation of science and its method of inquiry” (Zita 1988, 157–58). The early vibrancy of feminist interrogations of science was

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demonstrated in the late 1980s when the new journal Hypatia devoted two separate issues to “Feminism and Science.”7 A key early theoretical contribution to feminist studies was the distinction drawn between sex and gender. Introduced in the 1970s, the concept of gender came to denote the social construction of sex distinctions that reflects systems of social or sexual relations rather than biological categorizations of physical difference. In her influential 1986 article, “Gender: A Useful Category of Historical Analysis,” Joan Scott explained the importance of this distinction: “The core of the definition rests on an integral connection between two propositions: gender is a constitutive element of social relationships based on perceived differences between the sexes, and a primary way of signifying relationships of power” (Scott 1986, 1067, 1069). While Scott’s particular view did not go unchallenged, gender analysis nonetheless has been widely influential, especially in the history of women in biology (see especially Rose 1993; Keller 1995). As Evelyn Fox Keller and Helen Longino noted in the introduction to the 1996 volume Feminism and Science, the analytic value of gender extended “far beyond the minds and bodies of individual men and women. Using gender as a lens, or as an analytic tool for examining the implications of coding some spheres as masculine and others feminine, scholarship in the social sciences and humanities soon produced dramatic and persuasive reinterpretations of many familiar and well-studied matters (including institutions, practices, cultural artefacts, historic periodization, literary standards, etc.” (Keller and Longino 1996, 2). Gender analysis remains a dynamic component of science studies (Opitz 2015). Energized by the recognition of, and attempt to remedy, the long exclusion of women from science on the basis of biological differences between men and women, feminist science studies have been a particularly rich area of scholarship since the 1980s. Carolyn Merchant demonstrated the power of this approach in her pivotal 1980 book The Death of Nature (Merchant 1980). But it was her 1983 piece “Isis’s Consciousness Raised” that explicitly argued for the methodological value of bringing the theoretical and analytical categories of women’s studies into mainstream scholarship in the history of science. Asking whether feminist history of science can “contribute to a new perspective on our discipline,” Merchant both described current trends in feminist science studies and provided important future directives for developing such an approach (Merchant 1982, 398; reprinted in Kohlstedt 1999). Feminist history of science involves a female perspective on science, nature, and society, the study of female challenges to traditional scientific roles, and a female consciousness concerning the origins of women’s lower position and consequent exclusion from historical literature. A feminist approach to science and history can reveal hidden biases in a field that in recent years has considered itself free of the cultural assumptions of the present when treating the science of the past. Beyond this it can offer alternative interpretations of the rise of science, scientific professionalization, and the scientific world view, and it can create new synthesis in our field.

(Hypatia, “Feminism and Science,” 2, 3 (Autumn 1987) and 3, 1 (Spring 1988), including 17 articles and a bibliography of women in science).

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The program Merchant laid out soon began to flourish. While she herself developed ecofeminism, other historians and feminist theorists began to publish works demonstrating both the richness of feminist science studies and the severity of its critique of androcentric bias in science. This literature is vast, and it is only possible here to point to influential works and survey general positions advanced by feminist scholars. By the mid-1980s, a number of important works had appeared, many by scientists inspired by the women’s movement. In 1984, Ruth Bleier published Science and Gender: A Critique of Biology and its Theories on Women, followed 2 years later by the edited volume Feminist Approaches to Science (Bleier 1984, 1986). In 1985, Evelyn Fox Keller published Reflections on Gender and Science (a collection of six essays), and in 1986, Anne Fausto-Sterling’s Myths of Gender appeared, as did Sandra Harding’s The Science Question in Feminism, followed in 1991 by Whose Science? Whose Knowledge? Thinking from Women’s Lives (Keller 1985; Fausto-Sterling 1986; Harding 1986, 1991). These works laid out several areas of inquiry. Keller explored gender in science broadly, but especially through linguistic analysis; Fausto-Sterling closely examined the biology of gender and sexual difference; while Harding developed feminist epistemology, including gender analysis and androcentrism in biology (for an overview, see Crasnow et al. 2015). Harding also elaborated on standpoint theory, suggesting that viewing science through the lens of women provides a more adequate understanding of its development (Harding 1991, 1997; see also Bowell n.d.; Harris 2002). In 1988, Donna Haraway critically evaluated the concept of objectivity in science, developing the influential idea of “situated knowledges,” or the view that all knowledge is situated within specific bodies, locales, and time frames and, more importantly, that negotiation and selection are involved in what counts as “scientific” knowledge (Haraway 1988). The following year, in Primate Visions, Haraway explored primatology from a feminist perspective, introducing the notion that gendered assumptions can influence theory construction. This book spawned both vigorous discussions as well as fruitful new lines of investigation (Haraway 1989, 1991). The same year, Londa Schiebinger’s The Mind Has No Sex? Women in the Origins of Modern Science appeared, which applied feminist analysis to examinations of women’s participation in science (Schiebinger 1989). These and other stimulating works set the framework for feminist critiques of science in the next decade and beyond (Creager et al. 2001; Keller 2001; Keller and Longino 1996a, b; Koertge and Patai 2003; in addition to those mentioned, other leading scholars in feminist science studies include Adele Clarke, Joan Fujimura, Evelynn Hammond, David Hess, Ruth Hubbard, Emily Martin, Sue Rosser, Linda Layne, Richard Levins, Richard Lewontin, Rayna Rapp, Hilary Rose, Bonnie Spanier, S. Leigh Star, Sharon Traweek, and Nancy Tuana). By the 1990s, feminist science studies had moved into the mainstream of the history of science (Laslett et al. 1996). The 1991 gathering of historians of science to chart new disciplinary directions, the “Conference on Critical Problems and Research Frontiers in History of Science and History of Technology,” included two papers addressing “Knowledge and Gender.” Evelyn Fox Keller Keller (1995) provided a historical overview of feminist scholarship and suggestions about future lines of inquiry; Sally Gregory Kohlstedt (1995) provided a historiographical survey of extant literature on women in science. As Kohlstedt noted: “current research provides glimpses into how

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such studies might transform the history of science so that it incorporates not only the subject but also the methods and sensibilities we have uncovered together in our search for the position of women in science and scientific thinking” (Kohlstedt 1995, 58). In 1995 Kohlstedt and Helen Longino organized a conference at Minnesota that resulted in the edited Osiris volume “Women, Gender, and Science: New Directions” which specifically raised the question about the relationship between women in science and women and science studies (see Kohlstedt and Longino 1997). Four years later, considering whether feminism had changed science, Londa Schiebinger argued that indeed it had and would continue to do so in the future. “What is needed is a critical understanding of gender, how it works in science and society – and, while we have a better understanding of this now than we did just 20 years ago the question of gender in science remains a project for STS in the twenty-first century” (Schiebinger 1999a, b). Schiebinger’s prediction has proved accurate. Judging by recent scholarship, both feminist science studies and research on women in science have significantly benefitted from gender analysis (see Opitz 2015). The critique of potentially sexist assumptions in science has sharpened thinking in several areas of the life sciences, including cell biology, evolutionary biology, primatology, archeology, and developmental biology (Biology and Gender Study Group 1976; Keller 1996, 1997; Milam 2010; Fedigan 2001; Wylie 2001; Gilbert and Rader 2001; Montgomery 2013). Gender analysis has also provided probing reviews of the “woman question” and gender in nineteenth-century biology (Richards 1983, 1989, 1997, 2017) as well as twentiethcentury genetics (Satzinger 2009a, b, 2012, 2016). Moreover, it has opened new lines of inquiry. Following up the early analysis of the Biology and Gender Study Group (1976), for example, scholars exploring the ideology of biology have examined sexist connotations in language used to describe different aspects of sexual reproduction (Martin 1991; Fausto-Sterling 1986, 1989; Richardson 2013). Moreover, the power of gender analysis in exploring masculinity in science as well as women in science is demonstrated by recent works (Nye 1997; Milam and Nye 2015; Subramaniam 2014). Departing from previous tendencies to marginalize women’s roles in science, recent scholarship has begun to reinterpret the spaces available for performing science as well as to reevaluate ways women participated in science. An exciting new area of focus, especially given women’s near absence from past historical accounts, is on visual culture, which helps document women’s presence (Kohlstedt and Opitz 2002; Shteir and Lightman 2006). Another is the analysis of “domestic” science, which upends the previously disparaging connotations of women’s traditional locus and opens new lines of inquiry into spaces for producing knowledge (Lindsay 1998; Richmond 2006; Opitz et al. 2016). A subset of domestic science explores the scientific activities of aristocratic women (Opitz 2004, 2006, 2012). Studying new niches that women could fill has broadened our understanding of work in the life sciences. For example, women moved into new areas previously dominated by men (such as horticulture) and institutional positions outside the university (like museums) (Opitz 2013, 2014; Kohlstedt 2013; Mohr 2010). Scholarship thus continues to fulfill the goals of feminist science studies: to create depictions of science that more accurately reflect not only the lived experiences of all participants but also the historical record.

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Areas for Future Work in the History of Women in Biology Despite the many gains that have been made in understanding women’s participation and gender dynamics in science over the past 40 odd years, much remains to be done. The work of “recovery” continues. Women were present in biology in increasing numbers from the 1870s, yet we still lack a consistent understanding of their work and its social and intellectual context. It remains important to continue to identify individuals, but perhaps even more pressing is the need to place women within the institutional and disciplinary development of new experimental areas that emerged in the life sciences after 1900. As we have seen, women were present in the new laboratories of genetics, biochemistry, ecology, and other areas. Yet, despite the foundational work of social constructivism and science and technology studies, an androcentric perspective still persists. As Robert Nye pointed out, a masculine honor code was adopted when science professionalized in the late nineteenth century, and remnants continue both in scientific practice and historical and sociological interpretations (Nye 1997). Recent sociologists have shown that women do not fare well when competing within the hierarchical status systems in universities and other large bureaucracies (Smith-Doerr 2004; Gaughan and Bozeman 2016). Steven Shapin and Naomi Oreskes pointed out decades ago that much of the work in science is carried out by technicians and assistants, whose roles are frequently overlooked and yet their work may have been pivotal (Shapin 1989; Oreskes 1996). Given that in the late nineteenth and early twentieth centuries women’s work in science was generally categorized as that of assistants or technicians, this literature has helped open a way to view their contributions without the pejorative connotations of their official status (Oakes 2014). More recently the theme of “Invisibility and labour in the human sciences” has been explored in a collected volume (see the essays in Bangham and Kaplan 2016). While sociologists have examined the dynamics of collaborative knowledge production over the past century, less attention has been given to the complex organizational patterns and social interactions within research (see, however, Gerson 1998, Chap. 10). Indeed, sociologists as well as historians often focus solely on the head of the group (e.g., the one who receives a Nobel Prize) without interrogating important contributions made by other of members, including women. Yet, as scholars of women in science often find, status within institutes or laboratories often does not well correspond with an individual’s accomplishments (Smith-Doerr 2004). For example, protocols for assigning authorship frequently disadvantage women; the department or institute head who appears as first author may not even have participated in the study. Even more disheartening is to find that the activities and disciplinary prominence of particular women were openly acknowledged by their contemporaries and yet completely overlooked in historical accounts. It is important for historians as well as sociologists to take these lessons to heart in order to produce richer and more accurate historical accounts of biology. Another desideratum is the need for more international comparisons. As Hilda Smith noted over two decades ago, “Before we can produce a truly comparative women’s historiography, we must know much more of the facts and historiographical disputes among those concentrating on women in other countries . . .. To gain a

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comparative historiography of women, we cannot concentrate on theory or on broad categories which presumably organize scholarship in a number of nations: we must understand better why a particular set of scholars came to pose specific questions at one time out of their shared intellectual knowledge and concerns” (Smith 1989, 99–100). Although such comparisons require considerable work, it is only by comparing and contrasting different national and cultural traditions that general trends become apparent. In this way, different cultural and social norms within universities, institutes, and groups that either aid, hinder, or otherwise influence women’s participation in biology can be identified (for examples, see Canel et al. 2000). Margaret Rossiter pointed out long ago that the ultimate challenge posed by feminist critiques of science is to create “a better, more inclusive, more humane and relevant history of science” (Rossiter 1987, vii). A more inclusive history of biology would focus on the work of women as well as men to present more accurately the social and institutional structures as well as intellectual developments that have promoted the life sciences. It would also better capture the actual dynamics of knowledge production. The remedy is simple: all one needs to do is to ask “Where are the women, and what did they do?” Including women in historical accounts of biology might not always change the fundamental narrative, but at the very least it would enrich the story. To be sure, doing women’s history can be demanding. As many early practitioners noted in the 1970s, in comparison to other fields, women’s history was “the most challenging in substance, theory, and methodology” (Smith 1994, 15). Certainly, the growth of a robust body of literature, as well as the development of “tools of gender analysis,” has helped to guide scholarship (Schiebinger 1997; Fedigan 2001). However the main hurdle is not so much analysis but rather locating archival materials. Generally only women who achieved scientific prominence have dedicated archives, and even then the holdings are often not extensive. Scholars thus have to be creative, for example, tracing women through the men with whom they were associated. This requires pouring through correspondence and institutional records to find references to women’s activities. Undergraduate institutions are often good sources of biographical information. Alumni records often offer opportunities to trace an individual’s later career, given that after graduation women tend to keep in touch more consistently with their alma mater than institutions where they did graduate work. Because women generally changed their name upon marriage, this practice presents yet another hurdle in trying to follow someone throughout their career. Here the internet has been a great boon for women’s scholarship. Searching on someone using different names often turns up data or source materials that otherwise might have remained obscure. Women’s historians are often sophisticated sleuths in hunting down pieces of evidence and creating meaning from an incomplete puzzle. Pursuing women’s history, however, provides ample rewards. Not only has the body of literature created over the past four decades succeeded in expanding our understanding of women in biology – not simply identifying who the women were and what they did – but also providing insights into their passions, their insights, their collaborations, and the ways they discovered just to get on with their work. Finding the women and bringing their experiences in biology into the

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historiographical record certainly provides individual satisfaction, but more importantly, it also enriches and complicates our understanding of the history of biology itself. Women have long been part of the human endeavor to understand the living world and have participated in erecting the edifice of the modern life sciences. It is indeed important to include them in our histories.

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Contents Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Reflections on Terminology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A Distorted Past . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Race in America, Race in Science . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Some Future Directions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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Abstract

In the early and mid-twentieth century, race was widely regarded among physical anthropologists (and most Americans) as an essential, biological component of human identity. Within that consensus, however, there were serious debates over the nature of race, heredity, identity, classifications, and scientific methods. More recently, historians have begun articulating a more complex and contested picture of racialist theorizing among anthropologists, not least by Boas himself, who was a leading racial scientist and one of the foremost cultural anthropologists in America. Histories that address the complex interplay between cultural and biological theory offer a different assessment of the persistent contradictions and complexity of the American racial landscape and its scientific representations.

This essay was adapted from Tracy Teslow, Constructing race: the science of bodies and cultures in American anthropology. Cambridge: Cambridge University Press, 2016 T. Teslow (*) University of Cincinnati, Cincinnati, OH, USA e-mail: [email protected] © Springer Nature Switzerland AG 2021 M. R. Dietrich et al. (eds.), Handbook of the Historiography of Biology, Historiographies of Science, https://doi.org/10.1007/978-3-319-90119-0_18

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Introduction Much of the scholarship on the history of race and anthropology has perpetuated a narrative in which progressive scientists, chiefly Franz Boas and his students, championed a cultural understanding of human variation against pernicious essentialist, racist conceptions of difference (King 2019).1 In the early and mid-twentieth century, race was widely regarded among physical anthropologists (and most Americans) as an essential, biological component of human identity. Within that consensus, however, there were serious debates over the nature of race, heredity, identity, classifications, and scientific methods. Moreover, the somatic emphasis of racial anthropology did not preclude anthropologists from also understanding race in cultural and historical terms. More recently, historians have begun articulating a more complex and contested picture of racialist theorizing among anthropologists, not least by Boas himself, who was a leading racial scientist and one of the foremost cultural anthropologists in America. Histories that address the complex interplay between cultural and biological theory offer a different assessment of the persistent contradictions and complexity of the American racial landscape and its scientific representations.

Reflections on Terminology “Race” is a profoundly inadequate term. The failure of English speakers to devise a richer language to distinguish among the very many entities to which the term “race” has been applied suggests either a shocking lack of imagination or perhaps a rather sinister but unwitting ingenuity that casts the umbrella of one small word and its supposed conceptual clarity – an immutable natural kind – over a remarkably messy heterogeneity of objects. In the course of more than 400 years, race has referred to a family lineage; an animal or plant species or variety; a people or nation; large groups of people transcending tribal or national boundaries; the entirety of humanity; and even the class of all men or all women; in addition to the more common recent usage to denote inherited physical characteristics shared with a (often ill-defined) group and to denote a social class, often associated with particular physical features but not limited by that.2 A paltry few variations on “race” – “racial,” “racist,” “racialized,” 1

Charles King, a professor of international affairs and government, has authored one of the latest entries in the genre of popular histories that perpetuate this narrative, God’s of the Upper Air: How a Circle of Renegade Anthropologists Reinvented Race, Sex and Gender in the Twentieth Century (2019). The eager reception of his account by progressive critics is a good example of the allure of this recurring narrative, and the difficulty scholars have had getting traction for a more complex account of racial and cultural theory and practice. 2 The OED lists 62 adjectival permutations on the word “race” (“race line,” “race man,” “race question,” etc.). The term itself has six definitions, the earliest dating from 1547. Zoological definitions and application to identifiable groups of people (biologically or otherwise) date from the late sixteenth century. Use of the term to designate a supposedly physically distinct group of people dates from the early eighteenth century. Oxford English Dictionary (Oxford and New York: Oxford University Press, 1989).

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“racialist” – and “ethnic,” a twentieth-century innovation – represent the universe of terms. The advent of “ethnic,” which was supposed to clarify matters, has not. It has become as confused in its usage as “race,” simply replacing it without any change in intent,3 and often used as a polite way to refer to what is still also conventionally referred to as “race” when meaning essential, theoretically apparent, bodily differences, and their attendant individual and social ramifications. This usage is found in popular as well as academic contexts. Outram and Ellison offer an analysis of the conceptual issues involved in the persistent use of “race” and “ethnicity” among medical researchers, and the tendency to reify “ethnicity” (Outram and Ellison 2010). Carol C. Mukhopadhyay and Yolanda T. Moses discuss not only race and its cognates, but also classificatory terminology (Caucasoid, Negroid, Mongoloid, and the like), color language, and normative epithets like “fair skinned.” As Walter Benn Michaels noted in The Trouble with Diversity, faced with the fallacy of biological race, his students “just stopped talking about black and white and Asian races and started talking about black and European and Asian cultures instead” (Michaels 2006, 5–7). Artist Kara Walker, whose provocative phantasmagorical cutouts evoking nineteenth-century slavery raise complex questions about race, sexuality, gender, and American society, has argued that Americans do not really want to abandon the race concept. Echoing Ben Michaels, she has argued that Americans’ obsession with race is a form of identity. “I think really the whole problem with racism and its continuing legacy in this country is that we simply love it,” she said. “Who would we be without the ‘struggle?’” (Armstrong 1997). Historian Matthew Jacobson has argued that “ethnicity” was used as a tool to de-racinate whites by creating non-racialized groups among a broad “Caucasian” racial type, a trend that he argues was coincident with the mid-twentieth-century process of separating race and culture while reifying race along color lines (Jacobson 1998). Historically, and in practice, the biological and cultural remain intermingled, despite terminological and ideological attempts to sunder them. In most historical and popular uses, race has connoted both heredity, as somatic difference, and cultural difference, although the formulation “race and culture” suggests that culture resides only in the second term and that “race,” by binary opposition, denotes only bodies and ancestry. When meanings of “race” are taken apart, however, we see that “culture” is deeply embedded in nearly all its permutations. Cultural or social understanding of race often includes bodies and heredity, if only because the social construction of race was originally erected upon morphological – or purportedly morphological – differences. Since Ashley Montagu introduced “ethnic” in the 1940s as a way to denote groups without a biological component, there have been efforts to use race as a similarly purely sociological notion (Montagu and Ashley 1942, 1945). But this has been very hard to do without importing essentialist or hereditarian connotations. Race is a resilient, protean concept that is constantly redefined and renegotiated. It is one of the sturdiest, most promiscuous products of modernity, a persistent, frequently invidious tool

3

For an early discussion of this problem, see Montagu 1942 and 1945, where he advocated for replacing “race” with “ethnic group.”

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deployed in endless permutations and elaborations, repeatedly remade in shifting configurations of politics, law, economics, culture, and science. From a social and historical perspective, “race” is best understood not as a natural category or biological essence, but as a force in American life, an ideological system rooted in the pursuit of group and individual advantage that produces and is sustained by political, cultural, social, economic, legal, and scientific practices and their material consequences (Lopez 2005; Fields 1982).

A Distorted Past The historiographic treatment of Franz Boas is particularly emblematic of the lens through which racial science has too often been viewed. The picture of Boas as the father of cultural relativism and a champion against scientific racism that developed in the late twentieth century was a consequence of developments within anthropology in the postwar period, as well as in the early historiography of the discipline. It may come as a surprise to readers accustomed to accounts of Boas as the author of the modern, relativistic culture concept that in the years after his death in 1942 he was dismissed by many anthropologists, most notably his own students Alfred Kroeber and Clyde Kluckhohn, as a diligent data collector but no theoretician, someone who failed to systematize his work, and who, in his relentless, a-theoretical empiricism, actually stunted development of the culture concept and the field of anthropology more generally (Stocking 1963, 1966, 1968; Kroeber and Kluckhohn 1952). A broader view of the history of anthropology helps explain why Boas was first dismissed as insufficiently theoretical in the 1950s, and then later embraced as the critical figure in the development and promulgation of the culture concept. In the 1940s and 1950s, American anthropology experienced a resurgence of theories about human cultural and social formations that emphasized commonalities and universal qualities, functions, or processes, and was especially concerned to put anthropology on a fully “scientific” footing. Functionalism prospered at the University of Chicago; cultural ecology thrived at the Smithsonian Institution’s Institute of Social Anthropology under Julian Steward; the University of Michigan anthropologist Leslie White promoted an evolutionary culture theory that emphasized the discovery of universal cultural laws (Darnell 1998; Peace 2004). At odds with the evolutionists and others promoting a quest for universal laws of cultural development, Boasian anthropologists like Kroeber and Kluckhohn defended historical particularism by rooting the culture concept, not in Franz Boas’s articulation of it across his decades of work, but further in the past, in English anthropologist Edward Tylor’s 1871 definition of culture and its subsequent elaboration by a variety of anthropologists (Stocking 1966; Tylor 1871). Historian George Stocking’s work on Boas and his development of the culture concept profoundly changed the earlier, dismissive view. Even though Stocking was careful to characterize Boas as a “transitional figure” who “retained strong residual elements” of the nineteenth-century evolutionary “commitment to ‘progress in civilization,’” he nonetheless made the point that it was Boas, and not Tylor, who

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in the face of the enormous diversity of human traditions and social practices had articulated a systematic critique of cultural evolution. According to Stocking, it was left to his students to fully elaborate his discussion of “cultures” into the modern culture concept (Stocking 1966, 878–879). The irony was that, in response to the postwar resurgence of evolutionism and scientism in anthropology, some influential Boasian anthropologists had promoted cultural relativism and historical particularism in part by dismissing Boas’s critical contribution. Following Stocking’s intervention in anthropologists’ disciplinary history, and the eclipse of evolutionary and other universalizing theoretical orientations, Boas took on a founder’s mantle in American anthropology similar to that afforded Charles Darwin among modern evolutionary biologists. Anthropologists have been notably interested in their own disciplinary history, and in particular in examining the Boasian legacy. Numerous accounts of Boas’s anthropology and of the history of anthropology have been authored by anthropologists themselves since Boas died in 1942. These include A. Irving Hallowell, “The History of Anthropology as an Anthropological Problem” (1965); Robert Lowie, “Boas Once More” (1956a), and “Reminiscences of Anthropological Currents in America Half a Century Ago” (1956b); Alfred Kroeber, “The Place of Boas in Anthropology” (1956), and “A History of the Personality of Anthropology” (1959); Melville Herskovits, Franz Boas: The Science of Man in the Making (1953); Cole Franz Boas: The Early Years, 1859–1906 (1999); Margaret Mead and Ruth Bunzel, eds., The Golden Age of American Anthropology (1960), as well as Mead’s unconventional biography of Ruth Benedict, An Anthropologist at Work: Writings of Ruth Benedict (1959) and the later work, Ruth Benedict (1974), and her autobiography, Blackberry Winter (1972); Marvin Harris, The Rise of Anthropological Theory (1968); Leslie White, “The Ethnography and Ethnology of Franz Boas” (1963); and Stephen O. Murray, “The Non-Eclipse of Americanist Anthropology during the 1930s and 1940s” (1999). Regna Darnell has produced more than 30 books, articles, and edited volumes on the history of American anthropology, including Along Came Boas: Continuity and Revolution in Americanist Anthropology (1998), Theorizing the Americanist Tradition (1999), and Invisible Genealogies: A History of Americanist Anthropology, Critical Studies in the History of Anthropology (2001). The embrace of Boas as the father of the culture concept was accompanied by a pronounced lack of interest in his racial science. Following World War II and the humanist turn that prompted disciplinary organizations, as well as international organizations such as the United Nations Educational, Scientific and Cultural Organization (UNESCO), to speak out against racism, Boas’s antiracism was much more palatable, and more congruent with the politics of cultural relativism, than his racial science. By the 1970s, the treatment of Boas was enmeshed in the epistemological crisis that led humanists and social scientists, particularly anthropologists, to question positivism and claims to universal knowledge. Historian of anthropology Regna Darnell perhaps put it best when she noted: “The real Boas tends to disappear amidst the apotheosis of angst-ridden anthropological reflexivity” (Darnell 2000). For anthropologists and those who study the history of anthropology, the last quarter of the twentieth century saw a profound turn inward toward self-reflexiveness and an

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equally profound self-consciousness and discomfort with the traditional project of anthropology, its methods, and assumptions (Darnell 2000; Liss 1998; Clifford and Marcus 1986). For anthropologists, history has functioned in part as identity formation, reconstructing a lineage that justifies or supports current commitments, particularly for a discipline acutely uncomfortable with its participation, direct and indirect, in racial injustice and its legacy around the world. All history is written from a situated vantage point, but a discipline’s practitioners are especially burdened with the weight of their intellectual and methodological genealogy when attempting to reconstruct a fraught past (Haraway 1988). At the same time, cultural relativism and the multiplication of culture into cultures, from singular to plural – a part of the anthropological legacy – was so deeply and widely embraced that even the most selfreflexive scholars sometimes failed to see it as a historically contingent, evolving worldview. This is less true of anthropologists themselves, who continued to debate the nature and usage of the culture concept even while employing it. Darnell (1997) and Fox and King’s (2002) collection addresses recent controversies about the meaning, necessity, and utility of the culture concept, while Descola (2013) explores the idea of culture, its complex relationship to the idea of nature, and offers a way to move beyond the traditional dichotomy. The corollary to the embrace of “cultures” has often been an adamant, ironically nearly reflexive, rejection of biology or heredity as useful or interesting explanatory frameworks for understanding humanity. Those few who espoused such notions often have been dismissed as racists, part of a genuinely repugnant tradition of slaveholders, eugenicists, and white supremacists. Despite extensive historical and anthropological attention paid to Boas by scholars and anthropologists, until recently the scholarship (with some important exceptions) had not comprehensively – even adequately – addressed Boas as a physical anthropologist. Much of the initial historiography on physical/biological anthropology was created by anthropologists themselves in collections that traced the development of the discipline via the lifework and signal accomplishments of its practitioners, without much social or historical context (Spencer 1982, 1996, 1997; Little and Kennedy 2010). Boas is a key figure in the story of physical anthropology prior to World War II not only for the ways he and others defined the problem of race and its solutions, but also because in delineating his views and practices we confront prevailing historiographies, both visions of the past created by historians writing the history of anthropology, as well as in the history anthropology tells itself, that seem to have deliberately ignored a substantial portion of his life’s work. For many years, much of the historiography elided or distorted the actual character of his physical anthropology, and specifically his interest and belief in race. Much of the historical work that treats Boas fails to confront the full nature of his views, perhaps because it does not seem to comport with the kind of figure many historians, anthropologists, and others have wanted him to be, one crucial to a historiographical narrative about the ascendance of culture and the decline of scientific racism (Dominguez 1994; Reardon 2012). In introductory remarks to selections of Boas’s early writings on race, George Stocking made a similar point, noting that Boas has been recognized principally as a critic of racism, but that early arguments that “were conditioned by

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the racist milieu in which he wrote,” which brought together various aspects of anthropology and which revealed an early conception of culture embedded in his critique of racial determinism, had been overlooked (Stocking 1974; Allen 1989). Stoler (1997) offers an incisive analysis of the ways historians’ and other scholars’ own assumptions and desired narratives about race and racism shape histories of race. It used to be the case that Boas’s racial science received cursory treatment. There are now a number of important exceptions to this. George Stocking’s work is a prominent and crucial exception, along with more recent work by Regna Darnell (2000) and Julia Liss (1998). Moreover, there is a growing body of work that delves into the history of race in anthropology and the social sciences with nuance, including Lee D. Baker, From Savage to Negro: Anthropology and the Construction of Race, 1896–1954 (1998) and Anthropology and the Racial Politics of Culture (2010); Vernon J. Williams, Jr., Rethinking Race: Franz Boas and His Contemporaries (1996) and “‘What is Race’: Franz Boas Reconsidered” (2007); Nancy Leys Stepan, “Race, Gender, Science and Citizenship” (1998); Alice L. Conklin, In the Museum of Man: Race, Anthropology and Empire in France, 1850–1950 (2013); and John P. Jackson and David J. Depew, Darwinism, Democracy, and Race: American Anthropology and Evolutionary Biology in the Twentieth Century (2017); noteworthy too are Duster (2003), Armelagos (2004), Smocovitis (2012), Lindee and Radin (2016), Lipphardt (2015, 2017). Elazar Barkan discusses Boasian racial science and interwar racial science more generally at length in The Retreat of Scientific Racism: Changing Concepts of Race in Britain and the United States Between the World Wars (1992), although this work relies heavily on an unfortunate dichotomization of scientists into “racists” and “egalitarians,” a retrospective oversimplification that obscures more than it illuminates. An early, critical history of racial anthropology examined physician Samuel George Morton, one of Franz Boas’s precursors. In The Mismeasure of Man (1981), evolutionary biologist Stephen Jay Gould’s well-known analysis of American antebellum craniometry and its popularization, introduced broad public and academic audiences to the ways early physical anthropology constructed and perpetuated racial hierarchies. His detailed critique of how leading scientists’ racial prejudices overtly and implicitly shaped their practices and conclusions was an important early attempt to analyze the relationship of various human sciences to American race and racism. Some historians faulted Gould for focusing on the failings of individual practitioners rather than on the broader structures of science, and the racial categories and conceptualizations scientists employed (Allen 1984). Scientists and historians continue to spar over whether Gould was a perceptive investigator revealing the socially embedded nature of scientific practice, in particular the conscious and unconscious racial bias among past scientists and theorists, or an ideologue who misread or distorted evidence to support his perspective (Lewis et al. 2011; Weisberg and Paul 2016). Historian Donna Haraway’s early studies of racial science offered a dense analytical exploration of not only individual scientists and their practices but also the societal matrix in which the sciences she examined produced stories about human being. Her study of Carl Akeley and the African

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gorilla diorama at the American Museum of Natural History, and her account of Sherwood Washburn and the advent of the “new physical anthropology” in the early 1960s, along with meditations on the nature of knowledge production and biopolitics, were significant contributions to the increasing efforts of anthropologists, historians, and science studies scholars to grapple with the nature/culture dichotomy, and more nuanced approaches to the history of human variability (Haraway 1989, 1997). From the historians’ perspective, it is vital to understand the past of racial science as the full range of practice that it was, from work that was regarded at the time as wholly sound and legitimate to racist propaganda, and the relation of all of it to the society in which it was produced, which, in the twentieth-century United States was a society that embraced strict immigration restrictions, witnessed extensive racial violence, and contested Jim Crow segregation. Historians should not ignore scientific and intellectual contexts that existed in the past because they no longer seem to hang together as a legitimate whole, and retrieve only the science, the ideas that make the story coherent from a presentist point of view. Indeed, recapturing the fullest possible picture of how science and scientists functioned within society serves an epistemological and historical purpose in counteracting to some degree the powerful ideological and rhetorical force of science itself, which continually recasts itself, through reconstructed histories of its leading researchers and fundamental ideas, as a process outside of history and society. Anthropology, among the most self-conscious of the sciences, is surely less guilty of this than some. But even anthropology constructed histories that served current purposes and theoretical commitments more than the historical record. The failure to face unpalatable pasts squarely risks leaving us with incomplete or distorted accounts that ultimately fail to adequately explain past ideas and actions, and therefore also fail to illuminate how that past informs on our present and future. In gaining a deeper understanding of racial science, we can begin to see the outlines of a broader story about how race disappeared from twentieth-century anthropology in very specific ways, replaced by or transformed into other concepts and categories, and how the transformation of physical anthropology into biological anthropology both contributed to and reflected similar major shifts in modern social and political discourse in America and the wider world. By looking more carefully at the actual landscape of theory and practice surrounding the study of race, we can begin to see where the real disjunctions and actual continuities lie between various pasts and presents. For example, with a fuller understanding of Boas’s views, and of those who came after him and followed his lead, such as Harry Shapiro, Ruth Benedict, and Ashley Montagu, as well as those who did not, such as Henry Field, Arthur Keith, and Earnest Hooton, we can begin to see continuity with views about race that are more typical of the later twentieth century in the United States, the very views that seem to have made it difficult to lucidly comprehend the racial science Boas and others practiced. Anthropology was foundational to American racial formation precisely because it promoted both racial essentialism and cultural relativism. Taking the science of race seriously, understanding it as its practitioners did, in all its

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complexities, contradictions, and shifting emphases, illuminates not only how Americans used to think about race and culture, but helps us understand why and how race has remained a complex feature of modern life (Malik 1996; Tucker 2002; King 2004; Wailoo and Pemberton 2006; Brattain 2007; Schaffer 2008; Farber and Cravens 2009; Fabian 2010; Meyerowitz 2010; Hart 2011; Qureshi 2011; Hazard 2012; Yudell 2014).

Race in America, Race in Science In the 1990s, biological determinism linked to race seemed to have reemerged. Physicians promised better health via race-targeted drugs like BiDil. Companies offering DNA ancestry tests prompted genealogical odysseys as celebrities like Oprah Winfrey searched for their origins. Long-festering debates over race and IQ exploded back into public view with the publication of The Bell Curve (Kahn 2012; Wailoo and Pemberton 2006; Washington 2006; Gates 2007; Herrnstein and Murray 1994). By the first decade of the twenty-first century, race – or formulations that sounded remarkably like familiar racial categories – had re-emerged as a locus of debate among geneticists, anthropologists, historians, and other scholars, along with a broader public. While many consumers in the United States and Europe eagerly embraced genetic ancestry testing, many others who thought racial essentialism and biological determinism had been roundly discredited decades earlier viewed these developments with alarm, and puzzlement. From a longer historical perspective, however, we can see these developments as yet another stage in an ongoing dynamic in the United States between predominantly biological and predominantly social solutions to pressing problems. The discourses of nature and nurture, biology and society, have been consistently part of American culture for more than a century. Jenny Reardon has argued that, unlike past eras, when biological or sociocultural explanations for human variation and its associated attributes waxed and waned in oppositional trajectories, more recently our society has experienced these modes of explanation, and the sociopolitical tensions associated with them, virtually simultaneously. “Increasingly,” she argued, “politicization and de-politicization, control by the state and individual empowerment, the negation of biological concepts of race and their proliferation happen concurrently.” The effect is a sense of dissonance, and of perpetual crisis (Reardon 2008; Reardon et al. 2006). The complex mix of biologically essentialist explanations and historically or culturally grounded theorizing that one finds in the work of Franz Boas, Ruth Benedict, and Harry Shapiro in the interwar period is distinctive of its era but also remarkable in the ways it reaches across the supposed gulf of World War II (and the Evolutionary Synthesis) to illuminate tensions and connections between the biological and the cultural in the latter half of the twentieth century. Indeed, interwar racial science was much like postwar racial science in its complex brew of biology and society, the promiscuous intermingling of bodies and cultures. Rather than seeing the postwar reaction against race and racism, in which the UNESCO Statements on Race and a reoriented “new physical anthropology” are often framed as a decisive turning

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point, we might instead view the postwar era as part of an ongoing construction of human variation that has consistently involved a volatile, unstable mix of cultures and bodies (Barkan 1996; Brattain 2007; Müller-Wille 2007; Armelagos 2008; Hazard 2012; Selcer 2012; Tilley 2014; Bangham 2015; Smocovitis 2012; Haraway 1989). The construction of race in science has never been an either/or proposition. It has never been a question of race or culture, heredity or society, bodies or minds. The discourses of human variation since the eighteenth century incorporated visions of bodies, capabilities, and cultures as a means to explain diversity and justify hierarchy. That is not to say that there has been easy consensus. For an entity that supposedly encompasses a set of patently evident natural kinds, race has been a profoundly unstable scientific object, subject to constant contestation and in need of continual reconstruction. The science of race has been marked by persistent debates about methods, types, and implications. Although the nature of consensus clearly shifted away from racial essentialism and hereditarian racial typology between 1900 and 1970, this must be seen within a broader historical perspective of the waxing and waning of hereditarian, biologically deterministic, and essentialist views. The conviction that science will ultimately solve the riddle of human diversity is one that has persisted since at least the application of comparative anatomy to the “problem” of African Americans and Native Americans in antebellum America. As new scientific theories and technologies emerged, they have invariably been applied to the conundrum of race, from Johann Friedrich Blumenbach’s zoological classifications in the eighteenth century, to Samuel Morton’s craniometry and Francis Galton’s composite photographs in the nineteenth century, to genetics in the twentieth and twenty-first centuries. And yet, despite the growing authority of the sciences within American society, there has been no period of US history when essentialism, hereditarianism, and white supremacy have not been challenged. In periods when a hereditarian vision has been most widely embraced, we still find critiques and persistent challenges. Conversely, humanist, cultural accounts of human diversity never fully drive out biological essentialism or hereditarian claims. Indeed, from the perspective of a historian of racial science, what is striking is the unrelenting persistence of “racial” questions (Harrison 1995). Concerted efforts following World War II to reject scientific racism and promote cultural relativism (and later to promote an account of race as a social construction without biological meaning) can be seen as a moderately effective attempt to combat divisive biological, hereditarian essentialism by erecting a sturdy boundary between “race” and “culture” that had not existed earlier in American history and that began eroding again with the resurgence of genetics. The history of racial and cultural studies within anthropology is not one of successive Kuhnian paradigms, with Boasian cultural relativism displacing hereditarian, deterministic “scientific racism,” so much as tandem or enmeshed discourses with institutional and methodological histories that have been frequently allied as the practice of anthropology and other social and natural sciences developed in the United States. A variety of scholars have questioned the extent to which anthropology fits a Kuhnian paradigm of scientific development, and some have questioned the idea that

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“culture” replaced “race” in anthropology in any straightforward Kuhnian sense. In part this is attributable to the “four field” nature of American anthropology, in which the overall discipline of anthropology coalescing out of a variety of prior fields, including anatomy and medicine, philology, and natural history and philosophy, developed via four fairly distinct subfields: linguistics, archaeology, ethnology (later cultural anthropology), and physical (later biological) anthropology. Some also have argued that Kuhn’s theory of paradigms has been most fruitfully applied to the natural sciences, especially the “hard” experimental sciences (physics, chemistry, etc.), and less successfully to the human, social, and historical sciences, which some labeled “pre-paradigmatic.” These distinctions are still a matter of debate. Kuhn himself made no claim that his rubric applied to the social sciences (Kuhn 1962; Darnell 1998; Baker 1998; Kuznar 2008; Fuller 2000). Much attention has been paid by scholars to the nuances of the culture concept and cultural theorizing in anthropology; the racial science of anthropologists deserves a similarly careful examination (Darnell 1997; Fox and King 2002). As historian Bronwen Douglas has forcefully argued, we need more critical studies “grounded in rigorous vernacular reading of the original works of Euro-American thinkers whose broad, labile gamut of positions on human differences is often collapsed under the homogenizing rubric of racism” (Douglas 2008). Lumping together all anthropological racial science under the umbrella of “scientific racism” obscures rather than illuminates precisely the vexed concepts and practices we seek to understand. The fuller history of racial anthropology, and racial science more broadly, also requires attention to a wider cast of characters and institutions, including non-western, Black, indigenous, and other people of color, as practitioners, critics, and subjects. There is now a growing body of such work, long missing from the historiography (Graham 1990; Dubow 1995; Dain 2002; Penny and Bunzl 2003; Anderson 2006a; Kramer 2006; Braun and Hammonds 2008; Douglas and Ballard 2008; Saada 2012; Wade et al. 2014; Delgado 2020). Ann Laura Stoler’s theoretical interventions and historical studies have been highly influential in colonial and postcolonial studies across a number of disciplines, and foundational to work in the history of science, including her comparative analysis of the ways in which North American and colonial studies historiographies treated the “intimate domains” of sex, home, and child rearing implicated in the making of racial categories and imperial rule (Stoler 2001a; Cooper 1996). Two recent special issues of leading journals have also examined physical/biological anthropology in a wider, global context. A 2012 volume of Current Anthropology devoted to “The Biological Anthropology of Living Human Populations: World Histories, National Styles, and International Networks” brought together an array of anthropologists and historians to reexamine the history of race, anthropology, and genetics across the global south, Asia, Europe, and North America. A 2014 volume of Isis, “Focus: Relocating Race,” considered the problem of race and science from the perspective of a wide range of disciplines, locations, and periods, especially non-western and colonial ones. In the first four decades of the twentieth century, between “race” and “culture,” race was the more firmly established concept within and outside anthropology. For the American public, common sense held that bodies and societies were deeply,

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causally intertwined. “Civilization,” or the lack of it, was commonly viewed as a manifestation of race, and a given racial group’s inherent capacities. In this view, “race” and “ethnology” were merely different components of the “civilization” process. Franz Boas articulated a competing theory that stressed the fluid, historically contingent, environmentally conditioned process of human cultural formation, and rejected the idea of universally fixed stages of social, technological, and political development espoused by the evolutionists. The pluralistic culture concept propounded by Boas had been articulated by him before 1900, but was not fully assimilated into the practice of ethnology until the 1920s, and not into the rest of the social sciences and beyond until toward the end of the interwar period. The general public did not broadly embrace it until a good deal later. A broad public acceptance of the culture concept is tricky to pin down, but it seems to have taken a long while before the idea that diverse cultural forms were equally valid took hold. Lois Banner cites Margaret Mead’s assertion in 1928 that she felt she still had to explain it for her reading public. Ruth Benedict also went to great lengths in both Patterns of Culture in 1934 and Race: Science and Politics in 1940 to explain what she meant by “culture.” Arguably, it remained a matter of contention in 1955 when Edward Steichen mounted “The Family of Man” at the Metropolitan Museum of Art, a humanist vision of brotherhood illustrated by cultural variations on universal themes (Gilkeson 1991; Stocking 1992, 2001; Darnell 1998, xii; Banner 2003, 195; Mead 1973). In the nineteenth century and the first decades of the twentieth century, most of the American public and most scientists who studied human variation believed in racial typology. They thought race was something essentially biological and static, and at their most rigid adhered to the idea that not only groups of people but individuals could be accurately categorized. But even within this general consensus there were varied approaches. Whereas Henry Field at the Field Museum of Natural History in Chicago and his British mentor, Arthur Keith, promoted a traditional typological view of fixed racial types, Franz Boas and Harry Shapiro at the American Museum of Natural History in New York approached the problem of human variation with a more holistic view. Boas and Shapiro, like Field and Keith, believed that race was a feature of human biology. But Boas and Shapiro regarded it as malleable as well, associated with groups of people who moved through space and time and who existed everywhere in cultures that created environments that shaped the form of human bodies. By the late 1930s, many scientists, especially geneticists and physical anthropologists, objected to shoddy scientific and popular racial ideologies that trafficked in overgeneralization and what they viewed as folk knowledge or personal prejudice. Boas and Shapiro objected that race and racial characteristics were not biologically well understood, had been inadequately studied (in part because of serious methodological problems), and were improperly theorized (e.g., no one had any idea which characteristics were evolutionarily or hereditarily significant, nor just what the role of environment was, although Boas’s 1911 study of changes in the form of the head had demonstrated that there was one). The problem, from their point of view, was poor science, not a fundamental conceptual error in the object of study. They opposed racism, not race.

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By the late twentieth century many social scientists and humanists interested in human diversity, along with a significant body of biologists and geneticists, had rejected typological racial concepts altogether and came to see race as it had been theorized for much of the nineteenth and twentieth centuries as exclusively a cultural/social phenomenon that has no biological existence. But even this social constructionist definition often acknowledges that race is “real” in the sense that, as a social category, race remains a powerful factor in human relations and American society. By the 1960s, much work in genetics, anthropology, and human biology tried to redefine race in evolutionary terms, looking at population groups over time, not seeing race as static but rather as a function (as Shapiro termed it in 1939) of migration and environment. Although broadly speaking a generally biological vision of race predominated in the decades prior World War II, and a culturally constructed one in the latter twentieth century, a multiplicity of views have existed throughout the century, shifting in complex relation to changing concepts and methods in both the sciences and wider society. A wide range of investigators, using significantly different notions of “race,” have sought an answer to the same persistent question: why is there so much variety among human beings? Among cultural anthropologists, the slow diffusion of the culture concept into American society influenced the strategy they adopted in their attempts to combat prejudice and discrimination. By the 1930s, many anthropologists increasingly shared a sense that race had become distorted in the popular (and frequently in the scientific) mind. Misapprehensions about race had already led to all kinds of objectionable action, repercussions that only grew more dire as Nazis consolidated their power in Germany (Weindling 1993; Teicher 2020). Anthropologists felt an urgent need to address what they saw as gross distortions, misunderstandings, and misapplications. Most adhered to a liberal humanist, positivist belief that “truth” and knowledge would undo prejudice. Following profound theoretical interests in individual psychology and the relationship of individuals to their cultures, many cultural anthropologists believed that the root problem was individual racial prejudice. Thus, most of their solutions were focused on disabusing individuals of their biases and misinformation. They did not consistently offer an analysis of structures of racism, power, or wealth, and even when they did, such as Ruth Benedict’s work in Race: Science and Politics, they did not see those structures of racism as related to the “fact” of race, except through the political exploitation of bias. They did not view the race concept itself as inherently problematic, or likely to engender the disparities and discrimination they attacked. By the end of the interwar period, anthropologists were stressing the separate nature of race and culture, describing the “facts” of human physical variation on the one hand and the limitless diversity of culture, which varied independently of race, on the other. For Boasian ethnologists, interwar efforts to rethink race occurred simultaneously with efforts to change Americans’ perception of society and culture, shifting it away from social evolutionary hierarchies of “primitive” and “civilized” and toward the plural, relativistic culture concept. The effect, which would have far-reaching, unintended consequences, was to reify race while relativizing culture. The new postwar “common sense” held that race was merely a matter of inherited physical characteristics, while culture was an entirely separate process by which a group of people constructed a social life (Teslow 2016).

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George Stocking argued that the firm boundary erected by anthropologists between culture and race has been “defended” since it was first embraced in the 1920s for theoretical and methodological reasons, but also for ideological and political ones (Stocking 2001, 315). The challenge for the historian of race and racial science is not unlike that of postcolonial scholars attempting to provide subtle, nuanced accounts of an imperial world too easily cast into victims and villains, colony and metropole. Like the landscape of race and racial science in the United States, the landscape of race and power in colonial contexts is and was complex, and often cut against the tidy conceptual and historical categories that help scholars parse the past but which too often flatten or obscure the thing we most desire to understand. Race has always been a protean concept, slippery and flexible, used in contradictory and inconsistent ways, interwoven with politics, class, and economics, gender and sex, social relations, and material cultures, in myriad configurations. The historian is perpetually in danger of oversimplifying a complex story, imagining that the past was less complex than the present. Ann Laura Stoler captured the challenges and promise of historical work in these complex pasts when she argued that for the scholar of colonialism, “Replacing colonialism’s stick figures with actors who combined goodwill and sympathy for the dispossessed with racist beliefs underscores that colonial regimes were not less complex racially inflected social and political configurations than are ours today” (Stoler 2001, 896). The historian of racial science in the twenty-first century faces a similar challenge. The failure to remember The Races of Mankind exhibition at the Field Museum of Natural History, Ruth Benedict’s extensive exposition of race, Franz Boas’s decadeslong pursuit of racial science, and Harry Shapiro’s long and influential career in physical anthropology is not because these scientists and institutions were obscure, their works unknown. The Races of Mankind sculptures still grace the halls of the Field Museum. Benedict’s Patterns of Culture and Race: Science and Politics are still in print. Franz Boas published dozens of his papers on race in an anthology of his work shortly before he died in 1942; indeed, his last words were said to be, “I have a new theory about race” (Mead 1959, 355).. It seems that historians and other scholars have, until quite recently, had a blind spot for much of twentiethcentury racial science. Perhaps that is because the anthropology of race for much of the twentieth century contradicts a satisfying narrative of progressive antiracism in which physical anthropology was a practice vanquished and best forgotten. Like a narrative of World War II as the “good war,” the transplantation of cultural relativism where scientific racism once lurked provided a welcome resolution to a tumultuous, often violent and painful, past. The desire to forget the racial science of physical anthropologists is wedded to a liberal humanist ideology that once believed, and perhaps still hopes, that science is self-correcting, that “good” science will prevail over “bad,” just as freedom and equality will ultimately prevail over racism and other oppressive ideologies and practices. Indeed, cultural relativism’s triumph over scientific racism ranks with oxygen replacing phlogiston and the heliocentric universe as canonical case studies used to demonstrate that whatever might be the subjective limits to objective knowledge and however illusory the idea of progress, science can correct

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at least some profound misapprehensions. In these narratives, it is science itself, sometimes fueled by a progressive philosophy and often practiced by members of the oppressed classes, that forces repressive, reactionary ideologues to confront the inconsistencies, errors, and hypocrisy of racist, essentialist accounts of human variation that constructed the “other” as naturally, immutably less capable and deserving. Race is revealed as a social construct, not a natural discovery. The problem with this narrative is not that it is utterly wrong, but what it privileges and what it obscures. As Ann Stoler has argued, racial discourses are characterized by their “polyvalent mobility,” their remarkable flexibility and adaptability, and their ability to “draw on the past as they harness themselves to new visions.” Ideologically malleable, the concept of “race” has been employed by historical actors on all sides, not merely by those in power or those advocating a racist perspective. Scholars’ preoccupation with “debunking” race and the science of race, she argued, left us with a body of literature that relied on an oversimplified account (Stoler 1997, 190–192, 195–196, 198). Too often in such narratives, racial scientists become “stick figures,” and the complexities of nonlinear conceptual, social, political, and professional relations get flattened or ignored altogether. In a critique that still resonates, Stoler argued, “Histories of racisms that narrate a shift from the fixed and biological to the cultural and fluid impose a progression that poorly characterizes what racisms looked like . . . and therefore have little to say about what distinguishes racisms today”(Stoler 1997, 198). Since Stoler published her incisive analysis, much of the work on race and public health, eugenics, sociology, and psychology has adopted the more nuanced view she advocated, and has explored the “polyvalent mobility” of race that she identified (Anderson 2006a; Shah 2001; Wailoo 2001; Hammonds 2008; Reverby 2000; Reardon 2004, 2017; Stern 2005; Jackson 2001b; Weidman 2004; Tucker 2002). But the history of anthropology, especially the core narrative about culture and “scientific racism,” has largely resisted a similar analysis until very recently. It is this unwitting blindness, an “epistemology of ignorance,” to use Charles Mills’s evocative and apt phrase, born of a desire to be free of the odious burden of race and racism, that leads directly to the surprise and disgust with which The Bell Curve was greeted by so many (Mills 1997; Sullivan and Tuana 2007). The importance of capturing the complexities of a racial science that, whatever its currency in the mid-twentieth century, has been effectively marginalized for the last generation within anthropology, lies not only in correcting the historical record but also in improving our understanding of the present (Haraway 1989; Jackson 2001a; Brattain 2007; Zimmerman 2001; Mukhopadhyay and Moses 1997; Harrison 1999; Goodman et al. 2003; di Leonardo 1998; Kuklick 2008; Evans 2010; Conklin 2013; Lyons and Lyons 2004). Even a cursory acquaintance with everyday life in the United States should disabuse us about the efficacy with which culture replaced race. Race continues to intermingle with culture in ways that are virtually inexplicable had the ideas promoted by racial scientists been vanquished as thoroughly as the popular narrative suggests. An illustration of this conundrum has been discussed by Australian anthropologist Gillian Cowlishaw, who has written about anthropologists’ efforts to cope with race and racism in the modern world. Cowlishaw was trained in the early 1970s in Australia, a period when cultural relativism had become

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the consensus position in the field and had been widely embraced in Western societies. Although the analytical concept of “race” had been replaced by terminology such as “traditional”4 by the time she entered graduate school, Cowlishaw argued that categories such as “‘traditional’ filled precisely the same semantic space previously occupied by the Aboriginal race” (Cowlishaw 2000, 108). This semantic shift, she argued, did not lead to any diminishment of racism toward the Aboriginal people of Australia, but merely obscured the continuing significance of their racialized identity and hobbled anthropologists’ ability to respond effectively to the highly racialized and racist social, political, and economic world of contemporary Australia. In her view, the utter displacement of race with culture in anthropology “had the effect of confusing the interpretation of a world where race and culture, however conceptualized, had become so intertwined that attempts to separate their functioning were futile.” She argued that anthropologists simply accepted a biological definition of race and rejected it, leaving it uncontested (Cowlishaw 2000, 111). Cowlishaw warned that “while speaking of race may appear to reproduce racial categories” – a fear that motivated anthropologist Ashley Montagu and geneticist Julian Huxley to urge abandoning the term in the 1940s – “not speaking about race allows racial differentiation to flourish unchallenged” (Cowlishaw 2000, 101). It is not only modern anthropologists and other scholars who grapple with the meaning and implications of race and culture. Earlier anthropologists struggled to make sense of human diversity in the context of their times, too. Close reading of anthropologists’ texts, lectures, and imagery reveals scientists’ struggles to consciously create concepts and methods in accord with their discipline, the larger world, and their own instincts and proclivities, what Tom Holt has aptly termed a “sediment” of prior theoretical and personal commitments that constrain conceptual possibilities (Holt 2000, 23). Changes in the conception of race were sometimes a response by anthropologists to a disjunction between what they “knew” to be true and what their process gave them as “facts” about the world. In the late nineteenth century, Paul Topinard began to question the existence of race when measurements failed to yield categories and standards by which individuals and groups could be reliably classed, but retreated to a search for hypothetical pure races to explain the origins of such heterogeneous people, in part because his “common sense” experience of the world told him that races existed (Stocking 1968, 59; Jackson 2000). Similarly, Arthur Keith advised Henry Field that there was no reason to burden his Field Museum exhibition project with an enormous program of measurement and statistical analysis, as Harvard anthropologist Earnest Hooton urged, because it was plain to anyone who walked down the street that races existed.5 For scientists who studied race, it was a problem of how to make their work congruent with their

4 A term which itself has been critiqued as ethnocentric and racist. See, for example, Trouillot 2003, Pels 2008, and Bhabha 1994. 5 Reported to Malvina Hoffman by Stanley Field, September 9, 1931, Box 3, Malvina Hoffman Collection, 850,042–1, Special Collections, The Getty Center for the History of Art and the Humanities, Los Angeles, California.

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experience of and vision for the world, as well as a problem of how to orient their work with that of other sciences – and because the subject was race – also a social, cultural, and political problem, one whose complexities cannot be encompassed in a set of dichotomous epithets such as “racist” or “egalitarian.” How do we explain the growth of sociobiology, the appeal of evolutionary psychology, the return of IQ, the fascination with DNA analyses of ancestry for celebrities, commoner, and dog alike (Reich 2018; Wade 2014)? These trends suggest that ideas about race that had supposedly been put to rest with the ascendance of the anthropological culture concept endured, and that the broad embrace of cultural explanations for social phenomena has been incomplete. The intellectual framework with which Americans had narrated the undoubted shift toward culture over the course of the twentieth century was not a matter of rejection or replacement so much as transformation or ascendance. The desire to explain diverse bodies and cultures as a result of natural or biological processes looks like a persistent theme in American history, not an anomaly. For much of the twentieth century, anthropologists grappling with human diversity employed notions of race that included both naturalized, quantified bodies and concepts of culture, and both racialized peoples and efforts to understand diverse cultural and environmental contexts. The fact that this racial theorizing was not decisively replaced by cultural relativism but instead persisted in tandem with it for most of the twentieth century had profound consequences for popular understanding of human diversity. Despite concerted efforts by anthropologists to confine the notion of race to bodies, and to stress culture as the source of variation among societies and their denizens, race and culture remained intermingled, entwined in both professional and popular discourse. The failure to reject race itself along with racism left confusion about the nature and source of human variation, and hardened a biological definition of differences. The racial dimensions of World War II, particularly the horrors of Nazi racial policies, played a crucial role in the formation of race and the sciences of human variation in the twentieth century, but it was only one catalyst in a postwar shift away from racial essentialism. By the 1940s, a number of factors combined to create a climate in which it seemed politically, scientifically, and socially necessary to assert a humanist framework over and against a racialist, and frequently racist, one. The history of racial formation, and the salient place of anthropology and other sciences in that formation, long predate World War II, and persist after it in ways that intersect with global political, economic, and military events, but were not governed by them. The profound political, social, and economic shifts associated with World War II, civil rights activism, and postcolonial movements coincided with equally profound shifts in biological and anthropological theory and practice. Although racial essentialism was increasingly subordinated to a humanist cultural framework in the postwar era, it persisted in competing formulations of human diversity despite concerted efforts to eliminate it. The history of nineteenth- and twentieth-century racial essentialism and biological determinism is not one of long ascent followed by steep decline, but rather a story of waxing and waning in relation to complex societal conditions, not least the development of new scientific theories and tools with which to attack the persistent puzzle of human variation. This transformation of civilization

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into “cultures” has been regarded as a progressive story of the evolution of a scientific discipline. The story of racial anthropology is a messier, less appealing tale in which anthropologists and others, inside and outside the academy, persisted in attending to physical differences long after many had become uncomfortable with it. As a result, despite its important role in constructing race in the United States, for most of the twentieth century racial anthropology was often forgotten, dismissed, or mischaracterized. In recent decades, this neglect reflected the conflicted way Americans have thought about race, asserting that it does not exist or does not matter while still clinging to the reality and supposed significance of physical and, increasingly, genetic differences. There has been a profound dual consciousness about race in the United States, evident throughout the twentieth and now twentyfirst centuries. It is simply not true that racial science and race thinking were vanquished by the success of Boasian cultural anthropology, nor by the rejection of eugenics following its nadir under German National Socialism, nor as a result of African Americans’ persistent efforts to attain equality and civil rights, nor the success of postcolonial movements, nor a host of other things that were supposed to have contributed to a thorough rejection of scientific racism. Historians and others have begun to explore this complex history from new angles, using new conceptual insights, to better understand how race works in anthropology, in science broadly, and in society.

Some Future Directions In the twenty-first century, the “re-emergence” and “re-inscription” of race via genetics, along with the persistence of sociocultural and political contests over race and identity, has spurred a number of fruitful avenues for current and future research in history of science and science studies (Duster 2005; El-Haj 2007). Anthropologists, who once spurned or avoided race, as Gillian Cowlishaw and Faye Harrison had observed, have returned to the subject as well, in an effort to understand and speak to its continued usage, especially in public health, medicine, and genomics. The questions, concepts, and frameworks emerging in recent scholarship point to productive opportunities for future historical investigations and contemporary science studies. Broad interest among scholars in global, international, and transnational studies continues to inform the study of race and science, including anthropology, particularly where it intersects with health, medicine, and genomics (Reardon 2004). Our understanding of racial formations outside the United States and Europe and their relationship to anthropology, other sciences, and medicine, particularly in configurations that transcend national bounds, is a work in progress. Further investigation of the development and shape of racial sciences and anthropology outside the United States and Europe can build on work that has begun, particularly dealing with the global south, Asia, and Latin America (Anderson 2014). A promising angle in this work is a focus on particular localities, the movement of knowledges and practices across spaces and cultures as well as across time, and the distinctive forms and roles

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scientific knowledge and practice take in distinct locales. An example of this work is the aforementioned 2014 Isis volume on the theme of “Relocating Race” which aimed to reexamine racial sciences in disciplines, periods, and places beyond the traditional boundaries of western historiography. The essays consider “the ways in which new ‘locations’ required and inspired fundamentally different articulations of the ideas with which we have become familiar” (Seth 2014, 761). According to section editor Suman Seth, the collection aimed to “‘make strange’ our assumptions about the conditions of possibility for nineteenth-century race science in its oncecanonical form” and, citing Dipesh Chakrabarty, sought to “provincialize race science” (Seth 2014, 761). Chakrabarty (2000) discusses “provincializing” as a way of re-thinking modernity, colonialism, universalism, and knowledge production “from and for the margins.” The August 2011 volume of Hispanic American Historical Review devoted to “Science and Medicine in Latin America” took historian Nancy Stepan’s 1991 book on eugenics in Latin America as a point of departure to reflect on the state of the field and where further studies in science and medicine, including racial sciences, might lead (Stepan 1982, 1991). There is a robust literature on science in colonial contexts (Fullwiley 2011; Stern 2011; Sleeboom-Faulkner and Patra 2011; Hartigan 2013a; Chung 2014). A recent example is Veronika Lipphardt and Alexandra Widmer’s anthology, Health and difference: Rendering human variation in colonial engagements (2016). A growing literature on “decolonizing” science and the history of science brings together the study of race, science, imperialism, and postcolonialism with a more direct concern for indigenous knowledges, voices, and relations to power. Using the language of “decolonization,” this approach builds on and brings together scholarship examining the “situated” partial and multiple perspectives that inform scientific technologies, practices, and knowledge, including scientific networks in and across nation states and empires, with the extensive literature on empire and colonialism to create science studies, and histories, that attend explicitly to internal and external colonial contexts. While much of this work is concerned with current practice and active decolonization of pedagogy and scientific practice, there is also a place for historical accounts and pedagogy. One of the goals of this work is to re-orient a historiography of science that long privileged knowledge produced in “first world” nations and imperial centers toward a de-centered assessment of the ways knowledge, practice, and policy have been built, sustained, and challenged globally and locally (Dear 2005; Elshakry 2010; Harding 2011; Sturgeon 2007; Reardon 2009; Smith 2013; Raina 2019; Anderson 2018; Fischer 2018; Duara 2018). And to do this without undermining or co-opting indigenous, subaltern, and other historically marginalized voices, knowledge, and social justice activism (Tuck and Yang 2012). The late 1990s also saw a renewed interest among anthropologists in the problem of race, with concerted attempts to talk across their subdisciplines and with scholars in other fields. In a special issue of American Anthropologist, Faye Harrison convened specialists from inside and outside anthropology on “Expanding the Discourse on ‘Race’” (1999). It was a response to Carol C. Mukhopadhyay and Yolanda T. Moses’s call the previous year, in an essay published in the American Anthropologist, for cultural and biological anthropologists to “combine conceptual

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and methodological forces to cultivate a dialogue on race” (Mukhopadhyay and Moses 1997, 610). In her introduction to the volume, Harrison argued that “to effectively revive our discipline’s race-cognizance and deploy it in strategic arenas of public debate, policy formation, social action, and other loci of democratic practice,” it would be necessary to “reestablish race as a central issue for anthropological inquiry and analysis” (Mukhopadhyay and Moses 1997, 610). In 1999, the Wenner-Gren Foundation underwrote a symposium on “Anthropology in the Age of Genetics: Practice, Discourse, Critique” that brought together sociocultural and biological anthropologists, as well as evolutionary biologists, human geneticists, sociologists, and historians, to assess how anthropologists were “using, responding to, and studying genetic practices and discourses.” This included both the use of genetics to study human individuals and populations by biological anthropologists, and cultural analysis of genetic practices and discourses.6 The subsequent anthology, Genetic nature/culture: Anthropology and science beyond the two-culture divide (2003), captured the symposium’s cross-disciplinary effort to explore “a genetic borderland” of practices, resources, power relations, and meanings (Abby Lippman’s “geneticization”) in a way that investigated and transcended the usual nature/culture dichotomy to consider instead the “coconstitution of nature and culture and all their familiar iterations” (Lindee et al. 2003, 2; Lippman 1991, 1992). The collected essays represented “the fruits of a dialogue on genetics that brings together cultural studies of genetic knowledge production and natural scientific studies that foreground cultural-historical context” (Lindee et al. 2003, 2). Studies of contemporary genetics are accompanied by a growing body of historical work exploring race and earlier nineteenth- and twentieth-century genetics, especially human genetics. A series of anthologies and special issues in leading journals have continued an intensive examination of the vexing question, as one volume put it, of race in a genomic age (Fujimura et al. 2008; Koenig et al. 2008; Whitmarsh and Jones, Hartigan 2013). Among the most exciting conceptual developments in the study of race and anthropology, and race and biology more generally, is the use of “biocultural” or “biosocial” frameworks to rethink traditional dichotomies of culture and nature, race, science (Rabinow 1992, 2008). Instead of an oppositional relationship, “biosociality” or “biocultural” approaches invite us to examine the complex ways in which sociocultural, economic, and political forces are interwoven with biological factors, and the ways scientific practices produce knowledge about nature. Initially spurred by an interest in understanding how late twentieth-century genomic science was altering the way people thought about individual and collective identity, the concept has been taken up and adapted more broadly to the complexities of “race” and its resurgence in recent decades. The “return” of race has prompted scholars to re-examine the historical and contemporary trajectories of how human diversity has

Final Report, “Anthropology in the age of genetics: Practice, discourse, critique,” Wenner-Gren Foundation International Symposium #124, June 11–19, 1999, Teresópolis, Brazil, url: http://www. wennergren.org/history/anthropology-age-genetics-practice-discourse-critique 6

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been and is defined and deployed. Anthropology has been deeply engaged in this project. In their important 1997 essay, Mukhopadhyay and Moses noted that the traditional effort in their discipline to sever the connection between race and culture had failed to diminish the force of racial formations in society and also limited the ability of anthropologists to offer a more powerful explanatory framework. Mukhopadhyay and Moses argued that “it is time to consider relinking the cultural and the biological, but within a radically different paradigm” that would “situate human biodiversity within a sociocultural framework, in effect reuniting culture and biology by embedding biology in society and culture” (Mukhopadhyay and Moses 1997, 525–526). Advocating “unified biocultural approaches to race and human biodiversity,” they suggested researchers could more effectively address “the fluid, temporally, historically, and culturally specific nature of races and other social groupings in human history” (Mukhopadhyay and Moses 1997, 526). In 2006, in a formulation that would have sounded familiar to Franz Boas or Harry Shapiro, anthropologist Alan Goodman noted “biologicals are also culturals in that their study, development, and evolution involve interrelated ideological, social, and political economic processes that were there from the very beginning” (Goodman 2006, 225). Faced with the accelerating application of DNA analysis to questions of human ancestry and identity, Goodman employed the concept of biosociality to call for geneticists and biological anthropologists to better understand “the culturalness of biology or webs of power,” arguing “to understand why, when, and how biology and genetics are interwoven with the social, political, and ideological requires embracing complexity and a new biocultural anthropology” (Goodman 2006). Historians and others have begun to examine the relationships between genetics, identity, and the history of racialization, and the implications of that convergence, along similar analytical lines. Ian Whitmarsh and David S. Jones’s collection, What’s the use of race?: Governance and the biology of difference (2010), focuses on “the promise and dangers of genetics, race, and governance,” including the relationship of colonial histories and shifting racial/ethnic formulations to current matters of law, regulation, and medical practice (Whitmarsh and Jones , 2). Keith Wailoo, Alondra Nelson, and Catherine Lee’s anthology Genetics and the unsettled past: the collision of DNA, race, and history (2012) addresses the history of race in/and genetics and the use of genetics to resconstruct the past (Wailoo et al. 2012). Work that investigates race in bio-anthropological, biomedical, and biosocial contexts includes George T. H. Ellison, Duana Fullwiley, Amade M’charek, Marianne Sommer, and Rachel Watkins (Wailoo 2003; Watkins 2008; Ellison et al. 2008; Sommer 2010; M’charek et al. 2013; Fullwiley 2014). The voluminous historiography on race has begun to include significant research on the ways physical and biological anthropologists played an important role in the formation and employment of the idea of race. Trends in global and colonial studies, in the broad persistence of race in society – in public health arenas and medicine, as well as other corners of science such as forensics, and especially in genetics – along with conceptual insights that promise a move beyond dichotomy in thinking about nature or race and culture, suggest avenues for further fruitful examinations of race and anthropology, present and past.

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Montagu M, Ashley F (1942) Man’s most dangerous myth: the fallacy of race. Columbia University Press, New York Montagu M, Ashley F (1945) On the phrase ‘ethnic group’ in anthropology. Psychiatry 8:27–33 Mukhopadhyay CC, Moses YT (1997) Reestablishing ‘race’ in anthropological discourse. Am Anthropol 99:517–533 Müller-Wille S (2007) Race et appartenance ethnique: la diversité humaine et l’UNESCO Déclarations sur la race (1950 et 1951) [Race and ethnicity: human diversity and the UNESCO declarations on race (1950 and 1951)]. In: 60 ans d’histoire de l’UNESCO [60 years of UNESCO history]. UNESCO, Paris, pp 211–220 Murray SO (1999) The non-eclipse of Americanist anthropology during the 1930s and 1940s. In: Valentine L, Darnell R (eds) Theorizing the Americanist tradition. University of Toronto Press, Toronto, pp 52–74 Outram SM, Ellison GTH (2010) Arguments against the use of racialized categories as genetic variables in biomedical research: what are they, and why are they being ignored? In: Whitmarsh I, Jones DS (eds) What’s the use of race?: governance and the biology of difference. MIT University Press, Boston, pp 91–124 Peace WJ (2004) Leslie A. White: evolution and revolution in anthropology. University of Nebraska Press, Lincoln Pels P (2008) What has anthropology learned from the anthropology of colonialism? Soc Anthropol 16:280–299 Penny G, Bunzl M (eds) (2003) Worldly provincialism: German anthropology in the age of empire. University of Michigan Press, Ann Arbor Qureshi S (2011) Peoples on parade: exhibitions, empire, and anthropology in nineteenth-century Britain. University of Chicago Press, Chicago Rabinow P (1992) Artificiality and enlightenment: from sociobiology to biosociality. In: Crary J, Kwinter S (eds) Zone 6: incorporations. Zone Books, New York, pp 234–252 Rabinow P (2008) Afterword – concept work. In: Gibbon S, Novas C (eds) Biosocialities, genetics and the social sciences–making biologies and identities. Routledge, London, pp 182–192 Raina D (2019) The vocation of indigenous knowledge and sciences as metaconcept. In: Abu-ErRub L et al (eds) Engaging transculturality: concepts, key terms, case studies. Routledge, London Reardon J (2004) Race to the finish: identity and governance in an age of genomics. Princeton University Press, Princeton Reardon J (2008) Race and biology: beyond the perpetual return of crisis. NTM J Hist Sci Technol Med 16:373–377 Reardon J (2009) Anti-colonial genomic practice? Learning from Chacmool and the Genographic project. Int J Cult Prop 16:199–204 Reardon J (2012) The democratic, anti-racist genome? Technoscience at the limits of liberalism. Sci Cult 21:25–47 Reardon J (2017) The Postgenomic condition: ethics, justice and knowledge after the genome. University of Chicago Press, Chicago Reardon J, Dunklee B, Wentworth K (2006) Race and crisis. Social Science Research Council forum on “Is race ‘real’?” June 2006. http://raceandgenomics.ssrc.org/Reardon/ Reich D (2018) Who we are and how we got Here: ancient DNA and the new science of the human past. Oxford University Press, New York Reverby S (2000) Tuskegee’s truths: re-thinking the Tuskegee syphilis study. University of North Carolina Press, Chapel Hill Saada E (2012) Empire’s children: race, filiation and citizenship in the French colonies. University of Chicago Press, Chicago Schaffer G (2008) Racial science and British society, 1930–1962. Palgrave Macmillan, New York Selcer P (2012) Beyond the cephalic index: negotiating politics to produce UNESCO’s scientific statements on race. Curr Anthropol 53:s173–s184 Seth S (2014) Introduction. “Focus: relocating race”. Isis 105:759–763

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Tucker WH (2002) The funding of scientific racism: Wickliffe Draper and the Pioneer Fund. University of Illinois Press, Urbana Tylor EB (1871) Primitive culture: researches into the development of mythology, philosophy, religion, art, and custom. J. Murray, London Valentine L, Darnell R (eds) (1999) Theorizing the Americanist tradition. University of Toronto Press, Toronto Wade N (2014) A troublesome inheritance: genes, race and human history. Penguin, New York Wade P, Beltrán CL, Restrepo E, Santos RV (eds) (2014) Mestizo genomics: race mixture, nation, and science in Latin America. Duke University Press, Durham Wailoo K (2001) Dying in the City of the blues: sickle cell Anemia and the politics of race and health. University of North Carolina Press, Durham Wailoo K (2003) Inventing the heterozygote: molecular biology, racial identity, and the narratives of sickle-cell disease, Tay-Sachs, and cystic fibrosis. In: Moore DS, Kosek J, Pandian A (eds) Race, nature, and the politics of difference. Duke University Press, Durham, pp 235–253 Wailoo K, Pemberton S (2006) The troubled dream of genetic medicine: ethnicity and innovation in Tay-Sachs, cystic fibrosis and sickle cell disease. The Johns Hopkins University Press, Baltimore Wailoo K, Nelson A, Lee C (eds) (2012) Genetics and the unsettled past: the collision of DNA, race, and history. Rutgers University Press, New Brunswick Washington HA (2006) Medical apartheid: the dark history of medical experimentation on black Americans from colonial times to the present. Harlem Moon, New York Watkins RJ (2008) Knowledge from the margins: W. Montague Cobb’s pioneering research in biocultural anthropology. Am Anthropol 109:186–196 Weidman N (2004) Constructing scientific psychology: Karl Lashley’s mind-brain debates. Cambridge University Press, Cambridge Weindling P (1993) Health, race and German politics between National Unification and Nazism, 1870–1945. Cambridge University Press, Cambridge Weisberg M, Paul DB (2016) Morton, Gould, and bias: A comment on ‘The mismeasure of science’. PLoS Biol 14:e1002444 White L (1963) The ethnography and ethnology of Franz Boas. Texas memorial museum bulletin, vol 6. Texas Memorial Museum, Austin Whitmarsh I, Jones DS (2010) What’s the use of race? Modern governance and the biology of difference. MIT Press, Cambridge, MA Williams VJ Jr (1996) Rethinking race: Franz Boas and his contemporaries. The University of Kentucky Press, Lexington Williams VJ Jr (2007) ‘What is race’: Franz Boas reconsidered. In: Campbell JT, Guterl M, Lee RG (eds) Race, nation and empire. University of North Carolina Press, Chapel Hill Yudell M (2014) Race unmasked: biology and race in the twentieth century. Columbia University Press, New York Zimmerman A (2001) Anthropology and anti-humanism in Imperial Germany. University of Chicago Press, Chicago

Local, Global, and Transnational Perspectives on the History of Biology

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Ana Barahona

Contents Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . The Bipolar Distinction Between Center and Periphery . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A Different Landscape in the History of Science: The Transnational Change . . . . . . . . . . . . . . . . Circulation and Collaborative Networks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Decentering the Nation, Crossing Borders and the Transnational as a Unit of Historical Analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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The transnational perspective in the history of science has pointed out the need to reconstruct cross-borders narratives that account for how the knowledge produced in developing countries forms part of international knowledge as it circulates in networks of collaboration. This perspective has enabled the production of narratives that go beyond the national framework through analysis of transnational participants and processes and has allowed new ways of thinking about the history of biology in national and regional contexts. This manuscript will focus on the influence that George Basalla’s diffusionist model had on historians of science working in non-US-European science, the abandonment of the nation as the unit of historical analysis, and finally, on the recent trends on the history of biology. When proposed in 1967, the diffusionist model of science offered a new tool for historical analysis that was comparative and transcultural, in which the local element was seen as relevant. Nevertheless, despite having opened a new agenda for the history of science, this model has shown serious historiographical limitations. At the end of A. Barahona (*) Universidad Nacional Autonoma de Mexico, Mexico City, Mexico e-mail: [email protected] # Springer Nature Switzerland AG 2021 M. R. Dietrich et al. (eds.), Handbook of the Historiography of Biology, Historiographies of Science, https://doi.org/10.1007/978-3-319-90119-0_19

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the 20 th Century, the center-periphery perspective in the history of science was challenged, and recent studies have tried to introduce a different point of view in which the dynamic of scientific practices and the dismantling of imperial projects are the subjects of historical scrutiny. Recent scholarship has shown that the transnational perspective allows to escape from the tension between the local and the global, and to assume a wider narrative beyond national borders. This shift has allowed giving a richer account of knowledge on the move.

Introduction The field of science and technology studies (STS) has recently focused on the need to write connected transnational narratives based on a reciprocal treatment of global and local contexts that describe the dynamics of scientific practices. As historian Sanjay Subrahmanyam has pointed out, connected histories, as opposed to comparative ones, need to be written to shed light on local resistances and global trends. This transnational approach of STS abandons the nation as a unit of analysis in order to understand the development of science history. It also moves beyond Euro-US-centered narratives in order to explain the role of international networks and the circulation of knowledge, people, artifacts, and scientific practices. This new perspective, according to historian of science and technology Simone Turchetti, historian of physical sciences Néstor Herran, and historian and sociologist of science Soraya Boudia, could promote a novel understanding of science as historical phenomenon. The transnational approach that coalesced at the end of the Cold War has been subsequently influenced by the effects of globalization, multiculturalism, and the formation of circuits of practices, knowledge, and people, in which scientific developments go beyond nation-state borders, being the collaborative networks the units of historical analysis. This new focus draws attention to the flows themselves and moves away from mere international issues. Thus, recent debates regarding global and local contexts have called attention to circulation networks that explore interregional exchanges and transnational circuits that allow quicker crossborder transmission of scientific practices and a faster flow of people, ideas, and artifacts. In the case of recent historical studies of science in Latin America, Asia, and Africa, much of the research performed under this approach has indicated that despite their historiographical and epistemological importance, narratives on the national science perspective have revealed its analytical limitations. This research indicates the need to reconstruct transnational stories that account for how the knowledge produced in developing countries forms part of international knowledge as it circulates in these networks of collaboration. This perspective has enabled the production of narratives that go beyond the national framework through analysis of transnational participants and processes and has permitted new ways of thinking about the history of biology in national and regional contexts. Some of these narratives have insisted on more transnational and global histories that take into account the dynamics of scientific practices. This chapter is divided in two sections. In the first, I will begin with Basalla’s diffusionist model to show how the dichotomy center-periphery influenced historians of science working in non-European science. The amount of work that Basalla’s model generated was significant in

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bringing to the fore the local context, although in tension with the global dimension. In the second part, I will try to show how the transnational turn could solve the tension between the global and the local, pointing to the transnational perspective by using circulation and collaborative networks as historical categories. Also, I will address the abandonment of the nation as the unit of historical analysis inasmuch as it is conceived as a very restrictive category (paragraphs adapted with permission of the journal Dynamis).

The Bipolar Distinction Between Center and Periphery The first to use the center-periphery distinction was the American sociologist Edward A. Shils in 1961 in the context of the study of international relations and colonialism (Shils 1961, 1991).1 His approach to sociology led to his role as a bridge between the research traditions of European and American sociology. His studies were framed around the diverse ways of understanding society and the role of intellectuals in politics and public policy. As the editor of Minerva, he promoted the center-periphery distinction, allowing many authors to find a place to discuss this distinction as an analytical tool (Guillem-Llobat 2008). Among the many authors that published in that journal at the time was sociologist Joseph Ben-David. He used this dichotomy to compare nineteenth-century academic structures in some European countries and had a great influence in the field (Ben-David 1970).2 For Shils, “the pattern of center and periphery, which comprises inequality of achievement and status attests to the universal validity of science. The center would not be the center that it is in a particular field of intellectual activities if the periphery did not regard itself as inferior to it in the knowledge possessed by its incumbents and their works” (Shils 1991, p. 417). The asymmetry between center and periphery lies, then, in the universal validity of scientific truth. “The universal validity of scientific knowledge – the universality of science – is not the same as the universality of scientific activity” (Shils 1991, p. 409). The development of science in different regions is in terms of the traditions in those particular societies, and if consensus on the universality of scientific knowledge and its validity forms the basis for the development of scientific communities, then center and periphery necessarily have a different status. In this model, resistance to the dissemination of knowledge is a result of the irrationality of peripheral communities. According to Patiniotis, center and periphery were conceived by Shils as “structural elements of a society representing the relative density of institutions, authorities and symbols of unity. The center is the location where such dynamic is more concentrated exerting centripetal forces on the periphery. Periphery. . . is the wide area where less active individuals, followers and even disloyal dissidents are dispersed” (Pationiotis 2013, p. 364).

1

According to Raj, Argentinian developmental economist Raúl Prebisch conceived the same model independently (Raj 2010). 2 For other authors, see Nye (1975) and Singer (1982); also see Guillem-Llobat (2008).

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The use of this dichotomy in the 1960s and 1970s in economy, geography, and international policy paved the way for its promotion in the history of science. Many historians, especially those focused on non-Western and colonial science, referred to and often sought to implement the model proposed by historian of technology George Basalla (1967, 1993). Although British biochemist and historian of science Joseph Needham’s concept of the global context of scientific practice was also important in the 1960s, Basalla’s model had more impact. From his knowledge of Chinese science and technology, Needham raised the question of why Chinese science was surpassed by European science. His answer was that while Chinese science was built on local grounds, Western science was ecumenical. He was convinced of the universality of science as a human enterprise and thought that while modern science is uniquely Western in origin, it is culturally universal (Needham 1954; see Raj 2013). In 1967, Basalla announced a three-stage model describing the introduction of modern science into non-European nations.3 The paper “The Spread of Western Science” appeared in the influential journal Science, reflecting the importance of the topic and ensuring a wide readership in post-WWII America. This model, considered a typical product of the Cold War, helped the shaping of American national and foreign science policies and boosted studies that dealt with political and social dimensions (Raj 2010, 2013). As the title of the article announced, for Basalla, modern science, as a product of the Scientific Revolution, took place in a very restricted geographical area comprising Italy, France, England, the Netherlands, Germany, Austria, and the Scandinavian countries during the sixteenth and seventeenth centuries.4 These centers of knowledge production “were in a position to pass on their scientific heritage to a wider world, through military conquest, colonization, imperial influence, commercial and political relations, and missionary activity” (Basalla 1967, p. 611). This assumption reflects the fact that, as Renn has argued (Renn 2015), the history of science has been dominated by the history of Western and particularly European science, the Scientific Revolution of the sixteenth and seventeenth centuries being its paradigm. This revolution supposedly gave rise to modern science, which established a general method comprising of hypotheses and experimentation or observation as a way of testing them. According to this traditional narrative, this revolution began in astronomy and physics and then spread to other disciplines and geographies all over the world.5

3

In this paper the author acknowledged the influence of previous works. Among the most important are Thomas S. Kuhn, The Structure of Scientific Revolutions (1962), and Bernard Cohen, The new world as a source of science for Europe. 4 Although since WWII, the United States and Russia have become part of “modern science” producers. 5 This idea of the Scientific Revolution giving rise to modern European science is no longer held. Science is not the paradigm of universal rationality and is not distinguishable from other forms of cultural practices. Our most basic epistemological referents in which the classic image of science rests (proof, evidence, objectivity, etc.) have been historicized. See Renn (2015). On historical epistemology, see Rehinberger (1997) and Daston (2000). This historiographical and philosophical approach focuses on the local and temporal variation of scientific entities that articulates scientific practices, rather than on the evolution or development of scientific theories around immutable objects.

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For Basalla, there are three phases in the diffusion of science from the metropolis toward the periphery, that is, from Europe to non-European countries. These phases are linear, sequential, progressive, and necessary for transforming science in local or peripheral contexts and adapting it to European models. The first phase corresponds to the expansion of European science, during which scientists from the center of diffusion travel to regions in which there is no scientific tradition and use the local knowledge for their own benefit; this phase is an extension of geographical exploration and of classification and estimation of natural resources. “All the plant, animal, and mineral specimens collected in the foreign lands. . . were returned to Europe. . . for the benefit of its scientists. Phase-1 science may be scattered around the globe, but only nations with a modern scientific culture can fully appreciate, evaluate, and utilize it” (613). It is a process of globalization of science on the basis of polarization and concentration of scientific activity. In the second phase, which refers to the establishment of colonial science, scientific activity is developed by local researchers but is marked by its dependency on the metropolis for the formation and education of new scientists, the publishing of their results, and the acquisition of equipment; in this phase, colonized knowledge can be separated from metropolitan knowledge. For Basalla, the adjective “colonial” is used to denote dependent science based upon institutions and traditions of a nation where scientists are “dependent upon an external scientific culture and yet not a fully participating member of that culture” (613). Colonial scientists, native or settler, trained in European centers, live in asymmetrical conditions and seek to belong to European communities, but at the same time, they cannot become part of the Western science. For Basalla, colonial scientists are the heroes of peripheral science inasmuch as they use the resources of existing science to slowly develop a tradition of their own with the creation of national institutions which will provide an independent scientific culture. Finally, the third phase corresponds to the development of a local independent scientific tradition, characterized by intellectual, institutional, social, and financial independence vis-à-vis metropolitan European science. “The colonial scientist. . . is to be replaced . . . by a scientist whose major ties are within the boundaries of the country in which he works” (617), as he is inspired by a national sentiment. The result of this process is the establishment of independent scientific cultures and communities capable of generating science and technology on the basis of their own efforts. According to this Euro-centered model, which defines science as a Western phenomenon, modern science displaces local forms of knowledge, and not only undervalues local knowledge but also even ignores or supersedes it, considering peripheries as passive receivers of the scientific developments of the centers. Moreover, science is seen as one of the main instruments of central or imperial control over the colonies. This model of scientific diffusion was based on an internalist view of science that emphasizes the internal logic of knowledge as a system of formal propositions, and its essentialist nature being the motor of social and material progress; it also conceived sequential stages as the only possible scientific developmental process. Many studies were performed following this model. For example, MacLeod (1982) studied the institutional relations between Britain and Australia in the nineteenth century

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and considered the development of Australian natural sciences as following sequential phases; the visitors and collectors from Britain were seen as activists that valued the exploration of nature; then, they were in charge of the training of local naturalists, until the latter began to have independent scientific programs from the center (Britain). Following these studies, Inkster (1985), in his historical reconstruction of the nature and development of the scientific enterprise in Australia in the nineteenth century, offered a narrative where the spread of science from the center to the periphery allowed the establishment of local institutions such as the Agricultural and Horticultural Society of New South Wales (1826–1836) and the Australian Society (1830–1836), both to promote the growth and consumption of colonial products; these societies included geologists, botanists, zoologists, mineralogists, and astronomers, who were physically located on the Australian periphery but center trained; and these scientists emphasized the utilitarian, science-policy center-dominated. According to Inkster, it was not until the formation of the Australian Academy of Sciences and the Commonwealth Institute of Science and Industry in the 1880s that independent science began to be developed in Australia. In the case of the diffusion of evolutionary ideas in Latin America, Moreno de los Arcos analyzed the introduction of Darwinism in Mexico with emphasis on the diffusion of evolutionary ideas once they were received in the circles of Mexican intellectuals in the late nineteenth century, putting too much attention to physicians and anthropologists and omitting its impact on society and politics. This narrative, centered on the reception of Darwin’s ideas, exalted the role played by local scientists in their intention to develop a tradition of their own in the country (Moreno de los Arcos 1984; see also Glick 1974; for the Botanical Expeditions in Bogotá, Lima and México, see Glick 1991). Nevertheless, when proposed, this diffusionist model of science offered a new tool for historical analysis that was comparative and transcultural, in which the local element was seen as relevant. It also established a new agenda for historical studies of science that included epistemological and sociological considerations which yielded new interpretations on the nature of science and criticized the idea that the spread of science was not considered to be a historical object of scrutiny. Many studies influenced by the center-periphery dichotomy brought to the fore the tension between global context and local forms of knowing and addressed “the racial and economic discrimination involved in the workings of science in the colonial context and have, above all, convincingly shown that diffusion occludes the active processes of reception and appropriation on the part of receiving groups” (Raj 2013, p. 339). Thus, nationalist narratives tended to see Western science as a “hegemonic master narrative” (sensu Raj), imposed on different non-Western cultures, bringing to light the asymmetries of scientists working on the peripheries; also, colonial historians in their effort to globalize the field paid too much attention to local case studies (to name just a few: Varsasky 1969; Herrera 1971; Leite López 1972; Moreno de los Arcos 1984; Reingold and Rotherberg 1987; Glick 1992; Chambers 1993; Vessuri 1997).6

6

Carla Nappi, for example, thinks that the tension between the local and the global has influenced the practice and content of the history of science itself. For her, history looks different as practiced in different localities, be these institutions or geographies. Thus, decentering Europe in the history of

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These studies on the history of science stressed and validated a local and comparative focus for doing history of science. These studies have led to identifying key elements in the diffusion process and developing more accurate ideas about its complexity. They reflected that either the historian paid more attention to the process of transmission or to the process of reception and adaptation, validating the view that scientific practices are immutable entities produced in the center and received passively in the peripheries (Guillem-Llobat 2008; Patiniotis 2013). In the years that followed the implementation and critique of Basalla’s model, there were other proposals for the spread of modern science. In 1985, historian Lewis Pyenson proposed a different model for the diffusion of science from the metropolis to peripheral countries (Pyenson 1985a, b). Pyenson emphasized the utilization of science in the development of colonial empires than in the construction of national scientific traditions or how the reception and integration of modern science happened locally. He analyzed the transplantation of European scientists to the periphery, their practices, and their methods of scientific inquiry and identified three strategies of scientific expansion: functional, scientific, and economic.7 Although this model did not focus on the local context, it led historians of science to explore new relationships between science and imperialism and had thereby expanded and enriched the practice of the history of science in the last decades of the twentieth century.8 However, both localists/nationalists and imperialists agree on the idea that scientific knowledge is produced in confined spaces and then transferred to the rest of the world (see, e.g., MacLeod 1982; Kumar 1991; Petitjean et al. 1992; MacLeod 2000). A significant number of historical studies that acknowledged the complex interactions generated after contact between imported scientific novelties and local cultural traditions were written during the 1970s and the 1990s (Inkster 1985; Home and Kohlstedt 1991; Petitjean 1992; McClelland 1992; Palladino and Workboys 1993; Vessuri 1994). These studies framed the development of historical studies seeking to account for the development of science particularly in Latin American countries (Sagasti and Guerrero 1974; Stepan 1976; Chambers 1987; Lafuente and Sala Catalá 1989; Glick 1992). Others acknowledged the development of national institutions to give account of broader processes such as nation building or political legitimation (Agostoni 2003; Hochman 2009, 2011). None of these studies, however, acknowledged global or reciprocal connections, nor did they focus on the networks of collaboration that may help explain the construction of knowledge at both the local and the global level.

science will allow the historian to reexamine new categories of analysis (Nappi 2013). For a revision on educational contexts where it is conceived that creative versus expository sciences are divorced, just as peripheral local studies and creative centers, see Bertomeu-Sánchez 2015. 7 See also Petitjean 1992. 8 For a critic overview of this model, see Palladino and Worboys 1993. These authors have called attention to the traditional narratives that underline the unidirectional flow of knowledge from the centers to the peripheries; on the contrary they have acknowledged that imperialism has shaped metropolitan science itself.

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Although the center and periphery perspective in the history of science was instrumental in spurring productive exchanges among historians working in nonWestern science, this dichotomy has been challenged since the 1990s, and recent studies have tried to introduce a different point of view in which the dynamic of scientific practices and the dismantling of imperial projects are subjects of historical scrutiny (e.g., Chambers 1993).

A Different Landscape in the History of Science: The Transnational Change In the decades since the diffusionist model was published, historians of biology have developed a number of alternative approaches. Many historiographical accounts explore the “epistemological active role of the colonies as well as the dynamic interaction between metropolis and colonies in the exchange of scientific knowledge” (Gavroglu et al. 2008, p. 158). Although many historians have agreed that the abandonment of the center-periphery perspective with its reception studies and the diffusionist model does not intend to limit or deny the contributions of local studies, they have acknowledged that works done under that perspective have brought to the fore the intricate relationships of global science with local contexts and between science and sociopolitical factors. Patiniotis (2013), for example, calls for the moving beyond this artificial distinction, and instead of discrediting this bipolar perspective, “show the limitations of the historiography that serves this dichotomy and how new questions can emerge if research follows different trajectories.” In this way, scientific practices are not seen as immutable entities that travel and are adapted uncritically to the periphery but rather treated as cultural phenomena affected by local contexts. The emphasis should be, then, “the means employed by each receiving environment to incorporate the new ideas and practices into its established social, cultural and educational structures” (373). The transnational perspective in the history of science is very recent and can be traced to the 1990s, when globalization and regional integration became more important themes in public affairs, along with typical transnational problems such as climate change, mass migrations, or global epidemics that needed international cooperation and not national oppositions. These global problems required a new form of historiography. In the last decades, we have seen that the history of science has been influenced by the effects of globalization (including the globalization of the history of science itself), multiculturalism, and the formation of circuits of practices, organizations, objects, goods, knowledge, and people, in which scientific developments go beyond nation-state borders, the units of historical analysis being circulation and collaborative networks (formed by shared interests through which exchange is negotiated and in which the circulation of knowledge, people, and practices occurs). This new perspective emphasizes the flows themselves and moves away from simple international boundaries (Pestre 2012).

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Recent studies on the history of science in developing countries have confronted the accepted idea about the coexistence of the global circulation of people, practices, and techniques and the symmetries between scientists working in metropolitan centers and those working in less-developed ones, showing that not all science in the so-called periphery should be regarded as peripheral and criticizing those perspectives that see the diffusion process as unilateral and promoted by metropolitan centers and adapted in the peripheries. Global historians have pointed out the role played by scientists in international networks, not merely as ambivalent recipients of discoveries made in the West and the United States but as generators of scientific knowledge disseminated to the world (Espinosa 2013). The worldwide circulation of knowledge is now considered not just as a one-sided colonial or postcolonial diffusion process, but rather as an exchange in which each side is active and knowledge is shaped and transformed by appropriation (Renn 2015, p. 242). Moreover, some historians have proposed abandoning the terms center and periphery, inasmuch as they do not reflect the dynamics or circulation of the elites from less-developed countries who made outstanding contributions and participated in international networks (Kreimer 2010). Pationiotis for his part asserts that the rejection of this dichotomy could benefit historian’s attempts to give an account of how knowledge gained authority in a local context and not from a privileged standpoint (Pationiotis 2013). Other historians have introduced the term semiperipheral to acknowledge the fact that most countries do not easily fall into the center or the periphery (Bennet 2014). Transnational narratives should provide a clearer picture of the role of science in shaping modern history. The transnational perspective allows historians to engage in inherent transnational processes, such as globalization, regional integration, and industrialization (Van der Vleuten 2008). As some historians working on non-European science and technology have noticed, the traditional writing of the history of science and technology through the lens of European history has adopted a Western perspective in which the key players are just a few leading European countries, leaving aside the connected histories of Europe’s nations (Edgerton 2007).9 The new transnational perspective allows historians to depict not only national contexts but also subnational and international ones, in which “circulation (and control) of people, artefacts, goods, services and natural resources are research sites where Europe and technology were mutually constituted” (Van der Vleuten 2008, p. 976). According to Raj, recent studies on science and technology have paid more attention to the material, social, political, and cognitive aspects of scientific knowledge, aspects that are historically and geographically situated, while moving away from traditional narratives that see science as a system of formal propositions and criticizing the essentialist conception of science. “. . .Contingencies of place have thus come to acquire key importance in recent sociological and historical

9

Edgerton argues that the history of technology is an innovation-centered one that does not focus on use; the history of technology-in-use, rather than the traditional innovation/invention narrative, gives a global history inasmuch as it includes all places that use technology.

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studies of science. . . It has been convincingly shown that scientific propositions, artifacts, and practices are neither innately universal nor forcibly imposed on others. Rather, they are disseminated only through complex processes of accommodation and negotiation, as contingent as those involved in their production (Raj 2013, p. 341).” The actors in these narratives are no longer seen as local or global; they are seen as crossers of territorial disciplines and geopolitical regions. As Turchetti and colleagues have argued “transnational history has thus emerged as a key transition point in the recent past, typified by important changes in the international political landscape. The end of the Cold War and the rise of globalization in economic, cultural and military terms stimulated the search for novel explanatory frameworks. . .. Facing the emergence and consolidation of international organizations, the growing influence of corporate multinationals and the rising tide of environmentalism several historians introduced new methods to explore these issues. . . At the same time, the development of the internet, and electronic access to resources worldwide, created a new working space for the historian and significantly transformed the profession. Indeed, scholars began to use these tools to explore conceptually and geographically distant places, also considering the potential of adopting comparative perspectives or approaches transcending national borders” (Turchetti et al. 2012, p. 321). Nevertheless, these authors have highlighted the absence of the historian of science in this discussion; for them what is needed is that the historian of science takes the role of science in a global dimension seriously, “science being a typical object of study beyond borders” (323). What are the key characteristics of a transnational perspective? This historiography has focused on how knowledge circulates in widely different contexts and collaborative networks and has limited the role of the “nation” as the unit of historical analysis. To avoid the tension between locality and circulation, many works have focused on the problematic nature of circulation, paying attention to the effects locally produced knowledge exerts upon practices in motion and in its interaction with specific localities; in this view, circulation is a process of knowledge production, not only the moving of things. In the case of the nation as the unit of historical inquiry, many scholars have pointed out that decentering the nation can shed some light on the trajectories of things on the move across borders. They also have noted that “national boundaries” have been taken uncritically and without a serious epistemological analysis.

Circulation and Collaborative Networks Recent ideas on circulation can be traced to the first decade of the twenty-first century when historians realized the global situatedness and movement of science and the importance of circulation in knowledge production. Historian James Secord, for example, has agreed with many historians of science on the circulatory nature of knowledge and demonstrated that science can be understood as knowledge in transit, which has to cross national, temporal, and disciplinary borders owing to its social nature (Secord 2004). Even more recently, Diarmid Finnegan proposed four distinct

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types of place in which scientists operate: sites, regions, territories, and boundaries or circulation, the latter being used to understand the dynamics of knowledge that became universal by circulating (Finnegan 2008); for Robert Kohler (2012), these categories embrace the material and the social aspects of place, the local and socially constructed character of scientific knowledge. A few years before, David Livingstone had already drawn attention to the sites where science inquiry takes place, addressing the fact that produced in different locations, scientific knowledge is mobilized for different cultural and scientific purposes; thus, space and place matter. “As it moves it is modified; as it travels it is transformed” (Livingstone 2003, p. 4). Scholars who aspire to write more global accounts of science examining local contexts and their connections in producing science have focused on the circulation of ideas, people, practices, or artifacts to construct larger narratives by looking at the spaces that those circulations describe (Safier 2010; Nappi 2013; Raj 2010, 2013). This new perspective thoroughly emphasizes the interaction of experts from different countries and the transnational circulation of people, knowledge, and practices as an intrinsic part of knowledge production (Birn and Necochea López 2011). Thus, recent debates regarding global and local contexts have called attention to circulation collaborative networks that explore interregional exchanges and transnational circuits that allow quicker cross-border transmission of scientific practices and a faster flow of people, practices, ideas, and artifacts (Van der Vleuten 2008; Sivasundaram 2010; Safier 2010; Birn and Necochea López 2011; Hofmeyr 2013; Druglitrø and Kirk 2014). Sivasundaram, for example, emphasizes that in order for science to be successful, it has to travel, “studying networks fits well with global history because networks cross empires, nations, and regions,” and adds that a global historian of science must make the study of colonial science part of a larger tapestry (Sivasundaram 2010, p. 158). Instead of focusing on laboratories or confined spaces, this new approach is characterized by placing the emphasis on things on the move: scientific practices, artifacts, and people, addressing circulation as a historical category (Safier 2010). Although Secord (2004) stressed circulation as communication, for Raj circulation is more than mere communication, it implies the going and coming of things while transforming themselves. “By science we understand . . . the production of knowledge, practices, instruments, techniques, and services; and by circulation . . . not the “dissemination,” “transmission,” or “communication” of ideas, but the processes of encounter, power and resistance, negotiation, and reconfiguration that occur in crosscultural interaction” (Raj 2013, p. 343). For Raj, circulation is in fact a counterpoint of diffusion, dissemination, or transmission of the bipolar distinction centerperiphery, giving agency to all involved in knowledge production. But, as the author argues, there are some conditions for recognizing circulation; these could depend on the “exchange of favors, patronage, friendship, obligation, or just economic exchange to name but a few” (Raj 2013, p. 345). Another type of historiography that had been developing since the last decades of the twentieth century emphasized the role played by the creation of networks in the stabilization of scientific facts and focused on how local knowledge becomes universally accepted; this new focus contradicts the traditional idea that science is locally

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produced and then spread to the outside world; thanks to this scholarship, the primacy of universality over locality has been questioned and reformulated to become an object of historical analysis. Since the late 1980s, the bourgeoning field of history of science and technology has produced many works that have shown the complex interplay between biographical accident, tacit knowledge, institution building, and material culture in the production of knowledge. Important works that show the techniques by which scientists convinced their peers about their knowledge claims highlight the agency of local actors in the flow of knowledge (the literature in the field is enormous; here are just a few of them: Latour and Woolgar 1979; Rudwick 1984; Collins 1985; Shapin and Schaffer 1985; Latour 1988; Daston and Galison 1992; Shapin 1994; Porter 1995; Golinsky 1998; Daston and Galison 2007). This scholarship has been very important for the transnational perspective inasmuch as it has questioned the traditional narrative of the universality of knowledge and has drawn attention to the importance of circulation in the production of knowledge The idea of circulation allows us to elaborate on the journeys involved in knowledge production, in practices that are in search of reproduction and verification (Santesmases and Gradmann 2011). The term also allows us to study the paths through which scientific practices are introduced to and construct national scientific policies and “to reflect on the details of the travels, to pose and eventually answer questions regarding how knowledge and practices travel through geographical spaces and through time. The movements, shifts and travels are not taken for granted, but rather are included in the account as an agent: circulation is itself history” (Santesmases 2012). Since the beginning of the transnational turn, there have been historiographical initiatives that have stressed the need to produce new narratives constructed upon fresh ideas that highlight circulation and collaborative networks as key elements for historical analysis. For example, a group of scholars formed an international research group in the 1990s “Science and Technology in the European Periphery” (STEP) whose aim was to properly place supposedly marginal and provincial case studies into the mainstream of international historiography, that is, how to address the tension between particular local practices and the trends of globalization. Taking center and peripheries as flexible and dynamic categories, this group embarked on the idea that the study of circulation of knowledge may contribute to a new multicultural approach to “a truly European history of science in the future” (Agustí NietoGalán 2015, p. 69; see also Gavroglu et al. 2008). Detailed historical reconstruction of several journeys from the eighteenth century in countries such as Portugal, Spain, Greece, Turkey, Russia, Hungary, and Scandinavia showed that scientific and technological traveling became an analytical category for the history of science, contributing to the clarification of the processes involved in the appropriation of scientific practices, instruments, and ideas (Simões et al. 2003; see also Papanelopoulou and Kjaergaard 2009; Papanelopoulou et al. 2009). Hetch (2011) and Creager (2013) have studied the trading of radioisotopes and other materials that are controlled transnationally, calling attention to circulation as a transnational phenomenon. In this way “exploring the transnational circulation of knowledge thus becomes a key feature in the analysis of how hazardous trades have been reinterpreted, negotiated and relocated in undeveloped and less-developed countries” (Turchetti, p. 328).

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Decentering the Nation, Crossing Borders and the Transnational as a Unit of Historical Analysis The field of science and technology studies has recently focused on the need to write connected transnational narratives as opposed to comparative ones, which shed light on local resistances and global trends (Subrahmanyam 1997, 2005). The notion of “connected histories” as deployed in Subrahmanyam’s 1997 paper and in his two-volume work on the history of early modern Eurasia from the sixteenth to the eighteenth centuries, Explorations in Connected History, was intended to shed light on the intertwined balance of power and on the system of international political alliances between the Mughals in India and the Europeans. The displacement from a strictly local history to a connected one is not a simple shift of perspective; it meant “rescuing history from the nation, not only by bringing to the fore the local and the regional, or by scaling down as a form of bifurcation, but moving laterally. . . This lateral movement is not only an englobing one, but also one that stresses a certain sort of interaction. . . namely that of acculturation. . . that might take us then to a more complex topology, where the community and its history are not simply to be opposed to the . . .national history but. . . might transcend and encompass various national or regional histories” (Subrahmanyam 2005, pp. 11–14). For example, scholarly investigations of the history, biology, and medicine in Europe, Africa, and Latin America have moved beyond considerations of one-sided colonizer versus colonized encounters, or from the celebratory triumphs of the West, to the use of a broader analytical perspective of the global interaction to foster geographical and academic boundaries (for Europe see Santesmases 2006; for Africa see Parle and Noble 2014; for Latin America see Birn and Necochea López 2011, and Espinosa 2013). This transnational approach abandons the nation as a unit of analysis in order to understand the development of science history. It also abandons Euro-US-centered narratives in order to explain the role of international networks and the circulation of knowledge, people, artifacts, and scientific practices. This new perspective, according to Turchetti and colleagues, “could promote a novel understanding of science as historical phenomenon” (Turchetti et al. 2012). Recent historiographical studies have shown the complexity, and in many cases overlap, of the terms “national,” “international,” or “global” and “transnational.” According to Cueto and colleagues, “national” is understood to be that which constrains the nation as a unit, “global” is that which transcends nations, and “transnational” is that which transcends the nation without implying a global scope (Brown et al. 2006; Turchetti et al. 2012). Other authors mention that the term “transnational” has been vaguely defined in an attempt to describe the cross-border flow of people, goods, ideas, and processes. Van der Vleuten has argued that, since the popularization of the term in the 1990s, transnational history has been characterized as a fluid, broad term packed with contradictory meanings. He identifies at least three different, and sometimes overlapping, meanings; one, transnational history refers to the study of cross-border flows, in which circulation, connection, and relationship are the concepts of analysis. This perspective “may suit scholars working on globalization or regional integration better than those re-examining national or local history from transnational perspective” (Van der Vleuten 2008, p. 979). The second refers to the study of the role played by

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international nongovernmental or intergovernmental organizations that criticizes the nation-centric view of international affairs. The third meaning reflects an alternative to nation-state narratives in which the nation is conceived as the principal organizing category of historical analysis. These meanings offer a new perspective to historians that see the nation as a straitjacket (for further argument see Van der Vleuten 2008). Decentering the nation-state histories appeared as a response to comparative histories that reify the national narratives. Nevertheless, consensus has not been reached on this issue. For example, some historians have spoken on the need to abandon the nation as the unit of analysis in recognizing the importance of the local and the study of science where it is being produced (Pyenson 2002). Walker (2012) distinguishes national science from science in a nation and argues that seeing science as a “national activity” is not only a recurrent attitude but also an inevitable one. For other authors, abandoning the nation altogether could be a wrong decision due to the importance of nation-states in shaping modern history. “Advocates of the new transnational history did not advocate giving up that analytical category, but rather placing it in its proper historical context” but not abandon it (Van der Vleuten 2008, p. 983). Other historians have called our attention to the term “global” as a historical object, as a descriptor of time and space, something more than simple diversity in language, method, or approach. To uncritically use the term global to describe largescale phenomena can pose epistemological problems (Nappi 2013). Despite these problems, the agenda of the transnational perspective has inspired an impressive amount of scholarship. In the case of historical studies of biology and the life sciences, a lot of research performed under this approach has indicated that despite their historiographical and epistemological importance, narratives on the national science perspective have revealed its analytical limitations. This research has expressed the need to reconstruct transnational stories that account for how the knowledge produced in developing countries forms part of international knowledge as it circulates in international networks of collaboration. This perspective has enabled the production of narratives that go beyond the national framework through analysis of transnational participants and processes and has permitted new ways of thinking about science history in national and regional contexts. Some of these historians have insisted on more transnational and global histories that take into account the dynamics of scientific practices. For example, Müller-Wille (2003) describes how, in the 1730s in Europe, Linnaeus’ plant taxonomy was characterized by mobilizing and stabilizing specimens, a process that did not take place exclusively in the centers of knowledge production; it was rather a process in which the work of naturalists was dependent on local institutions, collections, and botanical gardens that were connected through global networks of translation and exchange, which allowed the global circulation of specimens. In the case of circulation of objects, the import of radioisotopes by two Spanish research groups, one in endocrinology and the second in molecular biology in post-WWII, was funded by foreign institutions that were instrumental in the establishment of national laboratories but at the same time were important pieces of radioisotope circulation in Europe; as Santesmases has put it, foreign policies toward scientific cooperation between the United States and Spain shaped the US and British programs of radioisotope distribution (Santesmases 2006). Camprubí, examining the

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circulation of laboratory standards, practices, and engineers in Spain between 1939 and 1959, has found that these agents were instrumental in the making of the Francoist regime while circulating for retooling the Spanish political economy; Spanish engineers mobilized scientific practices and the material culture unique to their specific and historical circumstances always taking into account Spain’s place in international cooperation networks (Camprubí 2014). Mathews (2011) has pointed out the efforts to produce a regime of transparent knowledge in the forests of Mexico in the 1940s, not by official declarations or international scientific projects, but from the texture of encounters between officers and their clients, the foresters, and indigenous people to affect the legitimacy of the state and the credibility of public knowledge. Cueto (2007) and Cueto and Palmer (2015) in their reconstruction of medicine and public health in Latin America have shown that important medical and healing practices, such as the use of quinine against malaria, arose in Latin America where indigenous populations incorporated the properties of chinchona bark into their rituals and then expanded to the Western tradition; in this way, science and medicine must be seen as a product of transnational networks and a creative interplay between “metropolitan” and “peripheral” actors. Cueto’s narrative on the development of Mexican physiology from 1920 to 1960 shows the asymmetries faced by Mexican physiologists when they return to Mexico after having worked at the Walter Cannon laboratory in Harvard University funded by the Rockefeller Foundation; these scientists inscribed their work in an international setting but faced adverse conditions when trying to set up a similar laboratory in Mexico; and although they acquired local influence and international prestige, financial resources remained in the United States (Cueto 2015). Quintero’s account on the American and Colombian naturalist ornithological expeditions in Colombia from 1910 to the 1940s calls attention to the complex but fruitful interaction between nationalist policies and imperial practices in the expansion of US political and scientific influence in Latin America (Quintero 2011). Other scholars have been working on the transnational character of human genetics in the aftermath of WWII, when there was growing interest in the genetic characterization of indigenous populations in many Latin American; these scholarship has shown the interplay of international and national institutions, the development of new laboratory techniques, and the instrumental importance of local actors in the reconfiguration of human genetics in the 1960s (Santos 2002; Santos et al. 2014; Barahona 2016). Jessica Wang (1999) and Krige (2006) have shown that the Americanization of science (which has been never self-sufficient) during the Cold War was a process in which both parties were involved; metropolitan and local scientists played very active roles. Now we have a myriad of case studies looking through the transnational glasses of the recent historiographical change. In Latin America see Birn 2006; Brown et al. 2006; Hurtado de Mendoza and Vara 2007; McCook 2009; Soto Laveaga 2009; Palmer 2010; Medina 2011; Escobar 2012; Podgorny 2013; McCook 2013; Duarte 2013; Barahona 2015; Cueto and Palmer 2015; in Asia see the works of Zuoyue Wang 2010; Subrahmanyan 1997, 2005; Kumar 2012; Raj 2013; Ganeri 2013; Phalkey 2013; for Africa see Hofmeyr 2013; Parle and Noble 2014; and the special issue of Medical History 2014; other works are those of Cañizarez Hesguerra 2001; Creager 2013; Achbari 2015.

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Conclusion The end of the Cold War not only affected the international geopolitics, diplomacy, and world economy but also had important influence on human culture; the important changes in the international political landscape that occurred alongside an unparalleled development of science and technology (that had been going on since the end of WWII, particularly telecommunications – Internet and electronic access) changed the international organization of science itself and its role in the increasingly globalized world. The history of science was not immune to the effects of these processes due to the globalization of many cultural aspects, regional integration, and the rising of international institutions for cooperation to face transnational phenomena. The transnational perspective dates back to the last decade of the twentieth century, when historians of science questioned traditional narratives which emphasized the colonial situatedness of knowledge and assumed, first, the nation as the unit of historical analysis and, second, the immutability of knowledge when moving from one place, usually the center of its production, to another, the dependent periphery. This new perspective has shown that the model proposed by Basalla in 1967, despite having opened a new agenda for the history of science, has shown serious historiographical limitations. As a typical product of the Cold War, the center-periphery bipolar distinction upon which Basalla’s model relied led to debates about the role of the local context and the universality of scientific knowledge to open. Scholarship produced in the 1970s and 1980s focused on colonial science in which dependency to the metropolis and the celebratory triumph of the West were highlighted. This picture was changed when other scholars pointed out the need to escape from the tension between the global and the local and to assume a wider narrative beyond national borders. In this way, historians of science shifted the perspective to a more transnational focus to give a richer account of knowledge on the move. The transnational turn thoroughly emphasizes the interaction of experts from different countries and the transnational circulation of people, knowledge, artifacts, and practices as an intrinsic part of knowledge production. I would like to thank Michael Dietrich who invited me to participate in this project back in March 2014 when I was expending a few days at his home in Amherst; also, to him, Mark Borrello and Oren Harman for putting this volume together; to M.C. Alicia Villela González for expert research assistance; and to David Bevis for his careful copyediting. This investigation has been funded by projects CONACyT CB-2012/178031, UNAM PAPIIT IN403718, and the Bioethics University Program of the UNAM.

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Historiography and Immunology

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Warwick Anderson and Neeraja Sankaran

Contents Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Early Histories of Immunology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . The History of Immunology Enters Philosophical Maturity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . “Denaturalized” Histories of Immunity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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Abstract

In this chapter, we provide an overview of attempts from different disciplines to treat immunology and immunity as objects in the history of science. Despite the science’s immense potential, there is a paucity of broad, synthetic historical scholarship, compared to writings about the philosophy and, indeed, historiography of immunology. Following a reprise of the origins of immunology as a field of investigation, we trace the pathways through which histories of immunology, and their entwined philosophies, have developed and matured over the past century. The recent output of expansive “denaturalized” histories, which has given rise to a variegated and relatively autonomous historical landscape, is surveyed. The chapter concludes with a call for fuller realization of histories of immunology and immunity, with scholarship that is more comparative, connected, and transnational, rather than internally focused on biography, institutional development, and the succession of ideas and techniques.

W. Anderson University of Sydney, Sydney, NSW, Australia e-mail: [email protected] N. Sankaran (*) Independent Scholar, Bangalore, Karnataka, India e-mail: [email protected] © Springer Nature Switzerland AG 2021 M. R. Dietrich et al. (eds.), Handbook of the Historiography of Biology, Historiographies of Science, https://doi.org/10.1007/978-3-319-90119-0_20

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Introduction For a field of inquiry that has generated so little actual historical analysis, immunology continues to excite disproportionate and often disheartening historiographic disputation. During the past 30 years, we have heard (and sometimes delivered) lectures on the “infancy” of the history of immunology, its subordination to disciplinary scheming or philosophical argument, and its tantalizing “internalism” or deplorable lack of “context” (see Judson and Mackay 1992; Söderqvist 1993; Anderson et al. 1994; Söderqvist and Stillwell 1999). Paucity of substantive historical writing is more readily explained than the peculiar hypertrophy of historiographic discussion. The scientific novelty of immunology, which has developed and proliferated mostly since World War II, partly excuses the hesitation, or allergic reaction, of historians who prefer to stretch out their analysis over a longer span. Technical and recondite aspects of immunological arguments have deterred many historians without training in biology or biomedicine. Also, the theoretic ambitions of the science, which claims to arbitrate matters of “self” and organismal individuality, often have proven philosophically diverting, leaving history as mere appendage to the lively contest of ideas. Consequently, rather than add to expostulation of the “how” and “why” of histories of immunology or amplify polemics over what the history of immunology is a history of, we are tempted simply to urge the field’s would-be historians to write some more history. Yet we remain intrigued by the potential – indeed, the “pluripotency” – of immunology and immunity as objects of the history of science (Cambrosio et al. 1994; Sankaran 2012). Moreover, we are fascinated by how a science of memory has managed until recently to resist its historicity. And so, exigently, we venture once more into the historiographic fray. Memoirs and biographies of leading scientists, many of them hoping to secure intellectual hegemony in the immunological field, have rubbed uneasily alongside normative philosophical disquisitions about where the field should be and what it ought to encompass (we will return to these below). In 1994, historians Warwick Anderson, Myles Jackson, and Barbara Gutman Rosenkrantz instead recommended an alternative approach that dismantled the naturalized walls erected around the science of immunology and opened up historical study of the dispersed cultures of “immunity.” They envisaged a narrative synthesis that might draw together immunological knowledge, broadly conceived, whether from the laboratory, the clinic, public health, philosophy, politics, literature, or ordinary experience. Avoiding the constraining categories of biography, institutional history, and narrow philosophical argument, this sort of history would entail registering concept work around immunity generally (see Anderson and Mackay 2014a). Evidently, the authors were reflecting on the postulates of historical epistemologist Georges Canguilhem, who thought that too often the history of science “is done as natural history, because science is identified with scientists and scientists with their civil and academic biographies, or even because science is identified with results and these results with their present pedagogical statement” (Canguilhem 2005 [1975], 203). In contrast, the proper history of a science, he continued, “is related not only to a group of sciences without intrinsic

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coherence but also to non-science, ideology, to political and social practice” (204). In this chapter, we want to chart the development of various forms of the natural history of immunology, before tracing the recent emergence of what Canguilhem might have called unnatural histories of immunity, observing along the way the many impediments to the broad tolerance that such narrative synthesis requires.

Early Histories of Immunology The earliest historical accounts of immunology may be found embedded in histories of bacteriology or microbiology, which is hardly surprising since the origins of the two fields were intricately entwined. Scottish hygienist William Bulloch, author of what is generally regarded as the first “authoritative standard work” (Foster 1970, ix) on the history of bacteriology, found the source of the discipline in the French – or Pasteurian, as derived from the school of Louis Pasteur – sector of bacteriology. Since Bulloch’s book, which grew out of a series of lectures delivered in 1936 at the University of London, ended as early as 1900, he left the nascent specialty clinging to the maternal discipline. The new specialization, according to Bullock, concentrated on the problems of the prevention of infective disease by artificial inoculation, and the processes involved in the recovery from infection [which in turn] led to the creation of that branch of science called Immunology, for although the latter now has implications far beyond the bacteriological sphere, it was in connexion with bacterial diseases that it first took its root. (Bulloch 1938, 255)

The publication record bears out Bulloch’s claims. Surveying articles and journals in the two fields from the mid-nineteenth century to the 1990s, historian Pauline M.H. Mazumdar noted that whereas articles specializing in immunology began to appear as early as the 1880s and 1890s, they were published in journals that featured bacteriological investigations in higher proportions. Well into the 1930s, in fact, the vast majority of immunology papers continued to appear in such journals, despite the establishment by then of three specialist immunology journals. It was not until a full century later that the number of stand-alone immunology journals overtook those of microbiology (Mazumdar 1989, 2–4). Bulloch’s book together with William Ford’s volume on Bacteriology for the Clio Medica series (1939) remained the only available chronicles of immunology for over three decades. Then another British pathologist, William Foster, noting that nothing new had appeared in the interim, proceeded to “bring these histories up to date” with a book that extended until 1938 (Foster 1970, x). The growing importance of immunology in relation to its parent discipline is clearly in evidence in Foster’s book – beginning with the appearance of its name in the title. Furthermore, in contrast to Bulloch’s book – where the single chapter devoted to the “history of doctrines of immunity” was the last of eleven – Foster allocated two entire chapters out of a total of eight to the subject, focusing first on its scientific basis (92–126) and then on its applications in medical practice (127–164).

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Foster’s treatment of immunology, being longer, is naturally more developed than that of either Bulloch or Ford, but all authors were largely in agreement with one another, both about the overall pattern of development of the field and the individuals whom they identified as important contributors. All observed the splitting of what they called “immunity science,” very early in its history, in two directions: a practical line directly arising from Pasteur’s work on immunization and a theoretical vein attempting to explain the mechanisms of immunity. These theories, in turn, led to the development of two distinct branches of immunology. One tendency, originating in the ideas of the Russian zoologist-turned-pathologist Ilya (Elie) Metchnikoff, derived from his observations of the phenomenon of phagocytosis,1 stressed the importance of cells in mediating immunity (Bulloch 1938, 259–260; Foster 1970, 92–100; see also Tauber and Chernyak 1991). No author identified a single counterpart of Metchnikoff to represent the rival, the so-called “humoral” theories of immunity, which attributed the property of immunity to protective substances, antibodies against bacteria, viruses, and sundry foreign material, in the blood and other body fluids (Bulloch 1938, 255–256). Not for a lack of choices, however, is such a figure missing, but rather, there were too many contenders: Paul Ehrlich (Silverstein 2001), Emil von Behring (Linton 2005), Jules Bordet, and Richard Pfeiffer, among many others, all made vital contributions to the early understanding and clinical applications of humoral immunity. Three of these men – von Behring, Bordet, and Ehrlich – received Nobel Prizes in Physiology or Medicine, specifically for their contributions to their branch of immunology that for a time was labeled “immunochemistry.” From the 1960s, memoirs supplemented these textbook digests and internalist accounts of the history of immunology. Both F. Macfarlane Burnet and Peter Medawar, who shared the 1960 Nobel Prize for Physiology or Medicine for their discovery of the mechanisms inducing the immunological tolerance of “self,” wrote accomplished and revealing autobiographies. In Changing Patterns: An Atypical Autobiography (1968), Burnet, a shy and introverted man, offered a survey of the changes in scientific aspects of medicine occurring over the course of his own life, including not just immunology but also virology and microbial ecology (see also Sexton 1991; Sankaran 2010b). In contrast, Medawar’s Memoir of a Thinking Radish (1986) cleverly skirted around his pioneering studies of transplantation immunology, preferring to display instead his consummate erudition, wit, and vivacity. Among the more revealing recent examples of this genre, Baruj Benacerraf’s From Caracas to Stockholm (1998) has described his life and times, placing his work in the genetics of the immune response in transnational contexts. A Venezuelan banker turned Nobel prize-winning immunologist, Benacerraf had a lot of material with which to work. Peter C. Doherty, who shared the 1996 Nobel Prize in Physiology or Medicine for his discovery of how the immune system recognizes foreign antigens in association with the body’s own tissue, or histocompatibility, antigens, told a similarly audacious story

1

Metchnikoff has been variously described as a zoologist, embryologist, immunologist, or medical scientist by different writers. His own self portrait was that of a zoologist who “suddenly became a pathologist” (as quoted in Foster 1970, 92–93).

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in A Beginner’s Guide to Winning the Nobel Prize (2006), which was heavy on scientific adventure and accomplishment, if light on personal detail. Understandably, most of these memoirs and tributes were designed to cement the place of the protagonists in the history of immunological thought and to entrench the boundaries of the discipline around them. Predicting the end of the history of immunology, the completion of inquiries, became a common trope. Burnet, for example, forecast in 1965 that immunology would soon run out of problems to solve. In retirement, he concluded that “most of the discoveries have been made and those that can be made are already discernable” (as quoted in Sexton 1991, 135). In 1967, Niels K. Jerne, who would receive the 1980 Nobel Prize in Physiology or Medicine for his cognitive model of the immune system, observed that “immunology will be completely solved within fifty years from now” (Jerne 1969, 347). Having labored hard in the immunological vineyard, Burnet and Jerne felt they could now “sit back, waiting for the End” (Jerne 1967, 601). In claiming their work as definitive and conclusive, Burnet and Jerne were not only securing their place in history, they were demarcating and validating what counted as immunology, or even as proper scientific research, and foreclosing on other, alternative styles of investigation. Thus, as historian Simon Schaffer (1991, 152) pointed out in his study of discourses on the “end” of physics, the strategic deployment of history can become part of the “theoretical technology of modern science.” Subsequent biographies of immunologists have proven more critical, sometimes even displacing them from their assumed central positions in the field. Thomas Söderqvist’s bold sally into the life of Jerne has proven particularly influential and controversial. In Science as Autobiography: The Troubled Life of Niels Jerne (2003), the Scandinavian historian of biology delved deep into the Danish immunologist’s psychological conflicts and disturbed personality as he attempted to show how existential strife might shape scientific research. Söderqvist believed that Jerne’s immunological research represented a flight from the guilt he experienced in his private life. But as historian Ilana Löwy (2004, 330) observed: “It is not clear how Jerne’s psychological profile explains his specific achievements as an immunologist or, alternatively, how his personality sheds new light on his work on antibodies or his talents as a scientific entrepreneur.” Söderqvist has remained a forceful advocate of biography as a means of understanding scientific creativity and as a way of making the history of technical matters more accessible, but few historians of immunology so far have followed his example.2

The History of Immunology Enters Philosophical Maturity Even when the science of immunology was somewhat inchoate, several of its active practitioners were thinking historically, in an analytically detached mode, about their newfangled enterprise. The Polish serologist turned philosopher of medicine Ludwik 2

See the Journal of the History of Biology 44 (2011) for a special issue on scientific biography in general.

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Fleck, for example, in the 1930s pondered the mysteries of the Wassermann reaction, a novel immunological test for the spirochete of syphilis. Genesis and Development of a Scientific Fact (1979 [1935]) traced the history of the Denkstil, the stylized or conditioned perception, which had rendered what was basically an allergic phenomenon, an indicator of host reactivity, into a test for an invasive microbe. Lost to obscurity until rediscovered and translated into English in the 1970s, Fleck’s treatise, in effect, presented the genealogy of a particular thought style in the emerging discipline, one that emphasized biological individuality and reactivity, rather than chemical specificity (Anderson and Mackay 2014a, 45–46; see also Löwy 1986; Moulin 1986). But as in many subsequent accounts of immunology’s development, the historical narrative was subordinated to philosophical argument, which in this case was advancing the sociology of knowledge, emphasizing the immunological thought collective’s “readiness for directed perception” when discovering and affirming scientific facts (Fleck 1979 [1935], 110). Like Fleck, most later historians of immunology have trained in the field: some quickly swerved into further education in the history of biology and medicine, whereas others waited to the end of their scientific careers before inclining toward historical reflection. A clinician tardily turned historian, Arthur M. Silverstein collected various essays on the history of immunological ideas in the synoptic A History of Immunology (1989), which covered mostly the exhilarating period of discovery between 1880 and 1930. Following Foster, he was particularly committed to differentiating the adherents to cellular (phagocytic) immunity, clustered around Pasteur and Metchnikoff in Paris, and the votaries of humoral (antibody-mediated) immunity, associated with Ehrlich and others in Berlin. Silverstein discerned three successive investigatory traditions or paradigms or thought styles: first, the early involvement with bacteriology, then the preoccupation with immunochemistry until the middle of the twentieth century, and later, the development of a biological mindset. As a clinician, Silverstein also proved attentive to pathogenic aspects of the immune response, especially autoimmunity – where the individual’s defense mechanisms are turned against normal body tissues – but also allergy and anaphylaxis. He described in meticulous detail the Berlin aversion to such dysteleological processes, contrasting this reaction to the peculiar conceptual terrain of early twentieth-century Vienna and Paris that was nurturing these, and other, disturbing notions. Silverstein’s fascination with Ehrlich was evident in his biography of the brilliant immunochemist (2002), which added immeasurably to our knowledge of the technical nitty gritty of research into antibodies and the convolutions of German institutional dynamics. In the second edition of A History of Immunology (2009), Silverstein expatiated on the period after World War II, examining the origins of the clonal selection theory of the humoral response, explanations for the generation of antibody diversity, Jerne’s network theories of the immune system, and so on – though steering clear of much recent molecular biology. Snippets of cultural and biographical context tantalized in the second section, which was misleadingly titled “social history,” for as philosopher of immunology Alfred I. Tauber (2010, 636) put it, “Silverstein’s reflections provide only glimpses of a highly competitive and highly rewarded discipline whose sociology has barely been scratched.”

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Anne-Marie Moulin embraced history of science not long after commencing her career in parasitology and immunology (see, for example, Moulin 1985, 1989). In the French tradition of historical epistemology, she was interested in how language framed scientific inquiry, particularly how metaphors and models such as the “immune system” shaped our understanding of embodiment and individuality. Le dernier langage de la médecine (1991) attempted a narrative synthesis of the history of immunological thought in the twentieth century, defining two distinct periods, the first focusing on the meaning of vaccination and the event of the antigen-antibody interaction, and the second concerning the concept of the immune system (see also Moulin 1989, 1993, 2000; Moulin and Cambrosio 2001). Moulin proposed that Metchnikoff’s idea of phagocytosis, dormant for much of the twentieth century due to the dominance of immunochemists, had prefigured the development of cellular immunology in the 1950s. From the 1940s onward, Burnet’s theories of antibody production and Peter Medawar’s research on tissue graft rejection increasingly implicated cellular processes in the immune response. According to Moulin, systemic thought in cell biology culminated in Jerne’s complex – and rather selfreferential – speculations on immunoregulation. Already Moulin (1986) had adroitly demonstrated the exchange of metaphor and model between the cognitive sciences and immunology during this period. But the binary logic of her book’s narrative was strained, and the argument for an epistemological cleavage in the 1950s proved difficult to maintain. Historian Peter Keating (1993, 611) admired Moulin’s historical ambition but regretted that the book “sometimes reads like an overwrought review article interlarded with potted biographies.” Nonetheless, Le dernier langage de la médecine remains the sentinel narrative history of immunology. The emergence of AIDS in the 1980s, seen as a devastating dysfunction or deficiency of the immune system, stimulated extensive historical and philosophical inquiry toward the end of the century into immunology and the defense of self – though rarely was the impetus explicitly acknowledged (but see Moulin 1991; Martin 1994). In Species and Specificity (1995), Pauline M.H. Mazumdar, previously a historian of human genetics, contrasted the ontological “specificity” of Berliners like Robert Koch and Ehrlich with “unitarian” biologists such as Carl von Nägeli, Max von Gruber, and the influential immunologist Karl Landsteiner, who emphasized physiological continuity and gradation (see also Mazumdar 1975). She saw this distinction as mapping onto the difference between structural chemistry and colloid chemistry. Mazumdar traced in particular the thinking of Landsteiner, following him from Vienna to the Rockefeller Institute in New York City, describing his interest in biological individuality and his efforts to redefine immunological specificity as the expression of graded quantitative affinity, culminating in his major work, Die Spezifizität der Serologischen Reaktionen (1933).3 As Warwick Anderson (1997, 560) put it, Mazumdar chose “to illustrate the power of ideas in biology, their transmission through teacherstudent relationships, and their refinement in dialogue.” The clinical, institutional,

3

Translated into English as The Specificity of Serological Reactions (1936).

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and cultural contexts were drawn faintly, if at all. A few years earlier, Mazumdar (1989) had edited a collection of essays similarly exploring conceptual histories of immunology, many of them engaging with Burnet’s clonal selection theory of antibody formation. Until the end of the twentieth century then, the history of immunology would struggle to escape the confines of biography or the genealogy of big ideas, as revealed in the scientific journals. The metaphysical imperative was felt most strongly in the influential studies of Alfred I. Tauber, an immunologist turned philosopher of science. Even in 1991, with Leon Chernyak, Tauber was championing Metchnikoff as a theorist of organismic integration and individuality, and not simply a discoverer of cellular defense mechanisms. They claimed Metchnikoff argued that evolutionary struggle between different cell lineages dynamically negotiated the organism’s integrity or self-definition (Tauber and Chernyak 1991). Discussions of the immunological constructions of “self” also figured prominently in a collection of essays that Tauber (1991) edited that year. In the foreword, evolutionary biologist Richard C. Lewontin (1991, xvi) suggested that the crucial question is: “How does the cell know the difference between you and me?” Typically, he emphasized the importance of the “dialectic” between self and other. Similarly, Chernyak and Tauber in their contribution criticized Burnet’s clonal selection theory for reducing self to a blank in the immunological repertoire – to that which is tolerated or whatever does not induce a reaction. Rather, they insisted, “the Other as non-self, not as another example of self, not as essentially ‘the same,’ but rather as ‘otherness’ must be an essential constituent of Self” (Chernyak and Tauber 1991, 128). An intellectually probing immunologist, Tauber had been inspired by the philosophical ambitions of Alfred North Whitehead and Burnet. “If science is not to degenerate into a medley of ad-hoc hypotheses,” Whitehead (1925, 24) wrote, “it must become philosophical and must enter upon a thorough criticism of its own formations.” Whereas, Burnet (1965, 17), a close reader of Whitehead, had asserted that “immunology has always seemed to me more a problem in philosophy than a practical science.” Tauber was happy to take Burnet at his word. In The Immune Self (1994, 7, 82, 83), Tauber offered “a careful delineation of the scientist’s theoretical development,” showing how the Australian took “diverse conceptual threads from virology, ecology, genetics, developmental biology and, of course, immunology,” weaving them during the 1940s “into a fecund theory [of self-tolerance] that remains the crucial conceptual foundation of the discipline today.” “Analogous to the apperception of the transcendental ego,” Tauber wrote, “Burnet’s self is based upon a notion of unity—a coherent whole—to account for the inner, fundamentally integrated organism” (291). Tauber felt that Burnet’s innovative “concern for accounting for personal identity provides the link to Metchnikoff” (7), though he failed to adduce much evidence for the bond apart from the perceived homology of ideas. Indeed, he realized, “in tracing the linkage between Metchnikoff and Burnet, I may be accused of over-stretching my thesis” (97). Additionally, Tauber wondered if Burnet’s recourse to “self” might indicate the impact of Freudian psychology on the immunologist (Tauber 1994, 97; see also Tauber and Podolsky 1994). If he had visited the extensive Burnet archive in distant Melbourne, Australia, Tauber might have noticed

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the scientist’s indifference to Metchnikoff and distaste for psychology, as well as his contrasting enthusiasm for Julian Huxley, George Bernard Shaw, Henri Bergson, and Whitehead, among others (Anderson and Mackay 2014b). He would have learned about the local and yet cosmopolitan intellectual world that Burnet actually inhabited. Thus, while Tauber’s studies conveyed the impression of historical research, they did not derive from the conventional methods of the discipline. The Immune Self (1994, 295) concluded with a recommendation that “we abandon attempts to pinpoint and define the self, allowing its definition by expression in its phenomenological address: the subject becomes defined in the process of its encounter with the world.” Tauber worried that in absorbing Burnet’s term “self,” immunology had “borrowed a philosophical term to approximate a language that is inadequate to that task” (295). A few years later, medical historian Scott H. Podolsky joined Tauber in telling the story of Burnet’s resistance to instructional models of antibody formation and his advocacy of cellular processes for recognizing and responding the foreign or nonself (Podolsky and Tauber 1997). According to Burnet, tolerance of self mostly, or most efficiently, was imposed in vertebrates through contact with “self-antigens” during embryonic life, which effectively knocked out some antibody-producing cells. Podolsky and Tauber praised Burnet’s biological insight, his awareness that self-tolerance was more interesting biologically than defense, and his application of Darwinian mechanisms in his theory of how encounters with foreign antigens might select and stimulate preserved clones of antibody producing cells. They went on to examine exhaustively the rise of molecular approaches in immunological research, assessing their influence on theories of immune selfhood. Concerned that “self” was dwindling into a fixed genetic signature, Podolsky and Tauber sought process-oriented and functional, or contextualist, accounts of immunological responsiveness. The “newer view of immune function,” they concluded, “is fundamentally formulated as self-seeking, self-organizing activity, whose structure is decentered from any bounded self” (Podolsky and Tauber 1997, 375). They implied that Burnet was responsible for the cognitive encumbrance of this bounded self, overlooking his immersion in Whitehead’s process philosophy and his propounding of dynamic, ecological thinking in infectious diseases research.4 Thus, Burnet had become a useful straw man in philosophical disquisitions, but this meant that, while accruing some value in philosophy, Tauber’s representation of the immunologist lost credit as history (see also Cohen 2000). Ironically, Tauber has functioned as the expedient historian of immunology for most philosophers and anthropologists seeking insight into the “science of self.” When philosopher Thomas Pradeu (2012, 47), for example, asked, “Why does Burnet introduce the concepts of ‘self’ and ‘nonself’ to immunology, and what exact definition does he give them?”, he turned to Tauber for the answers. Pradeu thus attributed to

Tauber (1999, 459) later observed that Burnet “sought a firm definition of the immune self” (see also Tauber 2004, 2008, 2016). But see Anderson (2004) on Burnet’s ecological vision. In a recent synthesis, Tauber (2017, 228) has attempted to reconcile the two Burnets, concluding redundantly that “immunology deserves much greater attention by philosophers of science.” 4

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Burnet the notion of a genetically determined self, which served to distinguish, misleadingly perhaps, his own “continuity” thesis (189). Similarly, anthropologist A. David Napier (2012, 130, 133) drew on an impoverished and distorted history of immunology when he claimed that Burnet’s clonal selection theory was based on a “culture-bound” assumption of an autonomous and static “self,” a “prior and persistent” identity. Instead, Napier imagined an immune system that engages with difference, rather than defends the body (Napier 2003). Such examples of metaphysical theories dependent on unreliable internalist histories of immunology could be multiplied indefinitely (see Anderson and Mackay 2014a, 144–51).

“Denaturalized” Histories of Immunity Reflecting in 1992 on the recent Ischia conference on the history of immunology, science journalist Horace Freeland Judson and clinical immunologist Ian R. Mackay lamented that historians and scientists seemed prone to talking past one another (459–61). Evidently, there had been some friction on the island in the Bay of Naples, which occasionally flared into “open confrontation between ‘the scientists’ and ‘the historians’” (Cambrosio et al. 1994, 375). According to Judson and Mackay, historians frequently had missed the point of the science, and those scientists trying to take up historical inquiries often were resorting to tedious “hyper-review articles,” rather than attending to “intellectual context, social settings, scientific lineages, styles and influences, failures, missteps and conflicts, the stuff of what scientists do” (460; see also Gallagher et al. 1995). Soon Söderqvist (1993, 565) responded indignantly, complaining that the agenda of the history of immunology at Ischia and elsewhere was “still largely set by practising scientists.” He was disappointed that Silverstein and Moulin in their monographs had merely alternated “between analyses of leading ideas, concepts and experiments, descriptions of institutions and portraits of individual scientists” (565). Instead, he recommended two distinct approaches. A “structural” explanation would “focus on the overall cognitive structure of the disciplinary discourse as it exists independent of the historical actors and their social context” (566). In contrast, a biographical exposition, which he favored, would “take the subjective and idiosyncratic self-understanding of individual immunologists as the point of departure” (566). Later, Söderqvist would shift his ground, charging that “so far there have been no sustained attempts to place the development of immunological thought and practice in the context of twentieth-century science and medicine, not to mention the wider social and political context” (Söderqvist and Stillwell 1999, 215). In the wake of the stormy Ischia meeting, science studies scholar Alberto Cambrosio and colleagues assembled another conference on “Immunology as a Historical Object,” as part of the 1993 series of the Boston Colloquium for the Philosophy of Science (Cambrosio et al. 1994). As editors of the subsequent special issue of the Journal of the History of Biology, they discerned two themes in the contributions: the role of “experimental systems” and technique in immunological practice (Stillwell 1994; Löwy 1994; Silverstein 1994; Keating and Cambrosio 1994) and how foundational models and metaphors might guide or inhibit research

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(Söderqvist 1994; Tauber and Podolsky 1994). The first theme, at least, represented a deviation from scientists or philosophers setting the scene of historical investigation. In the concluding article, addressing historiographic lacunae, Anderson et al. (1994) issued their manifesto for the “unnatural history” of immunity, a plea for historians to inquire beyond conventional, or ostensibly “natural,” delimitations or even typologies of the field. They wanted a more ecumenical or multidimensional history of immunity, one that might encompass the laboratory, the clinic, experiences of immunological disorders, distinctive research schools, national styles of investigation, manufacture of biologics, literary descriptions, social theory, politics, and philosophy. Thus, they hoped for critical and independent historical inquiry into the enormous variety of immunological worlds (see Haraway 1991; Martin 1990, 1994). Arguing against simple filiative models and linear narratives, they demanded dispersive studies that would account, in particular, for the social and material conditions of knowledge production and transmission. In response to Canguilhem’s (2005 [1975], 198) question, “Of what is the history of science a history?,” they were imagining – despite using the term “unnatural” – richly “ecological histories of immunity” (587). They yearned for a diverse “ecology” of immunological knowledge and practice (Rosenberg 1979). Some two decades later, Sankaran (2012) would return to this nagging issue, advocating similarly “pluripotent” histories of immunology. “The history of immunology,” she wrote (2012, 49), “seems akin to . . . pluripotent stem cells; while not yet committed to any single path of development, it is prolific nonetheless, brimming with the potential to grow in diverse directions.” Late in the twentieth century, a few historians of science had begun to chart such materialist and encompassing histories of immunity. Ilana Löwy, who trained in cellular immunology, was connecting the developing science with literature (1988), the clinic (1992, 2008), and the contemporary theories of biological individuality (2003), exploring boundary concepts and federative experimental strategies. Combining historical and ethnographic approaches, her monograph, Between Bench and Bedside (1997), was a pioneering study of the links between the immunology laboratory and clinical research in a French cancer ward. Peter Keating and Alberto Cambrosio specialized in tracking monoclonal antibodies between laboratory and clinic, especially in Biomedical Platforms (2003) which documented in impressive detail the use of this technique in the classification of leukemias and lymphomas, showing the influence of processes of standardization and regulation (see also Cambrosio and Keating 1995; Marks 2015). Having immersed herself in the archives, Sankaran (2008, 2010a, b, 2013) set about placing Burnet and his immunological research in a broader bacteriological and biological context, as well as a plausible institutional and national setting (see also Park 2009). Meanwhile, other historians surveyed the sensitive and reactive borderlands of immunological discourse, focusing on histories of “allergy,” an expansive field with an ambiguous relationship to the scientific discipline. Thus, Mark Jackson (2007) described the various scientific and clinical understandings of allergy since Clemens von Pirquet first coined the term in Vienna in 1906 to signify immunological hyperreactivity. A medical historian, Jackson, used the spectrum of beliefs about allergy – that new disease of civilization – to illuminate anxieties and insecurities about modern life

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(see also Jackson 2003, 2009; Jamieson 2010). As an environmental historian, Gregg Mitman (2007) preferred to regard allergy and asthma as sampling devices that might reveal American social inequality, habitat degradation, and political marginalization. Refreshingly, his attention to the clinical and civic dimensions of allergy and asthma included stories of the experiences of sufferers, not only analysis of the research conducted on them. The boundaries of what counted as immunological history were shifting. In A Body Worth Defending (2009), cultural historian Ed Cohen vividly depicted the passages of metaphor and model between the legal or political domain and the emerging immunological sciences, which shaped – or constituted perhaps – the meaning of modern personal identity. And so, at the beginning of the twenty-first century, we were gaining a clearer view of the historical heterogeneity of concepts of immunity. In Intolerant Bodies (2014a), Anderson and Mackay delineated the conceptual history of autoimmunity, or auto-allergy as it was once called, the excessive reactivity of the immune system against the body’s own constituents (see also Parnes 2003). It was, in a sense, the biography of a mode of pathogenesis, a historical inquiry into the longevity of notions of idiosyncrasy, diathesis, and individual variation in twentieth-century biomedicine. In trying to understand “how autoimmunity became thinkable” (4), they attempted to bridge the research laboratory and the hospital clinic, at the same time as they attended to the experiences of those suffering autoimmune disease, often articulated in literary forms, sometimes embedded in case histories. Additionally, they sought to adduce references to autoimmunity, usually figured as deconstruction of the self, in social theory and metaphysics, but their goal was to historicize philosophical speculation, not to use history to illustrate it. The book, they claimed, was “an attempt to describe the interpretation of disease—whether by scientists, physicians, philosophers or patients—as intellectual practice” (5). Importantly, they endeavored to situate, or contextualize, knowledge production about the reactivity and sensitivity of the modern self, focusing serially on the cultural settings and institutional matrices of Paris, Vienna, London, New York, and Melbourne. Once into the 1980s, however, the narrative became more schematic and less contingent, sometimes resembling one of those scientific review articles that Mackay had warned against. Tauber (2015, 385) generously regarded Intolerant Bodies as “perhaps the best overview of immunity (normal and pathological) available for the general reader,” but he chided the authors for their preoccupation with Burnet and his legacy, reproving them for brushing off Metchnikoff and limiting their philosophic ambition. Despite the book’s flaws, Anderson and Mackay hoped it might serve as a template for more elaborate and circumspect histories of immunity. During the past decade, a spate of histories of recent immunology has washed over the heads of the public. Mostly, these books manifest the desire of senior immunologists to instruct the common reader in the excitement and adventure of scientific discovery, but the density of detail contained in the texts, along with an emphatically teleological trajectory, has meant inevitably that their primary readership consists of medical doctors and neophyte investigators. Indeed, one author of such a work has declared his imagined readers are “research and clinical immunologists as well as students and teachers of immunology” (Nagy 2013, xv). Even when

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seeking to render esoteric knowledge exoteric and publicly accessible, as Fleck (1979 [1935]) would have said, these scientific authors have tended to revert, again, to extended reviews of immunological publications (Rees 2015; see also Brent 1997). The more appealing accounts of seminal events and major breakthroughs in recent immunological research function also as memoir, permitting the reader a sense of immediacy and human interest (Paul 2017). Too abstruse and Delphic to succeed as popular history, these chronologies of immunology will provide future historians of biology with ample material for less hermetic analysis.

Conclusion Conflict and disputation have marked the historiography of immunology, whether between scientists or clinicians and intellectual or cultural historians or between those favoring conceptual history or historical epistemology and those seeking historical cases that neatly illustrate philosophical speculations. Some historians have turned to biography to find an overarching telos, while a few historically inclined scientists have resorted to thinly contextualized literature reviews. Too often, historical research has been harnessed to scientific or institutional or philosophical agenda. The reasons for the narrow and dependent character of so much of the history of immunology remain enigmatic, but no doubt the novelty and extremely technical nature of the science have contributed to conceptual restraint and subordination. When so much effort is invested in attaining competence in the science, historiographic ambitions can become subsidiary – even as the desire for philosophical probing abideth forever, so it seems. Such historical reticence or timidity might pertain to other fields of biological inquiry, such as the neurosciences, but seldom has it been as severe or constraining as in immunology. Until recently, there was reluctance to engage with the social and cultural histories of immunity or to look beyond the rigid boundaries of disciplinary expertise. In the past decade or so, however, we have discerned several attempts to write richly situated and deeply contextual histories of immunity, rendering possible the emergence of more encompassing and resourceful narratives. These studies have revealed the scattered and fragmented geography of immunological knowledge and practice, showing us multiple heterogeneous sites of scientific work and influence, or “biomedical platforms,” many of them located far from conventional North Atlantic hotspots. Thus Melbourne, Singapore, Tokyo, and many other places have joined Paris, Berlin, Vienna, London, and New York as locales for immunological knowledge production. Our histories are gradually becoming as cosmopolitan as the science. Progress along this path so far has been tentative, and such densely situated and contextualized narrative histories of immunology or immunity remain rare. Nonetheless, there are, as we have suggested, some promising signs that the historiography of the investigatory and clinical enterprise is becoming less esoteric and less auxiliary to other modes of inquiry. One might say the history of immunology is moving away, slowly, from typological and subsidiary styles toward more entangled, situated, and connected – rather than derivative – ecological explanations.

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As the history of immunity belatedly gets lodged in particular cultures, institutions, and social structures, additional challenges arise. In emphasizing the multiplicity and dispersion of knowledges of immunity, we run the risk of writing a profusion of fragmented, isolated stories, which overstate the disunity of science. When scientists and philosophers dominated the history of immunology, this was never a problem: instead, they assumed the singularity of science and imagined the simple diffusion, or laminar flow, of scientific ideas around the globe. Now, we need more comparative and connected histories of immunity, critical studies that help us to understand how immunological knowledge and practice travel, how they are transacted, adapted, and contested at various sites, on different terrains (Anderson 2018). That is, we should be wondering what the transnational or postcolonial history of immunology might yet be. Acknowledgments Warwick Anderson is grateful for research support from the Australian Research Council (DP120100861).

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Sexton C (1991) The seeds of time: the life of Sir Macfarlane Burnet. Oxford University Press, New York Silverstein AM (1989) A history of immunology. Academic, San Diego Silverstein AM (1994) The heuristic value of experimental systems: the case of immune hemolysis. J Hist Biol 27:437–447 Silverstein AM (2001) Paul Ehrlich’s receptor immunology: the magnificent obsession. Academic, San Diego Silverstein AM (2009) A history of immunology, 2nd edn. Elsevier, Amsterdam Söderqvist T (1993) How to write the recent history of immunology – is the time really ripe for a narrative synthesis? Immunol Today 14:565–568 Söderqvist T (1994) Darwinian overtones: Niels K. Jerne and the origin of the selection theory of antibody formation. J Hist Biol 27:481–529 Söderqvist T (2003) Science as autobiography: the troubled life of Niels Jerne (trans: David Mel Paul). Yale University Press, New Haven Söderqvist T, Stillwell C (1999) Essay review: the historiography of immunology is still in its infancy. J Hist Biol 32:205–215 Stillwell CR (1994) Thymectomy as an experimental system in immunology. J Hist Biol 27:379–401 Tauber AI (ed) (1991) Organism and the origins of self. Kluwer, Dordrecht Tauber AI (1994) The immune self: theory or metaphor. Cambridge University Press, New York Tauber AI (1999) The elusive immune self: a case of category errors. Perspect Biol Med 42:459–474 Tauber AI (2004) Immunology and the enigma of selfhood. In: Wise MN (ed) Growing explanations: historical perspectives on recent science. Duke University Press, Durham, pp 201–221 Tauber AI (2008) The immune system and its ecology. Philos Sci 75:224–245 Tauber AI (2010) Review of A history of immunology, 2nd ed., by Arthur Silverstein. Isis 101:636–637 Tauber AI (2015) Immunology seen through the dark glass of autoimmunity. Metascience 24(3):385–391 Tauber AI (2016) Immunity in context: science and society in dialogue. Theoria 31:143–158 Tauber AI (2017) Immunity: the evolution of an idea. Oxford University Press, New York Tauber AI, Chernyak L (1991) Metchnikoff and the origins of immunology: from metaphor to theory. Oxford University Press, New York Tauber AI, Podolsky SH (1994) Frank Macfarlane Burnet and the immune self. J Hist Biol 27:531–573 Whitehead AN (1925) Science and the modern world. Macmillan, New York

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Contents Introduction: Question of Scope . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . The Varieties of Localization . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cortical Localization . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Specialization of Function in the Nerves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . The History of the Reflex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Canguilhem on the Reflex Concept . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . The Reflex Theory of the Brain and the Debate over Free Will . . . . . . . . . . . . . . . . . . . . . . . . . . . . The Origins of Computational Neuroscience . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Some Points of Contact with the History of Related Disciplines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Evolutionary Theory and the Brain . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . The Cell Theory and the Neuron Doctrine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Medicine and the Nervous System . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neurosciences, Mind, and Society . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . The Brain and Discrimination . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Technology In and Around the Neurosciences . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . The Brain as the Self . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Future Directions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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Abstract

This chapter includes within its purview the historiography of scientific and medical work that long predates the formation of the discipline now known as neuroscience. The unification of biological, computational, and medical sub-specialties that take as their subject matter the brain and nervous system of humans and other animals is very much a twentieth- century phenomenon (see, e.g., Smith (2000) and Casper (2014a)). But like psychology, neuroscience M. Chirimuuta (*) Department of Philosophy, University of Edinburgh, Edinburgh, UK e-mail: [email protected] © Springer Nature Switzerland AG 2021 M. R. Dietrich et al. (eds.), Handbook of the Historiography of Biology, Historiographies of Science, https://doi.org/10.1007/978-3-319-90119-0_23

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as we know it today has a long past, and, as I hope to show, there is a benefit to seeing how the history of the “neurosciences” is a strand in a larger story of the development of ideas and practices relating to mind and life within western European natural philosophy, at least from the seventeenth century onward.

Introduction: Question of Scope The topic that has received by far the most attention from historians is the issue of localization of function within the brain and nervous system. Indeed, many histories of neuroscience define their project just as the recounting of discoveries of the relationship between localized neural structures and particular functions and dysfunctions, from ancient times to the present. In the next section, I offer a brief and by no means exhaustive review of scholarship on localization. As I see it, an important task for historians of the neurosciences is now to forge connections between this body of work and the history of a range of other topics, including the concept of reflex, and the more general characterization of mechanistic approaches to the nervous system (section “The History of the Reflex”), as well as the synthesis of neurobiology and computational theory that occurred in the mid-twentieth century (section “The Origins of Computational Neuroscience”). In section “Some Points of Contact with the History of Related Disciplines,” I discuss work that makes evident the links with the historiography of other domains of biology such as evolution and the cell theory. I close in section “Neurosciences, Mind and Society” with a tour of studies that link the trajectory of brain science with trends in wider society, including technological innovations, scientific racism, and materialist conceptions of the self.

The Varieties of Localization The conception of localization of function has, historically, taken many forms. While neuroscientists today often now associate localization with the thesis that cognitive functions are supported by small, isolatable regions of the cerebral cortex in humans and other mammals, historically important treatments of localization concerned instead the contrasting functional roles of peripheral nerves or the gross functional difference between neocortex and the midbrain and brainstem structures. This illustrates how theories of localization were themselves dependent on the available conceptions of neuroanatomy: to the extent that localization presumes there are fixed structure-function relationships, the way that the brain and nervous system are parceled into structures will shape the possibilities for association with functions. For example, the physician Thomas Willis (1621–1675) is credited with many early discoveries in neuroanatomy and did not hesitate to assign different functions to the macroscopically observable structures of the brain. As Arikha (2006: 159) relates, Willis assigned, “the common sense to the corpus striatum, the imagination to the

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corpus callosum and memory to the grey matter.” One should not ignore the way, also, that localization theories tend to borrow the taxonomy of functions from available theories of mental operations. Willis’ tripartite division into faculties for imagination, memory, and common sense is reminiscent of the medieval scheme of faculties. Below, we will see how combinations of neuroanatomical and psychological1 schemata generate various kinds of localization theories. Another important dimension in the characterization of localization theories is in terms of what the localization thesis is put forward in antithesis to, for instance, against the conception of a brain consisting of functional networks rather than discrete “organs,” or in contrast to the notion that brain tissue is quite functionally homogeneous. This last option is sometimes called “equipotentiality,” but this is different from the definition given by Karl Lashley, the neurophysiologist and psychologist with whom the term is most associated: The term ‘equipotentiality’ I have used to designate the apparent capacity of any intact part of a functional area to carry out, with or without reduction in efficiency, the functions which are lost by destruction of the whole. This capacity varies from one area to another and with the character of the functions involved. It probably holds only for the association areas and for functions more complex than simple sensitivity or motor co-ordination. (Lashley 1929: 25)

The idea here is that brain tissue is not forever fixed in its operations but can compensate for the damage or loss of parts within a functional area, and it is interesting that this possibility of plasticity and compensation is in fact granted by the localizers of the nineteenth century, such as David Ferrier and Eduard Hitzig.2 Another point of reference in the background of discussions of localization is the changing conception of the relationship between the two hemispheres of the cerebrum, especially during the nineteenth century, whether asymmetrical or perfectly similar, as required by some metaphysical theories of the unity of the mind (Harrington 1987).

Cortical Localization The controversial work of the phrenologists, in particular Franz Gall (1758–1828) and Johann Spurzheim (1776–1832), has received much scrutiny from historians. Indeed, phrenology tends to dominate the conception of what localization is essentially about, such that critics of contemporary research on cortical localization and brain imaging cast the project as a “new phrenology” (Uttal 2001). Historian Laurent Clauzade (2018) relates the history of criticism of phrenology as a “false science,” and the polemical use of the history of the discipline is facilitated by phrenology’s having been taken up as a bogeyman among philosophers of science engaged in 1

Using the term anachronistically to describe any study of mental function I thank JP Gamboa for this observation.

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arguments about the demarcation of science from pseudo-science (e.g., Thornton 2018). The movement has received more nuanced treatments from historians of science concerned to understand the relationship between the phrenologists’ detailed anatomical pursuits and their aims as social reformers (Shapin 1979) or personal ambitions (Wyhe 2004), while other works have sought to assess the role of phrenology as a precursor to contemporary neuroscientific localization theories (Young 1970: chap. 1; Métraux 2018).3 Métraux’s work is also notable for its “close looking” at the visual productions of the phrenologists: drawings of skulls as well as the iconic phrenological heads. A number of studies pursue the theme of localization from the time of the phrenologists into the twentieth century, observing the various pendulum swings back and forth between conceptions of brain areas as more or less specialized, and noting the different ideas of what exactly is localized, be it the psychological faculties of Gall, or the simple sensations and muscle representations that were posited later in the nineteenth century (Tizard 1959; Hecaen and Lanteri-Laura 1977). One much-referenced work, which covers the period from Gall to Ferrier’s publications of the 1870s, is Mind, Brain and Adaptation in the Nineteenth Century by Young (1970). An important feature of this work, along with Smith (1973) and Danziger (1982), is the attention paid to the role of associationist psychology in Britain in shaping the course of neurophysiology and neurology in the anglophone world. Many of the figures associated with the genesis and unfolding of the research on cortical localization that occurred during the 1860s–1880s have been the target of biographies which offer narratives of their well-known observations and discoveries, such as Critchley and Critchley (1998) on John Hughlings Jackson, alongside Stanley Finger (2000) on Paul Broca, David Ferrier, and Eduard Hitzig. A different approach has been to focus on a particular phenomenon or pathology and compare how views on its neural substrate evolved within different contexts and research traditions, such as Greenblatt (1970) and Lorch (2008) on the “encounter” between John Hughlings Jackson and Paul Broca, regarding the interpretation of aphasia. It is also worth mentioning the existence of writings by the subsequent generation of neurologists and neurophysiologists in which favored interpretations of the localization theory are expounded and defended. Two examples are Ottfried Foerster (1936) and Sir Francis Walshe (1961) on the question of whether or not Hughlings Jackson was a “strict localizer.” When analyzing the late nineteenth and early twentieth century, there is an important question about how best to characterize and assess the motivations of the opponents of localization, who can no longer be characterized as anti-materialists and defenders of the status quo, as is the norm in accounts of the debates over phrenology. Rather, disagreements seemed to have occurred over the overall plan of functional organization of the brain and how it supports cognitive capacities. The most satisfying work on the localization debate in

See section “The Brain and Discrimination” on the relationship between phrenology, scientific racism, and the justification of colonialism in the nineteenth century.

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this era stands out for its treatment of these points of difference and the connections drawn across relevant views in psychology and biology. For instance, Anne Harrington (1996) analyses the “holistic” theories of the nervous system function put forward by neurologists Constantin von Monakow and Kurt Goldstein in the context of a broader opposition both to mechanistic physiology and associationist psychology, which had significant political and social connotations in Weimar Germany. In another rich study, Katja Guenther (2015) contrasts the localizationist Zentrenlehre of Carl Wernicke and Theodor Meynert in the 1860s and 1870s with the “connective” approach championed subsequently by Foerster and which, Guenther argues, is an important part of the background to Freud’s psychoanalysis.

Specialization of Function in the Nerves In contrast to the substantial literature on cortical localization, the early findings on functional specialization of the nerves have received less attention from historians. That said, the controversy during the 1820s between Sir Charles Bell and François Magendie over the priority of the discovery of the roots of sensory and motor nerves was examined in detail by Clarke and Jacyna (1987). In addition, a recent monograph on Bell by Carin Berkowitz (2015) adds important contextual information regarding institutionalization of medical training in early nineteenth-century London and the contrasting disciplinary cultures of anatomy (Bell) and physiology (Magendie). Edwin Boring (1950: chap. 5) credits Johannes Müller not as a discoverer of the functions of the nerves but as a theoretical systematizer of the recent findings, as he presented them in the “laws of specific nerve energies” in the 1838 edition of his Handbuch der Physiologie des Menschen. The reception and impact of the “laws,” and Müller’s influence on the course of the neurosciences more widely, is a fascinating issue that still calls for further study. Isaac (2019) argues that the laws of specific nerve energies were profoundly influential in the development of philosophical structuralism, via Helmholtz and later neo-Kantians (see also Patton 2018; Hatfield 1990). In Müller’s Lab, Otis (2007) analyzes Müller’s institutional role for the biological sciences in mid-nineteenth century Berlin. The book includes chapters on his mentees, Emil du Bois-Reymond and Hermann von Helmholtz, whose contributions to the physiology of the nervous system have in turn been the subject of numerous studies (see section “Technology In and Around the Neurosciences”).

The History of the Reflex The discovery of the separation of function of motor and sensory nerves is connected with the emergence of the modern conception of the reflex arc – a circuit involving a sensory nerve that is activated in response to an external stimulus and which elicits a predictable, involuntary motor response via activation of a corresponding motor

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nerve. Credit for the experimental results leading to this conception is given primarily to Marshall Hall (1833), with Johannes Müller taking a strong secondary role (Clarke and Jacyna 1987: chap. 4). That said, the concept of the reflex is usually said to long predate the nineteenth-century anatomical and physiological discoveries, with the physiology of René Descartes coming to the foreground as the site of a contested legacy. The case of the reflex is particularly interesting from the historiographical point of view because it is an instance of a historical narrative first constructed by scientific professionals later coming under the critical scrutiny of a historian of science, Georges Canguilhem. I will first discuss Canguilhem’s response to the standard narrative that traces the reflex concept back to Descartes, before then discussing recent scholarship on the Victorian debate over free will, which took place in the shadow of the ascendant reflex theory of the brain.

Canguilhem on the Reflex Concept Canguilhem’s La Formation du Concept de Réflexe aux XVIIe et XVIIIe Siècles (1955/2015) was one of the first monographs on the history of the neurosciences written by a historian and philosopher rather than a scientist. It has not yet received a complete translation into English,4 though it has attracted interest for its historiographical approach – the tracing of the formation of a particular concept – which is discussed at length by Schmidgen (2014b). Canguilhem’s work is a response to the book Reflex Action by Franklin Fearing (1930), an American professor of psychology, whose work is also discussed by Roger Smith (1992: 20), noting the historiographical legacy of Conrad Eckhard (1881). Canguilhem argues that the familiar narrative presented by Fearing systematically overlooks the contributions of research in the “vitalist” tradition concerning the reflex neuromuscular system and overstates the relevance of “mechanists” such as Descartes. To put it crudely, Canguilhem’s accusation is that a Whiggish impulse of later mechanists, such as Emil du BoisReymond, to find their own philosophical presumptions reflected in the history of their discipline is what led to this distortion. See in particular Canguilhem (1955/2015: 139–42), where the argument is based largely on the text of du BoisReymond’s (1887) memorial lecture for Johannes Müller. One of Canguilhem’s positive claims is that the seventeenth-century figure who can be credited with first formulating the reflex concept is Thomas Willis and, moreover, that the “iatrochemistry” that provided his theoretical outlook is continuous with the tradition exemplified in the eighteenth century by J. A. Haller and Georg Procháska, who also made important experimental and conceptual contributions, while departing from the Cartesian tenet that the inner workings of organisms are fundamentally no different from those of inorganic machines. Importantly, Canguilhem’s claims regarding the reflex are bound up with the historiographical view, expressed in his well-known essay, “Aspects of Vitalism” (Canguilhem 1965/2008a), that the history of the biological sciences is characterized

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by a sustained oscillation between what he names mechanist and vitalist tendencies. This stands in contrast with a narrative, popularized in the late nineteenth century by such figures as Thomas Henry Huxley in his public lectures (e.g., Huxley 1875), and still put forward by some philosophers of science (e.g., Craver and Tabery 2017), that the biological sciences since the seventeenth century have been on a linear trajectory in which the mechanistic conception of life is ever more dominant. Thus, the specific claims about the development of the reflex concept stand or fall with these broad commitments regarding the long arc of the history of biology. One might be wary of Canghuilhem’s advocacy of vitalism. However, it should be noted that he does not define “vitalism” as an ontological commitment to vital forces but as a kind of “positivism” in which biological phenomena, such as the “irritability” of nervous tissue, are taken at face value, not as demanding metaphysical exposition or reductive explanation in physical or chemical terms (Canguilhem 1955/2015: 113). That said, assessment of the thesis about the broad shape of the history of biology is beyond the scope of this chapter. I will mention, in passing, that one contemporary historian of science, Jessica Riskin (2016), does depict a comparable oscillatory history, which swings, in her framework, between “active” and “passive” versions of mechanism – the claim that the mechanisms of living systems do or do not have an intrinsic agency. In addition, a number of historians have taken the generational shift from the era of Johannes Müller’s Naturphilosophie-influenced research on neurophysiology to that of his students Hermann von Helmholtz and du Bois-Reymond, who stood self-consciously for an approach to physiology which aimed at assimilation with the physical sciences, to be of great significance for the history of the discipline (Otis 2007; Harrington 1996: chap. 1). Another historiographical perspective considers whether categories such as mechanist and vitalist should be reconceived as stances regarding the reducibility or autonomy of biology, as argued by Ernst Cassirer (who was Kurt Goldstein’s cousin) in his survey of the then recent history of biology (1950: part 2, chap. 11), or if such categories are too broad to be anything but misleading when applied to the detailed texture of particular schools and laboratories. The role of Charles Scott Sherrington in the story of the reflex concept merits much scrutiny. For instance, Casper (2014a) situates the notion of “integration” within the story of the formation of neuroscience as an interdisciplinary field during the twentieth century, while Fearing (1930) presents Sherrington’s Integrative Action of the Nervous System (first edition published in 1906) as the culmination point of experimental research on the reflex, elaborated into a comprehensive theory of the operation of the nervous system. It is worth noting that Sherrington’s use of the reflex, as both an experimental and theoretical device, is one of the main targets of Kurt Goldstein (1938) in his criticisms of the “atomistic” approach to the organism. As Wolfe (2015) points out, Goldstein was a key influence on Canguilhem, with Goldstein’s “holism” being closer to Canguilhem’s “vitalism” than any of the oft-declaimed vital-force theories.

The Reflex Theory of the Brain and the Debate over Free Will Until the mid-twentieth century, the spinal cord and peripheral sensory and motor nerves were vastly more accessible as experimental preparations than the brain itself.

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It should not be surprising, then, that the relatively more developed theory of peripheral function – encapsulated in the notion of the reflex arc – should have been taken as a key to the understanding of central function. This is the move made by the physician Thomas Laycock in his 1845 presentation of the reflex theory of the brain, which was endorsed by Hughlings Jackson in his 1875 pamphlet, “On the Anatomical and Physiological Localisation of Movements in the Brain” (Hughlings Jackson 1931/1985). Laycock (1845: 298) proposed as follows: the ganglia within the cranium being a continuation of the spinal cord, must necessarily be regulated as to their reaction on external agencies by laws identical with those governing the functions of the spinal ganglia and their analogues in lower animals.

The spinally mediated reflexes occur with a fatal inevitability, like the cause and effect chains in a simple machine. For this reason, the hypothesis that all the operations of the brain are essentially reflexes was taken by many to imply that this organ also operates in a deterministic fashion, ruling out free will. The Victorian debate over free will is treated in all its dimensions (including the philosophical and theological ones) in Free Will, an impressive study by Roger Smith (2016). Of particular relevance here is the 1870s controversy over the theory of “conscious automata.” In typically polemical style, Thomas Henry Huxley (1875) presents a case that experienced mental states are never causally efficacious and that the nervous system is an elaborate mechanism of nested reflexes. He appeals both to contemporary observations of the almost-normal behavior of decerebrated frogs and of a brain-damaged soldier undergoing episodes of unconscious “automatism.” As mentioned above, Huxley presents the reflex theory as the culmination of an illustrious history of mechanistic investigations into the nervous system, beginning with Descartes. Huxley’s account was almost immediately contested by the physiologist William Benjamin Carpenter, author of widely read textbooks, who reported that, nothing in the results of more recent researches [was found] to shake my early formed conviction of the existence of a fundamental distinction, not only between the rational actions of sentient beings guided by experience, and the automatic movements of creatures whose whole life is obviously but the working of a mechanism. (Carpenter 1875: 397)

William James published arguments against the automata theory both in (1879) and (1890: chap. 5). In a recent study of the place of religion in Victorian science, Matthew Stanley (2015: chap. 6) contrasts Huxley’s attack on free will with the physicist James Clerk Maxwell’s defense and explains the dispute as part of a wider contest in Britain between the waning “theological science” and the ascendant “naturalistic science,” which was itself a struggle for the control of institutions of scientific education between an establishment in thrall to the Anglican church and a movement of self-made men, often religious nonconformists, such as Huxley. One may contrast this with the case put forward by Chirimuuta (2017) that a shared commitment by physiologists in the 1870s to the doctrine of the causal closure of the physical (a metaphysical presupposition of the automata theory) was motivated, at least in part, by the practical expediency of defining the subject matter of neurology

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and neurophysiology in terms of causal mechanisms subject to experimental interventions.

The Origins of Computational Neuroscience Fearing (1930: vii) ranked the “reflex arc concept” as comparable in stature with the “fundamental explanatory principles of physics and chemistry.” Although the book concludes with the thought that the approach does not “yield a complete account of experience and behaviour” (1930: 315), there is no intimation that the classical reflex theory of the nervous system is about to be replaced, root, and branch, by a new alternative. Indeed, the displacement of the reflex theory turned out to be far more complete than the replacement of classical mechanics by modern physics. Classical mechanics never left the physics curriculum or disappeared from general use in applied science, whereas the reflex theory of the brain no longer appears in contemporary textbooks. The dramatic fall of the reflex theory is a matter that still awaits detailed historical treatment. One important question is how its decline may or may not relate to the rise of computationalism, in the years following the Second World War, as a theoretical framework for much neuroscientific research. Shepherd (2010) discusses the rise of computational theory in neuroscience in the 1950s as part of a larger picture of the coalescence of numerous other specialties to form the multidisciplinary field of neuroscience, as we know it today. It should also be appreciated that there was, in parallel, a surge of informational thinking within molecular biology at this time (Kay 2000). Cybernetics – the interdisciplinary science and engineering of intelligent or selfgoverning systems that emerged during the postwar years – has experienced a wave of historical interest, with recent monographs by Ronald Kline (2015) and Andrew Pickering (2010), and numerous other relevant studies. Since the home discipline of many of the protagonists within cybernetics was neurophysiology or psychiatry, much of this scholarly output is highly relevant to the history of the neurosciences. For example, Tara Abraham (2016) has published the first biography of the medically trained neurophysiologist, Warren McCulloch. The most recent edition of McCulloch’s collection, Embodiments of Mind, includes reflections by neuroscientists Jerome Lettvin and Michael Arbib on the significance of McCulloch’s work, which offer plenty of interesting though often anecdotal material about the formation of the computational theory of the brain (McCulloch 2016). Historical research has tended to focus either on the British or American networks of cybernetic research and the fora for interdisciplinary dialogue that occurred in those countries – e.g., the Macy conferences and MIT labs in the USA and the Ratio Club in the UK. The Ratio Club appears to have been an important conduit for the transmission of information theory and computationalism into British neuroscience, not least because its membership included Alan Turing, Donald MacKay, and Horace Barlow (Barlow and Husbands 2008; Husbands and Holland 2008). Interesting questions remain for comparative research, for example, were the principles and methods of the two “schools” largely shared, and can any divergences be

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accounted for in terms of the different nature of cold war politics in these two contexts? Moreover, it is important to consider activity beyond the Anglophone world, especially the Soviet Union. For example, Gerovitch (2002: chap. 5) argues that through adapting the language of cybernetics to his own purposes, the physiologist Nikolai Bernshtein was able to evade some of the charges of “vitalism” and “idealism” that had been directed to his non-Pavlovian (and hence politically transgressive) research on motor control. The term “cybernetics” is usually reserved for research produced during the 1940s–1960s. However, much of the “artificial intelligence” work of the 1970s and 1980s is continuous with cybernetics, as related in the very comprehensive history of AI by Margaret Boden (2006). Of particular relevance to our study is the development of neural network or connectionist modelling, which began with Frank Rosenblatt’s (1958) “perceptron” model. It is commonly recognized that this braininspired style of AI has coevolved with neuroscience. On this topic, the book Talking Nets (Anderson and Rosenfeld 1998) is a valuable resource – a collection of interviews with seventeen of the early developers of connectionist models, including neuroscientists Walter Freeman, Stephen Grossberg, and Jerome Lettvin,

Some Points of Contact with the History of Related Disciplines This section examines three topics in which the themes elsewhere in the history of biology intersect with research on the history of the neurosciences – evolutionary theory, cell theory, and the history of medicine. This does not, of course, exhaust the possible points of connection but is intended to highlight some areas in which important work has already been done and to give an indication of other avenues for future research.

Evolutionary Theory and the Brain The story of the sciences of brain and mind in the latter half of the nineteenth century cannot be understood without consideration of the contemporary rise of evolutionary biology. The incorporation of Darwin’s theory into accounts of mind and behavior in the Anglophone world is given a detailed treatment by Robert Richards (1987). Alongside Ernst Haeckel, Emil du Bois-Reymond was a major popularizer of Darwinism in the German-speaking world, as discussed by Finkelstein (2013: chap. 11). The incorporation (and alteration) of Darwin’s theory by psychophysicist and philosopher of nature, Gustav Fechner, is discussed by Heidelberger (2004: chap. 7), in an account which illustrates the philosophical richness and diversity of theorizing about the mind, nervous system, and its place in nature, which occurred at this time. Herbert Spencer’s theory of evolution – which, in contrast to Darwin’s, posited an inherent progressive tendency in which life forms evolved from homogeneous to more heterogeneous states and allowed for inheritance of acquired characteristics – had a large impact on the history of neuroscience. The significance of Spencer who, it should be

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noted, had an interest in phrenology at an early stage in his career, is the topic of Young (1970: chaps. 5 and 6). As argued by Feuerwerker et al. (1985), Spencer was a major influence on Hughlings Jackson, and through Jackson, on Charles Sherrington. Evidence for the significance of Spencer is abundant in Jackson’s “Croonian Lectures on the Evolution and Dissolution of the Nervous System” of 1884 (Hughlings Jackson 1932/1985). In this work, Jackson depicts the entire central nervous system in terms of an evolutionary hierarchy of lower to higher structures (the cerebral cortex in man being the most “evolved” structure). The notion of “dissolution” – the reverse process of evolution – is employed to explain a variety of neuropathological conditions. Arguably, this idea became disseminated within the twentieth-century neurology as the notion of “degeneration” of neural structures and pathways. Jackson was not the only one to conceive of neuropathology as a manifestation of dissolution. In the work of John Langdon Down, this had an overtly racial aspect with different kinds of cognitive disability being accounted for as a degeneration to the condition of a “lower” race – the most well-known being the characterization of the condition now known as Down’s syndrome as “the Mongolian type of idiocy” (Down 1866). Down is a particularly striking example, but the racial connotations of the evolutionary picture of the nervous system are at least there implicitly in the writings of other authors of this period (and see section “The Brain and Discrimination”).

The Cell Theory and the Neuron Doctrine Another landmark event in nineteenth-century biology was the development of the cell theory, following the microscopic observations of Henle and Schwann in the 1830s (Otis 2007: chap. 2). Subsequent to the formulation of the cell theory, the “neuron doctrine” is the thesis that the fundamental developmental, anatomical, and physiological units of the nervous system are the single neuron. Shepherd (1991) is a comprehensive study of the development of the neuron doctrine from observations of Jan Purkinje in 1837 to the forging of a consensus in favor of Ramón y Cajal’s version of the neuron doctrine around 1906. One useful feature of this monograph is that it includes, within the main text, long excerpts of primary sources in translation. The dispute between the anatomist Ramón y Cajal and Camilio Golgi (inventor of the staining method employed to great acclaim by Cajal), concerning Golgi’s advocacy of the reticular theory – which posits that the nervous system comprises one continuous network – is of particular interest because it draws attention to the way that a new microscopy technique could generate images subject to multiple interpretations bringing with them incompatible theoretical outlooks. The 1906 Nobel speeches of Golgi and Cajal are interesting documents, not least for their contrasting rhetorical maneuvers (Golgi 1967; Cajal 1967). Because the work of Cajal is so visually rich – he executed very attractive, detailed pen and ink drawings of different kinds of neurons, in various species and at different stages of development – and is valued as an aesthetic as well as a scientific achievement, Cajal has been the focus of scholarship on the topic of

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neuroscience and art. DeFelipe (2010) is one such work, and worth examining also is the catalogue of the 2018 exhibition, “The Beautiful Brain: The Drawings of Santiago Ramón y Cajal” at the Grey Gallery, New York University (Newman et al. 2018). Beyond the focus on Cajal, there are issues to be explored on the topic of neuroanatomy and scientific representation more generally, and understanding how social context, and the communicative ambitions of scientists, affects the choice of representational forms. Some notable works in this area are Carin Berkowitz (2015) on the anatomical drawings of Charles Bell, Poskett (2015) on the publication and reception of Samuel George Morton’s Crania Americana and Arminjon (2009/07/20), and Pogliano (2012) on Penfield’s “homunculus” (drawn by Hortense Cantile) – one of the icons of the twentieth-century neuroscience. The twentieth-century research on cellar neurobiology developed in many different directions, in tandem with discoveries in genetics and the invention of electron microscopy which made possible the detailed imaging of nanoscale structures such as the synapse (Shepherd 2010: Chaps. 2 and 5). The discovery of neural growth factor in the 1940s by Rita Levi-Montalcini forms a fascinating story of experimental science undertaken in the most adverse circumstances. The first stage of research, carried out on the nerves forming within chick embryos, took place in secret after the Mussolini regime banned Jews from holding research positions in Italian universities, with the underground laboratory moving a number of times in order to avoid the Nazi occupation. Subsequent research was carried out at Washington University in St. Louis and had a wide impact on the understanding of diseases not only of the nervous system (Chao et al. 2013).

Medicine and the Nervous System Much of the scientific research that I have included within the scope of the history of the neurosciences was performed by individuals whose primary training was in medicine or surgery and who practiced as doctors. To take some examples, Charles Bell was a surgeon, while Hughlings Jackson was a practicing neurologist and wrote about the different needs of theoretical and medical science, regarding classificatory systems (Chirimuuta 2017); Helmholtz trained originally as a physician but contributed also to physics and psychology (“physiological optics”). For this reason, there is much scholarship on the history of medicine that is relevant to our topic. The specialties of neurology and neurosurgery all have rich literatures concerning their disciplinary formation, within various geographical locations. I will mention a small sample of these.5 The historical writing on the nineteenth-century neurology has tended to be centered on the “flagship” hospitals of that era. The National Hospital for Diseases

5

I have omitted discussion of work on the history of psychiatry and clinical psychology, though much in this literature is relevant to the history of the neurosciences. See, for example, essays in Wallace IV and Gach (2008), and references therein.

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of the Nervous System including Paralysis and Epilepsy at Queen’s Square in London was founded in 1859 and was the workplace of a number of well-known neurologists, including Hughlings Jackson, Charles Brown-Séquard, Henry Head, and Francis Walshe. A monograph by Lekka (2015) brings to light the patients’ experience of the institution, whereas The Neurologists by Casper (2014b) focuses on the forging of the neurological specialist’s identity. On the other side of the Channel, the Salpêtrière Hospital was renowned for Jean-Martin Charcot’s Tuesday lectures in which patients, typically women suffering from “hysteria,” were presented to curious members of Parisian high society. Didi-Huberman (2003) offers a rich account of this phenomenon, with express interest in the photographic iconography that developed around Charcot’s work. An institutional history of neurology and related disciplines in Germany, during the late nineteenth century is provided by Guenther (2015). Moving into the early twentieth century, many of the famous cases published by neurologists were studies of war veterans. Following the Russo-Japanese war of 1904–1905, the ophthalmologist Tatsuji Inouye produced the first retinotopic map of the primary visual cortex, through observations of the visual field defects resulting from bullet wounds to this brain area (Glickstein 2014: 130–2). Geroulanos and Meyers (2016) compare the investigations of Henry Head and Kurt Goldstein in the years following the First World War, with a focus on the condition of aphasia. The case studies of Head and Goldstein were notable for their impact on contemporary philosophers (primary publications include Head (1920) and Goldstein (1942)). Most frequently discussed is the phenomenologist Maurice Merleau Ponty’s notion of the “body schema” [le schema corporel], derived from Head and Holmes (Paterson 2018), and his extensive discussion of Gelb and Goldstein’s case study of the veteran, Schneider (Merleau-Ponty 1945/2004: Pt. 1, Chap. 3). Similarly, Ernst Cassirer (1929/1957) devotes around 100 pages of his Phenomenology of Knowledge to a discussion of the findings of Head, Goldstein, and other neurologists. In case studies such as Gelb and Goldstein (1925), the interpretation of their patients’ symptoms as due to a deficit in the categorical attitudes or behaviors [kategoriales Verhalten] – the ability to understand how concrete objects stand in relation to abstract classes (e.g., of colors or tools) – bears important connections to Cassirer’s “philosophy of symbolic forms” (Métraux 1999; Matherne 2014). The discipline of neurosurgery began to form in the early twentieth century. Greenblatt and Smith (1997) relate the contributions of Harvey Cushing at the start of the century, who was based at Johns Hopkins and later Harvard University. Foerster’s development of surgical remedies for Little’s disease (a disabling pediatric disease) and tabes dorsalis (a condition of tertiary syphilis, affecting the spinal cord) is discussed in detail by Guenther (2015: chap. 4). Foerster was an early collaborator with Wilder Penfield on the surgical removal of cerebral scar tissue as a treatment for epilepsy. Penfield’s contributions to neurosurgery, and to the iconography of neuroscience (as mentioned at the end of section “The Cell Theory and the Neuron Doctrine”), are very much intertwined with the history of the Montreal Neurological Institute. This clinical and research setting are given a detailed depiction in the “biography” of the institute by Feindel and LeBlanc (2016). Another luminary of the MNI is Brenda Milner, whose case study of the postoperative amnesiac H.M. and

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discovery of the dissociation between episodic and procedural memory is often credited as a founding moment for the discipline of cognitive neuroscience (Zatorre 2018). Shepherd (2010: chap. 12) also discusses the work of Milner, and the McGill psychologist, Donald Hebb, whose theory of neuronal plasticity (“Hebbian learning”) has also been foundational to current cognitive neuroscience.

Neurosciences, Mind, and Society In this last section, I offer an overview of research that seeks to understand the history of neuroscientific ideas and practices in terms of wider social trends such as the dissemination of new technologies, the various prejudices against marginalized groups, and the changes in more abstract philosophical views about the nature of human subjectivity.

The Brain and Discrimination S. J. Gould’s (1996) The Mismeasure of Man is still a valuable reference point on this topic as it offers a comprehensive view on how craniometry in the mid-nineteenth century – in particular, the measurement of skull volume – was undertaken in order to establish a biological basis for the supposed inferiority of various groups. Targeted groups included nonwhite peoples, especially the indigenous nations of America, whose skulls were the subject matter of Morton’s Crania Americana of 1839 (Poskett 2015), women, and members of the economic underclass within Europe, deemed to be criminal and “degenerate.” It should not be forgotten that major figures within the history of science, not only Paul Broca but also Georges Cuvier and Francis Galton, are the protagonists here. This dimension of Broca’s research on the brain is more often than not omitted or skirted over in historical accounts elsewhere, which focus primarily on his localization of the speech defect. One finds this tendency, for example, in LaPointe (2014). Finger (2000: chap. 10) does write about Broca’s involvement debates over the relationship between brain size and intelligence but presents Broca in a far more favorable light than does Gould (1996) – where quotations from Broca (1873) speak for themselves. One of Gould’s theses is that Morton’s expectation that there would be a systematic difference in the skull volume across the races led him unconsciously to distort the measurements he made of his collection of human skulls. Lewis et al. (2011) attempted to vindicate Morton by re-measuring half of the skulls from Morton’s original set and by arguing that Gould, not Morton, was guilty of unconscious bias in his statistical analysis of Morton’s findings. Subsequently, Kaplan et al. (2015) have argued that while there were flaws in Gould’s methods, the errors in Morton’s dataset and logic of investigation loom even larger, such that Morton is in no way vindicated. In addition, Weisberg and Paul (2016) have proposed that Lewis and coauthors failed to address the substance of Gould’s claims, invalidating their project entirely. A matter that deserves further discussion is the relationship between phrenology and the biological-determinist argument employed by Samuel Morton and scrutinized by S. J.

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Gould. The appendix to Crania Americana is an essay by the prominent British phrenologist, George Combe, entitled “Phrenological Remarks on the relation between the natural Talents and Dispositions of Nations, and the Developments of their Brains.” Combe (1839: 270-271) argues, against Dugald Stewart, that the different “races” of humans are not uniformly endowed with mental capacities and that climatic and other environmental circumstances cannot account for the variable manifestations of human civilization. As Combe puts it, the hard data of craniometry offer a more satisfactory, biological explanation of the irrepressible disposition of Europeans toward civilization, industry, and conquest, in contrast to the chronic state of barbarity of the nations of America and Africa, and the plateauing level of achievement to be found in Asia: The phrenologist is not satisfied with these common [environmental] theories of national character; he has observed that a particular size and form of brain is the invariable concomitant of particular dispositions and talents, and that this fact holds good in the case of nations as well as of individuals. If this view be correct, a knowledge of the size of the brain, and the proportions of its different parts, in the different varieties of the human race, will be the key to a correct appreciation of the differences in their natural mental endowments, on which external circumstances act only as modifying influences. (Combe 1839: 274)

A corollary of the biological-determinist naturalization of cultural difference is the exclusion of the uncivilized peoples from conceptualization as properly human, and their co-categorization with the flora and fauna of the nonhuman, primeval wild spaces whose territory is retreating with the advance of Europeans. Combe quotes approvingly an unnamed writer for the Edinburgh Review: it now seems certain that the North American Indians, like the bears and wolves, are destined to flee at the approach of civilised man, and to fall before his renovating hand, and disappear from the face of the earth along with those ancient forests which alone afford them sustenance and shelter. (Combe 1839: 272)

Technology In and Around the Neurosciences It is often commented that the theoretical concepts at play in neuroscience are no more than mirrors of whatever technology is currently most advanced and impressive – from the hydraulically operated statues described by Descartes (Riskin 2016: chap. 2), and the telegraphy that proliferated from the 1850s (Otis 2002; Schmidgen 2003; Lenoir 1994) to the digital computers following the Second World War. While computational neuroscientists Daugman (2001) and Eliasmith (2003) consider the history of the obsolescence of such technological analogies in order to advise caution regarding the current computational framework, scholars such as Borck (2012) have examined the matter as a case study in how the production and the use of tools shape human self-conception and the theorization of biological systems (cf. Canguilhem 1965/2008b). Various studies have examined the issue of the transfer of technologies from other domains into neuroscience. This is of particular importance with respect to

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technologies for measuring the electrical activity of peripheral nerves or neurons and forms of imaging that trace neural activity indirectly such as PET, fMRI, and EEG (Borck 2008). Finkelstein (2013) provides detailed accounts of the efforts of du BoisReymond, along with the technicians and artisans whose skills were indispensable to laboratory research, to create a moving coil galvanometer sensitive enough to detect the tiny currents of electricity intrinsic to the nerves of dissected frogs’ legs, thus proving the existence of “animal electricity.” In addition, Dierig (2006) examines the material circumstances within the rapidly industrializing city of Berlin as the backdrop to du Bois-Reymond’s experimental activity. As a result of these observations, du Bois-Reymond is often named the discoverer of the “action potential” (e.g., LópezMuñoz and Alamo 2009), but such terminology was not current in his time. Frank (1994) gives an account of the technological developments coincident with the emergence of the action potential concept and “all-or-none principle” of nervous transmission during the first three decades of the twentieth century. Similarly, with a focus on post WW1 England, Garson (2015) argues that the wartime development of vacuum tube technology was the key innovation behind research in the 1920s in Lord Adrian’s laboratory, which generated visualizations of the electrical activity of nerves. In comparison to Helmholtz (see, e.g., Cahan 1993), however, little has been written on du Bois-Reymond, and none of his works have been translated since his lifetime. The magnum opus Untersuchungen über tierische Elektricität (volumes published between 1848 and 1884) was not translated into English other than a brief abstract. The public lectures are a rich source for studies of the emerging scientific culture in Western Europe (du Bois-Reymond 1912) but have received less scholarly attention than equivalent works by Helmholtz (e.g., Helmholtz 1995), and the majority are untranslated. Regarding Helmholtz and his collaborators, one should note Hennig Schmidgen’s accounts of the development of experimental procedures to measure the time course of nervous activity, looking both at the interdisciplinary context – spanning industrial instrument making, astronomy, physiology, and psychology – in which the relevant instruments were developed, and the cultural impact that such experiments made. Schmidgen (2003) focuses on Wilhelm Wundt, the student of Helmholtz, often referred to as a founding figure of experimental psychology. His account is a response to that of Schaffer (1988) who had argued, against Boring (1961), that astronomers had resolved the matter of the “personal equation” (the individual variability in observation of the time of astronomical events) independently of research by psychologists. Die Helmholtz-Kurven (Schmidgen 2009, 2014a) takes up the subject of Helmholtz’s experiments on the speed of nervous transmission, showing how the concept of “lost time” connects Helmholtz with the novels of Marcel Proust. The connection between physiology and turn of the century aesthetics is a theme also pursued by Robert Brain (2015) in The Pulse of Modernism.

The Brain as the Self The history of brain science is very much entangled with the history of conceptions of the mind. It is therefore not surprising that some of the influential early histories of brain science were written by professors of psychology (e.g., James 1890; Boring 1950). A

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number of social scientists, such as Jessica Pykett (2015),6 have commented on the increasing “neurofication” of discourses regarding human identity, behavior, and governance. In conjunction with the observation of this phenomenon in its current guise, some scholars have attempted to trace the historical path of this trend. An important work here is Homo Cerebralis by Michael Hagner (2000). Hagner’s central historiographical distinction is between the concept of the “organ of the soul” – the post-Cartesian notion (operative up to the end of the eighteenth century) of the brain as the organ by means of which the soul manifests its control of the body – and the brain, in its more familiar guise (from the early nineteenth century onward) as the lump of matter which by itself “secretes thought.” While one is reminded here of Carl Vogt’s infamous saying that, “thought is to the brain what the bile is to the liver, or the urine to the kidneys,” a strand of materialist thought among those discussed by Charles Wolfe (2016: chap. 6), Hagner’s distinction should not simply be taken for a contrast between dualist and materialist theories of the mind; for Hagner argues that the replacement of a unitary self with a mind fragmented into distinct capacities is more significant than the question of the substance of the mind (p. 20). The publication in 1796 of Über das Organ der Seele by Samuel Sömmering is presented as the inflection point between these two conceptions of the cerebrum,7 with Franz Joseph Gall as the first representative and popularizer of the new tradition. Vidal and Ortega (2017: chap. 1) present a “genealogy of the cerebral subject” in which John Locke’s theory of personal identity, presented in the Essay Concerning Human Understanding of 1694 is the founding conception of the human subject which makes possible the later identification of the person with his or her brain. This is an interesting proposal which hopefully will be explored by historians in the near future, though it should be noted that Bassiri (2016) has already argued that the fractured notion of personal identity to be gleaned from Hughlings Jackson’s account of brain pathology is the opposite of the Lockean one. The essays collected in Bates and Bassiri (2016) provide a wide range of angles on the topic of the historical formation of the “neural subject,” with one unifying theme being that if one considers the way that responses to the phenomena of neuropathology and neuroplasticity have shaped conceptual developments, the arc of the story is not simply that of a triumph of reductionism.

Future Directions The historiography of the neurosciences is currently about as diverse in its foci of interest and methods of investigation as the neurosciences, past and present, themselves are. For that reason, it is hard to determine specific trends or norms within the discipline, except to say that patient-centered studies are now well represented alongside the traditional practitioner focused ones and that institutional histories make up a large proportion of the current scholarly output. Many topics remain under explored, and it would be a positive development to see the history of the 6

See also essays in the volume Critical Neuroscience edited by Choudhury and Slaby (2012). Cf. Pecere (2016) on Immanuel Kant’s response to Sömmering.

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neurosciences further integrated into ongoing debates within the history of biology and medicine. More specifically, an important direction for future research would be an investigation of any connections between decline of the reflex theory in the early decades of the twentieth and the subsequent rise of the computational theory after WW2. One connecting thread between these two quite general frameworks for theorizing neural operations is the concept of “representation.” As I have argued, Hughlings Jackson’s introduction of the notion of “representation,” in the context of the reflex theory, facilitated a synthesis between two naturally antagonistic views of the nervous system – a mechanistic or reductionist one and a holistic one in which the function of motor regions of the cortex is to coordinate the movements of the entire body, requiring a global integration of “information,” and a high degree of context sensitivity (Chirimuuta 2019). It is plausible that the concept of neural representation, later deployed by computational theorists, has a similarly synthetic nature. In other words, it fills some explanatory gap – regarding the neuronal basis for the integration of sensory information and coordination of behavior – that would be left under an exclusively biomolecular approach to experimental neuroscience, an approach made popular with the development of tools for performing local and precise interventions on neural tissue. This line of investigation invites comparison with the rise of informational thinking within biology that coincided with the “molecularization” of the discipline following the discovery of DNA (Kay 2000).

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Historiography of Marine Biology

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Contents General Historiography of Biological Oceanography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Conferences and Edited Volumes: 1993–Present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Areas of Concentration in General Historiography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . History of Land-Based Research . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Historiography of Marine Stations 1910–1995 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Historiography of Marine Stations 2002–Present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Public Interactions with Marine Science . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . The State of the History of Marine Biology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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Marine biology is a difficult field to define. Those who work in the field come from a variety of subdisciplines, including chemistry, ecology, and biology. The history of this interdisciplinary field is equally as inclusive, encompassing historians of environment, oceanographies, fisheries, and culture. This chapter examines three distinct areas of historiography: biological oceanography, marine research stations, and public interactions with the ocean. A review of the literature shows that while there are distinct areas of study, there are few internal debates that bind the community. I identify new trends in the field and suggest avenues for future research. Marine biology is a difficult field to define. Someone claiming to be a marine biologist might work in chemistry, genetics, geology, ecology, microbiology, animal behavior, or any number of other fields. The term marine biologist did not exist until

S. Muka (*) Stevens Institute of Technology, Hackettstown, NJ, USA e-mail: [email protected] © Springer Nature Switzerland AG 2021 M. R. Dietrich et al. (eds.), Handbook of the Historiography of Biology, Historiographies of Science, https://doi.org/10.1007/978-3-319-90119-0_24

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the mid-twentieth century and even today is a loose form of identification, rarely used as a disciplinary designation. The lack of disciplinary definition in marine biology is not for lack of trying. In the post-World War II era, when government resources were flowing into marine endeavors, a group of prominent marine researchers sought to more concretely define the borders of marine biology. Many fields are defined by methodological similarities, but in the mid-twentieth century, marine biologists formulated “a definition for their field that prized the diversity of approaches and drew a geographic boundary (i.e., the marine environment) as the unifying feature of ‘marine biology’ and ‘biological oceanography’” (Ellis, 471). The result is that marine biology is defined primarily as exploration, experimentation, and observation on, near, or involving marine organisms and the marine environment. This incredibly loose definition encompasses what many call marine biology and the more specific designations of biological oceanography and fisheries biology. Applying the geographical definition of marine biology to historical literature would result in an expansive and unmanageable historiographical essay. Long histories of marine subjects almost always start with Aristotle and his observations of marine organisms on the docks of Lesbos. During this period, Aristotle described a variety of aquatic species, including the behavior of the paper nautilus and some of the earliest recordings and theorizing of bioluminescence. He is often whiggishly referred to as a marine biologist (or the “father of marine biology”) (Anctil 2018). In this vein, we can include many major figures over the course of the history of biology in this essay. Darwin’s work on the Beagle and his book on barnacles could easily lead us to label him a marine biologist (for an overview of Darwin’s marine research, see Stott 2003; Sponsel 2016, 2018). Ernst Haeckel’s work on invertebrates and iconic images of cnidaria could also place him squarely in this camp. A host of other researchers fall into this category because marine organisms have been consistently used for basic experimental research throughout the nineteenth and twentieth century due to the combination of their abundance and the desire for researchers to gravitate to the seashore in summer months. While it would not necessarily be wrong to place these researchers into the history of marine biology, and it might increase the visibility and prestige of my own field, a more structured definition of marine biology, and therefore when the historiography starts, is needed. For the purposes of this chapter, I will review the historiographical literature focusing on marine study from 1872 to the present. While earlier natural history can be considered in the marine biology tradition, many historians of marine science mark the voyage of the HMS Challenger (1872–1876) as the beginning of modern marine biology writ large. Before this, work was done on or near the ocean on marine organisms but without specific intent to understand the ecosystem more fully. The Challenger was an expedition launched specifically to study the marine environment and resulted in both a widespread knowledge of a large group of marine organisms and in the jumpstarting of more long-term studies of the marine realm. Using this as a starting point, the history of marine biology in this historiographical essay encompasses both the history of ship-based and land-based research after 1872 (Deacon 1971).

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The Challenger was a symptom of larger prevailing ideas about the growing importance for individuals and nation states to understand marine resources. Prior to the mid-nineteenth century, the ocean was a black-boxed, watery depth sailed and labored upon hesitantly. Travelers, sailors, and fishermen floated upon the marine waters, but the marine environment was not a common space for leisure or interest from those termed “landlubbers.” However, the mid-nineteenth century brought several major changes to this environment. The first is the turn toward the ocean as a salubrious place for health and leisure in the Western tradition. The linking of the seashore to inland communities via new forms of transportation and the rise of the beach vacation resulted in a greater interest in these spaces for natural history collecting and discovery. Second is the rise of industrial methods of fishing and the rapid realization that fish stocks were plummeting worldwide. The development of trawling and faster, wider ranging vessels meant that more fish could be caught; however, these developments also resulted in recognition that stocks were being rapidly depleted. While T.H. Huxley assured those at the International Fisheries Exhibition in London in 1883 that the oceans could not be overfished, nation states dependent on fisheries sought to understand their marine resources better (Nielsen 1976). This resulted in many countries developing fisheries programs that studied marine organisms and sought to breed and stock these organisms to increase abundance. These initiatives took biological examination and experimentation. Finally, the enhanced focus on vanishing coastal resources meant increased attention to staking nationalist claims to coastal waters and the resources in them. To do this, nations surveyed coastlines and studied the abundance and movement of their resources over longer periods of time, developing institutions close to the water to do so formed the impetus for the growth of marine biology in the last decades of the nineteenth century and throughout the twentieth and twenty-first century; the history of marine biology tracks onto these variables clearly. This chapter lumps marine biology, fisheries biology, and biological oceanography under the umbrella of marine biology because these fields are intertwined due to both the geographical definition of marine biology and the way that marine research arose. The first section will look at the historiography of biological oceanography broadly. This work looks at the history of ocean exploration starting with The Challenger expedition and extending into the present day; it is primarily the history of vessel-based marine science but includes a wide range of scholarship, including marine fisheries, gathered together through the development of several dedicated conferences. The second section reviews the historiography of land-based institutions, specifically marine laboratories. This literature intersects with larger themes in the history of biology and focuses primarily on institutional history of marine laboratories. The final section will review recent literature in the public culture of marine biology and animal studies. This is one of the newest forms of historical analysis and has great potential for expansion. Like the definition of the scientific discipline, the definition of marine biological history is one where subject transcends methodology. While there are traditional historians in this group, many of whom draw on extensive archival research for their studies, there are also sociologists, literary theorists, practicing scientists, and a

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whole host of other disciplinarily distinct methodologies included in this literature review. In this way, the historiographical boundaries of the field mimic the disciplinary boundaries nicely. If it is about the history of biological research done on or by the water, it’s marine biological history. The historiography of marine biology shows a very scattered literature, broadly and thinly spread. Many subjects have only one or two publications dedicated to their history. The largest concentration of literature is in the history of marine biological stations and even this literature is limited primarily to a few stations in Europe and the United States. The last decade has seen a surge in marine biological literature, but there remains a lack of focus. Marine biological historians now write about environmental and ecological history, non-Western institutions, and focus on a variety of social and cultural lenses. But this expansion does little to add coherency to the subdiscipline and the field remains loosely defined. It is unclear if this lack of definition is the strength or the weakness of the subdiscipline.

General Historiography of Biological Oceanography The history of ship-based marine research shows how difficult marine biology is to define. The Challenger expedition provides a generally agreed-upon date for the start of the field, but it does not produce an agreed-upon definition of the field in the historiography. Early ship-based marine research often involved national or commercial goals of mapping coastlines and seafloors. In addition, these research expeditions took detailed information on meteorological events and sought to track tides and currents. These endeavors fall under the disciplinary umbrella of physical oceanography. However, questions about physical oceanography were often entangled with biological theories. The earliest debates about the depth of the ocean involved not just sounding to understand dimensions but also describing what lived in those depths to understand physical pressures and conditions in that environment. To answer these questions, intensive biological surveys were conducted on these expeditions. While much of the biological materials were analyzed after the expedition, and therefore not done onboard, the collecting process and products jumpstarted larger ecological and biological studies of the marine realm. The earliest marine science expeditions combined physical and biological science, and the historiographical literature reflects this intertwining. There are no specific histories of shipboard marine biology; instead, there is literature that looks at the way that researchers used physical and biological methods to understand the ocean. Margaret Deacon’s Scientists and the Sea: 1650–1900 (1971) ushered in the history of shipboard science and identified the development of modern oceanography. Deacon traces the history of investigation of the oceans in the Western tradition from the ancient world to the years shortly after The Challenger expedition. Deacon is primarily concerned with physical oceanography throughout the book, looking at the history of the study of oceanic depths and tides. But her focus on The Challenger established this expedition as the beginning of modern marine biology and shows how shipboard research made it difficult to separate biological and physical studies

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in earlier periods. In her final two chapters of the book, Deacon transitions from talking about the physical oceanography of these expeditions to showing how these voyages jumpstarted modern marine biological research. According to Deacon, The Challenger jumpstarted oceanographic traditions and “the emergence of Edinburgh as a centre and as Wyville Thompson and his colleagues as arbiters for the world of marine science” during the last decades of the nineteenth century (1971, 370). Deacon focuses on how the transfer of survey materials from The Challenger set up networks of transatlantic biological conversations. While the majority of the book focuses on the history of physical oceanography, Deacon’s work is also the starting point of the history of marine biology. If Deacon covered the earliest development of shipboard marine biology, Eric Mills carried the narrative to the modern era with his Biological Oceanography: An Early History (1989). Mills does not pick up where Deacon left off but instead centers the development of biological oceanography in Germany and Scandinavia. If Deacon’s definitions of oceanography are murky, sometimes focusing on geological research and sometimes on biological identification, Mills gives a satisfying definition of biological oceanography: he centers the field around the combination of quantitative study of plankton and shipboard research. This definition makes it impossible to extend the book backward toward the ancients but places the start squarely in Germany in 1870. Mills traces the development of sampling and statistical methods in Germany and Scandinavia in the first half of the book. This portion of the book is incredibly important, as the larger histories of marine biology tend to focus on England and the United States, leaving out particularly important seagoing powers such as Russia, Scandinavia, Japan, China, and Canada. In the second half of the book, he centers the growth of new methods of biological oceanography in the mid-nineteenth century at the Biological Association of the United Kingdom’s marine laboratory in Plymouth, England, and in the newly founded Woods Hole Oceanographic Institution and the Scripps Institution of Oceanography (SIO) in the United States. Where Deacon linked Europe and the United States through the development of academic biology and the sharing of collections, Mills links Scandinavia and the United States through the development of fisheries biology and national interests in understanding fisheries research. While these books cover a large amount of time and space, they take very different approaches to the study of marine biology and do not necessarily form an unbroken chain of historical analysis from ancient Greece to the modern era. Instead, they show that focus on different definitions of marine biology produce vastly different histories of science. These two works ushered in the first generation of historians of shipboard science, and both Deacon and Mills became integral to mentoring the next generation of historians.

Conferences and Edited Volumes: 1993–Present Beginning in the 1990s, several conferences produced edited volumes, contributing to the general historiography of marine biology. The International Commission of the History of Oceanography, which originally met in Monaco in 1966, met at the

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Scripps Institution of Oceanography in 1993. The conference, co-chaired by Keith Benson and Philip Rehbock, included a wide range of international historians and oceanographers. The meeting resulted in an edited volume, Oceanographic History: The Pacific and Beyond (2002), that shows the permeable boundaries of the history of oceanography and marine biology. Subsections in the book focus on fisheries science, coral research, plate tectonics, technology, and a whole host of other subjects. Section 6 of the volume is entitled “North American Oceanography and Marine Biology” and includes an essay by Benson that looks at the bicoastal nature of marine biology and biological oceanography in the United States. Benson looks at the conditions that led to the development of biological oceanography and the founding of the SIO in California when the focus was primarily on surveying and biological experimentation on the East Coast during the first half of the twentieth century. This edited volume lacks coherence but demonstrates the international and disciplinary inclusivity of the historiography of marine biology. In addition to ICHO conferences, the Maury Conferences for the History of Oceanography, funded by the Office of Naval Research, led to a series of edited volumes. The first Maury Conference was held in Woods Hole Massachusetts, and the Third Maury Conference, held in Monterey, California, in 2001 centered on “oceanography’s role in understanding global environmental conditions and the application of technology to the project of understanding the ocean’s aquatic environment.” This conference resulted in the edited volume The Machine in Neptune’s Garden (Rozwadowski and Van Keuren 2004). While the Maury Conference had a narrower focus than the previous ICHO conference, the papers in the edited volume show an expansion of the definition of the history of marine biology. Christine Keiner’s article on the Chesapeake Bay Hydraulic Model brings together the history of modeling, environmental history, and the history of marine and coastal science. The paper looks at efforts to model the Chesapeake Bay with an expensive system that was abandoned almost immediately upon its completion; Keiner’s work engages more with the history of biological modeling than with other histories of marine biology (Keiner 2004). While this volume and the fourth Maury Conference volume on Arctic Science, Extremes, appear more coherent in theme, they still show a wide acknowledgment of what can and cannot be considered oceanography and marine biology (Benson and Rozwadowski 2007). In 2008, Helen Rozwadowski’s Fathoming the Ocean looked at the development of deep-sea oceanography from the American colonial period through The Challenger expedition. Similar to Deacon, Rozwadowski highlights the role of shipboard surveys on the development of scientific ideals about the ocean. But in an extension and expansion of the field, Rozwadowski clearly shows how the intertwining of cultural, national, and commercial interests in the deep ocean impacted the scientific work performed. According to Rozwadowski, “Scientific interest in the sea intersected with commercial interests, inspired by shipping and sperm whaling, as well as political interests, mobilized by submarine telegraphy, to define the ocean as important new territory” (214). Rozwadowski shows the intertwining of biological and commercial interests in the deep sea through the debate regarding the ability to sustain life at great depths. Sounding in the Atlantic showed how deep the water was

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in these spaces and dredging brought up organisms (always dead). Question arose about the ability for animals to exist in the high pressure of this environment. This was an important biological question for the European and American biologists interested in the biology and physiology of the ocean, but it was also important for commercial interests. The laying of the transatlantic cable pushed scientists to solve debates about what could and did live in these depths for the security of the endeavor. This historical debate allows Rozwadowski to show how these interests acted on each other to spur the exploration of the marine realm both physically and biologically. Rozwadowski was a member of the core group of historians of ocean science in the first ICHO and Maury workshops, along with Keith Benson, Philip Rehbock, David van Keuren, and Ronald Rainger, and she remains an important node in the network of historians of marine science. The most recent Maury Workshop was held in Halifax, Nova Scotia, co-organized by Helen Rozwadowski and Katherine Anderson. Anderson’s work focuses on the history of meteorology and exploration; she is currently researching William Beebe’s ocean expeditions. As with previous conferences, the result of this conference was an edited volume, Soundings and Crossings (Rozwadowski and Anderson 2016), that represents the status of the field of history of oceanography broadly. The essays span 170 years (1800–1970), and articles run the gamut from Megan Barford’s essay on early coastal mapping practices to Jennifer Martin’s paper on the history of the Scripps Model in scientific diving. Rodolfo Alaniz’s essay adds further nuance regarding the political and cultural shaping of the transatlantic debates by examining in-depth the role of major biological figures such as Darwin and Ernst Haeckel in the deep-sea biology debates first highlighted by Deacon and then Rozwadowski, making it one of the longest threads of research in the history of marine science. There is no debate in this thread but instead a sense of bricolage that continues to explore these historical debates and examines how they intersect with larger histories of physical oceanography, cultural history, and history of biology. In addition to the edited volumes on the history of marine science, there have been two edited journal volumes on the history of marine science. The first is an edited volume in Isis. The Focus section, entitled “Knowing the Ocean,” was co-edited by Rozwadowski and Michael Reidy. The section included a co-authored article between Reidy and Rozwadowski entitled “Science, Ocean, and Empire,” an article by Jennifer Hubbard on the development of fisheries biology, and another by Jacob Darwin Hamblin on the development and spread of the “Bergen” school of quantitative modeling in oceanography. Hamblin’s piece is particularly interesting because it pushes further the history of ocean modeling by looking at the history of quantitative analysis developed by the Bergen Geophysical Institute in Norway. Picking up on the earlier work by Mills, Hamblin shows how mathematical modeling spread through both physical and biological oceanography (still a Gordian knot of interdisciplinarity) throughout the twentieth century. This special focus section, which I will come back to later in the paper, shows that even choosing four papers on the history of marine science ranges widely in subject. In the same year, an edited volume on the history of marine science appeared in História, Ciências, Sau´de-Manguinhos. This volume, released in Portuguese,

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includes 13 papers covering the history of marine science in Portugal, Brazil, and Argentina. Maria Fernanda Rollo, Maria Inez Queiroz, and Tiago Brandao’s edited piece “The Sea as Science” traces the rise of marine science in Portugal from the turn of the twentieth century to 1974 (Rollo et al. 2014). The authors show how initial interest in studying the ocean came from national interests in naval power and fisheries use. Over time, these interests combined with commercial endeavors and cooperation with larger international scientific communities to build a large community of marine researchers in Portugal. The edition of Historia shows how much work is being done in the history of marine science in languages other than English and highlights how these narratives of the history of marine science fit together to form a wider narrative. It is important that more non-Atlantic, non-US work be done so that a fuller picture of the history of the study of the sea can be realized.

Areas of Concentration in General Historiography The study of the marine vessel as a space for biological research is one that has become more popular in recent years. Margaret Deacon and Helen Rozwadowski (Rozwadowski 2005) both focused on the work done in both early physical and biological oceanography onboard. Subsequent work was done by Dean Allard on the United States Bureau of Fisheries first collecting boat, the USS Albatross. Allard’s work looks at the way Spencer Allerton Baird, the first fish commissioner, employed the Albatross and the work that was done on the ship (1999). In a short article in PLOS One (2013), Niki Vermeulen argues that debates over the turn to “big science” post-World War II ignore the history of marine science. According to Vermeulen, marine science was the original big science, with the development of large shipbased voyages requiring the monetary cooperation of large communities with national or commercial interests. Anthony Adler (2014) has looked at the way that shipboard spaces have been shaped and reshaped by the needs of marine researchers. Adler looks at the progression of ships starting with The Challenger and states that a progression has occurred, from the “ship as instrument” to the “ship as a laboratory” where science is performed to the “ship as invisible technician” that gather data but where science is again no longer actively practiced. More recently, Penelope Hardy’s (2017) work on the long history of research vessels, including the changing structure and culture of shipboard oceanic research, shows that these spaces are important for understanding how ocean science has evolved. The second area of concentration in the history of marine biology is that of fisheries history. In the ICHO and Maury conferences, there is a group of historians of fisheries. The inclusion of fisheries history has been consistent at these events, although there is a much larger community of historians of fisheries, of which marine fisheries is only a small subsection. Arthur McEvoy’s The Fisherman’s Problem is both a legal and environmental history of California fisheries in the twentieth century. McEvoy uses the concept of the “tragedy of the commons” to look at how a once plentiful resource available to a wide range of individuals became commercialized and privatized (1990). Joseph Taylor’s work on salmon looks at

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the long history of salmon in the Pacific Northwest, looking at the history of fisheries from the earliest native fishing techniques to the modern fisheries biology that seeks to control a dwindling population (2009). Both McEvoy and Taylor’s work shows how the history of marine fisheries is one that requires a varied methodological approach that combines the history of biology, environment, and law. While these books look at the long history of dwindling stocks and point to mediation efforts and field research, another group of scholars have focused on the development and deployment of fisheries biology as a discipline. Jennifer Hubbard (2006) and Tim Smith (1994) published books on the development of fisheries biology in Canada and the United States, respectively. Both books look at the main players in fisheries research in these nations and how they used a combination of laboratory, field, and statistical methodologies to set national fishing policy. Carmel Finley’s All the Fish in the Sea (2011) continues this work by looking at the development of a single statistical model, maximum sustainable yield (MSY) between 1949 and 1955. This work and her subsequent book All the Boats on the Ocean (2017) show how scientific modeling sets fishing limits that affect national and international policy. Vera Schwach has written extensively on the history of fisheries biology and management in Norway and particularly on the impact of Johan Hjort’s work on both the development of fisheries biology and the international community of scholars working in ICES (Schwach 2002, 2012, 2014; Schwach and Hubbard 2009). Her work is particularly important in tracking the continued impact of Norwegian fisheries biology science and practice. Examining fisheries intertwines the historiographies of marine biology and environmental science. Frederick Davis’ book Archie Carr: The Man Who Saved Sea Turtles (2012) highlights the struggle for Carr, an ecologist working at the University of Florida, to understand and stop the causes of sea turtle mortality globally. Of particular interest is Davis’ discussion about Carr’s motivations for seeking fisheries prohibition over management in turtle conservation. In addition, Kjell Erickson’s work on the history of red tide in the Japanese pearl oyster industry is another look at fisheries biology. Erickson’s work looks at the history of the cultivated pearl industry and shows the way that Japanese oyster farmers explained and sought to alleviate the impact of red tide on their crops (2017). One area that has seen recent growth is the history of human interactions with marine cetaceans, especially whales. D. Graham Burnett’s The Sounding of the Whale (2012) is a work that highlights the changing relationship that humans have had with whales from the middle nineteenth century into the current era. This book uses the scientific work on whales in a given era to show how the relationship with whales has gone from viewing them as food to seeking to conserve their rapidly dwindling numbers. His narrative takes the reader through the wide range of scientific disciplines involved in this change, from fisheries to environmental biology. Etienne Benson’s “Endangered Science” (2012) shows how the passage of the Marine Mammal Protection Act impacted the way that biologists could track and study cetaceans. This work is one of few that explores the impact of modern environmental law on the study of marine animals. Most recently, Jason M. Colby’s Orca (2018) traces the history of human interactions with killer whales,

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from the earliest maritime myths of man-eating creatures to the current debates over captivity at SeaWorld. Colby’s work is especially important because it shows how quickly human perceptions of sea animals have shifted due to popular images of them in media. Recent research on marine fisheries intertwines marine biology, environmental history, and the history of the Pacific more generally. Todd Braje’s Shellfish for the Celestial Empire uses a combination of historical and anthropological methods to trace the rise and fall of Chinese abalone fishing colonies in California during the 1800s (Braje 2016). Jakobina Arch’s Bringing Whales Ashore is a social and cultural history of Japanese whaling. Her work examines the argument that whaling has a long and integral place in Japanese culture and should therefore be embraced as culturally relevant today. According to Arch, this is debatable. She uses a variety of methods, including following whales from ocean to processing and product movement inland, to show that Japanese whaling did not have the wide-ranging impact that Japanese officials now claim. Finally, Richard Ravalli’s work on the sea otter (2019) traces both the natural and economic history of the sea otter. Ravalli shows that political and social understandings of America were developed through exchange and argument over otter hunting grounds throughout the nineteenth century. All three of these authors’ work sits in a new area of marine biological history – one that uses animal studies to focus on the sea as interconnected with larger, geopolitical histories. Helen Rozwadowski’s work also examines larger geopolitical implications of marine research though her history of the International Council for the Exploration of the Sea (ICES). According to Rozwadowski, ICES has been successful as an international community of scientists “because of its uniqueness as an institution. Neither an academic nor a governmental body, it also had no direct connection to industry” (Rozwadowski 2002, 2). While work done on the history of fisheries biology would mark it as intrinsically political, Rozwadowski’s work reminds us that the creation and dissemination of marine knowledge operate better through international collaboration. More recently, Susana V. Garcia’s article “Commercial Fishing and the Study of Marine Fauna in Argentina, 1890–1930” (2014) looks at the impact of commercial fishing on the development of marine science. Garcia shows how important commercial networks were for the earliest Argentinian marine biologists. “At the end of the nineteenth century, each group of fishermen from Mar del Plata had an agent in charge of sales in Buenos Aires. The fishermen would notify him of the railroad shipments by telegram, and every two or three days the agent would telegraph them back with the selling prices of the products. The zoologists from Argentine museums would draw upon this communications and commercial network to supply themselves with samples and information”(4). Antony Adler and Erik Dücker’s recent work on the development of French deepsea microbiology in the late nineteenth century looks at the way that an emerging marine field combined methods and questions from microbial ecology with emerging deep-sea ecology. Adler’s “aim is to use the example of marine microbiology” to describe the birth of a scientific discipline as an emergent process by which one novel “field of enquiry – in this case, deep-sea biology – allowed the new

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combination of disparate existing disciplinary methods and research questions” (Adler and Dücker 2018, 108). All three of these papers look at the way marine science emerged and grew at the turn of the twentieth century. The historiography on the rise of marine science throughout the world has shown that pressures for the study of the marine environment can come from a variety of institutions, including national, commercial, and cultural. While the history covers a wide range of subjects, it continues to be relatively shallow in theory. Histories of ship board marine science often contribute to theoretical conversations that are not inherently marine based, including debates about the rise of big science and the role of the nation state in science building. Theoretical frames of histories of fisheries focus on larger arguments in environmental history, including perceptions of wilderness, conservation, and animal-human interactions. While all of these theoretical frames are useful, none originated in or has been drastically changed by narratives about the history of marine biology. It is in the historiography of land-based marine research that we see the most concentrated historiography and the development of theoretical arguments therein.

History of Land-Based Research Beginning in the 1860s, governments, universities, and private natural history groups interested in surveying and studying marine resources established permanent marine stations throughout the world. Russia, France, Japan, England, Canada, Germany, Italy, the Netherlands, Sweden, and America all established permanent stations by the end of the nineteenth century. Built by a local scientific society in 1867, the marine laboratory of Arcachon on the Bay of Arcachon in France was the first marine station. Russia’s privately funded Sevastopol Station, founded in 1871, was quickly followed by the Stazione Zoologica Anton Dohrn (1872) in Naples, Italy, and the Station Biologique de Roscoff (1872) in Brittany, France. Others swiftly followed, and new stations opened in Sweden (Kristiniberg, 1877), Japan (Misaki, 1887), Scotland (Gatty, 1896), England (Plymouth, 1888), America (Penikese Island, 1877 and USBF Woods Hole, 1888), Canada (New Brunswick, 1899), and the Netherlands (Helder, 1890) throughout the 1880s and 1890s. By the turn of the twentieth century, most of these countries had established multiple stations (Muka 2014). While there is great geographical diversity in marine stations, the historiography of these stations tends to focus on the Stazione Zoologica and the Marine Biological Laboratory. These two stations are similar: they were started by private communities (not Universities or government organizations), they received revenue from selling specimens to the science community and renting out table space to universities and private science groups, and research was predominantly based on the growth of late nineteenth-century German experimental embryology and morphology. Historians in the 1980s produced a robust historiography of these institutions that linked them to that biological tradition and showed that these laboratories were integral to the development, not just to marine biology but to experimental biology more broadly.

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However, a resurgence in popularity of this topic has occurred in the last 10 years with a wider focus on both the stations and science being studied. The more recent scholarship takes for granted that these spaces are accepted as sites of basic biology and asks how we can further understand the nuances of scientific practice, physically, politically, and socially by studying them. The historiography of marine stations draws on the larger history of marine biology themes and demonstrates the evolution of the field from a focus on individual and institutional history to a broader acceptance of social, cultural, and environmental history.

Historiography of Marine Stations 1910–1995 The earliest history of these stations was produced by Charles Atwood Kofoid for the American Department of Education (1910). Kofoid, a biologist by training, was tasked with touring the oldest stations in Europe so that the United States could develop a plan to build these stations within the United States. Kofoid visited stations throughout Europe, paying close attention to the history of their establishment and funding, how they operated, and the technological dimensions of each location. His history of these stations, many of which were somewhere between 25 and 50 years old at the time, remains some of the only work done on the Russian, Bulgarian, and many of the Swedish stations. Unfortunately, it is difficult to find any work on the Japanese stations although there is a brief history of the Misaki Station by H. Terayama in 1979. Kofoid’s work is the first in a long line of institutional histories of marine stations, focusing less on the science performed and more on the administrative infrastructure of these locations. In 1962, Paul Galtsoff, a Russian-born oyster researcher with the United States Bureau of Fisheries, wrote The Story of the Bureau of Commercial Fisheries Biological Laboratory. The Department of Interior Circular (the Bureau of Fisheries was part of the DOI during this period) gave an account of the biological laboratory founded by Spencer Fullerton Baird in Woods Hole, Massachusetts. This station, founded in 1885, focused on understanding the lifecycle of important fish used in the United States for commercial enterprises. Galtsoff details the work on oysters (his specialty) in this location. Dean Allard added to the history of the USBF marine laboratory in Woods Hole, although there was still little focus on the science performed in these spaces (1978, 1990). Douglass A. Wolfe, a historian of the National Oceanographic and Atmospheric Administration (2001), published a history of another important fisheries laboratory, established in Beaufort, North Carolina, in 1899. This space, previously the site of Johns Hopkins’ semipermanent marine station, was also dedicated to work on the history of commercially important sea life (including the black terrapin turtle). These two histories, both published by the US government, are the fullest accounts of the history of fisheries-based marine stations available to date and both fit into the historiographical narrative of exploring how the stations were developed and administered. While there is some information about the science performed, neither author goes far into the laboratory to examine the bench. The much more common historiography surrounds private- and university-based marine stations.

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In the 1980s, historians of biology became focused on comparative history of marine stations, and specifically the histories of the Naples Zoological Station in Naples, Italy, and the Marine Biological Laboratory in Woods Hole, Massachusetts. In 1980, Jane Oppenheimer, an embryologist who often worked at Woods Hole, published an article entitled “some historical background for the establishment of the Stazione Zoologica.” This article briefly outlines the development of the station by Anton Dohrn, including his ideas about funding and his vision for the station (Oppenheimer 1980). In 1984, Christian Groeben linked the Stazione Zoological with the MBL with a short article about the impact of the Stazione on the founding of the MBL. Groeben argued that the founders of the MBL looked to the Stazione for ideas about structuring their work spaces and funding. Both stations were operated by private groups, not universities. Groups or universities bought table space for researchers and these fees helped finance the stations (Groeben 1984). Other similarities included the way that these spaces focused on experimental embryology and morphology. One major difference which would be a focus of other historical accounts was that the MBL was a space for undergraduate and graduate students to take courses during the summer. The comparison of European and American stations jumpstarted a conversation about these institutions and their role in larger biological traditions. Between 1988 and 1995, a group of historians placed marine laboratories and marine biology at the epicenter of debates about the development of American biological traditions more generally. Jane Maienschein, Christian Groeben, Keith Benson, and Philip Pauly all linked marine biological stations into larger debates about the growth of German experimental biology during the late nineteenth century. Maienschein’s work on the growth of the German experimental program at Johns Hopkins expanded into looking at the way that the MBL was a space of transfer for that tradition from Naples to America (Maienschein 1981, 1985, 1987, 1988, 1989, 1991; Maienschein et al. 1991; Groeben 2006). Maienschein and Benson published articles detailing the differences between Naples and the MBL and sought to understand the reason that the MBL developed and became famous for coursework. Their research showed that American marine stations borrowed ideas from Naples but developed a system linked to universities for both financial and social reasons (Benson 1988a, b, c). Benson and Pauly both questioned how the cultural and social atmosphere of these stations popularized them. This group, whose work is seminal to the study of marine stations and American marine biology, identified the MBL and other American marine stations as spaces that created and sustained a culture of knowledge sharing – both between colleagues and from teacher to student. Pauly called special attention to the familial culture of the MBL and argued that these spaces functioned not just as research centers but as places of leisure for these researchers (Benson 2001; Pauly 1988, 2000). As Benson has stated, marine laboratories became the institutional identity of American marine biology (2001). In addition to the institutional histories, biographical memoirs and edited letters have highlighted the ubiquity of marine research in the careers of well-known biologists and shown the impact that these spaces had on scientific careers and larger biological movements. Many of these manuscripts take the form of biographies, including Philip Pauly’s biography of Jacques Loeb (1987) and Keith Benson’s

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dissertation on W.K. Brooks (1979). Christine Groeben’s edited letters between Karl Ernst Von Baer and Anton Dohrn (1993b) and between Charles Darwin and Anton Dohrn (1982) show a direct link between marine stations and the rise of evolutionary thought and Darwinism throughout Europe. In addition, the popular biography Black Apollo of Science (Manning 1985) about the life and work of Ernest Everett Just, one of very few African Americans to work in American biology during the first half of the twentieth century, shows the importance of these marine stations to his career trajectory. Finally, the newest biography in this group is that of Alfred Goldsborough Mayer, the director of the Carnegie Marine Station in the Dry Tortugas, Florida. Mayer’s work on jellyfish and coral jumpstarted several major research programs throughout the United States. The biography shows how Mayer shaped the Carnegie laboratory to suit his ideas about marine biology and, in turn, how those ideas spread throughout the scientific community through his use of space and community (Stephens and Calder 2006).

Historiography of Marine Stations 2002–Present One of the central arguments that has been picked up recently in literature about marine stations is whether these spaces represent researchers working within field biology or laboratory research. In Landscapes and Labscapes, Robert Kohler states that: Marine stations, despite their seaside location, were essentially extensions of campus labs, bound tightly by the web of teaching and supply to laboratory culture. In marine labs it was not the natural surroundings but cultural habits and customs that shaped practices most powerfully. Morphologists’ desire for fresh material was a harbinger of the ideal of a new natural history, but it was just a small step across the laboratory threshold. Microscopic morphology was a laboratory practice where it was performed, and its cultural geography is visible in the siting and spatial customs of marine labs. (Kohler 2002, 44)

Kohler’s argument was based on the research of those at the Marine Biological Laboratory, specifically Jacques Loeb. But the assertion that marine laboratories, and the scientific research performed therein, were divorced from the environment of the stations has been challenged recently. Raf de Bont’s 2014 book Stations in the Field shifts the focus of marine stations from the MBL and Naples to the French station at Wimereux. According to de Bont, the work done at Wimereux was based on the carefully chosen environment around the station itself; the founder, French biologist Alfred Giard, rejected overly laboratory-based methods for a more ecologically minded approach to marine studies. Wimereux was a quieter, simpler marine station focused on teaching undergraduates where studies of marine botany became popular. De Bont’s argument is that when you focus on a variety of stations in a wide range of locations, it isn’t clear that they were merely extensions of university laboratories (De Bont 2009, 2015). Samantha Muka chose a different lens to challenge Kohler’s assertions; instead of widening the scope of study to other stations, she instead suggests that focusing

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more fully on the experimental work done at marine stations can tell us much about the mix of laboratory and field work being done. In her article “The right tools and the right place for the job,” she looks at the history of neurophysiological research performed at marine stations between 1880 and 1930. Focusing on the MBL and the Carnegie Station in the Dry Tortugas off the coast of Florida, Muka argues that researchers working on neurophysiological questions during this period were forced to seek out marine stations that contained suitable organisms for this work. Both the MBL and the Carnegie laboratory had an abundance of acceptable jellyfish and therefore became epicenters for this research. Muka argues that “To work with these experimental organisms, one had to travel to them and immerse oneself in the environment and lifecycle of a specific organism” (Muka 2016b). Both de Bont and Muka argue that to truly understand the role of the field and laboratory in marine station research, it is important that we widen our subjects, be it by location or discipline. De Bont and Muka are part of a newer cohort expanding the historiography of marine stations by looking at a wider range of institutions and by identifying previously overlooked researchers in these spaces. Robert-Jan Wille’s work on Paulus Hoek and the transnational politics of biological oceanography at the Plymouth, Den Helder, and Helgoland marine stations opens new avenues of research into marine stations while linking them to the larger sociopolitical themes seen in the last section (Wille 2017). Christina Luk’s Forthcoming work on embryological research at Amboy marine station in China will similarly expand these conversations with a focus on new geographical and political communities (Luk Forthcoming). Both Wille and Luk work within the comparative framework to analyze the role of marine stations in the spread of biological theories and technologies, not just to America but around the globe. Another group of scholars is expanding on the social and cultural history of these spaces. Jenna Tonn’s work looks at the Bermuda Biological Research Station, run jointly by Harvard and NYU. In her research, Tonn examines the role of the BBRS as an extension of Harvard’s zoology program and shows how the colonial atmosphere of Bermuda shaped the racial and gendered characteristics of both the researchers admitted and science performed in these spaces (2015; 2018). Helena Ekerholm has done similar work on the Kristineberg Marine Zoological Station in Sweden; her work examines the role of support staff drawn from local communities in shaping scientific spaces at these institutions (Ekerholm 2015; Ekerholm et al. 2018). Both Tonn and Ekerholm approach the history of marine stations not just as a scientific space but as a domestic space that is permeable to not just the natural environment that surrounds it but also to the social and cultural environments. Marine stations both impact and are impacted by these social and cultural environments, and therefore, to understand the science, we must understand these milieux. A 2014 edited volume focusing on the institutional history of the St. Andrews Marine station in Canada shows that anchoring the history of marine biology at marine stations provides new avenues for studying the history of this scientific field. Edited by Jennifer Hubbard, David Wildish, and Robert Stevenson, the volume brings together scientists, historians, and administrators to review the first century of the station. Hubbard places the station into the historical context developed by the historiography

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outlined above and shows that the St. Andrews station fits into that narrative. But this volume also contains chapters on the role of women researchers at this station and the studies of shellfish toxicology and estuary ecology at St. Andrews. The attention to the technologies at the station and the studies performed there mark an expansion of the marine station narrative in the historiography. The book reminds us that these institutions can ground a wider exploration of marine biological fields that have not been explored in historical texts (Hubbard et al. 2014). New research has shifted the narrative from showing how integral marine stations have been in the growth of biological traditions to exploring more closely what it means to be doing marine biology more widely. The expansion of the historiography shows that marine research is only tethered by location, not discipline or methodology. Marine stations continue to be funded throughout the world and to operate as an integral part of government and university science systems. As these institutions persist, it is important that scholarship seek to understand what has made them so integral to the operation of marine biological science.

Public Interactions with Marine Science The final subcategory of the history of marine biology is the growing area of interest in the historical development of perceptions of and public interactions with the marine environment and marine biology. As Jakobina Arch states in her 2018 book Bringing Whales Ashore, “We also need to have histories of the ocean that include not just those people directly working in maritime spaces, but also all the other people who interact with it indirectly, even without having been there” (Arch 2018,19). To truly understand the expansion and direction marine biological research has taken over the last century, historians must look not just at institutional and national history but also at the cultural and social history of marine science. Public perceptions and interactions with the marine world can have obvious impacts on the way that marine science is done. The anti-vivisection movement of Victorian era changed the way scientific experiments were performed on terrestrial animals. Interestingly, it pushed some researchers into the use of marine organisms. George Romanes, an antagonist of anti-vivisectionists, stated in his 1885 book Jellyfish, Star-fish, and Sea Urchins that if you had a problem with vivisecting jellyfish, you should not eat oysters either. The public perception of the harm caused to animals in the laboratory pushed Romanes to find the least cuddly organism on which to work (Romanes 1885). In recent years, we have seen a surge of public interest in the welfare of marine mammals in captivity. Documentaries such as “The Cove” (2009) and “Blackfish” (2013) have focused public attention on the history and current practice of keeping animals in captivity. In turn, public backlash against institutions with captive mammals has had very real effects on the science performed in these spaces. For instance, the backlash against captive marine mammals has pushed mammologists to use innovative sampling techniques with wild caught specimens, resulting in advancements in a variety of field methods. This example shows that it is important to pay attention to perceptions of the marine realm more broadly. The history of

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public interactions with marine science can tell us much about how the field has been and is continuing to be shaped. The earliest work on public perceptions of marine science comes in the form of examining the history of collecting and aquarium keeping. In Victorian England, the rise of the railway popularized excursions to the seashore, leading to a “collecting craze.” David Allen’s essay “Tastes and Crazes” in the edited volume “Cultures of Natural History” highlights the importance of collecting seashells, seaweed, and other organisms from the coastline for both personal pleasure and scientific leisure (Allen 1996). The development of the personal aquarium and vivarium to keep these collections in the home, and the development of décor around these crazes, marks the beginning of the public’s fascination with the marine realm. Anne Secord’s work on botany and the popularity of botanical collecting and sharing includes work on seaweed collection. She explores the importance of visualization and aesthetic beauty in early popular texts on seaweed (Secord 2011). Ann Christie has written more in-depth about how the craze for marine collection intersected with fashion during this period, writing about the use of seaweed motifs on cotton material. In her work, we see how individual natural history pursuits impacted widespread perceptions of the ocean environment (Christie 2011). Molly Duggin’s work on seaweed collecting shows how marine interactions were more than mere fashion. She states that “Affordable and portable, albums played a significant, although often under-valued, role in the transmission of popular natural history pursuits, collecting patterns, aesthetic tendencies, domesticating initiatives, and performances of civility throughout the British colonies and, indeed, farther afield, circulating physically through global maritime networks and socially through gifting and exchange” (Duggins 2016, 16). All of these papers show how widespread the culture of seaweed collection was and how that craze impacted not just a small group of individuals who collected at the seashore but also more general understanding of what was beautiful and unique about the English and colonial coastal environment. Other historians have highlighted the role of personal aquariums in perceptions of the marine realm. Christopher Hamlin’s “Robert Warrington and the Moral Economy of the Aquarium” argues that the “balanced aquarium” – a self-perpetuating ecosystem developed by Robert Warrington in 1850 – became a teaching tool about the perfection of nature and the way that natural objects interact (Hamlin 1986). Aileen Fyfe’s work Science and Salvation contains a chapter entitled “Reading Fish” (Fyfe 2004). In this work, Fyfe builds on Hamlin’s work to show how Christian publications urged the keeping of aquariums to teach children about the wonder of God and the order of nature. These works highlight the importance of studying hobbies to understand the development of natural ideas in the larger public. The research on home aquariums has been inconsistent. For instance, Hamlin’s 1998 paper has no historical citations for the history of the aquarium (only primary sources), and until recently, Hamlin, Fyfe, and Allen remained the only sources for aquarium history. In 2005, Bernd Brunner published The Ocean at Home: An Illustrated History of the Aquarium. This is a popular work that contains wonderful images of the earliest Victorian aquariums but does not engage with historical themes about popular science, nor does it do much work on aquarium history in the modern era (Brunner 2012). More recently, Judith Hamera’s Parlor Ponds: The

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Cultural Work of the American Home Aquarium 1850–1970 uses performance theory to analyze the work that aquariums did in the American home during this period. Her thesis, that the act of collecting and curating the aquarium helped alleviate the stresses of modernity on masculinity, is an interesting perspective on the technology and its impact (Hamera 2012). However, more work needs to be done on the home aquarium hobby and its history to understand how the hobby impacts and is impacted by more traditional forms of marine biology. Other historians have sought to understand public perceptions of the marine realm through visibility and popular attractions. Susan Davis’ Spectacular Nature (1997) argues that SeaWorld and other and other amusement parks are not scientific institutions, but instead argues that they are amusement parks whose theme is “science.” Samantha Muka’s work on the history of conservation in public aquariums directly challenges Davis’ assertion that these spaces perform no conservation or science by tracing the role of these spaces in conservation from 1850 to the present (2018). Muka’s argument is that initial conservation initiatives in these spaces do not resemble those agreed upon by modern conservationists but that these spaces conveyed conservation to the public through their engagement primarily with fish stocking initiatives and public education. More recently, Kelly Bushnell’s work on the history of cetacean captivity in Victorian England adds nuance to the discussion of the intertwining of conservation and amusement in public aquarium spaces. Bushnell reads early aquatic spaces and looks at the way that “the whale” was constructed as an object for consumption by masses through the framing of the animals in tanks at amusement shows and aquariums. Her work bridges that of Davis and Muka by examining both the intent and outcome of the public display of marine animals (Bushnell Forthcoming-a, Forthcoming-b, Forthcoming-c). Gregg Mitman’s 1999 book Reel Nature looks at the history of dolphins on film and the way that water parks shape the idea of nature and present scientific research to the public. Jonathan Crylen’s 2015 dissertation on the history of underwater film goes further, stating simply that “We have, it seems, come to understand the ocean, and develop ecological ideas about it, primarily on account of films and the stunning images of marine life they contain” (Crylen 2015, 1). Michaela Jane Thompson’s 2016 dissertation on sharks shows how images of sharks in popular films such as Jaws impacted the research done on sharks internationally. Of particular importance, Thompson shows that fear of sharks has not only impacted the way that scientific research is performed but also how it is communicated back to the public (Thompson 2016). Dolly Jorgensen’s piece “Mixing Oil and Water” highlights the inclusion of oil rigs in public aquarium displays as a way for public aquariums to train the public’s eye and mind into perceiving the oil rig as a natural occurrence in marine spaces (Jørgensen 2012). All these pieces investigate how public spaces shape perceptions of the marine realm and seek to make “natural” a cultivated image of the marine space. Recent publications have shown how visualization of the marine realm is important for thinking through how marine biologists work. Lynn Nyhart and Christine Wessely have studied the development of ecological thinking in nineteenth-century Germany. Both historians trace the rise of ecological thought, and the idea of ecology more generally, to the use of these “watery milieus” that facilitated the visualization of community structures in the laboratory (Nyhart 2009; Wessely 2013; Huber and

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Wessely 2018). Samantha Muka’s work “Imagining the Ocean” looks at how scientists think about and portray marine organisms. Working with scientists, artists produced images that reflected scientific beliefs and trained public perceptions of spaces that could not be viewed directly (Muka 2014). Hannah Rose Shell’s film Things Under Water (Shell 2005) traces the history of locomotive underwater film from the work of Etienne-Jules Marey in 1898 to the present looks at how he sought to portray locomotion in this medium. Rodrigo Salvador and Barbara Tomotani look at how persistent myths of the kraken influenced the way that early marine researchers viewed giant squid and other marine organisms that resembled the famed monster. They show how the myth of the kraken influenced taxonomic identifications in molluscs, octopuses, and squids in the late nineteenth and twentieth century (2014). Natascha Adamowsky’s work on William Beebe and the visualization of the underwater realm explored how Beebe’s images in the popular work Half Mile Down impacted perceptions of the ocean. Her wider work on the underwater visualization and imagination shows how wonder and science intertwine to create ideas about the marine realm (Adamowsky 2015). All these publications seek to understand how scientists are shaped by and how they then shape the larger public imagination about the sea through visualization. Another area of research that shows how the public views marine science and the marine environment is the literature on exploration. Gary Kroll’s America’s Ocean Wilderness: A Cultural History of Twentieth-Century Exploration (2008) looks at episodes of exploration that were popularized in the American consciousness. Kroll argues that the sensational nature of ocean exploration is an extension of the “wilderness ethic” to the oceans; basically, the ocean is an unknown territory of the United States. With the settling of the West, the marine realm became the last frontier for exploration (along with space), and the narratives that are created around that exploration shape the science and vice versa. Kroll looks at some of the most recognizable explorers and naturalists, including Jacques Cousteau, Thor Heyerdahl, Roy Chapman Andrews, Robert Cushman Murphy, Eugenie Clark, Rachel Carson, and William Beebe. Public interest in these individuals continues to be high, with Brad Matsen writing biographies of Cousteau (2009) and Matsen (2006). Beebe did not necessarily want his biography written and disposed of most of his personal papers, but Carol Grant Gould was given access to his remaining estate and published The Remarkable Life of William Beebe in 2004. Kroll, Matsen, and Gould all highlight the way that science and sensation intertwine in the lives of marine explorers and, in turn, impact the way that the public perceives the importance of marine exploration.

The State of the History of Marine Biology The history of marine biology and marine science more generally remains undefined. In the Preface for the 2016 volume Soundings and Crossings, Eric Mills asks: Is there a history of oceanography? In their introduction to this volume Anderson and Rozwadowski don’t have much patience with that concept, preferring alternative phrasings such as “history of studying the ocean” and “history of ocean sciences.” “History of

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oceanography” appears to be shorthand for a complex of studies that range across all the subjects dealt with in this book and more besides. I agree with them. And yet, when asked what I do, I respond without hesitation that I study the history of oceanography, preferring the shorthand to the Byzantine discussion of what that term encompasses. There is something here that makes sense, despite the wide range of subject matter. (Anderson and Rozwadowski 2017, xv)

I am a historian of marine biology, but much of my work involves reading, writing, and thinking about a wider range of research on the history of marine science. The application of the title of historian of marine biology is as personal as that of the application of marine biologist – it tells almost nothing about what I research and more about the general community in which I fit. Our interests are with the ocean. This lack of boundary setting has resulted in a field of literature that is expansive and spread thinly. There are very few debates within the historiography, mainly because there is so much history yet to be done. This does not mean that the history of marine biology does not engage with current debates in the history of biology, but they are mostly concerned with adding nuance to a largely terrestrial focus in the history of science more generally. In her co-edited 2014 Isis Focus section, Rozwadowski states that there is a “modest existing tradition of the history of oceanography” but that new research is positioned to “draw the ocean itself into history” (335; see also Rozwadowski 2010). In the same volume, Naomi Oreskes makes a stronger call for the incorporation of the history of the ocean, and marine science specifically, into existing understandings of the history of science more generally. According to Oreskes, previous historians have ignored the marine realm “because we have viewed it as standing mostly apart from human societies and activities.” Oreskes says that general historians have traditionally ignored the sea as a void – a place where history mostly doesn’t happen. In addition, historians of science have also ignored the sea, mostly because we are “not so much historians of science but of scientists” (379). Oreskes recounts a roundtable at the American Historical Association in 2010 that drew a single audience member. The participants immediately moved the roundtable discussion to the more comfortable setting of the hotel bar. In this anecdote about the history of the field, we see that historians of marine science have not necessarily been internally debating as much as battling for their subject to be recognized as integral to developing a complete view of the history of science more fully. Recognition of the importance of this field may come, not only as the product of the hard work of historians of marine biology but because the world’s attention is rapidly shifting toward the marine realm. Climate change, dwindling marine fisheries, and increased plastic pollution in the marine realm have shifted conversations and focus onto the marine world. Historically considered an immutable and bountiful resource, the ocean is now being viewed as a shifting resource, capable of not just change but destruction. As the rising ocean begins to change coastlines, scholars’ attention will shift toward understanding the history of marine interactions, including the history of marine science. We’ll be waiting at the bar.

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Historiography of Physiology

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Contents Physiology-as-the-Study-of-Life-Itself: Histories as “Catalogues of Physiological Discoveries” (~1920s–1970s) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Physiology as “an Independent Science”: Histories of Institutes, Schools, and “National Styles” (~1940s–early 1990s) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bernard and Ludwig . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Two Trends: Vitalism/Mechanism and National Stories . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Physiology-as-Society: Political and Economic Histories of Physiology (~1980s–Present) . . . Histories of Physiology in Political and Economic Context: Some Examples . . . . . . . . . . . . . Histories of Physiology as the Study of “Labor” . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Two Opportunities: Gender and Race . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Physiology-as-Practice: Histories of the Instruments and Organisms of Physiological Experiment (~1980s–Present) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . The Death of the Queen? Histories of Physiology-After-1940 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Histories of Physiological Bodies in Place . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Future Directions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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Abstract

In 1978, Gerald Geison opened his ambitious biography of the celebrated British physiologist, Michael Foster, declaring a peculiar lack of scholarship on the history of physiology after 1850. Forty years later, Geison’s observation resonates. While historians of science since Geison have showed some interest in physiology through 1940, most historians of physiology have continued to study the nineteenth century. How to explain this historiographical trend? What hap-

A. Johnson (*) History and Sociology of Science, University of Pennsylvania, Philadelphia, PA, USA e-mail: [email protected] © Springer Nature Switzerland AG 2021 M. R. Dietrich et al. (eds.), Handbook of the Historiography of Biology, Historiographies of Science, https://doi.org/10.1007/978-3-319-90119-0_26

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pened in the middle of the twentieth century to physiology, the nineteenth-century “queen of the natural sciences” (du Bois-Reymond quoted in Lenoir 1988)? Did physiology – as an experimental enterprise, academic discipline, set of questions, career – die out over the course of the twentieth century, such that there wasn’t much to write histories of? Did the post-WWII “molecular gaze” in the life sciences completely eclipse the physiological “molar body” (Rose, 2007), with its organic systems that had captured the minds, eyes, and hands of physiologists like Claude Bernard [1813–1878] or L.J. Henderson [1878–1942]? Did the rise of genetics in the twentieth century leave no room for physiology other than as a one-off required course for medical students? Or, did physiology persist as a scientific discipline throughout the twentieth century, somehow failing to attract the attention of most historians of twentieth-century science? Motivated by these questions and more, this historiographical essay chronicles scholarship on “the history of physiology,” a phrase that has meant, for some historians, the history of the study of “life;” for others, something akin to the history of Western medicine; and for still others, the history of experimental investigations of vertebrate organisms only after 1800. The conclusion attempts to answer the simultaneously historiographical and historical question of what happened to physiology in the twentieth century. In the history of science, despite great ferment and activity in recent years, many large areas remain virtually unexplored. Conspicuous among them is the history of physiology after 1850. – Gerald Geison, (1978, xi), Michael Foster and the Cambridge School of Physiology.

In 1978, Gerald Geison opened his ambitious biography of the celebrated British physiologist, Michael Foster, declaring a peculiar lack of scholarship on the history of physiology-after-1850. Forty years later, Geison’s observation resonates. While historians of science since Geison have showed some interest in physiology-through-1940, most historians of physiology have continued to study the nineteenth century. How to explain this historiographical trend? What happened in the middle of the twentieth century to physiology, the nineteenth-century “queen of the natural sciences” (du Bois-Reymond quoted in Lenoir 1988, 139)? Did physiology – as an experimental enterprise, academic discipline, set of questions, and career – die out over the course of the twentieth century, such that there wasn’t much to write histories of? Did the postWWII “molecular gaze” in the life sciences completely eclipse the physiological “molar body” (Rose 2007, 5), with its organic systems that had captured the minds, eyes, and hands of physiologists like Claude Bernard [1813–1878] or L.J. Henderson [1878–1942]? Did the rise of genetics in the twentieth century leave no room for physiology other than as a one-off required course for medical students? Or, did physiology persist as a scientific discipline throughout the twentieth century, somehow failing to attract the attention of most historians of twentieth-century science?

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One goal of this essay is to answer the question: What happened to physiology in the twentieth century? A second goal is to report on what scholarship on the history of physiology does exist, a humbling task given that the history of physiology, for some historians of science, has meant the history of the study of “life;” for others, something akin to the history of Western medicine; and still for others, the history of experimental investigations of vertebrate organisms only after 1800. This essay accommodates, albeit incompletely, all of these definitions in a series of roughly chronological, but overlapping, sections. The only real epistemic foothold of this account is that the historians themselves identify their work as the history of “physiology.” First, histories of physiology-as-the-study-of-life-itself are unabashed catalogues of “great men” and “great discoveries,” from Aristotle [384–322 B.C.] to Andreas Vesalius [1514–1564] to Carl Ludwig [1816–1895]. These “progressivist” intellectual histories (~1920s–1970s) track mostly Western philosophical, anatomical, and experimental investigations into the source, location, and mechanisms of “life.” The historians of physiology represented here, with their sweeping catalogues and often unsuppressed admiration, bring ambition and humility to their task. Next, histories of physiology-as-an-independent-science explore whether and how nineteenth-century physiology became an “independent” discipline, independent either from anatomy, on the one hand, or from clinical medicine, on the other. Influenced by mid-century sociologists like Joseph Ben-David, historians of this approach (~ early 1940s–1990s) find themselves concerned centrally with questions of “national styles,” “schools,” and “institutes.” Within this approach, a focus on quantifying and describing the careers, institutions, and networks of nineteenthcentury physiologists is paramount, as is figuring out what nineteenth-century physiologists did with the soul. While this scholarship shines brightly on France and Germany, a smaller scholarly conversation erupts over the question of why British physiology “stagnated” during the nineteenth century and, to a lesser extent, what happened in the USA. Two other approaches became dominant in the 1980s and continue into the present: economic and political histories of physiology in particular times and places and histories of the spaces, instruments, or organisms of physiological experiment. The conclusion explores a small collection of histories of physiology-after-1940, attempting to answer the simultaneously historiographical and historical question of what happened to physiology in the twentieth century. This essay is not, cannot be, comprehensive, and I look forward to correcting oversights and including more scholarship in the years to come. Even with this caveat, there are two serious, specific limitations of my approach. First, this essay is not comprehensive with respect to the biographies of individual physiologists; there are too many. Second, due to my own limitations, I fail to review the scholarship of fellow historians of science who publish in languages other than English. English-language scholarship focuses on the work of European or American physiologists, a notable exception being the body of scholarship by historian Marcos Cueto, who for three decades has documented the history of

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Latin American physiologists. This language limitation amplifies the broader issue of how one defines the history of “physiology” to begin with. English language histories of non-Western ideas about the processes and mechanisms of life abound, from histories of elite and lay medicine or cosmologies to histories of botanical knowledge to histories of “the body” – but these histories are rarely framed by historians as histories of physiology, per se, in part because “physiology” or “physiologist” is either an inaccurate or unnecessary translation of the concerns or actors represented in primary sources. While there’s a tension in any historiographical essay produced by the recursively fragile and fictive linguistic, epistemic, and political bounds of its subject, this tension saturates the historiography of physiology perhaps more deeply than other sciences. One of the features of the current volume is that it will be available online, with the potential to be updated regularly. I would welcome coauthors who read in other languages to add to this limited account. Physiology, even if defined narrowly as a scientific discipline with origins in nineteenth-century France and Germany, became increasingly transnational over the course of the twentieth century. One can wonder whether the tendency for physiology-after-1940 to “remain virtually unexplored,” to appropriate Foster, is connected to its “transnationality.” But, first, the catalogues.

Physiology-as-the-Study-of-Life-Itself: Histories as “Catalogues of Physiological Discoveries” (~1920s–1970s) What we know and what we think is not a new fountain gushing fresh from the barren rock of the unknown at the stroke of the rod of our own intellect, it is a stream which flows by us and through us, fed by the far-off rivulets of long ago. – Michael Foster (1901, 1), Lectures on the History of Physiology during the Sixteenth, Seventeenth, and Eighteenth Centuries.

Before there was “biology,” coined in the nineteenth century, there was “physiology.” In the sixteenth century, French physician Jean Fernel [1497–1558] used the word physiologia, from the Greek physis, or “nature,” to connote the study of the nature of animal life (Nutton 2012; Olmstead 1944; Tansey 2013). The earliest histories of physiology made much of this linguistic lineage, tracing a continuous line from ancient studies of physis through to what the authors considered subdisciplines of modern physiology, such as endocrinology, cardiology, and neurology (Brooks and Cranefield 1959; Debru 1995; Foster 1901; Franklin 1949[1933]; Fulton 1930; Goodfield 1960; Hall 1969; Hodgkin et al. 1977; Mendelsohn 1964; Rothschuh 1973[1953]; Singer 1957[1925]; see also Duffin 2010[1999]; Tansey 2013[1993]; West 2015). Traversing centuries, these histories recount scholarly answers to questions like “What is life?” and “How do bodies maintain life?” These catalogues of physiological discoveries invariably include celebrated scholars like Aristotle [384–322 BC], who located life in the heart; Galen [AD 129–c.200], who described animal life as the result of a “vital spirit” that

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mixed with liver-generated blood during breathing; William Harvey [1578–1657], who studied the circulatory systems in a wide range of animals to conclude that continuously circulating blood maintains life in organisms; and von Haller [1708–1777], who defined all life, plant and animal, as being able to perceive (“sense”) and to respond (“irritate”). These histories of physiology, which take investigations of “life itself” as their subject, loosely consider “anatomy” to mean the study of the structures of life and “physiology” the study of the processes of life, but in practice these histories are not overly concerned with distinguishing between natural philosophy, anatomy, and physiology – especially for time periods before 1800. In his A Short History of Anatomy and Physiology from the Greeks to Harvey (first published in 1925 and then republished in 1957), pathologist-historian Charles Singer, for example, suggested that Vesalius’s anatomy was at the same time physiology, “essentially living anatomy.” Singer continued, “The parts [in Vesalius’s drawings] are not the subject of morphological treatment, but of investigation as contributing to the existence of that complex vital unit we call a Man” (Singer 1957, 116). Most historians who catalogue physiological discoveries prioritize studies of the processes of life in animals over studies of the processes of life in plants. (One exception is histories of respiratory physiology, e.g., West 2015; for a catalogue of discoveries in plant physiology, see also Pennazio 2005a, b). The emphasis on the history of animal physiology derives from the fact that the earliest histories of physiology were narrated by physiologists themselves, whose own teaching (though not necessarily their research) was oriented toward medical students. While the nineteenth-century German university system (which would become the model for many university systems in Europe and beyond) encouraged physiological research that was somewhat independent of direct practical applications to medicine, still, in Germany and elsewhere throughout Europe, it was medical students who filled physiologists’ lecture halls, worked in their laboratories, and justified their salaries (Latour 1992; Lenoir 1992; Olesko 1988; Warner 1980). Michael Foster’s Lectures on the History of Physiology (1901) are a series of lectures he gave to Cambridge medical students. Opening with the printing of Vesalius’ Fabrica Humani Corporis in 1543 and providing a lively chronicle of biographies central to physiological discoveries thereafter, Foster’s Lectures laid out a basic, continuous timeline of sixteenth- to nineteenth-century physiology-as-the-study-oflife-itself that remained relatively unquestioned for most of the twentieth century. In 1930, self-consciously supplementing Foster’s Lectures, Yale physiologist John Fulton compiled an ambitious “source book” of (translated) German, French, Latin, and English physiological tracts, mostly from the seventeenth to nineteenth centuries. Just over a decade later, self-consciously supplementing what he considered the English- and American-leaning histories of Foster, Fulton, and others, physiologist Karl Rothschuh repeated the basic story (including 775 individuals!). Beginning with his chapter on the Enlightenment, Rothschuh also noted the national context of an individual’s ideas. In doing so, Rothschuh called, in particular, for more attention to nineteenth- and twentieth- century German (and, to a lesser

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extent, Russian, Scandinavian, and Japanese) physiology and to the impact German physiologists had on the development of British and American physiology. Translated into English in 1973 by physician-historian Guenter Risse, Rothschuh’s catalogue of physiological discoveries became an “obligatory passage point” for the next generation of historians of physiology (though not without critique, see Bonah 1995). Physician-historian Jacalyn Duffin’s (2010) chapter on the history of physiology, “Interrogating Life,” brilliantly condenses all of this scholarship, putting into print a popular lecture she gives to her medical students, just as Foster did a century before. Two specific histories add significantly to this cataloguing of life itself. Mary Brazier’s (1988) A History of Neurophysiology in the Nineteenth Century carefully chronicles the great discoveries of 59 European physiologists, from Galvani’s finding in 1791 that muscles conduct electricity to various proposals for electrotherapy at the end of the nineteenth century. This monograph is worth spending time with for the reprints of physiologists’ original drawings alone. Second, in 1964, Everett Mendelsohn published Heat and Life. Focused on exploring theories of the source of animal heat from the ancient Greeks to nineteenth-century physicochemists, Mendelsohn was less concerned with tracing a continuous lineage and more concerned with investigating scientific change. Although framed as a history of “biology,” the text is chockfull of “physiologists,” “physiological explanations,” and “physiological phenomena,” which is consistent with other mid-twentieth-century histories of physiology that use the terms “biology” and “physiology” interchangeably (Any attempt to answer the question “what happened to physiology-after-1940” inevitably must wrestle with this discursive overlap). These two intellectual histories, similar in their goal of cataloguing studies of “life itself” but so different in focus, suggest that an essay on the historiography of physiology would maybe make more sense if it defined physiology more narrowly. One can imagine a different essay that excludes histories focused on time periods prior to 1800. Yet that would be a mistake. One recent history of physiology-as-the-study-of-life-itself doesn’t even consider the nineteenth century. The product of a 2009 interdisciplinary symposium on studies of bodily processes and functions from antiquity through the early modern period, Blood, Sweat, and Tears (Horstmanshoff et al. 2012) shows how scholars’ studies of bodily fluids and spirits were mediated by contemporaneous literary and artistic tools, scientific instruments and technical processes, and ideas about bodies and health. Bringing together the history of science, history of art, and comparative literature, the chapters illustrate how ideas about the functioning of living organisms borrowed from contemporaneous motifs and shared experiences (e.g., features of weather; every day activities, like making cheese; and objects, like sponges or houses). Blood, Sweat, and Tears, which wins for best title, challenges simplistic assumptions of continuity in the history of physiology. However, it also challenges blunt, exclusive periodizations or definitions of physiology that leave out studies of life prior to 1800.

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Physiology as “an Independent Science”: Histories of Institutes, Schools, and “National Styles” (~1940s–early 1990s) A glance at modern histories of the discipline of physiology or of experimental physiology—the two terms are confusingly used interchangeably nowadays—will reveal that historians agree that it was created new as a discipline in the very early nineteenth century in France. But they also, and at the same time, believe that it has a very long history which it is their duty to trace, and they take their stories back over twenty or more centuries. Both these stories cannot be true, at least not in the same sense. – Andrew Cunningham (2002, 636) “The Pen and the Sword.”

In 2002, historian of medicine Andrew Cunningham took stock of the history of physiology and argued that by tracing long histories of physiological discoveries from antiquity through the early twentieth century, historians of physiology-as-life-itself had been comparing apples and oranges. First, historians had been failing to appreciate a key distinction and relationship between anatomy and physiology prior to 1800: the anatomist, Cunningham, asserted cut while the physiologist thought. Physiology, that is, was “the speculative wing of anatomy” (King 2012, 20). More important to Cunningham, because of this oversight, historians had been failing to see that there was an “old physiology” and a “new physiology.” Before 1800, Cunningham explained, there were two kinds of “old” (thinking) physiology: natural philosophy and theoretical medicine. Through the sixteenth century, most answers to the question “What is life?” were provided by natural philosophers, who derived their ideas about the functions of organs, the movements of fluids, and the currents of spirits from observation, speculation, and reason. Then, in the seventeenth century, scholars’ studies of the nature of life began to be applied more toward understanding life processes in animal organisms, in particular (Fernel’s physiologia). These scholars, who identified as physiologists in the seventeenth and eighteenth centuries, drew upon anatomical investigations, as well as on studies of mechanics and optics, to participate in the “institutes of medicine” then taught in European universities. “Old physiology,” in this sense, was a medical discipline in the seventeenth and eighteenth centuries, Cunningham emphasized, but “not an investigative discipline, nor an empirical discipline, nor an experimental discipline. It was, by contrast, a thinking and talking discipline—a discourse” (645). If “old physiology” relied upon “the pen,” Cunningham suggested, “new physiology” relied upon “the sword.” That is, by about 1800, physiologists answered the questions “What is life?” and “How do bodies maintain life?” by cutting into living animals and studying the respiratory, nervous, digestive, and circulatory processes they encountered, as well as by studying the effects of the alterations physiologists themselves sometimes intentionally introduced into animals’ bodies. Around 1800, though many of the questions, facts, foci, and even techniques of physiology might have been centuries-old (again, see Blood, Sweat, and Tears, as well as Wolfe 2013), physiology became, first and foremost, experimental, laboratory-based, and material. On this, historians of physiology agree. So, whether or not historians of physiology trace their histories back to antiquity, whether or not historians of physiology emphasize Vesalius’ “living

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Anatomy” or Haller’s “animated anatomy” (i.e., “physiology”), and even whether or not historians flatly disagree with Cunningham and consider Galen’s or Harvey’s studies “experiments” (or the studies of other pre-1800 physicians, like Santorio [1531–1636], who slept and ate in a “balance chair” in order to measure his own ingestion, excretion, and metabolism, or de Graff [1641–1673] who, in the seventeenth century, created a fistula in a dog to study the properties of pancreatic juices), historians of physiology, regardless of approach, agree that during the nineteenth century, “physiology” became a thoroughly experimental science. (Whether, for physiology students, experiment meant participating in laboratory experiments or observing demonstrations has been debated; see Schmidgen 2004.) Historians also agree that it was around 1800 that physiology started becoming an “independent science” (Olmstead 1944). Physiology became independent from anatomy and (even though nineteenth-century physiologists had medical degrees and taught medical students) from practical medicine. Neurochemist-historian Tilli Tansey (2013) has summarized the ways in which physiology became “independent” as: “the creation of full-time research and teaching positions; the provision of adequately equipped institutions for research and teaching; the establishment of professional journals and societies; and wider opportunities for travel” (126). From roughly the 1940s through the early 1990s, nineteenth-century experimental physiology attracted the interest of scores of historians of science and medicine who drew primary source material from physiologists’ publications, personal notes, and correspondences, as well as the proceedings of newly formed scientific societies. Influenced by mid-twentieth-century sociology of science, these historians used their studies of the disciplinary formation of “independent” physiology to understand the history of science more generally. This approach to the history of physiology, then, focuses not only on narrating individual scientists’ accomplishments but also on quantifying how individual scientists trained and otherwise influenced others, such as through demonstrations or through the creation of disciplinary tools like journals, textbooks, and societies (Appel 1987; Borrell 1976; Butler 1988; Coleman 1985, 1988; Debru 1995; Frank 1987; Fye 1987b; Geison 1987a; Holmes 1963; Lenoir 1982, 1988, 1992; Schiller 1968; Werner and Holmes 2002). This approach also explores how nineteenth-century physiologists developed research “schools” (Amaral 2006; Geison 1978, 1981; Holmes 1988; Lipman 1967;), established “national styles” (Geison 1978; Larson 1979; Temkin 1946), and secured institutional support (Coleman and Holmes 1988; Fye 1987a, b; Gross 1979; Kohler 1982, 1985; Kremer 1992; Lesch 1988; Nyhart 1987; Olesko 1988; Sturdy 1992; Tansey 1992; Tuchman 1988, 1993; Turner et al. 1984). Chief among the concerns of historians of physiology-as-an-independent-science is how physiology related to other nineteenth-century disciplines, including anatomy (Nyhart 1987), chemistry (Gilman 1959; Holmes 1963; Schiller 1968; Werner and Holmes 2002), zoology (Gross 1979), clinical medicine or pathology (Geison 1979; Laszlo 1987; Maulitz 1987; Sturdy 1992; Wilson 1984), and natural history or biology more generally (Farber 1982; Maienschein 1987).

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To be clear, this institutional, sociological approach has been applied to periods outside of the mid-nineteenth century. In Harvey and the Oxford Physiologists (1980), Robert Frank describes the networks of physiologists he categorizes as “major scientists” (including William Harvey), “minor scientists,” or “virtuosi,” all associated with Oxford from the 1630s to the 1680s. Moreover, several examinations of the emergence of physiological subdisciplines in the early twentieth century take up concerns about the roles of institutional support and professional competition (Kohler 1982, 1985; Long 1987; Morgan 1990; Pauly 1987; Sturdy 1992). Nonetheless, the rest of this section briefly summarizes consensus on the rise of the discipline of physiology in the nineteenth century and then highlights two additional trends within this approach.

Bernard and Ludwig Histories of physiology-before-1800 touch down all over Europe – Greece, France, Italy, Germany, the Netherlands, England, and more. Histories of nineteenth-century physiology, in contrast, draw a map with two poles: Paris and Leipzig. In 1816, French hospital physician François Magendie [1783–1855] published his textbook of experimental physiology, describing animal experiments and the kinds of knowledge about the life processes of nutrition and sensation that they produced (Albury 1977; Lesch 1984; Pickstone 1981). Magendie’s assistant, Claude Bernard, elaborated Magendie’s experimental approach and revolutionized French physiology. Bernard’s techniques in animal surgery, experiments on the “vital processes” of “internal secretions” (digestion and metabolism), and lectures and laboratory demonstrations on animal organs and organ systems drew students and visitors from across Europe in the 1850s and 1860s. Along with developing experimental techniques – Bernard is the most famous “vivisector” in the history of physiology – Bernard became (posthumously) well known for his concept of the milieu intérieur (considered by historians and physiologists to be a precursor of American physiologist Walter B. Cannon’s [1871–1945] theory of “homeostasis.”) Bernard posited that living organisms (both animal and plants) balance their milieu extérieur and with their milieu intérieur, “the circulating organic liquid which surrounds and bathes all the tissue elements” (Bernard, quoted in Fulton 1930, 308; see also Holmes 1974). In mid-nineteenth-century France, institutional support for physiological research came from hospital medicine and veterinary schools, explaining in part why Bernard’s experiments centered on the study of animal (including human) life. Nonetheless, Bernard argued that the study of life involved being able to directly intervene in physiological processes, which required, he maintained, not a hospital nor a morgue but a laboratory (Coleman 1985; Lesch 1984; Schiller 1968; Tansey 2013). For Bernard, clinicians and pathologists were mere observers who “faced the vital phenomenon without arms” (Coleman 1985, 54). Bernard’s own laboratories were actually small and at times self-funded; he received imperial support for his research only toward the end of his career in the 1860s.

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Mid-century French physiologists had nowhere near the amount of money nor space nor personnel to conduct experiments as German physiologists (Lesch 1988). Historians of physiology-as-an-independent-science investigate the intellectual and institutional factors that led to the growth of German physiology between 1830 and 1870, including the creation of (and competition for) full professorships that promoted physiological experimental research in independent laboratories (Ben-David and Zloczower 1962, 54–57; Kremer 1992; Turner et al. 1984). The Investigative Enterprise (Coleman and Holmes 1988), for example, describes how several German states in the nineteenth century funded independent physiological institutes to support experimental research into the basis of life. Chapters in The Investigate Enterprise interrogate the origins of the German “research ideal” (Coleman and Holmes), the development of the first physiological institute in the Prussian city of Breslau (Coleman), the institutionalization of scientific education in the state of Baden (Tuchman), and the life of the Munich “school” of research on metabolism (Holmes). Published 4 years later, The Laboratory Revolution in Medicine (Cunningham and Williams 1992) also explores the scientific “institutional revolution” in nineteenth-century German states (Lenoir 1992, 16). For historians of nineteenth-century experimental physiology, only Carl Ludwig’s influence rivals Bernard’s (see Lenoir 1982, 1988). From his Physiological Institute at the University of Leipzig (established in 1865), Ludwig applied physical and chemical laws to investigate blood chemistry and the cardiovascular system, as well as renal and respiratory physiology. For Ludwig, physiology meant applied physics and applied chemistry. With instruments like his kymograph, a device that registered traces of blood pressure or muscular contraction over time, Ludwig pioneered the “graphical method” of physiology. Historians of physiology-as-an-independent-science trace how Ludwig’s physicochemical approach and his graphical method spread through physiologists’ networks across Germany, France, Russia, England, and the United States. Ludwig’s colleagues (members of the Berlin Physical Society) included Emil du Bois-Reymond [1818–1896], Ernst von Brücke [1819–1892], and Hermann von Helmholtz [1821–1894], all influential physiologists who fused mechanics, optics, and thermodynamics with their studies of the neuromuscular system, digestion and metabolism, and vision (Kremer 1990). Nineteenth-century German physiology is often characterized as being more reductionist than French physiology; nonetheless these physicochemically oriented physiologists worked within a “gesamte Physiology,” or “a discipline which formed a comprehensive and unified whole, in order to give a coherent picture of the animal machine” (Morgan 1990, 496).

Two Trends: Vitalism/Mechanism and National Stories Exploring the institutional contexts of physiology after 1800, historians of physiologyas-an-independent-science concentrate on two themes: vitalism/mechanism and national stories. In 1946, physician-historian Owsei Temkin concluded that nineteenth-century French and German physiology experimentalists were equally materialist and equally able to answer the question of “what is life?” without requiring a spiritual concept, such as “the soul.” Nonetheless, Temkin argued, French physiology remained more “vitalist”

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because of the emphasis on explaining life as a phenomenon of organisms that live within but are categorically distinct from the inorganic world. Germany physiology, in contrast, Temkin considered more “mechanist” because of the emphasis on explaining life as the product of physical and chemical properties and forces, mechanisms that apply to both organic and inorganic matter. Responding to Temkin, historians of nineteenth-century physiology have defined and redefined the concepts of “vitalism” and “mechanism,” debating who or what – and when – was vitalist (or “neovitalist”) or, conversely, whether the vitalism/mechanism distinction seems overblown (Brown 1974; Coleman 1971; Daston 1978; Galdston 1959; Goodfield 1960; Goodfield-Toulmin 1969; Hall 1969; Larson 1979; Lipman 1967; Mendelsohn 1964, 1965; Geison 1987; Rosenberg 1995; Schiller 1968; Sloan 1977; see also Schäfer 2012; Wolfe 2013). Here, historians of physiology draw as much from philosophers of science as they do from sociologists of science. Along with paying close attention to what nineteenth-century physiologists did with the soul, historians of physiology-as-an-independent-science define their histories by national context, as the discussion of “French” and “German” physiology already suggests. For example, taking stock of Japanese physiology in 1965, Chandler McC. Brooks, Kiyomi Koizumi, and the Committee on the History of Japanese Physiology compiled a brief history of physiology-as-an-independent-discipline in Japan, alongside a survey of Japanese physiologists’ research interests and institutional locations (Brooks 1965). From the late 1960s through the early 1990s, a number of historians of physiology, in light of the French and German poles of physiological research in the nineteenth century, asked “what happened?” to British physiology. They asked why (and to what extent, really) British physiology “fell” in the eighteenth century and then “rose” again at the turn of the twentieth century (Borrell 1976; Brown 1974; Butler 1988; French 1971; Geison 1978; Goodfield-Toulmin 1969; Guerrini 1985; Tansey 1992). As Stella Butler (1988) put it, “In 1860, experimental physiology was virtually unknown in Britain. . .By 1900, however, the physiology departments of Cambridge, University College London, and Oxford had become internationally recognized centers of excellence” (473). Butler explored this dramatic shift by comparing the histories of six British physiological departments and quantifying which factors institutionalized successful research “schools.” Butler (and others) described how British medical students went to Paris to study under Claude Bernard and to Leipzig to study under Ludwig and then, from 1870–1900, how Scottish anatomist-physiologist William Sharpey [1802–1880] established modern physiology at the University College of London, where he taught Michael Foster and others. Geison’s (1978) biography of Foster documents how Foster then trained a whole “school” of students at Cambridge to conduct research into the genesis and regularity of the heartbeat and to publish their research in scientific journals. Social and political support for the antivivisection movement provides one answer to the question of “what happened?” to British physiology in the nineteenth century. Experimental physiology developed in France due to, not in spite of, Bernard’s reliance on animal vivisection (Lesch 1988). In contrast, a strong antivivisection movement initially “stagnated” British physiological research in the nineteenth century. By the end of the century, though, British physiology was well

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defined and well respected, in part because physiologists had been forced to defend and clarify their experimental methods in light of an 1875 Royal Commission on Vivisection and subsequent 1876 Cruelty to Animals Act (French 1971; Richards 1992). It was in this defensive context that Foster formed The Physiological Society (1876), started the Journal of Physiology (1878), and co-founded the International Physiological Congresses (1889) – elevating British physiology. In 1987, coinciding with the centennial anniversary of the American Physiological Society (APS), a cluster of historians of physiology-as-an-independent-science also turned their attention to the United States. Late nineteenth-century physiological institutions, networks, and lineages within “the American context” received a flurry of attention (Benison et al. 1987; Brobeck et al. 1987; Fye 1987a; Geison 1987; see also DuBois 1950; Geison 1979; Warner 1980, 1986). A decade prior, Edward Atwater (1978) had documented the institutional and professional identities of American experimental physiologists through the Civil War, and Geison (1979) had questioned whether American physiology became “independent” because of institutional support from reform in medical education, as it had in other countries. Geison asked, in particular, whether American physiologists claimed practical clinical applications of their research (and whether practicing clinicians agreed), a theme picked up by several contributors to Physiology in the American Context, 1850–1940 (Geison, 1987; see also Warner 1986, 235–257). Both Alejandra Laszlo and Jane Maienschein investigated whether – and at which institutions – American physiology was a part of medicine or “a broadly biological discipline” (Laszlo, 68). Robert Frank showed that over half of the leading American physiologists of the 1880s had studied in Leipzig earlier in the century and brought back “the laboratory ideal,” and Bruce Fye (1987a, b) credited the institutionalization of the German “full-time faculty system” and “research ethic” in medical schools for the development of physiology as a discipline in the United States. Historians of physiology still map intellectual networks, historicize institutions, and measure physiologists’ influence nationally or internationally. For example, Isabel Amaral’s (2006) account of the history of late nineteenth /early twentieth-century physiology in Portugal applies the concept of a “research school” to the life and work of Mark Athias [1875–1946]. Kari Tove Elvbakken (2018) documents the institutionalization of physiology (and, later, nutrition science) in Norway between the 1840s and the 1920s. Galina Kichigina’s (2009) The Imperial Laboratory describes the institutional forms and networks of influence of nineteenth-century Russian experimental physiology. To the extent that Kichigina attributes the increase in Russian imperial funding for physiology to dramatic political, economic, and social reforms following the devastation of the Crimean war, The Imperial Laboratory also reflects a third approach in the history of physiology. In the next section, I describe how some historians approach the study of physiology as a way to study broader social context and change.

Physiology-as-Society: Political and Economic Histories of Physiology (~1980s–Present) In the mental life of the nineteenth century, work was at the center not only of society but of the universe itself. Social modernity, the project of superseding class conflict and social

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disorganization through the rationalization of the body, emerged at the intersection of two broad developments: the thermodynamic ‘model’ of nature as labor power, and the concentration of human labor power and technology of the second industrial revolution. – Anson Rabinbach (1990, 289, italics added) The Human Motor.

In his unique contribution to a 1959 edited volume on the history of physiology, physician-historian George Rosen did not “catalogue” physiological discoveries nor did he focus on the development of physiology-as-an-independent-science. Instead, Rosen argued that modern physiological knowledge about metabolism, including energy costs and exchanges, was rooted in two economic and technical features of nineteenth-century industry: “machine technology and rational business calculation” (1959, 245). Rosen claimed that physiological ideas about metabolism would be “unthinkable” without “a theory of the steam engine” (246). In addition, Rosen considered physiologists’ discussions of energy “costs” to draw from principles of accounting. Insofar as Rosen called attention to how the economic priorities of industrial capitalism shaped physiological knowledge, he anticipated one approach to the history of physiology that became more common in the 1980s: the history of physiology-as-society. Out of and alongside histories of physiology-as-an-independent-science emerged histories, such as Rabinbach’s The Human Motor (1990), which draw precise relationships between specific lines of physiological research and contemporary politics and economics. Historians like Rabinbach contend that political and economic life affords physiologists with language, metaphor, and inspiration, not least of which is social problems to solve. These histories show how physiologists borrow discourses, ideas, and tools from the political and economic contexts in which they work, as suggested by Rosen’s account of physiological research on metabolism. In the rest of this section, I first review a few examples of this political and economic approach. I then summarize the historical scholarship on the industrial context of nineteenth-century physiology, in particular, in which historians characterize physiology less as the study of life itself and more as the study of “work” or “nature as labor power.” I conclude highlighting two opportunities for further histories of physiology-as-society.

Histories of Physiology in Political and Economic Context: Some Examples One trend in this approach to the history of physiology is to describe how physiologists’ studies of the processes that constitute, maintain, and transform life borrow from their political views about the processes that constitute, maintain, and transform the state. John Pickstone modeled this approach in 1981 when he sought to understand anatomist and physiologist Marie François Xavier Bichat’s [1771–1802] ideas about tissue physiology. Pickstone showed that Bichat’s physiology reflected his tendency to “see the operations of wholes as bundles of separate functions carried out by constituents. . .There were no privileged levels, no ultimate units” (123). For Pickstone, Bichat’s “bureaucratic” tissue physiology borrowed from “the professional-bureaucratic ideal” of the state that was being publicly debated during an increasingly bureaucratized post-Revolutionary France.

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Historians of late nineteenth- and early twentieth-century specific subdisciplines of physiology also connect physiologists’ research with their politics. Steve Sturdy (1988) analyzed Scottish respiratory physiologist John Scott Haldane’s [1860–1936] scientific papers and political treatises. Sturdy described how, as part of the nineteenth-century Scottish philosophical idealist movement, Haldane publicly challenged political positions that society is merely an aggregate of individuals. Instead, Haldane outlined an “ethical realism which emphasized good works and social reform” – as well as active political citizenship (1998, 319–321). At the same time, Sturdy presented Haldane’s research on breathing rates, blood chemistry, and hemodynamics as demonstrating that respiratory processes respond to “the requirements of the organism as a whole” (1998, 328). Sturdy’s point, then, was that Haldane’s conclusions about the regulatory, holistic function of the respiratory system borrowed from his political views on the moral, holistic relationship between a state and its citizens. More recently, in his biography of du Bois-Reymond, Gabriel Finkelstein (2013) traces du Bois-Reymond’s life, studies with the great Prussian physiologist Johannes Müller [1801–1858], research into “animal electricity,” and political philosophy within the growth and upheaval of nineteenth-century Berlin. Finkelstein situates du Bois-Reymond’s scientific treatises and commitments to realism alongside his political engagement (and at times, cynical disengagement) with Prussian affairs. Late nineteenth- and early twentieth-century neurophysiologists and endocrinologists, too, borrowed language from social life to describe life inside organisms. In Inhibition (1992), Roger Smith explores late nineteenth- and early twentiethcentury neurophysiologists’ investigations of “inhibition.” Borrowing the term “inhibition” from economic and technical discourses, neurophysiologists’ investigations of mechanisms for maintaining physiological order intersected with broader concerns about mechanisms for maintaining political order and social control in industrializing Europe. Examining the history of endocrinology, Cheryl Logan (2007) traces the research of Austrian physiologist Paul Kammerer [1880–1926] from his laboratory experiments on rats to his hopes for the rebirth of Viennese society after World War I. She shows that Kammerer’s endocrinology, in particular his experiments demonstrating the transformative effect of environment on rat sexuality, supported his Lamarckian, anti-eugenicist, and pacifist political views. Kammerer’s “heat rat” experiments fueled his conviction that “genetic change through modification of the environment would be the basis for the ennoblement of the people and the progress of society” (Logan 2007, 710). Gregg Mitman and Anne Fausto-Sterling (1992) develop a similar argument about the political, social, and intellectual implications of the physiological research of American embryologist Charles Manning Child [1869–1954] during the first two decades of the twentieth century. Taken together, this scholarship suggests physiologists’ investigations of life borrow from political debates about the nature of social change as much from ideas about social stability. Historians of physiology-as-society specify further how physiologists borrow language for describing specific life processes or mechanisms from contemporaneous technical or engineering discourses. Projit Mukharji’s (2017) account of two schoolbooks on “human physiology,” introduced in Bengal in 1857 (the first decade

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of direct Crown rule), notes how descriptions of physiological processes borrowed from engineering infrastructure, apparatus, or tools. Both textbooks, Mukharji explains, drew upon the material culture of mid-nineteenth-century urban Calcutta to explain the mechanisms of human physiology. One textbook explained the circulatory system by referring a student to “the pumping station at Chandpal Ghat” (the heart) and “the humble mousetrap” (valves in blood vessels) and explained the muscular system by referring a student to “pulleys used to lift rafters” (577–578). Both textbooks depicted the nervous system, as the newly installed electric telegraph (578). Slava Gerovitch (2002) shows how Russian physiologist Ivan Pavlov [1849–1936] “regularly borrowed metaphors from contemporary technology,” calling “the digestive system a ‘chemical factory’” and comparing “the nervous system to a central telephone switchboard” (343). What’s most interesting about Gerovitch’s account is how he demonstrates that a subsequent generation of Soviet neurophysiologists rejected newer technological metaphors (the computer) out of a Cold War-fueled allegiance to Pavlov.

Histories of Physiology as the Study of “Labor” The biggest trend in the history of physiology-as-society centers on exploring how physiology emerged from, reflected, and responded to the changes wrought on urban life by industrial capitalism (Alexander 2008; Brain and Wise 1994; Blayney 2017; Derickson 1994; Di Guilio et al. 2006; Dierig 2003; Gillespie 1987; Lenoir 1992; Rabinbach 1990; Tuchman 1988; Wise and Smith 1989). Historians document the ways in which physiologists derived relevance from the political and economic contexts of living in rapidly industrializing cities. Arleen Tuchman (1988, 1993) explained just why one German state started providing institutional support for physiology research in the mid-nineteenth century. In the context of “a society responding to the early pressures of industrialization,” the Vormärz parliament in Baden funded physiology at the University of Heidelberg as part of its project of “creating a new type of citizen, and thereby a new driving force for the economy, and it was within this context that the scientific method and laboratory exercises grew in importance” (Tuchman1988, 66–67). Drawing on private correspondences as well as public philosophical and political treatises, Timothy Lenoir (1992), likewise, showed how physiology in German states “was self-consciously harnessed to the needs of a nascent, industrializing, capitalist economy” (16). Experimental physiology became institutionalized in medical education, not out of elites’ investment in knowledge for its own sake but rather as part of a vision of a new, professional, practically trained middle class open to technical change and capable of “rigorous methodical thinking” (16). Examining the careers of du BoisReymond, Helmholtz, and others associated with the Berlin Physical Society, Lenoir, like Tuchman, documented how moderate liberals held up laboratorybased science education as the key to ushering in a “distinctive German middle class” (18). German physiologists did more than reflect the priorities of industrial capitalism by teaching “rigorous methodical thinking” to increasing numbers of

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students. Historians of physiology-as-society also show how late nineteenth-century physiologists’ research participated in a science of “work.” What in France was science du travail became Arbeitswissenschaft in Germany and “industrial physiology” in Britain and the United States. Rabinbach’s groundbreaking The Human Motor (1990), for example, traces how the formalization of the laws of thermodynamics – that all natural forces (chemical, mechanical, electrical) add together to constitute a universal energy – enrolled experimental investigations of physiological processes into a “science of work” that was simultaneously physical, political, and economic. Thirty years on, Rabinbach’s thesis continues to bear fruit. Historians of physiology-as-society have demonstrated that physiologists like Helmholtz, Ludwig, and du Bois-Reymond studied life as work and nature as labor. Physiologists considered muscles as heat-producing engines and nerve fibers as telegraph wires, and across the physiological laboratories of Germany, Belgium, and France, experimental organisms became “thermodynamic machines.” For example, through a close reading of Helmholtz’s papers, publications, notes, and references, Robert Brain and Norton Wise (1994) demonstrate that, more than metaphor, Helmholtz quite literally borrowed tools, like self-registering “work-measuring” instruments and indicator diagrams, from Berlin’s civil and military engineers. Published shortly after The Human Motor, bringing together histories of technology, art, and science and drawing from newly available negatives, films, and letters, Marta Braun’s breathtaking biography, Picturing Time (1992), describes how French physiologist and inventor Etienne-Jules Marey [1830–1904] studied the “animal machine” by charting the movement of human and other animal bodies over space and time. Marey invented an astonishing number of instruments to graph and record animal movement: a myograph to measure muscle contractions, a sphygmograph to measure pulse noninvasively, a cardiograph to make tracings of heart activity, a chronophotographic gun to produce rapid fire photos (12 frames/second) of animals in motion, and numerous other instruments to inscribe animal movement. Marey intended his chronophotographic techniques and graphical methods, Braun explains, to help shed light on the mechanical requirements of different kinds of work and, ultimately, to help reduce worker fatigue and to improve the efficiency of human labor (Braun 1992, 320–348; see also Rabinbach 1990). Inspired by Marey and others, the Italian physiologist Angelo Mosso [1846–1910] sought to discover the thermodynamic laws of “fatigue” itself (Di Guilio 2011; Di Guilio et al. 2006). Reviewing scores of Mosso’s original publications, Camillo Di Giulio (2011) shows how Mosso intended his neurophysiological and respiratory research on fatigue to improve workers’ lives in an agrarian society that was increasingly industrial. According to Di Giulio, Mosso intentionally published his now classic La Fatica (1891) on the second anniversary of Labor Day, May 1, 1981. As the cases of Marey’s and Mosso’s interest in fatigue illustrate, these histories of physiology-as-society show that physiologists didn’t just borrow from industrial capitalism, they attempted to intervene in it. Proclaiming expertise on “the human motor,” physiologists positioned themselves to affect state policy on a number of crucial social issues of industrializing Europe and America. Weighing in on debates over the length of workday, wages, and productivity measures, physiologists’

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investigations of work proliferated well into the first two decades of the twentieth century (Rabinbach 1990; see also Alexander 2008). Leading the mid-1980s interest in American physiology, Richard Gillespie (1987) demonstrated that it was through the notion of “industrial fatigue” – at once socially relevant and conceptually flexible – that American and British physiologists successfully extended the boundaries of their discipline in the twentieth century. Alan Derickson (1994) examined how American Progressive Era corporate managers made use of physiologists’ insights. Mateo Munoz’s (2014) recent biography of American physiologist L. J. Henderson illustrates how, for Henderson, linking biochemical models of life with economic models of society was, in fact, his life’s work (see also Allen 1975; Parascandola 1971). Henderson co-directed one of the most important institutions in the history of American physiology, the Harvard Fatigue Laboratory [1927–1947], interestingly enough established within the Harvard Business School (on the Harvard Fatigue Lab, see Chapman 1990; Horvath and Horvath 1973; Oakes 2015; Scheffler 2011, 2015; Tipton 1998, 2014; and the 2015 special issue of the Journal of the History of Biology). Harvard Fatigue Laboratory physiologists received funding for their research from the Rockefeller Foundation, industries like the Corn Industries Research Foundation, and, during World War II, from the Office of Scientific Research and Development. Indeed, some histories of physiology-as-labor-power make the broader point that early twentieth-century physiologists’ questions and methods were shaped by the specific political and economic context of war. Steffan Blayney (2017) documents how British industrialists and politicians in the first two decades of the twentieth century invested in “industrial physiology” to increase worker and national productivity, particularly during World War I. In Blayney’s account, the object of physiologists’ inquiry, “life,” was fully reduced to “work” – even just one component of work, “maximum output,” and even just one particular kind of worker, munitions factory workers. Others have demonstrated how the “national stories” in the history of early twentieth-century physiology double as stories of military research on topics like the physiology of marching or of high-altitude respiration (Folk 2010; Gunga 2008; Kehrt 2006).

Two Opportunities: Gender and Race There are surprisingly few histories of physiology-as-society that emphasize gender, though there are enough to know we need more. In particular, we need to understand how elite, experimental physiology developed not only as an exclusive space for men, which many sciences did, but as a particularly misogynist branch of science. The neurophysiological concept of “inhibition” emerged from the discursive context of control and regulation in Victorian society, including control of “unruly” women (Smith 1992). The antivivisection movement in the United Kingdom – which put British physiologists on the defensive and in doing so precipitated the consolidation and professionalization of British physiology – was also a campaign against violence to women (Lansbury 1985, 83–129; Rose 1992, 332; on the politics of vivisection in

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American physiology, see Lederer 1995; Parascandola 2007). The scholarship of Toby Appel, historian for the APS, documents the persistence of physiology as a male-dominated field in the United States through the 1980s (1987). Appel’s (1994) study of American female physiologists between the 1860s and the 1930s draws on collections within the archives of women’s colleges to describe the careers and research programs of two generations of women physiologists. Appel shows how female physiology students responded both to the limited professional opportunities available to them by a male-dominated discipline and to a broader hygienic reform movement that constructed women’s bodies as both fragile and disruptive. Vanessa Heggie’s (2013, 2016) research on exercise physiology and environmental physiology (discussed more below) similarly points toward the gendered and classed constructions of “normal” physiology. And we can learn more about how lay people appropriated or contested the “laws of [gendered] life” being produced by physiological experts. Iwan Rhys Morus’ (2011) Shocking Bodies, more history of physics than physiology, provides an interesting roadmap for how to write the history of nineteenth-century physiology “from below.” April Haynes’ (2015) Riotous Flesh, which Haynes describes as “a prehistory of heterosexuality and liberal feminism,” examines how white and African American social reformers between the 1830s and 1860s used physiology “as a tool in their struggles against white and male supremacy” (25, 162). Only loosely appropriating contemporaneous experimental studies of digestion and nervous stimulation, Haynes’ “reform physiologists” formed lay physiological societies, solicited physiological “testimonials,” offered “cheap physiological lectures,” and shared ideas about “the laws of life” (6, 141), including the nature of “normal” female sexuality (on American antebellum popular physiology, see also Kohlstedt 1978, Rosenberg 1995). Historians of science could do more to problematize the gendered dimensions of the political and economic contexts of physiological research – not only in the popular lives of physiology but also in the experimental or elite academic lives as well. There are also surprisingly few histories of nineteenth- and early twentieth-century physiology-as-society that emphasize the racial, colonial, or imperial contexts of the production, appropriation, or circulation of physiological research – on labor or otherwise. Rabinbach (1990) mentions that the French Ministry of War in 1907 commissioned physiologist Armand Imbert to study “the working capacities of North Africa’s indigenous population,” after which Armand continued his physiological investigations of the work of filers, vine cutters, and glass polishers back in France (185). Again, though, the histories of physiology that have foregrounded these themes suggest there is more work to be done. Lundy Braun’s (2014) history of spirometry in Britain, America, and South Africa demonstrates that, since the nineteenth century, physiologists’ studies of respiration, including their statistical tools, have been saturated with assumptions about racial differences. With European and American industrial capitalism simultaneously imperial projects, historians who continue to research nineteenth- and twentieth-century physiology-as-society, even – perhaps especially – those who continue to emphasize physiology in Europe in the nineteenth and early twentieth centuries, could follow Braun’s lead in broadening the

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political and economic frames to consider the racial, colonial, and imperial contexts of European physiological research. Indeed, historians documenting the production, circulation, and appropriation of nineteenth- and twentieth-century physiological research beyond Europe and the United States leave little doubt that the history of physiology-as-society requires attending to the racial, colonial, and imperial (or antiracial, anticolonial, and anti-imperial) assumptions embedded in physiological investigations of life or labor (Anderson 2002). Ana Gomes (2012) describes how, in 1880, the French physiologist Louis Couty and Brazilian physician João Baptista de Lacerda looked to French physiology (i.e., vivisection, graphical methods) when building the Laboratory of Experimental Physiology of the National Museum of Rio de Janeiro. However, Gomes eschews any account of Brazilian physiology as merely derivative; on the contrary, Emperor Pedro II’s own imperial interests drove Couty’s and Lacerda’s research agenda. They studied the physiological effects of drinks and foods popular in Brazil – and increasingly profitable to export to Europe – such as herb mate, coffee, and dried meat (158–159). Gomes further shows how nineteenth-century Brazilian physiology developed in stark contrast to French physiology, comparing the epistemological status of the clinic to that of the lab in both Brazilian and French scientific communities. Gomes’ brief account of the ways the French elite contested Brazilian physiologist Lacerda’s claims about the efficacy of potash permanganate (for treating snake bites) demonstrates that geographically wide or complex stories of the circulation, appropriation, or contestation of physiological facts reveal as much as more local or geographically or socially constrained stories of the production of knowledge. Drawing on Rockefeller Foundation archives, Marcos Cueto takes institutes and nations as his units of analysis while at the same time remaining relentlessly attuned to transnational networks and colonial and nationalist imperatives. Cueto (1989, 1990, 1994, 2016) narrates the histories of the institutionalization of endocrinology, neurophysiology, and, above all, high-altitude and respiratory physiology, in Argentina, Mexico, Brazil, and Peru between 1910 and 1960. He describes how Latin American physiologists located their research within broader political conversations about indigeneity, authoritarianism, and the virtues or pitfalls of technical progress. For example, Cueto (1994) contextualizes the research of Argentinian physiologist, nationalist, and Nobel Laureate Bernard A. Houssay [1887–1971] within the political and economic history of Argentina in the twentieth century. Cueto shows how Houssay took advantage of the overcrowded classrooms of Argentinian universities in the 1920s and 1930s to create massive physiological experiments on metabolism that relied upon student labor and an “assembly line” approach to research that Cueto calls the monitores system. Elsewhere, Cueto (1989) highlights how Peruvian experimental physiologist Carlos Monge [1884–1970] participated in 1920s and 1930s indigenismo critiques of colonialism and elitism. Monge founded the Institute of Andean Biology (with Rockefeller support) based on his rejection of the notion that “normal” physiology occurred at sea level. “Where North American aviators ask for oxygen,” Monge chided, “Peruvians play soccer” (quoted in Cueto 1989, 647). Building on Cueto’s

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work, Jorge Lossio (2008) concurs that the development of physiological research in the central high Andes contributed to Peruvian scientific nationalism. At the same time, Lossio’s account of the physiological research conducted under the auspices of the Peruvian Ministry of Health and the Pasco Mining Corporation (the largest employer in Peru by the end of World War II) depicts physiology as an apology for industrial capitalism. Like vaccine research among “tropical workers” in South African gold mines in the first half of the twentieth century (Packard 1993), “high-altitude” physiology from the 1920s through 1950s both reinforced ideas of racial difference within Peru and provided an intellectual and discursive framework to attribute disease to maladaptation to “climate” rather than to the oppressive living and working conditions of the Peruvian copper and silver mines. In a final example of a recent history of physiology-as-society that attends to labor, race, and colonialism, consider Alison Bashford’s (2012) account of the physiology of early to mid-twentieth-century Indian economist and ecologist Radhakamal Mukerjee [1889–1968]. Addressing questions of overpopulation and global food insecurity, Mukerjee marshalled contemporaneous ideas of racial physiological difference toward anticolonial nationalist ends in the interwar period. Mukerjee argued that because white colonists were physiologically unfit for – and therefore could not labor in – “tropical” climates, their presence wasted potential agricultural productivity in many parts of the world (612–615). By contrast, he suggested, immigration laws should facilitate the movement of the more “adaptable” “Asiatic peoples” to large tracts of as-yet-unproductive land. There is ample room for more histories of physiology-as-society to chart just where, when, and how the study of life as gendered and racialized labor persisted or receded in physiology.

Physiology-as-Practice: Histories of the Instruments and Organisms of Physiological Experiment (~1980s–Present) From Max Weber to Jurgen Habermas to, most recently, Jean-Francois Lyotard, there has been a way of viewing the role of science in the modern world through the emergence and impact of rational skills, technical execution, or, in Lyotard’s language, of ‘performativity.’ This view encourages us to consider science not only in terms of its intellectual or technical products but also in terms of its activities: its training, its labor, and its workaday world in general. The largely unwritten history of these activities, all a part of the investigative enterprise of science, poses an intriguing challenge for the future. – Kathryn Olesko (1988, 324), “Commentary: On Institutes, Investigations, and Scientific Training.”

In her contribution to Physiology in the American Context, historian Merrilley Borrell (1987a) tells the story of Harvard physiologists Henry Pickering Bowditch [1840–1911] and William Townsend Porter [1862–1949]. In the 1870s, Bowditch returned from studies in Paris and Leipzig to Cambridge, where he established at Harvard a chemical laboratory, a space for microscope work, a table for experiment, and, as Bowditch’s student later recalled, “a tub for frogs” (296). Borrell charts how Bowditch expanded his laboratory in the 1880s to include room for six to eight students to study at once and to include a kymograph, respiratory apparatus, and a

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dark room. When Porter (who started the American Journal of Physiology) took over physiology at Harvard in the 1890s, he wanted all 208 of his students to be able to conduct laboratory experiments, leading to his investment in “reliable, well-made, inexpensive apparatus” (305), including kymographs, capillary electrometers, manometers, electrodes, and magnets. In 1904, Porter’s Harvard Physiological Apparatus spun off as its own company. Elsewhere, Borrell (1987b) explores the lives of the instruments of several mid-century physiologists: Ludwig’s kymograph, Marey’s multiple “–graphs,” and Helmholtz’s myograph. Borrell argues that experimental physiology became an “independent science” not only through the social and professional networks of physiologists and through the institutional support from governments or universities but also through the ways their tools traveled and shaped the experiences of their users. Recording instruments, in particular, transformed physiology, Borrell suggests, because they required students to practice and experience experimentation (rather than memorizing facts) and because they enabled physiologists to study multiple physiological events and systems simultaneously and over longer periods of time. By focusing on instruments, Borrell asks different kinds of questions of the historical record than approaches focused on cataloguing discoveries, mapping professional lineages, or contextualizing discourses. Borrell asks questions like, where did all those frogs come from? How easy was it to arrange for 208 student kymographs (or 104 if they worked in pairs) in Cambridge, given that at the turn of the century kymographs were only being made one at a time in Leipzig? How did constraints on production shape the tools available to study life? How can tracing material networks provide new insights into the history of physiology, not only into who physiologists were or what they thought but also what they did and the ways in which access to particular tools and materials shaped what could be known about life and who could know? Borrell’s research on physiological instruments and techniques reflected the “turn to practice” in the history of science in the 1980s and 1990s, so clearly called for by Kathryn Olesko in her contribution to The Investigative Enterprise (1988; see Landecker ▶ Chap. 12, “The Matter of Practice in the Historiography of the Experimental Life Sciences,” this volume). In the history of physiology, historians began to read old sources (physiologists’ publications and correspondences) in new ways. They also began to pay more attention to laboratory handbooks, students’ and assistants’ notes, and catalogues of scientific instruments as key sources. Chief among Borrell’s sources were the catalogues of the Harvard Apparatus Company. The history of physiology-as-practice received institutional support in the first decade of the twenty-first century from the Max Planck Institute for the History of Science. Hans-Jörg Rheinberger’s research group investigated “the history and epistemology of experimental practices,” one product being the Virtual Laboratory for Physiology (http://vlp.mpiwg-berlin.mpg.de/index_html). With a digital archive and collection of historical essays, the Virtual Laboratory remains an excellent resource for scholars interested in the practice of physiology from 1830–1930. Many historians of physiology, including Henning Schmidgen, Sven Dierig, Laura Otis, and Frederic Holmes, have been affiliated with this research group. Key to their method has been tracking how experimental setups change over time and what

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those changes in material practice can tell us about the sociocultural, institutional, and intellectual contexts of research, how each physiological experiment: is not a stable collection or a ‘thing’ in the etymological sense of the word, but a dynamic cooperation, a time-based assemblage involving the ‘together with’ and ‘against each other’ of artificial and natural organs as well as the sophisticated use of organic and nonorganic writing surfaces and styli. (Schmidgen 2005, 252–253)

(Otis’ (2007) intentionally Faulknerian biography of Johannes Müller is an interesting exception here in its emphasis on discourse over materiality.) Historians of physiology-as-practice from Borrell to Schmidgen examine when and by which instruments the “graphical method” took hold of experimental physiology (Brain 2008, 2015; Brain and Wise 1994; Braun 1992; de Chadarevian 1993; Frank 1988; Hankins 1999; Latour 1992; Lenoir 1986; Olesko and Holmes 1993; Schmidgen 2005; see also Hoff and Geddes 1959). What all these histories share is their refusal to take the development or the utility of the graphical method for granted. They want to understand how it made sense technically as well as within particular intellectual, cultural, and material contexts. Tracing the histories of instruments measuring the cardiovascular system between 1854 and 1914 (the kymograph, sphygmograph, and electrocardiogram), Robert Frank (1988) finds that attention to instruments and graphs answers old questions about the relationship between physiology “as an independent science” and clinical medicine. Frank shows that the instruments that traveled between physiology laboratories and clinical practices tended not to be those instruments that were simple to use but rather those that produced graphs that were simple to use. Soraya de Chadarevian (1993) situates Marey’s self-registering instruments within a broader disciplinary and cultural context. Marey’s innovation in producing selfregistering instruments, de Chadarevian suggests, can only be understood within a late nineteenth-century French cultural context obsessed with “the transformation and translation of different sensual qualities into one another” (279) and within a disciplinary context (again, in France) in which Marey had to find another way to study physiology if he didn’t want to participate in vivisection. Robert Brain picks up this theme in The Pulse of Modernism (2015), further exploring the impact of experimental physiologists’ graphical method and instruments on turnof-the-century European artistic and aesthetic theories and experiences. Brain’s is a story of technoscience in the service of art. Technologies of representation beyond graphs have their histories, too, from photography and projection to visual instruction. Schmidgen (2004, 2012) describes how late nineteenth-century and early twentieth-century physiological tools of visualization drew upon innovations in motion pictures, photography, lamps, and telegraphy, as well as on older architectural forms, like anatomical theaters or preparation rooms, adapted to newer forms, like Johann Nepomuk Czermak’s [1828–1873] “spectatorium” or Carl Jacobj’s [1857–1944] projection hall. Schmidgen’s work shows how enabling physiological students and publics to “visualize natural phenomena” required “highly artificial devices”

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(2004, 507) – such as Du Bois-Reymond’s “frog pistol,” which involved a recently severed, though still contracting, frog leg encased in a glass flask that could be held, “triggered,” and passed around during lecture. Along with focusing on histories of visualizing instruments and techniques, like the graphical method, other historians of physiology-as-practice have focused on the acquisition, breeding, and manipulation of experimental organisms (Dror 1999; Holmes 1993; Lederer 1992; Logan 2002; Richards 1986, 1992; Schloegel and Schmidgen 2002). For example, Adele Clarke (1987), studying early twentiethcentury physiology, argues that physiologists’ demands for live or “fresh” animals, rather than the dead animals of morphology, brought physiologists into highly coveted relationships with meat-packing facilities on cities’ outskirts, as well as with dog and cat pound owners, physicians who sent human embryos from miscarriages, or staffs at zoos. Historians of physiology-as-practice have contributed significant case studies of model organisms, from Planaria to frogs to dogs, to the history of biology (Holmes 1993; Johnson 2013a; Lederer 1992; Mittman and Fausto-Sterling 1992; Todes 1997, 2002). Some make the particular point that by not focusing on practice, previous historians of physiology took vivisection too much for granted. Analyzing a British laboratory methods textbook from 1873, Richards (1986) describes specific techniques of vivisection – excision, amputation, starvation, and the administration of anesthesia (or not) – and argues that the necessity to be cruel to animals presented a moral tension in physiology that was absent in other disciplines. Examining the laboratory notebooks of nineteenth- and early twentieth-century physiologists, Otniel Dror (1999) argues that physiologists selected animals for experiments depending upon assumptions about species’ emotions, such as rabbits as “sensitive,” cats as “angry,” and skunks as “fearless.” (This debate over animal emotions was certainly not new; in her 2009 book on the history of veterinary medicine, The Care of Brute Beasts, Louise Curth examines early modern ideas about specific physiologies of domesticated animals.) More importantly, Dror shows how entire research agendas in neurophysiology grew out of physiologists’ interest in trying to measure (and then to control for) these emotions of their experimental animals. Other historians find in the turn to practice, that is, in the close reading of laboratory notebooks, evidence of the emotions and aesthetics of the scientists themselves (Sattar 2013). A 2013 special issue of the Journal of the History of Biology uses close attention to the practices of vivisection to draw together experimentation across the sixteenth and nineteenth centuries. Most broadly, histories of physiology-as-practice show that local infrastructures generate not only the discourses or problems of physiological research but also its very material existence – not only instruments and animal organisms but also buildings and workers. At the same time as nineteenth-century European physiologists, for example, borrowed language from thermodynamics or tools from industrial engineering to study muscles as engines, some of their laboratories were factories (Latour 1992), complete with skilled and unskilled laborers, connections to city gasworks or waterworks, and mechanized work processes. Sven Dierig (2003) analyzes descriptions of experiment published by Bernard,

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Ludwig, and du Bois-Reymond, as well as laboratory notebooks and guidebooks for handling laboratory equipment, to construct detailed accounts of experimental practice. Dierig describes how Ludwig motorized different experimental setups in his labs by bringing in small engines powered through city gasworks and by advertising for assistants, whose job descriptions involved needing to be able to work with a variety of instruments and machines. By attending to practice, Dierig places the history of nineteenth-century physiology squarely within the history of cities. Daniel Todes’ account (1997, 2002) of Pavlov’s research on digestion embodies the history of physiology-as-practice. Analyzing laboratory notebooks, lectures, and publications, Todes shows that Pavlov’s laboratory at the Imperial Institute of Experimental Medicine in St. Petersburg was quite literally a factory. Todes describes Pavlov as an effective manager with a well-disciplined (military medical student) workforce. Todes shows how Pavlov’s factory style of production shaped his knowledge about digestion, future research interests, and commercial sale of gastric juices. This historical approach, then, is about getting at the infrastructures of physiology, the material infrastructures physiologists relied upon, and the new kinds of material, social, and economic relationships physiological research stimulated.

The Death of the Queen? Histories of Physiology-After-1940 A century ago, the discipline of physiology signified the search for causal explanations of body processes; by now that attitude is widely diffused through a range of disciplines which, in whole or part, might be regarded as physiology’s progeny. – John V. Pickstone (1990: 740-741) Although departments of physiology disappear or integrate, it does not mean that physiology is disappearing in teaching and research, but that it is less visible as a separate discipline.” – Westerhof (2011: 602).

Physiology did not die out after 1940. However, whether or not physiology as a discipline went into decline or flourished over the course of the twentieth century is a matter of perspective. Viewed in one way, physiology fragmented into “independent” biological sciences, including pharmacology, neurobiology, endocrinology, cardiology, psychology, and biochemistry. In the 1973 translation of his History of Physiology, Rothschuh appreciated the “impressive accumulation of details” stemming from “the gradual specialization of physiological research” (1973, 340). However he considered physiology “no longer a uniform and coherent field of investigation,” rather a field “undergoing in recent times a gradual process of fragmentation” (1973, 350). In 1990, John Pickstone observed that over the previous century across Europe and the United States, physiology’s “identity has become less clear, mainly through the differentiation of disciplines which were, at least in part, its offspring” (1990, 729).

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The American context is a case in point. “The first to peel off,” John Cook writes, “were the biochemists,” who formed the American Society of Biological Chemists in 1906, in turn becoming the American Society for Biochemistry and Molecular Biology 80 years later (1987, 436). That same year, 1906, the American Society for Pharmacology and Experimental Therapeutics formed. The American Association of Immunologists and the Association for the Study of Internal Secretions both formed a few years later (the latter becoming the Endocrine Society in 1952). By the 1940s, according to Geison, the number of new PhDs in physiology had declined both in absolute numbers and also relative to “new PhDs in other biological disciplines” (1979, 79). The Biophysical Society, the American Society for Cell Biology, the Society for Neuroscience, the Microcirculatory Society, and the Biomedical Engineering Society all “peeled off” from the APS in the 1950s and 1960s. At the same time, medical schools were requiring fewer and fewer hours of physiology instruction in lecture or in the lab, up to a 70% decrease from 1961 to 1971 (Warner 1980, 69). In the mid-1940s, the APS conducted a self-study to assess the state of physiology and concluded that physiology was experiencing an “inevitable but bewildering and doubtless deleterious fragmentation of physiology into subdivisions” (quoted in Kremer 2009, 359). The APS conducted other selfsurveys in the 1950s and concluded that physiologists were under “the threat of extinction” (Geison 1979, 80). Addressing the question of “the disappearance of physiology?” in 1990, historian of physiology Richard Kremer quoted a 1990 APS “White Paper on the Future of Physiology,” in which the Long-Range Planning Committee of the APS concluded “Physiology as a science and a profession does not in fact exist” (Kremer 1990, 361). Investigating physiology-as-an-independent-discipline toward the end of the twentieth century, Kremer synthesized quantitative data from American Men and Women of Science, the National Research Council, the National Academy of Sciences, and the APS, to demonstrate that relative to other fields, physiology doctorates had declined since the 1970s (361–366). Viewed differently, however, the twentieth century brought an “explosion of physiological sciences” (Tansey 2013[1993], 136). In 1950, physiologist Eugene Dubois considered the growth of physiology unbridled, even “acromegalic” (6). In the middle decades of the twentieth century, some physiologists’ work indeed became molecular, shifting foci “from that of the whole animal—vis a physiological system—to a specific organ or tissue, down to the cell, sub-cellular component, or molecule” (Tansey 2013[1993], 131). However, the molecular gaze was productive; the physiological approach to the study of life – experimental, mechanistic, and functionalist – became “widely diffused” across the biological sciences (Pickstone 1990, 741); physiology became a kind of “supradiscipline” (Kremer 1990, 366). So argues Tilli Tansey, historian for the Physiological Society, who examined research topics that earned Nobel Prizes in Physiology and Medicine during the twentieth century, attributing 39 Prizes to physiology – “even if they did not consider themselves primarily to be physiologists” (2013[1993], 136). Tansey concluded her catalogue of physiological discoveries (that started in antiquity), claiming “the physiological approach as the study of the function of living matter, has retained its identity, even if the boundaries of the subject itself are

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becoming increasingly indistinct” (150). Physiology “exploded” after 1940, though dispersing in the process. And what of the historiography of physiology-after-1940? For the most part, historians have followed suit, with the history of physiology-after-1940 fragmenting in one way and exploding in another. On the one hand, each of the offspring biological sciences has attracted its own historians, with explicit discussions of “physiology” fading quickly. On the other hand, the history of molecular biology and, in part, the history of the life sciences (see ▶ Chap. 9, “The Historiography of Molecular Biology” in this volume) are the history of physiology-after-1940. That being said, a small but growing number of histories of physiology construct a potential third narrative for the history of physiology-after-1940, a narrative where some midtwentieth-century physiologists carved out institutional and intellectual spaces to study the processes of animal and plant life in diverse conditions and environments.

Histories of Physiological Bodies in Place Robert Meurnier and Kärin Nickelsen (2018) suggest that the narrative of the history of twentieth-century biology need not only be “as a period that pushed reductionism to its limits” (9). In the case of physiology, alongside “the molecular gaze,” “the molar body” may have lived on in specific, peripheral disciplines for much of the twentieth century and even into the present. Some historians of physiology-after-1940 have documented the questions, spaces, and tools of plant physiology and of environmental and exercise physiology. These are histories of physiological bodies in place (Bashford 2012, 623). As indicated at the outset of this review, plant physiology rarely receives direct attention from historians of physiology. While many physiologists before and after 1800 who attempted to answer the questions “What is life?” and “How do bodies maintain life?” themselves studied plant as well as animal physiology, most historians of physiology highlight research on animal physiology. In addition, recent histories of twentieth-century plant biology and agriculture highlight the history of plant morphology, ecology, and genetics, but not physiology (e.g., Phillips and Kingsland 2015). Nonetheless, David Munns (2015) has pointed to the rise of plant physiology in the twentieth century. Most intriguing to me is the way Munns recounts the confidence plant physiologists had in their discipline and in their tools to, in their words, understand “the organism as a whole” and to develop “an integration of the parts rather than a cataloguing of them” (quoted in Munns 2015, 33). The holism invoked by these mid-century plant physiologists resembles the “organismic holism” described by Charles Rosenberg (2007) in his classification of four kinds of holism in twentieth-century medicine. For Rosenberg, “organismic holism” refers to physicians’ commitments to work with “the whole patient,” rather than with a specific disease or organ. This rhetorical and technical commitment to “organismic holism” pervades the work of other twentieth-century physiologists as well, who also invoke another of Rosenberg’s holisms, “ecological holism,” defined as understanding “the body in a particular social and physical setting” (Rosenberg 2007, 146). My own work looks at

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American exercise physiology (Johnson 2013a, b, 2015). Exercise physiology is the study of the physiology of bodies in motion, a subfield that grew out of industrial physiology and today overlaps with environmental physiology. To be sure, there are sub-subfields of exercise physiology concerned primarily with the molecular level of analysis; however, many exercise physiologists, in publications and interviews, aspire, as one exercise physiologist characterized the legacy of the Harvard Fatigue Laboratory, to study “the intact human being in all the conditions in which he lives and breathes and has his being” (Chapman 1990, 30). Like Munn’s mid-century plant physiologists, exercise physiologists regularly claimed, and still do, that they are not interested in understanding just one organ or just one physiological system but in understanding human beings as complex phenomena (Gibson 2018, 3). Reactionary rhetoric? Romantic posturing? We don’t have enough histories of physiology-after-1940 to know for sure how much these commitments to holism in these subdisciplines are last gasps at relevance or seeds for historians to identify alternate narratives in the history of twentieth-century biology, as Meurnier and Nickelsen anticipate. In the twentieth century, exercise physiology overlapped with environmental physiology and comparative physiology (the comparison of physiological processes across different organisms) because the scientists moved between the lab and the field to study continuities and differences in organismic function across diverse environmental conditions. Here, two kinds of scholarship buttress the history of physiology. First, recent scholarship on the history of field sciences draws attention to the spaces of research beyond the laboratory. Historians of exercise and environmental physiology attend to the ways twentieth-century physiologists moved between their laboratories and other places to answer the questions “What is life?” and “How do bodies maintain life?”. There is the previously mentioned work of Cueto, Lossio, and Bashford, all documenting mid-twentieth century-physiology as the study of physiologies in place. In addition, John West’s (1998) High Life provides an important intellectual and technical history of high-altitude physiology expeditions and experiments into the 1990s. West follows physiologists (including himself) to their field stations in the mountains of Peru, Chile, Nepal, Russia, the United States, Canada, the Alps, and Bolivia; describes experiments in simulated “environmental chambers;” and catalogues physiological teams and discoveries. Sarah Tracy’s biographical research (2012, 2018) on physiologist Ancel Keys [1904–2004], who led the 1935 High Altitude Expedition to Chile and the famed Seven Countries Study in the 1950s, unites the history of practice, place, politics, and constructions of physiological holism. Vanessa Heggie (2010, 2013, 2016) has published widely on the history of nineteenth- and twentieth-century physiology in extreme environments, describing the epistemological and social function of field stations, noting how these subdisciplines of physiology relied upon and perpetuated class-based masculinities, and exploring the intersections of science, sport, and constructions of “risky” bodies and places. Because the physiologists being chronicled in these histories work with whole organisms, including human subjects, across space, these histories attend, albeit unevenly, to themes of gender, race, colonialism, and constructions of physiological “normal.” Matthew Farish (2013), for example, recounts the role of Norwegian physiologist Kaare Rodahl, who, in the 1950s, headed the US Air Force-funded Arctic

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Aeromedical Laboratory’s Department of Physiology, overseeing research on “the physiology of the Eskimos” and “white controls,” including the study of thyroid function through the administration of the radioactive I-131 (15–16). In his history of a 1900s scientific expedition to Mount Everest, Philip Clements (2018) raises questions about the ethics and epistemologies of high-altitude physiological research during the second half of the twentieth century. An important tension in this scholarship, then, concerns the ways in which exercise and environmental physiologists directly stated or merely implied how physiological research on populations living in or traveling to different environments supported or undermined ideas about racialized physiologies and how this same research participated in Cold War insecurities. Histories of sport and sport science also buttress the history of twentieth-century exercise and environmental physiology (Delheye 2013; Heggie 2011; Wrynn 2010). Roberta Park and Jack Berryman have been at the forefront of this scholarship, describing when and where physiologists’ interests overlap with the interests of physicians, teachers, or other professions with stakes in exercising bodies (Berryman and Park 1992; Park 2007). Charles Tipton has edited two intellectual, transnational histories of twentieth-century exercise physiology (2003, 2014) that chronicle important developments within specific subfields of physiology over the twentieth century. John Hoberman’s (1992, 2005, 2017) histories of investigations into physiological limits examine science, medicine, and sport, pulling the history of Cold War physiology into a late twentieth-century public conversation about doping, politics, and gender. And Dick Kasperowski (2009) describes how exercise physiologists entered into debates over environment, training, the standardization of sporting spaces, and “fairness” during the 1968 Mexico City Olympics. Exercise physiologists still conduct experiments in the field and in the lab and play an increasingly visible role as arbiters and maintainers of boundaries between “natural” and “unnatural” physiologies; between “male” and “female” bodies; and between “abled,” “disabled,” and “superabled” performances. We could use many more histories of physiology-after-1940 that explain how “performance” became the new “labor.”

Future Directions This essay opened claiming that Geison’s 1978 observation – about physiologyafter-1850 remaining “virtually unexplored” – still resonated today. In one sense, my claim is absurd. Soon after Geison sent his biography to press, almost an entire generation of historians of biology cut their teeth writing the history of nineteenth- and early twentieth-century physiology as a discipline. Indeed, the history of nineteenth- and twentieth-century physiology continues to attract the interest of historians of science, who will not be finished with Leipzig and Paris for quite some time (attending, one hopes, more consistently to themes of gender, race, and colonialism.) Yet, in spite of this scholarship, I claimed his observation still resonates because the history of physiology-after-1940 remains relatively unexplored, not as molecular biology nor as independent offspring but as a transnational, still mechanistic, but non-reductive, physiology.

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Historians of physiology-as-society could ask their questions about the political and economic contexts and discourses of scientists articulating or rejecting commitments to physiological holism over the course of the twentieth century. In 1930, Fulton reprinted J.S. Haldane’s “The Organism as a Whole” (from his 1922 book on Respiration) because of Haldane’s striking optimism about the future of physiology as a “holistic” science. Interestingly, Garland Allen (1975) has argued that the spread of Marxism facilitated the early twentieth-century rise of “holistic materialism” in the research of physiologists like Haldane, Henderson, and Cannon (110–11; see also Arminjon 2016; Benison et al. 1987; Munoz 2014). How did specific political or economic commitments or implications shape the demise, fracture, or growth of post-War physiologies? According to Cueto, Argentinian physiologists remained steadfast in their holism at least through the 1950s. Around the same time, comparative physiology, in particular, became institutionalized in China (Cai and Du 2008) and in Serbia (Andjus et al. 2011). Hsiu-Yun Wang (2017) describes the institutionalization of physiology education in schools in Taiwan between the 1950s and 1980s as a Cold War biopolitical project to normalize and naturalize menstruation as an indicator of both individual and population “development.” Whether fragmenting or exploding, physiology became increasingly transnational over the twentieth century. By 1965, Brooks counted ten physiological societies existing in Japan alone. At the 24th International Congress on Physiology held in Washington, DC, in 1968 (the 1st was held in Basel 1889), 4,300 delegates represented 51 countries (Rothschuh 1973). Since then, the annual congress has been held in cities around the world, including Tokyo, New Delhi, St. Petersburg, and Rio de Janeiro. There is room for more national and transnational stories of the history of physiology-as-an-independent-science, before 1940 and after. These histories will need to attend to the relationship between genetics research and physiology. Mittman and Fausto-Sterling (1992) recount how, as “transmission genetics” using the Drosophila organism dominated studies of inheritance in the United States by the 1940s, inheritance research investigating “organism-environment interactions” using the Planaria organism moved to Japan. “Although planarian research was still going strong in the 1950s,” they write, “only 12 percent of the published papers came from U.S. laboratories [down from up to 94 percent before 1945]” (173). How did genetics researchers in different national contexts inherit or reject early twentiethcentury physiological questions concerning environment, nutrition, and adaptation? How did physiologists respond to the findings or techniques of geneticists? A recent joint report of the International Union of Physiological Sciences (the organizing body for the Congresses) and the Physiological Society remarked with enthusiasm that twenty-first-century physiology is the only scientific discipline with the “integrative” orientation and tools necessary to understand the “basic data” of “what genome sequencing has given us,” that is, to understand life (IUPS and TPS 2017, 2). Continued attention to the material culture of physiology will also offer new opportunities to write histories of physiology-as-practice before and after the nineteenth century. Taking the materiality of science as its core, the history of physiology-as-practice sheds concerns with locating and reinforcing disciplinary boundaries. Rather, histories of instruments, organisms, and techniques tend to be highly local and “micro” yet also to move between disciplines and subdisciplines

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with ease (Schloegel and Schmidgen 2002), to cut across national borders and individual lifetimes, and to construct longer histories of physiology without resorting to teleology. These histories of physiology-as-practice may best see through the screens historians have erected in the past 40 years separating physiological studies of humans, other animals, and plants. Finally, constructing histories of physiology-after-1940, self-consciously “holistic” or otherwise, could involve rereading older histories of physiology and putting them into conversation with newer ones. For example, in 1957, the State University of New York held a symposium on the history of physiology to dedicate a new science building, the result of which is a wide-ranging edited volume that includes sweeping papers on the history of physiological ideas, as well as dense literature reviews on specific topics. The resulting The Historical Development of Physiological Thought (Brooks and Cranefield 1959) is itself a rich primary source for historians of physiology-after-1940 interested in understanding the ways in which systems theory and cybernetics permeated biological sciences in the 1950s. Writing the historiography of physiology, one must, for the most part, bound the project by deferring to the historians’ themselves and the degree to which they consider themselves studying physiology. Historiographical journeys in letting go of the term “physiology” can mean quickly falling into rabbit holes deeply burrowed into histories of medicine, public health, anthropology, physics, chemistry, and biological sciences from ecology to neurology. Perhaps, then, the most pressing need within this scholarship is for historians who self-identify as working on the history of physiology, before and after 1800 and before and after 1940, across a range of national and cultural contexts, to clarify and redeclare the value of continuing to use this term “physiology.” And therein lies the potentially overdetermined but nonetheless strong seduction of calling not for less history of “physiology” but for more, more national and transnational stories of investigations into the processes of life itself, more histories attending to the gendered and racialized dimensions of the production of physiological knowledge, more histories of animal and plant physiological research both inside and outside the lab, and more histories of physiology-after-1940– sociological, intellectual, material, or otherwise – to construct an even richer picture of the intellectual commitments, institutional positions, and economic, political, and social concerns that unite and divide those who have found in the study of living organisms an orientation and tools to make sense of life.

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Contents Part 1: Plant Breeding and Agriculture in the Historiography of Biology . . . . . . . . . . . . . . . . . . . . . Darwin and the Breeders . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mendel and Genetics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eugenics and Biotechnology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Part 2: Plant Breeding and Agriculture for the History of Biology . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bio-agricultural Economics and Politics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bio-agricultural Knowing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bio-agricultural Field Science . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Part 3: Plant Breeding and Agriculture for Future Histories of Biology . . . . . . . . . . . . . . . . . . . . . . . Global Histories . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Environment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Biology and Technology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

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Abstract

There are unique opportunities that plant breeding and agriculture offer the historian of biology and unique ways in which the historian of biology can inform the history of plant breeding and agriculture. This chapter will first address what agriculture has in common with themes that cut across this handbook, before turning in Part 2 to issues, problems, and questions that stem from agriculture’s particular features and ending in Part 3 with paths for future work. There are unique opportunities that plant breeding and agriculture offer the historian of biology and unique ways in which the historian of biology can inform the history of plant breeding and agriculture (Harwood 2006; Phillips and Kingsland 2015). D. J. Berry (*) London School of Economics, London, UK e-mail: [email protected] © Springer Nature Switzerland AG 2021 M. R. Dietrich et al. (eds.), Handbook of the Historiography of Biology, Historiographies of Science, https://doi.org/10.1007/978-3-319-90119-0_27

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There are also of course questions and challenges that the study of agricultural sites shares with the study of other biological sites, such as those in medicine (Wilmot 2007; Woods et al. 2018), the environment (Agar and Ward 2018), and nonagricultural industries (Bud 1993). Indeed, in some instances the agricultural, medical, environmental, and biologically industrial will be one and the same. This is to say nothing of what agricultural sites share in common with histories of science beyond biology, but that is a broader discussion I can only mention in passing (Parolini 2015). This chapter will first address what agriculture has in common with themes that cut across this handbook, before turning in Part 2 to issues, problems, and questions that stem from agriculture’s particular features, ending in Part 3 with paths for future work. The chapter therefore treats the intersection of biology and agriculture as demanding its own integrated attention, the two parts making up a larger historiographical whole. There are a number of reasons to give agricultural sciences and technologies this kind of autonomy from the historiography of biology at large. First, it reminds us to question the nature, direction, and extent of influence that biological science and agriculture have had on one another. Second, it promotes a more symmetric understanding of the knowledges that have mattered for biological science and agriculture. This is particularly important because so much of the history of biological science in agriculture has been about establishing the authority of scientific expertise over agriculture, often in competition with other kinds of expertise distributed throughout farming. If we did not approach agricultural contexts symmetrically, we might end up recapitulating the very arguments we are meant to be analyzing. Third, it establishes a healthier and more distant vantage point for the historian, keeping the existing historiography of biology at arm’s length, allowing us to better observe its deficiencies and assumptions. Aside from giving autonomy to the agricultural in histories of biology, there is another broad historiographical point to make. Historians of biology and agriculture have to strike a balance between which historiographical lineages they dedicate their work to, or indeed, whether they see themselves contributing to both histories of biology and agriculture simultaneously. In some respects this issue is itself unique to agriculture, for if we look at the other topics in this handbook, only one or two other chapters are asked to compete with completely different sets of scholarly lineages in their telling, these including Tracy Teslow on Race and Ethnicity, Marsha Richmond on Women, and Ana Barahona on the transnational. Yes, other kinds of historian and scholar may make important interventions on the history of eugenics, Darwinism, and biotechnology, but when it comes to these topics, nobody is in a position to outbid the historian of biology. Agriculture is different, both in content, thanks to the variety of experts that it enrolls across a very wide range of potential specialist areas, and also in terms of the historiographical landscape in which it sits, because agriculture has indeed belonged to whole other kinds of historian, be they social historians, economic historians, or historians of agriculture and the environment. Ultimately all my talk of ownership and bidding is petty, and of course even in those topics that seem primarily the concern of the historian of biology other historical traditions and branches of scholarship are constantly being drawn in. What I mean to

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convey is that historians of biology have been late to agriculture; their insights have not always been understood as relevant or complementary to the history of agriculture; historians of agriculture seem to be getting on all too well without the historian of biology; and if the recent growth in interest among historians of science into the agricultural is to be maintained and consolidated, then interdisciplinary awareness is essential. Here historians of biology offer a suite of valuable opportunities for historians of agriculture, be it through all the techno-imagining that goes into broader agricultural debate or the chance to rethink social and economic relations on the farm, the meanings embodied in agricultural spaces, organisms, and communal practices or, as Jonathan Harwood has so brilliantly shown, through the issue of global food security (Harwood 2012). But agriculture also demands sensitivity and humility from the historian of biology, to know when multiple epistemologies are in play, multiple historiographies, and therefore how to translate any new historical understanding into a form that matters for defined audiences. These audiences should include not only historians of science but also those working on and in agricultural industries.

Part 1: Plant Breeding and Agriculture in the Historiography of Biology Agricultural history matters for an array of themes that cut across the history of biology. I will confine myself only to a select number of the topics covered by this handbook, but one could also write sections on agriculture and the mathematization of biology, agriculture and nutrition/food science, and agriculture and ecology/ environmental sciences. Indeed the latter is something I will return to in Part 3, as an issue that is becoming increasingly central to historians of biology, one in which agriculture can and should play a large role.

Darwin and the Breeders It is well known that plant and animal breeders were hugely important for the development of Darwin’s research (Secord 1985). Breeders were not only a key community in his expansive network of correspondents, but plant and animal breeding also provided evidence, theories, and resources. It is worth beginning here because if we can appreciate the extent of the community of breeders, along with the expansion and sharing of their knowledge and industry throughout the late eighteenth and early nineteenth century (Derry 2003; Matz 2011; Russell 1986; Wood and Orel 2001), then we are more adequately prepared for our later discussions of Mendelism and genetics, not because these histories connect linearly but because we need to understand the depth and breadth of hereditarian thought long before anything like Mendelian genetics entered the scene. Developing a good understanding of the relationship between agriculture and Darwinism is also an excellent way to acclimatize the historian of biology to the value added by taking agriculture seriously. For instance, if we failed to do so, then we might be left assuming that Darwin turned to breeding communities after the Beagle voyage

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because their selection practices made for a teachable analogy with selection events in nature. But this is not how his research actually unfolded (Hodge and Kohn 1985), and as Bert Theunissen has argued, the assumption that Darwin’s interests in domestic breeding mattered first and foremost for the making of such an analogy has masked both the variety of questions that breeding could actually illuminate and also Darwin’s selectivity toward the features of breeding practice he was prepared to recognize (Theunissen 2012). Our understanding of Darwin and his theorizing has been materially altered by giving priority to the context in which he worked and the importance of agriculture within it. The most significant feature of domestic breeding for Darwin was the evidence it supplied of the necessary preconditions for variation (Hodge and Radick 2003, 4–5).1 The changes in plants and animals achieved by breeders took place in environments outside of nature: in gardens, coops, stables, glasshouses, and on farms. It was the environmental setting of domestication, so Darwin came to argue, which increased an organism’s capacity to transmute. This increased variability would then become available to the breeder for management and manipulation (Winther 2000). From this vantage point, we immediately begin to see how very different were Darwin’s views on heredity in comparison to evolutionary theorizing today and that agricultural contexts were not just an analogical thought experiment but rather contained important information about biological development, physiology, and their interconnections with generation. The close relations between organism and environment (Baranski and Peirson 2015; Ritvo 1987; Woods 2017) have had multiple significances for biology and society, business, and politics and also for those maintaining an empire, areas that will be returned to in Part 2. A more general lesson that emerges from work on the history of breeding and Darwinism, one that is important to appreciate about breeding and breeders in any historical period, is that it is a pursuit steeped in epistemic anxiety (Lidwell-Durnin 2018). A breeder might be looking at the wrong parts of plants and animals, in the wrong ways, with the wrong apparatus, in the wrong kind of space, for the wrong length of time, with the wrong periodicity, and so on. But the history of breeding is not one of people fretting. Instead we find different kinds of actor (professional and amateur breeders, farmers, scientists) working in a wide range of social contexts (domestic private and public gardens, greenhouses, farms, universities), all developing prescriptions, theories, and techniques to help govern their practices. Many were also sufficiently convinced of the value of their findings and understanding that they shared them through meetings, correspondence, and publications. Anxieties, as I couched them, have typically only been drawn upon during public deliberations and disputes about what actually works and why. These are the kinds of argument that become acute when a new idea or technique seems to undermine widely held understandings and practices, devaluing all those earlier schemes as “misguided,” with implications for the socio-epistemic status of their initial promoters: if the new

Gregory Radick was the first person to suggest that I read Darwin on these terms, and I remain grateful for his sharing this insight.

1

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understanding is accepted, then erstwhile wizards can be turned into fools or charlatans. All of this remains true in the present and should be borne in mind throughout the rest of the chapter.

Mendel and Genetics The single most important topic prompting historians of biology to take an agricultural turn has been the origins, reception, and development of Mendelism and genetics. The number of comprehensive studies of agriculture and science that have been motivated by interest in Mendelism and genetics is formidable, at least in comparison to any other topic at the intersection of biology and agriculture. That genetics retains such a privileged position is perhaps by now problematic (MüllerWille and Rheinberger 2012; Wilmot 2007a), but it is nevertheless the case that early work on these questions by historians of biology in the 1980s and 1990s has gone on to provide a foundation that today supports more general enquiry into the relations between agriculture and biological science. As with Darwin, many historians now draw direct links between the context of Gregor Mendel’s research and industries horticultural, agricultural, and pastoral (Gliboff 2015; Müller-Wille and Orel 2007; Wood and Orel 2005). These histories have considerably revised earlier interpretations that had painted Mendel as a protogeneticist, one focused solely on discerning laws of nature through clever experimental design. Instead “it is much more likely that the direction of Mendel’s hybridization research was guided by the strong agricultural interests around him” (Allen 2003). As we shall see shortly, practical breeding and the commercial production of agricultural seeds would also go on to be essential for the firm establishment of genetics, a science which many of its founders famously traced back to Mendel, on grounds that are equally famously problematic (Olby 1979; Sapp 1990). If Mendel was responding to a wide ranging set of persons who at the end of the nineteenth century were developing means and methods for understanding several different aspects of heredity, then we had surely better expand our search, to emphasize the variety of questions pursued at the intersection of biology and agriculture over time, regardless of later gene-centered developments. Staffan Müller-Wille and Christina Brandt have made this argument in an introduction to their recent edited collection on heredity, in which agriculture features heavily (Müller-Wille and Brandt 2016; see also ▶ Chap. 6, “Gregor Mendel and the History of Heredity,” this volume). Once again, recognizing and attending to agriculture directly, rather than for the sake of appreciating Mendelism or some other candidate breakthrough, has substantially revised our histories of biology. When it comes to genetics and agriculture, the literature is vast. Geographically the discussion has primarily focused on the United States; a few European countries including France, Germany, the Netherlands, Italy, and the United Kingdom; and more recently China and Japan. Because all of this material can contribute to global histories of plant breeding and agriculture, I have compiled all the works that I know of in Table 1, organized according to primary geographical focus and a broad

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Table 1 Historical research at the intersection of agriculture and genetics, organized according to primary geographical focus and approximate period China

Denmark France Germany

International

Italy

India Japan Mexico

Netherlands

South Korea Spain Sweden United Kingdom

Early C20th Late C20th Early C20th Early C20th Early C20th Mid C20th Early C20th Early-late C20th Early C20th Mid C20th Mid C20th Late C20th Early C20th Early C20th Mid C20th Early C20th Early-late C20th Late C20th Mid C20th Early C20th Early C20th

Mid C20th

United States

Late C20th Early C20th Early-mid C20th Early-late C20th

Lavelle 2016 Jiang 2017; Schmalzer 2016 Müller-Wille 2007; Müller-Wille 2008 Bonneuil 2006, 2008; Bonneuil and Thomas 2010; Gayon and Zallen 1998 Gliboff 2016; Harwood 1997; Harwood 2000; Wieland 2006 Elina et al. 2005; Gausemeier 2010; Harwood 2010; Heim 2003; Saraiva and Norton Wise 2010 Berry 2014c, Bonneuil 2016, Campos and von Scherwin 2016, Harwood 2004, 2016; Onaga 2010; Saraiva 2016 Harwood 2012 Iori 2013 Saraiva 2010 Saha 2013 Baranski 2015a, b Fujihara 2018; Onaga 2016; Iida 2016 Barahona et al. 2005; Barahona and Robles 2001 Harwood 2009; Matchett 2006 Theunissen 2008 Maat 2001 Tae-ho 2018 Camprubí 2010 Ackerman et al. 1948; Åkerberg 1986; Müller-Wille 2005; RollHansen 1997; Roll-Hansen 2000 Berry 2014a, b, 2018; Brassley 2007; Button 2017; Charnley 2011, 2013a, b, c, 2016, Holmes 2017a, b, 2018a; Marie 2004; Olby 1989, 2000; Opitz 2011; Palladino 1993, 1994, 2002; Radick 2013 Palladino 1990; Parolini 2012; Peirson 2015; Wilmot 2007a, b; Holmes 2018b García-Sancho 2015; Myelnikov 2017 Allen 2000; Carlson 2005; Campos 2015; Cooke 1997; Curry 2016b; Kevles 1980; Kimmelman 1983, 1987, 1992, 1997, 2006; Tyrell 2015 Derry 2012, 2016; Fitzgerald 1990, 1993 Curry 2016a; Henke 2008; Kloppenburg 1988

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periodization.2 Though some of these histories address multiple countries, I have not repeated entries, so the table should only be used as a rough guide. Historical accounts clearly aspiring to an international perspective are collected under “international.” It is inevitable that my criteria for selection are imperfect. Perhaps the most important lesson we might take from this exercise and the growth of the historiography of agricultural genetics is that these works can and should now be used to explore biology in a more diffuse way, recognizing genetics as only one specialism, practiced around the world in ways that were not homogenous, a specialism which is by now perhaps most interesting as continuous or discontinuous with practices and knowledge that do not begin and end with genetics. In truth this is precisely how some of the histories in this table have already treated genetics, as an entry point. Within this literature I can highlight two key areas. The first is the creation of genetic expertise and its institutionalization, which was a fundamentally agricultural enterprise. The model study which broke this ground was Barbara Kimmelman’s investigation of American genetics (Kimmelman 1987), work which is equally significant for tying together agriculture, genetics, and eugenics, to be discussed in the next section. Kimmelman’s foundations have been subsequently built upon by an array of historians working on numerous geographical contexts (Berry 2014b; Bonneuil 2006; Charnley 2011; Fitzgerald 1990; Harwood 2012; Iida 2016; Iori 2013; Maat 2001; Olby 1991, 2000; Onaga 2016; Palladino 2002; Saraiva 2016). The availability of all of this material surely now demands a stronger international comparative approach. Harwood has also supplied an important summary and final assessment on the evidence for Mendelian genetics’ impact on breeding, which goes deeper than many earlier studies into the technical differences between breeding approaches and where Mendelian expertise could or could not have intervened. In the final analysis, so he argues, Mendelian genetics simply could not have had too considerable an impact on the practice of agricultural plant breeding, at least not until late into the twentieth century, by which point the extent to which genetics was “Mendelian” is itself debatable (Harwood 2016). The second key area that I have room to address here is agriculture’s significance for the history of intellectual property (IP) in biology, both before and subsequent to the emergence of genetics. Inspired by Daniel Kevles’ pioneering work (Bugos and Kevles 1992; Kevles 2007), which he was also asked to make suitable for international policy makers (Kevles 2002), and building on other essential interventions from legal scholars attending to biology and IP law (Dutfield 2003; Pottage 2006), historians have demonstrated the fundamental importance that agricultural plants and animals have had for the origins and development of intellectual property rights in biological things and organisms (Berry 2014a; Charnley 2013a; Charnley and Lawson 2015; Charnley and Radick 2013; Fullilove 2017; MacLeod and Radick

2

My considerable thanks to Berris Charnley, who helped make sure I missed as little as possible, and Jonathan Harwood, who many years ago shared with me his lists of historians working across agriculture and genetics. Also thanks to the History of Science Society’s IsisCB database which helps keep us all up to date.

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2013; Parthasarathy 2017; Radick 2013). The precise role played by genetics, geneticists, and legislators in the development of IP, and the role of IP in the development of genetics, remains an interesting topic of debate and offers historians of biology an excellent case study for exploring how scientific ideas are made in tandem with social and legal conventions.

Eugenics and Biotechnology Barbara Kimmelman was one of the first historians of biology to demonstrate the close connections between agriculture, genetics, and eugenics (Kimmelman 1983). Looking in particular at the membership of professional breeder associations and elite leaders of both genetics and eugenics at their origins, Kimmelman found considerable overlap between these groups, along with common cause. Historians of biology have since deepened the connections between eugenics and agricultural biologies, finding them in conservation movements (Allen 2013; Uekötter 2006), concern regarding national degeneration in food supply and people (Berry 2015a; Lovett 2007; Roll-Hansen 1989), and popular sports such as thoroughbred racing (Thurtle 2002; Tyrell 2015). Recent studies have demonstrated the importance of taking agricultural sites as seriously as those in human genetics if we are to understand how people and other organisms as materials of the state are remade for modernist agricultures and sciences (Bonneuil and Thomas 2010; Flitner 2003; Saraiva 2016). For the historian of biology working on agriculture, eugenics is an ever-present theme, in part because the analogies, metaphors, and evidence for eugenics have always been bound up with agriculture but also because when dealing with the improvement of agricultural animals and plants, the territory is potentially eugenic from the get-go. Rather than worrying about what is or is not eugenic per se, what we should take away from the historiography of eugenics is that reproduction, animal and plant stewardship, breeding programs, and so on, these are all inherently and inescapably political (Palladino 1987). The question then becomes: what arrangement do we want, and to benefit whom? Eugenics is also then, and much more straightforwardly, about economics. It should be well understood by now that biotechnology refers to a particular historical formation of biological science and technology emergent after the Second World War (Bud 1993). This has not stopped others from using the term to describe much earlier periods and indeed conjure up a long history of biotechnology from the ancient past to the present (Berry 2018). If there are indeed significant features of production, relations between humans and other organisms, breeding, and so on, which can be traced over thousands of years, then they need to be specified and thoroughly demonstrated. Otherwise the historical imagination is simply serving wish fulfillment. Within the historiography of biology, Sarah Franklin’s study of sheep breeding and cloning, with particular attention to the case of Dolly, has reinvigorated discussion of the links between science and the state and between research and policy (Franklin 2007). Where García-Sancho has emphasized the significance of the history of research at Edinburgh that eventually led to the birth

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of Dolly (García-Sancho 2015), Myelnikov has stressed the need to go beyond shallow politico-economic readings of the Thatcher government’s science policy and its impact on agricultural biotechnology (Myelnikov 2017). There are then multiple cultures of biotechnology, many having relations with or stemming from agricultural breeding, all of which are in need of appreciation, just as we appreciate multiple cultures of agricultural and biological science more broadly. To navigate them we will need, as Jean-Paul Gaudillière has argued, to become sensitive to new angles of analysis, one of his proposals being attention to historical “ways of regulating” that distinguish between professional, administrative, industrial, and consumerist/activist modes of regulation (Gaudillière 2009). Further historical works that matter for agricultural biotechnology, but which addresses politics and economics from a more macro level, are considered in Part 2.

Part 2: Plant Breeding and Agriculture for the History of Biology In addition to the above themes, there are important questions that can be imported or piped into the history of biology most efficiently through attention to plant breeding and agriculture. Some of the areas covered in Part 1 will reappear here, though in a different light. It is the opportunity to juxtapose or switch between seeing organisms, institutions, and actors as biological, technological, environmental, agricultural, and scientific that produces much of the added value gained by looking at the history of biology and agriculture together.

Bio-agricultural Economics and Politics The work of any biological scientist dedicated to agricultural questions takes place within a particular political and industrial setting. That setting provides its own incentives and is composed of multiple kinds of power relations. In order for the historian of biology to appreciate these aspects of their case, they need to be aware of some of the principle ways in which agricultural science and technology have been understood as influencing farming. For these purposes, one of the most significant research programs developed in the historiography of agriculture and biological science is that of Jack Kloppenburg in his now iconic First the Seed (Kloppenburg 1988). Analyzing the ways in which science and technology more generally intervene in agricultural contexts, his argument centers around the means by which seeds have undergone a process of commodification and the role of scientists within this process. Written around the same time, commodification was also a central feature of Deborah Fitzgerald’s history of plant breeding and the origins of hybrid corn (Fitzgerald 1990). These kinds of study, and others that have followed since, demonstrate the ways in which commercial opportunities have influenced the direction of scientific research and the long-standing shared interests of academic breeders with agribusiness. While this kind of argument is now commonplace in broader histories of biological science in the twentieth century, as, for instance, in biomedicine, the particularities of agriculture make their investigation all

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the more urgent. Unlike the majority of the products emerging from medicine, the products of agricultural science are not intended to have their use and circulation confined to a network of highly scrutinized institutions (hospitals, etc.) and professionals. Instead the products of agricultural science are intended for use within a broad, heterogeneous, loosely aligned set of communities, a set of communities that are already functioning and circulating extant products. I am suggesting that in terms of competition between different kinds of valuation and the differential power relations between the producers and consumers of science, the commodities of agricultural science have a more far-reaching impact than in those other places where biological science becomes biological business. Or at the very least, agriculture provides a more idealized setting for the historian of biologies’ investigation. In addition to commercial interest, other historians have focused on the ways in which agricultural scientists have become materially enrolled in the pursuit of nationalist agendas. This is clear in both histories of agricultural science under fascism (Bonneuil and Thomas 2010; Camprubí 2010; Gausemeier 2010; Harwood 2010; Saraiva and Norton Wise 2010; Saraiva 2016) and in democracies with long traditions of nationalism, paternalism, or technocracy (Charnley 2016; Olby 1991; Curry 2016c; Harwood 2012). An additional layer of significance for understanding biological diversity as a resource (Bonneuil 2019) is added by imperial contexts, be it for scientists whose work constitutes part of the imperial project, or for the administrators of colonial states, or for indigenous scientists and farmers (Baranski 2015b; Bonneuil 1999; Charnley 2013b; Kumar 2012; Maat 2001; Woods 2017). The role played by agricultural scientists within more general governmental strategies for control of food supply and aid, particularly throughout the Cold War, has also come to receive much needed attention (Perkins 1997). The period commonly referred to as the “Green Revolution” has come to receive substantial historiographical revision (Kumar et al. 2017). For instance, the impact and legacy of key figures such as Norman Borlaug, who is otherwise championed as single-handedly bringing an end to cycles of starvation, have been hugely overexaggerated. Not only did the dwarf varieties for which he became famous owe “their existence to the skill of Japanese farmers a century earlier” (Harwood 2018), but his real impact occurred at the level of influencing Indian political and agricultural officials (Baranski 2015b). Our discussion of the institutionalization of biological science within different kinds of political context, the creation and circulation of new commodities within and across empires, has quite naturally brought us to the question of knowledge. The historian of biology attending to plant breeding and agriculture cannot afford to concentrate solely on knowledge produced and promoted by people who identified as agricultural scientists. Here again then agriculture holds a wider lesson for the history of biology more generally.

Bio-agricultural Knowing Speaking historiographically, if there is one overriding feature of the biology of agricultural plants and animals that historians have found either promoted, or

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demanded, a particular kind of epistemological posture distinctive to plant breeding and agriculture, it is their tendency to vary. Variability between generations and during development appears time and time again as a fundamental fascination or epistemological problem. We saw this earlier with Darwin, but here I will first turn to genetics, before broadening our horizons. Paolo Palladino’s revisionist analysis of the knowledge claims made by geneticists in the context of commercial plant breeding has remained central to historiographical debate and discussion. He not only demonstrated that geneticists did not influence the schemes of breeders as immediately or straightforwardly as some historians of agriculture and agricultural science had made out but that they were more than comfortable asserting the value of their own knowledge of plant variability and the influences upon it (Palladino 1993, 1994, 2002). In addition, Kimmelman was early to argue that working with agricultural materials, and closely alongside commercial plant breeders and farmers, contributed to making agricultural genetics a distinctive community and strand of thought within genetics more broadly, one that took whole-organism variability more seriously than other kinds of genetics, a position that some agricultural geneticists themselves argued explicitly (Kimmelman 1992, 1997). To give two final genetic examples, it was plant variability in the field that according to Pauline Mazumdar eventually shook R.A. Fisher’s otherwise strong commitment to the insignificance of environmental influence on inheritance (Mazumdar 1992, 124). Likewise, Gregory Radick has shown how demonstrations of authority over the extent of variability provided excellent opportunities for geneticists such as William Bateson to publicly consolidate their new science and its productivity (Radick 2013). Numerous other examples could be listed. Moving beyond genetics, we have also learnt how important geographical biological variation has been for inspiring systematic analysis and mapping of economically significant plants and animals. Such was the aim of German oekonomie as has been explored by Denise Phillips in the eighteenth and early nineteenth century (Phillips 2015) and in nineteenth-century botanical distribution mapping (Güttler 2015). Indeed virtually all of the chapters in the collection from which these two examples are taken are to some extent dealing with biological variability. In addition, most if not all of these chapters also demonstrate how knowledge of variability was widely shared among scientists, farmers, local processors, and other actors, who could be called upon to provide specific knowledge regarding faraway corners, supply local labor for agricultural experiments, or corroborate analytical findings through their own testimony. The fact of variability and the fact of diverse lay and elite knowledges regarding biological variability are perhaps linked, because candidate explanations for variability are always necessarily limited and typically non-exclusive. Chauvinistic and scientistic understandings of variability may emerge from time to time, particularly when a new professional class or a new scientific discipline seeks to establish itself. But historians have nevertheless continuously found that personal knowledge and appreciation of organisms in local environments has always rushed back in, to the extent they ever really left. We should not have to render such knowledge natural, or bucolic, or innocent, or unscientific (especially given how centrally important it has been for the

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development of all agricultural biological science) in order to appreciate it. As it would therefore be inaccurate, not to mention patronizing, to distinguish between scientific and nonscientific epistemologies, how else might we analyze these kinds of knowing? Sometimes the categories of local knowledge or craft knowledge have been suggested, but neither of these adequately captures the body of knowledge communicated among practitioners and passed between generations. Helen Tilly has supplied one valuable alternative through attention to the history of colonial science in Africa. There she introduces the concept of “vernacular science,” which is meant to characterize local indigenous knowledge while avoiding historiographical treatments of such knowledge as to some degree romantic (Tilley 2011, 122). The long history of vernacular knowledge and its direct contributions to imperial and postcolonial science and industry attest to ways in which indigenous knowledge is sought and erased simultaneously (Bil 2018; Schiebinger 2004). It may be that one way in which such erasure would become less likely is if the concept of vernacular knowledge was applied equally to the farming and breeding communities found inside imperial centres, rather than only to those within colonized states. Doing so meshes nicely with Harwood’s analytical distinction between different plant breeding strategies as more or less “cosmopolitan” or “local,” a distinction which is intended to be equally applicable to programs in any national context. A cosmopolitan approach aims at producing a novel plant which can be marketed as suitable for growing in any environment. Meanwhile a local breeding strategy aims at refining a varieties’ suitability for a particular location (Harwood 2012, 45). We can therefore add that a local strategy would also be more amenable to vernacular knowledge than a cosmopolitan one. Building up our historiographical picture in this way provides a richer landscape in which to situate histories of biological science, directing our attention to features that we might otherwise fail to see. It also provides clear ways in which the work of agricultural scientists in the field is dependent on, and constitutive of, broader social relations.

Bio-agricultural Field Science Historical attention to agricultural science has the capacity to radically alter our conceptions of how biological research is conducted, by what experimental means, and with what relations between practice and understanding. One clear reason for this is that so much of agricultural science takes place in spaces designed differently from laboratories, which have otherwise been taken as the quintessential arenas of modern science. While this feature is not unique to agriculture, it is nevertheless the case that greater attention to agricultural experimentation has only just begun and is therefore due to receive much more systematic attention (Parolini 2015). Working with examples largely outside of the territory of agriculture, Robert Kohler has done the most to bring the field sciences to historians of biology (Kohler 2002). Precisely how the historian can and should understand the field sciences in themselves and in relation to other kinds of scientific space remains an interesting open question which historians of biology can more actively explore (Ekerholm et al. 2017; Vetter 2016). For instance, it is clear that historical actors

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have at times used and abused idealized notions of “field” versus “laboratory.” These kinds of idealization have typically revolved around putative differences in how a setting enables different kinds of interaction with nature. It is supposed, for instance, that a laboratory setting tries to make itself “nowhere” so that an experiment taking place in Helsinki could just as easily be taking place in Jamaica, while a field ideal aspires to make the most out of the specificities of place. It is also clear that holding up field and laboratory ideals and putting them in tension with one another has been a productive strategy for biological scientists looking to design new kinds of space, such as Frits Went’s midtwentieth-century phytotron (Kingsland 2009), an expensive kind of resource that often received direct financial support from agricultural industrialists and food processors (Munns 2017). But there are also good reasons to want to avoid making a typological distinction between lab sites and field sites, instead attending to how shared epistemic motivations can be enacted just as heterogeneously within laboratories and fields as between them (Berry 2015b). And though the questions of the importance of place specificity are more easy to appreciate in the field (Shavit and Griesemer 2009), they also feature in their own ways in biological laboratories (Shavit and Ellison 2017). Agriculture has a further benefit when it comes to historical attention to biology in the field, in that there is already a well-established and ongoing historiographical exploration of how direct engagement with a field site mattered for the epistemic and social authority of scientists, administrators, and farmers. To give two recent examples, Abigail Woods has defined aspects of this kind of interaction with the field as “learning by doing” in work on nineteenth-century veterinary research and administration (Woods 2013), and the historical literature on “book farming” and its tensions with the field has continued to expand (Fisher 2018). Lijing Jiang has also stressed the ways in which direct engagement between biologists and widely dispersed communities throughout China, modelled on social arrangements already developed by agricultural scientists, enabled not only collecting expeditions but also the opportunity to directly associate a biologist with China as embodied in its particular landscapes and natural history. “Chinese biologists regarded the physical labor involved in fieldwork and experimentation as providing an antidote to the old habit of book learning believed to be based on ‘empty philosophy,’ which was thought to have hampered China’s growth” (Jiang 2016, 165). In what ways have different kinds of space contributed to the making of sciences that are expected to bring about a material difference in farming? What cultures and traditions of experimentation have contributed to agricultural science, and how has the biological understanding thereby developed been associated with, or ignored by, more general biology? Any investigation of plant breeding and agriculture will necessarily encounter these issues.

Part 3: Plant Breeding and Agriculture for Future Histories of Biology In this section I highlight three particular areas of research that seem ripe for the attention of the historian of biology. All three have been subject to recent and rigorous interdisciplinary activity, opening up exciting potential future paths for work in the history of biology that either parallel, critique, or piggyback on these

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efforts. All three also happen to follow the lead of historians of technology, who have been among the most innovative in recent years when it comes to understanding agriculture and its relations with science and technology.

Global Histories We are still not in a position to draw upon historical investigations of biological science and agriculture from all continents (see also Barahona ▶ Chap. 17, “Local, Global, and Transnational Perspectives on the History of Biology,” this volume). The most obvious gap is that while we have accounts of administration in Africa, historians and anthropologists addressing plant breeding and agriculture in this continent typically have not pursued the role of biological science in particular but rather scientists and administrators in general (Hodge 2007). The exceptions are Bonneuil on Senegal (1999) and Saraiva on Ethiopia and Mozambique (Saraiva 2016), alongside some studies of the history of science and botany which from time to time stray into the agricultural (Drayton 2000). Nevertheless, even with these kinds of large gap still remaining to be addressed, it is heartening to see how much of the globe has by now received attention from historians of biology in agricultural mode. As with many other areas in the history of science and technology, historians of biology can and should now use these foundations in order to secure the resources needed for the production of global histories of plant breeding and agriculture. One way in which we could begin to conceptualize these efforts would be through the very recent historiographic innovation of “cropscapes,” as developed by historians of technology Francesca Bray, Barbara Hahn, John Bosco Lourdusamy, and Tiago Saraiva. While their full treatment is currently awaiting publication, and so it is somewhat premature to be discussing it here, the scope and scale of their project is significant enough for historians of biology addressing agriculture that ignoring it would be to make this handbook immediately outdated. Elements of their program can be gleaned from Bray’s recent co-edited collection on rice (Bray et al. 2015) and Hahn’s earlier investigation of tobacco (Hahn 2011), which will be drawn upon in a different context shortly. The following paragraph is based on a forthcoming journal article that the authors have been kind enough to share ahead of publication. While cropscapes constitute an analytical perspective that goes well beyond the typical bounds and ambitions of most histories of biology, the history of biology is nevertheless particularly well placed to contribute to them and be seen within them. To begin with, the authors define a cropscape as “firstly a concept encompassing the constellation of elements that are brought together to make a specific crop in a specific time and place, and secondly, as a tool to analyse the sum of movements and forces that produces and reproduces the working set of elements, as well as the capacity or incapacity of crops and their assemblages to travel” (Bray et al. Draft MS). The history of biology is implicated here in a number of ways. First, agricultural scientists and agriculture-facing biologists have been directly responsible for the making and unmaking of crops and the pests, symbionts, and cultivators – human and nonhuman – that go into making a specific crop. Second, the elements of

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different cropscapes around the world have been the locus of attention of agricultural research and extension. Third, and as these authors themselves highlight, agricultural plant scientists and breeders have been exceptionally dedicated to making organisms travel from their points of origin, finding ways to ensure they extend their crop life into new environments. Aside from these direct connections with cropscapes, some of these author’s programmatic prescriptions are also particularly attractive for historians of biology. It is not enough, for instance, for historians to continue pointing out how particular crops that now matter for western diets and industries were originally found through international exploration and extraction. Instead any history of biology that does not attempt to at least keep one eye on that crop in its original location, at the same time as investigating the new life of that organism in imperial centers, has not really told the history of that crop at all. To give an example, it is one thing to know how important were repeated trips to South America for research and breeding of potatoes throughout the twentieth century, but if we only focus on the results of those trips in, for instance, the United Kingdom and do not attend to ongoing maintenance and research in Peru throughout that same period, then we have been unhelpfully selective in our historicization of the plant. Such a prescription may also help us when delivering historical research into the present, complete as it is with governance instruments such as the Convention on Biological Diversity and the Nagoya Protocol. Lastly, some of our recent and most thoroughgoing accounts of crop movement and development in the history of biology may already be cropscapes avant la lettre. Bray et al. themselves point to Courtney Fullilove’s history of commercial wheat production (Fullilove 2017), and we might also add Phillip J. Pauly on horticulture’s role in colonizing the United States (Pauly 2007), or Rebecca Woods on the international circulation of sheep breeds (Woods 2017), or Berris Charnley on the significances of genetics for empire building (Charnley 2013b), or Sabine Clarke’s focus on sugarcane production for understanding the colonial and postcolonial history of the Caribbean (2018). Cropscapes just may well be the ideal vehicle for making global histories of biological science, plant breeding, and agriculture.

Environment Historians of biology invested in plant breeding and agricultural science cannot afford to be left behind as historians of the environment and technology set about co-producing a new scholarly discourse (Agar and Ward 2018; Jørgensen et al. 2013; Reuss and Cutcliffe 2010; Russell et al. 2011; Vetter 2016). No matter what level of the bio-agricultural sciences one looks at, be it parasitology, botany, microbiology, you name it, these can be realized as fundamentally environmental. Take Jon Agar’s recent suggestive “eight ways of combining environmental and technological history” (Agar 2018). These range from ways in which environments are inputs into technological systems (agriculture supplies essential examples), to the environment as something changed or damaged by technological processes (again, agriculture is key), to environmentalism and the environment as something untouched (you can

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see the point I am making). Situating biological work in such contexts can achieve many of the same ends that I highlighted above for global histories but at smaller and national scales (Uekötter 2014). A cropscape eye view is complementary with, if not part of, environmental history making. It might be worried that these approaches essentially replace history of biology with environmental history and histories of large technological systems. But this is not so. For as I explained at the beginning of Part 2, much of the value to be gained by looking at biological science and agriculture at one and the same time is the ability to chop and change between different historiographical perspectives. We might then wish to dedicate ourselves to understanding the development of, for instance, novel techniques for crop identification and then consider what interaction those who developed such techniques had with their immediate surroundings and the impact of their work on environments or industries at large. It is through biological materials that we can weave our historical accounts (Clarke and Fujimura 1992), provided we remember to make ample room for any given material’s multiple semiotics. That they will have multiple semiotics is one reason why we will never be able to replace history of biology, for biological science is one of the most important, exciting, and surprising machines for making meaning that we could hope to alight on.

Biology and Technology Completing the trifecta of future paths for the history of biology and agriculture that also happen to entwine it closer with the history of technology is the suggestion that historians of biology can productively integrate their work with the historiography of technology more generally. This runs all the way down from environments and technological systems, to organisms and the things they are made of. The motivations for making these suggestions are only tangentially related to the more recent history of agricultural science, in which certain actors claim to have remade biology as technology (Campos 2009; Scott et al. 2018). Being able to respond to such actors does not require that historians of biology understand technology better. But that some historical actors have at various times in the past and present interpreted their materials as much in terms of technology as of biology does inspire curiosity as to what elements of the historiography of technology may well serve our ends. More straightforwardly, the suggestion that an agricultural plant or animal also shares features with objects we would more readily identify as technologies is not in and of itself redundant or monstrous. Admitting this, it is perhaps no surprise that throughout this overview of the history of agriculture and plant breeding that I have often had need to call on the history of technology. Nor am I being particularly original in making this suggestion. For instance, Helen Curry has argued strongly that “it is impossible to understand early genetic technologies apart from the broader history of American technology and innovation. Genetic technologies were completely entangled with other areas of innovation, both in their material production and in the outcomes anticipated from them” (Curry 2016a, 3). This is a position that bears

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resemblances to Luis Campos’ findings regarding the proliferation of radium mutation research in biology (Campos 2015) and the modernism of agricultural organisms attended to throughout Tiago Saraiva’s Fascist Pigs (2016). Earlier Barbara Hahn had already demonstrated how attention to the various technologies of tobacco cultivation and processing could provide insights into the making of agricultural commodities, with implications for the value of botanical knowledge (Hahn 2011). And even earlier Robert Kohler had emphasized the potential need to understand how organisms can become technologies, in his case for the purposes of experimental practice (Kohler 1994, 6). My suspicion is that one reason why interest in agricultural science among historians of biology has tended to wax and wane is due to too ready a willingness to dismiss those parts of organisms that look technological rather than scientific. Even though historians of biology typically demur from admitting this selectivity because they are smart enough to know that hard and fast distinctions between the importance of science and technology are difficult to defend, I nevertheless suspect that a prejudice resides here. This prejudice betrays our historical material which is at once scientific and technological however we choose to define those terms or put them together. Those who wish to see investigation of agriculture and plant breeding continue to flourish among historians of biological science would therefore do well to collaborate with historians of technology.

Conclusion I should begin my conclusion with two apologies. The first is that I can only read and write in English. There is a world of excellent scholarship beyond my reach and which you can only learn about by following references to the various special issues and edited collections that I have cited. This deficiency no doubt warps my understanding of how this historiography has developed. The second is that agriculture here has been decidedly arable. Fisheries (Evenden 2004; Jiang 2017; Schwach and Hubbard 2009), scientific management of pests (Palladino 1996; Sayer 2017), forestry (Paskins 2018; Sivaramakrishnan 2008), and animal health and husbandry (Brown and Gilfoyle 2010) deserve more attention than they have been granted and will have to be faced another day. The autonomy that I gave agriculture at the outset might be thought only an annoying rhetorical tool, enabling rival historians to always ask for more no matter how diligently a historian of biology has pursued the agricultural. It also seems to demand that when choosing the scholarly lineage to which we each dedicate our work that we have to decide between the history of biology or agriculture, rather than something interdisciplinary. Might we not instead hope that through integration between history of biology and history of agriculture, some new kind of historian will emerge, one primed to tackle these issues from the outset and able to lead both fields in exciting new directions? But this might be a misguided hope. I do not imagine many of the scholars minted in such fashion would be particularly highly prized by either side, and they would probably struggle to find independent funding.

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Assuming this to be an accurate reading of the current institutional situation, it is therefore worth recapping the value of recognizing agriculture’s distinctiveness from the history of biology. First, we do not think of there being one history of biology unfolding over time, even as we offer up our work as part of that collective endeavor. Given this multiplicity, the existence of an additional historical tradition in parallel to the history of biology, sometimes overlapping, other times diverging sharply, is not only manageable but also fruitful, because the relationship sharpens shared questions about organic growth, care, knowledge, value, health, maintenance, situatedness, biological time, and a whole host of other problems provoked by living things. Second, business historians, economic historians, social historians, historians of rural life, and all those who have investigated farms and farming have been discussing biological things even when the biology of those things was not central to their own analysis. We can therefore think of all this scholarship as a reservoir of biological thought, perception, and evaluation, one which is available for the historian of biology to exploit or bring to their aid. Doing so might not be so easy if we were expected to find ways to assimilate these arguments into our own analytical framework, as would be one aim of interdisciplinarity. Third, history has witnessed attempts to render entire parts of the agricultural enterprise a science, be it a science of economics, of heredity, of chemistry, you name it. In some cases these ambitions have been totalizing – to make them a science and only a science. While I think historians of biology are perfectly capable of submitting such ambitions to critical analysis on their own terms, it might help to also have something called a “historian of agriculture” close by, providing a scrutiny of their own. Some historians of science can at times adopt and project the voice of scientific actors, just as historians of agriculture can sometimes wittingly or unwittingly speak on behalf of farmers. These predilections are sometimes healthy, other times pathological. Recognizing what might distinguish agricultural history questions from history of science questions, and retaining expertise in them respectively, is no doubt mutually beneficial and a way of mitigating the worst excesses, by providing a check on unreasonableness. This chapter has been designed as a primer, one that sensitizes newcomers to the intersection of the history of biology and the history of agriculture. Where I have explained that a particular theme or question has been thoroughly explored, I should not be read as arguing “to the point of saturation.” New cases and angles of analysis are emerging all the time providing ample opportunity for revisionism in even the most well-trod spaces. We have yet to see, for instance, where the integration of queer theory into ecocriticism and environmental humanities may lead us historiographically (Sandilands 2014; Gray et al. 2016). We are fortunate to have one clue in the shape of Luis Campos’ work on sexuality for understanding twentieth-century plant genetics (Campos 2010), but vastly more awaits to be discovered. I have made the case for the importance of three particular areas in the future of the history of biology and agriculture: global histories, history of the environment, and the intersection of biology and technology. If we are able to find the resources for global histories of biological science, then we can learn the who, how, and what has been

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taken to matter for different states independently and as a whole regarding agricultural life and production and the kinds of biological science that they have inspired. I have also highlighted the need for a fuller integration of the history of agricultural science and industries with the history of the environment. Historical claims regarding capitalization, commercialization, and biological technology look starkly different when approached from the history of biology and the history of the environment. Last, I have emphasized that agricultural contexts provide a suite of cases for fuller exploration and understanding of the relations between biology, biological science, and technology. There are deeply significant conceptual issues at stake regarding organisms, science, and technology, and we can champion the history of plant breeding and agriculture’s value for addressing them. Doing so also brings historical work into service in the present, for so much contemporary discourse is designed to swiftly conflate biology and technology only either to ignore plant breeding and agriculture’s past or invent one for it that serves narrow interests. Acknowledgments This chapter was written while the author was a Research Fellow on the Narrative Science project, www.narrative-science.org, and received funding from the ERC under the European Union’s Horizon 2020 research and innovation program (grant agreement No. 694732).

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Index

A Abir-Am, Pnina, 63, 67 A Body Worth Defending, 404 Acceleration technologies, 251 Adams, Mark, 136, 140 Agriculture and Horticultural Society of New South Wales, 376 A Guinea Pig’s History of Biology, 274 A History of Immunology, 398 Akeley, Carl, 304 Allen, Ann Taylor, 140 Allen, Garland, 89, 135, 136, 269 Altruism, 302 American Medical Association, 201 American phage group, 180 Amis, Martin, 291 Anderson, Warwick, 399 Anglicanism, 18 Anglophone historiography, 281 Anglophone scholarship, 269 Animal electricity, 426 Animal physiology, 486 Annals of Eugenics, 134 Annals of Science (1936), 110 Anti-Bolshevism, 133 Anti-Semitism, 133 Anti-vivisection movement, 450 Apical ectodermal ridge (AER), 97 Arabidopsis, 182 Arabidopsis thaliana, 276 Aretology, 294 Artisanal knowledge, 250 Ash, Mitchell, 136 Assembly line approach, 479 Astbury, William, 178, 180 Atomistic approach, 417 Aubin, David, 303 Australian Society, 376 Autobiography(ies), 178, 298

Autoimmunity, 398, 404 Automata theory, 418 Aveling, Edward, 17 Avery, Oswald, 179

B Bacon, Francis, 303 Bacteria, 180 Bacteriophages, 180 Baker, J.R., 134 Balanced aquarium, 451 Baldwin, Ernest, 71 Barkan, Elazar, 144 Barlow, Nora, 10 Basalla, George, 372, 374 colonial science, establishment of, 375 European science, expansion of, 375 implementation and critique of Basalla’s model, 377 local independent scientific tradition, development of, 375 modern science, 374 Bateson, William, 109 Baur, Erwin, 133 Bayh-Dole Act of 1980, 225 Beach vacation, 437 Beadle, George, 178 Beagle voyage, 10 Bechtel, William, 181 Beer, Gillian, 15 Behavioral ecology, 51 Bell, Alexander Graham, 131 Benacerraf, Baruj, 396 Ben-David, Joseph, 373 Bennett, Dorothea, 181 Bentley, H., 111 Benzer, Seymour, 180 Bernard, Claude, 180, 270

© Springer Nature Switzerland AG 2021 M. R. Dietrich et al. (eds.), Handbook of the Historiography of Biology, Historiographies of Science, https://doi.org/10.1007/978-3-319-90119-0

527

528 Between Bench and Bedside, 403 Biagioli, Mario, 303 Bibliometric tools, 196 Bigg, Charlotte, 303 Biocapitalism, 229 Biographers, 299–302 The Biographical Memoires of Fellows of the Royal Society, 306 Biography(ies), 178, 292 art of, 307–308 as form of eulogy, 306 future of, 308–310 science and, 305 of scientists (see Scientific biography) uses of, 302–307 Bioi, 294 Bioinformatics, 185 Biological oceanography, 439 land-based research, 445 ship-based marine research, 438 transnational politics, 449 Biology, 394, 397–400, 405 Biomedical platforms, 403 Biomedical research, as weapon in Cold War, 203 Biomedicine after Second World War, 197 concept, 197 definition, 195 distribution of activity in, 208–213 historiography, 196 physician’s degrees, 199 postwar, 199 postwar development, 201 researchers and policy-makers, 196 research funding, 207 20th century historiography, 210 of United States, 196 Biotechnology, 500, 506, 507 definition of, 218 first generation of scholarship, 223 historical trends in, 219 historiography of, 218, 219 popular conceptions of, 220–223 revolution, 235 second generation of scholarship, 219 third generation of, 236 third generation of scholars, 219 Biotechnology: The University-Industrial Complex, 224 Biotech Revolution, 223–228 Bio-typology, 139

Index Blacker, Carlos Paton, 134 Bleier, Ruth, 331 Blyth, Edward, 13 Bock, Gisela, 141 Body schema, 423 Bonner, John Tyler, 97 Boudia, Soraya, 372 Bowler, Peter J., 21, 115 Brachypodium distachyon, 276 Brain, Robert, 254 Brannigan, Augustine, 109 Brauckmann, S., 86 Brecht, Bertolt, 300 Brenner, Sydney, 180, 182, 266 British Association for the Advancement of Science, 131 British Broadcasting Company, 221 Britten-Davidson model, 181 Broca’s research on brain, 424 Bud’s etymological story, 226 Bulloch, William, 395 Burian, Richard, 176, 270, 272 Burkhart, Richard, 305 Burnet, Frank Macfarlane, 199, 396, 397, 399–402, 404 Butterfield, Herbert, 297

C Caenorhabditis elegans, 275 Cambrosio, Alberto, 231, 257 Canguilhem, Georges, 394, 403 Carnegie, Andrew, 22 Carnegie Trusts, 131 Carol, Anne, 140 Carr, E.H., 299 Cell biology, 181 legacy of, 181 Cell lineage studies, 91 Cell theory and neuron doctrine, 421–422 Cellular immunology, 399 Central dogma, 188 Challenger expedition, 438 Changing Patterns: An Atypical Autobiography, 396 Charles Darwin’s theory, 129 Chase, Allan, 141 Chemical embryology, 84 Chiang, Hsue-Hao, 72 Chinese science and technology, 374 Chromosomal engineering, 134 Chronophotographic techniques, 476

Index Churchill, Frederick, 88 Clarke, Adele, 267 Clause, Bonnie, 272 Cloned sheep (Dolly), 222 Coerced sterilization, 146 Cold Spring Harbor experimental station, 131 Cold War, 372, 374, 380, 385, 386 generation biography, 297 Cole, Francis Joseph, 84 Coleman, William, 89 Comfort, Nathaniel, 270, 301 Computational neuroscience, 419–420 Comte, August, 130, 296 Conflict thesis, 18 Connectionist modelling, 420 Cooke, Kathy, 280 Cortical localization, 413–415 Cowan, Ruth Schwarz, 130 Creager, Angela, 179, 230, 256, 276 Creative Couples in Science, 327 Creative maladies, 13 Creese, Mary R. S., 325 Crick, Francis, 177, 180, 188 Crystal Palace, 20 Cultural-historical poetics, 294 Cybernetics, 419, 420 Cytoplasmic inheritance, 270

D Darlington, Cyril Dean, 134, 302 Darwin, Charles, 8, 14, 113, 218, 305 bird breeders and fanciers, 16 Chicago meeting, 13 common context, 15–16 Darwin Correspondence Project, 17 Darwin’s theory-building, 11 evolution, 11 evolutionary synthesis, 14 hero of science, 9–10 natural theology, 17 Origin of Species, 17, 20 religion, 16–20 revolution, 20–21 social Darwinism, 22 The Descent of Man, 18 transmutation notebooks, 11 Victorian medicine, 13 Darwin, Francis, 9 Darwinian evolution, 37, 117 Darwinian revolution, 8 Darwinism, 300, 376, 500–502

529 Darwin’s Century, 298 Darwin’s theory of evolution, 86, 420 Davenport, C.B., 131 Dawson, Gowan, 16 de Beer, Gavin, 110 De Chadarevian, Soraya, 183, 233 The Death of Nature, 330 Deep-sea biology, 441, 444 Delbrück, Max, 178, 180, 266 Descartes, Rene, 295, 303 Desmond, Adrian, 298 Developmental biology, 82 embryology transition to, 94–95 evolutionary, 97 molecular aspects of, 95 non-molecular narratives in, 95–97 DeVries, Hugo, 277 de Vries’s law of dominance, 108 Dictionary of Scientific Biography, 297 Dietrich, Michael R., 62, 278, 303 Dinsmore, Charles, 91 Discours de la Méthode, 295 Discursive practices, 245 Dissolution, 421 Dohrn, Anton, 88 Doppler, Christian, 116 Down’s syndrome, 421 Doxographical writings, 294 Draper, John William, 18 Drosophila, 182 D. melanogaster, 266 Dubos, René, 176 E Eckart, Wolfgang, 137 Edmond de Rothschild Foundation, 179 Edna Suarez-Diaz, 185 Eighth Day of Creation, 270 Eimer, Theodor, 19 Electron microscopy images, 179 Electrophoresis, 71–72 Eliot, George, 15 Ellis, Joseph, 301 Embryology in 18th and early 19th centuries, 83–86 experimental, 88–89 in late 19th centuries, 86–88 Eminent Victorians, 307 ENCODE consortium, 186 Encomia, 294 Endersby, Jim, 274, 304 Environmentalism, 380

530 Environmental physiology, 487 Ephrussi, Boris, 178, 180 Epigenetics, 186 Epistemic Cultures: How the Sciences Make Knowledge, 248 Epistemology of the Concrete, 253 Equipotentiality, 413 Eugenicists, 130, 131 Eugenics, 23, 129, 506 critics of, 143–144 definition, 128 evolution and, 128–134 history of, 128 Nazi, 145 negative, 138–140 permeation of public health, 132 positive, 138 post WW1, 142 scientific critics of, 132 Eugenics Education Society, 138, 142 Eugenics Review, 134 Eulogies, 294 Euro-centered model, 375 Evans, Richard, 140 Evidence-based medicine studies, 209 Evo-Devo, 184 Evolutionary biology developmental biology, 40–43 ethology and behavior, 48–50 histories of selection, 45–46 long modern synthesis, 36–40 sexual selection, 46–48 sociobiology, 51–52 systematics, 43–45 Evolutionary developmental biology, 97 Evolutionary genetics, 162–163 Evolutionary synthesis, 36–40 Evolutionary theory and brain, 420–421 Experimental biochemistry, 253 Experimental embryology, 88 Experimental life sciences, 248 experimental systems and epistemic things, 250–251 model systems and biomedical platforms, 255–258 philosophers’ and historians’ concerns, 253 studies of visualization, 258–259 temporality and experimental systems, 251– 255 Explorations in Connected History, 383 Exploratory experimentation role of, 253 vs. theory-driven experimentation, 252

Index F Fantini, Bernardino, 270 Farley, John, 84 Farmelo, Graham, 305 Feminism and Science, 330 Finnegan, Diarmid, 380 Fischer, Eugen, 133 Fisher, R. A., 110 Fitzgerald, Deborah, 507 Fleck, Ludwik, 246–248, 398, 405 Florkin, Marcel, 71 Folia Mendeliana, 116 Fortun, Michael, 231 Foster, William, 395, 396 Foucault, Michel, 112, 121, 178 Fragmentation, 94 Franklin, Sarah, 236 French physiology, 479 Friese, Carrie, 280 From Caracas to Stockholm, 396 Fujimura, Joan, 228, 267

G Galton, Francis, 23, 113, 128 Galton Laboratory, 131 Gastrulation, 86 Gaudillière, Jean-Paul, 230 Gaukroger, Stephen, 295 Gaussian curve, 130 Gayon, Jean, 270 Geison, Gerald, 271 Gender analysis, 332 Gene(s) in action, 161–162 concepts, 159–161 in Drosophila development, 178 Genesis and Development of a Scientific Fact, 246, 398 Genetic(s), 503–506 after Second World War, 179 classical, 155–159 code, 183 and development, 161–162 evolutionary, 162–163 father of, 106 history and memory, 154–155 human, 163–165 Genetic Alchemy: The Social History of the Recombinant DNA Controversy, 224 Genetically modified organisms (GMOs), 222

Index Genetic engineering, 177 techniques, 181 tools, 182 Genome sequencing, 176 program, 184, 186 Genomics, 185 rise of, 177 German eugenicists, 141 Gesamte physiology, 470 Gilbert, Scott, 86, 276 Global Burden of Disease studies, 205 Globalization, 372 Goldschmidt, Richard, 303 Golgi’s advocacy of recticular theory, 421 Gopnik, Adam, 294 Gotto, Sybille, 140 Gould, Stephen Jay, 14 Govoni, Paola, 301, 326 Graham, Loren, 140 Graphical method of physiology, 470 Gray literature, 253 Green Revolution, 508 Gregor Mendel Memorial Symposium, 115, 116 Griesemer, James, 279 Griffith, Frederick, 179 Groeben, Christiane, 88 Guggenheim Foundation, 200 Gütt, Arthur, 139

H Hacking, Ian, 258 Haeckel, Ernst, 87 Hagiography, 309 Haldane, J.S., 304 Haller, Mark, 134 Hamburger, V., 93 Hankins, Thomas, 298 Haraway, Donna, 93, 247, 248, 331 Hardy, Thomas, 132 Harrison, Ross Granville, 85 Harwood, Jonathan, 135, 270 Hauptmann, Gerhart, 132 Hausen, Karin, 141 Hebbian learning, 424 Heimans, Jacobus, 110 The Helmholtz Curves, 251 Herbert Spencer’s theory of evolution, 420 Hereditarian ideas, 120 Heredity, 118, 129 Herran, Néstor, 372

531 Hertz, Friedrich, 143 Hessen, Boris, 116 Heterozygosity, 62, 70 Hilgartner, Stephen, 231 Himmelfarb, G., 13 Histories of Western science, 3 Historiography assessment of, 4 of biology, 4 description, 1 early, 346 and immunology (see Immunology and historiography) initial, 348 marine station, 445 ocean exploration, 437 robust, 445 of science, 361 traditions, 2 tragedy of the commons, 442 voluminous, 363 western, 361 History of biology, 2–4, 34, 35, 37, 42, 154, 165 growth and diversification of, 4 History of science Basalla, George (see Basalla, George) circulation and collaborative networks, 380–382 in developing countries, 379 effects of globalization, 378 epistemological and sociological considerations, 376 global historians, 379 international nongovernmental/ intergovernmental organizations, 384 international policy, 374 international relations and colonialism, study of, 373 localists/nationalists and imperialists, 377 nation-state histories, 384 Needham, Joseph, 374 non-European science and technology, 379 Pyenson, Lewis, 377 science and imperialism, 377 and socio-political factors, 378 Spanish engineers, 385 transnational history, 383 transnational perspective in, 378 universality of scientific knowledge, 373 worldwide circulation of knowledge, 379 Hofstader, Richard, 13 Holmes, Frederic Larry, 177, 272, 305 Holmes, Larry, 253

532 Holtfreter, Johannes, 92, 96 Homo cerebralis, 427 Hooker, Joseph Dalton, 16 Hopwood, Nick, 83 Horder, Tim, 83 Horowitz, Maryanne Cline, 321 HSS Women’s Committee, 322 Hughes, Sally Smith, 233 Human genetics, 163–165 Human Genome Project (HGP), 73, 222, 228, 231 Humoral immunity, 396 Huxley, Aldous, 132, 221 Huxley, Julian, 132, 134 Huxley, Thomas Henry, 10, 143 Hypothesis-driven experiments, 176

I The Immune Self, 400, 401 Immunity, 394, 395, 403, 406 cellular, 398 ecological histories of, 403 humoral, 396, 398 multidimensional history of, 403 science, 396 Immunodiffusion, 72 experiments, 66 Immunology and historiography Bulloch, William, 395 Burnet, F. Macfarlane, 396 Fleck, Ludwik, 398 Foster, William, 395, 396 Jerne, Niels K., 397 Mazumdar, Pauline M.H., 395 Medawar, Peter B., 396 Moulin, Anne-Marie, 399 Silverstein, Arthur M., 398 Tauber, Alfred I., 398, 400, 401 Immunophenotyping, 257 Individuality, 394, 398–400, 403 Industrial physiology, 477 Inkster, Ian, 376 Institute for Physical-Chemical Biology, 179 Intellectual property (IP), 505, 506 Interlaboratory constitution, 257 International agencies espoused eugenics, 142 International commission of the history of oceanography (ICHO), 439 International council for the exploration of the sea (ICES), 444 Interwar international agencies, 143 Intolerant Bodies, 404

Index J Jablonka, Eva, 14 Jacob, François, 112, 180, 251 Jefferson, Thomas, 301 Jerne, Niels K., 397 Jones, Greta, 136 Journal for the History of Biology, 272 Judson, Horace Freeland, 175, 270

K Kass-Simon, Gabriele, 318 Kay, Lilly, 72, 230 Keating, Peter, 231, 257, 399 Keller, Evelyn Fox, 93, 187, 269, 298 Kendrew, John, 180 Kevles, Daniel, 135, 221, 227 Kimmelman, Barbara, 505, 506, 509 Kimura, Motoo, 66 Kirk, Robert, 254, 257, 277 Kloppenburg, Jack Ralph Jr., 225 Kohler, Robert, 256, 272, 273, 381, 510, 515 Kohlstedt, Sally Gregory, 331 Koyré, Alexander, 297, 300 Krimsky, Sheldon, 224 Kříženecký, Jaroslav, 110 Kroeber, Alfred L., 112 Kühn, Alfred, 276 Kuhn, Thomas, 115, 246 Kundera, M., 293 Kushner, Tony, 144

L Labisch, Alfons, 139 Labography, 309 Laboratory Life: Construction of a Scientific Fact, 247 Laboratory organisms, 236 Lamarckian tradition, 180 Landecker, Hannah, 236 Landlubber, 437 Lapore, Jill, 301 Latin eugenics, 139 Latour, Bruno, 247, 277 Laubichler, M.D., 98 Lederberg, Joshua, 221 Lederberg, Muriel, 272 Lederman, Muriel, 272 Lemkin, Raphael, 138 Lenoir, Timothy, 86, 270 Lenz, Fritz, 133 Lenz, Widukind, 135

Index Leonelli, Sabina, 276 Levine, George, 15 Lewontin, Richard C., 62, 400 Lewontin historical thesis, 62 The Life of a Virus: Tobacco Mosaic Virus as an Experimental Model, 1930-1965, 256 Linnaeus, Carl, 113 Litchfield, Henrietta, 10 Literary poetics, 294 Livingstone, David, 381 Localization cortical, 413–415 functional specialisation of nerves, 415 theories of, 412 Locke’s theory of personal identity, 427 Logan, Cheryl, 277 Logic of Life: A History of Heredity, 112 Lords of the Fly: Drosophila Genetics and the Experimental Life, 256, 273 Lovejoy, Arthur O., 111 Löwy, Ilana, 397, 403 Lucas, Prosper, 118 Ludwig’s physicochemical approach, 470 Luria, Salvador, 180 Lwoff, André, 180 Lysenkoism, 116 Lysogeny, 180

M MacCord, Kate, 86 Mackay, Ian R., 402 Mackenzie, Donald, 131, 135 Magendie’s experimental approach, 469 Maienschein, Jane, 98 Malpighi, Marcello, 85 Malthus, T.R., 15 Mansfield amendment, 202 Marine biological laboratory (MBL), 448 Marine biology, 436 aquariums, 452 botanical collecting and sharing, 451 captive marine mammals, 450 colonial coastal environment, 451 embryological research, 449 fisheries biology, 439 fish stocking, 452 historiography, 438 kraken, 453 marine resources, 445 marine stations 1910-1995, 446–448 marine stations 2002-present, 448–450

533 Maury conferences, 440 ocean, 437 oysters, 446 public interaction, 450 realm, 452 salmon, 442 terrestrial animals, 450 turtle conservation, 443 water parks, 452 Marine invertebrates, 87 Marine mammal protection act, 443 Martin, Emily, 93 Maximum sustainable yield (MSY), 443 Mayr, Ernst, 13–14, 114 Mazumdar, Pauline M.H., 136, 395, 399 McCain, Katherine, 278 McClintock, Barbara, 266, 270, 298, 299, 302 Mckie, Douglas, 296 McKusick, Victor, 185 Medawar, Peter B., 221, 396, 399 Medicine and nervous system, 422–424 Mehler, Barry, 135, 136 Memoir of a Thinking Radish, 396 Memorials, 295 Mendel, Gregor, 106, 133, 503 Mendelian genetics, 271 Mendelian revolution, 119 Mendelian scholarship, 110 Mendelian theory, 252 Mendelism-Biometry debate, 110 Mendelsohn, Andrew, 273 Mendelsohn, Everett, 89, 320 Mendel, stuck in time, 112–120 Mengele, Josef, 133 Mental health research funding, 207 Merchant, Carolyn, 330, 331 Meselson, Stahl and the Replication of DNA, 253 Metabiography, 310 Metagenomic(s), 187 techniques, 252 Metaphysical concept, 96 Méthot, Pierre-Olivier, 179 Meyer, Arthur William, 85 Microbiology, 395 Micro-fixation technique, 69 MicroRNAs, 188 Microscopic morphology, 448 Mill, John Stuart, 130 The Mind Has No Sex? Women in the Origins of Modern Science, 331 Mitchell, Peter, 181 Mitman, Gregg, 404

534 Mitman’s approach, 50 Modern synthesis, 36–40, 98 of Mendelian genetics, 111 Molecular biologists, 180 Molecular biology, 181, 249, 253 characteristic of, 179 contributions to, 179 definition, 177 field of research, 179 historiography, 176, 177 history of, 176 structural face of, 182 transformations of, 182 Molecular evolution Abir-Am, Pnina, 63 biographical profiles and interviews of the main actors, 65 debates, confrontations, and negotiation, 67–70 Dietrich, Michael R, 62 exobiology and the early evolution of life, 64 heterogeneity, 63 informational molecules, 64 Lewontin, Richard C., 62 socio-professional dynamics, 65 technologies and techniques, 70–74 Molecular wars, 67 Momigliano, Arnaldo, 297 Monod, Jacques, 180 Monte Carlo experiments, 74 Morbidity, 207 of mental health, 208 Morgan, Lynn., 93 Morgan, Thomas Hunt, 91, 266, 269 Morsink, Johannes, 321 Mortality, 208 Mosse, George, 137 Moulin, Anne-Marie, 399 Muller, Hermann J., 221 Müller-Hill, Benno, 135 Mullins, Nicholas, 270 Multiculturalism, 372 Multispecies ethnographies, 282

N Naegeli, Carl, 110 Napier, A. David, 402 Napp, Cyril, 117 Nasar, Sylvia, 305 National and transnational narratives, 165–166 National Cancer Institute (NCI), 199

Index National Health Examination Survey, 205 National Heart Institute, 207 National Institute of Allergy and Infectious Disease (NIAID), 199, 207 National Institute of Mental Health (NIMH), 199, 207 National Science Foundations (NSF), 197 research budgets, 198 National scientific policies, 382 Natural selection, 10 Navy-sponsored research projects, 200 Nazi eugenics, 145 Nazi German sterilization, 139 Nazi hegemony, 132 Needham, Joseph, 84, 374 Negotium, 295 Neo-Darwinian synthesis, 36–40 Neo-Lamarckian tradition, 180 Neo-lamarckism, 19 Neutralist/selectionist controversy, 60, 67, 68 Neutral theory of molecular evolution (NTME), 60, 62 New Deal Public Health Service (PHS), 201 Newgenics, and resources, 146–147 Nietzschean ideas, 140 NIH grant success rates and budget, 202 NIH Institutes, budgets of, 198 Nisot, Marie-Thérèse, 134 Non-Western science, 378 Nowak, Martin, 306 Nucleic acid hybridization, 66, 72 Nucleic acid sequencing, 73 Nuttall, George Henry Falkiner, 70 Nye, Robert, 333 Nyhart, Lynn, 270

O Ochsner, Albert, 138 Ogilvie, Marilyn, 323 Ohta, Tomoko, 66 Olby, Robert, 176, 177, 182 Oldroyd, David, 12 O’Malley, Maureen, 187 Onaga, Lisa, 281 Oncogenes, 184 One gene–one enzyme (protein) relation, 178 Open Science movement, 268 Oppenheimer, Jane, 85–86, 88, 94 Orel, Vítězslav, 115 Oreskes, Naomi, 333 Organismal individuality, 394 Organisms, in experimental research

Index Anglophone histories, 280 biographies, research fields and national trends, 269–271 20th century biological science, 266 cloning, 280 in early 1990s, 267 early 20th century breeding research, 280 historical and sociological approaches, 279 historical research, 279 historiographical trend, emergence of, 272 history of experimental organisms, 268 Human Genome projects, 266 in late 1990s, 267 methodological innovation, 278 non-human organisms, treatment of, 267 non-Western settings, 281 observational and interventionist strategies, 266 philosophically-inspired approaches, 279 quantitative methods, 278 as research practices, 275–278 Origin of Species (1859), 107 Origins of Mendelism, 113 Otium, 295 Otter hunting, 444 Oxford Handbook of the History of Eugenics, 147

P Paley, William, 17 Pangenesis, 129 Paramecium aurelia, 276 Pasteur, Louis, 277 Pasteurization of France, 247 Patiniotis, Manolis, 373, 378 Paul, Diane, 135, 146 Pauly, Philip, 229 Perceptron model, 420 Perutz, Max, 180 Phage Group, 270 Philosopher vs. scientist, 294 Philosophy, 394, 400, 401, 403 Physical anthropology advent of, 350 character of, 348 influential career in, 356 prior to World War II, 348 progressive antiracism, 356 public and academic audiences, 349 transformation of, 350 Physical oceanography, 438 Physiological optics, 422

535 Physiology animal, 465 Bernard and Ludwig, 469–470 definition, 464 gender and race, 477–480 histories of, 463 as independent science, 467–472 as practice, 480–484 as society, 472–480 as study of life, 464–466 vitalism/mechanism and national stories, 470–472 Pickering, Andrew, 247 Plant breeding Mendelian experimental system and strategy, 252 microtechniques, 252 Plant breeding and agriculture bio-agricultural economics and politics, 507–508 bio-agricultural field science, 510–511 biology and technology, 514–515 Darwin and breeders, 501–503 environment, 513–514 eugenics and biotechnology, 506–507 global histories, 512–513 Mendel and genetics, 503–506 Plato’s notion, 130 Ploetz, Alfred, 130, 133 Pluripotency, 394 Podolsky, Scott H., 401 Political philosophy, 474 Population structure, 128 Post-genomics, 186, 187 era, 177 Post-war German genetics community, 135 Post WW1 eugenics, 142 Practices definition, 244 discursive, 245 plant breeding, 252 recombinant DNA techniques/cell fusion, 249 science-as-practice, 249 temporal, 251 value, 245 Pradeu, Thomas, 401 Pre-Galtonian eugenics, 130 Pre-Mendelian heredity, 118 “Presence/absence” paradox, 293 Primate trichotomy, 69 Prodger, Philip, 16 Protein finger printing, 70

536 Protein folding, 188 Protein sequencing, 64, 73 Protozoology, 84 Psychophysiological research, 276 Puericulture, 139 Purkyňe, Jan, 117 Pyenson, Lewis, 377 Pythagorean lifestyle, 294

Q Quality adjusted life year (QALY), 205 Quantitative analysis, 441

R Rabinow, Paul, 228 Race, 344 in America, 351–360 in science, 351–360 Racial hygiene, 130 Racial Hygiene Society, 142 Rader, Karen, 275 Ramsden, Edmund, 277 Ranke, Leopold von, 295 Rasmussen, Nicolas, 230, 234 Rassenhygiene, 136 Recombinant DNA, 221, 224 patents, 221 technologies, 218 Reflex, 416 Canguilhem’s work, 416–417 Regulatory RNAs, 188 Reiss, Christian, 277 Renaissance, 295, 321 Representing and Intervening, 258 Reworking the Bench: Research Notebooks in the History of Science, 253 Rheinberger, Hans-Jörg, 179, 250, 251 Richards, Dickinson, 199 Richardson, Jane S., 183 Ridley, Matt, 306 Rifkin, Jeremy, 222 The Right Tools for the Job: At Work in the Twentieth-Century Life Sciences, 272 Ritvo, Harriet, 278 Rockefeller Foundation, 132 program, 179 Rockefeller Foundation and Institute, 197 Röntgen, Conrad, 138 Rosenfeld, Albert, 222 Rossiter, Margaret, 324, 334

Index Roux, Wilhelm, 84, 88 Ruddle, Frank, 185 Rüdin, Ernst, 138, 139 Russell, Bertrand, 301 Russell, Edward Stuart, 85

S Saleeby, Caleb Williams, 138 Sanger, Margaret, 133 Sapp, Jan, 270 Sarton, George, 296, 319 Schaffer, Simon, 397 Schiebinger, Londa, 323, 331, 332 Schloegel, Judy Johns, 276 Schmidgen, Henning, 276 Schneider, William, 136 Schreiber, Bernhard, 135 Schrödinger, Erwin, 178 Science and Technology in the European Periphery (STEP), 382 Science and technology studies (STS), 372 transnational approach of, 372 Science as Autobiography: The Troubled Life of Niels Jerne, 397 Scientific biography, 292, 296, 304, 309 bibliographical essays, 293 biographical writing, 292 Brecht, Bertolt, 300 Comfort, Nathaniel, 301 Dictionary of Scientific Biography, 297 genre of, 293 Keller’s biography, 298 practitioners of, 302 skepticism of, 297 Thackray’s estimation, 296 Woolf, Virginia, 308 Scientific diving, 441 Scientific Revolution, 319, 374 Scopes Trial, 19 Scott, Joan, 330 Seaweed, 451 Secord, James, 380 Self, 394, 396, 400–402 Self-experimentation, 254 Serological taxonomy, 66 Seutonian vs. Plutarchian approaches, 308 Sexual selection, 46–48 Shapin, Steven, 254, 274, 333 Sheep breeding, 277 Shelley, Mary, 187, 221 Shils, Edward A., 373

Index Shipboard survey, 440 Shiva, Vandana, 222 Signs: A Journal of Women and Culture, 323 Silverstein, Arthur M., 398, 402 Situated knowledge, 247 Small interfering RNAs (siRNAs), 188 Social construction of science, 248 Social Darwinism, 22–23, 129 Social history, 2, 304 Social Spencerism, 22 Sociobiology, 51–52 Sociological approach, 469 Söderqvist, Thomas, 292, 296, 305 Speciation, 39 Species and Specificity, 399 Spemann, Hans, 92 Spencer, Herbert, 21, 22 Sperm whaling, 440 Spinally-mediated reflexes, 418 Spontaneous history, 109 Standard narrative, 98 Standing Committee on Women, 322 Star, Susan Leigh, 279 Stem cell technologies, 222 Stepan, Nancy, 136 Sterilization guidelines, 139 Stern, Curt, 110 The Strangest Man, 305 Strasser, Bruno, 71, 257 Strickland, Stephen, 201 Structural biology, 182 The Structure of Scientific Revolutions, 246, 297 Subrahmanyam, Sanjay, 372 Sunderland, Mary, 97 Sutherland, Earl Jr., 199 Synthetic biologists, 187 Synthetic biology, 187 Systematics, 43–45

T Talmudists, 302 Tatum, Edward, 178 Taub, Lisa, 294 Tauber, Alfred I., 398, 400, 401 Taylor, Gordon Rattray, 222 Temporal orientation, 251 Temporal practice, 251 Tennstedt, Florian, 139 Thackray, Arnold, 231, 296, 320, 322 “The Art of Biography”, 307

537 Thieler, Max, 199 Thompson, E.P., 274 Tickle, Cheryll, 97 Timofeev-Ressovsky, Nikolay, 115 Tolin, Sue, 272 Towards a History of Epistemic Things, 250 Transmutationism, 20 Trevor-Roper, Hugh, 135 Trinkaus, John P., 96 Truman, Harry, 201 Tuchman, Barbara, 308 Tumour suppressor genes, 184 Turchetti, Simone, 372 Twain, Mark, 292, 293

U Unger, Franz, 116 US Civil Rights movement, 134 U.S. National Institutes of Health, 266

V Value practices, 245 Venter, Craig, 304 Verschuer, Otmar von, 133 Vettel, Eric, 233 Victorian visual culture, 16 Vienna eugenics, 141 Viruses, 180 Vogt, Annette, 325 von Bertalanffy, Ludwig, 186 Voskuhl, Adelheid, 255 Voss, Julia, 16

W Waddington, Conrad H., 115 Wallace, Alfred Russel, 21 Wartime Committee on Medical Research, 201 Wassermann reaction, 246 Watson, Jim, 180 Weber, Marcel, 276 Weiling, Franz, 110 Weingart, Peter, 136 Weismann, August, 88, 113 Wells, H.G., 221 Werskey, Gary, 309 Western science, 376 Whitehead, Alfred North, 294 Wilderness ethic, 453 Willier, Benjamin, 88

538 Wilson, Adrian, 255 Women in science, 323 academic programs, 320 Aristotle’s biological view, 321 biographical dictionaries, 325 biographical studies, 326–327 biology education and teaching, 328–329 Creese, Mary R. S., 325 feminist studies of biology, 329–332 gender dynamics, 333 higher education, 327 late 1960s, 320 life sciences, 327, 328 nature writers, 328 Rossiter, Margaret, 324 scholarship on women, 323–324 scientific work, participation in, 318 second-wave feminism, 319 in twentieth century, 319 women’s movement and history of biology, 321–323

Index Women Scientists in America: Struggles and Strategies to 1940, 324 Woolf, Virginia, 298, 307, 308 Woolgar, Steve, 247

Y Years of Productive Life Lost before age 65 (YPLL65), 206 Young, Robert M., 15, 135, 298

Z Zallen, Doris, 270 Zita, Jacqueline, 329 Zola, Emile, 132 Zollschan, Ignaz, 144 Zone electrophoresis, 72, 75 Zone of polarizing activity (ZPA), 97 Zymotechnology, 226