George C. Williams and Evolutionary Literacy (Literatures, Cultures, and the Environment) 3031116496, 9783031116490

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LITERATURES, CULTURES, AND THE ENVIRONMENT

George C. Williams and Evolutionary Literacy

Michael P. Cohen

Literatures, Cultures, and the Environment Series Editors

Ursula K. Heise Department of English University of California Los Angeles, CA, USA Gisela Heffes Rice University Houston, TX, USA

Literatures, Cultures, and the Environment focuses on new research in the Environmental Humanities, particularly work with a rhetorical or literary dimension. Books in this series explore how ideas of nature and environmental concerns are expressed in different cultural contexts and at different historical moments. They investigate how cultural assumptions and practices, as well as social structures and institutions, shape conceptions of nature, the natural, species boundaries, uses of plants, animals and natural resources, the human body in its environmental dimensions, environmental health and illness, and relations between nature and technology. In turn, the series makes visible how concepts of nature and forms of environmentalist thought and representation arise from the confluence of a community’s ecological and social conditions with its cultural assumptions, perceptions, and institutions.

Michael P. Cohen

George C. Williams and Evolutionary Literacy

Michael P. Cohen Professor Emeritus Southern Utah University Reno, NV, USA

Literatures, Cultures, and the Environment ISBN 978-3-031-11649-0    ISBN 978-3-031-11650-6 (eBook) https://doi.org/10.1007/978-3-031-11650-6 © The Editor(s) (if applicable) and The Author(s), under exclusive licence to Springer Nature Switzerland AG 2022 This work is subject to copyright. All rights are solely and exclusively licensed 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. Cover credit: Valerie P. Cohen, Watercolor This Palgrave Macmillan imprint is published by the registered company Springer Nature Switzerland AG. The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland

To Sarah Blaffer Hrdy, who kept me going and Ronald M. Lanner (1930–2022), who taught, “Never give up.”

Acknowledgments

Decades ago, I had the chance to discuss the task of teaching scientific literacy with John W. Little. He and his colleague Roy Parker provided a guide for graduate students in biochemistry, entitled “How to Read a Scientific Paper.” I realized that this skill should also be inculcated in teachers and students in the environmental humanities, including myself. My engagement with environmental historians, and especially with Donald Worster, Richard White, and William Cronon, furthered this interest in integrating scientific literatures into the writing of the humanities. How indeed should one become scientifically literate? In this book I suggest a model or procedure—certainly not the only one, but a start. For some time, I had been interested in the literature that emerged from the so-called modern synthesis of evolutionary theory. About twelve years ago, when I began to focus specifically on literature on animal behavior, I was encouraged and facilitated especially by Ursula Heise, who kindly invited me to speak to her seminar at Stanford University. At home, Cheryll Glotfelty at the University of Nevada has been steadfast in furthering this work. Animal behavior is such a vast subject that I needed a foundation for my study’s compass. As I read contemporary sources, it was clear that much of what counted as evolutionary literacy was grounded in the late twentieth-­century by the writings of George C. Williams. I was drawn to his books because they modeled critique in the biological sciences and because of the understated clarity of his prose style. I began to correspond with some who had known him, contemporaries and students who had vii

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worked with him, and also conferred with some of my closest colleagues in the humanities. By a remarkable accident, I discovered that Sarah Blaffer Hrdy, who supplies our family with walnuts, read this manuscript for another press. She re-invigorated the project and led me to Randolph Nesse. This reading of Williams has also been encouraged by Laura Betzig, SueEllen Campbell, Steve Frank, Douglas J.  Futuyma, William F.  Harms, Tim Lewins, Jonathan Kramnick, Kurt Schwenk, Vassiliki Betty Smocovitis, and Stephen Stearns. Doris Williams has patiently answered many of my questions; she, along with Karen Leibowitz and Kristen J. Nyitray at Stony Brook University, helped me acquire appropriate photographs of George C. Williams. Sarah Blaffer Hrdy, Randolph Nesse, Jon Christensen, and Michelle Niemann read, commented on, and facilitated improvements for the complete manuscript. I am in their debt, but of course any errors are mine alone. This book would not have been possible without the guiding hands of my editor, Michelle Niemann, on whose patience, perspicacity, precision, and good sense I have come to depend. While this book was being edited, I read J. Arvid Ågren’s The Gene’s-­ Eye View of Evolution (Oxford University Press, 2021), an assessment of this tradition that centers on Richard Dawkins. Ågren’s book does not explore the meaning of critique as a methodology or the literary shape of evolutionary genres, but his exposition is of great value in sorting out the ideas of Williams, Dawkins, and William Hamilton.

Contents

1 Introduction: What the Imagination Can Accept  1 2 Overture:  The Williams Critique, Trajectories of Lives and Groups 29 3 Proper Discourses of Design and Natural Selection 71 4 Opening  Sociobiology While Avoiding Aesthetic Distractions 99 5 Sex, Death, and the Language of Sociobiology133 6 How  Scientific Reductionism Leads to Evolutionary Explanation: A Defense165 7 Evolution  and Human Ethics: An Expansion from Sociobiology189 8 History, Contingency, Constraint: A Book of Nature227 9 Repairing Human Natures: Why We Get Sick265

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10 The Dark Side of Biology: Plan and Purpose in Nature293 11 A Retrospective: How Shall a Human Face the End?323 Index353

About the Author

Michael  P.  Cohen  explores the ways scientific conceptions inform and create languages and literatures of environments. He works at the intersection of literary theory and nature writing. In such books as Pathless Way: John Muir and American Wilderness (1984), A Garden of Bristlecones: Tales of Change in the Great Basin (1998), and Granite and Grace: Seeking the Heart of Yosemite (2019), his narratives often engage the personal with the professional, the subjective with the scientific. As a literary critic, he has published critical essays, notably “Blues in the Green: Ecocriticism Under Critique” (2004) and “Reading After Darwin” (2007).

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List of Figures

Fig. 1.1 Fig. 1.2 Fig. 1.3 Fig. 2.1

Fig. 2.2

Fig. 8.1

George C. Williams. Photograph courtesy of Stony Brook University; reproduced with permission 2 George C. Williams in 1962. Photograph courtesy of Doris Williams; reproduced with permission 8 George C. Williams in his office. Photograph courtesy of Stony Brook University; reproduced with permission 21 A diagram of the Weismann barrier, showing the one-way direction of inheritance, where the soma (body cells or organism) cannot influence the germ plasm (genes). Ian Alexander, August 31, 2017, Wikimedia Commons, https://commons.wikimedia.org/wiki/File:Weismann%27s_ Germ_Plasm.svg, Creative Commons license, CC BY-­SA 4.0 50 A diagram of Crick’s central dogma of molecular biology circa 1958. Wikimedia Commons, https://commons.wikimedia. org/wiki/File:Crick%27s_1958_central_dogma.svg. Public domain 51 These dendrograms illustrate “natural selection of codices,” B being an enlargement of A. Thickness indicates numerical abundance. C and D illustrate a pattern of imagined punctuated equilibria, and lack indication of abundance. This figure appears in George C. Williams, Natural Selection: Domains, Levels, and Challenges (New York: Oxford University Press, 1992), 14. Reproduced with permission from Oxford University Press 234

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Fig. 8.2

Dendrograms generated by random branching and character changes. As Williams explains, “A shows a completely random phylogeny while B shows random changes opposed by normalizing clade selection.” This figure appears in George C. Williams, Natural Selection: Domains, Levels, and Challenges (New York: Oxford University Press, 1992), 133. Reproduced with permission from Oxford University Press 235 Fig. 10.1 “Payoff matrix for the sex-ratio game.” Figure in George C. Williams, The Pony Fish’s Glow: And Other Clues to Plan and Purpose in Nature (New York: Basic Books, 1997) 303

CHAPTER 1

Introduction: What the Imagination Can Accept

George C.  Williams is the most important twentieth-century American evolutionary biologist unknown to the lay public. This book explores the literary shape of six of his short and seminal books. These texts, normally read by biologists as technical monographs, are stylistically distinctive in their own right. I call them nature writing for this reason. Though readers tend to imagine that nature writing is about nature, a closer look reveals that Nature is literally created in these texts—and nowhere is this more startling than when biologists write. Unlike most contemporary scientists, Williams chose primarily to write not articles, but book-length arguments. Aside from the explication of Williams’s prose works, my purpose is to define evolutionary literacy—that is, the ability to read natural processes and articulate their principles (see Fig. 1.1). The biological career of George C. Williams is unimpeachable. More importantly, he was a commanding writer. He changed minds. And to compare great things to small, reading Williams’s texts on evolutionary theory changed my mind. Literacy consists of this kind of encounter, for the writer with thoughts and a reader with text, as both ponder the question “What can be said and what should be said about nature?” Not, mind you, that I ever doubted the truth and validity of evolutionary laws, but reading Williams’s books reshaped the way I think about the processes of adaptation, natural selection, sexual reproduction, and the evolution of

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 M. P. Cohen, George C. Williams and Evolutionary Literacy, Literatures, Cultures, and the Environment, https://doi.org/10.1007/978-3-031-11650-6_1

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Fig. 1.1  George C. Williams. Photograph courtesy of Stony Brook University; reproduced with permission

biological behavior. His texts on evolutionary theory confront the grounding realities of life while recognizing that ideas about these change. A reader of Williams comes to understand how evolutionary processes seem to shape biological entities and their ecology. In the same way that atomic theory is at the bottom of chemistry, so too are biological systems relational and capable of change. There is, however, as the physicist Niels Bohr argued, no escape from abstract thought.1 A reader might ask where I come from that I needed advice from Williams. And the short answer is that as a wilderness advocate and author of books about environmental challenges, I study the literature of environmental matters. Speaking in the first person, I should not forgot to include that I graduated from UCLA with a major in physical chemistry and a minor in math in 1966. Williams had completed his doctorate there a dozen years earlier. Indeed, as I write this book I begin to wonder whether my reading of this fellow alumnus could be a coded memoir. But my thinking and activism emerged from an American romantic tradition

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that begins with Ralph Waldo Emerson and Henry David Thoreau and encourages a fixed spiritual relationship to nature. For much of my life, I was convinced by Emerson’s argument that humans ought to be in harmony with nature, and that nature itself is timelessly harmonious. Williams, as this book will show, thought otherwise. I reconsidered my own naïve point of view over a period of time—by critiquing my own ideas. Most obviously, this is what I attempted in “Blues in the Green: Ecocriticism under Critique” (2004).2 As a kind of terrible learning, critique terrorizes received ideas, scrapes away intellectual detritus, turns the self-evident into the questionable, the eternal into the historical. Yet many readers continue to confuse this methodology with the curmudgeonly character of the writer. This is unfortunate. When considering how evolutionary explanations are constructed, one sees the importance of literacy, which requires the written word. Given a focus on critique, I briefly place two contemporaries—Welsh writer Raymond Henry Williams (August 31, 1921–January 26, 1988) and American biologist George Christopher Williams (May 12, 1926– September 8, 2010)—alongside each other. The writings of each complement those of the other: the first is known for providing a rigorous critique of theories of culture and the second of theories of nature. As Raymond Williams writes in Keywords, “Nature is perhaps the most complex word in the language.” Of three areas of meaning he distinguishes, the second is most pertinent here: “the inherent force which directs either the world or human beings or both.”3 Critique is essential to clear thinking. It also constitutes a literary genre. The wonderfully styled cultural analysis of Raymond Williams, for instance, was crucial for my progress. One might say he invented the interdisciplinary field of cultural studies, which is concerned with the role of social institutions in shaping culture—or to put it more simply, with how a society creates and shares meaning. His arguments, especially in The Country and the City (1973) and Keywords (1976), are pivotal. Specifically, he explored and complicated commonly held ideas (and my own sense) of any ideal human relationship to nature. As colleague William Cronon has reminded environmental historians many times, Williams showed that “the idea of nature contains, though often unnoticed, an extraordinary amount of human history.”4 Yet I did not find that this view was sufficient for my purposes. I believe the sciences of life do confront certain bedrock realities, though these realities change as they undergo close inspection. In the study of literary theory, one learns that it is not possible for a critique

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of ideology to stand outside ideology; there is no being outside of ideology. We are always going somewhere when we think, though we imagine ourselves as the still point in a changing world. George C.  Williams was also a master, and when he dismantled the ideas of nature commonly held by biologists in the 1960s, he terrorized my own ideals. Like most of my friends who grew up in the early 1960s, I believed that an organic, communal ideal of ecology was foundational for understanding the processes of life and for building an environmental ethic. George C. Williams’s texts undermine this aesthetic ideology, at first by implication and then directly. Like most of my colleagues, I resisted the kinds of biological theory that interfered with my sentiments. As an eminent ecologist himself, Williams dismissed the harmonious aesthetic perspective we preferred as frankly distracting. He stressed, quite elegantly, that ecosystems are a consequence of evolutionary processes, that these are far less systematic than many wished to believe, and that biology is more about conflict than it is about cooperation. Or to put it another way, ideas of natural balance, homeostasis, or unity are probably untenable. For Williams, I learned, it was not possible to understand ecology without thinking about its evolutionary underpinnings, because good scientific writing does not aim to provide answers but to explore a process of rational thinking and test it through empirical experiments. Following the literary structures of Williams means following the literary structures of his thought. But of course, George Williams’s major works are classics not only because of what they say or imply, or what one can learn from them, but also by virtue of their unique style. The style of a writer is simply the way in which he approaches his medium—words. This is where my expertise enters. From my perspective as a literary critic, every writer comes out of a literary tradition and works within a set of conventions. Literary classics are not timeless, but they outlive their times. I did not go to Williams because I wanted to stay up to date with “current” evolutionary theory, or because I wished to apply his ideas to other literary works—as has become popular of late—but because I wanted to see how evolutionary theory was written and reconceived in the mid-twentieth century. Though some sociobiologists would like to study literature as if it revealed biological phenomena, I have preferred to study biological texts as literature.5 It was a time of scientific crisis, backgrounded by the blinding light of the atom bomb and foregrounded by the elucidation of the structure of DNA. I do not exaggerate when I say that George C. Williams reshaped

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evolutionary theory in the late twentieth century by rewriting it. I take his often understated rationalism, couched in a plain style, to be a model of discourse for times of crisis.

Reading George C. Williams I arrived at the texts of Williams in this way. I began by assessing the tradition he came from. At first, because I wanted to better understand evolutionary theory—to clear my mind, so to speak—I embarked on a reading program, beginning with Stephen J. Gould’s The Structure of Evolutionary Theory (2002). I admire Gould’s writing. But what struck me later in Gould’s final book was his respect for Williams. I also realized that I would have to read the primary texts, and not just Darwin. I worked my way through the classic works of the so-called modern evolutionary synthesis that unified Darwinian theory with modern population genetics, especially the four classic texts: Theodosius Dobzhansky’s Genetics and the Origin of Species (1937), Ernst Mayr’s Systematics and the Origin of Species (1942), George Gaylord Simpson’s Tempo and Mode in Evolution (1944), and G. Ledyard Stebbins’s Variation and Evolution in Plants (1950). Dobzhansky in particular hooked me, just as his writings had seduced Williams decades before. These classic texts of the modern synthesis provide convincing evidence that the writing of twentieth-century biologists constitutes a literary tradition. This tradition has engendered the most important nature writing of my era. I also began to wonder about literary conceptions of nature writing. Joyce Carol Oates famously announced, in “Against Nature” (1986), that nature “inspires a painfully limited set of responses in ‘nature writers’—REVERENCE, AWE, PIETY, MYSTICAL ONENESS.” These are not the kinds of responses one finds in Williams, who was convinced that population genetics, the processes of adaptation, and the modes of natural selection revealed a less comforting dynamic, as conflict, simple statistics, and blind trial and error created a frightening historical drama of life where each individual fended for the chance that its genetical heritage would be passed on to the future. If Oates is troubled by a set of texts that provide a limited conception of nature writing as a genre, my purpose is to enlarge both that conception and the set of texts understood as nature writing. I also wanted to understand, as well as I could, how an evolutionary theorist thinks. In this sense, I found the forceful and accessible writings of Williams compelling. One has to choose a teacher, and one has to judge

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one’s teacher. George C. Williams reinforced my sense that the technical literature of biology can be more accessible to a good generalist reader with a rudimentary knowledge of its conventions than is generally conceded. When I began to explore the body of Williams’s work, I could see immediately how his prose would allow me to test my claim. In that sense, this book is an experiment. To be fair to the reader, let me say that I have had a fortunate preparation for reading evolutionary theory through my undergraduate background in physical and geochemistry—though I have lost most of my mathematical skills. Though I have written as a literary critic and an environmental historian, in my day job I taught people how to read and write. My methodological premise is not specialized. Under the rubric of the environmental humanities, there have been many recent calls for humanists to learn from scientists and vice-versa—with the promise that public policy might be enlightened. On one hand, this book directs itself to humanists, arguing implicitly that the careful reading of primary scientific texts, within their scientific context, is both a worthwhile and essential task. On the other hand, it argues explicitly to scientists that scientific writing is subject to literary analysis, as nature writing. The rhetorical strategies scientists use when they write about nature are often most visible to humanists. This can open a potential dialogue. My approach demonstrates one form a constructive engagement might take. Environmental humanities calls for interdisciplinary exploration of cultural understandings, interpretations, and definitions of nature in order to clarify the cultural meanings associated with environmental issues. Close literary reading of scientific texts is fundamental to this enterprise and central to its aims because scientific literacy underlies both political and technical responses to environmental crises. A reading of Williams provides a case study: he makes sophisticated and entertaining arguments, using different genres for different audiences. He writes neither for an academic nor a popular audience, but for both, exploring the ethical dimensions of biological thought and engaging a range of humanistic issues. In short, environmental humanists can and should read primary—not just popular—works by scientists, and not only by the scientists who support their own ideology, but also by the scientists who represent consensus. Scientists need to understand that their style, which is to say the literary qualities of their writing, matters. More important, public policy

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should not be made on the basis of the congeniality or popularity of ideas, but on their substance and efficacy. Reading George C. Williams closely has been a richly rewarding literary experience, and what’s more, following the history of his ideas in published texts has given me a sense of the trajectory of his thought—and the thoughts of his generation. What I liked best was following the way he changed his mind as he reassessed what he had previously written. The progression of his books constitutes a wonderful scientific memoir, focused, as I will argue, on science itself, on the scientific life as a multidimensional journey. I consider his later, more philosophical ideas to be direct consequences of his scientific understanding. I do not suppose it is appropriate to use his personality, character, or biography to explain his thinking or writing. I do not presume to judge him as a curmudgeon or as a confrontational personality. So my premise is that Williams is a great scientific writer who deserves a wider and historical literary reading, informed by knowledge of his tradition. This is what I have tried to do. As I wrote to Doris Williams, George’s wife, who facilitated and encouraged this work, “I can certainly tell you that I learned a great deal and was forced into serious reassessment of my own ideas. As a result of reading his texts closely, I will never be the same. In the end, I argue broadly that he instituted an era of critique for biological thought, and the dignified, restrained, and precise means by which he did so calls for universal respect and admiration.” I read these books as literary texts, but I do not pretend to be intimate with the man. For one thing, Williams is an intimidating intellect. Indeed, most biologists cannot imagine speaking of him by his first name, and I will not do so again. Yet every report I have read describes the man as straightforward, warm-hearted, generous, and humane—a “good helper,” in Doris Williams’s phrase.6 I believe this to be true. His literary texts seem straightforward in style, but when explored seriously, that is revealed as a deception. They are sophisticated, multilayered, often multivalent, and sometimes subversive as a result of their startling juxtapositions and significant changes in perspective (see Fig. 1.2). When one enters Williams’s prose, it is as if one encounters the power of ice for the first time, in its crystalline angularity, in its mysterious properties, in its hardness, in its penetrating coldness, and in its relentless momentum. This cold and precise prose—which reflects the conscious intentions of an exceptional mind—flows glacially, unsparing, even fierce, in its rationality. Yet all rationality can turn upon itself. I believe that the

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Fig. 1.2  George C. Williams in 1962. Photograph courtesy of Doris Williams; reproduced with permission

strategy of understatement so characteristic of Williams expresses this coldness of intellect, and undercuts itself as well, by doubting itself. Williams was always willing to question his own thoughts. He changed his own mind. The results, in a series of short, precise texts, constitute a compelling narrative of the mind testing itself against rock-bottom realities of life. I think humanists would be well advised to follow his example. I have been guided in the spirit of this enterprise by the restrained, impersonal, and understated tone Williams chose for his own writing. He was never effusive about biology. No doubt there are plenty of readers for books about the wonderful, awe-inspiring secrets of biological evolution, books that explain what “scientists” have “discovered,” that aim to enchant their audience, that are designed to buttress the faiths in nature’s goodness and in human progress that are so essential to early

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twenty-first-­century complacency. This is not such a book, and I am quite sure that George C. Williams would not want it to be that book.

Adaptation Williams focused on the process of adaptation. Though he is sometimes introduced as an “adaptationist,” that term, as we will see, has become highly controversial. It is enough to say that, for him, a chief goal of evolutionary biology is to understand the evolution of adaptations and their relation to natural selection. He considered adaptation and natural selection to be the chief processes in the evolution of organisms, necessary for constructing evolutionary explanations, and central to the goal of research on evolution.7 A research agenda, however, entails more than creating hypotheses. By definition, critique seeks grounds and I follow Williams’s continuing search for the grounds and implications of evolutionary theory as his critique operates upon instituted practices, discourses, and especially traditional institutional ways of thinking. Critique is not simply fault-finding.8 His close inspection of evolutionary theory in Adaptation and Natural Selection: A Critique of Some Current Evolutionary Thought (1966) revolutionized thinking in a great many disciplines. Let me make one thing clear from the beginning: evolution is true.9 The devil, however, is in the details, in how one thinks about processes. “Reading George Williams,” according to Daniel C. Dennett, author of the widely praised book Darwin’s Dangerous Idea, “showed me for the first time how hard it is to be a good evolutionary thinker, and how easy it is to make simple mistakes.”10 Dennett’s book relies heavily on Williams, and his admiration is so complete because “this is a harder game to play than any of us realize, and George plays it better than anybody else in the world.” George C. Williams investigates the realm codified by the simple and complex word nature, announcing at the onset that it is “difficult to dispense with belief in a moral order in living nature,” though it must be done. Nature is conflict, he writes, and he considers the ecological environment as a “strategy employed by Nature against an organism which, in turn, replies with a strategy of its own.”11 Williams makes a coherent and compelling case for ideas I did not believe for decades—indeed, for most of my career. I was what he would have called a “group selectionist,” distracted by aesthetic considerations

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and imagining natural processes in terms of what can be called “organic” beneficence. Like others, I found the term “mechanism” as used by Williams and biologists of his era, like E.  O. Wilson, insulting. Why? Because it implies that organisms are machines, and machines function in predetermined ways. Well, don’t they?12 Mostly, I am convinced now, he is right. I mention this because many of my friends and colleagues believe that all good environmentalists should consult with and be guided by scientific thought, but usually only by texts that buttress their own sentiments. I now believe that we should not consult scientists because they are our environmental allies, but listen to the scientists who think clearly, even if they contradict our views. We environmentalists have used the sometimes misleading language of scientists with whom we agreed because their language suited our aims, and ignored the language preferred by scientists with whom we disagreed. I am guilty of that sin. Much of this book is devoted to the public language scientists use when they speak to each other, and especially to the diction and syntax of Williams. Consequently, I have attempted to quote him directly as much as possible, to avoid adding my own language, metaphors, and analogies to his, even when paraphrasing. Paraphrase has been necessary for compression, as has my own principle of selection from his texts. Who is this man? George Christopher Williams received his PhD in biology from the University of California at Los Angeles in 1955. His actual biological specialty was always marine vertebrate zoology. He studied the evolutionary biology of fish, enjoyed teaching courses in marine biology, and spent most of his career teaching at the State University of New York at Stony Brook, where, as was typical of his self-effacing manner, he “always introduced himself to incoming undergraduate and graduate students … as a fish biologist.”13 When he acquired broad notoriety for disassembling the concept of “group selection” in Adaptation and Natural Selection (1966), he had already published a number of influential essays in the 1950s. He was a steadfast reductionist in biology. His work on the evolution of sex, his views on evolutionary ethics, and his interest in evolutionary medicine emerge from his early methodology. He received plenty of awards late in his life. In 1989, the Ecological Society of America named him an Eminent Ecologist, recognizing “an outstanding body of ecological work or of sustained ecological contributions of extraordinary merit.” He was a member of the National Academy

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of Sciences, which in 1992 awarded him its Elliot Medal because of his contributions to current evolutionary thought, especially the importance of natural selection and adaptation and the understanding of sexual reproduction, social behavior, senescence, and disease. In 1999, he received the Crafoord Prize for Bioscience jointly with Ernst Mayr and John Maynard Smith. Scientific American called him “one of the great evolutionary biologists of this century.” He was also a Guggenheim Fellow, devoted years to editing the American Naturalist and the Quarterly Review of Biology, and served as the president of the Society for the Study of Evolution. He was an exemplary collaborator. As even the notorious literary agent John Brockman noted, “few laypeople have heard of Williams. Yet nearly all evolutionary biologists, even those who do not agree with him, admire him.”14

An Atomic Theory of Natural Selection There is no such thing as an original relation to the universe: all see through the eyes of teachers, even when they rebel against inherited vision. Ralph Waldo Emerson, in “Nature,” claims the opposite: “the foregoing generations beheld God and nature face to face” but “we, through their eyes. Why should not we also enjoy an original relation to the universe?”15 Because Emerson’s suggestion is impossible in practice, it is best to choose teachers well. Williams began to formulate his ideas in 1954 and 1955—only a few years after the major texts of the modern evolutionary synthesis had been published by Julian Huxley, R.  A. Fisher, Theodosius Dobzhansky, J. B. S. Haldane, Sewall Wright, Ernst Mayr, George Gaylord Simpson, and G. Ledyard Stebbins.16 Some believe that Williams’s 1966 monograph constitutes a major turning point in the ideological history of the modern synthesis.17 His four major books are concise, plain of style, straightforward, accessible, and engaging. They are Adaptation and Natural Selection: A Critique of Some Current Evolutionary Thought (1966), Sex and Evolution (1975), Natural Selection: Domains, Levels, and Challenges (1992), and Plan and Purpose in Nature (1996). In addition, he wrote influential essays about reductionism in evolutionary biology, evolution and ethics, and the evolution of senescence and gerontology; he also coauthored a book with Randolph Nesse, Why We Get Sick: The New Science of Darwinian Medicine (1995).18 Each of his books focuses on natural selection and

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biological adaptation—in particular the adaptation of behavior—and each re-evaluates inherited thinking and methods of evolutionary theory. The deliberate plainness of titles reveals his unassuming style, and because of that style his writings have been immensely influential. Late in Williams’s career, in 1995, he was interviewed or wrote a response for a project by John Brockman in which he recalled that his interest in evolution was sparked during the summer of 1947. He was twenty-one years old and spent six weeks with the paleontologist Sam Welles in the Painted Desert, where, “For the first time in my life, people—real biologists, real scholars—were willing to sit and listen to my opinions.”19 He went to Berkeley and took a course with Ledyard Stebbins, a central figure of the modern synthesis and author of the classic Variation and Evolution of Plants (1950). There he was introduced to Dobzhansky’s Genetics and the Origin of Species: “Stebbins was great, but Dobzhansky’s book was what got me interested in natural selection as a process.”20 Yet in many ways Williams is not like Dobzhansky—not in his beliefs and, more pertinently, not in his professional aspirations. As every teacher knows, questions shape inquiry. Williams frames precise questions. He inculcates literacy by exploring the implications of biological questions and how certain constraints limit answers to them—that is why foundational questions interested him. What is an individual, or a group? How do individuals age, and what has aging to do with reproduction? What is an adaptation, and what is not? Who is kin, and what might altruism be? How does one describe the life history of a species? Why do any species reproduce sexually? Out of these, more philosophical questions emerged. Is Mother Nature beneficent, indifferent, or grossly immoral? What does Darwinism have to offer medicine? How can evolutionary knowledge allow humans to improve the quality of their lives, particularly in their declining years? Williams’s sometimes tentative answers to these questions impinge directly on human ethics. Sex and death are never far from his queries. Indeed, sex and death are a more universal focus for evolutionary studies than might be imagined by casual readers. Kim Sterelny and Paul Griffiths underline this emphasis, entitling their introduction to the philosophy of biology Sex and Death (1999).21 Williams endorses a particular part of the Darwinian tradition that stresses what has come to be called the “gene’s-eye-view of selection.” He, along with William Hamilton, created the perspective made popular by Richard Dawkins as the “selfish gene.” Early controversies over Dawkins’s

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book—which are not the subject of this book—include Mary Midgley giving Williams credit in 1979, when she wrote that Dawkins “hastily adopts a general definition for ‘gene’ which he attributes (rather surprisingly and without reference) to George Williams.” A more elaborate genealogy of the selfish-gene theory by Stephen J. Gould foregrounds Williams’s influence on Dawkins more precisely.22 To draw an analogy for my colleagues in the environmental humanities, Williams is to Dawkins as Aldo Leopold is to David Brower. Adaptation and Natural Selection concludes with a scientific testament: “Perhaps today’s theory of natural selection, which is essentially that provided more than thirty years ago by Fisher, Haldane, and Wright, is somewhat like Dalton’s atomic theory. It may not, in any absolute or permanent sense, represent the truth, but I am convinced that it is the light and the way.”23 As the analogy with Dalton suggests, when Williams offers a kind of atomic theory of Darwinian natural selection, he is acknowledging a tradition whose central figures include Ronald Fisher, who in large part initiated it; William Hamilton, Robert Trivers, and Williams himself, who developed it; and Richard Dawkins, who advocated it. Those inside biology mark Williams’s approach as rescuing and reinvigorating modern evolutionary theory because, as his subtitle indicates, it is a critique of inherited evolutionary thought and a program for going forward. It is a truism of scientific literature that classic scientific articles are often more cited than read. People actually read Williams. The light of Williams is not like the light of Theodosius Dobzhansky, who writes in 1973—to biology teachers—that “Nothing in Biology Makes Sense Except in the Light of Evolution.” Dobzhansky, the great unifier, sees through the light of evolution a satisfying and inspiring science: “Without that light it becomes a pile of sundry facts, some of them interesting or curious but making no meaningful picture as a whole.”24 Williams questions the origin of the “light,” and asks what “way” might best shape mid-twentieth-century evolutionary thought, characterized by powerful deterministic mechanisms, ubiquitous competition, and principles of selfishness ruling an eternally mindless evolutionary process. The tone of this milieu comes not from Dobzhansky, but from George C. Williams. Writers create a public world. Readers may engage that world or choose not to. Yet nobody who uses evolutionary explanations for anything can ignore George C. Williams because his texts—and I shall speak of them as literary texts—create situations. The conclusion of a text is only what the

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reader encounters at the end. Williams challenges readers with premises; he situates and engages readers, and then forces them to admit that situation. Williams creates a reasoned world that expects a reasoned response. And reasoning, as he says many times, is onerous. His plain style values reasoning over feeling or intuition—though it is far from simple, since biology is complex. His plain style is personally opaque, just as physical facts are, but his reasoning is not obscure. One cannot often see inside his private mind. His sentences are declarative. He reports what is in order to ask what happened and sometimes to ask, by testing, what might or ought to be. Testing his written world against one’s own reality seems to begin as a scientific enterprise, yet it turns into an ethical and finally a political act. Testing his written world against his personal world would call for a biographical study. I defer to Williams and attempt neither here. I am interested in what he called the “publicly demonstrable.”

Evolutionary Literacy and Evolutionary Theory Literacy is a public discipline. One must know the letters and be able to read and write. One must acquire evolutionary literacy by great labor; it is “onerous” to learn the tradition. I define evolutionary literacy as just this faculty: skill in reading and writing biology from a perspective sometimes called Darwinian. Evolutionary literacy is a disciplined way of thinking within this tradition about the processes that constitute life and its changes in a realm sometimes known as nature—a way of thinking that impinges on virtually everything else worth considering. Critique is essential to literacy because it always uncovers and investigates the grounding of a method. First literacy must face its grounding; applications come later. Evolutionary literacy cannot be acquired by accumulating information and facts about natural history, or even by classifying them. Instead, it entails interpreting information as part of a process, assessing significance, and applying that signifying process to a range of information. This way of thinking and writing, as George C.  Williams argued, must be onerous. There are many contemporary and historical versions of evolutionary literacy, in part because of the many processes that are entailed, and there are multiple versions of what might be most pertinent in understanding the whole. Reading any version of evolutionary narrative depends on assessing the perspective of the writer, the choice or selection of

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information, the evaluation of authorities, and the applications to which the narrative is put. A simple mode for acquiring evolutionary literacy might come from studying a reputable and comprehensive textbook like Douglas J. Futuyma’s Evolutionary Biology (1998) or Mark Ridley’s Evolution (2004). I use these textbooks as references.25 One can become knowledgeable about the history, philosophy, principles, and central controversies that comprise what Steven Jay Gould claims in his magnum opus to be The Structure of Evolutionary Theory; one can examine the major classic works and articles; one can even study the biographies of major figures—but at bottom being literate means being able to read and write with a firm evolutionary grounding. Modern evolutionary theory investigates the grounds for its own methods and modes. Evolutionary theory also constructs a disciplined, systematic analysis of how evolutionary thinking can be used—or abused—by professionals and by the general public. Moreover, evolutionary literacy intends to ground a great many other disciplines, including environmental literacy. So I argue that George C.  Williams explores evolutionary literacy, establishes theory through critique, and writes to clarify. It is a truth generally acknowledged that many foundational works of evolutionary theory are obscurely written. To write like Williams is to write with compression and clarity, yet even those writers who recall his analogies and anecdotes often rewrite them poorly. That is to say, his writing is literary. As a literary critic I am neither competent to nor desirous of judging the precision or longevity of his ideas; I aim only to judge the precision and elegance of his expression of these ideas. Williams’s texts follow, sequentially, a classical rhetorical pattern. A classical argument is divided into five parts: the exordium or introduction prepares the reader’s attention; the narratio or statement of facts sets forth the relevant events; the partitio or division assesses agreements and disagreements and announces the author’s main points; the refutatio or refutation destroys the adversary’s arguments; the confirmatio or proof presents both argument and corroboration; and the peroratio or conclusion is more artistic in style and meant to move an auditor to action. Although tradition dictates that the elements of classical oration proceed according to the order above, it is unnecessary to imagine that they, as used by modern writers, must follow such a rigid structure. Of Williams’s major works, the early essays can be imagined as an introduction, Adaptation and Natural Selection constitutes the narration, partition occurs first in Sex and Evolution and later in Natural Selection, refutation

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in his essays in defense of reduction, confirmation in Natural Selection, and peroration in Plan and Purpose in Nature and the essays on evolutionary ethics. The coherence of the whole of his writings is revealed by this rhetorical coherence. His style changes gradually over the course of his career, while his ideas change substantially. Consequently, I undertake a longitudinal study of his thoughts and especially their expression in his writing, primarily through his books. By definition, a longitudinal study comprises repeated observations of the same variables over a period of time, typically decades. I do not attempt to correlate his ideas with those of other biologists unless that is necessary for context or pertinent to his motives in writing. For instance, Williams is principally responsible for much of a central late twentieth-­ century transformation from the study of animal behavior—called ethology by Niko Tinbergen—into what came to be called sociobiology, behavioral ecology, life history, evolutionary psychology, and other related fields. Theory is made by a process whose opportunities are constrained by time and place. Williams is scrupulous in indicating these. The preface to Adaptation and Natural Selection indicates that the book was begun in 1963, when Williams used the library at the University of California in Berkeley. Natural Selection was researched in 1981–82 at Stanford University and at Queens University in Kingston, Ontario, in 1988–89.26

Style as Development and Expansion A sonata announces its theme at its inception in an exposition; a scientific text announces its theme by abbreviation, in an abstract. Musical form can be imagined as a “series of strategies designed to find a successful mean between the opposite extremes of unrelieved repetition and unrelieved alteration,” and so too with the form of scientific writings.27 The pleasure of reading scientific prose, like that of listening to a symphony, consists not simply in recognizing the principal theme, but in following the theme’s development, which puts a reader in the midst of complexity. Williams once remarked: “If one has Bach, who needs anyone else?”28 In The Genetical Theory of Natural Selection (1930), R.  A. Fisher attempted to become the Bach of evolutionary theory, setting out the framework for themes that George C.  Williams would later explore. As Fisher wrote, “Natural Selection is not Evolution. Yet, ever since the two words have been in common use, the theory of Natural Selection has been

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employed as a convenient abbreviation for the theory of Evolution by means of Natural Selection, put forward by Darwin and Wallace.”29 Williams’s prose becomes a strategic mode for developing or, as he wrote, “expanding” this tradition of thought. This writerly strategy is by no means the principal mode of expression used by evolutionary biologists of his generation. He thinks with and through language, structuring his exposition with problems to be solved and questions to be answered; he engages in analogies and thought experiments—which a reader might call stories. When he tells these stories, he reveals the perspectives and methods of human observers: that is where his introspection comes into play. He reveals not the consensus in the literature of evolution, but its controversies and conflicts. Contrarian perspectives, in a kind of intellectual counterpoint, often structure his essays and books. Consequently, the burden or pleasure of elucidating his texts also calls for revealing the contexts of these writings, especially in the central texts of other biologists who shape the tradition in which he works. In his biological thinking and literary style, Williams follows the rules and methods of reductionism. He prefers to explain complex biological processes and phenomena in terms of the laws of physics and chemistry. As he writes in Adaptation and Natural Selection, “This book is based on the assumption that the laws of physical science plus natural selection can furnish a complete explanation for any biological phenomenon, and that these principles can explain adaptation in general and in the abstract and any particular example of an adaptation.”30 As we will see, he alters this view later in his life, recognizing that history and contingency prohibit any complete reductionism to principles.31 One notices how Williams places short narratives—or stories—strategically within his arguments. They have this in common: they more often than not assess the perspective of a scientific observer. He prefers to think deductively. He begins with definitions and proceeds with explorations of their implications. Big arguments are studded with small ones, little gems, and close observations juxtaposed with general premises. Often, it is more rewarding to read a portion of an argument and by congruence understand how the whole argument works. Many of his deductive trains of thought begin with a paradox. His arguments climb up a ladder of reasoning from the micro to the macro and shape his larger literary structures. In this sense, reduction employs minimalist literary strategies to build larger structures from the ground up— and to question them.

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When I treat Williams’s texts as literary works, I read for the most part according to his stated intentions, not against the grain, as I seek to understand their literary style and structure. Often these literary strategies suggest more than his stated intentions. In that sense, these are imaginative texts—a premise he would likely deny. Like many American writers, Williams hides his subtlety and his sense of humor by pretending innocence and practicing understatement. When he coins the term Sisyphean genotype, for instance, he briefly tells the story of Sisyphus and simply defines it according to his dictionary as “requiring continuous redoing.”32 As Mark Twain remarks in “How to Tell a Story,” “The humorous story is told gravely; the teller does his best to conceal the fact that he even dimly suspects that there is anything funny about it,” and indeed “the teller will divert attention from that nub by dropping it in a carefully casual and indifferent way.” Williams follows this strategy. Unlike many of his contemporaries who sought scientific celebrity, he is not much of a showoff. Quite the opposite. These are imaginative texts also because they reveal a strange and singular enterprise rooted in an all-consuming desire to know or imagine the origins of species, including ourselves. “Why are we here?” is a silly question; “How did we get here?” a better question; “What are we, that we are here?” more to the point, but most important is “Where indeed are we?” Only recently has devoting one’s life to studying these kinds of questions become a profession for which one gets paid.

Design and Literary Tradition At the root of Williams’s inquiry is the meaning of design. And it is not flippant to claim that Williams has designs on his reader. Using Williams’s favorite term, I ask, what is the function of a book on evolutionary theory, and what does its function reveal about its design? As Alexander Pope claimed in “An Essay on Criticism,” “True wit is nature to advantage dress’d; / What oft was thought, but ne’er so well express’d.” What does the writer expect to accomplish? Why, in the end, does Williams so frequently suggest that his own thinking is incomplete and imperfect? Throughout his career, Williams places his writing in traditions larger than the neo-Darwinian one he overtly suggests in naming Fisher, Haldane, and Wright. The origin of his interest in design may be revealed when he consults Williams Paley’s Natural Theology: Or, Evidences of the Existence and Attributes of the Deity, Collected from the Appearances of

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Nature (1803).33 Paley is best known for the watchmaker analogy by which he made a teleological argument for the existence of God: design implies a designer. The watch is too perfect not to have been designed, writes Paley, and so is the human eye; the upshot is a so-called argument from design. Paley’s argument appears in nearly every book by Williams, often cited through long extracts. In Adaptation and Natural Selection, Williams recites Paley’s description of the human eye to his reader as a classic example of “recognizing functional design from careful observation.”34 In Natural Selection, he returns to Paley’s description and confronts and deconstructs the theological argument from design, demonstrating that the human eye is far from perfect, and for historical reasons: its flaws must be a result of contingencies—of trial and error.35 In Why We Get Sick, he and Nesse not only explore in detail the design and function of the human eye, but also transform Paley’s image of the found watch into an image of the imperfect and mutable design of the human body.36 In Plan and Purpose, while continuing to admire Paley’s observational skills, he introduces his own analysis to reveal not only “functionally elegant features, but also as a kind of cumulative historical burden, the arbitrary and dysfunctional features of organisms.” As he concludes in 1997, “there is no evidence that God has any engineering expertise.”37 To recognize possible design, one must observe closely. However, there is no designer. Evolutionary processes create what appear to be designs by trial and error, and cannot go back to correct early problems. Designs, if that is what they are, are constrained by previous events. Within this context, Williams creates his own argument(s) from design while working with traditional means and materials. Design dominates his thinking and creates a coherent method and worldview. The rules for his procedure are as follows: (1) never assume any motive for design in the structure or behavior of natural organisms; (2) examine the organism itself and compare its designs for survival with those of other creatures; (3) seek the lowest possible level of explanation. Eyes and observations—lights and ways—become critical for this reasoning and recur in his writings. The eye is thus both subject and tool. Williams is acutely aware of this recursiveness. What later writers like Daniel Dennett and Steven Pinker would call “reverse engineering” is the very method Williams adopts in the 1960s. Williams uses Paley, not Darwin, as a stylistic model. As he writes in 1992, “Adaptation is demonstrated by observed conformity to a priori design specifications,” and he finds this method followed by Galen, Paley,

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and some modern writers.38 As a result of such demonstrations, Williams allows the reader to see how he follows and resists traditional perspectives. What the tradition of scientific writing means to Williams is revealed by his scrupulous crediting of ideas to other observers; a kind of extinction of personality or even a depersonalization can be seen in his writing. Williams rarely claims that his ideas, even ones later attributed to him, are original— quite the opposite—and he often praises those who later clarified ideas he had attempted to articulate. That is to say, he recognizes his own writings as part of a historical process. He gives them no special status and often denies them lasting value.

The Light and the Way In “Nothing in Biology Makes Sense Except in the Light of Evolution,” Dobzhansky concludes by extolling the faith of Pierre Teilhard de Chardin, a faith that Williams would never be able to countenance. As Dobzhansky writes of himself and Teilhard, Christianity was the cornerstone of his worldview. Moreover, in his worldview science and faith were not segregated in watertight compartments, as they are with so many people. They were harmoniously fitting parts of his worldview. Teilhard was a creationist, but one who understood that the Creation is realized in this world by means of evolution.39

Unlike Dobzhansky, Williams seems uninterested in showing any confluence between evolutionary doctrine and religious faith. As Williams notices in the 1980s, he follows T.  H. Huxley, who thought cosmic evolution might be historically useful, but was “incompetent to furnish any better reason why what we call good is preferable to what we call evil than we had before.”40 Williams wishes to demolish the authority of traditional views of nature and of traditional cultural dicta that make normative judgments from or with nature. Neither does he expound on the unity and diversity of life, nor create a synthesis. Nevertheless, belief is involved in his self-assessments. Later in life, Williams defined himself not as an atheist but as an antitheist. In this, he performs not so much as a unifier as a detective, ever on the lookout for contradictions, inconsistencies, omissions, and paradoxes in evolutionary thought. As a theorist, he is more interested in the perspective of the observer than in creating a “meaningful picture.”

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The final sentence of Adaptation and Natural Selection—“I am convinced that it is the light and the way”—uses the first person to create a tension between conviction and belief, “the light and the way.” This shape of writing and of thought emphasizes, as Dobzhansky’s does not, that theorizing natural selection is fraught with personal danger, and natural selection can be at best only a fragile human construct. Williams believes that this conception must continue to change because all representations are subject to revision and cannot in any sense be absolute or permanent. Theory is always contraindicated by exception. Adaptation and Natural Selection focuses on the means of natural selection, and Williams is not alone in his critical outlook or in his faith. Others convinced of “the light and the way” include especially his contemporaries and colleagues, principally David Lack, John Maynard Smith, William Hamilton, Robert Trivers, Richard Dawkins, and occasionally E. O. Wilson (recent controversies to the contrary).41 In his many students and followers, George C. Williams is very much alive through multiple traditions, and so is his light and way (see Fig. 1.3).

Fig. 1.3  George C. Williams in his office. Photograph courtesy of Stony Brook University; reproduced with permission

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In certain respects, Williams’s critique, as it first appears in 1966 and as he develops it, finds a strong analogue in Niko Tinbergen’s ethology, the study of animal behavior. Tinbergen’s manifesto on the “biological study of behavior,” published as “On the Aims and Methods of Ethology” (1963), also appraises the perspective of the observer.42 He means three things by “biological study”: first that ethology is inductive, beginning with “observable phenomena,” second that it uses “a particular method of study (the biological method),” and third that it is shaped by “the kinds of questions we ask.” Tinbergen believes the four major questions for biology are causation, survival value, evolution, and ontogeny; he wishes to apply more rigorous “biological thinking” to behaviors as “part and parcel with the adaptive equipment of animals.”43 According to his colleague John R. Krebs, Tinbergen was striking in four ways: the “breadth of his approach” to biological problems, “his insistence on great precision of thought,” “his total lack of hierarchy and pomp,” and “his athletic skills.”44 Tinbergen told Julian Huxley in 1959: Now projecting ourselves into other human beings is perhaps allowable. We all belong to the same species … [but] It is very hard to imagine what a starfish feels when it is angry, if it ever gets angry. Many persons argue that this is a difference of degree, but many zoologists (and I am one) think that this is a matter of principle with no compromise possible.45

Biologists can never assume motive when thinking about the behavior of organs or organisms, nor can they assume motive with regard to natural selection. So too with Williams, who follows Tinbergen’s “matter of principle” rigorously, resisting what he calls vitalism and its subcategory, mentalism. As Williams writes, “I have no inclination to deny the mental realm or belittle its philosophical importance. I am inclined merely to delete it from biological explanation, because it is an entirely private phenomenon, and biology must deal with the publicly demonstrable.”46

Critique and Conflict Williams’s central theme is: “Evolutionary adaptation is a special and onerous concept that should not be used unnecessarily, and an effect should not be called a function unless it is clearly produced by design and not by chance.”47 Onerous is a term central to Williams, who believes that it is

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burdensome, troublesome, and tiresomely difficult to establish specific qualities of organisms as “adaptations.” Always skeptical of easy answers or easy explanation, he is a wary thinker, and this begins with critical thinking. But also, facing onerous tasks constitutes the central challenge of being a human. What is onerous requires great or extreme bodily, mental, or spiritual strength. So too with its synonyms: arduous, backbreaking, burdensome, demanding, difficult, effortful, exacting, exigent, formidable, hard, heavy, laborious, oppressive, rigorous, rough, severe, taxing, tough, trying, weighty. He chooses the toughest challenges because they are inescapable if he is to follow his own calling. According to the preface he wrote for its thirtieth anniversary, Adaptation and Natural Selection emerged from an academic culture—from the conflict between established science and a younger generation’s reconsiderations. Williams’s subject included conflicts between youth and age in a natural population, as well as “pervasive inconsistency in the use of the theory of natural selection.” As Williams wrote thirty years later, “the triggering event may have been a lecture by A.  E. Emerson, a renowned ecologist” that “dealt with what Emerson termed beneficial death, an idea that included August Weismann’s theory that senescence was evolved to cull the old and impaired from populations so that fitter youthful individuals could take their places.” In 1954, on the approximate date of the lecture, Williams was 28. A.  E. Emerson was about 60  years old. And as he wrote later, “My reaction was that if Emerson’s presentation was acceptable biology, I would prefer another calling.”48 So Williams’s forte is rebellious yet conservative from the beginning, as critical thinking about one’s elders and one’s own calling. It is a truism among cognitive psychologists that skills for critical thinking are not the same as a disposition to deploy them. Critical thinking involves both ability and a commitment to guiding the behavior of others. Williams exercises skills and disposition.49 Yes, conflict has always marked discussions of evolutionary theory and has been a mode of refining it. As he enters a highly contentious era of evolutionary theory—one that continues—Williams establishes characteristic modes of framing arguments. John Maynard Smith and George C. Price wondered in 1973, “In a typical conflict between two male animals of the same species, the winner gains mates, dominance rights, desirable territory, or other advantages … Consequently, one might expect that natural selection would develop maximally effective weapons and fighting styles for a ‘total war.’”50 But this does not happen. Some “natural”

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process imposes limitations on conflict. How and why? the biologist wonders. So too with human conflict and human choices, but for very different reasons. As Williams writes, “The opposition arises, as Darwin himself observed, not from what reason dictates but from the limits of what the imagination can accept.”51

Notes 1. Niels Bohr, creator of a famous model for atomic structure, was once reported to have said, “There is no quantum world. There is only an abstract quantum physical description. It is wrong to think that the task of physics is to find out how nature is. Physics concerns what we can say about Nature.” As quoted in Aage Petersen, “The Philosophy of Niels Bohr,” Bulletin of the Atomic Scientists 19, no. 7 (September 1963) 8–14: 12. This is generally not regarded as a direct quote, but as Petersen’s paraphrase of Bohr’s viewpoint. 2. Michael P.  Cohen, “Blues in the Green: Ecocriticism under Critique,” Environmental History 9, no. 1 (2004): 9–36. 3. Raymond Williams, Keywords: A Vocabulary of Culture and Society (New York: Oxford University Press, 1985). 4. Raymond Williams, Culture and Materialism: Selected Essays (Verso, 2005), 67. 5. See Barash, David P., and Nanelle R. Barash. Madame Bovary’s Ovaries: A Darwinian Look at Literature (Dell Books, 2008). 6. Jim Bull, Eric L. Charnov, and Teresa Carson, “In Memoriam: George C. Williams (1926–2010),” The American Naturalist 177, no. 1 (2011): v–vii. 7. Steven Hecht Orzack and Patrick Forber, “Adaptationism,” The Stanford Encyclopedia of Philosophy (Winter 2012 Edition), edited by Edward N.  Zalta, http://plato.stanford.edu/archives/win2012/entries/ adaptationism/. 8. Judith Butler, “What is Critique? An Essay on Foucault’s Virtue,” in David Ingram, ed., The Political: Readings in Continental Philosophy (London: Basil Blackwell, 2002). Reprinted: http://eipcp.net/transversal/0806/butler/en. 9. Jerry A.  Coyne, Why Evolution is True (New York: Viking Penguin, 2009). There are many popular treatments of this theme. An easy read is Ernst Mayr’s What Evolution Is (New York: Basic Books, 2002). 10. Daniel C. Dennett, Darwin’s Dangerous Idea: Evolution and the Meaning of Life. (New York: Simon & Schuster, 1996). For ­Dennett’s appreciative comments, see http://www.edge.org/discourse/george_williams.html.

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11. George C.  Williams, Adaptation and Natural Selection: A Critique of Some Current Evolutionary Thought (Princeton, NJ: Princeton University Press, 1966), 4, 67. 12. See Ingo Brigandt and Alan Love, “Reductionism in Biology,” The Stanford Encyclopedia of Philosophy (Fall 2008 Edition), edited by Edward N.  Zalta, http://plato.stanford.edu/archives/fall2008/entries/ reduction-­biology. 13. Steven P. Ferraro, “Red-Letter Days,” the Quarterly Review of Biology 80, no. 1 (2005): 13–17, 13. 14. John Brockman, The Third Culture (New York: Simon & Schuster, 1995), 34. 15. Ralph Waldo Emerson, “Nature,” in Selections from Ralph Waldo Emerson, edited by Stephen E.  Whicher, 21–56 (Boston: Houghton Mifflin, 1957). 16. Four books published by Columbia University Press are normally cited as constituting the classic American literature of the modern synthesis: Theodosius Dobzhansky, Genetics and the Origin of Species (1937); Ernst Mayr, Systematics and the Origin of Species (1942); George Gaylord Simpson, Tempo and Mode in Evolution (1944); Ledyard Stebbins, Variation and Evolution in Plants (1950). Many might also include Julian Huxley, Evolution: The Modern Synthesis (New York: Harper & Brothers, 1942), since this volume names the movement. J.  B. S.  Haldane’s The Causes of Evolution (1932) and R.  A. Fisher’s The Genetical Theory of Natural Selection (1930) are also essential to the tradition. See Vassiliki Betty Smocovitis, Unifying Biology: The Evolutionary Synthesis and Evolutionary Biology (Princeton, NJ: Princeton University Press, 1996). 17. See “In Conversation with George C.  Williams,” in The Evolutionist: http://www2.lse.ac.uk/CPNSS/projects/darwin/publications/evolutionist/williams.aspx. 18. George C. Williams, Adaptation and Natural Selection (Princeton, NJ: Princeton University Press, 1966); Sex and Evolution (Princeton, NJ: Princeton University Press, 1975); Natural Selection: Domains, Levels, and Challenges (Oxford: Oxford University Press, 1992); Plan and Purpose in Nature (London: Weidenfeld & Nicolson, 1996); and George C. Williams and Randolph Nesse, Why We Get Sick: The New Science of Darwinian Medicine (New York: Random House, 1995). Plan and Purpose in Nature was published in the United States as The Pony Fish’s Glow: And Other Clues to Plan and Purpose in Nature (New York: Basic Books, 1997). I use this 1997 American edition. 19. George C. Williams, “A Package of Information,” Edge, edited by John Brockman (May 1, 1996), https://www.edge.org/conversation/ george_c_williams-­chapter-­1-­a-­package-­of-­information.

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20. Williams, “A Package of Information.” 21. Kim Sterelny and Paul E.  Griffiths, Sex and Death: An Introduction to Philosophy of Biology (Chicago: University of Chicago Press, 1999). 22. Mary Midgley, “Gene-juggling,” Philosophy 54 (October 1979), http:// www.royalinstitutephilosophy.org/articles/article.php?id=14. See also Stephen Jay Gould, “The Evolutionary Definition of Selective Agency, Validation of the Theory of Hierarchical Selection, and Fallacy of the Selfish Gene,” in Rama S. Singh, Costas B. Krimbas, Diane B. Paul, and John Beatty, eds., Thinking about Evolution: Historical, Philosophical, and Political Perspectives 2 (Cambridge, UK: Cambridge University Press, 2001): 208–34. 23. Williams, Adaptation, 273. 24. Theodosius Dobzhansky, “Nothing in Biology Makes Sense Except in the Light of Evolution,” American Biology Teacher 35, no. 3 (1973): 125–29. 25. Douglas J.  Futuyma, Evolutionary Biology, 3rd ed. (Sunderland Massachusetts, 1998); Mark Ridley, Evolution, 3rd ed. (Malden, MA: Blackwell, 2004). 26. Williams, Adaptation (1966), xv; Williams, Natural Selection (1992), “Preface,” pages unnumbered. 27. Percy A. Scholes, “Form,” in The Oxford Companion to Music, 10th ed. (Oxford: Oxford University Press, 1977). 28. Frank C.  Erk, “George C.  Williams: Personal Reminiscences,” the Quarterly Review of Biology 80.1 (2005): 7–11, 10. 29. R.  A. Fisher, A Genetical Theory of Natural Selection: A Complete Variorum Edition (1930; Oxford: Oxford University Press, 2003), vii. See also A. W. F. Edwards, “The Genetical Theory of Natural Selection,” Genetics 154 (April 2000): 1419–26. 30. Williams, Adaptation, 6–7. 31. See Ingo Brigandt and Alan Love, “Reductionism in Biology,” The Stanford Encyclopedia of Philosophy (Fall 2008 Edition), edited by Edward N.  Zalta, http://plato.stanford.edu/archives/fall2008/entries/ reduction-­biology/. 32. Williams, Sex and Evolution, 46. 33. William Paley, Natural Theology: Or, Evidences of the Existence and Attributes of the Deity, Collected from the Appearances of Nature (Bridgewater Treatises, Faulder, 1803). 34. Williams, Adaptation, 258–9. 35. Williams, Natural Selection, 72–73. 36. Williams, Why We Get Sick, 127–9, 234–5; 272 note. 37. Williams, The Pony Fish’s Glow, 6–12, 10, 153. 38. Williams, Natural Selection, 41.

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39. Dobzhansky 1973, 129. 40. T. H. Huxley, Evolution and Ethics (1893): 79–80. 41. See Samir Okasha, “Maynard Smith on the Levels of Selection Question,” Biology and Philosophy 20, no. 5 (2005): 989–1010. 42. Niko Tinbergen, “On Aims and Methods of Ethology,” Zeitschrift für Tierpsychologie 20, no. 4 (1963): 410–33. 43. Tinbergen, “On Aims and Methods of Ethology,” 411, 430. 44. Richard Dawkins and Timothy R.  Halliday, eds, The Tinbergen Legacy (London: Chapman & Hall, 1991): 60, 61. 45. Sol Tax and Charles Callender, eds, Evolution after Darwin, vol. 3 (Chicago: University of Chicago Press, 1960), 185. 46. Williams, Natural Selection, 4. 47. Williams, Adaptation, v. 48. George C.  Williams, “Preface (1996),” in Adaptation and Natural Selection. Reprint. Princeton Science Library Edition (Princeton, NJ: Princeton University Press, 1996), ix. 49. D. Alan Bensley, Critical Thinking in Psychology: A Unified Skills Approach (Pacific Grove, CA: Brooks/Cole, 1997), 6. 50. John Maynard Smith and George Robert Price, “The Logic of Animal Conflict,” Nature 246, no. 2 (1973): 15–18, 15. 51. Williams, Adaptation, 3–4.

CHAPTER 2

Overture: The Williams Critique, Trajectories of Lives and Groups

Stephen Stearns writes of George Williams that “the major themes of his life in science were all there in 1957, expressed when he was 31  years old.”1 This chapter explores those themes, which are encompassed by disciplines now known as life history and sociobiology. In particular, assessment of his intellectual development begins early with problems interesting to him, how he articulated his inquiries, and in what context. These early essays begin to explore life’s limits and limitations; they focus on the mechanisms that control timing and the location of life’s events, of reproduction and birth, place or niche, home and schools, family relations, senescence, and death. In following the trajectory of these universal themes of life as he works them out over the years, one discovers that neither his ideas nor their expression are unchanging because he is his own most scrupulous critic. For reasons I hope Williams would approve, I foreground his penchant for critiquing biological ideas, including his own. I would not like to say that his thinking evolves, but that it changes and adapts—especially to a changing environment for his thought. What follows is an assessment of his mode of biological critique, his salient themes, and a careful reading of his early essays for their import.

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 M. P. Cohen, George C. Williams and Evolutionary Literacy, Literatures, Cultures, and the Environment, https://doi.org/10.1007/978-3-031-11650-6_2

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Williams and the Era of Critique When Williams almost single-handedly initiates what I would call an era of critique in the biological sciences, his analysis encompasses, first, the judgment and reasoning of biologists generally; second, their assumptions about levels of selection, as adaptation occurs in individual organisms and groups; third, theories about adaptive advantage and maintenance of sexual reproduction and sexual behavior; fourth, the role of history and contingency in adaptive function; fifth, the normative value of evolution in human ethics; and sixth, medical practice and medical education. Williams’s most important contribution to evolutionary theory is in the realm of imagining and testing the stories biologists tell. By evaluating stories, by analyzing traditional narratives, he re-grounds evolutionary literacy. According to him, literacy requires more than accumulation of information, empirical confirmation, or a coherent narrative. Literacy entails acts of judgment, most of all the judgment of one’s own thoughts—thoughts that, to a great extent, are embodied in inherited narratives. Here might be Williams’s place in a literary history of biology and evolutionary theory: If mid-twentieth-century evolutionary thought made deterministic mechanism powerful, competition ubiquitous, selfishness the rule, and the stupidity of the evolutionary process eternal—and many grew up with this view—the language and tone for this milieu comes from George C. Williams, and later from Richard Dawkins and others who were influenced by him. Yet, from everything I have read, I believe that Williams attempts to constrain speculation, not overextend it. This is why he repeats his thesis, namely: “The ground rule—or perhaps doctrine would be a better term— is that adaptation is a special and onerous concept that should be used only where it is really necessary.”2 I read this doctrine as prescribing not only what adaptation is but also what biologists are entitled to say—or not say—about the changes they observe in organisms over evolutionary time. Williams is interested not only in what actually happens “out there” but in the kinds of questions that should be asked, the stories that should be told, and as a result he creates a style of writing that ought to impinge on— humanists would say police—how biologists represent what happens. Style, then, is not ornamentation for Williams, but substantial and integral to modes of thought. As a result of the power of his writing, he successfully alters the discourse of his discipline, opens up sociobiological research,

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and perhaps also fosters intelligent general public discussion of the biological origin of sexual and social roles. For Williams, critique is an unselfish activity whose purpose is to advance clear thinking in one’s discipline and especially to encourage it in young researchers. Williams’s thought led Sarah Blaffer Hrdy to look for evidence of selection at the individual level when she studied infanticide in Hanuman langurs, and Mary Jane West-Eberhard to look for individual-­ level adaptations when she studied dominance interactions in paper wasps. As Sarah Blaffer Hrdy remembers, “Williams’ early influence was transmitted via Robert Trivers. Trivers however was very explicit about his debt to Williams on this. I’ve often referred to Trivers as Hamilton’s bulldog, but he was also Williams’s, leading sociobiologists in Cambridge in the early seventies to read Adaptation and Natural Selection.” As she points out, when Williams altered discourse in his discipline “it was not always directly.”3 Often enough it was. Despite recent announcements that the Dawkins and Williams era is over—as in David Sloan Wilson and Edward O. Wilson’s “Rethinking the Theoretical Foundation of Sociobiology”—an acute critic must recognize that the discourse Williams taught is alive as long as there are certain “tainted words” or modes of expression that scientists, and others, cannot use.4 Williams made it uncomfortable if not impossible for a scientist to say that individual biological organisms might “jeopardize their own interests for group welfare,” or that “individuals adaptively died for the benefit of the population.”5 A generation inherited, internalized, and practiced a Williams kind of language—and avoided other kinds of language. The way one talks about natural selection shapes thinking about life. How do you learn to unthink habits that were prescribed over a lifetime? You historicize the thinking by careful literary reading, so that the ideas become visible as indissolubly embodied in the language that articulates them. One must not lose sight of the tradition Williams inherited. As early as 1930, Ronald Fischer observed that natural selection had become a metaphor for evolution as a whole, and “a convenient abbreviation for the Theory of Evolution by means of Natural Selection.”6 Fischer’s “convenient abbreviation” consequently became a manner of speech and a focus of inquiry.

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Opening a Door to Sociobiology and Evolutionary Psychology With the 1966 publication of Williams’s Adaptation and Natural Selection, which explored the link between the two processes, he closed some doors of expression while opening others. Because people actually read the book, it changed their thinking, and marks, as more than one historian believes, the end of one era of evolutionary theory and the beginning of another. Most importantly, it legitimized for biologists and others a new rigor in discussions of the evolution of behavior. Ways of thinking are born or die because people select them or ignore them, just as they open up certain books and leave others closed. In the Triumph of Sociobiology (2001), John Alcock, a follower of the adaptationist program, gives Williams primary credit for opening up the methods of sociobiology.7 As he puts it—triumphantly—this was a “revolution in evolutionary biology that led to the emergence of sociobiology as an active, vibrant discipline.” According to Alcock, Williams redirected ethology’s primary focus on proximate concerns to the asking of ultimate questions, moving from Niko Tinbergen’s proximate view of how an individual organism’s structures function to an evolutionary or ultimate view of why a species evolved the structures or adaptations it possesses. Alcock sees how this transition to “why” questions led to what came to be called “just-so stories” by biologists, sociobiologists, evolutionary psychologists, and popular writers. He finds that Stephen J. Gould’s phrase “just-so stories” has become “one of the most successful derogatory labels ever invented.”8 Nevertheless, these stories have been powerful and influential. One should remember that E. O. Wilson coined the term sociobiology nearly a decade after the publication of Adaptation and Natural Selection in his monumental tome Sociobiology: The New Synthesis, defining it “as the systematic study of the biological basis of all social behavior.”9 But Alcock is not alone in seeing Williams’s guiding hand. Ronald de Souse, a Canadian philosopher, writes, “In the narrow sense, sociobiology is a specific set of doctrines, based on gene selectionism, or at least rejection of group selection. In this sense its originator is not so much E. O. Wilson (1975) as George C. Williams (1966).”10 As he argues, Williams specifically grounded sociobiology in the technical concepts of inclusive fitness and parental investment, among others. What was group selection, that Williams took up arms against it? Group selection is a simple but virtually untestable premise that natural selection

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operates between groups of organisms, rather than between individuals, and results in adaptations that benefit the group rather than the individual.11 Williams saw every sign that group selection was impotent; he believed that W. D. Hamilton’s theory of inclusive fitness (or kin selection) was more likely to explain the problem of altruism through calculation of costs and benefits to individuals, an ideology that has come to be called “the selfish gene” or the “gene’s eye view” of evolution. Such inclusiveness was rare without close genetic relations, and adaptation was a blind process achieving no progress. We have been living with the ideology that has come to be called “the selfish gene” for a generation now, perhaps to our sorrow. Evolutionary psychologists have claimed Williams as a progenitor and to some extent Williams accepted—even welcomed—their claims. According to evolutionary psychology’s “Santa Barbara School,” the strain of theoretical evolutionary biology that started in the late 1950s and early 1960s, with the work of George Williams, William D. Hamilton, and John Maynard Smith, became the first building block of evolutionary psychology (1982).12 Evolutionary psychologists like to recall that Williams once wrote, “Is it not reasonable to anticipate that our understanding of the human mind would be aided greatly by knowing the purpose for which it was designed?” But this statement is preceded by another that perhaps cuts against evolutionary psychology’s basic assumptions: “Despite the arguments that have been advanced (e.g., Dobzhansky and Montague, 1947; Singer, 1962), I cannot readily accept the idea that advanced mental capabilities have ever been directly favored by selection.”13 Discussions of the genealogy of sociobiology and evolutionary psychology as they have emerged incrementally from Tinbergen’s ethology continue, with some commentators attributing additions to Williams, Hamilton, E. O. Wilson, and Trivers. Dawkins calls himself an ethologist and was mentored by Tinbergen. Daniel Dennett probably oversimplifies when he describes evolutionary psychology as merely a marriage of sociobiology and cognitive psychology; in all probability its literature has been inspired as much by the theoretical work of Dawkins and Williams as by E. O. Wilson and his followers.14 It is one thing to claim, as I do, that Williams opened the way to sociobiology by making Hamilton’s work accessible in a language biologists and social scientists could understand. It is quite another thing to trace the way Williams himself came to think “like” a sociobiologist. As I will argue, a set of thoughts, perhaps occasioned by Hamilton’s work, led Williams to

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reassess his views on sexual reproduction as it relates to social behavior. After all, sociobiology is all about sexual reproduction. As always, he would go to ground, starting his book Sex and Evolution by re-evaluating the concept of meiosis. Yet Williams’s sociobiology is not E. O. Wilson’s. This puts him in a strange rhetorical situation in the late 1980s, defending or ramifying the implications for ideas his work seems to substantiate, but as publicized and extended by others, sometimes beyond where he would reach. While Wilson argues that “the time has come for ethics to be removed temporarily from the hands of the philosophers and biologicized,” Williams argues quite the reverse—that human ethics must not be bioligicized.15 In other words, as Williams spells out his commitment to a convergence—but not a consilience—of sociobiology, evolutionary theory, and ethics, he cannot avoid implicating himself in a worldview. With current knowledge about sociobiology, he asks, what behaviors ought humans to practice? One must see Williams’s essays, including “Evolutionary Ethics,” through his own lens, his assessment of evolutionary processes, which has rarely been sufficiently done. One must also see him in the context of his readings. Most of all, one needs to see where he positions himself among a range of philosophers, historians, and other evolutionary biologists— especially Thomas Henry Huxley. Williams’s Huxley-inspired essays of the 1980s constitute a first step toward applying an understanding of evolutionary processes to make discussions of public policy rational. He continues to do this by engaging with medical practice in Why We Get Sick (1995), a book coauthored with Randolph Nesse. Williams makes this leap—or maybe it is not a leap—when he writes increasingly explicit discussions of human behavior and human illusion. It is not surprising that he chooses to continue the satirical mode of the Huxley essays on evolutionary ethics in Plan and Purpose in Nature (1997), which he published as an emeritus professor, after he had retired from formal teaching. How and why Williams decided to take up these public positions, and more importantly, to modulate from an ironic to satirical tone for these discussions, is something of a mystery. But he did, and as he wrote increasingly on the biological processes of human life, the tone of his writing approached dark comedy. As he repeatedly announced, the evolutionary process prunes populations through conflict between competing juveniles, including siblings, and precipitates heavy losses, inefficiencies, and cruelties. Natural selection operates through stupid and blunt methods; it does not seek to perfect optimum design, or provide peace for any organism,

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but permits only what is sufficient for genetic survival. Its designs are often jury-rigged, being constrained by the historical contingencies of an unreachable past. As a process that cannot look forward, natural selection shapes the experiences of all organisms. This situation might be either sad or ridiculous. Nevertheless, Williams was first of all a biologist. Before asking about his view of culture, one should explore carefully the processes that he found in nature and know what he thinks understanding these processes can do for our culture. What is distinctive about his evolutionary theory is how he imagines pattern in these processes. Williams may not believe that there is plan or purpose in nature, but there is plan and purpose in his writing. As he once said, “In biology, when you’re talking about things like genes and genotypes and gene pools, you’re talking about information, not physical objective reality. They’re patterns.”16 An ecocritic might notice that the very pattern of this talking—or more precisely, writing— comes to embody a structure of biological thought that reveals more than itself. There may be no plan or purpose in nature, but there is literary structure in human thought about patterns.

Doctrine as Literate Evolutionary Theory Scholars in the humanities are in need of a more nuanced conception of what evolution means, particularly for a working evolutionary biologist. Call this conception evolutionary literacy, best gained by reading the texts evolutionary biologists write for their academic and professional peers. Those who are literate in evolutionary theory must be able to think with— if not like—an evolutionary biologist. Williams set out three doctrinal bases for successful biological research: 1. Mechanism (as opposed to vitalism) 2. Natural selection (trial and error as opposed to rational plan) and 3. Historicity As Stephen Stearns puts it, “All three explain the features of organisms, not natural selection alone.”17 As a writer, Williams aims to create an evolutionary literacy grounded in these doctrinal bases, including modern theories of natural selection and adaptation—not controversies about them, but what he understands to be the consensus of the scientific literature. This mode of thinking has been influential among biologists,

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sociobiologists, and evolutionary psychologists, but the care of Williams ought to rein in the looser discourses of sociobiology as they enter the public domain. Williams consistently cautions that evolutionary theory is imperfect, and will always be so, because: 1. Scientific thinking, especially adaptationist thinking, must create inexact stories. 2. As a result it constructs imperfect, though increasingly accu rate, theories. 3. Consequently evolutionary theory can never come to a single comprehensive conclusion. In other words, evolutionary theory converges on the truth but also branches. Evolutionary literacy similarly converges and branches. Or, to put it in a Williams way, evolutionary theory is always evolving, with no perfect end in sight, no master plan, and no guiding hand. Implications continue to sprout from what Williams taught, and drawing diagrams of these can go on forever. Yet any branch points back to firm doctrinal roots: whatever ideas humans may have of themselves or their fellow creatures, all will have to accommodate themselves to basic evolutionary causes, processes, and results and to confront “why?” questions. Evolutionary literacy always implies these questions, and its essential recursiveness is inescapable. Although it may be true that illiterate humans are not influenced by theories of natural selection, for Williams, certain aspects of evolutionary theory inexorably shape a literate human’s more-than-scientific sense of the world: 1. Humans may desire to attribute higher functions to natural processes, or desire to believe in a plan or purpose. To do so is to enter a world of illusion; as Darwin wrote, it is “to enter into the realms of miracle, and to leave those of science.”18 As the New Critics from whom I learned argued about texts, so Williams argues about natural selection: “Never assume a motive.” Seek the lowest level of explanation first. 2. Humans cannot and should not aspire to live according to the “laws of nature,” and not only because the laws of nature are cruel and ruthless. Humans have no choice. They must invent their own laws, as against natural laws.

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3. The immediate present shapes all basic conditions for life. Life can look neither backward nor forward and is constantly constrained by an unredeemable because historically constrained past, and an unpredictable future. Evolutionary processes can have no conception of the past or the future. But humans have conceptions; they can look backward and forward, even if imperfectly. These conceptions can be perfected. 4. Like all forms of life, humans have limitations and imperfections of design. Literacy requires facing up to the processes by which humans and other creatures have been made, recognizing that the ways in which organic beings have been designed contain both possibilities and limitations, and that these are the result of a long history that continues to shape thought and behavior. Evolutionary literacy becomes itself onerous because it entails using and disseminating public scientific knowledge rigorously, while investigating its methods and appropriate discourse. This responsibility extends beyond the professional discourse of scientists to all those who use science in public discourse. 1. To be literate requires using evolutionary theory without making slips, falling into errors, or choosing language that is indecorous or obfuscates. 2. Literacy includes the ability to speak and write clearly when applying and explaining this knowledge, especially in the public realm. 3. Literacy requires the use of parsimony for clarity of thought and for style of expression. The most straightforward and plainest explanation should be matched with the most straightforward and plainest expression. Literacy is itself a method, a process. It does not suffice to acquire a body of knowledge or a theory, though those are both necessary for literacy. More information or data may help, but data does not create literacy, whose aim is to use information. Williams was a master of this ability. He presents a coherent worldview that a reader can approach by understanding his search for grounds of explanation. Such books as Ernst Mayr’s What Evolution Is (2001) or Jerry Coyne’s Why Evolution is True (2009) speak for a sense among evolutionary biologists that they are besieged by ignorance.19 This is true. Many biologists

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write with the hope that if only evolution is depicted as a coherent, complete, and thoroughly tested theory, it will be believed. Consequently it is also normally assumed that the scientist’s public role is to explain how evolution works, leading to a kind of contest for the position of best explainer of evolution. As Friedrich Schiller is said to have written, “Against stupidity, the gods themselves contend in vain.” But resistance may come from other sources than questions of accuracy, validity, or truth. Resistance to theory may come from those who do not want to think. Thinking and writing about evolution is not simply an entertainment. What makes Williams rewarding is that his plain style and his aesthetic distance from his subject entail questioning his own views. He presents the doctrine, but he also interrogates it. He is always grounding, grounding, grounding. Critique uncovers grounding, and literacy begins with grounding. Later come the applications. Williams says, don’t pick the fruit without examining the roots of the tree of thought.

Naturalist as Theorist, Biologist as Popular Nature Writer Sex and Evolution (1975) demonstrates through its use of examples— plants and animals, terrestrial and aquatic—the extensive breadth of Williams’s knowledge. In this book, he is most saliently a naturalist. To be a naturalist is to use the comparative method, to appreciate diversity. No doubt the comparatist must admit to an aesthetic motive. A biological comparatist, like a literary one, not only appreciates diversity, but also brings abstract order to the chaos of forms—as Williams attempts to do with the chaos of reproductive strategies and behaviors. Evolutionary and ecological theorists are first of all biologists and often begin their careers as naturalists. Hamilton and Maynard Smith have been sure to remind interviewers that they were first of all naturalists. It may be necessary but not sufficient for a biologist to be a naturalist, as Lee Alan Dugatkin argues in Altruist Equation: Seven Scientists Search for the Origins of Goodness (2006). Nevertheless, for those who are most rigorous and perceptive, a “professional” interest in natural history has, in our time, been superseded by an interest in evolutionary and ecological theory. Evolutionary theorists work within a unique literary tradition, yet chronological histories of science writing often ignore how texts continually return—along with their ideas, or paradigms, to foundational issues,

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as in Robert May’s somewhat tongue-in-cheek description of “Ecology’s 12  year cycle.”20 Nature writers may nuance the world but they do so recursively, creating in their writings a nature according to certain paradigms, and these paradigms are inevitably aesthetic. What evolutionary theorists think and feel about what they know also matters, but there is no shortcut to understanding this—it is not apparent from reading about them or reading interviews with them. A critic cannot discuss the style of an evolutionary text without learning to follow its scientific content and the various views of the larger scientific community, the paradigms under which scientists work. As Hamilton pointed out, citing an article or book is not the same as understanding it.21 There must be an encounter, or else why not write only bibliographies? Certainly Williams attempts to write for a general public—to become a popular writer—with Plan and Purpose in Nature, but he resists the task too. A list of his previous publications, particularly their titles, does not offer much promise that Williams originally intended to be a popular writer. But by what criteria? Robert J. Richards has delineated aspects of “the popular” in his biography of Ernst Haeckel, The Tragic Sense of Life, and one can test Williams’s texts against Richards’s criteria.22 The first is that “the author intends the book for a general audience,” which Williams no doubt did not attempt until the mid-1990s. Yet by many other criteria, Williams always attempted to write popular books. For instance, many popular writers often direct their work initially at or to a university audience. Williams’s books have disseminated not just information but a way of thinking to a wide set of disciplines. The popular reaches, then, not toward an average Joe on the street, but toward scholars from different disciplines within the university. Avoiding jargon, another of Richards’s criteria, has always been a goal for Williams’s texts, though they do not always reach it. Nor can a popular work “burden its readers with a larger critical apparatus.” Some popular writers, like Haeckel, used media other than prose. Williams rarely did. According to Richards, “popular works of science will often strive for intimacy with the reader” by using the first person. Williams does this constantly. Richards decides that Haeckel is a prototype for authors like Stephen J. Gould, Richard Dawkins, E.  O. Wilson, and Steven Pinker, where “little excavation is required to reveal the hard philosophical-­ political-­ social core upon which rests the surface discussions.”23 For Williams’s early work, it is hard to find a “hard philosophical-political-­ social core.” In his later work, excavation reveals considerable complexity.

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Using Richards’s criteria for a short survey of Adaptation and Natural Selection reveals that George C. Williams intended to be a popular writer, but not exclusively so. Certainly the book meets many of Richards’s criteria and flirts with others. Williams wrote monographs, but not exclusively, as William Hamilton did. A monograph is, by one definition, “a detailed written study of a single specialized subject or an aspect of it,” and as Richards argues, the narrow scope of subject matter may be a hindrance to popularity—unless that subject becomes a source of public interest. That explosion of public interest happened to Williams, to his surprise. The popular is about thought and expression, changing minds and modes of discourse, not about numbers of books sold or numbers of citations. By all these criteria, in retrospect, Williams’s first book is his most popular. The best measure of popularity must be a work’s influence on disciplines inside and especially outside the author’s specialty. Williams’s influence on evolutionary psychology is especially revealing, in that more than one reader was hooked by the single question quoted above, about the purpose for which the human mind was designed.24 Ideas are not disembodied. Humans write the world, and inevitably they write into the world their own sense of their place and of human nature. Whatever nature is, it is created, fabricated one might say, by texts and enabled by bodies and minds. Nature writing, as traditionally or conventionally defined, is popular because it pleases the naturalist impulses in the reader. Evolutionary biology and related fields have written and fabled a world that we call nature. Its classic texts shape contemporary perceptions; for this reason, I call Williams a nature writer. In particular, scientific texts have cultural authority on one hand and their own traditions on the other. For instance, it is certainly too simple to say that John Maynard Smith or William Hamilton invented “kin selection” or “inclusive fitness,” or that Richard Dawkins invented the “selfish gene.” This language has been embodied not simply by empirical authority, theoretical rigor, knowledge, an aesthetic, or arguments, but also by modes of thinking and attitudes. Most obviously, writers turn various kinds of thinking into language. For Randolph Nesse, the problem with Williams’s early writing was its failure to reach for popularity. As Nesse argues, [Doris and George Williams’s] joint paper in 1957 on the evolution of altruism came close to describing inclusive fitness, but they veered away and relied instead on a variation of group selection. In typical fashion, they explicitly avoided the engaging word ‘altruism’ and instead advocated using

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the more neutral terms ‘social donors’ and ‘social non-donors.’ As memes, these were non-starters; selfish genes displaced them completely.25

But as in any tradition, scientific themes return and double back on themselves, even while terminology metamorphoses. Science imagines itself as progressive and cumulative, but texts, as T. S. Eliot once claimed, are co-­ extensive and work intertextually, containing and reinterpreting each other. Hamilton thought of his scientific papers as haiku, on the model of Basho!26

Narrative Confabulations: Life Histories as Bookkeeping The use of first person is considered acceptable but frequently avoided in scientific writing. This stylistic choice could be seen as a form of reduction; using first person forefronts an individual writer’s mental state, including both reason and emotion. But using first person also permits the narrator to position himself, and this is why one notices its appearance in the writings of Williams. Like scientists, humanists make up stories about fantasies of escape from the first person and perhaps from anthropomorphism. The strategies are different, but the motives may be similar. I illuminate this contrast by introducing Italo Calvino, who finished five out of his Six Memos for the Next Millennium (1988) before he died in 1985. The fifth memo, published as part of the posthumous collection, is entitled “Multiplicity.” Calvino writes, that “although science interests me just because of its efforts to escape from anthropomorphic knowledge, I am nonetheless convinced that our imagination cannot be anything but anthropomorphic.”27 In this regard, Calvino was particularly interested in the limitations of the first person, as pronoun and perspective: Think what it would be to have a work conceived from outside the self, a work that would let us escape the limited perspective of the individual ego, not only to enter into selves like our own but to give to that which has no language, to a bird perching on the edge of the gutter, to the tree in the spring and the tree in fall, to stone, to cement, to plastic.28

From the beginning of his writing career, one might argue, Williams attempted to “conceive from outside the self.” He was interested in life histories for that reason, first in terms of homing fishes, then kin selection,

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then senescence, then sexual histories, and finally in terms of evolutionary medicine. His goal was not to enter into selves, but to perceive similar and dissimilar shapes of the lives of others and ourselves, as if life history could be abstracted and theorized. As Williams writes of this aspiration, he uncovers a story of his own relationship with the narratives he inherits, some accepted and some not accepted, and of the difficulty of trying to re-tell them or get them right. Use of the first person, ever-present throughout his writing, reveals the extent to which Williams was aware that he was writing his own intellectual biography. Consider the advent of life history theory as it impinges on animal and human biology, psychology, and evolutionary anthropology. One might begin with the kind of thinking by duality that was traditionally framed in terms of r-selection versus K-selection theory—r representing reproduction, and K indicating competitive population or carrying capacity of the environment.29 In this frame, an r-selection strategy entails production of a large number of offspring with few survivors as early in life as possible, while a K-selection strategy entails producing smaller numbers of offspring with higher survival rates and more parental investment. Or consider the overarching analogy created by R. A. Fisher, who used actuarial tables to study mortality—so-called life tables that allow the actuary a clear notion of what is meant by “death rate at any age,” “expectation of life,” and so on—in order to show the “important biological aspects of vital statistics.”30 Imagine that many physiological traits and behaviors of individuals can be understood in terms of the timing of maturation, reproduction, and senescence. The course of any life can be measured in terms of age at weaning, of sexual maturity or puberty, adult body size, reproductive schedule and behavior. Any set of lives can be tabulated with mortality schedules, times of maximum fecundity, first sexual activity or mating, first reproduction, duration of gestation, litter size, intervals between births. So a reduction of lives to a cost-benefit analysis is possible: the costs of reproduction versus energy being diverted away from the routine repair and maintenance of the individual who reproduces. Life history becomes a matter of trade-offs, and these trade-offs and strategies can be compared across species. Consider further the tradeoffs between parent and child as worked out by Robert Trivers.31 Individual humans normally think about these traits and behaviors by creating and inspecting narratives. Humans often imagine these narratives as predictive. Life history theory suggests that, rather than our telling our

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stories, evolution tells its stories through us. A reader is faced with a question of authorized stories. Who has them, who holds them? How do people live within the context of life history stories? Which stories get dismissed as confabulations? (I can use my arm, says an injured mental patient, I just prefer not to!) I once had a colleague about whom not a few graduate students were heard to mumble, “Everything is about him.” And I wondered, in light of evolutionary theory, how it could be otherwise. Everything is about us, as individuals. There is always an analogy in human thinking, the anthropomorphic analogy, and facing it is onerous indeed for all of us, including the sociobiologists. Stories, like genes, are always present: they inhabit us, but they do not entirely rule us.

Design as Shape of Thought: The Typewriter In the largest sense, Williams staked out his interest in the permanent and the temporary in the history of life, in its flow: his fascination was almost like that of Heraclitus. In 1998, he expressed his so-called lament: “Natural selection maximizes short-sighted selfishness, no matter how much pain or loss it produces. There are far more losers than winners, and great losses often arise from trivial gains.”32 The coordinates of his world are time and space, home and school, reproduction and death. As he writes about these, he creates a universe. Humans are part of that universe, but the individual humans whom we know as phenotypes are ephemeral—like Socrates, for instance: Socrates consisted of the genes his parents gave him, the experiences they and his environment later provided, and a growth and development mediated by numerous meals. For all I know, he may have been very successful in the evolutionary sense of leaving numerous offspring. His phenotype, nevertheless, was utterly destroyed by the hemlock and has never since been duplicated. If the hemlock had not killed him, something else soon would have.33

This version of Socrates would surprise humanists but is a consequence of a general principle. We recognize individuals, but as phenotypes they are extremely temporary manifestations, each being a result of an interaction between genotype and environment.

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While Williams was eternally interested in the perspectives of human observers, he also scrutinized the manifest perceptions or senses of biological entities. What any organism thinks or does can best and perhaps only be determined by what it perceives and what it might imagine others see. Perspective has vectors through time and space; perspective binds or releases relations between bodies and within bodies. Williams wants to know not only what a fish can perceive and what a gene sees, but also what natural selection “sees,” what it can operate upon, and what it cannot touch. He wants to know how directional processes of biology work, how any organism’s capabilities are influenced by the past. Sometimes he wishes to predict, by asking what can be supposed about the future from the present. These explorations are cumulative because they indicate the shape and the connections of his thoughts. He begins with fish. He is a fish biologist. Frank C. Erk, who was a close colleague in Chicago, remembers that Doris Williams facilitated the flow of George’s ideas during the writing of his thesis for UCLA. “George found that his ideas were coming too fast to have to worry about changing the paper in his typewriter. So, at Doris’s suggestion, he bought a roll of shelf paper and put one end in the typewriter.”34 Erk thinks this might have been the first thesis in modern times to be written on a scroll, but the shelf-paper strategy was not a new one. Kerouac used it in 1951 while typing On the Road, because, according to him, speed facilitated the uncovering of truth. As he wrote to Neal Cassady, when he rolled his scroll out on the floor, it looked like a road.35 The purpose of this book is to follow the Williams road, and for that it is necessary to begin at the beginning.

Homes In 1952 and 1953, as a 26-year-old zoology student at UCLA, Williams spent much of his time observing the intertidal migrations of two species, the wooly sculpin and the opaleye, on the rocky shoreline between Redondo Beach and San Pedro, primarily on the Palos Verdes peninsula. Their homing behavior in tide pools became the subject of his doctoral dissertation. He believed that nothing but natural elements, principles, and relations were required to understand the fish’s world, sometimes by mathematical modeling. In several senses of the term, George C. Williams began as a naturalist. But like nature, naturalism, according to Raymond Williams, is a complex word, a tangle of slippery scientific, religious, philosophical, and literary

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meanings.36 Being a naturalist, shortened from “natural philosopher,” was originally synonymous with being a scientist, but more recently, the term speaks of those non-specialists who engage a broad range of interests in the study of natural history. Consequently, it now includes those who are amateur observers of nature and those who teach others to appreciate the wonders of their natural environment. A naturalist is often engaged in a quest to find order in nature. In a second, related sense of the term, nineteenth-century literary naturalists strongly drawn to Darwinian conceptions of natural selection often attempted to accurately depict life as determined by biological and social forces, emphasized indecorous themes such as sexuality, violence, and disease in their writings, and sometimes focused on human depravity and misery. Third, philosophical naturalists oppose their perspective to that of supernaturalism by attempting to understand the world in scientific terms, without recourse to spiritual or supernatural explanations. Indeed, naturalism can be described vigorously, as Thomas Nagel puts it, by a modern theistic philosopher like Alvin Plantinga, as “the view that the world describable by the natural sciences is all that exists, and that there is no such person as God, or anything like God.”37 According to this third sense of the term, most evolutionary biologists are naturalists. How else could the biological world be described by a scientist if not in naturalistic terms? But the natural sciences describe the world in diverse ways, and there are conflicting versions of evolutionary description, storytelling, and theory. After all, evolutionary scientists do not describe a static or homogeneous world that exists “out there” but rather sets of processes in different places and times, where humans find themselves immersed not at the origins, but in media res, as biological and intellectual observers, participants, and sometimes victims. According to Williams, restraint is called for; it is not the role of every evolutionary biologist to ask ultimate questions, but only to investigate processes available to humans, in our world, where it is possible to see and think. There is nothing grand about this investigation. It is often assumed that every evolutionary theorist begins by imagining himself as a “born naturalist,” as Darwin and later William Hamilton did. But this is not true for all. One might describe a scientist’s choice of an occupation as “niche picking.”38 A few years before Williams pursued his fish experiments, in another niche Watson and Crick were building a model, racing toward the publication of their discovery of the structure of DNA on April 25, 1953.

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In his study of the “Homing Behavior of California Rocky Shore Fishes,”39 Williams introduces the habitat, describes his experiments, and narrates the behavior of the wooly sculpin, Clinocottus analis (Girard), and the opaleye, Girella nigricans (Ayers). This, he believes, “is the first detailed description of the periodic migrations of intertidal fishes.” He observes considerable movement and well-developed homing behavior, but his aim is “to explain this behavior in adaptational terms” (GCW 1957 c, 249). The three-inch-long, large-headed robust sculpin lives in very turbulent water. Williams observes that it has no air bladder and its tissues are much heavier than seawater. Sculpins rest heavily on the bottom when not actively swimming, poised. He notices how they move by “short hops” and the patterns their black-and-white coloring make on the gravel below them. Though the wooly sculpin is the dominant intertidal species, the opaleye is the most conspicuous fish in the rocky shores near Los Angeles because it is “a free swimming, mid-water animal that moves more or less constantly … readily identified by its uniform dark grey color, blue eye and prominent white spots in the middle of the back on either side of the dorsal fin.”40 At low tide, both fish were, in Williams’s era, abundant in stranded pools, and since defense against predators requires movement, becoming accidentally stranded at low tide was dangerous, particularly for the wooly sculpin. Crabs, for instance, ate them. Williams needed to catch fish to sample and tag them, and did so sometimes by fishing with what he calls a “gangen-frame” that he designed, and sometimes with an anesthetic. “After a number of fish were tagged,” as Williams writes, “most of my time in each experiment was spent in quiet observation, frequently from behind natural blinds such as large boulders, and the fish were disturbed as little as possible.” Most observations entailed standing on shore or perching on top of a rock, but when the incoming tide made the water above the pools deeper than three feet, “the turbulence generally required that I get under water with a diving mask to see the fish.”41 So imagine a man with a research program that begins with wooly sculpins elegantly described in their tide pools. Williams is not simply staring at the fish, their habitat, and their learned behavior but exploring an intersection between evolution and ecology. In the 1950s, biologists who engaged in this kind of observation called themselves ethologists. Williams does not, though he refers to Tinbergen’s classic work of ethology, The Study of Instinct (1951).42 Nor is this merely nature study: Williams

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designed experiments to study behavior. What Williams can learn about what fish learn, about their adaptations and ecology, is also framed by the time and space of biological study. These variables shape his career. Of course he might have chosen another niche. He might have been designing experiments to study birds like his friend David Lack, whose ideas Williams later “refined.”43 These niches had certain things in common. Young biologists who inherited the so-called modern evolutionary synthesis, which included a research program that began with “population thinking,” used its distinction between proximate and ultimate causes. As Elliot Sober remembers, Ernst Mayr in particular argued that “Darwinian theory discredited essentialist modes of thought and replaced them with what he has called ‘population thinking.’”44 As Mayr put it, population thinking emphasized the uniqueness of individuals and the critical role of individuality in evolution. “Variation, which had been irrelevant and accidental for the essentialist, now became one of the crucial phenomena of living nature.”45 Mayr also contributed the distinction between ultimate and proximate questions. The proximate question is “what mechanism enables the organism to exhibit the behavior?” and the ultimate question is framed in terms of evolution: “how and why did that behavior come to be?”46 Observing populations of organisms in the wild where “the inshore migration is probably an essential feature in the ecology of both species,” Williams observes “a striking tendency for fish distribution to anticipate the future drainage pattern for the area.” Though any mechanism that allows fish to avoid being stranded in impermanent pools is not obvious, surely homing plays a role. How can they know? Since large numbers of both species dwell in small shallow pools and are always at risk of being left in pools at low tide, Williams expects “special adaptations to these circumstances” because “an animal reacts to what might be called direct influence, such as temperature and presence and absence of suitable cover.”47 But he suspects that more easily measured variables, such as depth or elevation, are largely indirect in their influences. Homing is governed by direct influences. As a scientific term, homing has many meanings, and Williams discusses these at some length. Homing may mean “return of an animal to its home range” or “return of a fish on spawning migrations to its natal stream.” But for his study, Williams takes the term to mean “periodic return of a certain animal to a certain area that is small compared to the total home range.” Fish do this: the same fish goes to the same pools at low tide. He

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postulates, therefore, that an individual fish “actively seeks that particular pool on an outgoing tide.” Fish also follow the same routes up and down the slope as tides change. So, when he finds the same fish in the same pool on two successive observations during two different low tides, he calls that homing, but if he finds the fish in two different pools, he calls that “straying.” His observations are complicated because capture and tagging at low tide disturbs the fish and “strongly effect[s] subsequent behavior.”48 Fish learn to avoid ichthyologists. “Homing in an active mid-water fish such as the Opaleye can only be explained by postulating that the fish recognizes its home pool,” so he attributes to the fish “an accurate spatial familiarity with its home area.” Things are more complicated with the sculpins because tagging disturbs their behavior. Nevertheless, Williams believes he can speak in terms of “the fish’s attitude toward its home pool.” A sculpin must remember these narrowly defined routes to its home pool and must actively seek this pool on outgoing tides.49 He finds no indication that woolly sculpins or opaleyes of a pool “move as a group,” and observes that members of any pool’s population move independently of each other and not to any special environment. Year-round observations reveal that homing is not seasonal. Williams characterizes homing according to two essential features: “repeated occupation of the same small area at low tide” and “extensive movement over neighboring areas between low tides.” These would be difficult to quantify: “To say, for instance, that the Opaleye homes 80 percent of the time (or any other percent) would be a statement as much about the techniques and definitions of the investigator as about the fish.” Nevertheless, homing must be dependent on “a precise behavior mechanism that is inherent in the fish themselves.”50 His conclusion suggests many questions. What is the survival value of homing? Is this a special adaptation to life in the littoral zone, that area close to the shore? Does this adaptation come about specifically because a species frequently exposed to the risk of being stranded “would develop the ability to avoid the traps and find suitable low-tide habitat”? Surely homing plays some part the selection of habitat, but “the question is whether homing is the primary mechanism for avoiding dangers of being left in an unfavorable habitat, and whether this danger was the primary cause for the development of homing behavior.” To these proximate and ultimate questions, Williams has no answers. All he knows for sure is: “As

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long as its home area remains a favorable low-tide habitat, the fish continues to home to it.”51 Williams remained an ichthyologist for his entire life, largely working with marine fish. In fact, a 1991 edition of Rachel Carson’s The Sea Around Us includes an afterword that tells a story of Williams’s study of Iceland eels.52 Williams did extensive research on Icelandic eels. Carson was amazed by the story of migration, reproduction, and hybridization that he discovered. One can think of home in a slightly different and more domestic way. Consider Williams’s fondness for cichlids. Cichlids constitute a diverse and very large family of freshwater fishes including at least 1300 species. They are commonly used in hobbyists’ aquariums. Tinbergen and his students studied the behavior of sticklebacks, cichlids, and gulls, creating a kind of patchwork quilt of animal behaviors.53 Late in his career, Williams contributes a foreword for his friend George W. Barlow’s book on cichlids.54 As Williams writes, “they and other readily available aquarium fish turned me into an ichthyologist at a tender age” and became “a cherished aspect of my fish world.” In particular, as a teenager he was “deeply involved in the sex life and parental behaviors of the firemouth.” Two aspects of cichlids interest Williams. First, he admires the design, the “subtle and intricate mechanism of cichlid jaws,” because they provide a lesson to “someone who still thinks of evolution as a story of steady progress from amoeba to man.” Secondly, he has great interest in comparing the “parental motives” of cichlids with “similar human motivations,” particularly because “cichlid parents seem to show greater love for their offspring as they grow and approach independence.”55 Home and straying.

The One-Way Street: Direction and Perfection The ability of any organism to adapt is rooted in its genes and how they operate. To understand what a gene is to Williams, one must have some knowledge of the so-called central dogma of molecular biology, which— according to its pioneers, James Watson, Francis Crick, Jacques Monod, and François Jacob—entails a one-way flow of information.56 The idea of one-way or single direction inheritance is a legacy of August Weismann’s The Germ Plasm: A Theory of Heredity (1893): germ cells produce somatic cells and cannot be influenced by what somatic cells might learn or any abilities they might acquire. According to Weismann, genetic information

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Fig. 2.1  A diagram of the Weismann barrier, showing the one-way direction of inheritance, where the soma (body cells or organism) cannot influence the germ plasm (genes). Ian Alexander, August 31, 2017, Wikimedia Commons, https:// commons.wikimedia.org/wiki/File:Weismann%27s_Germ_Plasm.svg, Creative Commons license, CC BY-­SA 4.0

cannot pass from soma to germ plasm and on to the next generation; this is referred to as the “Weismann barrier.” In modern and informal terms, hereditary information moves only from genes to body cells, and never in reverse (see Fig. 2.1). Biologists are likely to warn that the Weismann barrier is not the same as the central dogma of molecular biology. Francis Crick coined the phrase “central dogma” in 1957 and published it in 1958. As he wrote, “The central dogma of molecular biology deals with the detailed residue-by-­ residue transfer of sequential information. It states that such information cannot be transferred back from protein to either protein or nucleic acid.”57 No sequential information can travel from protein to DNA or RNA. The central dogma maps a one-way street for genetic transmission. Since it governs evolutionary processes, it provides a linear trajectory for evolutionary history. The first point is that genetic information transfer is sequential, unidirectional, “one-dimensional,” and remarkably isolated within a reproductive process in a species (see Fig. 2.2). A great deal follows from the central dogma, especially in interpreting the constraints on evolutionary change and imagining the perfection of design that evolution is capable of achieving, or prohibited from achieving, as a result of constraints and limitations. In his autobiography, What Mad Pursuit, Crick writes about his choice of the word dogma and some

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Fig. 2.2  A diagram of Crick’s central dogma of molecular biology circa 1958. Wikimedia Commons, https:// commons.wikimedia. org/wiki/ File:Crick%27s_1958_ central_dogma.svg. Public domain

of the problems it caused him. A later exchange that Horace Freeland Judson records in The Eighth Day of Creation is illuminating in this regard: “‘My mind was, that a dogma was an idea for which there was no reasonable evidence. You see?!’ And Crick gave a roar of delight. ‘I just didn’t know what dogma meant.’”58 The central dogma gave a much more precise meaning for the distinction between phenotype, the “outward, physical manifestation” of the organism, and the genotype, the “internally coded, inheritable information” carried by a living organism. Later Williams argued that, though made of DNA material, “The gene is a package of information, not an object. The pattern of base pairs in a DNA molecule specifies the gene. But the DNA molecule is the medium; it’s not the message. Maintaining this distinction between the medium and the message is absolutely indispensable to clarity of thought about evolution.”59 In 1957 and beyond, Williams is interested in what is ephemeral and what is permanent in this process that embodies and transfers information, and he worries about human illusions about this process. As he writes in Adaptation and Natural Selection, the “necessary condition” for the process of adaptation is that “the selected entity must have a high degree of permanence and a low rate of endogenous [inherent] change.” A genotype encoded in DNA would seem to fit the bill, but “the natural selection of phenotypes cannot in itself produce cumulative change, because phenotypes are extremely temporary manifestations.”60 As John Maynard Smith later wrote, If the central dogma is true, and if it is also true that nucleic acids are the only means whereby information is transmitted between generations, this has crucial implications for evolution. It would imply that all evolutionary

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novelty requires changes in nucleic acids, and that these changes—mutations—are essentially accidental and non-adaptive in nature. Changes elsewhere—in the egg cytoplasm, in materials transmitted through the placenta, in the mother’s milk—might alter the development of the child, but, unless the changes were in nucleic acids, they would have no long-term evolutionary effects.61

This strict separation between gene and soma in turn is related to an early idea of population genetics. If one assumes that genes are visible to selection and manifest themselves in phenotypes, it is possible to come to an oversimplified notion that a mutation in one gene predictably leads always to the same change in the phenotype. This view has been called the “one-­ gene one-trait” perspective, deprecated by the phrase Ernst Mayr coined in 1959, as “bean-bag genetics.”62 Nobody takes the one-gene one-trait view any more, and Williams probably never did. Complex relations among multiple genes at multiple sites and relations between genes and environment are now posited, but the evolution of genotypes remains linear and one-way.

Schools Williams uses the gene to think about the properties of schools of fish in “Measurement of Consociation Among Fishes and Comments on the Evolution of Schooling” (1964), where the phenomenon of schooling reveals animal behavior in general. He begins with a paradox, as he often does. Superficially, schooling seems like a simple phenomenon that should lend itself to quantification and causal analysis. But “there are no really convincing ethological, ecological, or evolutionary explanations.”63 First of all, there is no apparent “purpose” for schooling. The assumed purpose is safety, but consider what happens when swordfish or other predator fish attack schools, “swinging their two-edged weapons.” Williams describes the “wastefulness of the attacks” in graphic terms and debunks the idea that schools protect “by rapid satiation of the predator.” After reviewing a set of experiments and observations, he lays down the gauntlet in a typical Williams style: “It is incumbent on those who believe that schools are protective adaptations to show that the structure and working of a school are such as to achieve mass protection in a functionally efficient way.” He can see “no evidence that schools do have such properties.”64

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So what is schooling for? He draws his conclusion, in part, from a set of experiments in tanks inside a laboratory. Williams proposes this central insight, “that a school is not an adaptive mechanism itself, but rather an incidental consequence of adaptive individual behavior.” He reasons in this way. If schooling were adaptive, the school would have to have some functional organization. Such schooling would be based on four factors: “(1) The recognition of conspecific individuals, (2) the desire to approach them, (3) the giving of species-recognition cues, and (4) the initiation of defensive recognition cues when such reactions are detected in other members of the school.” Yet he has observed no alarm signals, as one sees in flocks of birds that are functionally organized. Indeed, he says that any evidence for these in schools of fish would invalidate his position.65 He guesses that schooling occurs when fish seek cover and should be most evident in habitats lacking cover. So he has found, generally. Schooling could be expected in any population that aggregates, precisely because “peripheral or isolated individuals would be especially liable to predation” and one would expect “genetic tendencies to avoid such positions.” He also expects that individuals would be adapted to “react appropriately” to any appearance of flight or distress by other members of the school. But that is not a function of the whole group. He concludes that “the regularities of the properties of schools are those of statistics, not of a functional organization of the whole.”66 Over a decade later, W. D. Hamilton, by then a close friend, writes what he calls “a model of predation in two dimensions” and generalizes many of Williams’s findings. In “Geometry for the Selfish Herd,” he credits Williams’s “admirable study and discussion” of the theory of fish schooling as his inspiration.67 As Hamilton builds on Williams’s work, he dramatizes how well and frequently ideas passed between them, and how they both believed empirical investigations could be used for more inclusive and more mathematical analysis. Hamilton discusses the similarities between the schooling of fish and the herding of land animals to reveal how Williams’s work as a fish biologist can be generalized. Hamilton can extend the ideas of Williams because of their common grounds. Williams, he writes, “points out that schooling is particularly evident in the fish that inhabit open waters. This fits with the view that schooling is similar to cover-seeking in its motivation.” Indeed, “when predation is relaxed gregarious instincts would be selected against.”68 What Hamilton was trying to say was that cover-seeking and gregariousness were related to

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predation, and when predation decreased, gregariousness diminished. A cold view of gregariousness. Hamilton has a second and inverse source for his thinking, namely Sir Francis Galton, best known as a eugenicist who coined the phrase “nature versus nurture,” or so we have been told. Hamilton has some fun with “Galton’s perhaps over-persuasive words,” and includes in “Geometry” the following rather romantic extract from an article he wrote in 1871: To live gregariously is to become a fibre in a vast sentient web over-­spreading many acres; it is to become the possessor of faculties always awake, of eyes that see in all directions, of ears and nostrils that explore a broad belt of air; it is to become the occupier of every bit of vantage ground whence the approach of a lurking enemy might be overlooked.69

Hamilton enjoys responding that there is “nothing in the least altruistic in keeping alert for signs of nervousness in companions as well as for signs of the predator itself, and there is, correspondingly, no difficulty in explaining how gregariousness on this basis could be evolved.”70 It is clear that Hamilton and Williams find Galton’s idea of a “vast sentient web” ridiculous.

Genotype, Phenotype, Fitness Why do Williams and Hamilton think this way? Williams works at the critical intersection of several longstanding biological concepts that are best illuminated by defining a few basic terms, and then examining their relations. Biology and population genetics normally define a genotype as the genetic constitution of an organism as specified by its alleles, an allele being one of a pair of genes in a sexually reproducing species. The phenotype defines the characteristic bodily expression of the organism’s genotype in a particular environment. This relation can be rendered by a scheme:

genotype  environment  random variation  phenotype.

Or more simply

genotype  environment  phenotype.

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A phenotype within an environment generates fitness, upon which natural selection operates.71 At a lower level, as Mark Ridley’s textbook Evolution explains, a gene consists of a sequence of nucleotides coding for a protein or a part of it. A gene may exist in alternative forms called alleles. Allele comes from the German term allelomorph—a term that combines the Greek words allos, meaning other or alias, and morphe, meaning form, shape, or outward appearance. More generally, a gene is the unit of heredity. An allele, then, would be “a variant of a single gene, inherited at a particular genetic locus; it is a particular sequence of nucleotides coding for messenger RNA.”72 As Ridley has written, “Williams defined the gene to make it almost true by definition that the gene is the unit of selection”: the gene is “that which segregates and recombines with appreciable frequency.”73 Biologists generally define evolution in terms of the sum total of the genetically inherited changes in the individuals who are the members of a population’s gene pool. Though individuals feel the effects of evolution, only the population as a whole actually evolves. According to a fundamental principle in population genetics, called the Hardy-Weinberg principle, the genotype frequencies and gene frequencies of a large, randomly mating population remain constant provided immigration, mutation, and selection do not take place. Williams also asserts that “genes are potentially immortal” because there is no physiological limit to their survival; they reproduce faster than external agents can destroy them, and they are stable, rarely mutating. In evolutionary theory, Williams continues, a gene can be defined as “as any hereditary information for which there is a favorable or unfavorable selection bias equal to several or many times its rate of endogenous [internal] change.” Therefore, the stable prevalence of these entities within populations becomes a measure of the importance of natural selection.74 Phenotypes are not immortal. They are bodies; they are soma. Indeed, Williams later credits the British biologist Tom Kirkwood with inventing the term “disposable soma,” but many believe it is Williams’s idea. “If you have a disposable soma, that soma will undergo senescence. If you don’t, it won’t.”75 Williams’s most striking analogy in this regard comes in Natural Selection (1992), where he urges that one consider the soma a material entity but not confuse this observation by claiming that it is a static object:

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On a scale of microseconds a candle flame would seem to be an object. On a scale of seconds it is a place where certain processes occur. Likewise a human soma is an object if viewed for seconds or hours, but not over a period of years or decades … Like a candle flame, it is a region where substances enter, play various roles in various processes, and later depart, usually in altered chemical forms. The persistence of somatic pattern over years and decades is not a material persistence like that of a robot; it is the persistence of information, partly genetic and partly taken from the environment.76

One might read this image as a response to a famous passage about genes in Richard Dawkins’s The Selfish Gene: “Now they swarm in huge colonies, safe inside gigantic lumbering robots, sealed off from the outside world, communicating with it by tortuous indirect routes, manipulating it by remote control.”77 It is revealing that Dawkins images a biological body as a lumbering robot, while Williams imagines a flickering candle flame. The connection between disposable soma and nearly immortal genes appears when one considers the relations among genotypes, phenotypes, and fitness. Regarding various approaches for studying the genetics of adaptation, as a recent review puts it, “Ideally, an allele is designated as adaptive only when connections are made between its genotype, phenotype and fitness.”78 What, then, is fitness? By a standard definition, the fitness of a genotype measures its relative ability to reproduce itself compared to other genotypes. Williams writes, “A thorough grasp of the concept of a gene’s mean phenotypic effect on fitness is essential to an understanding of natural selection.”79 This question leads to a discussion of the temporal duration of any organism’s fitness. Fit for what? Fit for how long?

The Inevitability of Generalized Deterioration: Pleiotropy and Senescence As becomes obvious, Williams’s theoretical interests transcend his focus on fish. From 1957 onwards, a foremost theme in his writing is that living beings age and die. His article “Pleiotropy, Natural Selection, and the Evolution of Senescence” has become a classic.80 It does not speak of death but of the process that leads to it. Its theme is and is not a memento mori, and has a tone somewhat like Emily Dickinson’s lines, “Because I could not stop for Death, / He kindly stopped for me.” The thematic order of Williams’s title is reversed in the order of his presentation.

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Senescence is a widespread problem neglected by non-medical biologists, he writes, because students of natural populations rarely find senile individuals in the wild. He disposes of generally unsatisfactory and harmful theories of aging and the emotional difficulties associated with them. “The most injurious of these,” he writes, “is the identification of senescence with the ‘wearing out’ that is shown by human artifacts.” On the contrary, consider that “the senescence of human teeth consists not of their wearing out but of their lack of replacement when worn out.” Though an artifact is mechanical and static, an organism consists of “an open system in a state of material flux.”81 This would seem to contrast with his mechanical analogies for organisms elsewhere. Williams reviews previous literature, particularly by Peter Medawar, who would win the Nobel Prize in physiology or medicine in 1960, and whom Richard Dawkins called “the wittiest of all scientific writers.” Medawar had presented senescence as “An Unsolved Problem in Biology” in a lecture in 1951 that was published the next year.82 “However,” Williams writes, “the relationship of age to selection has never been precisely formulated.” One sees one of Williams’s characteristic strategies as he thinks through this essay, revealed in this passage: I shall assume initially, therefore, that senescence is an unfavorable character, and that its development is opposed by selection. To account for its prevalence, therefore, it is necessary to postulate another force that favors its development in such a way that the observed variations in senescence reflect variations in the balance between these two forces. I believe that this other force is an indirect effect of selection, and results from the selection of genes that have different effects on fitness at different ages.83

Williams works outward from this contradiction, paradox, or set of opposing forces: “Why senescence?” he asks. An evolutionary explanation of senescence sees inevitable “generalized deterioration” and a system-wide decay of organisms as they age. There must be a correction to theory that can explain this. Williams lists his premises, or factors as he calls them. First, he distinguishes between soma—the non-reproductive parts of the body essential to reproductive success but not passed on in either sexual or asexual reproduction—and the process of natural selection of alternative alleles in a population. Then he postulates “pleiotropic genes of a special sort,” defined to “have opposite effects on fitness at different ages, or, more accurately, in different

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somatic environments.” And lastly but most importantly, he posits “decreasing probability of reproduction with increasing adult age.”84 It is surely no surprise that “natural selection may be said to be biased in favor of youth over old age whenever a conflict of interests arises.”85 But how? What if “a group of adaptively unfavorable morphogenetic changes” of senescence arise as “side effects of otherwise favorable genes, and which have only been partly expurgated by further selection”? A selective force, he thinks, “acts to increase the rate of senescence by favoring vigor in youth at the price of vigor later on.” Yet there is another “force of direct selection that acts to reduce or postpone the ‘price’ and thereby decrease the rate of senescence.” For any particular species, the rate of senescence would depend on the balance between these opposing forces. The term Williams uses for this phenomenon, pleiotropy, means generally, according to the Oxford English Dictionary, “the production by a single gene of two or more apparently unrelated phenotypic effects,” though it is not in widespread public use. It is not likely that pleiotropy will ever become a household term, but George C. Williams is the source of its modern use by biologists. More particularly, the balance between these opposing forces allows for the concept of antagonistic pleiotropy. Though Williams prefers the term balance and does not use the word antagonistic in his article of 1957, the idea has become pervasive in the literature about aging. Pleiotropy’s value to biologists is that the process provides a perfect example of evolution at work. Its etymology means, literally, “many changes.” The idea can be found in the writings of Hamilton and Dobzhansky too. In essence, pleiotropy means that, as life grows more complex, it does not grow more coherent in space, plan, or time. There is always something else coming that could not be expected. After all, there is some mystery to the multiple effects of genes. Different detectives offer varying solutions. Mark Ridley instructs students that, “pleiotropy is the condition in which a gene influences the phenotype of more than one part of the body” and “exists because there is not a one-to-one relationship between the parts of an organism that a gene influences and the parts of an organism that we recognize as characters.”86 But that is not quite what Williams means. To think like Williams about pleiotropy, you must think not only about adaptation but also about maladaptation. One can make a simple definition of Williams’s idea of antagonistic pleiotropy for lay readers by posing this question: How could something be good for an organism when it is young and reproducing, but bad for it when the organism grows older?

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Later in his career, Williams considers the example of the human male’s prostate. Or to open a larger analogy, as evolutionary scientists sometimes do— an always dangerous process—how could something be useful to the human species in that mythical or hypothetical Pleistocene savanna, but become troublesome in the modern age? One is inescapably plunged into the language of human function and design. Can an organism change its biology? asks the biologist. No. “Can people change?” asks Grace Paley in “A Conversation with My Father,” one of a collection of stories entitled Enormous Changes at the Last Minute. The daughter says yes, but the eighty-six-year old father doubts. It depends on perspective. What kinds of changes are available to a biological or social entity? As many have noticed, and as Stephen C. Stearns notes in his precise biographical memoir for the National Academy of Sciences, Williams immediately “went on to derive nine striking predictions that have shaped research on the evolution of aging ever since.”87 Some people speak of these predictions in terms of “trade-offs,” and the conception of trade-offs became central to the study of the evolution of life history. But Williams prefers to speak of costs and benefits in the relationship between reproduction (reproductive success, fitness, vigor) and longevity. Williams spells out implications and exceptions, the most devastating ones coming, perhaps, with numbers five and six: (5) “Senescence should always be a generalized deterioration, and never due largely to changes in a single system.” (6) “There should be little or no post-reproductive period in the normal life-cycle of any species.”

Number five, as he writes, “banishes the ‘fountain of youth’ to the limbo of scientific impossibilities.” There is no consolation, and his conclusions are disappointing, but he thinks perhaps such research might seek more “fruitful directions.”88 Regarding number six, anyone might notice for humans and domestic animals a long period of life after reproduction, sometimes even longer than the period of reproduction, but unfortunately, “these observations lose much of their seeming importance when it is realized that they are largely artifacts of civilization.” Williams does clarify his view. As we all know, “no one is post-reproductive until his youngest child is self-­ sufficient.” Human offspring, for instance, experience a “very long period

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of dependence after conception,” so that any parent “caring for dependent offspring is acting in a way that promotes the survival of his own genes and is properly considered a part of the breeding population.” With women in particular, “at some time during human evolution it may have become advantageous for a woman of forty-five or fifty to stop dividing her declining faculties between the care of extant offspring and the production of new ones.” If this is the case, he argues, “it is improper to regard menopause as a part of the aging syndrome.”89 Williams is generally given credit for being the first to claim that menopause may be an adaptation; this has come to be called the “grandmother effect.” With this article, Williams opens up what came to be called “life-history evolution.” Just as “reproductive maturation is the most important landmark in the life-cycle for the evolution of senescence,” so it would also follow that “senescence may theoretically begin right after this stage in development.” And perhaps the most astonishing conclusion he draws is number nine: “Successful selection for increased longevity should result in decreased vigor in youth.” He elaborates: “I would predict that no human being who is over a hundred years old was unusually vigorous as a young adult.”90 And this from a man at the age of thirty-one years. Surely, Williams thinks, natural selection is responsible for aging and death not through some “specific death-mechanism designed by natural selection to eliminate the old, and members of a population.”91 The teleology of such a theory is loathsome to him. But the trajectory towards death is interesting. Axel Meyer recalls that, when Williams was fifty-­ two years old, he began to record his own senescence: how long it took to run 1700  meters around the track at Stony Brook each year. “Williams presented the graph of his 12  years of slowing speed at his acceptance speech for the Crafoord Prize.”92 Yet he carefully masks himself in the published essay, writing, “I conducted such an investigation on one individual,” and labeling the figure as “age-related decline in speed of a male human subject.”93 A good guess would be that Williams was doing these kinds of experiments on himself all along.

The Language of Altruism According to Stephen Stearns, Williams’s questions directed him from the beginning toward “issues related to adaptation and its limits—aging, altruism, reproductive investment, sex, and medicine.”94 In retrospect, “the question” or “the problem” of the late 1950s and early 1960s is

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succinctly described by those who gave Williams, Ernst Mayr, and John Maynard Smith the Crafoord Prize in 1999 (Hamilton had won it in 1993): “The problem remained that of defining how natural selection functioned—selection among individuals or within a population, or selection at some higher level (‘for the good of the species or group’).”95 This problem became the subject of Williams’s first book. Before launching into that book, one must think of the essay he and Doris Williams wrote about the good any biological individual might be capable of doing for another. Altruism is the theme of Williams’s first published article, coauthored and facilitated by his wife Doris, and entitled “Natural Selection of Individually Harmful Social Adaptations among Sibs with Special Reference to Social Insects”—a roundabout title for reasons that the authors explain.96 Competing forces, costs and benefits, shape the thinking of this article. To begin with, social adaptations can be harmful to individuals if they prevent them from reproducing. To speak of social adaptations and relations is to explore the evolution of behavior. After all, “the adaptive nature of most of the normal relations between members of one species is usually apparent.”97 Active participants receive benefits from their behavior as they cooperate, form packs, and so on, with the benefits to individuals always to be measured in terms of their own reproductive success. But social adaptations also have costs. George and Doris Williams start with an example. Consider the costs to a pregnant mother who not only sacrifices food energy, but also becomes more vulnerable to predation. Yet even sacrifices resulting in the death of the mother “cannot be regarded as deleterious in an evolutionary sense” because such sacrifices are also essential to reproduction. Obviously parents make individual sacrifices and often endanger their own wellbeing for the benefit of others; these sacrifices “may be regarded as the price paid for attendant reproductive advantages, and no special explanation is required.” Relations between parents and offspring are not like relations between siblings. In a larger sense, the Williamses are interested in “the effect of natural selection on genes that influence individually harmful but socially advantageous characters expressed in a sibling group.”98 As a famous legend has it, in a pub one evening, J. B. S. Haldane told his friends that he would risk his life by jumping into a river to save two brothers, but not one, and that he would jump in to save eight cousins, but not seven. (Another version gives Haldane’s quip as a reply when asked if he would give his life to save a drowning brother: “No, but I would to save two brothers or eight cousins.”) Lee Allan Dugatkin has studied the history of

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thought about altruism and points out that “this particular quip makes sense only in terms of gene counting and hints at the logic that would eventually underlie inclusive fitness theory.”99 It also makes sense only if a human acts according to genetic dictates. Doris and George Williams write that “the sorts of characters that we are dealing with are difficult to discuss objectively.” They attribute the term for an altruistic adaptation to Haldane, but demur, saying, “but we prefer to avoid terms so burdened with value judgments and emotional flavor.” They recommend the name social donors for “individuals that sacrifice themselves for the good of others.”100 By extension, there are also social non-donors. What follows is a discussion more mathematical than any other Williams would publish, perhaps because the language of altruism requires an abstract distance. In fact, as Doris Williams acknowledges, “I did the math for the 1957 Evolution paper, although most everything else is attributable to George including the term ‘social donor.’”101 Her mathematical model shows, among other things, that “competition within the sibship will always place the donors at a disadvantage,” but competition between sibships favors donors and so “the two types of selection might balance.”102 Why would an ichthyologist be interested in insect societies? First of all, because “most notable examples of sibling donorism are found there,” and secondly because one can often observe there “total sacrifice of reproductive capacity by the social donors.” Thirdly, these so-called insect servile castes—a troubling term used widely by entomologists including E. O. Wilson, which George and Doris Williams inherit but would prefer to replace with the idea of social donors—“exhibit many striking structural and functional adaptations relating to their donor functions.” Ants are polymorphic; that is, a population includes ants of multiple forms. Wilson explored the origin and evolution of these forms in 1953 when he was a junior fellow at Harvard.103 Morphological and behavioral adaptations create reproductive and so-called “servile castes.” If one speculates that these insect societies developed from family groups when young of different ages were cared for simultaneously by parents, then older sibs were in a position to aid the parents and their aid could result in more efficient reproduction. However, “it might seem unreasonable to expect that genes causing a large proportion of their carriers to be sterile would ever be favored by selection in a breeding population.” Yet if such an arrangement provides one sibling group with an advantage over another, selection could favor it. Surely the process of selection involves more than a single

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genetic change, and George and Doris Williams imagine the advent of sterile castes beginning with a small delay in sexual maturity that “could evolve towards a condition of permanent sterility.”104 That seems to be the case with termites. A key question about this process involves the level of selection. Does selection operate on the level of individuals, colonies, or populations? It would appear, at least in ants and probably in other social insects, that multiple queens within a colony are either sisters or mother and daughters, and social organization is still based entirely on the family. Natural selection reveals itself in changes of populations, but those population changes are created by the reproduction of individuals and accumulate over time. Factors include: How many queens are there, and how are they related? How many offspring are there, and how are they related to others in the colony? George and Doris Williams find no convincing evidence that “intraspecific social organizations of insects ever normally go beyond the family level,” and thus it is “unnecessary to postulate selection at a higher level than the family to explain the evolution of insect societies.”105 This article opens a door to “inclusive fitness theory” that Bill Hamilton, generally taken to be its originator, steps through. Well known for his literary allusions, Hamilton in 1963 recasts the opening sentence of Jane Austen’s Pride and Prejudice—“It is a truth universally acknowledged, that a single man in possession of a good fortune, must be in want of a wife”—in the first sentence of his first published article: “It is generally accepted that the behaviour characteristic of a species is just as much the product of evolution as the morphology.”106 The next year, 1964, he refines and elaborates his theory substantially, creating a more complete and rigorous mathematical model in what becomes the central text of the theory of inclusive fitness. First of all, he argues, “there is nothing special about the parent–offspring relationship except its close degree and a certain fundamental asymmetry. The full-sib relationship is just as close. If an individual carries a certain gene the expectation that a random sib will carry a replica of it is again one-half.”107 So what is called “inclusive fitness” begins exactly with the consideration of sibling relationships that George and Doris Williams undertake. In 1999, when George C. Williams reflects on four decades of discussions of altruism for The MIT Encyclopedia of the Cognitive Sciences, he begins with a familiar caution: “In biology, altruism has a purely descriptive economic meaning: the active donation of resources to one or more

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individuals at a cost to the donor. Moral values or conscious motivations are not implied, and the ideas are as applicable to plants as to animals.”108 What appears to be altruism could be imagined as being facilitated by processes of kin selection, reciprocity, manipulation or deception, and lastly but least likely group selection. These factors favoring altruism are developed by William Hamilton in 1964, Robert L. Trivers in 1971, Robert Axelrod and Hamilton in 1980, David Sloan Wilson in 1980, and others later.109 Yet, as Williams continues to believe, and as his survey of biology reveals, a “greater prevalence of human altruism would be more likely to result from culturally transmitted than genetic differences” (GCW 1999b, 13). He finds little value in seeking sources or models for the human practice of altruism in biological processes. Humans need better reasons or models than inclusive fitness or kinship for their own ethical decisions. Humans are on their own.

Notes 1. S.  C. Stearns, “George Christopher Williams 1926–2010. Biographical Memoir.” National Academy of Sciences (2011), 8. http://www.nasonline.org/publications/biographical-­memoirs/memoir-­pdfs/williams-­ george.pdf. 2. George C. Williams, Adaptation and Natural Selection (Princeton, NJ: Princeton University Press, 1966), 4. 3. Personal communication. 4. David Sloan Wilson and Edward O. Wilson, “Rethinking the Theoretical Foundation of Sociobiology,” the Quarterly Review of Biology 82, no. 4 (2007): 327–48. 5. George C.  Williams, “Preface (1996),” in Adaptation and Natural Selection. Reprint. Princeton Science Library Edition. (Princeton, NJ: Princeton University Press, 1996), x. 6. Ronald Fischer, The Genetical Theory of Natural Selection (Oxford: Clarendon Press, 1930), vii. 7. John Alcock, The Triumph of Sociobiology (Oxford University Press, 2001). 8. Alcock, Triumph, 93, 64. See, for instance, Stephen J.  Gould, “Sociobiology: The Art of Storytelling,” New Scientist 80, no. 1129 (1978): 530–33. Gould and Lewontin speak of “adaptive stories” in their 1979 paper, and Williams is not mentioned. 9. E. O. Wilson, Sociobiology (Cambridge, MA: Belknap/Harvard University Press, 1975), 4.

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10. Ronald de Sousa, “The Sociology of Sociobiology,” International Studies in the Philosophy of Science 4, no. 1 (1990): 271–83. http://homes.chass. utoronto.ca/~sousa/sociosoc.html. 11. Mark Ridley, “Group Selection.” A-Z Browser for Evolution, Third Edition. Blackwell website for the book. http://www.blackwellpublishing.com/ridley/a-­z/Group_selection_.asp. 12. John Tooby and Leda Cosmides, “Conceptual Foundations of Evolutionary Psychology,” in The Handbook of Evolutionary Psychology, ed. David M. Buss (John Wiley and Sons, 2005): 5–67; 8. The Center for Evolutionary Psychology (CEP) at the University of California, Santa Barbara, publishes a great deal of its research online at http://www.cep. ucsb.edu/. 13. Williams, Adaptation, 14, 16. 14. Leif Edward Ottesen Kennair, “Evolutionary Psychology: An Emerging Integrative Perspective within the Science and Practice of Psychology,” The Human Nature Review (January 15, 2002) 2: 17–61, http:// human-­nature.com/nibbs/02/ep.html; Daniel C.  Dennett, Darwin’s Dangerous Idea: Evolution and the Meaning of Life (New York: Simon & Schuster, 1996): 488–91. 15. Wilson, Sociobiology, 4, 27. 16. George C. Williams, “A Package of Information,” in The Third Culture: Beyond the Scientific Revolution, edited by John Brockman (New York: Simon & Schuster, 1995): 39–47: 43, http://www.edge.org/documents/ThirdCulture/h-­Ch.1.html. 17. George C. Williams, Natural Selection: Domains, Levels, and Challenges (New York: Oxford University Press, 1992), 3; Stearns, “George Christopher Williams,” 14. 18. Charles Darwin, “Chapter 7, Miscellaneous Objections to the Theory of Natural Selection,” in The Origin of Species, 6th edition. http://www.literature.org/authors/darwin-­charles/the-­origin-­of-­species-­6th-­edition/ chapter-­07.html. 19. Ernst Mayr, What Evolution Is (New York: Basic Books, 2001). Jerry A. Coyne, Why Evolution is True (New York: Penguin Books, 2009). 20. See Gregory J. Cooper, The Science of the Struggle for Existence: On the Foundations of Ecology (Cambridge, UK: Cambridge University Press, 2003), 66–67. 21. W. D. Hamilton, Narrow Roads of Gene Land vol. 1: Evolution of Social Behaviour (Oxford: Oxford University Press, 1996), 20. 22. Robert J. Richards, The Tragic Sense of Life: Ernst Haeckel and the Struggle over Evolutionary Thought (Chicago: University of Chicago Press, 2008), 264–68. 23. Richards, Tragic Sense of Life, 268.

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24. See Carl Zimmer, “Stretching the Limits of Evolutionary Biology: A Profile of George Williams,” Science (May 28, 2004): 1235–36. 25. Randolph M. Nesse, “George Williams, 1926–2010” (Friday, September 10, 2010), http://skepticaladaptationist.com/2010/09/george-­ williams-­1926-­2010.html. 26. See a recent attempt by Stuart West and colleagues to clean up the semantics of altruism and related terms: Stuart A.  West, A.  S. Griffin, and A.  Gardner, “Social Semantics: Altruism, Cooperation, Mutualism, Strong Reciprocity and Group Selection,” Journal of Evolutionary Biology, 20 (2007): 415–32: 416. 27. Italo Calvino, Six Memos for the Next Millennium (Cambridge, MA: Harvard University Press, 1988), 90. 28. Calvino, Six Memos, 124. 29. Robert MacArthur and E. O. Wilson, The Theory of Island Biogeography (Princeton, NJ: Princeton University Press, 1967), 149 and ff. 30. R. A. Fisher, “The Actuarial Treatment of Official Birth Records,” The Eugenics Review 19, no. 2 (July 1927): 103–8, 103. 31. Robert L. Trivers, “The Evolution of Reciprocal Altruism,” the Quarterly Review of Biology 46, no. 1 (March 1971): 35–57. 32. Frans Roes, “A Conversation With George C. Williams,” Nat. Hist. 107 (May 1998): 10–13. 33. Williams, Adaptation, 23. 34. Frank C.  Erk, “George C.  Williams: Personal Reminiscences,” the Quarterly Review of Biology 80, no. 1 (2005): 7–11, 8. 35. Howard Cunnell, “Fast This Time: Jack Kerouac and the Writing of On the Road,” an introduction to Jack Kerouac, On the Road: The Original Scroll, edited by Howard Cunnell (New York: Penguin, 2007): 1–52, 1. 36. Raymond Williams, “Naturalism,” Keywords: A Vocabulary of Culture and Society (New York: Oxford University Press, 1988): 216–19. 37. Thomas Nagel, “A Philosopher Defends Religion,” a review of Alvin Plantinga, Where the Conflict Really Lies: Science, Religion, and Naturalism (New York: Oxford University Press, 2011), in NYRB (September 27, 2012). Plantinga makes an overall claim that “there is superficial conflict but deep concord between science and theistic religion, but superficial concord and deep conflict between science and naturalism” (ix). 38. Frank J.  Sullaway, Born to Rebel: Birth Order, Family Dynamics, and Creative Lives (New York: Pantheon Books, 1996), 95. 39. “Homing Behavior of California Rocky Shore Fishes,” Univ. Calif., Publ. Zool. 59 (1957): 249–284. 40. George C.  Williams, “Homing Behavior of California Rocky Shore Fishes,” University of California Publications in Zoology 59 (1957): 249–84, 249, 250.

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41. Williams, “Homing Behavior,” 261, 253. 42. Niko Tinbergen, The Study of Instinct (New York: Oxford University Press, 1951). 43. See Cooper, Science of the Struggle for Existence, 58. 44. Elliott Sober, “Evolution, Population Thinking, and Essentialism,” Philosophy of Science (1980): 350–83. 45. Ernst Mayr, One Long Argument (Cambridge, MA: Harvard University Press, 1991), 42. 46. Ernst Mayr, “Cause and Effect in Biology,” Science 134, no. 3489 (November 10, 1961): 1501–6, 1503. 47. Williams, “Homing Behavior,” 256, 257, 258. 48. Williams, “Homing Behavior,” 259, 262, 264, 265. 49. Williams, “Homing Behavior,” 273, 275. 50. Williams, “Homing Behavior,” 278, 279, 280. 51. Williams, “Homing Behavior,” 280–81. 52. Rachel Carson, The Sea Around Us (New York: Oxford University Press, 1991), 231. 53. Niko Tinbergen, The Study of Instinct (New York: Oxford University Press, 1951), 179. 54. George C. Williams, “Foreword” to George Barlow, The Cichlid Fishes: Nature’s Grand Experiment in Evolution (New York: Basic Books, 2000), ix–xi. 55. Williams, “Foreword” to The Cichlid Fishes, ix–xi. 56. See Horace Freeland Judson, The Eighth Day of Creation: The Makers of the Revolution in Biology (New York: Touchstone, 1979). 57. See Francis Crick, “The Central Dogma of Molecular Biology,” Nature 227 (August 8, 1970): 561–63. 58. Judson, Eighth Day of Creation, 337. In his autobiography, Crick writes, “I had already used the obvious word hypothesis in the sequence hypothesis, and in addition I wanted to suggest that this new assumption was more central and more powerful.” But as it turned out, the word dogma created its own problems. “Many years later Jacques Monod pointed out to me that I did not appear to understand the correct use of the word dogma, which is a belief that cannot be doubted. I did apprehend this in a vague sort of way but since I thought that all religious beliefs were without foundation, I used the word the way I myself thought about it, not as most of the world does, and simply applied it to a grand hypothesis that, however plausible, had little direct experimental support.” Francis Crick, What Mad Pursuit: A Personal View of Scientific Discovery (New York: Basic Books, 1988), 109. 59. Williams, “Package of Information,” 43. 60. Williams, Adaptation, 23.

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61. John Maynard Smith, Evolutionary Genetics, 2nd ed. (Oxford: Oxford University Press, 1998), 10. 62. See Ernst Mayr, “Darwin and the Evolutionary Theory in Biology,” in Evolution and Anthropology: A Centennial Appraisal, edited by Betty J.  Meggers (Washington, DC: The Anthropological Society of Washington, 1959): 1–10. 63. George C. Williams, “Measurement of Consociation among Fishes and Comments on the Evolution of Schooling,” Michigan State University Museum Publications, Biology Series 2 (1964): 351–83, 351–2. 64. Williams, “Measurement of Consociation,” 375. 65. Williams, “Measurement of Consociation,” 376–77. 66. Williams, “Measurement of Consociation,” 378, 380. 67. William D.  Hamilton, “Geometry for the Selfish Herd,” Journal of Theoretical Biology 31, no. 2 (1971): 295–311, 298, 306. 68. Hamilton, “Geometry,” 298, 306. 69. Galton quoted in Hamilton, “Geometry,” 309. 70. Hamilton, “Geometry,” 309. 71. Richard Lewontin, “The Genotype/Phenotype Distinction,” The Stanford Encyclopedia of Philosophy, edited by Edward N.  Zalta (Spring 2007). http://plato.stanford.edu/archives/spr2007/entries/ genotype-­phenotype/. 72. Mark Ridley, Evolution (Malden, MA: Blackwell Publishing, 2004), 308. 73. Ridley, Evolution, 308; Williams, Adaptation, 24. 74. Williams, Adaptation, 24, 25. 75. Frans Roes, “A Conversation With George C. Williams,” Nat. Hist. 107 (May 1998 ): 10–15; See also T. B. Kirkwood, “Evolution of Ageing,” Nature 170 (November 24, 1977): 201–4. 76. Williams, Natural Selection, 17–18. 77. Richard Dawkins, The Selfish Gene (New York: Oxford University Press, 1976), 19. 78. R.  D. H.  Barrett, and H.  E. Hoekstra, “Molecular Spandrels: Tests of Adaptation at the Genetic Level,” Nature Reviews Genetics 12, no. 11 (2011): 767–780. 79. Williams, Adaptation, 25. 80. George С. Williams, “Pleiotropy, Natural Selection, and the Evolution of Senescence,” Evolution 11, no. 4 (1957): 398–411. 81. George C. Williams, “Pleiotropy, Natural Selection, and the Evolution of Senescence,” Evolution 11 (1957): 398–411, 398. 82. Richard Dawkins, The Oxford Book of Modern Science Writing (New York: Oxford University Press, 2008): 179; Peter Medawar’s 1952 essay, “An Unsolved Problem in Biology” (H. K. Lewis, London) is reprinted in his book, The Uniqueness of the Individual (London: Methuen, 1957): 44–70.

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83. Williams, “Pleiotropy,” 399. 84. Williams, “Pleiotropy,” 406, 400. 85. Williams, “Pleiotropy,” 401. 86. Ridley, “Pleiotropy,” A-Z Browser, Evolution, http://www.blackwellpublishing.com/ridley/a-­z/Pleiotropy.asp. 87. Stearns, “George Christopher Williams,” 7–8. 88. Williams, “Pleiotropy,” 406, 407. 89. Williams, “Pleiotropy,” 407, 408. 90. Williams, “Pleiotropy,” 409, 410. 91. Williams, “Pleiotropy,” 398. 92. Axel Meyer, “George C.  Williams (1926–2010): Incisive Thinker Who Influenced a Generation of Evolutionary Biologists,” Nature 467 (October 14, 2010): 790. 93. George C. Williams, “The Tithonus Error in Modern Gerontology,” the Quarterly Review of Biology 74, no. 4 (December 1999): 405–15, 411. 94. Stearns, “George Christopher Williams,” 8. 95. The Royal Swedish Academy of Sciences, “The Crafoord Prize 1999,” The Crafoord Prize website. https://www.crafoordprize.se/press_ release/the-­crafoord-­prize-­1999. 96. George C.  Williams and Doris C.  Williams, “Natural Selection of Individually Harmful Social Adaptations among Sibs with Special Reference to Social Insects,” Evolution (1957): 32–39. 97. Williams and Williams, “Natural Selection,” 32. 98. Williams and Williams, “Natural Selection,” 32. 99. Lee Alan Dugatkin, “Inclusive Fitness Theory from Darwin to Hamilton,” Genetics 176 (July 2007): 1375–80, 1376. 100. Williams and Williams, “Natural Selection,” 32–33. 101. Personal communication, November 9, 2012. 102. Williams and Williams, “Natural Selection,” 34. 103. Edward O.  Wilson, “The Origin and Evolution of Polymorphism in Ants,” the Quarterly Review of Biology 28, no. 2 (1953): 136–56. 104. Williams and Williams, “Natural Selection,” 36, 37. 105. Williams and Williams, “Natural Selection,” 38. 106. William D.  Hamilton, “The Evolution of Altruistic Behavior.” The American Naturalist 97, no. 896 (1963): 354–56. 107. William D. Hamilton, “The Genetical Evolution of Social Behaviour. I and II,” Journal of Theoretical Biology 7, no. 1 (1964): 1–52, 1–2. 108. George C. Williams, “Altruism,” in The MIT Encyclopedia of the Cognitive Sciences, edited by Robert A. Wilson and Frank C. Keil (Cambridge, MA: MIT Press, 1999), 12–14. http://ai.ato.ms/MITECS/Entry/williamsg.html.

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109. In addition to Hamilton’s “Genetical Theory of Social Behavior,” Williams cites a set of “seminal” titles in 1999, including these: Robert Axelrod and W. D. Hamilton. “The Evolution of Cooperation,” Science 211 (1980): 1390–1396; D.  J. C.  Fletcher and C.  D. Michener, Kin Recognition in Animals (New York: Wiley-Interscience, 1987); Matt Ridley, The Origins of Virtue (New York: Viking Press, 1996); T. D. Seeley, “The Honey Bee as a Superorganism,” American Scientist 77 (1989): 546–553; P.  J. B.  Slater, “Kinship and Altruism” in P.  J. B.  Slater and T. R. Halliday, eds., Behavior and Evolution (Cambridge University Press, 1994); Robert L. Trivers, “The Evolution of Reciprocal Altruism,” the Quarterly Review of Biology 46: 35–57; David Sloan Wilson, Natural Selection of Populations and Communities (Boston: Benjamin/Cummings, 1980); David Sloan Wilson and Elliott Sober, “Re-introducing Group Selection to the Human Behavioral Sciences,” Behavioral and Brain Sciences 17 (1994): 585–654; Richard Wright, The Moral Animal: Why We Are the Way We Are (New York: Vintage Books, 1994); V. C. WynneEdwards, Animal Dispersion in Relation to Social Behavior (London: Oliver and Boyd, 1961).

CHAPTER 3

Proper Discourses of Design and Natural Selection

The ideas expressed in Adaptation and Natural Selection: A Critique of Some Current Evolutionary Thought (1966) were primarily a result of the collaboration of contemporaries Bill Hamilton and George Williams, though others, like David Lack and Colin Pittendrigh, contributed. Williams and Hamilton had literary skills, but Hamilton tended to mathematical solutions for problems, and Williams engaged in thought experiments. Not as full throated in style as Dawkins’s The Selfish Gene, which came out a decade later, this short book announces that it examines the interior of biological thought. Williams’s discourse is, in other words, meant to be thoughtful, not arrogant. I am interested in not only the content of this manifesto, but also its rhetorical stance, how the author approaches his material. Though Dawkins and others believe that Williams was “correct,” and that scholars of this literature should be “getting it right,” critique is not simply a matter of corrections, but of opening discussion, generating dialogue. For that reason, I offer a close reading of Williams’s rhetorical structures and use his language as much as possible, along with that of his critics and his sources, as if the reader were witnessing a discussion. For reasons that will become clear, also I divide my reading into two parts, corresponding to the first and second halves of Adaptation and Natural Selection. In the first, Williams examines the traditional concept of

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 M. P. Cohen, George C. Williams and Evolutionary Literacy, Literatures, Cultures, and the Environment, https://doi.org/10.1007/978-3-031-11650-6_3

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adaptation and in the second he opens up newer explorations called sociobiology and life history. As his title suggests, he grounds evolutionary theory on the process called adaptation, which is a result of changing genetic information in populations. So anyone who asks what evolution is sooner or later must ask how adaptation occurs, what drives it, what it can accomplish, and where it is a strong or weak tool for the design of living beings. Though selection is a blind process that has neither goal nor plan, selection must surely occur between adaptations. This entails a statistical process. Although humans may have predilections toward agile, fast, articulate, strong, colorful, and clever forms of life, may value the diversity of wild beings and feel themselves to understand creation, selection has no aesthetic. Though humans have created appealing ideas and useful domestic creatures, as Darwin noticed, evolution, because it is a statistical process, has no such aspirations. When discussions of evolution turn to purpose, they are about the human discussants themselves.

Maintaining Adaptation Through Analogy To simplify somewhat, a biologist speaking of a specific adaptation refers to some trait whose current functional role in the life history of an organism has evolved, and says it is assumed also to be “maintained” by natural selection, as Williams puts it, “as a matter of definition.” To cause some existing state to continue, to repair, persevere, sustain. Thus natural selection maintains adaptations, which is to say, “A species maintains itself by the activities of its members despite destructive external influences,” and Williams goes on, “so does a nation. But does a species have anything at all akin to a spirit of nationalism?”1 And by analogy, humans maintain positions in discussions or arguments. Williams uses the word “maintain” thirty-seven times in his book, usually in the former sense, but also, and sometimes strenuously, in the latter. Does he mean to dramatize a continuing and dynamic process in his own thought? In any case, as Ernst Mayr says, “Adaptation, insofar as such a concept existed, could no longer be considered a static condition, a product of a creative past, and became instead a continuing dynamic process.”2 There are proper and improper, accurate and inaccurate, ways of speaking of this dynamic, proper because these ways or discourses lead to clear thinking about and understanding of (1) evolution’s processes and the organisms that result, and (2) concepts that biologists use, how they are articulated, and how they can be tested or modified. Williams

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recommends to biologists what he considers a necessary maturity of discourse—which might also be described as an austerity of discourse. He pinpoints preconceptions that cause difficulties. In general, he advocates higher standards of rigor and discipline in discussing these matters, particularly for biologists who speak to the public. He speaks in the first person, stressing the importance of biological means and ends, because “such a conceptual framework is the essence of the science of biology.” How does one investigate means and ends, when the genetical theory of natural selection consists of a statistical bias in relative rates of survival of alternatives, of genes and individuals? How can he speak as if the “ultimate essence the theory of natural selection” deals with “a cybernetic abstraction, the gene, and a statistical abstraction, mean phenotypic fitness”?3 Because his “ultimate essence” sounds like an abstraction of an abstraction, he is led to depend upon analogies and narratives for explanation. What many people imagine about evolution is not what scientists think: It is difficult for many people to imagine that an individual’s role in evolution is entirely contained in its contribution to vital statistics. It is difficult to imagine that an acceptable moral order could arise from vital statistics, and difficult to dispense with belief in a moral order in living nature. It is difficult to imagine that the blind play of the genes could produce man.4

All people use analogies when speaking or writing of adaptation, and in this sense, Williams creates a perhaps onerous argument about the use of one of his analogies that a reader can easily follow: “A frequently helpful but not infallible rule is to recognize adaptation in organic systems that show a clear analogy with human implements.” We recognize convincing analogies between bird wings and airship wings, between bridge suspensions and skeletal suspensions, between the vascularization of a leaf and the water supply of a city. In all such examples, for Williams, “conscious human goals have an analogy in the biological goal of survival.”5 Sometimes, he both approves such analogies and undercuts them. For instance, one might think about the progress of human designs using the analogy of propeller-driven aircraft being replaced by jets, but closer inspection reveals that progress “need not imply that the improvement involves an increase in complexity.” For instance, vertebrates with vertical biting devices—that is, jaws—almost entirely replaced jawless creatures, presumably because they were more effective fishes. And flowering plants whose seeds are enclosed by ovaries or fruits largely replaced the

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gymnosperms with naked and enclosed seeds, like pines, “presumably because they were more effective at making their own food and fixing carbon.” But when during the Pliocene a mass extinction of many of the higher mammalian carnivores, ungulates, and primates occurred, most primitive mammals and lower groups were little affected, which is to say, “the game can be played both ways.” Therefore, one cannot cite as evidence of adaptation “selected examples of the supposed operation of a process, such as the dominance of recently evolved types over the more ancient.” And this is his point: analogies linking adaptation to improved design are likely to be fallible.6 Because anthropomorphism is ubiquitous in all forms of analogy, “There may also be a desire, unconscious in many and expressed by a few, to find not only an order in Nature but a moral order,” and such uncritical analogy with self-conscious human organizations, though probably not the whole explanation for humanization of groups of organisms, satisfies such desires at the expense of accuracy. “There is a rather steady production of books and essays that attempt to show that Nature is, in the long run and on the average, benevolent and acceptable to some unquestionable ethical and moral point of view.”7 So a desire to find a benevolent process in Nature is fostered by a desire to analogize between human societies and populations of species, and may in fact be the germ of the idea of group selection. Williams asks whether a species could have a collective will to avoid extinction and answers, “No modern biologist has explicitly proposed that such factors are operative in the history of a species, but I believe that biologists are unconsciously influenced by such thinking, and that this is true of some distinguished and capable scholars.”8 By creating its own analogies, religion also creates certain problems, and Williams thinks that “biology would have been able to mature more rapidly in a culture not dominated by Judeo-Christian theology and the Romantic tradition. It might have been well served by the First Holy Truth from the Sermon at Benares: ‘Birth is painful, old age is painful, sickness is painful, death is painful.’”9 When Williams speaks of pain, but not his own, he reveals what he will and will not include in his discussions. The biologist recognizes pain; because he has no way to assess it in others, and perhaps because of Tinbergen’s dicta, he cannot speak of the pain experienced by other lives concretely. And what I would say is that the lack of the personal reveals Williams’s well-known lack of vanity. For one thing, he chooses to keep his own personal experience private, perhaps in order to avoid the confusion certain

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analogies create. But there is something more here. Williams may be fond of fish and fascinated by their mouths as he has admitted throughout his texts. That is personal. But it is quite another thing to be fond of your own theories, or proud of them. I define that impulse as vanity.

Pattern Versus Design An observer is mesmerized by a compelling pattern when watching a school of fish. Surely such a seemingly coordinated dance of living beings is striking. However, by definition a pattern must have a model. But for Williams only cause-and-effect relationships are patterns. Some observers describe patterns made by groups as clues to “swarm intelligence” that suggest the group has some overall emergent structure or design. Nevertheless, Williams’s fundamental question remains. What in any pattern benefits the individual? How does an individual sense cues and consequently behave to its own benefit? A cue, as identified by Tinbergen and ethology generally, triggers a behavior. A cue does not require consciousness from the organism, but only a response observable to the scientist. Cues may also be thought of as signals or signs. Mark Hauser wrestles with this issue at the beginning of The Evolution of Communication.10 Tinbergen, following Lorenz, speaks of behavior being “released” by certain “sign stimuli” in an animal’s environment, to which it reacts in a particular way. This discussion has become more complicated of late. Some ethologists speak of signals and cues as stimuli that contain information, but distinguish signals that were molded by natural selection from cues that were not. Tinbergen’s idea of “sign stimuli” is at the root of what Williams means by a cue, and the way this idea relates to behavior allows him to speak of a pattern. To avoid confusion, an observer must first “distinguish between response to environmental stimuli and susceptibility to environmental interference,” because responses and susceptibilities “both conform to a pattern of cause-­ effect relationships involving organisms and their environment.” For Williams, “a response shows the unique biological property of adaptive organization,” while “susceptibility results from the absence or deficiency of this property.” Response requires not only the ability to sense specific aspects of the environmental situation but also some means of resisting effects from the environment, while susceptibility occurs when an environment produces an effect “in spite of any responses that may be activated.”11 He illustrates this principle by contrasting a human with a reptile:

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the former’s metabolic rate responds to changes in temperature, while the latter is susceptible to these changes. When Williams seeks patterns of response, he goes beyond Tinbergen’s focus on instinct to indicate that elaborate systems of behavior, particularly reproductive patterns, “will usually be a blend of learned and instinctive elements.” Nevertheless, instinct is at the root of his discussion of pattern. In his 1966 book, he focuses on sensory and motor patterns necessary for the feeding and growth that maximize the next generation as well as patterns of behavior, patterns of interaction, mating patterns, patterns of parental behavior, patterns exhibited by larger populations, and patterns shown by his own graphs. Overall, he emphasizes widely prevalent patterns of self-seeking behavior in the animal kingdom and finds them “easily attributed to selection for competitive efficiency in genetic survival.”12 Note how his syntax separates the pattern from the actor. This leads him to conclude that “schooling would be based on active behavior patterns and on passive exhibition of cues.” Williams recapitulates this view repeatedly: analysis of cause and effect must ask “whether the observed effect is caused by environmental interference or by the organism acting in response to some cue.” Not all cues result in an adaptive response, but “If an environmental cue evokes what is thought to be an adaptive response it should be so identified, and the nature of the postulated adaptation explained.” For instance, the behavioral patterns of predators like swordfish may be interpreted as “adaptations designed to exploit the schooling behavior of prey.”13 He advises great care because this task of interpretation is by no means straightforward. “Even in groups in which environmental cues determine sex, for instance the reptiles, the responses may make little adaptive sense.”14 The point, however, is that recognizing the morphological and behavioral patterns of a group must lead to causal explanation in terms of the response of individuals. To speak of causal explanation entails speaking of design. Pattern can reveal design. From the onset, Williams announces “I will use terms appropriate to human artifice and conscious design,” because— and he lapses into passive voice—“designation of something as the means or mechanism for a certain goal or function or purpose will imply that the machinery involved was fashioned by selection for the goal attributed to it.” Design is a term central to the Williams lexicon. The word’s history is Latinate, and constructed from the word signare. To design is to mark out, to make a sign. As any dictionary will reveal, the term design has experienced many modern metaphoric extensions. Being highly

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susceptible to attributing design, humans often reveal their own predilections. “The central biological problem is not survival as such, but design for survival.”15

Critique and Causality So much for pattern, analogy, and design. We are now ready to read Williams’s first sentence in Adaptation and Natural Selection: “Evolutionary adaptation is a special and onerous concept that should not be used unnecessarily, and an effect should not be called a function unless it is clearly produced by design and not by chance.”16 This first sentence of the annotated table of contents has become the most frequently extracted from the text, and for good reason. Williams repeats it with variation four more times. Underlying this statement, two skeptical premises shape his discourse. The first is that every feature observed in a modern organism cannot be assumed to be an adaptation. The second is that the physiology and behavior of an organism cannot be assumed to be good for any group to which it is imagined to belong. These premises have widespread implications for the grounding of biological thought because they call for a set of re-­ evaluations. Steven Stearns notices that Williams also introduces other “novel interpretations,” among them: (1) Selection for evolutionary adaptability cannot be prescient; (2) “Species do not have high fecundity to compensate for high mortality, as had often been supposed; rather, high mortality is the consequence of high fecundity in ecologically limited populations”; (3) Delayed reproduction may or may not be individually advantageous, depending on costs and benefits, risk of the mother’s death versus her increased fecundity later in life; (4) As with schools of fish or flocks of birds where an individual seeks gain by placing itself inside a group and puts others between itself and sources of danger, so “All the processes observed in ecosystems are the aggregate consequences of features, such as photosynthesis and predation, that are advantageous to individual organisms.”17 Thus, Williams engages in what literary critics call an intervention. Most of his interpretations challenge and often invert the thinking about means and ends used in what he calls “current evolutionary thought”— the phrase itself signaling dissatisfaction with professional discussion of the relationships between adaptation and change, fecundity and death, individuals and groups. He intervenes—that is, stands between long-held but

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untenable ideas and what he understands to be clear thinking on the subject. His reading of these ideas becomes a form of rewriting; his interpretations desacralize and demythologize. Most of all, Williams interrogates standard biological discourse by providing an incisive critical analysis of the questions biologists are not asking. He shapes a way to more rigorously articulate theories of natural selection, his aim being to provide grounding for more productive discussions. Consider his acute dissatisfaction with the idea of beneficial death that I mention in the introduction. This old idea was that death evolved to cull populations so that fitter individuals could take their rightful places. “My reaction was that if [A. E.] Emerson’s presentation was acceptable biology, I would prefer another calling.”18 One should not make light of the expression “another calling,” since this kind of characteristic introspection not only reveals the central authorial questions of American literature— “What do I have to present to the world?” and “What is my calling?”—but also reveals the constant self-questioning that marks Williams’s writing. In general, Williams feels called upon to disassemble a set of imagined phenomena attributed to what A. E. Emerson and others called “group selection,” where adaptations recognized by professional biologists were properties of the group, which “often required individual members to jeopardize their own interests for group welfare.”19 These biologists believed that groups regulated their own populations, exerting control above the level of the individual; they conceived ideas like beneficial death, even asserting that as a rule animals practice effective birth control, as suggested by V.  C. Wynne-Edwards in a variety of publications. Wynne-­ Edwards believed in a set of group-directed controls of a variety of demographic phenomena, among them regulation of the ratio of males to females and the origin and maintenance of sexual reproduction.20 In 1971, Williams finds the idea that natural populations practice birth control ridiculous, and he is no doubt aware that when ecologist Paul Ehrlich published The Population Bomb three years earlier, he turned the idea of birth control into an explicit political movement, by insisting that biological possibility could or should not determine human choice.21

Parsimony and Style Life is bracketed by conception and dissolution, but reproduction connects them as a single process, as Williams had proposed in “Pleiotropy” (1957). His continuous and coherent verbal exploration of means and ends, cause and effect, in biology constitutes “one long argument” framed

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by sex and death. It begins with early contributions, is capped by “The Tithonus Error in Modern Gerontology” (1999), and includes, at the center of his career, the monographs Adaptation and Natural Selection (1966), Sex and Evolution (1975), and Natural Selection: Domains, Levels, and Challenges (1992). Yet he often portrays himself a refiner of ideas rather than an originator of them, as evidenced by the title of one article: “Natural Selection, the Costs of Reproduction, and a Refinement of Lack’s Principle.”22 He was also a prolific reviewer of scientific literature. The trajectory of his writings also demonstrates how, recursively, he takes his own critique and those of others to heart, constantly questioning his own thoughts, reconsidering, and refining them. His writing is concise and restrained. There is no nonsense or hyperbole; metaphors are distinctly subdued; no taxonomies appear; no purple prose like the conclusion to Darwin’s Origin adorns his books. This suggests that he did not seek fame. As he writes of Adaptation and Natural Selection, “I fully expected that the perspective I urged would ultimately be accepted as orthodox; however, I did not at that time expect it to prevail so soon.”23 A writer who aims for parsimony—who aims to “recognize adaptation at the level necessitated by the facts and no higher”—would, as a matter of course, likely approach a minimalist style.24 A more complicated question is, what kind of voice would be appropriate to explore a powerful process that operates at very low levels by blind and selfish chance at the expense of its designed products, purely by trial and error? Williams often uses concrete examples. Such a story, this one of flying fish, appears early in Adaptation and Natural Selection. A fish has just left the water. It cannot survive long in the air. As Williams comments wryly, it is “a matter of common observation that an aerial glide normally terminates with a return to the sea.” Because termination of flight is not a mechanism for getting the fish back into the water, there is no reason to invoke the principle of adaptation. Gravity is quite sufficient. “The real problem is not how it manages to come down, but why it takes it so long to do so,” and “it would be absurd to recognize an adaptation to achieve the mechanically inevitable.” Then comes the argument’s turn: “I believe, however, that this is essentially what is done by those who propose that gene mutation is a mechanism for ensuring evolutionary plasticity”.25 After all, just as the flight of the fish is terminated by mechanical law, so genes mutate according to mechanical, not forward-­thinking laws. So too, when one understands mutation as mechanical, the flight of those who propose that mutation looks forward in time crashes down to earth.

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Occam’s Razor as Mirror or Window One can explore evolutionary theory without factoring in styles of writing, but to do so is to miss its literary tradition that includes Darwin’s plots and metaphors. In this instance, one enters the portal of Adaptation and Natural Selection only when one recalls the almost universally accepted principle of parsimony, sometimes known metaphorically as Occam’s razor, a principle that shapes Williams’s logic, thought, explanation, and writing style. Williams follows this principle implicitly, announcing that it would be a violation of parsimony to assume multiple levels of explanation when one will suffice. This principle impinges on but does not determine all explanations of adaptation. Though one might recognize levels of adaptive organization from the subcellular to the biospheric, “the principle of parsimony demands that we recognize adaptation at the level necessitated by the facts and no higher,” and that it “need almost never be recognized at any level above that of a pair of parents and associated offspring.” This premise seldom has to rest on appeals to parsimony alone, as he notes, “but is usually supported by specific evidence.”26 When Williams invokes the principle called Occam’s razor, he defines the shape of reduction—the process he understands as epitomizing scientific thought. Note that he also invokes or embeds a story or narrative, of a pair of parents and an offspring. Occam’s maxim is itself written. It is normally taken to be entia non sunt multiplicanda praeter necessitatem (entities must not be multiplied beyond necessity), though what he actually wrote was apparently numquam ponenda est pluralitas sine necessitate (plurality must never be posited without necessity). The poles of his maxim are entities and necessity. Philosophers continue to suffer over Occam’s principle. As Ludwig Wittgenstein, writing in his famously compressed style, understood, “Occam’s razor is, of course, not an arbitrary rule nor one justified by its practical success. It simply says that unnecessary elements in a symbolism mean nothing. Signs which serve one purpose are logically equivalent, signs which serve no purpose are logically meaningless.” For Wittgenstein generally, “The procedure of induction consists in accepting as true the simplest law that can be reconciled with our experiences.”27 To think clearly, logical entities are not to be multiplied needlessly. In its most

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radical form, as Wittgenstein points out, the maxim suggests that unnecessary signs not only serve no purpose, but mean nothing, are logically meaningless. This is exactly what Williams means when he deprecates the sign “group selection” by explicit reference to Occam.28 Some wish to assert that Occam’s razor is really a practical aesthetic, leading to a style (of writing) that values parsimony, economy, succinctness. Though this may be debated, it does remind us that kinds of logic are related to writing styles. To be specific, biology’s styles, and Darwin’s in particular, have often multiplied examples and possible causes, valuing diversity of exposition as a way to embody in prose “endless forms most beautiful,” and have consequently tended toward aesthetic elaboration. Elliott Sober, an eminent philosopher of biology at the University of Wisconsin, responds to Williams and his principle in “Let’s Razor Ockham’s Razor.” Though Sober recognizes Williams’s 1966 concern that various “group” hypotheses were the result of sloppy thinking and appreciates his desire to obtain a “properly rigorous Darwinism,” he focuses on Williams’s analogy or allegory of the flying fish by noticing that Williams follows Lloyd Morgan’s rule that “lower-level explanations are preferable to higher-level ones.” Yet there is no need to invoke parsimony for this phenomenon, he argues, and constructs an alternate argument, positing unlikely dual ancestral populations of flying fish, one heavier and the other lighter than air. As Sober writes, it is enough to doubt that there could be a population lighter than air. No need for parsimony here.29 Sober pursues Williams by entering another of his narratives, of “wagon training” by musk oxen, where the seemingly group behavior turns out to be simply a statistical summation of individuals acting out of self-interest. Sober notes that Williams’s book of 1966 repeatedly deploys this pattern of reasoning, and Sober wishes to break out of it, seeking new ground from which to explore possibilities of group selection. His well-reasoned response signals an opening for explorations that he has published over several decades, well into the twenty-first century, but also reveals the power of Williams’s argument, which after four decades still engages the critic by putting him inside the stories or parables Williams composed.30 Though he was notoriously laconic in person, writing became the medium of Williams’s careful thought, as his supporters and critics have observed over and over. His style was shaped by reflection and response. Evolutionary theory must be written, though not always in a parsimonious style. No doubt one can learn and express evolutionary theory through other media, some mathematically rigorous and others—like

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YouTube—popular, but writing is foundational to Williams’s thinking. Indeed, Sober’s critique of Williams dramatizes the extent to which the respondent must enter the writing, and especially the narratives or stories, and attempt to rewrite them from within.

What Is Fitness? (If Not Design for Survival?) As Sober’s responses demonstrate, Williams does not merely take aim at group selection, but also seeks a clearer way to see adaptation. Along the way he clarifies what biologists mean—or ought to mean—by many common terms. Chief among them is fitness—a term not so much along the path of thinking as a vestibule or portal to it. From the perspective of the scientist, fitness suggests a goal of adaptation. Adaptations are, by definition, functions that promote fitness. Williams does not mean only that “natural selection commonly produces fitness in the vernacular sense,” which people generally understand as “mechanisms leading to an increase in health and comfort and a decrease in danger to life and limb,” because from an evolutionary perspective, reproductive survival is the most important aspect of fitness.31 Though fitness is never absolute, adaptation must be understood as a dynamic process. As Williams had previously written, any individual entering a population may be said to have a “reproductive probability distribution” that is zero before maturity, reaches a peak at some point in the life course, and then declines during the descent toward death.32 Behind Williams’s definition of fitness as reproductive probability is Fisher’s pronouncement, sometimes called “Fisher’s Fundamental Theorem,” namely that “the rate of increase in fitness of any organism at any time is equal to its genetic variance in fitness at that time,” which foregrounds the genetic possibilities the population contains.33 So fitness can be measured by the genetic diversity in a population, not by individual adaptations. Expressed in Fisher’s quasi-mathematic terms, “The rate of increase in the mean fitness of any organism at any time ascribable to natural selection acting through changes in gene frequencies is exactly equal to its genetic variance in fitness at that time.”34 Commentators agree that Fisher’s theorem does not mean that the average fitness of a population will increase. Nor is it clear that this theorem is as important as Fisher seemed to think; he hoped it was a law that might hold a supreme position in the biological sciences. If fitness as reproductive power has a trajectory, the idea of fitness also has a history.

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Function, Mechanism, Instinct To understand why recognizing adaptation and testing claims about it is an onerous task, one must understand what Williams means by function, mechanism, and—to the extent that he was influenced by Tinbergen— instinct. To repeat, an adaptation is a mechanism that has a function. The process of natural selection produces it. As one commentary puts it, because “definitions of adaptation are often confounded with definitions of natural selection, rendering them somewhat circular,” Williams introduced a definition that avoids such tautology; he developed a strategy for testing adaptive claims as against chance as an alternative explanation for complex design.35 Speaking of a mechanism implies goal, function, and purpose, as Williams explained, and “will imply that the machinery involved was fashioned by selection for the goal attributed to it.” Here is his apple analogy: “I would say that reproduction and dispersal are the goals or functions or purposes of apples and that the apple is a means or mechanism by which such goals are realized by apple trees.”36 As for words appropriate to merely fortuitous relationships, he prefers the terms cause and effect. In 1992, Williams distinguished mechanism from vitalism, dismissing vitalism’s doctrine that functions of living organisms are due to some vital principle, distinct from biochemical reactions, that would make processes of life inexplicable by the laws of physics and chemistry alone. It must have been disappointing to him that it would be necessary to do so at so late a date. Because mechanism implies that only physicochemical processes are at work in an organism, every vital function is performed by material machinery and in principle can be understood by investigating physical and chemical processes.37 The modern view is that “vitalism now has no credibility.”38 But discrediting it seems a lifelong task. Ernst Mayr, for example, finds in 1982 that vitalism “virtually leaves the realm of science by falling back on an unknown and presumably unknowable factor.”39 Yet this was not a dead issue in 1966, nor is it now. In 1966, Williams argues that mechanism cannot be incidental. It must lead to a specific function: “Any biological mechanism produces at least one effect that can properly be called its goal: vision for the eye or reproduction and dispersal for the apple.” Yet in the 1996 Preface to Adaptation and Natural Selection, he reminds the reader that “in evolution it is easier to lose elaborate mechanisms … than to acquire them.”40 According to any dictionary, we use the word mechanism for a mechanical appliance, an

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arrangement of connected parts in a machine, or a system of parts that operate or interact like those of a machine in a physical or mental instrument or process by which something is done or comes into being. In biology, mechanism refers to the involuntary and consistent responses of an organism to a given stimulus; in psychology, to a usually unconscious mental and emotional pattern that shapes behavior in a given situation or environment; and finally in philosophy, to the doctrine that all natural phenomena are explicable by material causes and mechanical principles. Williams means all of the above. The dictionary dramatizes how mechanism escalates in our language. Mechanism is itself an analogy, as Williams is well aware: “A frequently helpful but not infallible rule is to recognize adaptation in organic systems that show a clear analogy with human implements.” Used in this way, analogies may help a physiologist create hypotheses. Even when the purpose of a mechanism may not be initially apparent, seeking its goal motivates further study. One must examine the machinery to decide about its purpose in terms of means and ends, and this decision “cannot be based on value judgments of actual or probable consequences.”41

Explanations: Organic and Biotic, Proximate and Ultimate Because it is unlikely that mechanisms apply to groups, Williams distinguishes between what he calls organic and the biotic explanations. Organic adaptations “function to maximize the genetic survival of individuals,” while biotic adaptations “would be designed to perpetuate a population or more inclusive group.” By making a sharp distinction between the organic and the biotic, he confronts this rhetorical situation by careful redefinition with a structure and strategy that looks like this in retrospect: (1) “adaptation was pervasive in biology, essentially defining the subject,” (2) “natural selection could explain all examples of adaptation,” and (3) “adaptations, with few exceptions, were the properties of individual organisms and not groups thereof.”42 This is the shape of his program. A concept that William Hamilton introduced in 1964 intervenes in this distinction. Hamilton’s own definition of “inclusive fitness” is elaborate and entangled, with roots, as I mentioned, in the 1957 article by Doris and George Williams and their idea of “social donors.” Hamilton’s “inclusive fitness” measures an individual’s relative genetic representation in the

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gene pool of the next generation, including the contributions of kin. In 1964, Hamilton defined his term in nearly impenetrable prose: “Inclusive fitness may be imagined as the personal fitness which an individual actually expresses in its production of adult offspring as it becomes after it has been first stripped and then augmented in a certain way.”43 What follows in his writing is even more tortuous. One can see why, in 1992, Williams uses a diagram of pedigrees to illustrate the concept of inclusive fitness.44 In 1966, Williams writes that an organic adaptation, as a mechanism designed to promote the success of an individual organism, can be measured by the extent to which it contributes genes to later generations of the population. Citing Hamilton, he says that “it has the individual’s inclusive fitness … as its goal.” In contrast, a biotic adaptation must be defined as “a mechanism designed to promote the success of a biota, as measured by the lapse of time to extinction.”45 Many errors await many thinkers. It is clearly possible to ascend too far in time and space and in levels of thinking, ascribing functions so that everything becomes, at least potentially, an adaptation. It is also possible to confuse levels of thinking. For instance, biologists commonly assume that mechanisms relating to respiration, nutrition, and so on are likely organic adaptations promoting individual survival, but reproductive processes are often imagined to relate to the survival of the species.46 Williams finds the second unlikely, but in 1966 he has not yet worked out the details. Mistakes are not only frequent in the perceptions of biologists but can also be observed in the actual performance of sexual functions. Consider that when parents aid offspring—that is, engage in behavioral mechanisms—“there will inevitably be times when such aid is provided ‘by mistake’ to unrelated individuals.” Hybrids, too, exemplify “malfunctioning of the mechanisms of reproduction.”47 Observers can err, but so can biological entities. Consequently, one must always remember “the unquestionable principle that natural selection can often produce mechanisms of extreme precision, but never of perfection.” All mechanisms, including those of behavior, are surely subject to occasional malfunction and often have less-than-perfect systems of timing. Finally, death is neither a mechanism nor an evolved adaptation.48 Given all these pitfalls, Williams asks how one is to ultimately ascertain the function of a biological mechanism. “In this book I have assumed, as is customary, that functional design is something that can be intuitively comprehended by an investigator and convincingly communicated to

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others.” Intuition may be a poor tool, and Williams stresses that, for any biological mechanism, there are no established principles and procedures for answering the question, “What is its function?” Yet generally speaking, “We do not need weighty abstractions to help us decide that the eye is a visual mechanism.” He is not about to discard the many helpful parallels between natural and artificial mechanisms, because they are “so convenient as to be inevitable.”49 It is easy to see how one can get from Williams’s framework to Noam Chomsky’s expression “language organ” and Steven Pinker’s “language instinct.” For Williams’s followers, like Randy Thornhill, a zoologist at the University of New Mexico, there may or may not be general-purpose adaptations, but following Williams, his followers prefer to focus on the adaptive specialization of specific mechanisms. Arguments continue about the distinction between the terms mechanism and instinct, especially over what Darwin regarded as the instinct for sociability. The terms instinct and mechanism consequently overlap in the discussions of biologists. Tinbergen describes instinctive or genetically preprogrammed patterns of behavior as mechanisms—sequences of events that include a stimulus or releaser, an innate releasing mechanism, and a fixed-action pattern.50 Instinct is efficient. For instance, as Williams argues, for any animal, human beings included, “fear of the edges of precipices would be universally adaptive,” and consequently “we can expect such fear to be instinctive, rather than learned.” For any organism, instinct is more efficient than learning. Though there are things that have to be learned, such as the individual characteristics of a particular mate or the location of a nest, Williams writes that natural selection prefers instinct: “All elements that can be instinctive … will be instinctive. Instinct costs less than learned behavior, in the currency of genetic information.”51 Williams needs a principle for recognizing a hierarchy of adaptations, or at least a way of specifying the subordination of one function to another, and he thinks such a system could follow Tinbergen, who classified instincts on the basis of the nature and generality of the purposes served. Williams prefers to imagine these classes as beginning with adaptations so basic that they are found in all organisms. Consider that each organism has mechanisms for its own nutrition, for processes relating to growth, differentiation, reproduction, and the completion of the life cycle. Every organism also has defense mechanisms.52 The plain title of Adaptation and Natural Selection is deliberately understated, not the result of any lack of imagination. So too Williams

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engages in remarkably understated yet arresting dismissals. Though the idea of species may be a central taxonomic and evolutionary concept, it “has no special significance for the study of adaptation. It is not an adapted unit and there are no mechanisms that function for the survival of the species.”53 Though he certainly discusses the characteristics of species, he does not use phrases like “benefit to the species” or “benefit of the species” in 1966 and speaks negatively of these kinds of functions in his retrospective preface written in 1996. Randolph Nesse uses language borrowed from Mayr and Tinbergen to summarize Williams’s schemes of adaptation and show how they are relevant to medicine: “The framework grows from the fundamental principle that all biological traits need two kinds of explanation, both proximate and evolutionary.” Though Williams observes proximate explanations, his focus is on the more universal mechanisms that evolution seems to have allowed.54

Names and Structures in Adaptation and Natural Selection The annotated table of contents of Adaptation and Natural Selection takes up three and a half pages, using a unique design with each chapter abstracted into three or four pithy sentences. None of these sentences are questions, though a question embedded in the abstract of the final chapter is “What is its function?” where “it” is “biological mechanism.” The contents, chapter by chapter, reveal the process of reduction, moving from adaptation and the naming of functions in Chap. 1—when should a scientist call something a “function”? when it is “produced by design and not by chance”—to questions of selection, how it works, how adaptations are maintained, and questions of progress or the lack thereof in selection in Chap. 2. Chapter 3 concerns larger issues of ecology and development, Chap. 4 the problems with the concept of “group selection,” Chap. 5 the mechanisms of reproduction, sexual and asexual, Chap. 6 the relationship between reproduction and behavior of organisms, and Chap. 7 matters of behavior, generally. Chapter 8 is a classical refutation placed in the traditional order for an oration, before the concluding Chap. 9, and it offers a solution for the problem that “there are no established principles and procedures for answering the question, ‘What is its function?’” As the table of contents concludes, the entire endeavor seeks “a suitable name for this

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special field of study,” because names are necessary “for the study of adaptation as a general principle.”55 Stearns finds that the book is broken into two parts, the first four chapters constituting “a powerful critique of group selection and group-level adaptation with lasting impact on evolutionary biology,” and the second part, Chaps. 5 through 9, treating “adaptations of the genetic system (e.g., mutation), reproductive physiology and behavior (e.g., reproductive rate), social systems (especially cooperation), and others such as senescence, multicellularity, the self-regulation of population size, and the supposed integration of communities or ecosystems.”56 I will argue that Chaps. 5 through 8 constitute a derivation of sociobiology from the ground up. After Williams confronts his reader by naming biological entities or processes such as function and mechanism and distinguishing between organic and biotic adaptations, he turns to ground rules or doctrines about the use of terms like progress, specialized, advanced, higher. He is not entirely prescriptive. Yet many of these terms, like progress or advance, slip toward describing the rhetoric of biologists and their arguments, as in the “advancement of knowledge.”57 Names call for care because Williams wishes to establish “adaptive mechanisms worthy of the name,” and he knows that “attempts to demonstrate the benevolence of Nature often take the form of name changing.” He reminds the reader that a generation of social Darwinists saw “nature red in tooth and claw” in the killing of deer by mountain lions; yet for a more recent generation this killing “has become Nature’s kindness in preventing deer from becoming so numerous that they die of starvation or disease.” Williams concludes: “The simple facts are that both predation and starvation are painful prospects for deer, and that the lion’s lot is no more enviable.”58 One should never forget that biologists like Williams work under the generalizing name of natural selection. For instance, he finds that when biologists study population size and density, the ratios of age groups, and the rates of birth and death, they employ the same body of concepts and problems “whether the data derive from human, biological, or hypothetical populations.” He must name as he goes along because, though it is not always efficacious to use the same terms for human and non-human species, he finds demography appropriate for all.59 At the center of Williams’s inquiry is the name design, with the history of ideas bundled into this term, especially the “argument from design” made by William Paley (1743–1805). What design means is more often

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assumed then established, as Williams writes, and is often in the eyes of the beholder. For instance, a frequently cited example of design is the vertebrate—or human—eye. But is it well designed? “For the same reason that it was once effective in the theological ‘argument from design,’ the structure of the vertebrate eye can be used as a dramatic illustration of biological adaptation and the necessity for believing that natural selection for effective vision must have operated throughout the history of the group.”60 Later in his career, Williams will argue that the vertebrate eye is “stupidly designed,” and as a result “embodies many functionally arbitrary or maladaptive features,” and people suffer over his view.61 But more to the point, the design of the vertebrate eye—and the discussion of its design— is historically constrained, and the structure of the eye and discourse about it reveals its evolutionary and cultural history. History is a messy, contingent one-way street—though contingency plays less of a role in this, Williams’s first book, than it will later in his career. So how can this 40-year-old biologist—or evolutionary theorist—use the term design without falling into the argument from design? Once he has decided to “use terms appropriate to human artifice and conscious design,” though it may be easy in practice to intuit functional design, he recognizes disagreements about whether “certain effects are produced by design or merely as by-products of some other function.” It is hard to prescribe objective criteria, so Williams relies on informal arguments, asking “whether a presumed function is served with sufficient precision, economy, efficiency, etc. to rule out pure chance.” One informal mode of argument is storytelling. Williams tells a story about how a fox heading for the hen house right after a heavy snowfall has initial difficulties, but after this the furrow the fox made the first time will make the trip easier. Because a path through the snow saves time and food energy used in subsequent journeys, “such savings may be crucial for survival. Should we therefore regard the paws of a fox as a mechanism for constructing paths through snow?”62 We shall not! Williams’s fox is unlike Thoreau’s human in Walden: for the latter, the surface of the earth is soft and impressible, so the paths that the mind travels result in deep ruts of tradition and conformity. Fox paws were designed before the advent of henhouses or trails to them. His propensity for this kind of storytelling would earn Williams the name “adaptationist,” a complimentary name for some biologists and an insult for others—but stories like this are meant to remind a reader that everything is not an adaptation. Randolph Nesse, Williams’s last

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collaborator, described what he calls “a cognitive anomaly” in the “Williams Vision” “that made it very difficult for him to blind himself to evidence that contradicted his ideas.”63 He could not stay in the furrows or ruts. There are consequences for those with this kind of vision. Nesse mixes his metaphors a bit, calling what Williams writes “the way” or “route,” but certainly the Williams vision “lacks the pleasures of seeing the world as a wondrous whole and calls attention to the dark side of nature.”64 Perhaps humorously, Nesse plays on Williams’s own aim, to destroy conceptions of optimization or “naive group selection.” Nesse writes that the “‘Williams Vision’ may not yield a net benefit to the possessor, but it is invaluable for the species.” Nesse dearly wants to keep Williams out of the highly publicized battle of words between adaptationists and anti-adaptationists—a battle that Williams himself entered infrequently, cautiously, and with restraint. “If anything, he is a maladaptationist,” Nesse writes. “He has spent his entire career preoccupied with one apparent maladaptation after another.”65 Stories change over time; and as Williams reflects on his own, he changes his evolutionary explanations. These differences matter. Strangely enough—or maybe not!—this person who changed the evolutionary story most for the generation that read biology in the late 1960s was himself a kind of changeling. The story of those changes is interesting in itself, and perhaps may provide a kind of object lesson revealing the rewards and dangers of reading and rereading and writing and rewriting evolutionary theory. Williams’s 1966 introduction to Adaptation and Natural Selection claims: “This book is based on the assumption that the laws of physical science plus natural selection can furnish a complete explanation for any biological phenomenon, and that these principles can explain adaptation in general and in the abstract and any particular example of an adaptation.”66 By 1985, he includes a third variable—history with its contingencies. Devils will emerge from these historical details.

Levels of Selection: Genes, the Concrete, and the Abstract Williams opens his discussion with what evolutionary biologists call “levels of selection” because adaptations, with few exceptions, must be properties of individual organisms and not groups thereof. Yet he also asks himself repeatedly what a group or herd or mob is, and how it operates. To do so

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is to turn a critical eye on adaptations that appear to be social. If he does not approve loose talk of “group selection,” what mode of exploration or explanation does he approve for the naming of groups of organisms? As an analogy, consider schools of fish, for instance—keeping in mind that there are also schools of thought. Williams refines his article of 1964 on the schooling of fish for Adaptation and Natural Selection, creating a parable about groups and individuals and about the perspective of the scientific observer: A school in its typical development is a striking phenomenon. It is remarkable for its compactness and the way it moves about as a single unit. As with a well-prepared military drill, it is the regularity and precision of the group that excites interest, not the behavior of the individual participants. If one does look at individuals, one finds that they are all of the same species and of almost identical size, and that each fish always swims with very nearly the same speed and direction as its immediate neighbors. It is easy to lose interest in such individuals; if you have seen one you have seen them all. It is the school as a whole that persistently excites our curiosity. I would maintain, however, that an understanding of schooling can be achieved only by counteracting this intuitive reaction. One should concentrate first on the individual and seek an understanding of the adaptive aspects of its behavior. If this inquiry is successful, one can then ask how many of the phenomena of the school can be explained as simply the statistical summation of individual adaptations.67

Schooling allows Williams to illustrate exactly the difference between an effect and a function. Schooling is an effect, but not a function. His parable is also a paradigm that favors the statistical—the summing up of individual behaviors—over intuition about groups. He does not include much math or do much calculating in his books, but arguments like this one accumulate, as incremental additions that create a kind of statistical summation. Every close look at a group or herd reveals, using his codical phrase as a conclusion, that a certain behavior “can be explained as simply the statistical summation of individual adaptations.” He uses this phrase six times in his seventh chapter. This kind of rhetorical emphasis is salient in Williams as a writer, but apparently not observable in Williams as a talker. Though he appeared to some notoriously taciturn, reserved, aloof, distant, and perhaps shy, he was, as Sarah Blaffer Hrdy observes, “extraordinarily generous in giving credit to others, he was also exceedingly supportive and warm to graduate

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students and younger colleagues,” and “along with this warmth there was also a kind of quiet impishness and intellectual playfulness.”68 He listened, maybe mulled over what he heard, allowed his ideas to build up, and then released his responses, increasingly in books rather than articles. Later, in the 1990s, when famous, he was interviewed repeatedly. Even in the transcripts of those interviews, it is at least partially apparent that he continued to have an extended conversation with himself about what he thought of what he had written. He would be known throughout his career for this first book, which advanced a gene-centric view of evolution and created the “evolutionary concept of the gene.” Genes have received many definitions, some very general, as in the phrase Mark Ridley borrows for his textbook: “that which segregates and recombines with appreciable frequency.”69 But Williams can also take credit for developing the abstract, cybernetic definition of the gene. His understated warning about relations between abstractions and concrete entities notes that “in its ultimate essence the theory of natural selection deals with a cybernetic abstraction, the gene, and a statistical abstraction, mean phenotypic fitness.” That kind of theorizing appeals to those capable of using cybernetics and statistics. However, as he playfully reminds, “Fruitful applications of this theory will also require a detailed knowledge of biology.”70 In later writings, Williams would continue to remind biologists of the relationships between the abstractions they use and the concrete entities they study. As the wit of the previous extract suggests, though Williams entered the public eye as a critic and an iconoclast, he was always a professional collaborator, and increasingly so over the course of his career. As such, he became a kind of “interior model” of professional style and perspective within biological discourse, particularly because his approach is also accessible to non-biologists. This makes his writings a vestibule into the idea of what clear thinking about natural selection was, especially in the period that followed the modern synthesis.71 But in the 1950s and early 1960s, he worked alone, or sometimes with Doris, his wife.

Evolutionary Style as Stimulus and Artifice A stimulus is, according to the Oxford English Dictionary, “Something that acts as a ‘goad’ or ‘spur’ to a languid bodily organ; an agency or influence that stimulates, increases, or quickens organic activity.” One purpose for writing—I would say its highest purpose—is to stimulate the brain,

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that languid bodily organ, through the use of language, and incite people to think outside their ruts. Williams goads and challenges the very structure of means, ends, and causality as practiced by evolutionary scientists. I use style to describe his strategy for the undertaking. A style is an author’s approach to his subject through the medium of language. In literary terms, “Style combines two elements: the idea to be expressed and the individuality of the author. From the point of view of style, it is impossible to change diction or syntax and say exactly the same thing; for what the reader receives from a statement is not only what is said, but certain connotations that affect the consciousness.”72 As a literary critic and environmental historian, I have wondered about the style of the major works of the modern synthesis by Mayr, Stebbins, Dobzhansky, and Simpson. I am considering here the style of the work that comes after theirs—that of Williams. I prejudice my own case somewhat by assuming that all of them are authors of literary texts, that these authors use a sophisticated rhetoric, that their texts are designed to teach others how to engage in “population thinking” and also model the proper use of language for the task. As to the individuality of the author, since evolutionary studies have interested themselves in self-expression, self-­ deception, and consciousness as defined in terms of self, the matter of style is recursive. Who is the self that speaks of the evolution of self? If style is personal, how can it relate to the impersonal features of evolutionary processes? Further, the idea of the writer as narrator, storyteller, and creator of narrative structure must be squared with processes of evolution that stimulate writing—processes that seem distinctly not structured like human narratives since they seem to be without a genre or plot, like tragedy or comedy, as humanists traditionally understand them. This is not to say that a story of how evolutionary tales are told lacks a plot. As I will note along the way, writings by George C. Williams are also frequently allusive, not simply to the works of other scientists or classic writers like Paley, Galen, and T.  H. Huxley, but also to literary works, mythology, and detective fiction on occasion. Williams was not only a prolific reviewer of books; he was also known to devour histories and other humanistic literature. He studied Icelandic and was said to have written an Icelandic saga. It is highly unlikely that the literary allusions found in his works are unintentional, but even more interestingly, they sometimes are ironic and even double-edged. It is generally assumed that people read science as nonfiction, the purpose being to receive truth. I assume that the purpose of reading

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evolutionary theory is to see how scientists think and express themselves, partly so I can understand what kind of intellectual world they inhabit. I write as a reader. You will not receive a definitive view of evolutionary theory from what I write, but you will learn how George C.  Williams wrestled with expressing problems he considered important—how, through language, he attempted coherent explorations of these problems. In any case, Williams achieved a mature style in Adaptation and Natural Selection (1966). Generally, it is reasonable to think of this book as cultivating Richard Lanham’s “three central values” of the plain style: clarity, brevity, and sincerity.73 Williams sometimes hedges the last, as we shall see. To achieve its aim, Lanham argues, the plain style ought to be maximally transparent and minimally self-conscious. In the interest of transparency, Williams consistently proceeds by questioning assumptions and making assumptions, as in “The unwarranted assumption of organic evolution as an explanation for biotic evolution dates at least from Darwin,” a passage that no doubt requires an emoji.74

Notes 1. George C.  Williams, Adaptation and Natural Selection (Princeton, NJ: Princeton University Press, 1966), 25, 253. 2. Ernst Mayr, The Growth of Biological Thought (Cambridge, MA: Harvard University Press, 1982), 483. 3. Williams, Adaptation, 11, 22, 33. 4. Williams, Adaptation, 4. 5. Williams, Adaptation, 10. 6. Williams, Adaptation, 50, 51. Chapter IV, “Natural Selection; or the Survival of the Fittest,” of the definitive edition of Charles Darwin, The Origin of Species by Means of Natural Selection or The Preservation of Favoured Races in The Struggle for Life, sixth edition (London: John Murray, 1873), deals with “advance in organization” (62–105) and elaborates the analogy of the “tree of life” (104–5). 7. Williams, Adaptation, 245. 8. Williams, Adaptation, 253–4. For Emerson and the history of group selection, see Chapter 6 of Gregg Mitman, The State of Nature: Ecology, Community, and American Social Thought, 1900–1950 (Chicago: University of Chicago Press, 1992), 110–45. 9. Williams, Adaptation, 255. 10. Marc D. Hauser, The Evolution of Communication (Cambridge, MA: MIT Press, 1997), 9–10.

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11. Williams, Adaptation, 76. 12. Williams, Adaptation, 83, 193. 13. Williams, Adaptation, 90, 91, 213, 214. 14. George C. Williams, Natural Selection: Domains, Levels, Challenges (New York: Oxford University Press, 1992), 150. 15. Williams, Adaptation, 9, 159. 16. Williams, Adaptation, v. 17. Stephen C.  Stearns, “George Christopher Williams, 1926–2010” (Washington, DC: National Academy of Sciences, 2011), 11–12. http:// www.nasonline.org/publications/biographical-­memoirs/alphabetical-­ listing/memoirs-­w.html. 18. George C.  Williams, “Preface (1996),” in Adaptation and Natural Selection. Reprint. Princeton Science Library Edition. (Princeton, NJ: Princeton University Press, 1996), ix. 19. Williams, “Preface,” ix, x. 20. George C. Williams, “Introduction,” Group Selection, edited by Williams (Chicago: Aldine-Altherton, 1971), 6. According to Oren Solomon Harman, what Williams needed to kill group selection he “found buried in the pages of [Fisher’s] The Genetical Theory of Natural Selection” (Harman, The Price of Altruism: George Price and the Search for the Origins of Kindness [New York: Vintage Books, 2011], 170). See Williams, Adaptation, 21. Fisher presents a dialectic that assures the symmetry of sex ratios, and Harman believes that Fisher’s discussion involves Fisher unknowingly in game theory. There are many kinds of games and not all the games biologists conceived came out of John von Neumann’s game theory. 21. Williams, “Introduction,” Group Selection (Chicago: Aldine-Atherton, 1971), 6,10. Paul Ehrlich, The Population Bomb (New York: Ballantine Books, 1968). 22. In Letters to the Editors, American Naturalist 100, no. 916 (Nov.–Dec., 1966): 687–90. 23. Williams, “Preface,” ix. 24. Williams, Adaptation, 19. 25. GCW, 1966a, 11, 12. 26. Williams, Adaptation, 18–19. 27. Ludwig Wittgenstein, Tractatus Logico-Philosophicus, translated by C.  K. Ogden (1922), parallel edition including the German text on the ­facing page to the English text, prepared with assistance from G. E. Moore, F.  P. Ramsey, and Wittgenstein (Routledge & Kegan Paul, 1981), 5.47321, 6.363. 28. Williams, Adaptation, 230. 29. Elliott Sober, “Let’s Razor Ockham’s Razor,” Chapter 7 of From a Biological Point of View, 136–157 (Cambridge, UK: Cambridge University Press, 1994), 142–3. D. S. Wilson believes that parsimony is

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the true Williams’s principle, that “Adaptation at a level requires that there was selection at that level.” Speaking for himself and Elliott Sober in a blog post, he writes, “In our opinion, this is the most enduring part of George’s legacy, which should be taught to everyone learning about evolution. George earned a permanent place in history not for showing that group-level adaptations never evolve, but for showing how to make the judgment call.” http://scienceblogs.com/evolution/2011/01/ homage_to_george_williams_and.php. 30. Sober, “Let’s Razor,” 144. 31. Williams, Adaptation, 26. 32. Williams, “Pleiotropy, Natural Selection, and the Evolution of Senescence,” Evolution 11 (1957): 410. 33. R.  A. Fisher, The Genetical Theory of Natural Selection (Oxford: Oxford University Press, 1930), 35. 34. See Anthony W.  F. Edwards, “The Fundamental Theorem of Natural Selection,” Biological Reviews 69, no. 4. (1994): 443–74. 35. P.  T. Ellison and G.  Jasienska, “Constraint, Pathology, and Adaptation: How Can We Tell Them Apart?” American Journal of Human Biology 19 (2007): 622–30, 622. 36. Williams, Adaptation, 9. 37. Williams, Natural Selection, 3. 38. “This is sometimes credited to the view that vitalism posits an unknowable factor in explaining life; and further, vitalism is often viewed as unfalsifiable, and therefore a pernicious metaphysical doctrine,” as Robert C. Richardson and William Bechtel put it in “Vitalism,” in The Routledge Encyclopedia of Philosophy, edited by E. Craig (London: Routledge, 1998), http://mechanism.ucsd.edu/teaching/philbio/vitalism.htm. See also Hilde Hein, “The Endurance of the Mechanism-Vitalism Controversy,” Journal of the History of Biology 5, no. 1 (Spring 1972): 159–88. 39. Mayr, Growth of Biological Thought, 52. 40. Williams, Adaptation, 8; Williams, “Preface,” xiii. 41. Williams, Adaptation, 12. 42. Williams, “Preface,” vi, x, ix. 43. W.  D. Hamilton, “The Genetical Evolution of Social Behaviour. I,” Journal of Theoretical Biology 7, no. 1 (July 1964): 1–16, 8. 44. Williams, Natural Selection, 19. 45. Williams, Adaptation, 96, 97. 46. Williams, Adaptation, 160. 47. Williams, Adaptation, 193, 205. 48. Williams, Adaptation, 139, 145–46, 208, 225–26. 49. Williams, Adaptation, 260–61. 50. Niko Tinbergen, The Study of Instinct (London: Oxford, 1951).

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51. Williams, Adaptation, 83. 52. Williams, Adaptation, 263. 53. Williams, Adaptation, 252. 54. Niko Tinbergen, “On Aims and Methods of Ethology,” Zeitschrift für Tierpsychologie 20 (1963): 410–33; Ernst Mayr, “Cause and Effect in Biology,” Science, New Series, 134, no. 3489 (Nov. 10, 1961): 1501–6; Randolph M. Nesse, “Evolution: Medicine’s Most Basic Science,” Lancet 372 (2008): S21–27S, S27. But see Kevin N. Laland et al., “Cause and Effect in Biology Revisited: Is Mayr’s Proximate-Ultimate Dichotomy Still Useful?” Science 334 (2011): 1512–16. 55. Williams, Adaptation, viii. 56. Stearns, “George Christopher Williams, 1926–2010,” 10, 11. 57. Williams, Adaptation, 48, 49, 14, 15. 58. Williams, Adaptation, 119, 255. 59. Williams, Adaptation, 66. 60. Williams, Adaptation, 6. 61. Williams, Natural Selection, 73. 62. Williams, Adaptation, 9–10, 12–13. 63. Randolph M.  Nesse, “George C.  Williams, 1926–2010” (September 2010), http://skepticaladaptationist.com/2010/09/george-­williams-­ 1926-­2010.html. 64. Randolph M. Nesse, “Maladaptation and Natural Selection,” the Quarterly Review of Biology 80, no. 1 (2005): 62–70, 64. 65. Nesse, “Maladaption,” 62, 64. 66. Williams, Adaptation, 6. 67. Williams, Adaptation, 212–13. 68. Personal communication. 69. Mark Ridley, Evolution (Malden, MA: Blackwell Publishing, 2004), 308. 70. Williams, Adaptation, 33. 71. For his collaboration, see, for instance, Sarah Hrdy and G.  C. Williams, “Behavioral Biology and the Double Standard,” in Social Behavior of Female Vertebrates, edited by S. Wasser (New York: Academic Press, 1983), 3–17. Williams also collaborated on long projects with Randolph Nesse and James Paradis. 72. William Harmon and Hugh Homan, A Handbook to Literature, 9th Edition (New Jersey: Prentice Hall, 2003), 491. 73. Richard A. Lanham, Analyzing Prose (New York: Charles Scribner’s Sons, 2003), 1–10. 74. Williams, Adaptation, 99–100.

CHAPTER 4

Opening Sociobiology While Avoiding Aesthetic Distractions

There is no talk of beauty in Adaptation and Natural Selection, or for that matter of any other aesthetic distractions that universally and insidiously posit certain unwarranted “higher order” processes for natural selection and reveal themselves as distorting predispositions for an observer who desires to understand how organisms behave. Instead, Williams expresses hope that this book will “help to purge biology of what I regard as unnecessary distractions.”1 For Williams, higher order and aesthetic are so closely related that both might be classified as names for a version of wishful thinking. When watching the behavior of schools of fish, what strikes an observer as an ensemble does not necessarily lead to accurate perception of the phenomenon. Like others who knew him, Stephen Stearns found Williams “utterly lacking any romanticized view of nature.”2 This is not to say that Williams lacks an aesthetic. For instance, he was extraordinarily fond of the music of Johann Sebastian Bach. He considers it difficult for a biologist to think clearly while admitting romantic conceptions of beauty into discourse, yet what seems rational and bleak has its own aesthetic. It is best to say that while Williams’s plain style for conveying naturalistic explanations and arguments about adaptation and its products lead him to dismiss any digressions toward traditional questions of beauty or ornament, it also reveals its own aesthetic. Whether one aesthetic is better than another becomes a historical question.3

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 M. P. Cohen, George C. Williams and Evolutionary Literacy, Literatures, Cultures, and the Environment, https://doi.org/10.1007/978-3-031-11650-6_4

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The particular aesthetic predisposition Williams calls out is “biotic adaptation,” where organisms play roles that “would subordinate their individual interests for some higher value, as in the often proposed benefit to the species.”4 This is precisely the romantic idea of beauty expressed by Samuel Taylor Coleridge, for whom the beautiful is that in which the many become one. Using the principle of parsimony, Williams inverts this view from the very beginning and finds that those who posit mechanisms for group benefit misinterpret, because “higher levels of selection are impotent and not an appreciable factor in the production and maintenance of adaptation.” Thus he says that the higher-level theories “should be used only as a last resort.”5 There is also an aesthetic to the concept of “progress.” Williams doubts that many biologists “subscribe to the view of evolution as a deterministic progression towards man, but there is widespread belief in some form of aesthetically acceptable progress as an inevitable outcome of organic evolution.” He quietly and perhaps facetiously traces notions of progress to the central institutional conventions of naming and systematics used by biologists: “I suspect that evolutionary progress and the inevitability of man may seem like scientific ideas only because of our heritage of such orthogenetic terms as ‘higher’ or ‘advanced’ organisms and the fact that a list of taxonomic categories has to have a beginning and an end.”6 Whenever biotic adaptation is postulated, he writes, “its immediate or ultimate effect is the improvement of the situation from a traditional aesthetic point of view.”7 All aesthetic perspectives—truth, beauty, goodness—are of a higher order, are likely to be anthropocentric, and are certainly anthropomorphic. Many aesthetic assumptions are ecological, not only because they imply the beauty of diversity, but because they suggest some higher order in the assemblage of organic beings and the notion that some adaptations are better than others. Williams finds these assumptions reprehensible for biological clarity, as he indicates in a stunning catalogue of animal behavior near the end of Adaptation and Natural Selection: It is assumed that: a population of vigorous individuals under heavy predator pressure is better adapted than one that is sickly and chronically starved; a population that divides its resources into stable individual territories is better adapted than one in which there is a chaotic scramble for resources; a population in which territory or social position is held by threat-display and recognition by neighbors is better adapted than one that maintains the

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social structure by frequent combat with effective weapons; a population with stable density, stable age distribution, etc., is better adapted than one in which such factors fluctuate widely; a population with limited fecundity and low juvenile mortality rates is better adapted than one with high fecundity and high juvenile mortality rates; a population in which the old and dominant individuals regularly yield to promising youths is better adapted than one dominated by a stable regime of fecund but slowly displaced oligarchs; populations in which individuals, such as worker bees, often jeopardize their own well-being for a larger cause are better adapted than those whose members consistently act only in their own immediate interests; those in which individuals normally live in peace or active cooperation and mutual aid are better adapted than populations in which open conflict is more in evidence; on the other hand, when active mutual destruction must take place, infanticide is preferable to the killing of peers. I submit that the only consistency found in such propositions is that they all conform to prevailing aesthetic concepts of what organisms ought to be like.8

Beyond these aesthetic distractions are others. Many traits of organisms are not adaptations, but simply incidental byproducts of natural selection, perhaps including even the much-vaunted faculty of language in humans. It has been argued that Williams provides the first critique of broadly attributing all products of evolution to adaptation, often accompanied by “just-so stories.” Because he writes that adaptations must exhibit evidence of design, not chance, the language of adaptation must not be used for certain processes that are not adaptive, like pleiotropy—broadly defined as multiple traits inherited together, through single genes, with sometimes sequential and seemingly unrelated effects.9 Many of these processes pass on non-adaptive features. Sometimes these processes have been named “genetic hitchhiking,” “genetic drift,” or “spandrels”—yet these non-­ adaptive traits are sometimes associated with later development of other adaptations.10

Emotional Restraint While Witnessing Behavior By seeking to avoid aesthetic distraction, the plain style may be most notable for its restraint—for what the speaker will not express. When discussing behavior, Williams is most reticent to speak of motive, mental state, and especially emotions of organisms. Consequently, like Tinbergen, he sees behavioral adaptations from the outside. He follows this style of perception when discussing friendships and animosities exhibited only by

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certain rare and intelligent animals like humans, who live in social groups. “Primitive man lived in a world in which stable interactions of personalities were very much a part of his ecological environment. He had to adjust to this set of ecological factors as well as to any other.” Like Hamilton, he limits discussion of friendship to observable exchanges of favors and reciprocations. As Williams and many others note, Darwin writes of “the ‘lowly motive’ of helping others in the hope of future repayment.” In contrast, Williams sees it as “necessary that help provided to others be occasionally reciprocated if it is to be favored by natural selection. It is not necessary that either the giver or the receiver be aware of this.” Gratitude may or may not be an emotion, but Williams is quite certain that the interactions between hens in a barnyard are “adequately explained without postulating emotional bonds between individuals.” The only mention of conscious emotion in his later book, Natural Selection (1992), is negative: “The tearful eyes of a sea turtle laying eggs on a beach result, not from any emotional impact of motherhood, but from her use of lachrymal glands to secrete a concentrated brine to keep her internal electrolytes at the usual vertebrate level.”11 Plain, of course, is at the center of explain. The prose style of George C. Williams is memorable for both. That is why other biologists discover that his stark narratives and examples have been inscribed in their own memory. A Williams statement often takes this form: “The opposition arises, as Darwin himself observed, not from what reason dictates but from the limits of what the imagination can accept.”12 Such a sentence uses grammatical parallelism, applies historical depth to the situation, and moves from reason to imagination. Generally, when he uses such rhetorical devices, they are not just rhetorical devices. As a sidelight, one notices that books by Williams are visually austere, never supplemented by spectacular illustrations or sidebars, and rarely use other visual aids. Yet the plain style of Williams is strangely, or perhaps naturally, also reminiscent of the Western biblical style. The Bible is, after all, made of words and only words. A capacity for elegant understatement may explain why Richard Dawkins remembers that Williams reminded people of Abraham Lincoln: “George was a tall, shy, diffident scholar, rather like W D Hamilton in character, his legendary wisdom seeming somehow enhanced by a physical resemblance to Abraham Lincoln.” Perhaps Dawkins meant to notice not just his physical appearance, but also a distinctive style of expression. “One must remember that style is not just a constellation of aesthetic properties

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but is the vehicle of a particular vision of reality,” Robert Alter writes in Pen of Iron. Alter also notes that stylistic plainness “is a quality that Lincoln labored to perfect over time, especially against a background of American oratory that favored highly wrought ornamentation.” So too one might argue with regard to Williams. Also, as Harold Bloom has claimed, “A test for great poetry and prose is an aura of inevitability in the phrasing.”13 Inevitability is a sense that comes from witnessing precision of statement— this is why biologists refer not only to Williams’s ideas, but to the forceful language in which they are expressed. Consider this sentence: “Whatever the gene may be, it is part of the mundane world and therefore not perfect, in its self-duplication or in any other features.” Here we have a version of Williams’s loose style that warns against tight connections. Then there is the balanced style: “Selection has nothing to do with what is necessary or unnecessary, or what is adequate or inadequate, for continued survival.” The tenor of Williams’s sentences is almost always cautionary, as in “I would regard it as a mistake to say that anything is a basic characteristic of life. In an organism we see only the basic characteristics of matter and the results of billions of years of adjustment to a changing environment.” Such cautions frequently include rhetorical questions: “Bird songs function as aids in the maintenance of territory, but what is the function of territory?”14 Williams’s questions—and he asks a lot of them—are nearly biblical, as if they came from some Old Testament—and one might say he writes about the oldest testament. The first and foremost question is “what is a function?” or “what is its function?” When one thinks of biblical language, as in the King James Bible, one thinks of the Lord answering Job from out of the whirlwind: Who provideth for the raven his food? when his young ones cry unto God, they wander for lack of meat. (Job 38:41) Knowest thou the time when the wild goats of the rock bring forth? or canst thou mark when the hinds do calve? (Job 39:1)

“Question” and “can” are words that appear copiously in Williams’s texts: “There is no reason to believe that a hen can harbor grudges against or feel friendship toward another hen. Certainly the repayment of favors would be out of the question.”15 He focuses on what is and avoids speaking of what ought to be. The word “ought” appears only once in Adaptation and Natural Selection, and then in a pejorative way, as in the

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introduction to the previous catalogue. What “ought to be” is associated with the idea of progress and imports an unwarranted assumption of teleology, an assumption inherited from the elaborate plots and metaphors of Darwin. But can biologists live without the rhetorical style that made teleology so powerful?

From Chemistry to Behavior: Design and the Grounds for Sociobiology Sociobiology always grounds itself in reproductive behavior, though it is often taken to be the generalized biology of social relations. The intersection of sexual behavior and culture is culturally dangerous ground. In biology restraint is necessary to rid discussions of teleology and aesthetic preconceptions. Williams builds a structure of thought in Chaps. 5 through 7 of Adaptation and Natural Selection, deriving and elaborating a set of principles of sociobiology, from cellular process of reproduction through sexual and social behavior. By ascending this ladder of reason, he posits theoretical principles of sociobiology from the ground up. His argument is ordered in a straightforward manner, tightly reasoned by means and ends—with no references to Malthus, Darwin, or any nineteenth-­ century authorities. As he later indicates, his work launched from that of Fischer, Haldane, and Wright and is mediated by that of David Lack and William Hamilton, who made editorial changes to Chaps. 6 and 7 of the book. Williams would find his materialist and economic argument confirmed in later writings of Hamilton, Ghiselin, Maynard Smith, Trivers, and others.16 Although his argument progresses, it is not an argument about progress but about consequences. I find it best to follow his ascent and leave out many details, including most of his sources, but try to use as much of his language as possible to reveal salient points of his argument’s structure, emphasizing his light and way. When treated as “design for survival,” behavior must “make it likely to contribute a more than average number of genes to future generations.” So Chap. 6, “Reproductive Physiology and Behavior,” announces Adaptation and Natural Selection’s second half. The general rule remains that fitness of any group depends upon “summation of the adaptations of its members.”17 Williams believed that, as W. D. Hamilton put it, the characteristic behavior of a species is “just as much the product of evolution as the morphology.”18 This major thesis shapes the design of Williams’s later

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chapters and is meant to be more than a summation. And as his preface indicates, the text is directed toward advanced students and biologists, not a group of specialists.19 So, to borrow from Williams, the book is meant to have “effective design for reproductive survival” in more than the sense of publication of multiple copies. The thinking within is meant to be reproduced by others.

The Machinery of Sexual Reproduction Williams anticipates a transition to behavior in Chap. 5, “Adaptations of the Genetic System,” by taking the machinery of sexual reproduction in higher animals and plants as an unmistakably evolved adaptation, because it is “complex, remarkably uniform, and clearly directed at the goal of producing, with the genes of two parental individuals, offspring of diverse genotypes.”20 He defines a Mendelian population as one that, by virtue of at least occasional sexuality, possesses a common gene pool. In the making of diploid organisms from haploid gametes—the process of meiosis, as we remember—Mendel determined that his peas had two sets of alleles, one from each parent. When the gametes containing these sets joined, they formed diploid organisms, with two (“di”) sets. Not all organisms reproduce in this way. Biologists very commonly regard sexual reproduction as a biotic adaptation, so that “natural selection itself [is] regarded as an adaptive mechanism whereby species avoid extinction or exploit the possibilities of adaptive radiation,” but Williams finds the fact that this entails a teleology reprehensible, since it suggests foresight. In more recent evolutionary studies, he finds a healthy tendency to “search for evolutionary forces in immediate circumstances rather than in future needs.” When arguments seem to work as well at one level as another, so that “acceptance of one level and rejection of the other might seem a matter of taste and of little consequence,” he still insists in precisely such situations that the observer must use Occam’s razor. If any phenomenon can be explained as an organic adaptation, it is simply not permissible to explain it as a biotic adaptation.21 Surely widespread existence of genetic recombination and a molecular basis of sexuality must be an ancient evolutionary development, and he thinks “sex is at least as ancient as DNA,” and though “it is most unlikely that DNA dates from the earliest stages of organic evolution,” the DNA molecule “has all the appearances of an evolved adaptation” because of its

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precision and effectiveness in coding information. Genes too are stable, but the gene is a very specialized chemical entity. “The only kind of change it can tolerate, and still function as a gene, is a very limited number of substitutions among rigidly specified chemical groups.” This would suggest that he believes sexual reproduction to be nearly as old as life. That mutations to chromosomes continue to occur after billions of years requires no special explanation, since the unquestionable principle—natural selection can often produce mechanisms of extreme precision, but never of perfection—means “It is not so much that mutations occur that requires the attention of biologists, but that they occur so rarely.” On this lowest level, the chemical level, “evolution takes place, not so much because of natural selection, but to a large degree in spite of it.”22 Williams dismisses notions that the “dominance” of normal genes over mutants has evolved, and sees no serious attempt to formulate a theory of dominance as a biotic adaptation. Such thinking appears only in general summaries of evolutionary principles. So, “There is no support for a general conclusion that heterozygosity [where an organism has dissimilar pairs of genes for a hereditary characteristic] has value as a biotic adaptation, either by allowing rapid evolutionary responses to changing conditions, or by increasing tolerance to long-term changes, or in any other way.” Nor does he believe that “introgressive hybridization”—a process that spreads the genes of one species into the gene complex of another due to hybridization between numerically dissimilar populations and extensive backcrossing—can be an adaptation. At large, because the whole reproductive machinery of the higher plants appears directed toward the successful crossing of members of the same species, “To postulate that this machinery is intended to fail a part of the time is completely gratuitous.”23 Williams asks why there should be separate sexes in some groups and not in others, and some with hermaphrodites, to which he has no answer. He is forced to proceed using a strictly sexual population and the common circumstance of one male being able to fertilize the eggs of a large number of females, so that he can ask “what would constitute an optimum sex ratio for population survival.” What follows is an elaborate set of calculations on sex ratios and sex-determining mechanisms. The calculations lead him to theorize that in strictly sexual populations an equal sex ratio will prevail “to the extent that natural selection, on the basis of individual reproductive success, is the determining factor,” but there are complications, the most salient being dependence of any offspring upon parents after conception. Nevertheless, in close conformity to his theory, no

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convincing evidence shows that sex ratios ever behave as a biotic adaptation. He wonders if “sex ratio provides the curious situation in which natural selection, instead of producing an adaptation, abolishes its function” and is somewhat frustrated because R. A. Fisher finds that sex ratio is not an organic adaptation and evolved for the species rather than for individual advantage. Though each individual may appear to be designed to produce equal numbers of male and female offspring, he wonders if this might be “an incidental consequence of the evolution of the sex chromosomes themselves,” where sex may be a historical legacy from the chemistry of life.24 He also notes that sex determination adjustments might be caused by external and demographic environments. Note that Williams has been speaking of populations that require a male and female to produce a new individual, and he expects a different outcome in a species where a female can produce viable offspring parthenogenetically, which is to say without fertilization. “This capacity would be especially important in what have been termed fugitive species … which frequently invade new habitats with an extremely small number of colonists.” This does happen, but more to the point, when conditions change to favor sexual reproduction, so too the picture changes rapidly and females readjust physiologically and behaviorally as sexual reproduction becomes advantageous. “Males are produced in large numbers whenever changed conditions make sexual reproduction adaptive.”25

Behavior of Sexual Machinery These premises about physiological and behavioral readjustment open the door to the mediation on “Reproductive Physiology and Behavior,” where Williams cites a definition of fitness from Peter Medawar, who had spoken of the “pricing of endowments” or “a genetic valuation of goods” for past individuals. Medawar had at least focused attention on reproductive success, but lacking clairvoyance, one cannot tell how fit a future individual might be, and luck may enter the equation. Instead, it is best to focus only on adaptations that relate directly to reproduction. For Williams, we remember, fitness may be defined as “effective design for reproductive survival”—reproductive survival being the central biological problem— and the more fit an individual is, the more likely that its adaptations, plural, allow it to contribute genes to future generations. Somatic survival is favored by reproduction only when necessary for reproductive survival. For instance, heart failure and mammary failure have “exactly equivalent

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effects on the fitness of a female mammal (except in the one bottle-­ manufacturing species).”26 To put it less nicely, bodies are disposable while their adaptations are necessary. Put in terms of means and ends, survival of a species is one result of reproduction, but this does not constitute evidence that survival of a species is a function of reproduction. Surely interactions between parent and offspring are crucial to reproduction. These may be considered adaptations and limit fecundity for this if for no other reason: “the maximization of individual reproductive success will seldom be achieved by unbridled fecundity.” This alludes to David Lack’s work, which shows the limitation of avian clutch size to be determined by nutritional sources because, as Lack complains, “there has been no end of uncritical thinking based on the assumption that reproduction is adjusted to the needs of populations.” When correlating fecundity and mortality rate, Lack finds that “the cause-effect relationship is the opposite to that usually recognized.” Obviously, no matter how well adapted and successful a species is, it cannot continue to increase indefinitely in a limited environment, so when a population approaches or reaches the carrying capacity of the environment, “a high fecundity merely increases the number of individuals that must die without issue.”27 Individual parents or pairs have limited resources. From the beginning, for instance, a fish egg has yolk reserves. Smaller eggs, like smaller seeds of plants, have greater mortality rates. To generalize, because the limited amount of nourishment that parents can provide must be divided up among the young, the only result must be “increased mortality as an ecologically inevitable consequence of the increased fecundity.” Williams finds no evidence for relationships between parental care and fecundity in fish, where fecundity is limited by food supply, successful spawning, and “ultimately, by the amount of space available for the storage of eggs in the body of a female.” Elsewhere, natural selection might reduce fecundity in relation to parental care because there is a clear advantage in limiting energy output for the production of gametes so that the parent might be better able to care for the eggs and young immediately afterwards.28 Regarding prenatal care, “it is only the female that expends a major part of her resources in the production of gametes.” For her, it is not hard to see, the optimum number of young will be related to the optimum mass of young. “If, at the time of spawning, food for the young will continue to be in low supply for some time, I would expect the eggs to be large and yolky, and therefore slow-developing and few.” Because optimum fecundity is related to the possibility of making an investment of energy, it is also

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limited by sources of energy. Once internal fertilization evolved, viviparity—bringing forth living young, rather than depositing eggs—would likely follow, not as teleology, but as an expected adaptation. Williams calls this an “optimum shift towards viviparity” because it allows improved viability of offspring, but he notices too that greater material sacrifices are required of mothers who produce advanced juveniles, compared to the energy required to produce the same mass of eggs. For this reason, he finds “absence of viviparity among the birds most mysterious.”29 If one assumes an optimum expenditure of materials by the mother, one must also see a larger problem, namely the amount of effort an organism must expend to reproduce and the degree of exposure to danger it faces in the course of reproduction. As Williams knows, “parental sacrifices are sometimes enormous, and sometimes very slight,” so he can restate fitness in this way: “A well-adapted individual is one that will engage in reproductive activities only when the chance of success is at some peak value and the probable costs and hazards at some low point.” With optimal reproduction, there is always an appropriate “if clause.” Reproduction sometimes entails extreme sacrifice, as with Pacific salmon that only breed once, and as he puts it, “among them we find the expected emphasis on reproductive functions to the detriment of the parental somata.”30 That is, parents die before offspring emerge. Where this sort of thing happens, one finds belligerence or pugnacity in the males, along with bright coloration. This is only one extreme in a spectrum that includes a broad range of types of life cycle. So Williams has prepared a way to speak of “reproductive effort.” He expects it to be greater among smaller organisms than among larger ones with lower mortality rates. As he indicates in a numbered list, the effort or hazard will be (1) “greater when a species spawns a large mass of eggs” relative to its size, (2) greater if it spawns several times in a breeding, (3) greater with “conspicuous breeding coloration” and “we would assume that the more conspicuous the color, the greater is the somatic hazard,” (4) “greater in species that engage in conspicuous courtship,” (5) greater in species where members of the same sex are combative, “especially when the competition includes actual fighting with effective weapons,” (6) greater in territorial than in nonterritorial species, (7) “greater in species that provide some special protection for their eggs,” and lastly, (8) “greater in viviparous than in oviparous species.”31 Further, an “optimum reproductive effort should not only vary among species, it should also change

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with time in any given species,” especially those that become more fecund for several seasons after becoming sexually mature. Such reasoning allows Williams to derive and express certain commonly observed—and perhaps clichéd or stereotypical—patterns of sexual behavior in terms related purely to energy expenditure and hazard. Males show “greater readiness for reproduction” than females as a consequence of the “greater physiological sacrifice made by females.” While a male mammal’s role may end with copulation, for the female “copulation may mean a commitment to a prolonged burden.” So one would expect that while the male is only involved, the female is committed. “Natural selection should regulate her reproductive behavior in such a way that she will assume the burdens of reproduction only when the probability of success is at some peak value that is not likely to be exceeded.” Hence “the traditional coyness of the female,” which leads Williams to speak of “female choice” and “discrimination.” “Inevitably there is a kind of evolutionary battle of the sexes,” he writes: males use “skilled salesmanship,” pretend to be fit, and show greater promiscuity, while females demonstrate greater caution. Females are at risk because “it is almost always true that females contribute the greater amount of material and food energy to the next generation.” One sees “development of the masculine emphasis on courtship and territoriality or other forms of conflict with competing males.”32 Sexual roles are simply a result of energy expenditures! Exceptions to these gender stereotypes—like pipefish and seahorses, tinamous and phalaropes—prove the rule. Williams finds overwhelming evidence that “promiscuity, active courtship, and belligerence toward rivals are not inherent aspects of maleness,” but are shaped by the allocation of material contributions. Given the energy investment, generally herds of gregarious mammals break up into pairs in foaling seasons. To state that more strongly, “I know of no evidence that the adaptive organization of subhuman mammalian reproduction ever normally transcends the interactions between a pair of parents and their offspring.” The principle is always the same, where all aspects of the reproductive behavior and physiology of a species—intensity, timing, ontogeny—and all physiological and behavioral mechanisms can be expected to be precisely designed to maximize individual reproductive performance.33 What then of “Social Adaptations,” the subject of his seventh chapter? Because Williams derives behavior from chemical, cellular, and the energy expenditure of reproductive functions, incidental interactions between unrelated individuals can only be incidental. The characteristic and

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uncompromising darkness of Williams’s view would disturb some of his colleagues. Like Hamilton, he notices that normal behavioral or physiological relations between parents and their own offspring are “benign and cooperative,” at least some of the time, “but interactions between unrelated individuals normally take the form of open antagonism, or, at best, a tolerant neutrality.” Williams insists that most of the conspicuous patterns of animal behavior between unrelated individuals are manifestly competitive, and must be attributed to competitive efficiency in selection for genetic survival. “To provide benefits to a genetically identical individual is to benefit oneself.” The nature of such behavior guarantees that natural selection will “rarely favor genes that cause their bearers to expend resources to benefit their genetic competitors.” Exceptions would include cooperation between closely related individuals, as illustrated by Haldane’s quip. Also, because there are behavioral mechanisms by which parents aid their offspring, it is inevitable that such aid may be provided “by mistake” to unrelated individuals. The possibility of parental mistakes also allows for the possibility of taking advantage of such mistakes. Williams summarizes Hamilton’s treatment of the genetic evolution of social behavior, and what it amounts to is that “the relative genetic homogeneity of a Mendelian progeny is a factor that significantly softens the competitive interactions among brothers and sisters.” This is true widely, for insects and for California woodpeckers. Only when reproductive behavior or function is misplaced does cooperation or self-sacrifice occur between unrelated individuals. At its maximum, “Benevolent behavior towards unrelated individuals should never be more intense, and should usually be less intense, than the same behavior towards offspring.”34 Williams entertains views contrary to his own, citing the production of hybrids, which he sees as an example of the malfunctioning of the mechanisms of reproduction due to looseness of timing or imperfect execution. He recognizes “accessory reproductive behavior” like warning calls and the rare but significant presence of “helpers,” unmated individuals that assist breeding pairs. He finds these most easily explained by closeness of genetic relationship. “Helpers appear to be birds that have their own reproduction frustrated in some way so that they must find some other outlet for their parental instincts.” So “the helper phenomenon can be attributed to selection pressures for the maintenance of a certain pattern of parental behavior, with a less-than-perfect system of timing mechanisms for regulating this behavior.”35

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Why Are There Groups At All? Since “the important initial problem is why animals should exist in groups of several to many individuals,” Williams seeks a parsimonious explanation of “why one individual would expend its own resources or endanger itself in an attempt to aid another.” Why are there groups at all? He reminds the reader of two misconceptions or false assumptions. The first serious error comes with “the assumption that when one demonstrates that a certain biological process produces a certain benefit, one has demonstrated the function, or at least a function of the process”—a clear fallacy because “demonstration of a benefit is neither necessary nor sufficient in the demonstration of function.” One must show that the process is designed to serve a function. The second misconception is “that to explain the functional aspects of groups, one must look for group functions.” As he is fond of saying, there is a difference between a mob and a functional group, and “the behavior of the mob is easily understood as the statistical summation of individual adaptation.” Consider the retention of warmth as a group benefit: “there is no more reason to assume that a herd is designed for the retention of warmth than to assume that it is designed for transmitting diseases.” For instance, “The huddling behavior of a mouse in cold weather is designed to minimize its own heat loss, not that of the group.”36 Though his favorite example of social grouping comes from schooling behavior in fishes, Williams finds that social groups among mammals differ because mammals “never more than partly abandon their roles in the drama of reproduction.” It is true that bulls are somewhat belligerent at all seasons, calves are dependent for several seasons, and cows may retain their attachment to calves while in herds. Wolves attack large game in packs out of individual need, but as he says in the first person, “I am not aware, however, of any evidence of functional organization of wolf packs,” and he does not find any functional organization in the “dominance-­ subordination hierarchy shown by wolves and a wide variety of vertebrates and arthropods.” Individuals may make compromises in competition for food, mates, or other resources: “each compromise is adaptive, but not the statistical summation.” All in all, he discounts the ubiquitous anecdotal reports of functional organization in groups of mammals: “Many of these may be more a product of romantic imagination than of careful observation.”37 He concludes that any possibility that such groups really do show functional organization warrants further research by students of

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gregarious animals, but he also notes that no studies have shown this to be so. He has thrown down the gauntlet. The challenge is open, but given the terms in which Williams frames it, the argument is powerful and leads in only one direction. It is no wonder that Williams would focus in his next book on the reasons for the prevalence of sexual reproduction, the process being foundational for patterns of behavior.

Teleonomy and Intuition: Of Clocks, Lights, and Eyes The conclusion to Adaptation and Natural Selection spells out a program for the study of adaptation. First, “the most urgent requisite for the success of any science is that it have a name.” Colin Pittendrigh, best known as “father of the biological clock” because of his pioneering work on circadian rhythm, also established the notion of teleonomy. Williams borrows the term to signify a concept that “would connote a formal relationship to Aristotelian teleology, with the important difference that teleonomy implies the material principle of natural selection in place of the Aristotelian final cause.”38 Teleonomy is a strange term to advocate that biologists adopt for general use. It is hardly suggestive of Williams’s plain style, revealing something of the problems a biologist faces in writing about adaptation. Ernst Mayr explains why it was invented in 1958. As Pittendrigh wrote to Mayr in 1970, I was haunted by the famous old quip of Haldane’s to the effect that ‘Teleology is like a mistress to a biologist: he cannot live without her but he’s unwilling to be seen with her in public.’ The more I thought about that, it occurred to me that the whole thing was nonsense—that what it was the biologist couldn’t live with was not the illegitimacy of the relationship, but the relationship itself.

As Pittendrigh thought, for biologists “teleology would not do, carrying with it the implication that the end is causally effective in the current operation of the machine. Teleonomic, it is hoped, escapes that plain falsity which is anyhow unnecessary.”39 Consequently, a great tangle of teleology and teleonomy enlivens the literature of biology. In 1958, Pittendrigh noticed the verbal scruples that

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led biologists to say “a turtle came ashore and laid its eggs” rather than “a turtle came ashore to lay its eggs.” Such utterances “were intended as a rejection of teleology but were based on the mistaken view that the efficiency of final causes is necessarily implied by the simple description of an end-directed mechanism… . [but] recognition and description of end-­ directedness does not carry a commitment to Aristotelian teleology as an efficient causal principle.”40 In any case, when Williams adopts Pittendrigh’s term to be used strictly to designate the study of adaptation, he also insists, “Teleonomy would not be a branch of the study of evolution. Its first concern with a biological phenomenon would be to answer the question: ‘What is its function?’”41 Within the great many arguments about teleonomy and teleology lie a few pretty good jokes. As usual, Ernst Mayr attempted to clarify by examining the use of teleonomic syntax: Let us take, for instance the sentence: ‘The Wood Thrush migrates in the fall into warmer countries in order to escape the inclemency of the weather and the food shortages of the northern climates.’ If we replace the words ‘in order to’ by ‘and thereby,’ we leave the important question unanswered as to why the Wood Thrush migrates. The teleonomic form of the statement implies that the goal-directed migratory activity is governed by a program. By omitting this important message, the translated sentence is greatly impoverished as far as information content is concerned, without gaining in causal strength.42

One ought not be astonished when Williams insists that study of adaptation ought not be a branch of the study of evolution, because his aim is to recognize functional design. He places himself in that tradition, writing, “I know of no better illustration of this process of recognizing functional design from careful observation than that provided by Paley (1836) in an answer to the suggestion that the eye just happened to be suitable for vision.”43 This strange authorial choice would seem to bring teleology back into the room—a room that it has never left. Williams continues, “Any such plausible demonstration of design in relation to a goal would provide the answer to the teleonomist’s prime question.” Then would come questions about mechanism, and others. He finds “too optimistic” Pittendrigh’s assumption that a biologist can discern the proximate end of an adaptation “fully by direct observation,” not only because functions, such as the function of the singing of birds, have

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been insufficiently discerned, but also because they have often been erroneously discerned. “How, ultimately, does one ascertain the function of a biological mechanism?” We remember his equivocation, that functional design “can be discovered intuitively and convincingly communicated to others.” Surely some biological problems can be approached intuitively: “We do not need weighty abstractions to help us decide that the eye is a visual mechanism.” Beyond that assumption, as he says, the study would require that intuition and communication through language might or might not someday be replaced by “standardized criteria.” Teleonomic understanding, or study of adaptation, would be aided by such tools as “a recognized hierarchy of adaptations, or at least a way of specifying the subordination of one function to another,” following “Tinbergen’s hierarchical classification of instincts on the basis of the nature and generality of the purposes served.” Only historical considerations could disclose what is functionally inexplicable, so “a teleonomic analysis would not proceed far without the use of historical data.” And in one final distinction, “I would like to nominate one more principle for initial inclusion in the science of teleonomy. This principle is that the nature of the stimuli that initiate and regulate a response may be no indication of the function of the response.”44 Pittendrigh illustrated this last principle by a striking demonstration. The timing of certain activities of wild populations of fruit flies seems to be controlled or regulated by visual cues from the daily cycle of illumination, but it turns out that the function of the timing is not an adjustment to conditions of illumination but to changes in humidity, because the insects “adapt to an important, but poorly perceived environmental factor, humidity, by reacting to a closely correlated factor, light, that is unimportant in itself but reliably perceived.”45 This is important because what an organism can perceive becomes proxy for what it cannot. The clock and the eye are sometimes adjusted, but they may respond to another variable that cannot be directly perceived. Pittendrigh sees that an organism’s adaptations to a specific pressure of selection will take different forms, and he announces that perception of “adaptive organization in its true light” will not be as of “Paley’s perfection demanding an intelligent designer, but rather as a patchwork of makeshifts pieced together, as it were, from what was available when opportunity knocked, and accepted in the hindsight, not the foresight, of natural selection.”46 In this world that evolutionary biologists like to imagine as a tinkerer’s workshop, where Williams and his colleagues work, teleonomy meets reality.

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Analogies—Mundane and Epic, Inflationary and Deflationary An investigation of design allows a teleonomist a certain freedom of expression: “The close analogy between the lens of a camera and the lens of an eye makes the term lens appropriate for both.” But transfers of terminology are appropriate only “when there is a real functional analogy between what man’s reasoning (and trial and error) can produce and what natural selection can produce.” Teleonomy is better than teleology, but cautions must be followed. Analogies may also sometimes be made between scientific developments themselves, as Williams does between contemporary theory of natural selection and “atomic theory of two centuries ago.”47 One might call this analogy between biology and physics the closest Williams gets to an epic simile. Yet there are no epic similes in Williams. Consider by contrast Loren Eiseley’s epic simile in Darwin’s Century (1959), where “[The Origin’s] mass of accumulated evidence had the weight of a boulder. Criticism flowed around and over it but the boulder in all its impenetrable strength remained.”48 But Williams celebrates neither Darwin nor Darwinian science generally. He is not celebrating at all. Steve Jones, himself a notable geneticist, once wrote that “evolution is to analogy as statues are to birdshit. It is a convenient platform upon which to deposit badly digested ideas.”49 The Steve Jones dictum contains, pithily, an argument for stylistic restraint. A biologist’s use of fancy and elaborate analogies is itself onerous: the aspiration to literary elaboration imposes an onerous burden, an obligation, so that the analogies themselves may outweigh their advantages. If scientists wish to present themselves as dignified or clear-thinking, they are best advised to use restraint in their style. One might say that when Williams, in 1966, presented a scientific view as an onerous task, using a flamboyant style would have been inappropriate unless there was a special reason to do so. Williams practices what he preaches. Some have argued that he was, as a result, naïve about the business of publishing. But his writing is largely coherent, and his attention to restraint may paradoxically reveal why he preferred to write complete books.

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Analogy and the General Reader In Adaptation and Natural Selection, as I have noted, Williams—like Darwin—provides an expanded table of contents. Unlike Darwin, Williams spells out his argument in complete and concise, well-structured sentences, so deftly that the most frequently cited sentence of the book comes from its table of contents. Earlier, in Chap. 2, “Natural Selection, Adaptation, and Progress,” Williams used an extended analogy about pink polar bears that demonstrates his idea of appropriate discourse for the general public. As elsewhere, he ranges between third, second, and first person narration. We might indulge at this point in the fanciful act of rendering all present polar bears and their descendants a bright pink. We can now be sure that the species will not henceforth survive as well. Its numbers will suddenly decline and its geographic and ecological range rapidly contract, but we cannot be sure that this decrease will proceed all the way to extinction. Each polar bear, after meeting unaccustomed frustration in its hunting, will adapt by hunting for longer periods of time. Some may learn that they can hunt more successfully at night than by day. These and other adjustments might enable the species to continue in those regions where pinkness is, for one reason or another, less of a handicap than in others. Needless to say, there are many obviously necessary adaptations. If, instead of depriving the bear of its whiteness we deprived it of its lungs, it would immediately become extinct. Such examples, however, do not invalidate the conclusion that the mere presence of an adaptation is no argument for its necessity, either for the individual or the population. It is evidence only that during the evolutionary development of the adaptation the genes that augmented its development survived at a greater rate than those that did not.50

In this analogical or perhaps allegorical narrative, Williams establishes that the mere presence of an adaptation is no argument for its necessity and shows why this is the case. Considered in the light of so-called progress, “It is mainly when biologists become self-consciously philosophical, as they often do when they address nontechnical audiences, that they begin to stress such concepts as evolutionary progress. This situation is unfortunate, because it implies that biology is not being accurately represented to the public.”51 This may indeed be a warning to himself; Williams becomes much more “self-consciously philosophical” in later writings.

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Life History: Lack’s Principle and Williams’s Principle Out of social behavior comes life history. What do lineages do? They are born, mature, and reproduce according to certain principles and schedules, as they have always done, constrained by certain calculable costs and benefits. Yet, as he commented much later, “Most evolving lineages, human or otherwise, when threatened with extinction, don’t do anything special to avoid it.”52 The process called life history is the focus of an emergent discipline for which David Lack and George C.  Williams built the foundations. One needs Lack’s principle(s) because Williams derives an important corollary to Adaptation and Natural Selection from Lack. David Lambert Lack (1910–1973) was an ornithologist and, like Williams, was interested more generally in evolutionary theory. The Natural Regulation of Animal Numbers (1954) is his most important book for Williams’s purposes in 1966, though he is best known for Darwin’s Finches (1947). As Ernst Mayr observed, David Lack revived “interest in the ecological significance of species” and demonstrated that “the process of speciation is not completed by the acquisition of isolating mechanisms but requires also the acquisition of adaptations that permit co-existence with potential competitors.”53 Lack’s interest in the acquisition and maintenance of adaptations, and his insistent focus on “individual fitness” as exhibited by sexual reproduction, paralleled that of Williams. In 1999, Barry Sinervo argues that the classic paradigms developed by Lack (1947, 1954) and Williams (1966) “provide the selective context for understanding the evolution of reproductive allocation.” As Sinervo notes, Lack established that avian clutch size is limited by the amount of care that parents provide, and this can be talked about in terms of “selective premium accrued by investing in additional offspring” versus “decreased fledging success in large clutches owing to limitations on parental effort.” As Sinervo also notes, production of large numbers of small offspring in organisms without parental care is “balanced by the high survival of a few large offspring.”54 This is a cost and benefit analysis. Sinervo claims that Williams contributed a second paradigm of life-history biology, “trade-­ offs involving costs of reproductive allocation,” which “provides the selective context for understanding selection during adult phases of the life history.” Williams observed that by investing in current reproduction, individuals defer costs to future reproductive success, with a consequence

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that “lowers parental survival, fecundity, or growth, which might reduce body size and fecundity.” The choice is to pay now or discount the future. To simplify, the theoretical foundation of Lack’s model is that natural selection favors neither reproductive strategies that maximize fertility nor those that minimize offspring mortality: it favors those that yield the maximum number of surviving offspring. This sometimes comes at the expense of less than maximal parental investment per offspring, and hence less than maximum chances of survivorship. Another way to put this is to say that intermediate clutch size will be favored by natural selection when offspring mortality increases with clutch size, as it often does. These constraints operate on all life histories. When Williams begins to work on Adaptation and Natural Selection, as he writes in the preface of 1996, “Right from its opening paragraph, Lack’s ‘The Evolution of Reproductive Rates’ was a sublime encouragement. I had found a biologist who believed, as decisively as I did, that natural selection is a real scientific theory.”55 Williams published “Natural Selection, the Costs of Reproduction, and a Refinement of Lack’s Principle” as a letter in American Naturalist later in the same year that Adaptation and Natural Selection appeared. Here he constructs an elaborate argument using a language of “cost and benefit” that runs from Fisher through Lack to Williams himself (though it has been claimed that Lack was unaware that Fisher worked out this problem).56 Because the salient question comes from Fisher, his text constitutes the epigraph for Williams’s letter: “It would be instructive to know not only by what physiological mechanism a just appointment is made between the nutrient devoted to the gonads and that devoted to the rest of the parental organism, but also what circumstances in the life-history and environment would render profitable the diversion of a greater or lesser share of available resources to reproduction.” Williams explores what Fisher’s question implies, turning Fisher’s awkwardly written prose into something clearer but still rather stiff. First, “mass selection of alternate alleles in a population is ordinarily the only force tending to maintain or improve adaptation.” If this is true, then a general solution to Fisher’s problem is possible, namely, the immediate costs of reproductive processes must be “in functional harmony with each other” and worth the long-range costs to reproduction, and “the uses of resources for somatic processes is favored to the extent that somatic survival, and perhaps growth, are important for future reproduction.”57 In this single passage, as Stearns argues, Williams encapsulates “the evolutionary forces that make soma disposable.”58

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Bodies are disposable. Williams writes to formalize what Lack had advocated, that “reproductive processes serve individual interest” but not the reproducing bodies, and he uses the distinction between organic and biotic adaptation he has developed. “An individual of a certain age and sex,” he writes, “has a reproductive value” that can be at least quasi-­ quantified, and “a reproducing organism is faced with a series of choices: defend or abandon a threatened mate or litter or territory, make more ovules, more pollen grains, or a vegetative branch, etc.” What follows is somewhat mathematical, but quite straightforward. Think of “reproductive value” as an investment ϕ: “The barely justified cost [of reproduction] is thus seen to be directly proportional to the magnitude of the threat or opportunity (a or b), or to their sum if both are involved, and also to the size of the investment at stake (ϕ).” But an organism also possesses a “residual reproductive value” that depends not only on such things as flight versus fight thresholds, but “also such quantitative characters as fecundity and development of secondary sex characters.” As a matter of broad applicability, one can construct a theory that “allotted resources will be distributed among the component processes in a way that maximizes the rate of reproductive profit on the investment.”59 Robert E. Ricklefs believes that “Lack was the dominant personality in the development of life-history thinking,” and his central idea was that “evolutionary responses to natural selection maximize individual fitness.”60 In Lack’s own words, “clutch size is considered to be ultimately determined by the average maximum number of young for which the parents can find enough food… . Clutch size increases with increasing latitude and daylength because, in general, a longer day enables the parents to find more food per day, and so to raise more young at one time.”61 Lack had shown that because of the limitations on food gathering, any birds laying more eggs than the average for a species may end up producing fewer fledglings. So Williams infers, “there must be some point of diminishing return.” As a result, it should be possible to rank aspects of reproducing species so that “the sum of its rankings for several aspects of reproduction could serve as a species index of reproductive effort.” Lastly, for species with low annual mortality, and especially those that increase fecundity from year to year, “the parent’s lifetime reproductive interest may demand a low reproductive effort in any given season, much lower than that which would maximize reproduction for the season.” In conclusion, he finds it not necessary, as Lack (1955) himself had urged, to

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confine Lack’s principle of maximization of individual reproductive effectiveness to comparisons within species.62 Consequently, Williams’s refinement is in fact an expansion. To understand why Williams calls it a refinement one must consider costs and benefits and “discounting.” Lack understood how certain constraints on the adaptation of life history “arise from allocation trade-offs between number versus size of offspring,” to use Sinervo’s phrase, and current versus future reproductive success. Williams realized that this was a form of “discounting.” As Sinervo puts it, “an enhancement of one fitness trait is expected to lead to changes in other traits that can have negative effects on fitness.”63 Later commentators speak of this process as the “tradeoff.” Mart R.  Gross generalizes the Williams principle of life history as “investment into the present comes at the cost of investment into the future.” Therefore natural selection will, subject to this constraint, always favor behaviors that maximize reproductive success over a lifetime. Because the cost of investment made into current progeny is traded off with investment made into future progeny, “natural selection will optimize the allocation of investment into the present relative to the future.”64 Gross believes that in 1972 Robert L. Trivers was implicitly using this principle. Bobbi Low used it later, in 1978. Williams’s way of articulating tradeoffs becomes infectious, especially “the notion of tradeoffs for understanding parental care.” Peter Singer has spoken of an “escalator of reason,” arguing that “once reasoning has got started it is hard to tell where it will stop.”65 This is perhaps the kind of escalator Lack and Williams set in motion, and that Robert Trivers, Bobbi Low, Sarah Blaffer Hrdy, Stephen Pinker, and other sociobiologists have stepped upon. A few years later, in Sex and Evolution, Williams coined the term “Sisyphean genotypes”—those that require continuous redoing.66 He is also supposed to have wisecracked that evolution of new species takes place despite natural selection. As Williams reiterates in 1992, “Every organism will show a long list of characters that make no adaptive sense but record past adaptations.”67 This way of thinking not only makes the term progress difficult, but also makes any program of reverse engineering rather difficult.68

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Signs, Choice, and “Selfishness” Organisms make choices, whether conscious or not. As we have seen, Williams has proceeded down a path that allows him to imagine tentatively the costs and benefits of behavioral choices, but not their progress. What he said has become the subject of cultural contention. Regarding female choice, Williams reasons in Adaptation and Natural Selection as follows: first, “the optimum reproductive effort should not only vary among species, it should also change with time in any given species,” and second, “the optimization of effort, combined with the different primary roles of the sexes, may explain the different secondary roles of the sexes.”69 Speaking in terms of “vernacular fitness,” Williams notes, or perhaps even jokes, “Unusually fit fathers tend to have unusually fit offspring.” Since courtship is a form of advertisement by a male, “a male whose general health and nutrition enables him to indulge in full development of secondary sexual characters, especially courtship behavior, is likely to be reasonably fit genetically.” There are other signs of “vernacular fitness” by which natural selection functions, including a choice nesting site, good territory, and even the power to defeat or intimidate other males. Any female choice is dictated by a dynamic of dominance and submission, because “in submitting only to a male with such signs of fitness a female would probably be aiding the survival of her own genes.”70 Being the result of several considerations, any “choice” of roles in the evolutionary battle of the sexes is not fixed. As usual, exceptions reveal the rule, for in birds like the tinamous and phalaropes the male customarily assumes all or most of the burden of incubating the eggs and feeding the young. Here, “as expected, the female is the aggressor in courtship, more brightly colored than the male, and inclined to polyandry in both groups.” As a result, The evidence strongly supports the conclusion that promiscuity, active courtship, and belligerence toward rivals are not inherent aspects of maleness. They will be developed in whichever sex cannot effectively increase its production of offspring merely by increasing its material contribution, either by an augmented production of gametes or supply of food energy to the support of young.71

Sarah Blaffer Hrdy began to work on a much more sophisticated model of female choice, and when she did so she was supported by Williams.

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Leigh Van Valen (1935–2010) expresses concern about Williams’s adaptationist program, or the dynamic that Gross calls the Williams Principle of Life History. In 1973, Van Valen posits a “law of constant extinction,” stating that the probability of extinction for a species bears no relation to how long it may have already existed. He is most famous for the “Red Queen hypothesis,” using a metaphor from Lewis Carroll that expresses a never-ending struggle for existence, where no species or lineage can cease to change. As the Red Queen told Alice, “Now, here, you see, it takes all the running you can do to keep in the same place.” Natural selection becomes, metaphorically, an “arms race”—and “arms race” becomes a “concept” in the writings of Richard Dawkins.72 In one of his last essays, “How Ubiquitous is Adaptation?” (2009), Van Valen reminds the contemporary reader that he reviewed processes known for non-adaptive evolution in 1960, and then in 1966 Williams “demolished some proposals on optimization for which there seemed to be no viable mechanism.” Williams was followed by Gould and Lewontin who “made an analogous refutation of some more recent overenthusiastic work.” Van Valen concedes that “both these critiques were valuable in paring down excesses,” but he argues that “both can now be seen as overextending themselves and inhibiting productive diversity that has later managed to emerge.”73 From everything I have read, I believe that Williams attempted to constrain speculation, not overextend it. As a result he creates a style of writing that ought to impinge upon—humanists would say police—how biologists represent what happens. His style is not ornamentation but serves to encourage parsimony and attention to evidence in the next generation. He led Sarah Blaffer Hrdy to look for evidence of selection at the individual level when she studied infanticide in Hanuman langurs, and Mary Jane West-Eberhard to look for individual-level adaptations when she studied dominance interactions in paper wasps. As Blaffer Hrdy points out, this book altered discourse in his discipline, though “it was not always directly.”74 Often enough it was. Doris Williams remembers, “He was also a wonderful husband and helper through our lives together.”75

Thinking of Trees in 1966 Because Williams encourages cold knowledge, a reader must step back and pay attention to his sly analogies from the beginning. Everyone knows the story of the apple, knowledge, and the fall. Consider a story of an apple

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falling from a tree, viewed from the perspective of the laws of physics plus natural selection. All one needs to explain the trajectory of a falling apple are its mechanical properties, initial position, and velocity. Then the principles of mechanics would be sufficient, as adequate for the apple as for a rock. “If, however, we were asked how the apple acquired its various properties, and why it has these properties instead of others, we would need the theory of natural selection, at least by implication.” Then what? Close observation of the apple reveals that it is covered with waterproof wax and contains dormant embryos, and “an impressive list of structural details and processes of the apple can be understood as elements of a design for an efficient role in the propagation of the tree from which it came.” Surely, as he says, an apple’s contribution to man’s economy is not relevant for a biological discussion. “Thus I would say that reproduction and dispersal are the goals or functions or purposes of apples and that the apple is a means or mechanism by which such goals are realized by apple trees. By contrast, the apple’s contributions to Newtonian inspiration and the economy of Kalamazoo County are merely fortuitous effects and of no biological interest.”76 But an apple can never be a casual image. Allusions to the tree of knowledge are prominent in texts by Williams. Elsewhere in Adaptation and Natural Selection, Williams presents a shark that “was said to bite off mouthfuls of the school in a manner that suggested a person eating an apple.” A quince root system “may supply water and nutrients to an apple trunk.” He discusses a spider monkey spending most of its time in a tree, birds nesting in holes in trees, trees as safe retreats, trees used for storage and settlement by California woodpeckers, the fusing of the root systems of trees.77 His stories often have trees in them, but of particular sorts, and perhaps not the species one would expect in a book about evolution. The image or model of the tree is an abstraction closely associated with thinking of the relationships between species—as it also is with thinking on original sin—and, for biologists, trees are most important for phylogenetic thinking. Darwin’s central metaphor, after all, is of the tree of life. The apple is a structure whose purpose is to proliferate life, and that process is essential for the branching tree of life. A dangerous knowledge of the apple’s function is also implied. At the same time, other organisms live in trees in ways that are biologically relevant. A subversive tree grows more complicated in Williams’s Natural Selection (1992). He refers to a 1989 article entitled “Inclusive Fitness in a Nutshell” that he calls a “provocative discussion of the conceptual

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challenge presented by any fruit or even a single seed.” He writes, “A nut on a nut tree at first sight might appear to be a part of the tree, and on closer consideration an individual in its own right. Still closer attention shows it to be a composite of several genetically distinct individuals of varying kinship, each of which must be selected to optimize its interactions with each of the others.” He comments in passing that one can predict “the range of trunk diameters of trees harvested by beavers as a function of distance from the pond.” And he marvels, “mammals and insects and trees are so endlessly complicated, from the grossest down to the most microscopic structures, and so strikingly different from each other.”78

Natural Selection as Allegory Any written text becomes interpretable through simile, as an allegory or allegorical narrative. Sometimes such interpretability is humorous, and the humor, though not explicitly encouraged by a scientific writer, ought to be factored in. Take Darwin’s much storied introduction of the simile of the tree, which begins, “The affinities of all the beings of the same class have sometimes been represented by a great tree. I believe this simile largely speaks the truth.” Darwin’s simile includes “growing twigs [that] have tried to branch out on all sides, and to overtop and kill the surrounding twigs and branches, in the same manner as species and groups of species have at all times overmastered other species in the great battle for life.”79 Currently, there is much discussion about the demise or uprooting of Darwin’s simile of the tree of life and the notion that species could master one another.80 For one thing, Joel C. Velasco of Caltech observes that the image of the tree has always been linked to images of ladders or chains.81 The image of the ladder is a form of the great chain of being, which prompts one to imagine some kind of hierarchy of traits, with lower creatures possessing basic ones and higher creatures more advanced ones. Darwin’s narrative assumes a unity of life that Williams and others find dubious. Readers who do not believe in Darwin’s unity see Darwin’s narrative as an allegory and a literary artifact. And Williams asks us to go another layer lower. He is not interested in Darwin’s orders, families, or genera. Despite the silliness of such a simile, there remains a certain humor in working it out in detail.

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Sometimes the allegory grows, along with its humor and even its ridiculousness, as the writer casts and recasts a narrative. The narrative turns into an allegory by the process of abstraction and eventually becomes its own reduction to absurdity. The allegory remains even when the hypothesis is no longer considered valid, and even as the absurdity shadows it. This set of processes in literary history do not auger a good thing or a bad thing, but reveal simply how language and narrative work upon attentive readers, leaving their traces in thought. The contemplation of any narrative, as representation, offers an opportunity to turn it into a simile or an allegory. No narrative suffers complete demise. Yet “tree-thinking” has been considered for many years a central teaching tool in biology. As Robert J. O’Hara writes, “The ability to analyze evolutionary ‘why’ questions in this way comes from what I call ‘tree thinking’”—as after Mayr’s “population thinking”—and he considers this metaphor absolutely necessary for answering almost all evolutionary “why” questions.82 When in 1966 Williams deliberately avoids engaging in phylogenetic tree-thinking, this is particularly interesting because he does so a few years later in Sex and Evolution, depicting and labeling two diagrams as “random phylogenies” and “the evolution of hypothetical character states.” I take his phrasing as a way of reminding the reader that these diagrams are entirely suppositional.83 In 1992, he uses these diagrams more avidly, exploring the use of dendrograms to model a variety of evolutionary phenomena. He does not call these cladograms—similarities between phylogenies—as many other biologists do. A dendrogram is a branching figure, usually “rooted,” and must show evolutionary relations. Either figure may or may not be “evolutionary.”

What Progress? There is a danger in “tree-thinking” because the tree, as an aesthetic object, seems to reach into the sky, suggesting ascent and progress. This is why Stephen Jay Gould has suggested that evolution is a bush, not a tree. An ever-present analogy lurks between change, progress, and ascent. Williams seems to shudder when he reports that “long-term evolutionary determinism is still detectable in some discussions of progress in evolution.” Though he finds a widespread belief among biologists “in some form of aesthetically acceptable progress as an inevitable outcome of organic evolution,” he does not find it. “Once a certain level of complexity is evolved, selection will maintain adaptation by occasionally substituting

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one adaptive character for another, but this will not result in any of the kinds of cumulative progress that have been envisioned.” While he allows that science may progress, he maintains quite specifically that “there is nothing in the basic structure of the theory of natural selection that would suggest the idea of any kind of cumulative progress.”84 Ideas of progress saturate the tradition, and more importantly saturate the human view of its own species as higher and its behavior as more benevolent than that of “lower” creatures. In the writing of Julian Huxley, who named the modern synthesis in which “the force that drives and guides evolutionary progress is said to be natural selection,” Williams finds “an excellent example of how one can abide by the outward forms of the theory but violate its spirit.”85 Williams often expresses his most abiding fears by creating poetic conceptions and then undermining them with satire, because he believes that all biologists should fear “descending into teleology.” Consequently, much of his own style of writing is meant to demonstrate ways of avoiding abrupt and disastrous descents by recognizing how teleological assumptions sneak up from behind. Williams supposes that “belief in a deterministic succession of evolutionary stages” was put to rest by George Gaylord Simpson’s book of 1944.86 Simpson demolished a notion sometimes called “orthogenesis,” that life has some internal, “mysterious inner force” driving it to progress, leading to horses and men. Williams says that Simpson’s book “can be taken to symbolize the end of orthogenetic interpretations of paleontological data.” How could it be, then, that so many concepts of evolutionary progress still “adhere to the forms of an earlier orthogenetic doctrine even though the doctrine is almost unanimously discredited”? Yet people continue to suppose that evolution, having produced an organism like a man or a horse, means that progress can be “arbitrarily designated as any change in the direction of man or horse.”87 “Biologists who are especially zealous in avoiding such concepts as evolutionary progress will sometimes use the term specialized instead of advanced.” It may be reasonable to use such terms as progress and advance to “designate conformity to common phyletic trends or approach to an arbitrarily designated final stage, but unfortunately the acceptance of the term in this sense can disguise its use in other senses.” Progress is part of the linguistic heritage and cognitive structure humans use to sort things, part of stories used to make sense of the world. Williams suspects that evolutionary progress leading to “the inevitability of man” only seems to be a scientific idea “because of our heritage of such orthogenetic terms as

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‘higher’ or ‘advanced’ organisms and the fact that a list of taxonomic categories has to have a beginning and an end.”88 Nevertheless, he does not expect the ideology of progress to disappear from scientific or public discourse. His point about progress relates directly to the study of behavior. In Williams’s view, adaptations tend to be additive, not advancing toward any goal. Consequently, animal behaviors do not “improve” or refine themselves. The behavior of an organism does not revolutionize itself. Behavior suffices. In this sense, one can say that the sexual and social behavior of all organisms meet similar challenges of risk and call for similar effort: therefore, one would expect them to be similar for all species, given the constraints. Human behavior is more like than unlike the behavior of other species facing similar reproductive tasks, so that sociobiological analysis should be applicable across many varieties of sexually reproducing species.

Notes 1. George C.  Williams, Adaptation and Natural Selection (Princeton University Press, 1966), 4. 2. S.  C. Stearns, “George Christopher Williams 1926–2010. Biographical Memoir.” National Academy of Sciences (2011), 10. http://www.nasonline.org/publications/biographical-­m emoirs/memoir-­p dfs/williams-­ george.pdf. 3. See Owen Flanagan, “Varieties of Naturalism,” Chapter 26  in Oxford Companion to Religion and Science (Oxford University Press, 2006): 430–52. 4. Williams, “Preface,” in Adaptation and Natural Selection, Princeton Science Library Edition (Princeton University Press, 1996), xii. 5. Williams, Adaptation, 8, 11, 19. 6. Williams, Adaptation, 22, 49. 7. Williams, Adaptation, 232–33. 8. Williams, Adaptation, 233. 9. Williams, Adaptation, v. 10. On “genetic hitchhiking,” see J. Maynard Smith and John Haigh, “The Hitch-Hiking Effect of a Favourable Gene,” Genetics Research 23, no. 1 (1974): 23–35. On “genetic drift,” see Motoo Kimura, “Solution of a Process of Random Genetic Drift with a Continuous Model,” Proceedings of the National Academy of Sciences of the United States of America 41, no. 3 (1955): 144. On “spandrels,” see Stephen Jay Gould and Richard C. Lewontin, “The Spandrels of San Marco and the Panglossian Paradigm:

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a Critique of the Adaptationist Programme,” Proceedings of the Royal Society of London. Series B. Biological Sciences 205, no. 1161 (1979): 581–98. 11. Williams, Adaptation, 93, 94, 111. Williams, Natural Selection: Domains, Levels, and Challenges (Oxford University Press, 1992), 139. 12. Williams, Adaptation, 3. 13. Dawkins quote in “George C.  Williams (1926–2010), A Remembrance with Robert Trivers, Richard Dawkins, Daniel C. Dennett, Niles Eldredge, Randolph Nesse,” Edge, September 9, 2011, https://www.edge.org/conversation/robert_trivers-­r ichard_dawkins-­d aniel_c_dennett-­n iles_ eldredge-­randolph_nesse-­george-­c. Robert Alter, Pen of Iron: American Prose and the King James Bible (Princeton University Press 2010): 4, 12. Harold Bloom, The Shadow of a Great Rock: A Literary Appreciation of the King James Bible (New Haven, CT: Yale University Press, 2011), 17. 14. Williams, Adaptation, 120, 31, 260, 269. 15. Williams, Adaptation, 95. 16. Williams, “Preface,” xi. 17. Williams, Adaptation, 158, 159, 17. 18. William D. Hamilton, “The Evolution of Altruistic Behavior,” American Naturalist 97, no. 896 (1963): 354–356, 354. 19. Williams, Adaptation, xv. 20. Williams, Adaptation, 125. 21. Williams, Adaptation, 127, 129, 130–31. 22. Williams, Adaptation, 133–34, 138, 139. 23. Williams, Adaptation, 143, 144, 145. 24. Williams, Adaptation, 147, 150, 151, 152, 153, 157. 25. Williams, Adaptation, 155, 157. 26. Williams, Adaptation, 158, 159. 27. Williams, Adaptation, 160, 162, 163. 28. Williams, Adaptation, 164, 165, 166. 29. Williams, Adaptation, 166, 167, 170. 30. Williams, Adaptation, 172–3, 175. 31. Williams, Adaptation, 177–9. 32. Williams, Adaptation, 183–4, 185. 33. Williams, Adaptation, 189, 191–2. 34. Williams, Adaptation, 193, 163–5, 203, 204. 35. Williams, Adaptation, 205, 208. 36. Williams, Adaptation, 212. 37. Williams, Adaptation, 217, 218, 220. 38. Williams, Adaptation, 258.

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39. Ernst Mayr, “The Multiple Meanings of Teleological,” in Towards a New Philosophy of Biology (Cambridge, MA: Harvard University Press, 1988), 38–66, 65, 66. 40. Colin Pittendrigh, “Adaptation, Natural Selection, and Behavior,” in Behavior and Evolution, edited A. Roe and George Gaylord Simpson (New Haven: Yale University Press, 1958), 390–416, 394. 41. Williams, Adaptation, 258, italics mine. 42. Ernst Mayr, “Teleological and Teleonomic: A New Analysis,” Boston Studies in the Philosophy of Science 14, no. 1 (1974): 91–117. 43. Williams, Adaptation, 258–9. 44. Williams, Adaptation, 260, 261, 263, 269. 45. Williams, Adaptation, 269. 46. Pittendrigh, “Adaptation,” 400. 47. Williams, Adaptation, 261, 271. 48. Loren C. Eiseley, Darwin’s Century: Evolution and the Men who Discovered it (New York: Anchor Books, 1961), 195. See also Mary Ellen Pitts, Toward a Dialogue of Understandings: Loren Eiseley and the Critique of Science (Lehigh University Press, 1995), 131, 132. 49. Steve Jones, “Go Milk a Fruit Bat!” New York Review of Books 44, no. 12 (July 17, 1997). A review of Why Is Sex Fun? The Evolution of Human Sexuality, by Jared Diamond. 50. Williams, Adaptation, 28–9. 51. Williams, Adaptation, 55. 52. Frans Roes, “A Conversation With George C. Williams,” Natural History 107 (May 1998): 10–13. 53. Ernst Mayr, The Growth of Biological Thought (Cambridge, MA: Harvard University Press, 1982), 274–5. 54. Barry Sinervo, “Mechanistic Analysis of Natural Selection and a Refinement of Lack’s and Williams’s Principles,” The American Naturalist 154, supplement (July 1999): S26–42, S27. 55. Williams, “Preface,” x. 56. See David Lack, “The Evolution of Reproductive Rates,” in Evolution as a Process, ed. J.S. Huxley, A.C. Hardy, and E.B. Ford (London: Allen and Unwin, 1954), 143–56. 57. Williams, “Natural Selection, the Costs of Reproduction, and a Refinement of Lack’s Principle,” Letters to the Editors, American Naturalist 100, no. 916 (Nov-Dec 1966): 687–90, 687. 58. Stearns, “George Christopher Williams,” 12. 59. Williams, “Natural Selection,” 687, 688, 689. 60. Robert E. Ricklefs, “Lack, Skutch, and Moreau: The Early Development of Life-History Thinking,” Condor 102 (2000): 3–8, 3.

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61. David Lack, “The Significance of Clutch-Size,” Ibis 89 (1947): 302–52, 331. 62. Williams, “Natural Selection,” 689. 63. Sinervo, “Mechanistic Analysis,” S26. 64. Mart R. Gross, “The Evolution of Parental Care,” the Quarterly Review of Biology 80, no. 1 (2005): 37–45, 39. 65. Peter Singer, The Expanding Circle: Ethics and Sociobiology (Farrar, Straus and Giroux, 1981), 113–14. 66. Williams, Sex and Evolution (Princeton University Press, 1975), 45 and ff. 67. Williams, Natural Selection, 7. 68. See Chris Haufe, “Perverse Engineering,” Philosophy of Science 75 no. 4 (2008): 437–46. 69. Williams, Adaptation, 182, my emphasis. 70. Williams, Adaptation, 184. 71. Williams, Adaptation, 186. 72. Leigh Van Valen, “A New Evolutionary Law,” Evolutionary Theory 1 (1973): 1–30. 73. Leigh Van Valen, “How Ubiquitous is Adaptation? A Critique of the Epiphenomenist Program,” Biology and Philosophy 24 (2009): 267–280, 268. 74. Personal communication. 75. Personal communication, May 23, 2013. 76. Williams, Adaptation, 6, 8, 9. 77. Williams, Adaptation, 69, 91, 108, 201, 215, 223, 238. 78. Williams, Natural Selection, 20, 65, 127–8. 79. Charles Darwin, The Origin of Species, 6th edition (1872), 104–5. 80. W.  Ford Doolittle, “The Practice of Classification and the Theory of Evolution, and What the Demise of Charles Darwin’s Tree of Life Hypothesis Means for Both of Them,” Philosophical Transactions of the Royal Society B 364 no. 1527 (August 12, 2009): 2221–28; W.  Ford. Doolittle, “Uprooting the Tree of Life,” Scientific American 282, no. 2 (2000): 90–95. 81. Joel Velasco, “The Future of Systematics: Tree-Thinking Without the Tree” (in Philosophy of Science Dec. 2012). Available at http://joelvelasco. net/Papers/. 82. Robert J. O’Hara, “Homage to Clio, or, Toward an Historical Philosophy for Evolutionary Biology,” Systematic Zoology 37, no. 2 (1988): 142–155, 151. http://rjohara.net/cv/1988-­sz. See also David Baum and Stacey Smith, Tree Thinking: An Introduction to Phylogenetic Biology (Greenwood Village, CO: Roberts, 2012). 83. See Williams, Sex and Evolution, 165, 163, illustrations 2.1, 2.2, 2.4, and 3.2.

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84. Williams, Adaptation, 21, 22, 5, 34. 85. Williams, Adaptation, 22. 86. George Gaylord Simpson, Tempo and Mode in Evolution (New York: Columbia University Press, 1944). 87. Williams, Adaptation, 21, 47. 88. Williams, Adaptation, 47–9.

CHAPTER 5

Sex, Death, and the Language of Sociobiology

Whoever understands reproduction unlocks a key mechanism of evolutionary change—but nobody understands reproduction perfectly. We remember that sexual reproduction produces offspring with new combinations of the parental genes by means of “machinery designed to produce that result.” Consequently, sexual behavior is essential to Williams’s thought and, in 1966, he writes confidently that, in higher animals and plants, sex is “unmistakably an evolved adaptation.”1 If only it were so simple! On the one hand, reproductive mechanisms evolve; on the other hand, evolution is driven by reproductive mechanisms. In 1966, Williams requires that general trends in evolution must be “a by-product of the maintenance of adaptation.” Because survival depends on reproduction, no matter how much any organism’s appearance changes, reproduction “would still show the uniquely biological property of adaptation, and it would still be precisely adjusted to its particular circumstances.”2 Yet even as he writes, Williams knows what continues to be the case as late as 2001: “theory has so far failed to provide a universal and satisfactory explanation for the adaptive advantages of sex in Darwinian terms.”3 Williams prophesizes that his argument of 1966 will fail, but not from lack of trying. He tries again in 1975, in his book Sex and Evolution, where two aims are linked: because “prevalence of sexual reproduction in higher plants and animals is inconsistent with current evolutionary

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 M. P. Cohen, George C. Williams and Evolutionary Literacy, Literatures, Cultures, and the Environment, https://doi.org/10.1007/978-3-031-11650-6_5

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theory,” the theory must be modified “to account for the persistence of so seemingly maladaptive a character.”4 Paradoxes abound here. Why should higher organisms not behave like bacteria and use the machinery of parthenogenesis? Parthenogenesis is defined as reproduction without fertilization; partheno- means virgin, so this term evokes virgin birth. By 1975, Williams believes that parthenogenesis or its equivalent always replaces sexual reproduction in vertebrate populations when that is possible, so by his definition sexuality in these organisms must be maladaptive.5 If it is maladaptive, why would sexual design be so universal, so omnipresent? Typically, Williams does not predict answers to such questions, but their likely currency. “Prophesy is a hazardous exercise in science,” he writes in the introduction to Group Selection (1971), an anthology he edited about the controversy known by that name, but he believes evolutionary theory “will center increasingly on the phenomenon of sexuality.”6 Teachers publish anthologies for their students and include their “favorite” writers. Williams’s 1971 anthology reveals his views, as all anthologies do, by juxtaposition of texts. He buttresses his own thinking with that of his colleagues, highlighting perspectives that he thinks advanced students should learn and explore. He also sets their perspectives against those of others, including essays on the subject of sex from many major players of the early 1970s. In retrospect, their contributions initiate a sociobiological tradition. One should include in this group at least Williams himself (b. 1926) on adaptation, John Maynard Smith (b. 1920) on optimization and game theory, William Hamilton (b. 1936) on inclusive fitness, and Robert Trivers (b. 1943) on reciprocal altruism. The three elders of this select group are well represented in Group Selection. Most prominently, Hamilton’s “The Genetical Evolution of Social Behavior, I & II,” in slightly revised form, comprises over a third of the anthology. This set of manifestos by these writers creates a language in a campaign that places biology within an argument about adaptation and sexual reproduction and expands that argument. The terminologies for these investigations change, but the issues are closely intertwined and call for unpacking. These writings about sex become both collaborative and contentious, and though initially the most important discussants are Williams, Hamilton, and Maynard Smith, Trivers produces a set of articles in the early 1970s that contribute to what Williams has to say. Robert Trivers and John Maynard Smith later “worked through the whole manuscript” of Sex and Evolution.7 Writing after Williams’s death in 2010, Laura Betzig, who was a graduate student at the time, remembers this key discussion as created by the

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big three: “Back in the day, there were three big names in evolutionary biology: Bob Trivers, Bill Hamilton and George Williams.”8 She also remembers Trivers admitting that “the importance of sex role-reversed species was first suggested by George Williams (1966), and I certainly read it there, but I forgot where I had learned it and imagined that I had made this extension myself.” When Hamilton wrote about inclusive fitness, senescence, selfish herds, or sex, he credited Williams for anticipating his work, saying that Williams had a “characteristic knack of seeing a problem from a totally fresh point of view.” When Richard Dawkins and E. O. Wilson published later, their first debt, Betzig thinks, was also to Williams because he taught “fundamentals of the levels of selection and adaptation” that “defined and clarified the problem.” One must always remember that the purpose of critique is to clarify, reveal possible avenues of thought, and especially to help young researchers. Betzig thinks that the field’s empirical debt to Williams is enormous because his work opened up sociobiology. I believe she is right. But that is to get ahead of the story.

The Language of Sociobiology When Williams engaged in an extended conversation with Robert Trivers, John Maynard Smith, and William Hamilton by writing Sex and Evolution, this conversation invented a language for sociobiology. In other words, this community created a discourse, a way of discussing certain problems, in order to gain power over them. I borrow the term discourse from Michel Foucault, who means by it, in simplified terms, the rules, systems, and procedures that are part of a social “will to knowledge” and that create and institute a discrete realm that he calls the order of discourse, “the conceptual terrain in which knowledge is formed and produced.”9 Williams, Trivers, and many others locate the origin of sociobiological discourse, or kinship theory, in Hamilton’s 1964 paper, which defines four major categories of social interaction—namely altruistic, selfish, cooperative, and spiteful.10 These behaviors must be understood in terms of their consequences, specifically their “effects on the reproductive success of the two individuals involved.”11 Hamilton institutes a language of behavior that does not ask about internal motives, but instead scrutinizes costs and benefits, acts and “payoffs.” Trivers introduces three relevant concepts as part of kinship theory. One is “reciprocal altruism,” where unrelated individuals will engage in

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altruistic behavior if they can expect future benefits from strangers that surpass the short-term cost of helping. The second is “parental investment,” where the behavior of parents who invest time, energy, resources, or risk increases the probability of offspring survival at the cost of the parents’ ability to produce or care for more offspring and themselves. The third, “parent-offspring conflict,” arises in sexually reproducing species because parents and offspring are not identical, but only related by one-­ half. Incomplete genetic overlap creates a conflict of interest whose implications are worked out in terms of the amount of time and energy the parent invests versus what the offspring seeks. In his famous paper of 1971, “The Evolution of Reciprocal Altruism,” Trivers uses the term “cheating” to denote failure to reciprocate, and consequently observes that a mechanism must evolve for detecting cheaters. He also shows briefly that the discourse of costs and benefits used in the discussion of reciprocal altruism is “exactly analogous to what game theorists call the Prisoner’s Dilemma.”12 Williams engaged with this language when, as Trivers remembers, he received that paper and praised it.13 Williams mentions Trivers’s ideas often in Sex and Evolution, but not always with approval. Consider the language of the prisoner’s dilemma, a classic of game theory in which each of two isolated prisoners must decide whether to confess (defect) or protect (cooperate with) the other. Robert Axelrod and William Hamilton offer a definitive description of the game in an evolutionary context: “In the Prisoner’s Dilemma game, two individuals can each either cooperate or defect. The payoff to a player is in terms of the effect on its fitness (survival and fecundity). No matter what the other does, the selfish choice of defection yields a higher payoff than cooperation. But if both defect, both do worse than if both had cooperated.”14 The prisoner’s dilemma has become a metaphor for the difficulty of achieving cooperation among rational, self-interested individuals, and some think it accounts for the generally pessimistic tone of evolutionary biologists who have written on the subject of altruism and cooperation. As a discourse, it begins with cooperation and defection and relies on analyzing the costs and benefits of rational choices. The origins of cost-benefit analysis are hazy, but it begins in engineering and was perhaps invented by Jules Depuit (1804–66), a French engineer and economist. John Maynard Smith was an engineer, and Williams studied engineering briefly. The dilemma takes varying forms, illustrated by various versions of a payoff matrix.15

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The Cost of Sex In an original article for Williams’s 1971 volume “The Origin and Maintenance of Sex,” John Maynard Smith wrote, “At the cellular level, sex is the opposite of reproduction: in reproduction one cell divides into two, whereas it is the essence of the sexual process that two cells should fuse to form one.”16 These processes underlie the many forms of sexual and asexual reproduction. Members of a sexually reproducing population must produce both sexes, but only half of them—that is, only the females— can produce offspring. But all offspring of an individual from an asexually reproducing population can reproduce. Therefore, any asexually reproducing population will grow at twice the rate of a sexually reproducing population. This, quite simply, is the twofold cost of sexual reproduction. Given the cost of sex, then unless there are other factors, it is hard to see also how sexual reproduction can increase Darwinian fitness.17 But Williams and Maynard Smith each accounted for this exactly twofold cost differently. In 1971, Williams saw it as a result of genome dilution, which was dismissed by the late 1970s as the source of the twofold cost of sex in species with distinct males and females.18 Maynard Smith saw the cost of sex as the cost of producing males. It is a matter of perspective. Mary Jane West-Eberhard puts it this way: “George saw that [an individual female] would profit more by only duplicating her own genes. The extra burden on females is ‘the cost of meiosis.’”19 Williams frames the problem in terms of a dynamic between selfishness and sharing. Imagine a population with two kinds of females. One produces fertilizable eggs, and the other skips meiosis, producing eggs that match its own genetic makeup. All other things being equal, the second one would be twice as well represented in the next generation and, as Williams puts it, “she shares her reproductive success with no one.” So how could meiosis, “in which an individual actively reduces its genetic representation in its own offspring,” continue?20 It would appear, as Williams explains in a metaphor he would later change significantly, that “sexual reproduction is analogous to a roulette game in which the player throws away half his chips at each spin,” a strategy that makes no obvious sense. Such a game would be fair as long as everyone behaves in this way, “but if some do and some don’t, the ones who keep their chips have an overwhelming advantage and will almost certainly win.” This analogy illustrates the first paradox, the paradox resulting from cost. All reproduction is gambling, but who are the players?

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If biologists see sexual reproduction as an adaptation and wish to recognize group-related adaptations in social behavior, then Williams’s paradox must be relevant to the problem—an adaptation for what and to whose benefit? “This sequence of developments will someday be recognized as a curious feature of the history of biological thinking in the twentieth century.”21

What Is the Use or Benefit of Sex? Williams’s motive for writing Sex and Evolution (1975) is revealed first in Group Selection (1971). To understand the function of sex, one must find (1) a reason why sexual reproduction began, (2) a reason why it is maintained, and (3) a reason why it continues to work. In many ways, Williams is also interested in establishing and prolonging a discussion of this problem, for which he is required to show (1) the reason discussion began, (2) the reason it should continue, and (3) why the discussion is still productive. Most immediately, because sexuality seems universal, it is “exactly the sort of thing for which group selection must be postulated.” So, lacking any acceptable explanation of how it could be favored by individual selection, sexual reproduction will continue to offer “a powerful argument in favor of group selection.” To put it another way, if group selection “can produce the machinery of sexual reproduction, it ought to be able to do many other things as well.”22 Williams finds this conclusion difficult to accept, and this is part of his motive for writing Sex and Evolution, though not the entirety of it. Given the cost, “What Use is Sex?” This is the question Maynard Smith asks in another article of 1971. In 1975, Williams uses this question for the first chapter of Sex and Evolution, titling it “An Important Question, Its Easy Answer, and the Consequent Paradox.” He considers a species capable of reproducing sexually and asexually and finds that “sexual reproduction will occur where ecological difference will be greatest between two generations.” This is what he calls “the simple answer”: sexual reproduction can be imagined as “a parental adaptation to the likelihood of offspring having to face changed or uncertain conditions.” Reproduction consists of dispersal through time and space. One must therefore compare the sexual and asexual processes as they emerge in time and space, in terms of the size, frequency, dispersal, development, environment, mortality, and consequently selection of progeny. Yet a conclusion that sex is an

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adaptation to changed conditions is not very satisfying “until we understand why it is adaptive to changed conditions.”23 According to an acute analysis by Joachim Dagg, Williams began to conceptualize the evolutionary cost of sex as the cost of reducing the genome during meiosis, a process of cell division in sexual reproduction that cuts the number of chromosomes from diploid to haploid, or half the original number. Working in 1971 with an argument known as genome dilution—because 50 percent rather than 100 percent of the genome is transmitted in sexual reproduction—Williams, as Dagg puts it, “ends up allowing sociobiological notions of cheating to slip in.”24 By 1973, in his second foray into this controversy, Williams thinks that this 50 percent loss of genetic material and other disadvantages are partially offset by “possible minor advantages in genetically diverse, rather than uniform progenies, but what might be termed the cost of meiosis must be the overwhelming consideration.”25 By 1975, he writes, “The primary task for anyone wishing to show favorable selection of sex is to find a previously unsuspected 50% advantage to balance the 50% cost of meiosis.”26 Dagg concludes, “the 50% reduction of the genome has become a 50% hazard of genes coding for sex (the gene’s eye view).” Williams continues to pursue the cost of meiosis into the 1980s. By 1980, he wonders what an individual in a sexual population would gain by cheating—that is, by eliminating meiosis and fertilization while remaining otherwise the same. In this same article, he speaks of parents “subsidizing the reproduction of the other” and of meiosis as a “sexual game.”27 By 1988, Williams imagines that it is only a matter of time before “someone discovers (or invents in the laboratory) an all-male species” that can inseminate the eggs of a related species. “The point of the story is that any male of any species that refrains from such egg piracy is paying a cost of meiosis as a direct result of the haploid and cytologically cooperative behavior of its sperm.”28 The language of cost, subsidizing, gaming, cheating, and piracy suggests the evolution of Williams’s discourse. This language meets resistance not just from social scientists, as Trivers notes, but also from scientists like Lynn Margulis.29 Hamilton quotes a version of her complaint in Narrow Roads. “The incorporation of ‘cost-benefit’ analysis methods borrowed from insurance practices has led to the biologically puerile numerology,” Margulis argues, writing that “such aseptic language dominates current evolutionary theory” and ignores chemistry and molecular biology. In response, Hamilton supposes that it depends on the researcher’s interest,

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considers those “worthy disciplines” not germane, and comments somewhat obliquely, “Who wants to hear about wood pulp when discussing economics, or chemistry when discussing altruism or sex-ratio strategy?”30 Williams uses the terms cost and benefit frequently. One might almost say that from the beginning he consistently reduces biology to economics or engineering, and his views might be contrasted with those of the environmentalists of his era who wished to subsume economy under ecology.31 Nevertheless, his argument slides toward ecology. Cost-benefit analysis is not exactly game theory. Game theory models social behavior as calculated self-interest, and this way of thinking can arise out of cost-benefit analysis. Yet when Williams refers to game theory in 1966, he seems relatively unimpressed.32 “Nature,” he writes, “is by convention assumed to have no strategy at all in the vernacular sense, but to play at random, that is, without regard for whether she wins or loses.” In game theory an organism’s ecological environment is presented as abstract nature. Surely “environmental ploys are a more effective strategy than would ever be produced at random,” but in a real ecological environment other organisms have effective strategies of their own. “Only a lifeless Earth would really behave as Nature is supposed to in game theory, and a dead body in such a world would last for an immensely long time.”33 In spite of Williams’s skepticism, and probably as a result of his discussions with game theorists like Trivers and Maynard Smith, language associated with game theory abounds in Sex and Evolution. First of all his diction includes terms like cost or cost-benefit relations, cost and profit, cost to the mother, invest or investment, and this terminology flows toward game, game of life, payoff, cheat, cheater, cheating, and cooperation. A few years later, Williams is still wondering about the value of game theory.34 The best way I can define his situation is that he partakes of this discourse while he suspects it. In retrospect, this language may trap him in a narrative. As usual, Williams prefers to ground any discussion of behavior from the lowest level. His discussion of the cost of meiosis leads to a consideration of other physical and chemical costs of reproduction, such as in the production of sperm and eggs, and then the costs of sexual behavior and parental care.35 Williams begins a key chapter—“Why are Males Masculine and Females Feminine and Occasionally, Vice Versa?”—by asserting that these differences follow from the contrast in size between egg and sperm and lead to behavior “resulting from partly conflicting male and female strategies.”36

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The Paradox and the Lottery Though sexual reproduction seems disadvantageous to individuals for a variety of reasons and ought to disappear, this clearly has not happened. “Paradoxically it coexists in evolutionary equilibrium with asexual [reproduction] in many life cycles, in which its occurrence is related to ecological change and uncertainty.”37 If it is not good for the individual, is sex good for the group or species? Remember that R.  A. Fisher, who generally opposed any recognition of biotic adaptation and was a guide for Williams in many matters, regarded sex as “the one exceptional adaptation that was ‘evolved for the specific rather than for the individual advantage.’”38 This second paradox—Fisher’s exception—troubles Williams because it leads to yet another paradox, this one between theory and empirical observation. Astonishingly, Williams writes in 1975, current evolutionary thought seems certain that sex cannot be an “immediate reproductive adaptation in higher organisms”: “Yet this conclusion must surely be wrong. All around us are plant and animal populations with both asexual and sexual reproduction.” This is the third paradox. His approach to this paradox is to “accept the comparative evidence and to try to revise the analysis.”39 In 1975, Williams reconsiders why sexual reproduction must be adaptive, though it seems that it shouldn’t be, by revising his analogy of the roulette wheel into an analogy of the lottery—an analogy now well known among all who study sexual reproduction. He replaces the roulette wheel for reasons of precision. Suppose, he asks, you were offered a choice in a lottery: “either you could have several different tickets, or you could have the same number of copies of the same ticket.” So he begins the second chapter of Sex and Evolution. Of course nobody would buy a large number of lottery tickets that all have the same number on them. It would be as bad as asexual reproduction, producing little or no genetic variety in offspring. In fact, it would be like limiting your chances of winning, which is to say, surviving. He asks the reader to imagine that “sexually produced offspring may be analogous to lottery tickets, and those asexually produced analogous to redundant copies of the same ticket.”40 If asexual reproduction does not produce genetic variation, it gives offspring little ability to adapt quickly to a changing environment. The popularity of Williams’s analogy of the “lottery of survival” is related to its explanatory power. Williams uses the analogy four times and includes it in his index as the “lottery analogy.” He extends it as he writes of the “game of life” with its “prizes.” Though he is often sparing in his

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use of analogies, these terms indicate that something larger is going on, namely Williams’s engagement with game theory.41 His main premise, that sexual reproduction evolved because of the benefits gained by organisms in fluctuating physical environments, is often called the “bet-­hedging” hypothesis or the “tangled bank” hypothesis. After all, Williams’s book is entitled Sex and Evolution, not The Evolution of Sex, and his argument ramifies the implications of sex within ecologies, toward what is now known as biodiversity. The book has thirteen chapters. The early chapters introduce a set of models of sexual behavior; Chap. 6, “Natural Selection in High Fecundity Populations: Theory,” functions as a kind of fulcrum; the following chapters discuss viability and fertility in high- and low-­ fecundity populations, dwelling on patterns of sexuality and sexual behavior; and the book concludes by considering sex as a factor in organic and biotic evolution. As in Adaptation and Natural Selection, an expanded table of contents outlines the argument precisely.

Models of Reproduction: Aphid, Rotifer, Strawberry, Coral, Elm, Oyster In Chaps. 2, 3, and 4 of Sex and Evolution, Williams creates three models by comparing the life histories of what he calls his “authorities”: (1) the “aphid-rotifer model,” (2) the “strawberry-coral model,” and (3) the “elm-oyster model.” I will call these models narratives. The first thing one notices is that his comparative method pairs terrestrial and aquatic species for each model. The behaviors of these species offer three major modes of sexual and asexual reproduction, and that is why they constitute a kind of table of authorities. Each also offers a separate ecological model because, as Williams believes, the advantage of sex is connected with environmental uncertainty. Rotifers, by the way, are bilaterally symmetrical microscopic aquatic animals that some consider wonders of miniature design.42 Williams uses the “aphid-rotifer” model to initiate an explication of his “genetic lottery.” He constructs what has been called a “balance argument” that begins with the assumption that if sex were maladaptive at the individual level, it would disappear from any species that could alternate between sexual and asexual reproduction. In the short term, parthenogenesis is favored when the environment is stable. But when the environment changes, sex provides offspring with greater fitness than parthenogenesis does. Also, sex is favored especially where selection is intense and

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competition between siblings occurs. As Matt Ridley suggests, one can imagine aphids multiplying during the summer on a rosebush and rotifers reproducing in a street puddle. At the end of summer, they change, and the last generation of aphids or rotifers reproduces sexually. This transition would appear to increase their chances in the lottery. Under stable conditions, they produce offspring as fast as possible, asexually, but under uncertain conditions parents produce more costly but diverse offspring.43 Put in more technical terms, sex can be “periodically advantageous” when it maximizes the chances of producing the most locally fit genotypes in a “heterogeneous and unpredictable environment.” Most of the subsequent models Williams introduces reveal the advantage of producing more genetically diverse offspring as a consequence of recombination. “This also applies to the trendy ‘Red Queen’ hypothesis,” as one commentary puts it, “which emphasizes the necessity to evolve fast in a fast-evolving environment.”44 Turning to the second model, the strawberry is a species that Williams finds enlightening and authoritative in itself. There is some humor in comparing its sexual and asexual strategies as if he were consulting an authority on the subject—and so, in a sense, he is.45 Like animals that build coral reefs, strawberry plants are stationary for their whole lives. They produce runners, like coral branches, and this allows an individual plant and its clones to extend into surrounding space. As Ridley explains in The Red Queen, strawberries also produce sexual seeds and corals produce sexual larvae called planulae: “The seeds are carried away by birds; the planulae drift for many days on the ocean currents.” To Williams, Ridley explains, this “looks like a spatial version of the lottery.”46 Those seeds or larvae traveling the farthest would be most likely to encounter different conditions in which the genetic variability of offspring could matter. Why, to turn to the third model, should species like elm trees and oysters disperse millions of sexually produced seeds? Williams first notices that there are few clearings in an elm forest and few vacancies on an oyster bed because elms and oysters have saturated their living space already. Any opening elicits intense competition: “Sex gives variety, so sex makes a few of your offspring exceptional and a few abysmal, whereas asex makes them all average.”47 One notices how wasteful this system of dispersal is, since it is driven by attrition. As in a lottery, so with sexual reproduction—most tickets are worthless.

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High Fecundity and Theory Williams carefully chooses species with life histories nothing like those of humans for his models. His sixth chapter, subtitled “Theory,” argues on a priori grounds “that the intensity of selection in a population has a positive relation to its fecundity.” This is a necessary intervention, he asserts in 1975, because ecological genetics have been biased toward organisms with low fecundity, like humans. “I believe that if evolutionary theorists had not been so preoccupied with low-fecundity organisms with simple life cycles, they would have long ago seen the significance of sexuality.”48 When thinking about species that disperse through many environments, such as starfish and cod, Williams introduces the term “Sisyphean genotypes,” or emerging genotypes that occur with very low frequency in very large populations of sexually reproducing species. Given the extraordinary fitness of these individuals, their direct or indirect descendants will subsequently constitute most of the population.49 In fact, Williams replaces Dobzhansky’s original term for this phenomenon, “genetic elite.” But as Williams’s alternative suggests, the process must repeat itself over and over because recombination breaks down these elite genetic combinations. Sisyphus pushes the boulder up the slope until, “on the verge of reaching the peak, it goes out of control,” and he can only watch it descend again. As Williams puts it, “an individual at the top end of a fitness distribution has achieved its near maximum of fitness by an only momentarily effective combination of genetics and individual history,” and so, in accord with the definition of Sisyphean he finds in his dictionary, the process requires “continuous redoing.”50 It is a long and repetitious uphill battle, and it is likely that Williams chooses the mythical narrative for its tone. Jeffrey Mitton, one of Williams’s coauthors, emphasizes the effect of environment, writing in 1997 that Williams proposed low heritability of fitness from generation to generation because “the fitness of the genotype is highly dependent on the environment in which it was tested, and environments are rarely constant. Because the optimal genotype varies with environmental conditions, natural selection, like Sisyphus, is never done.”51 To understand this phenomenon in 1975, Williams uses his cod-starfish model.52 Sexual species of this type are so mobile that there is no close association of siblings; given their enormous fecundity and high mortality before reproductive age, he further assumes that most individuals must not be viable when faced with a sequence of ecological demands. He asks his reader to forget “the abstract genetic meaning of fitness” and frames

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fitness instead as “an immanent physiological property of an organism in relation to immediate problems.” Imagine that half a population under severe stress will die. Surely there is some genetic difference between the survivors and the dead. Williams compares, for instance, Pleistocene humans and codfish. Man and Cod, if you will. With humans, let’s say, a series of stresses each removing 10 percent of the juveniles would allow half to survive to puberty. As he comments, “it would take 7 such stresses to produce the requisite killing.” Requisite indeed. And leaving the math aside, with the highly fecund cod and requisite deaths, “selection could accomplish in one cod generation what would take perhaps 25 human generations.”53 Death facilitates selection. As T. S. Eliot wrote, “I had not thought death had undone so many.”54 Here, Williams writes, one has a classical Darwinian argument—indeed, an argument that Darwin himself makes in The Origin—about the “struggle for existence.” In this comparison, “the struggle is surely greater in the cod, where one in ten million can survive, than in man, where the odds are about even.” According to this way of thinking about sex and death, more reproduction just means more death. When selection is intense, it works via the massive waste of attrition. Low fecundity does not reduce the strength of selection, but allows the deaths to spread out over a longer time. Further, in all processes of culling, “it would be more correct to say that natural selection is mainly concerned with preventing evolution, not causing it.” What does he mean? Most obviously, natural selection primarily eliminates genotypes with low fitness. Though he does not count these many deaths, he notes that “it takes many deaths to produce an appreciable amount of selective change.”55 This leads directly to Williams’s discussion of “Fitness for the Environment and Fitness of the Environment.” Earlier he has argued that sexual populations are more likely to survive changed conditions than asexual ones, but for reasons different than those commonly assumed. He finds internal and external forces working in parallel here. First, in marine fishes like cod great differences in survival are caused by environmental variation, and these variations are so extreme that in some years all zygotes are likely to die when faced with a changed environment. “If ordinary environmental variation can produce such widely different rates of survival,” he asks, “is it not reasonable to suppose that ordinary genotypic variation can do the same?” Survival, in any case, may depend upon the Sisyphean tail end of a genotypic distribution within a population. In fact, he finds contributions to fitness are equally efficacious whether they come

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from genotype or environment. For cod or elms, the size of the next generation will be “almost independent of present numbers or anything else in recent history and almost entirely determined by the current size and suitability of habitat.” Further, in cod there is a high degree of independence between the size of spawning stock and the numbers that will survive from a given spawning. Consequently, “a given environment will select its Sisyphean few from the vast pool of available genotypes.” Not only should genetic changes among the inhabitants of the changing environment track environmental fluctuations closely, but also “the amplitude of genetic fluctuation should parallel that of environmental fluctuation.”56 Analogies with the story of Sisyphus or with a lottery cannot create a complete theory, though Williams’s lottery has reappeared over the years under many names with many twists. One must not lose sight of the larger shape of Williams’s argument. And it is the shape of his argument I am interested in, not its correctness, in part because, as Stearns showed in 1985, other comparative evidence “does effectively rule out the idea that sex is primarily an adaptation to unpredictable environments.”57 It is difficult to generalize about fecundity because the fertility of eggs and seeds is widely variable. In statistical terms, fitness in high-fecundity populations can be approximated by what Williams calls distributions of high variation, plotted logarithmically, where only a few Sisyphean genotypes contribute to the next generation. A lognormal distribution is defined as a continuous probability; logarithmic distribution of a random variable like fitness can be graphed as a standard bell-curve with its tails. The effect of this distribution overshadows the cost of meiosis, which can only be “a minor part of a commonly manifold difference in fitness among genotypes.” Consequently, the benefits of genetic diversity outweigh the advantages of parthenogenesis. But this situation would seem to be reversed in low-fecundity populations. “If and when any form of asexual reproduction becomes feasible in higher vertebrates, it completely replaces sexual.” Exclusive reliance on sexual reproduction must be inherited from some high-fecundity ancestor and, “in these forms sexuality must be a maladaptive feature” that cannot be removed.58 Williams begins to write of maladaptation as a trap laid by history in Sex and Evolution, and it becomes a major theme in his later works, including the 1996 Preface to Adaptation and Natural Selection. Why does Williams call sexual reproduction maladaptive? According to the OED, the term maladaptation appears in English usage in 1877, where “silly

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maladaptations” are imagined as “steps toward improvement or discarding of imperfect organs.” This is the opposite of what Williams means. He means the retention of an irremovable, imperfect mechanism. There are obstacles to any transition from sexual to asexual reproduction. What might be adaptive for the embryo, like reproduction by fission, twinning, and such, is not adaptive to the mother who has already paid the price of meiosis. And “the conflict of generations seems generally to be resolved in favor of the mother, for reasons that are not immediately apparent.” Also, this kind of transition requires multiple cellular changes to occur “simultaneously and completely.” Yet when parthenogenesis is possible, sexual reproduction is “immediately lost in any vertebrate population.” This, in turn, must give “decisive evidence of the maladaptive nature of sexuality in these organisms.”59 There are many patterns of sexuality. Many commentators, as Williams notes, just assume that asexual reproduction has historical priority and speak of primitive sexuality as if the simple way would come first. He imagines a narrative that goes like this: bacteria under promising conditions would have no reason not to reproduce the same genotype, but a gene will be selected because it maintains precision of cloning under optimal conditions but permits recombination when conditions are not optimal, so that “as soon as cloning was possible in the history of life, sexual reproduction would be confined to times of change or stress.”60 The terminology becomes elaborate because the patterns of sex and asex are so various. Key sexual patterns include isogamy, found in some algae and other protist groups, a pattern characterized by the fusion of gametes that are not distinct in size or structure, and anisogamy, a union between two gametes that differ in size or form. A gamete, we remember, is a haploid reproductive cell, and two unite to make a new cell called a zygote. Differences in their size or form have large consequences. Williams explores these consequences in “Comparative Sexuality of Higher Organisms,” where he begins by chiding the early Darwinists for not seeing that “the organization of life cycle is a product of natural selection.” As he insists, evolutionary biology must be devoted to explaining diversity not only of anatomy, but also of life cycles. Certainly “various modifications of sexual reproduction form a key element in the organization of any life cycle,” and life cycles are enormously diverse. A naturalist notices that plants are rarely dioecious—etymologically, “from two houses,” meaning species with distinct individuals that produce either male or female gametes—while this is common for animals. In contrast, it is

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common for plants to be monoecious, where one individual produces both male and female gametes, while animals are rarely hermaphroditic. R. A. Fisher, as Williams remembers, found that there can be no advantage of one sex over the other in dioecious organisms, and this has implications in terms of parental investment. Williams follows those implications to the whole population, where male and female functions must be equally expensive for hermaphroditic plants, whose individuals must spend equal resources on pollen and on ovules. Mind you, “there is nothing to suggest that male functions are especially cheap.” It is precisely the wide variation of reproductive patterns that makes it difficult to find any optimized strategy for them.61

Low Fecundity and High Conflict Though Williams puts it as a question—“Why are Males Masculine and Females Feminine and Occasionally, Vice-Versa?”—one thing is certain and indeed almost a cliché for the more familiar higher animals: “The essential difference between the sexes is that females produce large immobile gametes and males produce small mobile ones.” Big eggs; small sperm. If the ultimate goal for both is “maximal genetic representation in the same population,” as it must be, their goals in reproduction seem different, to judge by their behavior. This contrast provides “a prima facie difficulty for evolutionary theory.” These strategies, Williams argues, must emerge from the difference in gamete size and lead him to scrutinize three phases of the sexual comedy: gamete production, achievement of fertilization, and care for offspring. One must have a sense of humor about what follows, for it is not possible to read it all with a straight face. For example, males “take a more active role in courtship, are less discriminating in choice of mates, more inclined toward promiscuity and polygamy, and more contentious among themselves.”62 In sessile or spatially fixed marine mammals, Williams finds that reproduction requires little beyond production and release of gametes and, consequently, “a minimum of contrasting masculinity and femininity.” Mobile animals, in contrast, “achieve fertilization by close coupling and consequent reduction of sperm wastage,” as he puts it. If you compare sizes of gonads (ovaries and testicles), you find that they relate to this process. For mammals, and to a lesser extent for birds, costs borne by females come almost entirely after fertilization, as every mother has told her daughter. David Lack demonstrated how female fertility “may be well below that

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which is physiologically achievable.”63 Fertilization requires synchronization, timing in release of gametes, and cues. These cues are paramount for “sexual selection,” defining the set of criteria first introduced by Darwin, upon which mates choose each other.64 As Williams summarizes, “mate quality would seldom be as important to a male as their number,” and monogamous males should prefer larger, younger females because they offer maximum reproductive value over the long term. As in big female fish. In the midst of this, Williams makes an astounding leap to human behavior, to what men and women want. This is the only place in his book where the word woman or women appears. Though he finds no data on the subject, his impression that a “strong male preference for women who are sexually mature but youthful is universal in all cultures, perhaps more so than any other standard of beauty”—upon which so much biological and cultural advertising is based, one supposes. A male’s age is less important for female choice, he continues. He qualifies only slightly, arguing that individual variation in sexual preference indicates the “great plasticity of human behavior, but in no way impugns the evolutionary significance of the norms.”65 And so evolutionary psychology was born. Williams backed away from this theme, perhaps because he knew it was a dangerous misstep, before the feminists attacked, and there was good reason for them to do so. The year 1975 was when Susan Brownmiller published Against Our Will. On the other side of the sexual divide, “mate quality rather than number must be the dominant consideration in female strategy.” In some fishes, for instance, “conspecific matings are the rule only because females reject males of all but their own species,” whereas some male fish seem indiscriminate in this matter and require a long period of learning. Williams credits Trivers (1972) with recognizing this difference, where males try to maximize quantity of fertilization while females try to optimize its quality. For this reason, Williams approves Trivers’s view that “the seemingly cooperative aspects of family life may be misleading.” Trivers is famous for relating his findings directly to human behavior and speaking of adultery, desertion, and other kinds of cheating. As Trivers points out, courtship and related behavior in birds can be seen as “attempts to preserve one’s options and restrict those of the mate.” Why not humans too? Williams revises his earlier concurrence with the common view that female discrimination favors the genetically fittest male as expressed by his phenotype. “I now feel that this would require an unrealistically high heritability of fitness.” Instead, he decides, “the important female adaptation in relation to

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courtship is an ability to predict future resources for her offspring from the appearance and circumstances of the courting male.” Her criteria include considerations of territory.66 All of this stereotyping can be verified with contradictions, by comparison to reversed strategies. He cites egg holding by male seahorse-pipefish as an example and predicts at large that “behavioral masculinity and femininity will loosely parallel variation in parental burdens of male and female.” Cost and benefit associated with sexual strategy cannot be attributed to the species but to the individuals involved. He considers this “a choice illustration of the logic of the comparative method,” where “it is the exception to the rule of masculine males and feminine females that proves the theory that explains both rule and exceptions.”67 Williams supposes that with external fertilization, “kin selection may promote the evolution of parental care of the young,” when male territoriality and external fertilization “preadapts a species to parental care by the male.” Some male fish tend and protect large numbers of eggs in their territories. In cichlids, both sexes are territorial and that leads to sharing of care. With internal fertilization, as in land mammals, the female is associated with the young for a longer period of time, and one sees maternal behavior; if she remains in the male’s territory, “care of the young by both parents may evolve.” At fertilization males have invested little more than gametes, but Williams thinks some male fish may continue their investment because a suitable hard-to-win territory is a long-term resource when held jointly with a cooperative female. Males die younger than females, and following Trivers’s argument, Williams writes that “the sex with the greater variance in fitness should have the greater mortality,” for there is a wide range in the reproductive success of individual males, with many not reproducing at all, while the most successful reproduce many times.68 This suggests that patriarchs live longer than bachelors. The analogy of the lottery cuts both ways, as Trivers’s language in Williams’s mouth allows. “At every moment in its game of life the masculine sex is playing for higher stakes.” This is true in terms of possible reproductive “winnings” and the “ultimate empire of wives and kind,” while there are also “possibilities for immediate bankruptcy (death)” and “permanent insolvency,” as in celibacy. Males experience “greater susceptibility to psychosomatic damage,” are “less tolerant to environmental stress,” and “must gamble at odds in the effort to attain the upper tail of the fitness functions,” while females need only to avoid malfunction.69

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The Nirvana of Parthenogenesis The most important behavior of any organism is reproductive behavior. Consequently, it is fitting that this remains a great mystery in its origin, maintenance, and variety of strategies. Hamilton believed it to be “biology’s most outstanding problem” when he reviewed Sex and Evolution in 1975, and he reflected on his own review in 1996. As he remembered, his thoughts ran on lines fairly parallel to those of Williams, “whose evolutionary ideas have always seemed convergent like those of a twin or older brother from whom I was parted at birth.”70 In terms of human reproduction, twinning itself would appear to be an interesting puzzle. Williams notices elsewhere that “twins are usual for bears, and even this litter size may form a kind of evolutionary trap from which bear species are unlikely to escape,” but that “human twinning was maladaptive in advanced societies as late as the eighteenth century” due to high mortality for mother and children.71 Which is to say that Hamilton’s comment about his brotherliness with Williams constitutes a complex bit of wordplay, troubling and a possible trap, though also meant quite sincerely. Like every biological performance, reproductive strategies, in all their strangeness and awkwardness, in all their embarrassing glory, must have evolved and must continue to do so. They must surely be adaptations of some sort, but of what sort is puzzling. Humans, who have sex on their minds a good deal of the time, find the most important of their own behaviors to be sexual, but do not like to think of them as biological. For other creatures, sex is not on the mind. For some, as Williams shows, reproduction is sometimes sexual and sometimes not sexual, depending on the occasion. So too the offer of parthenogenesis, or virgin birth, enters the purview of Hamilton and Williams. As Hamilton writes in his 1975 review of Sex and Evolution, the “silly rigmarole of sex” is troublesome and constitutes a set of “intellectual antlers” that offer to sink humans in a bog. Williams believed this rigmarole might offer to disappear, and as Hamilton writes, “Williams seems to have proved in his book that man’s extremely low fecundity gives him no evolutionary strength to object to or withstand this offer when it comes.”72 The offer is not coming soon.

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Math and Sex Sex is also a mathematical puzzle, especially for Hamilton and Maynard Smith. Math is in many ways a mode for distancing the observer from the observed, and therefore safer than suppositions about behavior rendered in prose. Comparative evidence allows Williams to announce at the outset, “my reasoning is much less rigorous than Maynard Smith’s thorough mathematical treatment, but I think I have the more reliable conclusion. For answering questions on function in biology, comparative evidence is more reliable than mathematical reasoning.” Williams jokes, “fortunately, it is easier to reason from a set of premises to a valid conclusion when you know the conclusion in advance.”73 He means that one can see the success of sexual reproduction all around and, by comparing various sexual strategies, one can construct various mathematical designs. Because it involves division and multiplication, the very language of reproduction is mathematical. Surely it is necessary to understand reproduction mathematically, but is mathematical understanding sufficient? Williams is not immune to mathematical reasoning—and he certainly does not ignore it in the work of Hamilton, Maynard Smith, or Trivers. He tallies his own debts to the mathematical reasoning of R.  A. Fisher, J. B. S. Haldane, and Sewall Wright in 1966, speaking of the gene as a cybernetic abstraction and a phenotype’s fitness as a statistical abstraction. Williams also argues that the study of natural selection “has little appeal to those who are not in the regular habit of using mathematical abstractions in their thinking about organisms.”74 This constitutes a certain distancing by which a biologist puts himself above the non-mathematical reader. Maynard Smith remembered Haldane saying, “if you are faced by a difficulty or a controversy in science, an ounce of algebra is worth a ton of verbal argument.”75 But that is not quite right. Mathematics is a rhetoric with its own way of creating “predictive narratives,” as Williams called them. These narratives take a form something like this: If one variable like the size of gametes is important and has a relation with another variable like the costs and benefits of parental care, then the result can and should be transformed into an expression of probability revealing the frequency or likelihood of a particular constellation of behavior. Mathematics is one mode of expression available to a biologist. The philosopher Samir Okasha reminds us that when Fisher, Haldane, and Wright created the field of population genetics in the 1920s and 1930s, their work paved the way for the modern synthesis. When they integrated the principles of Mendelian

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genetics with Darwinian natural selection, they approached the task by manipulating cybernetic and statistical abstractions.76 Since then, as evolutionary biology theorizes about populations, its principles are often best expressed by extensive mathematical manipulation. Population geneticists consider changes in the frequencies of alleles at a small number of gene loci; Fisher and Wright derived quantitative genetics from infinitesimal changes at a large number of gene loci. Maynard Smith was Haldane’s student. W. D. Hamilton and Williams were heavily influenced by Fisher’s calculations and followed them into behavior. How could reproduction not be a matter of math? Sex is math, or sums, as Maynard Smith preferred to say. Division and multiplication. Just as sexual reproduction manipulates genetic material, so too mathematics manipulates information. Mathematics models phenomena and in this way embodies biological relationships. Often in modern mathematical biology, a biologist works with a mathematician. This was not so much the case for Fisher, Haldane, Wright, Hamilton, or Maynard Smith, who were their own mathematicians. Williams remembers that when he was a young biologist, “few people read such mathematical discussions as Haldane’s and Fisher’s. We got our theory in words from Dobzhansky, Mayr, and Simpson, who were deeply concerned with natural selection, but gave little attention to its levels of operation.”77 Why? Dobzhansky confessed to a lack of mathematical skills. Williams knew the math and followed Fisher, but did not write like a mathematician. Hamilton admits, in commentary written for his collected papers in 1996, that when he began to write of senescence, “I had found I had forerunners in the theory of senescence in P. B. Medawar and G. C. Williams, and that what I had been doing was little more than to put a mathematical gloss to their ideas,” but math was necessary precisely because they “had not done a very complete job.”78 Consider also the truism that professional papers are more frequently cited than read, and in particular “that Hamilton’s 1964 paper is much more frequently cited than read.”79 Alan Grafen, a professor of theoretical biology and Hamilton’s close colleague at Oxford, thinks that generations of biologists learned about inclusive fitness from secondary sources and that Williams was the “prime secondary source.” Williams believed, as his writing demonstrates, that sense and nonsense can be measured and distinguished in various ways, and that one of the best ways to do so is via clear prose. The person who writes convincingly changes minds.

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Mathematics has become a road to clarity for most but not all modern biologists. Alan Grafen, for instance, thinks of Williams as eminent among “verbal and conceptual thinkers.”80 I once engaged Steven A.  Frank, a biologist at the University of California, Irvine, in a short email conversation about Williams. He studies evolutionary genetics, host-parasite interactions, and social evolution. Frank’s website allows open access to all his research.81 He develops “mathematical, computational, and conceptual models to study complex phenotypes.” He reminisces that early in his career when he studied “how evolutionary and genetic processes shape reproductive and behavioral traits,” “It was necessary at that time to treat as a black box many of the genetical and physiological details that determine phenotypes, and to focus in a general way on how natural selection influences phenotypes over very broad assumptions about underlying mechanisms,” but now it is possible to see “below the surface of complex phenotypes to the biochemical and genetical mechanisms that control those characters.”82 I stated that it was my sense that Williams took it as a primary task “to translate what is now called mathematical biology into prose,” to which he replied that Williams’s aim was “Perhaps: to express concepts in a clear and direct manner. The word ‘mathematical’ puts too much emphasis on math instead of concepts.” In answer to my sense that his articulation of principles of adaptation without recourse to mathematical reasoning was perhaps no longer possible, Frank answered that “Perhaps he used the tools over which he had greatest command and which felt natural to him.”83 He considered this a matter of preference.

Williams Reviews Maynard Smith Three years after he published Sex and Evolution, in a review of John Maynard Smith’s The Evolution of Sex (1978), Williams revealed what he thought of his own work. He was convinced that Maynard Smith’s book was important, yet—as he might have said of his own book—“I believe many of its proposals may fail the test of time. Its lasting value will lie not so much in the validity of answers suggested as in questions formulated and confronted.” After listing some of those questions, Williams expresses on a rather personal note, “my enthusiasm for this small volume may be based partly on its recognition of my role in developing some of the ideas and on its consistently charitable treatment of such earlier groping.” He also confesses that his enthusiasm “is based even more on the realization

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that it makes my own work … so nearly obsolete.” Then he recites a list of his own confusions that required correction, some of which “I now wonder why it took me so long to appreciate.”84 After praise, he makes two important statements. “I still favor almost entirely environmental causes of extinction and evolution despite Maynard Smith’s criticism of this position.” More globally, he criticizes Maynard Smith’s concept of an evolutionary stable strategy (ESS), a part of Maynard Smith’s contribution to game theory. An ESS is a strategy that, if adopted by a population of players, cannot be replaced by any alternative strategy that is initially rare. Williams objects to the limitations of what he calls Maynard Smith’s economic argument, saying that the use of the concept of ESS “shows an unfortunate disregard for the limitations of this form of reasoning.” This critique is somewhat peculiar, since Williams himself makes an economic argument. He is still troubled and believes that “the seemingly discriminating mate choice by females of species in which males do nothing for the offspring is a serious (fatal) difficulty for the whole theory of sexual selection.” It is, Williams concludes, “a field of controversy and conceptual flux,” and as one might argue, he continues his role, even in this review, of articulating, formulating, and confronting serious questions.85 About one thing they certainly agreed—namely that, as Maynard Smith would argue near the end of the twentieth century, “A central idea in contemporary biology is that of information.” Developmental biology allowed biologists to study “how information in the genome is translated into adult structure,” and evolutionary biology allowed them to study “how the information came to be there in the first place.” Yet he still insisted, as Williams always did, that “differences due to nature are likely to be inherited whereas those due to nurture are not; evolutionary changes are changes in nature, not nurture.”86 Sexual behavior in humans may be molded by nurture, but is grounded in nature, they both believed, and this is the essence of their sociobiology.

Lottery How is reproduction like a lottery? It is called reproduction because it happens over and over, pretty much in the same ways and pretty much with the same results—or so a naïve human might think. There is also the cultural reproduction of arguments, sometimes with difference.

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Williams is characteristically modest in making his argument and in crediting himself with any final success. As evidence of uncertainty, one notices that he continues to rephrase his premise in Sex and Evolution. For instance, toward the end, he writes almost hesitantly that variety of genotype “provides a margin of safety against environmental uncertainty” and “may be more important to population survival than is precise adaptation to current conditions.” Citing three theories about genotypes faced with environmental uncertainty and consequent extinction, he asserts that “the facts of biology … offer little encouragement on any of these three theories.” He is not satisfied with his argument—or analogy of the lottery— but says, “I would perhaps claim that the information is somewhat less embarrassing to my own theory than for either of the others.” By the end of the book he can only say, in characteristic Williams fashion, “I am sure that many readers have already concluded that I really do not understand the role of sex in either organic or biotic evolution. At least I can claim, on the basis of the conflicting views in recent literature, the consolation of abundant company.”87 Such a statement is not mere humility. It is meant to stimulate further thought. Provoked by Williams, Hamilton worked on the problem for some twenty years, from the mid-1970s until the end of his life.

Sex as a Factor in Biotic and Organic Evolution Sex and Evolution concludes with a parting shot on the efficacy or importance of organic over biotic mechanisms of evolution. Williams reminds his reader of the basic and important role of sex in evolution, formally separable from the role of sex as a character shaped by selection. Ever the contrarian, he quotes the published views of Mayr and Dobzhansky, that “ample genotype variety may be a safeguard against extinction,” and he agrees, but his reasons are the opposite of theirs. Though they speak for many other evolutionary thinkers, claiming that the process of recombination allows organisms to respond to selective pressure and adapt more quickly, Williams argues in his concluding chapters that (1) “sexual reproduction usually opposes the effect of selection,” and, (2) by retarding adaptive evolution, “may produce long-term group advantage.”88 This is an argument about rates of change. Obviously, organic evolution is change in the genetic constitution of a population. But the rate of change is sometimes mysterious. For instance, “despite the reasons why man’s evolution ought to be slow—small populations, low mutation rate,

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long generation rate—he evolved with unusual rapidity.” Williams finds that textbooks mislead their readers by showing “factors capable of producing sufficiently rapid change,” when they instead ought to show “why populations can so effectively resist change over such amazingly long periods.”89 We remember too that biotic evolution is change in the composition of a biota. Extinction, an absolute form of change, “occurs not because an organism loses its adaptation to an ecological niche, but because its niche becomes untenable.” To translate this into contemporary terms, there is no point in speaking of biodiversity unless you know how it is produced, by global origins and extinctions during eons of geological time. In this flux of time, the variety of genotypes provides what Williams calls “a margin of safety against environmental uncertainty” that may be more important for a population’s survival than “precise adaptation to current conditions.” Yet he finds “little respectable evidence that extinction has any consistent biases” because “consistently recognizable causes of extinction are seldom, perhaps never demonstrable.”90 There is no force driving extinction. Williams views the usual textbook explanations as “part of the folklore of scientists, not science.” He avoids the problem of anthropogenic causes of extinction that would be discussed a decade later in a national forum.91 In this context, Williams once again brings up group selection. As he insists, “profitable discussion of group selection must relate to the question of importance, not reality,” or to put it more dramatically, “The problem is not: Is it there? But rather: Has it produced important effects?” A population might, for instance, “adapt more and more perfectly to a disappearing habitat,” and this phenomenon, the disappearance of an ecological niche, goes beyond “destruction of a bit of geography.”92 Further, it is a tautology to suggest that various groups became extinct because they were overspecialized or had “inferior” structures, like the wings of pterodactyls or the naked seeds of gymnosperms such as conifers. “This sort of speculation is empty of scientific meaning,” not only because of questionable judgments about the inferiority or superiority of adaptations, but also because it is impossible to assess their roles in any extinction. “Extinction occurs because there is not corrective feedback between dangerously low population size and forces of evolution.” There is no foresight, correction, or adjustment possible; Williams quotes Marjorie Grene, who writes of a species doomed to extinction, and unconscious of its fitness, as it “continues happily to perform its traditional rites.”93 In this

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sense, a species is powerless to act on its own behalf, and further, whatever sexual strategy it exhibits, though part of evolutionary change, offers no hedge against ultimate failure.

Sexual Humans Haldane’s quip—“Would I lay down my life to save my brother? No, but I would to save two brothers or eight cousins”—does not map the boundary of human responsibility, nor does it explain what it means to save a brother, cousin, or any living being. Humans choose their acts. Humans also choose to think about the world in certain ways. They accommodate themselves to the symmetries and asymmetries of the world and sometimes call them beautiful, or even, as with human sexuality or sexual attraction, a masterpiece. Yet humans also sterilize most domestic animals and pets and control their own sexual reproduction. Sex is an embattled territory, and one finds a great many writers on human sexuality appropriating authority from the writings of George C. Williams. Sometimes the appropriations are deeply troubling. In a disturbing instance, Randy Thornhill and Craig T. Palmer argue that ideologies associated with human reproduction are rigid, complicated, extraordinarily rules-oriented, powerful—and yet surely a veneer over something biological and perhaps pathological.94 This is why, they assert, you cannot stop rape by shouting, “Just say no!” Their argument appears in the preface to their controversial book The Natural History of Rape, entitled “Rape and Evolution: A Reply to Our Critics.” This idea, that culture is never more than a veneer over biology, is pervasive among sociobiologists. But what can you do if you refuse to see biological forces as sources of sexual behavior? One thing is certain, sex makes humans what they are, and makes humans do many things that seem reprehensible. Students of Williams like Bobbi Low can write, simply, that “Sex Matters.” More radically, another, Laura Betzig, writes, “I, personally, find ‘culture’ unnecessary.”95 Tellingly, Bobbi Low appropriates Hamilton’s use of an interchange between the characters Temple and Stephen Daedalus in James Joyce’s Portrait of the Artist as a Young Man.96 Hamilton places this passage as an epigraph to his personal introduction for “The Moulding of Senescence by Natural Selection.” As Joyce wrote, “The most profound sentence ever written, Temple said with enthusiasm, is the sentence at the end of zoology. Reproduction is the beginning of death.” And then he asks Stephen,

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“Do you feel how profound that is because you are a poet?” It seems that Stephen does not feel its profundity, but Temple’s nexus is profound enough to be worth exploring.

Notes 1. George C.  Williams, Adaptation and Natural Selection (Princeton, NJ: Princeton University Press, 1966), 125. 2. Williams, Adaptation, 54. 3. Paul Schmid-Hempel, “Wondering About Sex: W.  D. Hamilton’s Contribution to Explaining Nature’s Masterpiece,” Behavioral Ecology 12, no. 3 (2001): 266–68. http://beheco.oxfordjournals.org/content/12/ 3/266.full. 4. George C.  Williams, Sex and Evolution (Princeton, NJ: Princeton University Press, 1975), v. 5. Williams, Sex, 106. 6. George C. Williams, editor and author of “Introduction” and headnotes, Group Selection (Chicago: Aldine-Atherton, 1971), 12. 7. Williams, Group Selection, v. 8. Laura Betzig, “Laura Betzig on George Williams,” The Evolution and Medicine Review (September 15, 2010), http://evmedreview. com/?p=441. 9. Michel Foucault, “The Order of Discourse,” in Untying the Text: A Post-­ structural Anthology, edited by Robert J.  C. Young, 48–78 (Boston: Routledge & Kegan Paul, 1981). 10. Robert Trivers’s most important early papers are: Trivers, “The Evolution of Reciprocal Altruism,” the Quarterly Review of Biology 46 (1971): 35–57; Trivers, “Parental Investment and Sexual Selection,” in Sexual Selection and the Descent of Man, 1871–1971, edited by B.  Campbell, 136–79 (Aldine-­ Atherton, Chicago, 1972); Trivers and D.  Willard, “Natural Selection of Parental Ability to Vary the Sex Ratio of Offspring,” Science 179 (1973): 90–92; Trivers, “Parent-offspring Conflict,” American Zoologist 14 (1974): 247–62. 11. Robert Trivers, Natural Selection and Social Theory: Selected Papers of Robert Trivers (New York: Oxford University Press, 2002): 6. 12. Trivers, Natural Selection, 23, 20. 13. Trivers, Natural Selection, 61. 14. Robert Axelrod and William Hamilton, “The Evolution of Cooperation,” Science, New Series 211, no. 4489 (March 27, 1981): 1390–96. 15. See Stuart A. West, A. S. Griffin, and Andy Gardner, “Social Semantics: Altruism, Cooperation, Mutualism, Strong Reciprocity and Group

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Selection,” Journal of Evolutionary Biology 20, no. 2 (2006): 415–32. Also see Steven Kuhn, “Prisoner’s Dilemma,” The Stanford Encyclopedia of Philosophy (Spring 2009 Edition), edited by Edward N.  Zalta, http:// plato.stanford.edu/archives/spr2009/entries/prisoner-­dilemma/. The standard introduction is Robert Axelrod, The Evolution of Cooperation (New York: Basic Books, 1984). For a readable introduction, see William Poundstone, Prisoner’s Dilemma (New York: Anchor, 1993). 16. John Maynard Smith, “The Origin and Maintenance of Sex,” in Group Selection, edited by George C.  Williams (Chicago: Aldine-Atherton, 1971), 163. 17. Joachim Dagg, “The Paradox of Sexual Reproduction and the Levels of Selection: Can Sociobiology Shed a Light?” Philosophy & Theory in Biology 4 (2012): 1–16. 18. Yet one still encounters this explanation in the literature; see Jussi Lehtonen, Michael D. Jennions, and Hanna Kokko, “The Many Costs of Sex,” Trends in Ecology & Evolution 27, no. 3. (2011): 172–78, 172. 19. Mary Jane West-Eberhard, “The Maintenance of Sex as a Developmental Trap due to Sexual Selection,” the Quarterly Review of Biology 80, no. 1 (2005): 47–53, 48. 20. Williams, Group Selection, 13. 21. Williams, Group Selection, 13, 14. 22. Williams, Group Selection, 13, 161. 23. Williams, Sex, 3, 7. 24. Dagg, “Paradox of Sexual Reproduction,” 4, 5. 25. George C. Williams and J. B. Mitton, “Why Reproduce Sexually?” Journal of Theoretical Biology 39 (1973): 545–54. 26. Williams, Sex, 9–10. 27. George C.  Williams, “Kin Selection and the Paradox of Sexuality,” in Sociobiology: Beyond Nature / Nurture, edited by George W. Barlow and James Silverberg (Boulder, CO: Westview Press, 1980), 371, 372, 377. 28. George C.  Williams, “Retrospect on Sex and Kindred Topics,” in The Evolution of Sex, edited by R.  E. Michod and B.  R. Levin (Sunderland, MA: Sinauer, 1988), 287–98, 294. 29. See comments in Lynn Margulis, and René Fester, eds., Symbiosis as a Source of Evolutionary Innovation: Speciation and Morphogenesis (Cambridge, MA: MIT Press, 1991), 6. 30. William Hamilton, Narrow Roads of Gene Land: The Collected Papers of W. D. Hamilton, vol. 1 (New York: W. H. Freeman, 1996), 267, n. 11. 31. Consider, for example, the discourse of Gary Snyder in Earth House Hold (New York: New Directions, 1969). 32. See Williams, Adaptation, 67–70. In 1966, he cites in his references Richard Lewontin, “Evolution and the Theory of Games,” Journal of

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Theoretical Biology 1 (1961): 382–403. For a definitive review of evolution and game theory, see John Maynard Smith, Evolution and the Theory of Games (Cambridge University Press, 1982). Also see J.  McKenzie Alexander, “Evolutionary Game Theory,” in The Stanford Encyclopedia of Philosophy (Fall 2009 Edition), edited by Edward N. Zalta, http://plato. stanford.edu/archives/fall2009/entries/game-­evolutionary/. 33. Williams, Adaptation, 68. 34. George C.  Williams, “When Does Game Theory Model Reality?” Behavioral and Brain Sciences 7, no. 1 (1984): 117. 35. Williams, Sex, 9, 10, 101, 136, 114, 126. 36. Williams, Sex, ix. 37. Williams, Sex, vii. 38. See Williams, “Adaptations of the Genetic System,” Chapter 5  in Adaptation and Natural Selection (1966), 157. R. A. Fisher, The Genetical Theory of Natural Selection. A Complete Variorum Edition, edited by J. H. Bennett (Oxford: Oxford University Press. 1999), 280, an addendum of 1958. 39. Williams, Sex, 11, 12. 40. Williams, Sex, 15. 41. Williams, Sex, 67, 138, 53. An initial step into evolutionary game theory is John Maynard Smith and George R. Price, “The Logic of Animal Conflict,” Nature 246 (1973): 15–18. 42. Williams, Sex, 26–43. For rotifers, see the webpages of Wim van Egmond: http://www.microscopy-­uk.org.uk/mag/indexmag.html?; http://www. microscopy-­uk.org.uk/mag/wimsmall/rotidr.html. 43. Matt Ridley, Red Queen: Sex and the Evolution of Human Nature (New York: Penguin, 1993), 56. 44. C.  Rispe et  al., “Models of Sexual and Asexual Coexistence in Aphids Based on Constraints,” Journal of Evolutionary Biology 11, no. 6 (1998): 685–701, 686. 45. Williams, Sex, 26–43. See also West-Eberhard, “Maintenance of Sex,” 51. 46. Ridley, Red Queen, 58. 47. Ridley, Red Queen, 58. 48. Williams, Sex, 62, 63. 49. E. O. Wilson picks up this term in 1978, but when he applies it to humans he misunderstands Williams’s model. See Wilson, On Human Nature (Cambridge, MA: Harvard University Press, 1978), 168. 50. Williams, Sex, 45–46. 51. See Jeffrey Mitton, Selection in Natural Populations (Oxford: Oxford University Press, 1997), 157–66. 52. Williams, Sex, 44, 45, 52, 59, 61, 83. 53. Williams, Sex, 63, 64.

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54. T. S. Eliot, The Waste Land (1922), line 63. 55. Williams, Sex, 64, 65, 67. 56. Williams, Sex, 68, 69, 72, 73, 75, 76. 57. Stephen C.  Stearns, “The Evolution of Sex and the Role of Sex in Evolution,” Cellular and Molecular Life Sciences 41, no. 10 (1985): 1231–35, 1233. 58. Williams, Sex, 101, 102–103. 59. Williams, Sex, 103, 105, 106. 60. Williams, Sex, 111, 112. Williams prefers to classify modes of reproduction into recombinational and nonrecombinational, rather than “the more facile terms sexual and asexual” (114). I find the facile terms easier to use. 61. Williams, Sex, 117, 119, 120, 121, 122. 62. Williams, Sex, 124. 63. Williams, Sex, 125, 126, 127. 64. Helena Cronin, The Ant and the Peacock: Altruism and Sexual Selection from Darwin to Today (Cambridge, UK: Cambridge University Press, 1993). 65. Williams, Sex, 128, 129. 66. Williams, Sex, 129, 130. 67. Williams, Sex, 133, 134. 68. Williams, Sex, 135, 137, 138. 69. Williams, Sex, 138, 139. 70. William Hamilton’s retrospective introduction to his review of Sex and Evolution, “Gamblers Since Life Began: Barnacles, Aphids, Elms,” the Quarterly Review of Biology 50, no. 2 (1975): 175–80, is found in Hamilton, Narrow Roads, vol. 1, 354, 357. 71. George C.  Williams, Natural Selection: Domains, Levels, and Challenges (New York: Oxford University Press, 1992), 79. 72. Hamilton, Narrow Roads, vol. 1, 367. 73. Williams, Sex, 7. 74. Williams, Adaptation, 33. 75. John Maynard Smith’s obituary for J. B. S. Haldane is in Nature 206, no. 4981 (1965): 239–240, 239. 76. Samir Okasha, Evolution and the Levels of Selection (Oxford, UK: Clarendon Press, 2006). See also Samir Okasha, “Population Genetics,” The Stanford Encyclopedia of Philosophy (Fall 2008 Edition), edited by Edward N. Zalta, http://plato.stanford.edu/archives/fall2008/entries/ population-­genetics/. 77. Williams, Natural Selection, 47. 78. Hamilton, Narrow Roads, vol. 1, 89.

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79. Alan Grafen, “William Donald Hamilton 1 August 1936–7 March 2000,” Biographical Memoirs of Fellows of Royal Society London, 50 (2004): 109–32, 116. 80. Alan Grafen, “The Intellectual Contribution of The Selfish Gene to Evolutionary Theory,” in Richard Dawkins: How a Scientist Changed the Way We Think, edited by Alan Grafen and Mark Ridley (Oxford University Press, 2006), 73. 81. http://stevefrank.org/. 82. http://stevefrank.org/interests.html. 83. E-mail January 24, 2012; see Steven A.  Frank, “Natural Selection. III. Selection versus Transmission and the Levels of Selection,” Journal of Evolutionary Biology 25 (2012): 227–43. 84. George C.  Williams, “The Mysteries of Sex and Recombination,” the Quarterly Review of Biology 53, no. 3 (Sept. 1978): 287–89. 85. Williams, “Mysteries,” 287–89. 86. John Maynard Smith, “The Concept of Information in Biology,” Philosophy of Science 67 (2000): 177–94, 177, 189. 87. Williams, Sex, 154, 169. 88. Williams, Sex, 140. 89. Williams, Sex, 146, 147. 90. Williams, Sex, 147, 154, 155, 156. 91. See E.  O. Wilson and Frances M.  Peter, eds., Biodiversity (Washington, DC: National Academy Press, 1988). 92. Williams, Sex, 157, 158. 93. Williams, Sex, 159, 160. 94. Randy Thornhill and Craig T. Palmer, A Natural History of Rape: Biological Bases of Sexual Coercion (Cambridge, MA: MIT Press, 2000). See “Rape and Evolution: Authors’ Reply to Critics”: http://mitpress.mit.edu/ books/thouh/thornhill-­preface.pdf. 95. Laura Betzig, ed., Human Nature: A Critical Reader (New York: Oxford University Press, 1997), 17 n. 49. 96. Hamilton, Narrow Roads, vol. 1, 85; Bobbi Low, Why Sex Matters (Princeton, NJ: Princeton University Press, 2000), 282 n. 63.

CHAPTER 6

How Scientific Reductionism Leads to Evolutionary Explanation: A Defense

A central text in the Williams canon is the short but vigorous essay “A Defense of Reductionism in Evolutionary Biology,” published in 1985 as the lead article for the second volume of Oxford Surveys in Evolutionary Biology, edited by Mark Ridley and Richard Dawkins. According to its publisher, the essay is meant to stimulate discussion and outline progress.1 Williams writes his “defense” as a personal essay that looks backward and forward; as a narrator, he is introspective and prophetic. To simplify a great deal, for a biologist like Williams reduction begins with the ontologically materialist idea that any biological system consists of molecules and their interactions. A methodological assertion follows: that these biological systems are best investigated at the lowest possible level. Further, there is an epistemological principle at work: the general idea that knowledge about higher-level processes can best be understood by seeking a lower and more fundamental level.2 It is no accident that a dictionary definition of reduction includes the phenomenon of meiosis, where the first meiotic cell division reduces the chromosome number by half. Williams makes a reductionist argument and buttresses it with Dawkins’s work in The Selfish Gene (1976). Such a nod to the man who popularized Williams’s own ideas suggests that there are degrees of deference to celebrity even among scientists. Williams’s text absorbs Dawkins’s terminology incompletely, at the same time as it resists his terms, perhaps because © The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 M. P. Cohen, George C. Williams and Evolutionary Literacy, Literatures, Cultures, and the Environment, https://doi.org/10.1007/978-3-031-11650-6_6

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Williams is ambivalent about passing on the mantle of reduction. Yet he also plays the role of an American “hired gun,” providing authority for the “adaptationist program” in this introductory essay to a collection in a British journal. “From this selfish-gene theory of adaptation comes a reductionist methodology known as the adaptationist programme,” he writes, whose “practitioners imagine that they understand a studied organism well enough to recognize certain of its features as components of some special problem-solving machinery.” Prevalent genes must be those that “have had the most favorable phenotypic effects for their own replication,” Williams argues, using Dawkins’s language while mixing metaphors of bookkeeping, replication, and problem-solving machines.3 It is a literary critic’s job to un-mix these metaphors. Williams chose John Dalton’s atomic theory as a paradigm for the conclusion to Adaptation and Natural Selection in 1966, and we can assume that he did so because Dalton is generally considered a prime example of a reductionist. The question of reduction in biology is “where, primarily, does natural selection act, on species, groups, individuals, or genes?” Samir Okasha’s sophisticated analysis, Evolution and the Levels of Selection (2006), indicates that the meaning of reductionism can be difficult, but “Williams’s (1966) injunction that Darwinian explanations should always be sought at the lowest possible level is often called ‘reductionistic.’” Okasha also reminds us that Williams’s original thesis was “based on an empirical belief about the types of selection process most likely to be found in nature, not on methodological considerations.”4 Williams uses an analogy to illuminate his paradigm. Before Dalton, “the concept of matter as fundamentally particulate had been used in an undisciplined fashion at least since the days of Democritus”; it was “invoked, as natural selection often is today, whenever it seemed expedient to do so, but nothing was ever really demanded of the theory.” Real tests were required and “theory had to be stated in an explicit, quantitative, and uncompromising form before it was possible to recognize logical implications or to demand that there be a precise congruence between the theory and observation.” Dalton, by analogy, proposed six theoretical postulates about the nature of atoms, and because his statements “took no refuge in vagueness,” they could be tested empirically. As Williams observes, “The theory survived in a modified form, even though every one of Dalton’s six postulates turned out to be wrong, or at least inaccurate.”5 It is not the role of reduction to be right, but to further the progress of a science. In accordance with this paradigm of testing theory against observation, nearly two decades after Adaptation and Natural Selection, Williams

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continues questioning the ultimate value of his own postulates and explanations. However, a proximate reason for his “Defense of Reductionism” is Stephen Jay Gould and Richard C. Lewontin’s well-known paper “The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme” (1979). Their very title, by using the word “critique,” implicates Williams’ Critique of Some Current Evolutionary Thought. Nevertheless, their paper is most clearly directed toward the adaptationist program as practiced by sociobiology generally, occasioned in part by the publication of E. O. Wilson’s Sociobiology (1975) and his On Human Nature (1978).6 As late as 2007, Lewontin continued to speak of sociobiology as “an extreme example of the adaptationist program.”7 For this reason, there is value in disentangling the sociobiological program of Williams from that of E. O. Wilson.

Grounds of Sociobiologies The relationship between the reductive foundations of sociobiology and the problem of sexual reproduction is key to Williams’s transition, as the German biologist Joachim Dagg argues: “The rejection of group selection as a mechanism of sociobiology in the 1960s turned the maintenance of sexual reproduction into a major anomaly or paradox.” As Dagg believes, Williams’s discussions of sexual recombination led him increasingly to investigate the foundations of sociobiological conceptions. He began with the premise of Hamilton’s inclusive fitness, which in turn spotlighted the paradox of sex. As Dagg puts it, “the sociobiological conception of the cost of sex seems to correspond more closely to the root problem that initially perplexed these pioneers.”8 This is precisely the argument Williams makes in 1980, at a conference that includes both Gould and E. O. Wilson, when he asserts that the near universality of sexual selection “remains a major unsolved mystery,” especially “in relation to kin selection, on which so much of sociobiological thought depends.”9 Kin selection is another name for Hamilton’s inclusive fitness theory, that individuals have evolved to favor those who are genetically related to them. This theory is the basis for the sociobiological definition of altruism. Williams realizes, as Hamilton later writes, that “the problem of sex and the problem of recombination are closely related, but they are not simply two names for the same thing. Given that a population reproduces sexually, there still remains the vexing question as to why its recombination rates do not evolve downward toward zero.”10 In this context, Williams

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wonders in 1980, “What would be gained by an individual, in an otherwise sexual population, who cheated by eliminating meiosis and fertilization from its production of an offspring, but remained otherwise the same? Or the question may be put in the other way: what is the cost of not cheating?”11 With this kind of question, Williams “arrives at an entirely sociobiological conception” by 1988.12 Sociobiologists pursue the phenomenon of cheating when considering reciprocal altruism, and cheating is defined by Robert Trivers in a well-­ known article of 1971, but he is specifically interested in human interactions—that is, interactions above the genetic level. Trivers writes: “Altruistic behavior can be defined as behavior that benefits another organism, not closely related, while being apparently detrimental to the organism preforming the behavior, benefit and detriment being defined in terms of inclusive fitness.”13 Examples include visual and vocal warning signals given by birds and other animals. In 1966, when Williams considers these as social adaptations, he finds them incident to “well-developed parent care” and assumes that they would continue even when no offspring or conspecifics are present. “In principle,” as he puts it, “it might be well to dispense with both the behavior and the markings whenever there are no young to be protected. In practice, it is not worth burdening the germ plasm with the information necessary to realize such an adjustment.”14 Altruism becomes an essential question for sociobiology, but by diverting the question to sex, Williams brings the discussion of cheating to a lower level, to the natural selection of reproductive processes. As I have argued, he was attempting to construct some kind of grounding. Meanwhile, E. O. Wilson continued to define sociobiology in a more haphazard way. At an important symposium held at the 1978 American Association for the Advancement of Science annual meeting in Washington DC in February 1980, Wilson described sociobiology as “the systematic study of the biological basis of social behavior and the organization of all kinds of societies in all kinds of organisms, including human beings.”15 More than one sociobiology is possible, Wilson suggested, as he delineated “a discipline and not a particular theory.” Williams noted at the same symposium that, while Wilson’s Sociobiology (1975) may have focused attention on kin selection, reciprocal altruism, and parental manipulation in explaining social behavior, at a lower level, such functional concepts also describe genetic interactions between individuals, between mammalian mother and fetus, and between male and female gametophytes in the

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ovary of the flower, “not merely social behavior.”16 This precisely dramatizes his reductionist approach. It would be useful if there were a single theory or set of observable and agreed-upon grounds. Consequently, for this 1978 symposium that later became an edited volume, Williams contributed “Kin Selection and the Paradox of Sexuality.” His premise is that “an understanding of the cycle of meiosis and fertilization, and its genetic, ecological, and social consequences, is basic to the understanding of sociobiology.” According to Williams, sociobiology can then be defined not only as “the study of social behavior,” but also as “a set of observable phenomena.” Kin selection may be important to sociobiology, but it is complexly related to “variation in degrees of relationship arising from sexual reproduction” and the chromosomes. “Without this chromosome cycle, all the coefficients of relationship would be one or zero (complete genetic identity or total independence), and much of the complexity of interactions among organisms would presumably disappear.” Apparently, as Williams admits, his introduction of the cost of meiosis in 1975 led to discussions that “make it clear that the subject is more complex and confusing than I had thought.”17 A story of a hypothetical cheater who eliminates the reduction of meiosis calls for players. In such a story or narrative, If the fusing gametes are equal, neither parent seems to be subsidizing the reproduction of the other. In a more important sense this conclusion is wrong. Given that one cell fuses with another, we can then ask about the consequences of whether it plays the sexual game (meiosis) in its next cell division, or cheats (mitosis). It then becomes clear that the 50% cost of meiosis is still very much a reality.18

A few years later, Williams elaborates on this thought experiment, and one can see how he reduces the problem to cheating and piracy on the level of reproductive function. As he imagines, “Perhaps it is just a matter of time before someone discovers (or invents in the laboratory) an all-male species.” Such an invention could produce diploid sperm and inseminate eggs of a related species. This would be piracy, clearly, and the point is that “any male of any species that refrains from such egg piracy is paying a cost of meiosis as a direct result of the haploid and cytologically cooperative behavior of its sperm.” He further notes that while “a cost of meiosis is readily recognizable in my all-male species, a cost of males is not, at least not as an aspect of selection at the individual level.”19

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This kind of storytelling is both recursive and risky for a reductionist. After all, these examples of cheating or piracy are precisely not observable, as Williams reminds the reader in 1980. The mystery remains. Once again, when Williams scrutinizes grounds, he uncovers unanswered questions. His engages in reductionist reasoning that seems satisfactory; yet what he believes should happen does not happen. In a wider context, any structure for sociobiology’s grounds must be built on the mysterious foundation of the evolution and maintenance of sexual reproduction. By definition, a mysterious foundation is not a firm foundation. Under the circumstances, one way to seek or construct a grounding is to imagine testable stories.

Testing Stories As I have argued, George C. Williams’s most important contribution to evolutionary theory is in this realm of imagining and testing stories, because he establishes a role for critique in modern biology and makes it an essential grounding for evolutionary literacy. Hamilton may have established inclusive fitness, but Williams tests these stories or narrative explanations, developing the methodology essential to literacy. Stories are testable on empirical grounds, but they are also testable because they have plots and therefore embody theories of causality. Critique tests not just for empirical “truth,” but also for plausibility, for representative relevance. Critique asks how likely various sorts of stories are to happen: never, sometimes, mostly, often, or always? Those who test narratives must also offer replacement narratives. After all, if one story does not work as a representation, then another must be more representative. Critique must engage in its own storytelling, which is both a risk and an obligation for it. The new stories offered are then subject to further scrutiny, as Williams so frequently argued. We are told that stories and storytelling are natural and essential to human nature— our “evolutionary niche” according to pop writers like Jonathan Gottschall—but stories, even so-called scientific stories, often begin with prejudice and end with self-congratulation about some human essence. Caution is called for here because what happens in the processes of natural selection is predominantly not about humans. The natural sources for human stories are not so interesting or crucial for understanding our world as the cultural shaping of these stories, a process that must also be subject to critique. If humans are to sift the wheat from the chaff of narrative interpretations of the world, they cannot trap

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themselves in the tautology that humans tell stories because they are storytelling animals. In this sense, hierarchies are a prominent part of a set of tautologies associated with storytelling. People inescapably create hierarchies in stories about human activities and the activities of other living creatures. These stories often presume that more “natural” inclinations are more admirable, or higher, or more essential and good. For that reason and others, people, including some scientists, desire to believe that altruism, cooperation, and subordination to teams or groups are valuable because, in part, these motives are rooted in the natural. But people also create hierarchies of stories that value individual achievement and pass down tales of ascent as if this part of natural processes were most valuable. Most of all, many stories insist that whatever adaptive process produced the myriad forms of life must be beneficial for life as a whole and should elicit wonder. In a strange case, E.  O. Wilson argues, in the Social Conquest of the World, that ants—like humans—must exhibit social cooperation and altruism. Wilson’s is a bizarre kind of anthropomorphism, since after all ants are only programmed creatures, and unlike humans, for instance, lack any kind of individual will. Ants are mechanisms, chemically formed into castes. Surely Wilson doesn’t believe that humans are only chemically formed mechanisms. Humans may be programmed, but by more than deterministic mechanisms. A better image of the complexity of the human world is exhibited in Aldous Huxley’s Brave New World. George C. Williams finds hierarchies everywhere and scrutinizes their structure. He is suspicious of hierarchies in stories and of stories, but most of all he is skeptical of the desires that motivate the telling of success stories. He is especially suspicious of the notion that human acts shall be imagined as admirable because of, or in proportion to, their grounding in natural processes.

Disposable Soma and Indispensable Reduction To read Williams with any degree of engagement, one must entertain reduction as critique and as both a literary and a scientific strategy. Consequently, even if it runs against many literary sensibilities, an investigation of evolutionary theory must entertain reduction as a real biological phenomenon, a mode of thought, a method of inquiry, and a style of writing too.

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Reduction is a key step in sexual reproduction. In meiosis, we remember, the number of chromosomes is reduced. But meiosis is a form of reduction in more than one sense. Meiosis produces daughter cells not identical to the cells of parents. As Williams has argued, the process is costly. Another cost of this process that generates genetic diversity, one might say, is identity itself. Identity is biologically ephemeral. Consider those interested in their own genealogy, for instance, or those who purchase a reading of their own DNA: they may be investing in a kind of iconography and engaged in a narcissistic task. Meiosis marks the onset of extinction of identity; it leads toward death. In the long view of life history, death becomes a tautological kind of ultimate reduction. All mammals senesce and die because their soma is, from the perspective of natural selection, disposable, as Williams has argued. This idea encounters resistance from individualists and specifically from romantics. In the methodological sense, reduction is the name of the likely most successful method for understanding evolution. In the coda to his “Defense of Reduction,” Williams writes: The history of science shows the inevitability of widespread emotional dissatisfaction with reductionist interpretations of the world. I suppose there are people who think the theory of sexual selection robs the grossbeak’s song of its music, or that kin selection makes a mockery of brotherly love. There used to be those that thought that Newtonian optics destroyed the beauty of the rainbow.20

Williams’s attack on romanticism has been broadened and politicized by others. One can find Thomas DeGregori writing on the Cato Institute’s website about the fiftieth anniversary of Watson and Crick’s deciphering of the double helix structure of DNA: “To some, the molecular biology of DNA is a symbol of the much-despised, modern, Western, reductionist science that is blamed for every conceivable ill, from imperialism and male domination of women to environmental degradation.”21 But let us assume that conversation with and between working scientists must respect what David L. Hull calls the internalist premises of reason, argument, and evidence, which buttress the integrity and validity of their methods. Hull has written extensively on Williams (and Dawkins) as “gene selectionists” and, as he argues, “gene selectionism is an extreme form of selectionism.”22 But that statement of 1988 appears before Williams completes his work.

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So how does Williams speak of gene selectionism as genetic reductionism in 1966? He accepts Haldane’s 1957 premise that “the principal unit process in evolution is the substitution of one gene for another at the same locus.”23 But Williams acknowledges problems, writing, “Obviously it is unrealistic to believe that a gene actually exists in its own world with no complications other than abstract selection coefficients and mutation rates.” As a complex unit, a genotype subordinates individual genes to each other and to their surroundings. Consequently, what Williams calls “the one-locus model of natural selection” would seem to be invalid. Yet this does not bear on the basic postulates, for “no matter how functionally dependent a gene may be, and no matter how complicated its interactions with other genes and environmental factors, it must always be true that a given gene substitution will have an arithmetic mean effect on fitness in any population.” One can do the math to treat individual genes algebraically by assigning “selection coefficients” to them. Unfortunately, “although this theory is conceptually simple and logically complete, it is seldom simple in practice and seldom provides complete answers to biological problems.”24

Enlarging Reduction with History One imagines a young man studying biology, and in the midst of his work he is influenced, maybe even interrupted, by the advent of DNA as understood by Watson and Crick—DNA as model, machine, and one-way street, the ultimate biological reduction. But by 1985, when he is nearly fifty years old, he reconsiders in “A Defense.” “Reductionism,” begins Williams in 1985, “is the seeking of explanations for complex systems entirely in what is known of their component parts and processes.” For evolutionary science, “the current most important reductionist device is to regard gene frequency change as the essence of evolution.” One may think of this device through an analogy: “Such change is a bookkeeping process that records past reproductive success and predicts what is likely to succeed in the future.” Nevertheless, bookkeeping is not prediction and, though critics of evolutionary reductionism may complain, “the idea that a theory of evolution must predict future evolutionary change is unrealistic.” It is one thing to read a book and another to predict how it will be changed, revised, or rewritten in time. Clearly, Williams tempers his vigorous defense by recognizing its limits, and he takes the term bookkeeping from some of the critics of his own practice of

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“gene-centered reductionism.”25 The “Defense” is also the most overt entry Williams makes into the well-publicized controversy over levels of selection and the elaborate debates about the “adaptationist program.” As the analogy to bookkeeping suggests, the “Defense” introduces Williams’s version of a Book of Nature—a trope with a long history. Literary critics of my generation, for instance, were introduced to the discipline through a reductionist methodology called “the New Criticism” that treats any work of literature “as if it were a self-contained, self-­ referential object.” The New Critic interpreted texts without recourse to a reader’s response, an author’s stated intentions or biography, or any relation to historical and social contexts. The New Critic performed a close reading, concentrating on relationships between words within the text, to show how language creates a distinctive character or form. In particular, New Critics read poems as internally creating expectations and either meeting them or deliberately not doing so.26 I find a kinship to this mode of analysis in the early writings of Williams. Anyone who wishes to understand how Williams judges his contributions to evolutionary science would be well advised to follow his defense closely, for here he assesses his lifelong method of study, explains what he thinks he is studying or “reading,” and reveals where he finds himself in his tradition. Thus, when he writes that “a biological explanation should invoke no factors other than the laws of physical science, natural selection, and the contingencies of history,” the third factor comes as a surprise to an attentive reader. In Adaptation and Natural Selection (1966), he argued that “the laws of physical science plus natural selection can furnish a complete explanation for any biological phenomenon.”27 By including history as a factor in 1985, Williams changes his perspective significantly, and steps carefully, if incompletely, away from radical gene selectionism. When Williams includes historicity in explanation, he complicates his mission and complicates his argument. “I once insisted,” he writes later in the “Defense,” that “‘the laws of physical science plus natural selection can furnish a complete explanation for any biological phenomenon,’ … I wish I had taken a less extreme view.” Even physicists “need to reckon with historical legacies to explain any real-world phenomenon,” and “history is an even more obvious element in the biological world.” In 1985, he introduces chance, contingency, drift, and the stochastic nature of the mutation process repeatedly. Here is where bookkeeping comes in: “I suggest that the term evolution should be used only in this sense, of a process

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determined by chance events plus the persistent biases of natural selection.”28 An explanation is dependent upon certain factors, or to put it another way, explanation factors in certain forces. When Williams factors in historical legacies, when he admits contingency into his narrative, he also admits environmental forces. As he writes, “no biological theory could have predicted the global climate changes and environmental disruptions of the Pleistocene.”29 This constitutes a transition in his thinking toward macroevolution—or evolution on a large scale, toward the big history of life, including the stability, change, rise, and extinction of lineages. This is traditionally the realm of paleontologists, from George Gaylord Simpson to Stephen Jay Gould. There are some perspectives that Williams will not entertain. He proposes that “only confusion can arise from the use of an animal mind concept in any explanatory role in biological studies of behavior.” There are, he asserts, no shared parameters between the mental and the physical. The awkwardness of his language indicates, perhaps, that Williams chafes against a role that calls for arguing not only against Gould’s “pluralism,” but also against Donald R. Griffin’s “mentalism” in The Question of Animal Awareness (1976) and Niles Eldredge’s “hierarchical concepts.” “Perhaps my efforts here will help to reduce the frequency of future misunderstanding,” he writes. “And perhaps not,” he continues, tongue in cheek.30 Williams spells out his case, or really a set of wishes. He wishes for a “formal separation of the informational (genetic) aspects of natural selection from its concrete ecological aspects,” by which he means its interactions with the environmental variables it faces—a separation that “may help in avoidance of semantic distractions.” He wishes to distinguish between “selection” and “response to selection,” or as in Hull’s terms, between replication and interaction. He prefers Hull’s term interactor to Dawkins’s term vehicle.31 This is not a minor terminological issue, since Dawkins’s vehicles are like robots being driven by genetic drivers. Hull’s distinction is this: a replicator is “an entity that passes on its structure directly in replications,” but an interactor is “an entity that directly interacts as a cohesive whole with its environment in such a way that this interaction causes replication to be differential.”32 One might say that this is what Williams means by “ecological.” Williams asks his reader to consider “natural selection as a history of success and failure, and a record of that history for various possible levels

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of selection,” or a “genetic book,” if you will. Because bookkeeping “has been taking place in the past,” because a “gene is not the molecule, but the information coded by the molecule,” and because “it just so happens that in genetics the archival medium is always DNA,” the gene must be imagined as “an ideal entity of bookkeeping,” given the “precisely engineered replication processes of DNA.”33 Yet with each and every generation, precise genetic text is lost, “effectively erased” during the shuffling of genotypes through sexual reproduction, so that phenotypic expression of a gene is “thoroughly context-dependent.” Bananas, he says, always look the same, but they are not. “There can also be a compelling familiarity in the faces that look out at us from centuries-old portraits of organisms that did no cloning.” Nevertheless, though there seems to be a “persistence of type or pattern” among phenotypes, “the term ‘persistence’ is misleading.” What one sees, in fact, is recurrence: “The distinction is fundamental and only confusion can result from labeling as persistent or permanent that which merely recurs.”34 History, one might say, is not about permanence, but about recurrence and change.

Prediction and Bookkeeping At the center of this essay, Williams encounters questions of prediction, some of greater and some of lesser importance. As he argues, prediction is dependent upon bookkeeping. If one imagines a population, “a long-­ lasting collection of individuals, typically hundreds or more, which maintains itself for many generations by its members’ reproduction,” then by definition “the bookkeeping consists of the proliferation of some sets of gene-frequency values and the disappearance of others.” To extend the analogy, “a gene pool too can be a suitable medium for writing a record.” He simplifies somewhat, speaking of two kinds of bookkeeping and researchers who follow one or the other. The first book reveals an “importance for producing and maintaining adaptation” and the second an “importance for determining the properties of the earth’s biota.” Williams simply admits that “my interests have always been in the first kind,” through the approach Gould and Lewontin called the adaptationist program. “I view an organism as a participant in a contest and it is as such that I wish most to understand it. Other biologists no doubt have other interests.” In the game of life, as Williams understands it—and he traces this

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view back to prehistory and the writing of Galen—each organism participates as an individual contestant, “not as a member of a team.”35 This kind of bookkeeping goes on within individuals, not within groups. Though removing a species from a community, for instance, “can have drastic consequences for those that remain,” Williams insists that, “for most of the many species that inhabit a community without accounting for a major share of its biomass or metabolism, the extirpation of one will make little difference to most of the others.” Contrary to the views of so-called systems ecologists like Eugene Odum, Williams argues that there is no order in ecosystems: “Functional subordination of component species is not normally found in natural communities,” and other possible kinds of order are too difficult to detect. But there are, Williams continues, “many kinds of selfishness,” including some that may be self-­ destructive. All in all, “it may be that the adaptationist programme based on selfish-gene reductionism will have little to contribute for an understanding of macroevolution.” But “perhaps a chaotic sort of historical contingency is a more important macroevolutionary factor than group selection.” He concedes that critiques of the adaptationist program by the philosophers Elliot Sober and William Wimsatt and geneticist Richard Lewontin are “entirely correct in their reasoning” when they assert that “reductionism is incapable of predicting future gene frequencies at even a single locus.” In fact, he considers that only short-term predictions might be possible, and that they are the “less important kinds of prediction in evolutionary biology.”36 On one hand, Williams says that many less important predictions of evolutionary change are “routinely made” by following short-term genetic change, sometimes using experimental populations and sometimes natural populations, as he has done with eels. Yet he admits, “we had no hope of gaining information on what eels would be like in the remote future, or even of identifying environmental factors responsible for the changes.” He notes that applied biologists make predictions for economically important species, and they do so more from experience than from any models. A second kind of prediction, “as old as the idea of evolution,” comes from reconstructing phylogenies, and can “predict characteristics not yet investigated.”37 Though archaeologists seek common ancestors of humans and chimpanzees, the required fossil may be a long time in coming. On the other hand, the more important and worthwhile predictions are meant to illuminate the methods of biologists. “The primary role of theory in evolutionary biology, as in any other science, is to predict the

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outcome of investigations not otherwise predictable.” This is what prediction does: it constructs human narratives and furthers the human ability to test them. At the center of his “Defense,” Williams arrives at the following thesis, or to put it another way, at an explanation for his particular mode of thinking: The important predictions in evolutionary biology derive from a concept of adaptation based on a selfish-gene model of natural selection and take this form: given that all organisms throughout their history have been subject to this kind of bookkeeping, any given organism is expected to have certain properties and not others. The properties are expected to form a closely-­ optimized strategy for maximum proliferation of the genes that directed the development of the organism. No compromise with any other goal, like the survival of the species, is expected. Inspection of the organism will disclose whether it does or does not have the predicted properties.38

Williams consistently reasons this way, comparing the predictions of theory to the actualities exhibited by organisms. He clarifies this distinction in terms of what is called optimization, a term with a very different meaning than that used by modern creators of computer-based simulations. As he argues later, in 1992, “organisms are never optimally designed,” but consist of “legacies from the past” on which natural selection works, often by adjusting the numbers of mutually exclusive designs until only one design “excludes alternatives.” Or, natural selection can “optimize a design’s parameters so as to maximize the fitness attainable with that design under current conditions. This is what is usually meant by optimization in biology.”39 All too often, scrupulous inspection by a researcher discovers problems in optimization and failures of theory to predict accurately. What should result from “the rules of the game being played” and “other necessary constraints” is not always found. In particular, “most evolutionary biologists seem to recognize that optimization is valid only in relation to local optima.”40 Theory often allows one to see how the process of natural selection leads to imperfections— failures to achieve optima. One problem in using the concept of optimization to understand adaptation comes about because “it is necessary to distinguish ‘strategies,’ which organisms try to optimize, from ‘winnings,’ which they try to maximize.” Unfortunately, “there is no simple rule for distinguishing strategies from winnings.” Such a distinction can only be gained by “a detailed

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familiarity with the natural history of the organism under study. This difficulty is further evidence of the immaturity of the study of adaptation.” Conflict is often the driving force for adaptive change or stasis and is better analyzed by game theory than optimization, at least according to Maynard Smith. “It is especially clear for contests with other organisms that strategies favoured by selection may be quite different from ideal adaptations that we might imagine.” Yet Williams mentions thousands of attempts to “understand adaptation via a thoroughly reductionist theory of natural selection” and takes authority from Karl Popper, who endorses the richly rewarding reductionist program that “attempts to understand organisms as problem-solving complexes” by studying the “origin and maintenance of the problem-solving machinery.”41 By definition, such study calls for understanding and telling stories.

Factors of History and Mind in Natural Selection: Just-so Stories Williams insists on “the methodological identity of evolutionary biology with other sciences,” particularly because they all create hypotheses and attempt predictions—sometimes by telling stories. But there are some kinds of stories he will not countenance. Consequently, the question of prediction leads to a discussion of storytelling. Williams needs to refute claims that adaptationists simply create “just-so stories” without losing sight of his doctrine or ground rule of 1966, that “adaptation is a special and onerous concept that should be used only where it is really necessary,”42 It is onerous to follow the bookkeeping, read its history, and devise stories—narratives with direction but without predictability—to explain the designs of biological machinery. As Williams explores the relevance of history in explaining evolutionary change, he justifies this affirmation by invoking a history of Darwinian explanation. When he endorses Darwin’s materialism, he dismisses the many other explanatory factors—natural and supernatural—that have been proposed. He pokes some fun at Gould and Lewontin when speaking of their proposals in the same satirical sentence that dismisses “thinly-­ veiled mysticism with erudite names, like aristogenesis and the Gaia hypothesis.” His use of the first person reveals personal commitment as he articulates his own sense of the Darwinian tradition: “I accept with enthusiasm that for anything to evolve it must do so by modifying that which

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came before, and I heartily agree with Darwin’s insistence that every stage in the evolution of any useful feature has to be useful to its own possessors, not merely of potential use later.” But also, he confesses that in the past he did not give enough emphasis to a “need to reckon with historical legacies to explain any real-world phenomenon.” Taking heed of chance events and contingencies, like mutation and drift, requires some redefinition of evolution itself to include “a process determined by chance events plus the persistent biases of natural selection.”43 One version of persistent bias, termed canalization by Conrad Hal Waddington, had been useful to Williams since 1966. As one review of the subject puts it: “In the 1940s, Conrad Hal Waddington coined the concept and term ‘canalization’ to describe the robustness of phenotypes to perturbation.”44 As a specialized term, canalization describes a preponderance of normal self-regulatory developmental processes leading to nearly identical phenotypes in populations. One knows that genetic variation maps onto phenotypic variation. Why, then, are some traits phenotypically invariant despite apparent genetic and environmental changes? This has remained a major puzzle. Williams concedes a variety of kinds of canalization—a more canalized phenotype alters less in response to changes in environment or changes in genetic background. He thinks individual ontogeny must be canalized by control mechanisms “that consistently correct perturbations from a prescribed course of development.” To a lesser extent, he notes, some believe ecological succession can be canalized, by feedback from densities of populations or by the activities of separate species interfering with each other. As he puts it, “I see no evidence that evolution is subject to such control, but others often do.”45 The biggest story is that of natural selection, which will be the title of his next book. “There is no biological generalization so general that it can serve as principle of the same rank as natural selection.” Natural selection is mindless, of course, and including the idea of mind as a factor only confuses issues. In particular, “mind may be self-evident to most people, but I see only a remote possibility of its being made logically or empirically evident.” He adds that “there is no such evidence for mind as an entity that interferes with physical processes.”46 He is speaking of Animal Minds (1981) by Donald R. Griffin and a 1983 article by Daniel Dennett.47 Like Tinbergen, Williams remains very skeptical of these perspectives and writes in that tone: “I feel intuitively that my daughter’s horse has a mind. I am even more convinced that my daughter has. Neither conclusion is supported by reason or evidence.” Adducing defenses by Tinbergen and

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T. H. Huxley, he continues, “Lewontin’s solution to the non-objectivity of mind is to abandon reductionism. Mine is to exclude mind from all biological discussion.”48 In this remarkable passage where a father claims to be engaged in biological discussion, he objectifies and demeans his daughter by perceiving her in the same context as a horse. One is stunned. Williams declares that “the whimsical recognition of mentalism is a frequent obfuscation in discussions of adaptation and especially animal behavior,” which is why he uses botanical examples when teaching undergraduates about kin selection, mate choice, and life history. This strategy “makes it easier for students to discuss analogous zoological examples as biology rather than soap opera.” The only mind in question, when thinking about biology, is the mind of the human observer. In a nice touch, Williams borrows a story about animal intelligence from Gould, of Clever Hans the horse everyone knows, who was no mathematician and was “unable to use Arabic numerals or German for any sort of analysis or communication.” Yet, as Williams adds to the story, “every time he trotted across a field he rapidly performed a series of calculations far more subtle than long division.”49 It is best to ask the right questions in biology by making calculations based upon “component parts and processes.” It is best to advocate this strategy for understanding “flesh-and-blood (or cellulose) organisms” in the first person. “I find the inclusive fitness concept, directly derivable from reductionist, selfish-gene theory of natural selection, a useful device.” Yet as he argues in his conclusion, “I depart from Dawkins” because he “would restrict the concept of fitness to the gene and not try to salvage its applicability to their interactors.”50 Williams is still mulling this problem over a decade later when he writes “A Package of Information” for Brockman’s The Third Culture (1995). The Dawkins view, he insists, confuses things by defining a replicator as a physical entity “duplicating itself in a reproductive process,” and though Dawkins is not alone in this, he “was misled by the fact that genes are always identified with DNA.”51 When one follows the perspective of Williams’s narration in 1985, one sees that the message becomes increasingly personal, and as with earlier distinctions between strategies and winnings, he attributes to himself only small suggestions or improvements.

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From Reduction to Life History: The Book of Nature Reduction, as Williams argues, entails thinking not on the level of the biota, species, herd, group, individual, or even gene, but thinking on the level of information. For instance, an early version of life history theory is the r/K selection theory devised by E. O. Wilson and Robert MacArthur in The Theory of Island Biogeography.52 By their definition, r-selected organisms mature rapidly and reproduce early. In a short lifespan, they have many offspring, most of whom die. They invest little in parental care. K-selected organisms mature slowly, reproduce at a later age, live longer, and have fewer offspring spread over their reproductive period; their offspring are more likely to survive, partly as a result of parental investment. At the extremes, there are eternal reproducers like pine trees on one hand and, on the other, species of plants and animals whose life histories entail parental death after first reproduction. Contemporary commentators see that the dualistic and deterministic scheme of r versus K reproduction strategies does not adequately reflect the range of life histories or the many genetic traits and environmental constraints that influence life histories of species.53 In 1977, Stephen C.  Stearns, who submitted his early ideas to Williams and became the expert on the subject, decided that “we do not yet have a general and reliable theory of life history evolution, and the crux of the problem is: What will be an empirically sufficient set of parameters in which to couch the theory?”54 Implicit in Stearns’s conclusion is a desire for reduction and for a complete narrative. Life history can be broken down into traits, and the traits can be categorized as parameters, as information. Life history theory then aims to construct narrative models which predict what sorts of traits will be favored in what sorts of environments. This is exactly the method Williams attempts and accomplishes in 1957 when he writes about the evolution of senescence. Implicitly, he begins to think about a set of rules that can illuminate allocations of energy and time for any creature and the tradeoffs between reproduction and growth. Fertility, mortality, growth, and survival: life history theory is not only a theory of life cycles, but a theory of fitness. It is tied to environment, so there are an increasing number of studies attempting to predict the effects of climate change on the life-history traits of plants and animals, birds and trees.

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But in 1985 Williams recognizes two kinds of history—and they are not congruent. The first kind he has been doing all along, and it is amenable to prediction via reduction. The second kind of history requires archival information, like fossils and so on, and is subject to contingency. Because prediction depends upon smooth, uniform environmental change, and because contingent environmental change is not gradual, this second kind of history is not amenable to prediction. These two versions of history influence his style of writing. In such introspective discussions, the author cannot disappear as does the artist in James Joyce’s novel, who, “like the God of the creation, remains within or behind or beyond or above his handiwork, invisible, refined out of existence, indifferent, paring his fingernails.” On the contrary, for Williams in 1985, a first-person narrator must appear. He is neither infinite nor above. As a reader of the Book of Nature, he is in medias res. * * * Murray Krieger, literary critic, theorist, and my teacher, taught that a romantic symbol, as Samuel Taylor Coleridge suggested, denotes the natural world imagined as a Book of Nature, at the same time as it partakes of the Word of God. Coleridge writes: True natural philosophy is comprised in the study of science and language of symbols. The power delegated to nature is all in every part: and by a symbol I mean, not a metaphor or allegory or any other figure of speech or form of fancy, but an actual and essential part of that, the whole of which it represents.55

Krieger considered Coleridge’s “monistic conception of metaphor” as “a romantic reversion to the sacramental union put forth in Christian theology,” as embodied in his famous definition, in Biographia Literaria, of the primary imagination as “a repetition in the finite mind of the eternal act of creation in the infinite I AM.”56 But for any reader, these two kinds of “writings” call for two kinds of interpretation. The word of God is deterministic and predictive. The Book of Nature is more complicated, more multivarious than Coleridge imagined, and requires empirical evidence.

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Notes 1. George C.  Williams, “A Defense of Reductionism in Evolutionary Biology,” in Oxford Surveys in Evolutionary Biology, vol. 2, edited by Richard Dawkins and Mark Ridley (Oxford: Oxford University Press, 1985), 1–27. For a definition of reductionism in biology, see Ingo Brigandt and Alan Love, “Reductionism in Biology,” The Stanford Encyclopedia of Philosophy (Summer 2012 Edition), edited by Edward N.  Zalta, http:// plato.stanford.edu/archives/sum2012/entries/reduction-­b iology/. Daniel Dennett’s more animated discussion, indebted to Williams, can be found in the conclusion to Chapter 3, “Universal Acid,” under the subheading, “Who’s Afraid of Reductionism?” in Daniel C. Dennett, Darwin’s Dangerous Idea: Evolution and the Meaning of Life (New York: Simon & Schuster, 1996), 80–83. 2. Brigandt and Love, “Reductionism.” 3. Williams, “Defense,” 1. 4. Samir Okasha, Evolution and the Levels of Selection (Oxford, UK: Clarendon Press, 2006), 139, 141. 5. George C.  Williams, Adaptation and Natural Selection (Princeton, NJ: Princeton University Press, 1966), 272, 273. 6. See Stephen Jay Gould and Richard C. Lewontin, “The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme,” Proceedings of the Royal Society of London, Series B, 205, no. 1161 (1979): 581–98; E.  O. Wilson, On Human Nature (Cambridge, MA: Harvard University Press, 1978). See also Ullica Segerstråle, Defenders of the Truth: The Sociobiology Debate (Oxford: Oxford University Press, 2000), 284–91, and Elisabeth Lloyd, “Units and Levels of Selection,” The Stanford Encyclopedia of Philosophy (Winter 2012 Edition), edited by Edward N. Zalta, http://plato.stanford.edu/archives/win2012/entries/ selection-­units/. Lloyd has published an earlier version with the same title in Evelyn Fox Keller and Elizabeth A. Lloyd, eds., Keywords in Evolutionary Biology (Cambridge, MA: Harvard University Press, 1994), 334–40. 7. Richard C.  Lewontin, “Sociobiology as an Adaptationist Program,” Behavioral Science 24, no. 1 (2007): 5–14. 8. Joachim Dagg, “The Paradox of Sexual Reproduction and the Levels of Selection: Can Sociobiology Shed a Light?” Philosophy and Theory in Biology 4 (January 2012): 2, 5. http://hdl.handle.net/2027/ spo.6959004.0004.001. 9. George C.  Williams, “Kin Selection and the Paradox of Sexuality,” in Sociobiology, Beyond Nature/Nurture?: Reports, Definitions, and Debate, edited by George W. Barlow and James Silverberg (Boulder, CO: Westview

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Press, 1980), 371–84. This volume is based on a symposium held February 12–17, 1978, at the AAAS National annual meeting in Washington, DC. 10. J. Seger and W. D. Hamilton, “Parasites and Sex,” in The Evolution of Sex, edited by R.  E. Michod and B.  R. Levin (Sunderland, MA: Sinauer Associates, 1988), 177. 11. Williams, “Kin Selection,” 372. 12. Dagg, “Paradox,” 5. 13. Robert L. Trivers, “The Evolution of Reciprocal Altruism,” the Quarterly Review of Biology 46, no. 1 (1971): 35–57, 35. 14. Williams, Adaptation, 206–7. On biological altruism, see Samir Okasha, “Biological Altruism,” The Stanford Encyclopedia of Philosophy (Winter 2009 Edition), edited by Edward N.  Zalta, http://plato.stanford.edu/ archives/win2009/entries/altruism-­biological/. On cheating, the prisoner’s dilemma, game theory, and sociobiological explanation, see Harmon Holcomb and Jason Byron, “Sociobiology,” The Stanford Encyclopedia of Philosophy (Fall 2010 Edition), http://plato.stanford.edu/archives/ fall2010/entries/sociobiology/. 15. E.  O. Wilson, “A Consideration of the Genetic Foundation of Human Social Behavior,” in Sociobiology: Beyond Nature /Nurture, edited by George W.  Barlow and James Silverberg (Boulder, CO: Westview Press, 1980), 295–305, 296. 16. Williams, “Kin Selection,” 371. 17. Williams, “Kin Selection,” 371, 372. 18. Williams, “Kin Selection,” 377. 19. Williams, “Retrospect on Sex and Kindred Topics,” in The Evolution of Sex, edited by R.  E. Michod and B.  R. Levin, 287–98 (Sunderland, MA: Sinauer, 1988), 294. 20. Williams, “Defense,” 24. 21. Thomas DeGregori, “DNA and Reductionist Science,” Cato Institute (April 3, 2003), http://www.cato.org/pub_display.php?pub_id=3045. 22. David L. Hull, Science as a Process: An Evolutionary Account of the Social and Conceptual Development of Science (Chicago: University of Chicago Press, 1988), 10, 422; see also 408–17. 23. J. B. S. Haldane, “The Cost of Natural Selection,” Journal of Genetics 55, no. 3 (1957): 511–24, 511. 24. Williams, Adaptation, 56–7. 25. Williams, “Defense,” 1. See, for instance, William C.  Wimsatt, “Reductionistic Research Strategies and their Biases in the Units of Selection Controversy,” in Scientific Discovery, vol. 2: Case Studies, edited by Thomas Nickles (Dordrecht, Holland: D. Reidel, 1980), 213–59. 26. See John Crowe Ransom, The New Criticism (Westport, CT: Greenwood Press, 1979), and Frank Lentricchia, After the New Criticism (University of

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Chicago Press, 1981). For a simple definition of the New Criticism, see http://bcs.bedfordstmartins.com/virtualit/poetry/critical_define/crit_ newcrit.html. 27. Williams, “Defense,” 1, my emphasis; Williams, Adaptation, 6–7. 28. Williams, “Defense,” 20, 21. 29. Williams, “Defense,” 10. 30. Williams, “Defense,” 2. Donald Redfield Griffin, The Question of Animal Awareness: Evolutionary Continuity of Mental Experience (Rockefeller University Press, 1976). On the question of continuity, see Niles Eldredge, “Evolution and Prediction,” Science 212, no. 4496 (May 15, 1981): 737. 31. Williams, “Defense,” 3. 32. David L.  Hull, “Individuality and Selection,” Annual Review of Ecology and Systematics 11 (1980): 311–32, 318. See also a slightly revised version in Hull, Science as a Process, 408. 33. Williams, “Defense,” 3, 4. 34. Williams, “Defense,” 5. 35. Williams, “Defense,” 7, 8. 36. Williams, “Defense,” 8, 9, 10, 11. 37. Williams, “Defense,” 11. 38. Williams, “Defense,” 12. 39. George C.  Williams, Natural Selection: Domains, Levels, and Challenges (New York: Oxford University Press, 1992), 56. 40. Williams, “Defense,” 12, 13. 41. Williams, “Defense,” 13, 14, 15. 42. Williams, Adaptation, 4. 43. Williams, “Defense,” 1, 19, 20, 21. 44. See Mark L.  Siegal and Aviv Bergman, “Waddington’s Canalization Revisited: Developmental Stability and Evolution,” Proceedings of the National Academy of Sciences 99, no. 16 (2002): 10528–32; Thomas Flatt, “The Evolutionary Genetics of Canalization,” the Quarterly Review of Biology 80, no. 3 (2005): 287–316. 45. Williams, “Defense,” 21. 46. Williams, “Defense,” 21. 47. Daniel C.  Dennett, “Intentional Systems in Cognitive Ethology: The ‘Panglossian Paradigm’ Defended,” Behavioral and Brain Sciences 6, no. 3 (1983): 343–55. 48. Williams, “Defense,” 21, 22. Lewontin writes specifically, “If it were our purpose in this chapter to say what is actually known about the evolution of human cognition, we could stop at the end of this sentence.” Richard Lewontin, “The Evolution of Cognition: Questions We Will Never Answer,” in An Invitation to Cognitive Science: Vol. 4 Methods, Models, and

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Conceptual Issues, 2nd edition, edited by D. Scarborough and S. Sternberg (Cambridge, MA: MIT Press, 1998): 107–32, 107. 49. Williams, “Defense,” 23. 50. Williams, “Defense,” 24. The famous line in Richard Dawkins’s The Selfish Gene (1976) is: “We are survival machines—robot vehicles blindly programmed to preserve selfish molecules known as genes” (vii). See David Hull and John S.  Wilkins, “Replication,” The Stanford Encyclopedia of Philosophy (Winter 2008 Edition), edited by Edward N.  Zalta, http:// plato.stanford.edu/archives/win2008/entries/replication/. 51. John Brockman, The Third Culture (1995), 42, 43. http://www.edge. org/documents/ThirdCulture/h-­Ch.1.html. 52. Robert R. MacArthur and E. O. Wilson, The Theory of Island Biogeography (Princeton, NJ: Princeton University Press, 1967). 53. Steven S. Stearns, The Evolution of Life Histories (Oxford: Oxford University Press, 1992). 54. Steven S. Stearns, “The Evolution of Life History Traits: A Critique of the Theory and a Review of the Data,” Annual Review of Ecology and Systematics, 8 (1977): 145–71, 168. 55. Coleridge quoted in Denis Donoghue, “Murray Krieger versus Paul de Man,” in Revenge of the Aesthetic: The Place of Literature in Theory Today, edited by Michael P.  Clark (Berkeley: University of California Press, 2000), 102. 56. Murray Krieger, Words about Words about Words: Theory, Criticism, and the Literary Text (Baltimore, MD: Johns Hopkins University Press, 1988), 276, 277.

CHAPTER 7

Evolution and Human Ethics: An Expansion from Sociobiology

In one of their roles, scientists try to get the “public” to face realities and offer strategies to deal with those realities, not just technically, but through policy. In this context, Williams speaks for contemporary biology and its suspicion of a normative view of nature in his essays on T.  H. Huxley. Williams later calls suspicion “a fortunate human tendency” that allows arguments to be grounded in “reasoned analysis prompted by suspicion.”1 As he writes repeatedly, “I account for morality as an accidental capability produced, in its boundless stupidity, by a biological process that is normally opposed to the expression of such a capability.”2 The process can be subverted—as, leaning on Huxley’s perspective, Williams will argue—if human ethical nature, though entirely unrelated to the selfishness responsible for evolutionary changes, can emerge “incidentally” from biological nature. In a variety of venues, and most particularly in Zygon: A Journal of Religion and Science, Williams argues, “my most basic conflict is with the idea that evolution is good in general or on the average.”3 He explores the relationship between evolutionary knowledge and ethics in a set of “expansions” in the late 1980s and early 1990s.4 He takes an epigraph for several of these essays from Thomas Henry Huxley’s Evolution and Ethics (1893–1894): “Let us understand, once and for all, that the ethical progress of society depends, not on imitating the cosmic process, still less in running away from it, but in combatting it.” The first sentence of most © The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 M. P. Cohen, George C. Williams and Evolutionary Literacy, Literatures, Cultures, and the Environment, https://doi.org/10.1007/978-3-031-11650-6_7

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versions of Williams’s essays is, repeatedly, “I agree with Huxley’s statement, but others do not.”5 When he entitles his own central essay “A Sociobiological Expansion of Evolution and Ethics,” he means that he expands Huxley’s views in light of a modern “expansion” of Darwinism into sociobiology.6 One might also argue that he expands his questioning of the benevolence of nature at the end of Adaptation and Natural Selection, asking whether nature is “an appropriate guide for devising ethical systems and for judging human behavior.”7 How a more general public readership responds to this question is a different matter. The occasion for Williams’s “Sociobiological Expansion” is his editing of T.  H. Huxley’s Evolution and Ethics. The text includes Huxley’s Romanes lecture of that title, given at Oxford University on May 18, 1893, and an additional “Prolegomena” completed in June 1894. Essays by Williams and his co-editor James Paradis frame a complete facsimile of both essays in the original 1894 publication.8 In other words, Williams gives presence to a work nearly 100 years old and reintroduces Huxley’s argument to readers, while Paradis explores the historical context, or the pastness, of Huxley’s ideas. I have noted a surprising shift in Williams’s perspective when he expands upon the manifesto of 1966. He no longer promises that “laws of physical science plus natural selection can furnish a complete explanation for any biological phenomenon,” but admits “the contingencies of history.” Now working within a specific literary-historical context, he correspondingly identifies these three themes—physics, natural selection, and history— sourcing them in the suspicions of Huxley, who recognized “the hostility of the cosmos to human life and aspirations,” undertook “a moral evaluation of the evolutionary process,” and doubted “the evolutionary origin of the human moral impulse.”9 Two versions of his first essay, “Huxley’s Evolution and Ethics in Sociobiological Perspective” and “A Sociobiological Expansion of Evolution and Ethics,” deal with the second and third themes in Huxley’s work. Meanwhile, in 1989, Williams reviews Robert J. Richards’s Darwin and the Emergence of Evolutionary Theories of Mind and Behavior; the book includes a postscript entitled “A Defense of Evolutionary Ethics” with which Williams respectfully disagrees.10 Three years later, Williams publishes “Gaia, Nature Worship and Biocentric Fallacies” as “commentary” in the Quarterly Review of Biology (1992). This essay is nearly identical to the one entitled “Mother Nature is a Wicked Old Witch” included in the collection Evolutionary Ethics (1993). The title of one is slightly less

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inflammatory than that of the other, but one can hardly doubt that Williams aims to provoke. When he disperses these essays among various publications, he also finds it useful to descend or reduce his argument to the lower, more grounded level by addressing the physics of global environmental conditions. At the same time, he expands his argument toward broader audiences with whom he can dwell on the cultural import of current sociobiological thought. Williams indicates that his most elaborate essay, the “Sociobiological Expansion,” was written in 1981–82 at Stanford University.

What Is Evolutionary Ethics? By definition, evolutionary ethics is situated in a dangerous territory between ethical theory and evolutionary theory.11 Those who explore this territory, or expand into its abyss, may seek a sociobiological source for human morality and ethics in the process of natural selection, perhaps even claiming that altruism as an adaptation improves social structures and improves the fitness of members of social groups. Morality can be imagined as a useful adaptation for increasing the fitness of its holders, providing a selective advantage. This appears to be what Edward O. Wilson means when he says that “scientists and humanists should consider together the possibility that the time has come for ethics to be removed temporarily from the hands of the philosophers and biologicized.”12 Philip Kitcher, a generally incisive philosopher and critic, responds to what he believes is a set of confusions by E. O. Wilson. Kitcher works out the logic of Wilson’s position in “Four Ways of ‘Biologicizing’ Ethics” in 1993. As he puts it, sociobiology claims to explain how people “have come to acquire ethical concepts, to make ethical judgments about themselves and others, and to formulate systems of ethical principles.” It also claims “to derive normative principles that we had not yet appreciated” and to “settle traditional questions about the objectivity of ethics.” Sociobiology thus aims to “lead us to revise our system of ethical principles … by teaching us new fundamental normative principles.” In which case, as Kitcher points out, “sociobiology is not just a source of facts but a source of norms.”13 He finds it a flawed source. There are plenty of pitfalls for evolutionary ethics. A primary one is known as the “is-ought problem,” which most writers say was first articulated by David Hume (1711–76). After he watched philosophical texts slide from description to prescription when dealing with moral matters,

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Hume reminded his readers that saying what is the case is unrelated to saying what ought to be the case. You can’t get easily to there from here. The “is-ought problem” is closely associated with the “naturalistic fallacy,” which involves attempting to define what is “good” by reference to other natural, empirically verifiable properties.14 George E. Moore, who named this fallacy in 1903, did so to point out a weakness in the social Darwinist thinking of Herbert Spencer, arguing that Spencer and his followers fallaciously maintained that evolution, “while it shews us the direction in which we are developing, thereby and for that reason shews us the direction in which we ought to develop.”15 Kitcher does not believe the principle of social Darwinism has lost its sway. When he finds E. O. Wilson claiming that “the naturalistic fallacy has lost a great deal of its force in the last few years,” he thinks Wilson’s claim “will ring hollow unless the type of argument leading from biology to morality is plainly identified.”16 Closely related to these issues is what Stephen Jay Gould and Richard Lewontin dubbed the “panglossian paradigm” in 1979, naming it for the absurd optimism of Pangloss, a character in Voltaire’s Candide who believes that this is the best of all possible worlds. Gould and Lewontin find the panglossian paradigm central to the adaptationist program, which wants everything to be specially adapted to suit specific purposes. For Gould and Lewontin, the adaptationism is panglossian because it posits “the near omnipotence of natural selection in forging organic design and fashioning the best among possible worlds,” so that adaptation “becomes the primary cause of nearly all organic form, function, and behavior.”17 When Williams combines evolution and ethics in the same sentence, he does not mean what E. O. Wilson means or what Herbert Spencer means. He is certainly no panglossian. Like Huxley, he speaks of evolution and ethics; he separates and distinguishes the two. Yet Williams may be caught between the horns of a dilemma when he argues not from but against the moral authority of natural selection, because showing what ought not to be constitutes an antithetical kind of naturalizing. A critique of naturalizing, as antithesis, has to grant status to that which it opposes. A specific target of Williams’s original critique in 1966 was a set of ecologists at the University of Chicago. In these essays of the late 1980s and early 1990s, his target has not changed, but he now aims for a different aspect of their thinking. As historian of science Gregg Mitman has argued, the same ecologists who created a biology that incensed Williams in 1966 also took nature to be “normative,” by which they meant that “nature’s economy” could be a source of healing and hope, a moral

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prescription for human society—that nature could teach humanity about morality. Generally speaking, a normative statement makes a claim about how things ought to be. Alfred E. Emerson of the University of Chicago, one of Williams’s targets, was a central figure in what came to be known as the Chicago school of ecology. Emerson coauthored Principles of Animal Ecology (1949) with Warder Clyde Allee and Thomas Park, a book that demonstrated the principle of group selection. The authors even criticized Weismann’s idea that germ plasm was completely isolated from the soma and from all outside influence, and were apparently willing to ignore other Darwinian theories of evolution.18 Gregg Mitman reads a moral from his history, that “nature cannot solve our problems, it cannot give us guidance, it cannot give us direction. Humans, not nature, are the sole arbiters of their fate.”19 His views are predated by those of Williams and David Lack, who wrote in 1969 that the world of birds, for instance, seems idyllic to humans until one looks closely and sees how cruel it is, for “we would not enjoy a society in which one-third of our adult friends and over four-fifths of the teenagers die of starvation each year.”20 Nevertheless, the ideas of the Chicago school have had a long life, lasting well beyond Williams’s 1966 critique, as Mitman has argued elsewhere: “Throughout the 1980s, scholarship in environmental history had a strong normative edge.”21 I have been one of those scholars. In the twenty-first century, controversies over the normative role of nature have become bioethical in ways Williams probably never imagined.22 Williams followed other biologists who had successfully refuted the scientific views of the Chicago school, yet in 1966 and beyond, their philosophical views and normative ethical values accounted for their popularity, and their take on ecology became “firmly entrenched,” as Mitman puts it. Either it is valuable to ask how human social values emerge from the biology of social behavior, or it is not. Williams confronts this emergence directly by drawing a sharp distinction between biological behavior and human ethics. Given his refusal in the “Defense” to factor in “theories of mind,” his discussions must focus on acts, not motives. From early in his career, Williams had been interested in social behavior. Recall that his first publication, coauthored with Doris C. Williams, was about the social adaptations of insects. He also carried out experiments in fish schooling.23 He found no evidence of any functional organization in schools of fish, which leads him to distinguish these kinds of groups from those of insects, where he believes he can find it. His interest in schooling became a

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keystone in the argument of Adaptation and Natural Selection, as Carl Zimmer and others have noted.24 Williams always focuses on the evolution of social behavior. In Adaptation and Natural Selection, the term behavior is mentioned on fifty-nine of the 274 pages of text; in Natural Selection (1992), the term appears on twenty-eight of the 151 pages of text. He pays especially close attention to the complexity of reproductive behavior, both sexually dimorphic and homosexual, as exhibited in courtship, spawning, nest building, parental care, and feeding, treating it sometimes as adaptive and sometimes as mistaken. But does “mistaken” imply a moral or quasi-moral judgment? He documents behavior that takes advantage of the behavior of other species, behavior as seasonally and even diurnally cyclical, including dominance hierarchies, group-related acts, predatory strategies, and learned tactics. His principle, expanded from Tinbergen, is that “instinct costs less than learned behavior, in the currency of genetic information.”25

“Incompetent to furnish any better reason”: The Contemporary Context Surely any version of evolutionary ethics that concerns Williams is of a certain historical moment; the position he advances comes at a point in the scientific history of knowledge of human behavior, and like T. H. Huxley’s, his perspective is limited by certain gaps. Some would say that Williams, like E. O. Wilson, invades the realm of philosophers, where he does not belong. Yet it is by no means clear that he thinks biology can replace philosophy or ethics, as Wilson does. He also invades the consciousness of his contemporaries. As one of Margaret Drabble’s characters says to himself, “He did not like the selfish gene, the spiteful gene, the inevitability of what he, in his old-fashioned way, could still call wickedness. He yearned still for the possibility of the generous gene, the sacrificial gene.”26 One should also remember that Huxley’s evolution is not Williams’s evolution, partly because of the late twentieth-century expansion of Darwinism, a more complete working out of Darwinian theories by modern biologists. Indeed, sociobiology itself can be defined as “an expansion of standard Darwinian evolutionary theory (which traditionally explains morphological adaptation) to a new domain: namely, animal sociality.”27

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E. O. Wilson sometimes frames sociobiology as an “extension of population biology and evolutionary theory to social organization.”28 Who exactly cares about these questions? Williams speaks and writes in sometimes strange company. The first version of his essay on Huxley appeared in 1988  in Zygon, an academic journal that seeks dialogue between scientists and theologians. In Evolution and Ethics, where it appeared in essentially the same form, Williams collaborated with a respected Victorian scholar to edit T. H. Huxley’s writings on the subject, contextualizing these historically as Victorian texts and scientifically in terms of modern biology. Williams wrote his Evolutionary Ethics essay for a 1990 symposium in Chicago that included noted biologists, philosophers, and historians. There he responded particularly to Robert J. Richards, whose book he had reviewed. Williams’s review of Richards’s book is telling in a variety of ways—and forms a bridge between what Williams thinks and what he finds in humanistic literature. In that book, Richards quotes and comments on a suggestive passage from Adaptation and Natural Selection where Williams recruits Darwin to argue for an evolutionary advantage for any individual “who maximizes his friendships and minimizes his antagonisms,” speculates that “selection should favor those characters that promote the optimization of personal relationships,” and imagines that “this evolutionary factor has increased man’s capacity for altruism and compassion and has tempered his ethically less acceptable heritage of sexual and predatory aggressiveness.” Yet, as Richards points out, Williams pointedly does not note “that Darwin thought the principle cause of moral behavior would be group selection.”29 Williams has high praise for Richards’s text, calling it “a work of outstanding merit and importance.” He approves its central question, “how is objective science to come to terms with the phenomenon of mind?” But he expresses surprise at Richards’s sympathy with William James’s idea that “the brain [is] a machine controlled by the mind in the way that a captain controls a ship,” a position rife with problems. Williams notices that Richards is “willing to accept without question that mind is something that evolved, and therefore that its development must have been favorably selected.” But “how does one document the evolution of mind?” Williams sees this as a major problem for materialist explanation: “Until mind can be shown to have parameters in common with matter, theories such as Darwinism, which are designed to deal with matter, can never serve for dealing with mind.”30

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Richards credits Darwin as founder of group-selection theory and claims that “Darwin thought the principal cause of moral behavior would be group selection.”31 Williams disagrees in his usual way: “He was the peerless giant of nineteenth century biology, but he did leave a few problems unsolved for the attention of his followers.” When Richards offers “A Defense of Evolutionary Ethics” as an appendix to his history, Williams finds it “a debacle of ideas decisively refuted by Huxley almost a century ago and perhaps by Hume a century before that,” and he accuses Richards of muddling the discussion.32 As a parting gesture to Richards, a sort of tit for tat, Williams quotes a passage by Huxley, who disparages those who look for the origin of “moral sentiments” in the process of evolution: I have little doubt, for my own part, that they are on the right track; but as the immoral sentiments have no less been evolved, there is, so far, as much natural sanction for the one as the other. The thief and the murderer follow nature just as much as the philanthropist. Cosmic evolution may teach us how the good and the evil tendencies of man may have come about; but, in itself, it is incompetent to furnish any better reason why what we call good is preferable to what we call evil than we had before.33

Truly, any discussion of altruism cannot have things both ways. Yet, his review of Richards and his essay in Zygon may be Williams’s way of beginning a discussion, not ending it as Huxley had claimed to do. In later book projects, Williams increasingly calls attention to widening contexts for his ideas in literary traditions, and he attempts to speak to those contexts, to demonstrate their value, by including extracts from classic writers, like Galen, William Paley, and others.

Thomas Henry Huxley’s Garden in a Contemporary Context In the meantime, the writing of T. H. Huxley provides a perspective that Williams finds useful as he engages in his “expansion.” Apparently, T. H. Huxley coined the term “Darwinism” for Darwinian ideologies, but he was certainly no social Darwinist. Yet advocating Huxley’s perspective comes with a risk—it could encourage some sociobiologists to create norms or propagate a new form of social Darwinism, especially among those who do not understand the science or get it secondhand in what Williams refers to as “printed tracts.”

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Countless arguments about the relationship between social Darwinism and sociobiology emerged as a result of E. O. Wilson’s On Human Nature (1979). Mary Midgley (1919–2018), the influential British moral philosopher, is particularly and perhaps representatively acerbic. She finds sociobiology to be defined by the libertarian tradition of Spencer: “Herbert Spencer, elaborating that crude interpretation to suit his extreme libertarian views, had produced that very unDarwinian thing ‘Social Darwinism’— the idea that competition between individuals was the central law of life and must never be interfered with.”34 Richards argues that this is a misrepresentation of Spencer. In any case, Williams chooses to follow Huxley precisely because Huxley allows him to argue for the opposite course from the one Midgley caricatures; Williams advocates interfering with the war-­ like central law of competition, and analogizes his position to that of a pacifist. By providing an expansion of Huxley’s ethics, Williams is identifying himself as opposed to at least one form of social Darwinism, though he does not use that expression. The contentious context of the so-called sociobiology debate also leads Williams to argue for the relevance of T. H. Huxley’s Evolution and Ethics. He writes, “to me the important message in Huxley’s superb essay is its development of the nature-as-adversary theme, which no subsequent work has improved upon.”35 In a much-discussed metaphor, Huxley imagines the garden as a work of art or artifice, as an antithesis to the works of nature. But of course Huxley’s garden, an elaborate metaphor made of words, is itself an artifice. James Paradis explores this trope and observes that the argument of Evolution and Ethics is “nested in the fable of Jack and the beanstalk,” a story to which Huxley alludes. Huxley likens his ideal garden to an imperial colony where colonists “clear away the native vegetation, extirpate or drive out the animal population, so far as may be necessary, and take measures to defend themselves from the re-­ immigration of either.” They introduce English crops, English dogs, English livestock, English men; they “set up a new Flora and Fauna and a new variety of mankind, within the old state of nature.”36 These gardeners select, include, exclude, cull, and uproot what they do and do not prefer. As Paradis points out, Huxley’s artificial world is “none other than English industrial civilization.”37 About this imaginary garden, Williams is mute, but I cannot be. Administered as an “earthly paradise, a true garden of Eden,” Huxley’s garden is designed for “the well-being of the gardeners,” who exclude the “cosmic process,” that “coarse struggle for existence of the state of

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nature.” Within, everything is “adapted to human wants, and would perish if human supervision and protection were withdrawn.” Even human denizens are selected for their efficiency and function. Huxley therefore insists in his conclusion that social progress requires “checking of the cosmic process at every step and the substitution for it of another, which may be called the ethical process,” in order to insure not the survival of the fittest, but of “those who are ethically the best.” Huxley finds the cosmic process and state of nature hostile to the art of the garden and to the principles of horticulture. He wishes to abolish the state of nature and replace it with this state of art, but he knows, perhaps like Williams, that it cannot be so. He foresees instead an eternal battle, where “the state of art, here and elsewhere, can be maintained only by the constant counteraction of the hostile influences of the state of nature.”38 To my knowledge, Williams does not imagine any Edens, nor does he praise gardens in any writings. But, like Huxley, he wonders whether humans can garden themselves. It would have been impossible for Huxley and perhaps even for Williams to imagine that humans might someday believe that they could garden their own genome, for, as Huxley wrote, “the theory of evolution encourages no millennial anticipations.”39 Huxley remains a perplexing figure, as James Paradis observes, “a man of two visions, the one filled with hope and wonder, the other dominated by a sense of futility and doom.”40 Paul Lawrence Farber writes the story of Huxley’s two visions as a story of his decline to pessimism: “as he aged, his optimism increasingly was overcome by pessimism concerning the condition of man,” and so, “late in life, in such a mood, he wrote his major work on evolution and ethics.” According to Farber, Huxley’s “liberal politics made him uncomfortable with the conservative views of those who sought to justify a laissez-faire, noninterventionist state policy by reference to natural selection in the realm of morality and social institutions.” There is much evidence to suggest similar motives led Williams to this position—images in Huxley’s writing of demagoguery, of ignorance from the pulpit, of corruption and the abuse of power. According to Farber, Huxley’s “ample political motivation for his antagonism to naturalistic ethics” was behind his public analysis of “the intellectual weaknesses of evolutionary ethics.” And as Farber points out, Huxley is a perfect example of how one could be “an ardent supporter of evolution and scientific naturalism, and not embrace evolutionary ethics.”41 “Incompetent to furnish any better reason”: this is exactly the sort of phrase Williams would notice in Huxley, and it becomes his own premise.

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Out of temperamental or intellectual affinity, Williams latched onto Huxley’s paradox and its language. Why examine the charnel house of evolution so minutely? The point is easily made. Williams wishes to demolish the authority of traditional views of nature, especially the authority of traditional cultural dicta that make normative judgments from nature.

How Could Nature Be Normative? The Expansion Itself Williams claims his purpose is merely to “update Huxley’s message and to characterize the enemy more clearly than was possible in Huxley’s time.” Yet he goes one step further when he offers an “updated program for the betterment of the human condition” as “a twofold attack on the natural enemy and any institutional enemies favored by cultural evolution” in the age of a nuclear arms race. The twofold attack is embodied in the central analogy of the “Sociobiological Expansion” and comes early. Huxley’s phrase is the “moral indifference of nature,” but Williams writes, “I am inclined to substitute gross immorality for moral indifference,” attributing “the weakness of Huxley’s version mainly to weakness of his grasp of natural selection.” Williams continues, “my use of immoral is like that of a pacifist who maintains that war is immoral.” No matter why wars start, a pacifist would not speak in terms of moral indifference, but would rather confront immorality directly, just as “a pacifist regards war as immoral because it frequently and systematically produces results that he finds morally unacceptable.” This is to take seriously the military metaphors so often used by evolutionary biologists like Dobzhansky, metaphors that are no doubt rooted in Tennyson’s “Nature red in tooth and claw.” When such metaphors morph into terms like “arms race” in a nuclear age, Williams says, “I hope that my discussion will show, at least to those who would concede that war is or may be immoral, that organic evolution is worse than traditional forms of warfare, and worse than Huxley imagined.”42 Williams finds it necessary to teach reductionist evolutionary theory to his audience. Take Hamilton’s rule, which Williams calls “an expansion of Darwin’s concept of fitness to include the ability to get one’s own genes represented in future generations, whether these genes are present in one’s own cells or present, for genealogical reasons, in relatives.” Hamilton’s rule is typically expressed in textbooks as C > R × B, where C is the cost in fitness to the actor, R the genetic relatedness between the actor and the

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recipient, and B the fitness benefit to the recipient, and the sign > means “greater than.” Sometimes, as in Hamilton’s original article (1963), the expanded mathematical equations are formidable. Williams turns what normally has been expressed as mathematical formulas into prose: “[Hamilton] termed this broader concept inclusive fitness. His key formulation was that assistance to a relative will be favored if the benefit to the relative, times the coefficient of relationship, exceeds the cost to the donor. Costs and benefits are always measured in the currency of reproductive success.”43 Williams continues to use Hamilton (and Trivers) to argue that biologists usually find “altruistic behavior is limited to special situations in which it can be explained by one or more of three possible factors”: nepotism, manipulation, and reciprocity. Therefore he does not consider it profitable to look for the origin of cultural conceptions of altruism in natural science. The phenomenon of manipulation plays a peculiar role in this argument, as it slips from part of the evolutionary process into a realm of modern social life that is often dramatized by the presence of Melvillean confidence men. “Anyone who makes an anonymous donation of money or blood or other resources as a result of some public appeal is biologically just as much a victim of manipulation as the snapper in the jaws of the anglerfish.” Biologists like Williams are supposed to reveal realities about the natural, biotic world, but this kind of statement crosses different kinds of realities. As for reciprocity, for instance, “in nature [it] is strictly limited by the necessity of safe-guards against cheating,” but those realities impinge upon any public view of social perspicacity.44 Manipulation and deception are integral to cultural intercourse, possibly to the evolution of language, and require suspicion on the part of the individual. He says that philosophers like Peter Singer miss these aspects of supposedly altruistic biological behavior.45 At the same time, Williams speaks directly against “normative biologism,” using a term coined by the psychologist and sociologist D. T. Campbell. Campbell rejects “the view that what is biologically natural is normatively good. Such a normative biologism underlies much of current enthusiasm for sociobiology,” he argues. “It is implied, for example, even in E. O. Wilson’s popular accounts of sociobiology … in spite of explicit disavowals in his more formal presentations.”46 Williams thinks that anyone who looks closely will see a litany of misery in the process of natural selection, though language used in public hides what humans refuse to see. Those who object to “accusing animals of practicing slavery,

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adultery, rape, or other sins” appear to have no problem using terms like courtship or singing, and “even highly disciplined biologists would rather study what they find appealing than what they find less so.” No wonder that “biologists spend more time studying defense against predators than against parasites, even though animals spend more time scratching at small enemies than fleeing from large ones.”47 These texts on evolution and ethics work out more completely and more publicly what Williams implies in Adaptation and Natural Selection (1966): “Attempts to demonstrate the benevolence of Nature often take the form of name changing.”48 In 1966, he had little hope that his ideas would be noticed. Now that they have been, he carries them into a broadly public realm and provides science without mind-numbing math. Williams continues to be an iconoclast. His “Sociobiological Expansion” is compressed, dense yet nuanced, concise not only in the way he frames the argument, but in decisive topic sentences. His expansion iterates a litany of biological conflict: fights over mates and with mates, fights between offspring, adultery. Life in the waters is ravenous, but so too on land and among the “higher beasts.” He documents cuckoldry and rape and escalates through examples of conspecific destruction, cannibalism, and infanticide. He reports the tears and anger that reputable scientists like Sarah Blaffer Hrdy experience while observing langur family life and alludes to the desire to “cry shame” among amateur naturalists like Annie Dillard. He documents excess carnage by predators who kill more than they can eat. Williams discusses krill, whales, and the violence of the deeps, perhaps for the same reason that Herman Melville dwelled upon pilot fish. For instance, “the stomachs of large walleyes contained smaller ones, which had eaten still smaller ones, for at least a fourfold cycle of cannibals within cannibals.”49 What follows is something like an epic simile: “I will end this section with a thought experiment to give final emphasis to the world’s wretchedness that Dillard deplored.” Imagine, he says, a “device of great beauty and utility” like an “artfully designed watch,” and imagine a large number of them, all reliable and accurate, and then imagine someone who “to secure some minor gain” smashes each with a hammer. “For me, a personally attractive wielder of the hammer would make the deed worse, not better. If the act by the hammerer were compelled by some cosmic process, it would not absolve the blame, but only transfer it.” So “replace each watch with something far more worthy of admiration, a krill (Euphausia superba) … a beautifully streamlined animal of shades of red,

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with optically precise photophores on eyestalks and abdomen.” Then “on comes the blue whale, to me an extremely appealing animal,” that nets hundreds of thousands of krill, so that “a force of many tons ends hundreds of thousands of lives.” This process is repeated many times a day. “The life history of one whale is merely a microscopic detail in the carnage of the seas.”50 To compare great things to small, the evolutionary process is immensely powerful and oppressive, but unlike Job’s God, Williams writes, it is “abysmally stupid.” Williams cannot, like Huxley, depict the Book of Job as “ancient tragedy,” with its “unfathomable injustice of the nature of things.”51 Tragedy is not compatible with victimization by “stupid forces,” for there is no dignity to be found in such a drama. The trouble with naturalizing morality by antithesis—as Williams does when he depicts the natural world, or God’s world, as immoral and cruel—is that such a move does not absolve what is also patently stupid or cruel human behavior. In an era when human consumption is so obviously thoughtless, destructive behavior is no more admirable than that of the blue whale, and may be worse because potentially ruled by conscious decisions. But do humans have a choice? Suggesting that they do or don’t—both seem risky moves for a sociobiologist. No wonder humans are so suspicious. On two facing pages, Williams speculates about the origin of suspicion and turns it to his purpose. In its origin, suspicion “aided the receivers of messages in deciding” whether they gain or lose personally by complying or defying. “These can be problems of great subtlety,” but, as a questioning ability, suspicion is “designed to spread selfish genes”: “In its boundless stupidity, this evolutionary process incidentally designed machinery capable of answering other sorts of questions, such as, Is the message one of help or harm for what I really want for the world?” And answering such questions “might well serve morality rather than selfishness” because “we can use abilities developed for petty intrigue to deal with sermons coming from the pulpit, printed tracts by sociobiologists, or manipulative arguments from politicians on television screens.” “I have some hope,” Williams writes, that the process might “provide the humane artifice that can save humanity from human nature.”52 Humans are different from other creatures, though why they are different is as much a mystery to Williams as to Huxley. How might a species arise that would respect and help fellows, and be devoted, in Huxley’s words, “not so much to the survival of the fittest, as to the fitting of as

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many as possible to survive”? The answer surely lies in the public and cultural world. In the concluding section of his essay, called “Prospects for Morality in an Immoral World,” Williams seconds Peter Singer’s principle that “the human capacity for reasoned argument constantly imposes a need for the public justification of personal action.” One should suspect everything, as Williams argues, including sociobiological tracts. Yet also, as he cites and quotes from Peter Singer, “a biological explanation is often a way of debunking the lofty status of what seemed a self-evident moral law.” As he writes, “the sociobiological imperative is thus a negative one: beware of manipulation by selfish individuals, or selfish institutions, or our own selfish genes.”53 This negative sociobiological imperative must also apply to Williams. He enters the debate, pretends to stay aloof, and mentions that Huxley, being an Englishman, was prone to understatement. It is harder to know about Williams himself, because he is prone to deliberate paradox. Huxley was the first to command his readers to “refuse any longer to be the instruments of the evolutionary process.” As Williams nears his conclusion, he writes, “In Dawkins’ more modern terms, we must ‘rebel against the tyranny of the selfish replicators.’”54 It is interesting that he uses the same commandment by Dawkins in both essays and concludes the Zygon essay but not the “Sociobiological Expansion” with it. Surely he endorses Dawkins’s call of Non Serviam, but what of Dawkins’s reasons? Were his views congruent with those of Dawkins or was Dawkins shadowing his? In A Devil’s Chaplain, Dawkins sets out an array of perspectives: “I prefer to stand up with Julian’s refreshingly belligerent grandfather T. H. Huxley, agree that natural selection is the dominant force in biological evolution and unlike [George Bernard] Shaw, admit its unpleasantness unlike Julian [Huxley], and, unlike [H.  G.] Wells, fight against it as a human being … That is G. C. Williams’s recommendation today, and it is mine.”55 One must read Williams’s Natural Selection (1992) closely to see exactly where they are in agreement and where they are not.

Call and Response Williams’s essay on Huxley received four commentaries that were published simultaneously in Zygon. They were written by sociobiologist Sarah Blaffer Hrdy, professor of history and philosophy Michael Ruse, the journal’s founding editor Ralph Wendell Burhoe, and professor of theology John B. Cobb. Williams responded to them vigorously. Like later writers,

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commentators in Zygon seem puzzled by Williams’s pessimistic view of the prospects for naturalizing morality and prefer to distance themselves by characterizing him as old-fashioned—as, at the age of 63, “the grand old man of gene selectionism.”56 When Williams depicts evolution as “the tenacious and powerful enemy” of all morality and goodness, his stance is sometimes seen as an expansion of Huxley’s “gladiatorial” approach against nature as a whole. This is an interesting reversal of Huxley’s position, which advocates self-restraint in place of ruthless self-assertion. Huxley wants individuals to not merely respect but help their fellows, to facilitate “the fitting of as many as possible to survive,” and in consequence, his position “repudiates the gladiatorial theory of existence.”57 Williams encourages no belief that an organism can be designed for any purpose other than the most effective pursuit of its own self-interest. For him, only culture can emancipate us from the dictates of our genes precisely because, powerful and oppressive as the evolutionary process might be, “it is abysmally stupid.” Evolution can only “maximize current selfishness at the level of the gene,” it is blind to any future, and “it does not have the sense to realize that mechanisms evolved for practicing unfair nepotism or making self-seeking deals with others can be subverted in the interests of broad altruism.” This is the opening Williams seeks. The process can be subverted, as Huxley too argues, so that ethical nature, though “entirely unrelated to the selfishness responsible for the changes,” can emerge “incidentally” from biological nature. “The helping hand of the good Samaritan and the motivation for its use raise no question of the malice or power of natural selection. They merely show that this persistent and powerful enemy is a mindless fool.”58 This view is so deeply American that one wants to shake one’s head at any respondent’s surprise. What literate reader would not remember Thoreau in “Higher Laws,” at the center of Walden, vilifying the selfish sensual impulses to “eat, or drink, or cohabit, or sleep sensually” and insisting that “Nature is hard to be overcome, but she must be overcome.”59 The question of beneficence of the evolutionary process is always related to the imagined source for human altruism. In Zygon, the subject predictably jumps to the question of human nature.60 Sarah Blaffer Hrdy believes that “No informed person could sensibly turn to nature for templates in moral matters, and no one makes this point more persuasively than Williams does.” “Indeed,” she argues, “we would be hard put to come up with a species-neutral or truly universal moral system.” For that reason, she questions “whether Williams or anyone else can come up with

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a broadly acceptable moral system that will not in some respects be constrained by the legacies of selfish and kin-selected replication.”61 Ruse is less affirmative when he takes issue with the language Williams uses. They argue over rattlesnakes—or other snakes in the garden. Williams distinguishes between being struck by lightning and being struck by a rattlesnake: “Only the intellectually dead could fail to see that the snake has what are clearly weapons, precisely designed and used to produce a victim.” As Williams says, any biologist recognizes “the major difference between a morally accountable human mind and a snake mind.”62 Ruse argues that “Williams is insensitive to the metaphorical use of such terms as weapon and design,” and he is surely right that Williams creates a metaphor by conjoining the terms weapon and design. Ruse continues less convincingly that, “if we grant that the snake is striking blindly, there really is no moral difference between it and the lightning. Otherwise, why do we grant such defenses as: ‘Not guilty by reason of mental incompetence?’”63 Williams answers this confusing set of assertions and comparisons decisively, but surely he is wrong when he asserts that “there is nothing metaphorical in my recognizing that the snake’s fangs are designed as weapons.” He tries to explain how, when the snake poisons its victim, “in endless details the observable features of the machinery conform to the predictable design requirements for an envenomating apparatus.” The mechanical and behavioral design of the snake guarantees that it does not strike blindly, but on the contrary, precisely. “What else can one mean by functional design other than such mapping of observable engineering onto functional requirements? Nothing is implied about the nature of the designer”64 Williams does revise and qualify his tone as a result of Ruse’s commentary, and this process reveals a great deal about difficulties in writing about animals. In the “Expansion,” Williams omits his quip about “the intellectually dead” and concedes much more, including “the greater evil of murder by a human being over predation or lethal self-defense by a rattlesnake and the absurdity of holding the reptile accountable for its actions.”65 Ruse, who always considered himself a friend of Williams, concludes, “We agree in being evolutionists, Darwinians, and—a point on which we part company from well over ninety percent of my fellow philosophers—that in our biological origins lie both the questions and answers about our moral natures. Ultimately, that is what is really important.”66 It is not clear that Williams would agree to such a statement, but Ruse and Williams continue their friendly disagreements for decades. In 1994, for instance,

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Williams argues with Ruse about Christianity in Zygon, where he says that he is “in general agreement with Ruse on most religious and scientific issues,” but he is troubled by what he sees as Ruse’s “partial return to Christianity,” to the point of torturing his own syntax: “His recognition that religious establishments have been responsible for much personal stress and many of history’s great tragedies is understated.”67 Williams prefaces his responses of 1988 by noting, “my most basic conflict is with the idea that evolution is good in general or on the average, as stated or implied by everyone except Hrdy.” His rejoinder to the theologically minded is vigorous and sly. For instance, he puts his thesis into the mouth of a critic, John B. Cobb, writing: “Even in some of the criticisms there is much that I find agreeable, most notably John Cobb’s recognizing that an unremitting effort is required to expand the circle of sympathy for others. This effort is in opposition to much of human nature and is surely the struggle proposed by Thomas Huxley and seconded by me.”68 He is adamant.

The Groundless Gaia Hypothesis Following the “Sociobiological Expansion,” Williams turns to the physical environment, critiquing the popular idea of Gaia in two nearly identical, though differently titled, essays. Although “Mother Nature is a Wicked Old Witch” (1993) is surely his most notable title, I will explore “Gaia, Nature Worship and Biocentric Fallacies” (1992) because the latter text includes a more complete argument.69 James Lovelock and Lynn Margulis advanced the Gaia hypothesis in the 1970s. Lovelock is often described as an independent scientist, environmentalist, and futurologist, and Margulis as a distinguished microbiologist. An assessment of their ideas finds that their Gaia proposal consists of three central propositions: “(1) that biologically mediated feedbacks contribute to environmental homeostasis, (2) that they make the environment more suitable for life, and (3) that such feedbacks should arise by Darwinian natural selection.” James Kirchner, professor emeritus at UC Berkeley’s Department of Earth and Planetary Science, argues that “these three propositions do not fare well under close scrutiny.”70 Because Williams is more aggressive in his critique, he adopts a satirical style. In retrospect, one can see how the ironies that characterize Williams’s early style move closer to satire in some of his later works, particularly in this essay. The shift in tone continues in Plan and Purpose in Nature

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(1996). Because these works were written for a wider public with whom he does not agree and for whom he often has little respect, it is not surprising that Williams should turn to a satirical mode, especially when one understands the traditional function of satire. It is nearly redundant to speak of intellectual satire. As an authorial strategy, it is associated with detachment. The satirist speaks with the voice of an outsider to pose as an outsider. In Anatomy of Criticism, Northrop Frye shows how irony and satire exemplify a mythos—or plot—of winter and cold reason, where “the central principle of ironic myth is best approached as a parody of romance.” Frye notices a chief distinction between irony and satire, namely that “satire is militant irony: its moral norms are relatively clear, and it assumes standards against which the grotesque and absurd are measured.” Frye argues that only when a reader is in doubt about an author’s attitude does one see irony with little satire. And in the opposite case, satire becomes “irony which is structurally close to the comic.” Scientists often use satire as a traditional genre to combat superstition. Indeed, satire probably began with Greek silloi, satirical poems which Frye calls “pro-scientific attacks on superstition.” This way of defining satire seems to fit Williams’s aims. Further, Frye also notices that “the romantic fixation which revolves around the beauty of perfect form, in art or elsewhere, is also a logical target for satire.”71 Williams’s satirical tone reveals some of his impatience with pseudoscientific thinking and may even unmask his own desperation as an outsider. Williams knew that satire was a natural choice under the circumstances, whether he began with the aim of satirizing or drifted into this tone. Satire is often the tone of critique. “The idea that the universe is especially designed to be a suitable abode for life in general and human life in particular,” Williams writes, is an old one that “had to be abandoned in its early forms with the triumph of Copernican astronomy.” That geocentrism has returned in various forms disturbs Williams, particularly its “main modern manifestation,” the Gaia concept put forth by Lovelock and Margulis.72 Williams disapproves of two particular essays. Richard A. Kerr’s “No Longer Willful, Gaia Becomes Respectable” is a portrait of Lovelock, and Charles Mann’s “Lynn Margulis: Science’s Unruly Earth Mother” is a puff piece.73 What Williams finds offensive is not only that they turn scientists into celebrities, but they present a totally uncritical view of the science itself. He descends to the lowest level to show how the beneficence of nature, as posited by the Gaia hypothesis, may or may not be exhibited by the

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physical environment. For instance, it is often claimed that life is “closely dependent on the special properties of water,” but rather than showing the benefits of this situation, Williams looks for its adverse effects. The high specific heat of water is “clearly disadvantageous for any reptile or insect, or other animal that depends on basking to reach a favorable temperature on a cold day.” He entertains the idea that only a moderate effort would be required to demonstrate the “thesis that our physical environment is diabolically perverse.” Snow has the wrong color for thermal efficiency; any outside observer or extraterrestrial visitor would notice that freezing water kills by mechanical disruption, and sunlight contributes the “lethal nature of both the ultraviolet and visible wavelengths,” while the high concentration of oxygen “would force the conclusion that all materials at the Earth’s surface must be oxidized (i.e. dead).”74 The earth is a hostile environment, Williams insists, in which “living organisms are elaborately adapted to their particular ways of life in the environments in which they evolved. There is no evidence for any other kind of adaptation.” Indeed, “adaptation is always asymmetrical: organisms adapt to their environments, never vice-versa.”75 For Williams, the mechanics of evolution don’t work any other way. This view is contrary to Richard Lewontin’s argument, in The Triple Helix, that all organisms alter their environments. Nevertheless, reductionism being the opposite of holism, Lewontin too finds the “extreme holism” of the Gaia hypothesis sadly lacking.76 Williams thinks that Lovelock’s concept of Gaia takes mutualism, the idea that species obtain mutual benefits while paying individual costs, to a level of absurdity. This kind of idealism about biotic relationships creates a public scandal that calls for a satirical response. In ecosystems, as Williams always insisted, “the really good examples of mutualism are relatively rare.” Scientists look for mutualism when it isn’t there for ideological reasons, as he has said over and over—thus the steady production of books and essays claiming to show a benevolent and acceptable nature as a guide to ethics and human behavior. “In some cases it would appear that ‘love thy neighbor’ must stand or fall according to whether mutualism or parasitism is the more prevalent phenomenon.”77 Gaia, Williams writes, is the “modern form of the idea that adaptation is symmetrical.” He continues, “its logical structure is not entirely clear to me, but it seems rather similar to the invisible hand in economics. If each organism freely pursues its own interests, the average result over the ecosystem will be favorable to organisms.” The trouble is that “discussions in

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support of Gaia are often unclear as to what sort of interests organisms are supposed to be pursuing.” Gaians are “overtly group-selectionist” as one would expect, and though Williams does not deny the possible but minor influence of the phenomenon of group selection, “if the Gaia concept depends heavily on group selection, it should not be represented as being in harmony with modern Darwinism.” Lovelock advances the idea that earth is a self-regulating system and that Gaia, a “total planetary being,” maintains stability and “practices homeostasis.” But that’s absurd, writes Williams, because “homeostasis requires a feedback loop” and “should be recognized only if special sensory and control machinery can be shown.” Stability cannot be “uncritically attributed to homeostatic machinery.”78 Williams writes in the first person. Though he says he doesn’t know whether Adam Smith’s invisible hand can manipulate economies in the way he proposed, “I do know that the biological equivalent often manipulates ecologies in maladaptive ways.” He gives an example of trees that grow in close proximity with resultant fires; he shows how the productivity of marine communities is subverted by a process that creates insufficient nutrients where there is light and insufficient light where there are nutrients. The lack of fit in marine environments would seem to indicate Gaia’s destructive influence, but Williams wonders, “is it really Gaia’s role to maximize marine productivity?” Lovelock believes that Gaia prefers more productivity, and Williams recalls Lovelock saying that “the health of Gaia is measured by the abundance of life.” But, Williams objects, “if so Gaia must be sick indeed over most of the earth, where the biological community is a desert and life’s abundance at a paltry level.”79 “What would a really fit environment be like?” Williams wonders. Apparently, Lovelock believes that Gaia maintains early habitats between 0 and 30 degrees Centigrade (32–86 degrees Fahrenheit) “because that is the most favorable range for organisms.” Such an assertion begs the adaptive question or turns it upside down. Organisms can tolerate extreme conditions and do so, as Williams argues: “organisms will be most abundant under the most common and persistent conditions, where they have had the greatest opportunity to adapt.” The most diversified communities are simply the oldest, or so he argues. He may be right, in the sense that they are most resilient in the face of disturbance. But he wonders if the preoccupation of Lovelock and others “with the maintenance of the narrow range of conditions normally experienced by human life seems a little parochial” and not simply geocentric but even anthropocentric.80

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By and large, neither evolution nor ecology works by mutualism. If one thinks in terms of the biomass of the earth, what portion of life practices mutualism, and how might that be related to how many moral animals there are? Surely the principles by which any process of evolution proceeds cannot be deduced from the sameness or difference of humans from primates and other mammals. Williams’s distinction between cooperation and reciprocal altruism is conceded even by some, like Frans de Waal, who dislike Williams’s analysis. As de Waal writes, “Unlike simultaneous co-­ operation or mutualism, reciprocal altruism involves exchanged acts that, while beneficial to the recipient, are costly to the performer.”81 Williams believes that popular anecdotal examples for biological mutualism are usually examples of reciprocal altruism. There is little cooperation to be found in the evolutionary process. Though Williams summarizes some previous materials in his “Mother Nature” essay, and does not do so in the “Gaia” essay, both include this allusion to Huxley’s condemnation of those “who look to Nature for an ‘exemplar for human conduct’”: “The ‘moralizing of sentimentalists’ continues to work its mischief in the use of biological material intended for the lay public.”82 To sum up, because “the universe is hostile to life in general and human life in particular,” and “the hostility of the universe is abundantly shown by its general physical properties,” natural selection can be characterized by “moral perversity.” Therefore, it is well to follow “Huxley’s call for combat against the cosmic process.”83 Why indeed would Williams portray “mother nature” as a “wicked old witch” in 1993—rather than rejecting the concept of “mother nature” altogether, and simply separating biology from the moral world? The easy answer is that he is just being satirical, but the issue is complicated by realities. In 1999, Sarah Blaffer Hrdy refuses to separate biology from morality in her brilliant book on mothering, Mother Nature: A History of Mothers, Infants, and Natural Selection. Instead, she confronts Williams’s view directly, saying, “Once we understand that natural selection has neither morality nor values, a concept like ‘Mother Nature’ ceases to be shorthand for romanticized Natural Laws that are more nearly wishful thinking than objective observation of creatures in the world around us.” And so her initial question is: What do we mean by “maternal instincts,” and how are they associated with familial love and “shaped by processes that by current standards can only be viewed as inexorable?”84

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Morality as a Veneer: On Pickup Trucks and Fires Much of what Williams publishes later in life may be inherent in Adaptation and Natural Selection. As he spins out down-to-earth philosophical implications in the sociobiological essays, or Huxley essays as I prefer to call them, he also emphasizes a greater role for historical contingency. But, Richard Alexander quips, “as George Williams once noted, if your pickup truck does not start, you don’t generally feel it necessary to start doubting the fundamental laws of physics.”85 In later writings, Williams seems fond of analogies that use pickup trucks, smoke alarms, pocket watches, and other ordinary mechanisms, as if these might bring his readers down to earth by reducing cosmic processes to mechanicals. His critics usually prefer to deal with cosmic questions in the abstract. It is comforting to imagine that design by nature is good, and so Williams enumerates historically constrained designs by nature in ordinary objects. In Why We Get Sick (1995), he writes about a popular line of pickup trucks whose gasoline tanks are located outside the frame, so that the trucks catch fire when struck from the side. “But to locate the tanks within the frame would require a major redesign of everything now there, which could cause new problems and require new compromises. Even human engineers can be constrained by historical legacies.”86 It has been said that biology became the new philosophy after Darwin, or after the modern synthesis. Since then, every generation has discovered biology anew as the new philosophy. Certainly biology has always been viewed as philosophy and psychology. The new sociobiology of E. O. Wilson, the new psychology of Stephen Pinker, Richard Dawkins, or Daniel Dennett, and possibly even the new ethics of Richard Alexander have all been criticized as the new ideology by Richard Lewontin.87 Nevertheless, journalists announce that neuroscience is the next new philosophy. Williams can take part of the credit for the increased frequency of these kinds of sayings in the late twentieth and early twenty-first centuries; yet, he also wants to say that biology cannot assume any such role until it goes beyond speculation. He also asks a more difficult ethical question— “is biology destiny?”—and his answer is both complicated and ambivalent. Yes, he says in one regard—for instance, we all die—and no, he says in another regard: humans need not live according to their biological origins, or value their origins as beneficent. Following Williams, Richard Alexander has argued since the early 1980s that “the culture-biology [or culture-genetic] dichotomy derives

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from continued misunderstanding and misstatement of certain aspects of biological theory.” The first three misunderstandings are a result of a complex that includes belief and fear: 1. Fear that to admit an influence of genes on behavior will lead to genetic determinism. 2. Belief that learned behavior—and by extension, culture—has no connection to genes. 3. Fear that accepting a potency for individual level selection suggests justification for selfish behavior.88 Fear number three has led to alarm by primatologists who fear not only justifying selfish human behavior, but especially justifying views of primate behavior as selfish. In this context, Frans de Waal inveighs against what he calls “veneer theory,” which posits a very thin layer of morality in humans and other primates overlaying less noble characters. He sources the veneer theory in Huxley and Williams; he watches it spread into a shape of thought; he believes this premise demands refutation.89 His is one of many complaints about evolutionary theory’s usurpation of the realms of humanistic disciplines. In all of these complaints, one sees conflicting “levels” of discussions of evolutionary processes and evolutionary products, a problem exacerbated by selecting only certain processes or certain products. To repeat Williams’s question, in terms of the earth’s total biomass, how many moral animals are there? One thing is sure. The mass of life on the earth is not made up of moral beings, and most life has no moral or ethical consciousness. Surely, the process of natural selection cannot be inferred from narrow studies that posit the sameness and differences of moral consciousness in primates, or even among mammals. Williams asks for a wider view. He suspects the desire to see ourselves and our fellow inhabitants of the earth as “good natured,” as the title of Frans de Waal’s book has it. Young dogs, as we know, learn to moderate the strength of a bite, learn bite inhibition from their mothers as part of what observers see as juvenile socialization. When in the company of mother and siblings, a pup learns that biting too strongly ends play, nursing, or feeding, and also is indoctrinated into a so-called dominance hierarchy. But it would be a serious error to assume that this process is universal in carnivores, that it is generally representative of socialization throughout the biological or even the canine world, or that it illuminates the origin of

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human forms of socialization. Our domestic animals allow us to mislead ourselves terribly. It is reasonable, in this context, to remember Darwin’s definition: “A moral being is one who is capable of comparing his past and future actions or motives, and of approving or disapproving of them. We have no reason to suppose that any of the lower animals have this capacity.”90 However, humans are moral beings, and with that comes obligations to the other beings, as I believe and probably Williams believed. If Nature is good for humans, we will have to enact certain behaviors, many of which we have not been evolved to value. Our consciousness is not continuous with that of the others, as Paul Shepard calls them.91 We must make the connections. Darwin’s definition of a moral being is embedded in a long discussion of the continuities and discontinuities between humans and other living beings. Kitcher stresses Stephen Jay Gould’s distinction, that “in any evaluation of our evolutionary history you can emphasize the continuities or discontinuities.”92 There is a huge difference between (1) an author like Williams indicating how evolution works, and consequently how unlikely it is that moral consciousness could grow out of its processes, and (2) typical ethical statements about how humans ought to relate to the (usually mammalian and mostly primate) products of the evolutionary process. Beneficent or not, the world, or some corner of it, constitutes all a human or any living creature has. Humans can choose to value the world for that reason alone: it is where we are and what we are, whether or not we love it and whether or not it loves us. But that is no reason for averting one’s gaze from its horrors. The British science writer Marek Kohn has written, “Hamilton’s arguments take their examples from nature and seethe with tiny monsters.”93 American literature has often preferred larger monsters, such as Herman Melville’s white whale. In a well-known poem, Melville pointedly portrays a Maldive shark, the “phlegmatical one,” whose “Gorgonian head,” holds a “port of serrated teeth” that Melville likens to the “jaws of the Fates.” Hiding this head, “The sleek little pilot-fish, azure and slim / How alert in attendance be.” Melville’s pilot fish becomes an image of beauty hiding the voracious shark, “the dotard lethargic and dull / Pale ravener of horrible meat.”94 In a parable where the silent azure pilot fish hides carnage, and beauty hides violence, his poem reveals this veneer of beauty as only a predatory illusion. There is surely something satirical, something grotesque, about Melville’s Maldive shark. So too are Williams’s narratives

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sometimes satirical and grotesque parables. But while Melville focuses on a striking and frightening individual, Williams draws multiple images of unfortunate behavior, images that reduce behavior to self-interest and greed.

Seal Ethics and Mob Behavior From the beginning, Williams is dismayed by public perceptions that imagine and expect, against scientific sense, wildlife to behave beneficently. He dramatizes this by describing a film of elephant seals: Amid the crowded but thriving family groups there was an occasional isolated pup, whose mother had deserted or been killed. These motherless young were manifestly starving and in acute distress. The human audience reacted with horror to the way these unfortunates were rejected by the hundreds of possible foster mothers all around them. It should have been abundantly clear to everyone present that the seals were designed to reproduce themselves, not their species. I know of no evidence that the adaptive organization of subhuman mammalian reproduction ever normally transcends the interactions between a pair of parents and their offspring.95

People who live with animal pets in their homes, whether under human domestication or their own self-domestication, know that the claim with which Williams opens this anecdote—that “females of all species suckle their own young only”—is not always true. Some people may find the final scene where Rose of Sharon nurses the sick old man in John Steinbeck’s The Grapes of Wrath “unnatural.” But Ma’s moral is interesting: “Use’ta be the fambly was fust. It ain’t so now. It’s anybody. Worse off we get, the more we got to do.” But Williams’s point is that, generally speaking, society is an artifice and must be distinguished from mob behavior. For anyone who has been mobbed by terns must acknowledge “that mob, rather than team or task force” describes their behavior. Though each individual is undoubtedly bolder in the attack when it is accompanied by other terns, there is “no organized strategy of attack and no division of labor,” so that the dynamics of a mob is “not a factor for which biotic adaptation need be postulated.”96 It is certainly more comfortable to imagine that we are “good natured,” as Frans de Waal puts it, or that nature is good. But we would not wish to live in what the novelist Robert Stone calls “the social

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dynamics of a coral reef.”97 We are good natured to those beings closest to us because we imagine we recognize ourselves in them. But it is unclear to Williams that the processes of evolution can be deduced from the sameness or difference of selected “higher” species. One does not have to belabor the concept of selfish genes to recognize selfishness and its attendant violence, or to engage with it. As Williams grows older, he often grows impatient. As he writes when he is 62, morality is accidentally produced by a stupid biological process “normally opposed to the expression of such a capability.”98 Further, human morality is confused by self-deceptions about reproduction and death rooted in ideologies and theories of life. From the beginning, he explores death as it attends reproduction. Like Robert Trivers, who also writes extensively about self-deception, he asks what can be fixed, and what cannot be fixed. Birds don’t go to old-age homes. How do they die? People must decide how far their loyalties extend. Williams allies himself with certain literary figures, such as Tennyson and especially T. H. Huxley, but others as well. He is often curmudgeonly and confrontational, and sometimes he himself seems cruel and ruthless, as when he responds in an interview, I’m probably being unfair, but I would say that Lynn Margulis is very much afflicted with a kind of ‘God-is-good’ syndrome, in that she wants to look out there at nature and see something benign and benevolent and ultimately wholesome and worth having. Whereas I look out there with Tennyson and see things red in tooth and claw. In other words, it’s a bloody mess out there.

He thinks her desire to “see cooperation and things being nice to each other” leads to the idea of Gaia, which is only a kind of goddess worship. She sees what she wants to see. Williams plays with the satirical and ironic when he speaks about gods and belief in this interview. “I think ‘God is evil,’ and I look out there at His creations and see nothing but evil. Time will tell, and will show that my approach is more fruitful in generating predictions about discoveries we’re going to make.”99 Marek Kohn wonders, “What is it about the form of the shark, the crocodile or the monkey-puzzle tree that has continued to pass natural selection’s scrutiny when so many other forms have come and gone?”100 He reports that he asked Williams about this. “‘That is a very interesting question,’ Williams declared, but elaborated no further.” Kohn sees this as “Williams’s inclination to reserve his insights for his peers.” I believe it is

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from another cause. For someone with such strong opinions, Williams is often taciturn, “disposed to be silent”—from the Latin word tacitus, “silent” or “that is passed over in silence, done without words, assumed, silent.” And who shouldn’t keep silent when such questions are asked? But there also comes a time when one has to shout fire in a burning theater. Darwin wonders “what a book a devil’s chaplain might write on the clumsy, wasteful, blundering, low, and horridly cruel works of nature.”101 In the end, Williams turns the tables on ethics in many ways. I’ll choose six, though there is the promise of many more: 1. Evolution works one death at a time. Adaptation is not only about survival, but is also about extinction. 2. This is not to say that it is appropriate for humans to wipe out the products of natural selection and evolutionary change. 3. But it is certainly to say that the currency of evolutionary change is death, whether human collusion exists or not. 4. Most obviously, nature is itself unethical and not a model for human ethics. 5. Human guilt about interfering with natural process is thus diminished in many ways. 6. Human responsibility and human guilt must depend upon the human judgments of human acts, and those judgments cannot be based upon the perceived a priori goodness or badness of natural objects or the processes that create them.

What is a Naturalist, or Naturalism? Given his dark view of nature, how then can one speak of Williams as a naturalist? As a twentieth-century philosophical theory, naturalism seems willing to embrace science as the most reliable source of knowledge and the scientific method as the most effective route to knowledge about nature. Otherwise, it would only be faith or religious doctrine. For someone like Williams, Hamilton, Trivers, or E. O. Wilson, that means using reductionist strategies of inquiry and analysis. These biologists also conceive naturalism as a corrective social force, but correction can come from at least two directions. Some who call themselves naturalists, like E.  O. Wilson, also embrace and express more personal rewards from nature, and their writing strategies allow them to express feelings like

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enchantment, a sense of wonder, awe, and joy in their love affair with life. Wilson spells these feeling out in two books, Biophilia and Naturalist.102 Many naturalists still believe that natural processes constitute a model for better human behavior. The argument of The Closing Circle (1971) is emblematic of this model. There, the biologist Barry Commoner famously announces four laws of ecology as a kind of credo for naturalists. The laws are (1) “Everything Is Connected to Everything Else,” (2) “Everything Must Go Somewhere,” so there is no waste in Nature, (3) “Nature Knows Best,” and (4) “There is No Such thing as a Free Lunch.”103 These are not laws Williams can respect, at least not the first three. He considers them the product of mysticism. Further, one does not have to love nature to be interested enough to study it. One can be a naturalist without accepting the natural process as an ethical norm. All naturalists are interested in natural history. Amateur naturalists have been taken by some as proxies for more professional scientists and derogated by others as nature lovers and haphazard collectors. According to the latter view, naturalists have archaic interests; they continue to be engaged in assembling evidence for a great chain of being; they fill cabinets and books with their personal discoveries and narratives. Amateur naturalists join social clubs and associations, attend meetings, and encourage children to go out in the woods to observe the natural world. Literary naturalists record their experience in a notebook, keep a journal, join conservation organizations, take naturalist courses, and preserve natural parks. Nevertheless it is not necessary that one who studies nature must love it, or be a nature lover, in order to be a naturalist. There is commitment, but to what? This becomes quite complex with sociobiology. Frans Roes sheds light on this with a set of interviews with sociobiologists, many of them originally published in the Human Ethology Bulletin and now posted on his website, “Speak, Darwinists!”104 The interviews with Hamilton and Williams of 1997 and 1998 reveal how they think of naturalism as a demythologizing force. For them, evolutionary science reveals many social, political, and religious ideas about human life—including those about human nature—as illusory. Hamilton has written that evolutionary ideas “turn out to have, or are perceived to have, the unfortunate property of being solvents of a vital societal glue.”105 As he explains in his interview with Roes, “The glue that I am thinking of is various myths that tend to hold societies together.” He thinks that these myths “helped people to be more cooperative than they would be if proceeding with a full understanding of evolutionary theory.”

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He thinks the threat of evolutionary theory is real, though dimly perceived by religious people who exaggerate the danger. As he concludes, “I think they exaggerate the danger but they don’t exaggerate a nothing. There is a danger of that kind.”106 This is the danger of sociobiology—that it offers to ground sociology in biology and then undermines the cultural forces that hold societies together. Williams also has interesting things to say about sociobiology and religion in his interview. When prodded by Roes, he admits to admiring “the explicit recognition in Buddhism that things are not naturally good. There is a lot of pain and suffering in the world, and that is because that’s the way the world is.” He finds this view opposed to some Christian and Jewish traditions “in which everything is for the best to some extent, because it is God’s will that things be this way.” For instance, he finds the message of blind obedience in the Book of Job repellent and calls the book “a really sick piece of literature in that respect.”107 The British philosopher Galen Strawson agrees when he insists that we know with certainty that the Christian God, standardly defined as a being who is “omniscient, omnipotent, and wholly benevolent,” does not exist. “The proof lies in the world, which is full of extraordinary suffering.” What is worse, “a genuine belief in such a God, however rare, is profoundly immoral: it shows contempt for the reality of human suffering, or indeed any intense suffering.”108 In 1998, when asked if he is a sociobiologist, Williams says he is willing to be so labeled, but “one of the problems with this sort of labelling is that the term is identified with one individual—E. O. Wilson. And although I like him immensely and certainly admire him, I don’t want to be labelled as a disciple of his.” I would say that Williams defers the label in part, but not entirely, because his ethical allegiances, unlike E. O. Wilson’s, are not grounded in a normative view of nature. As I understand, George C. Williams has spent his entire career studying natural processes that he believes are evil. His own values have always been allied primarily to those who believe that human ethics, pursued within a cultural context as a cultural construct, must counteract that evil. It may be valuable to ask about the biological origin of human behavior, but seeking roots for human ethics in biology is poorly advised.

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Notes 1. George C.  Williams, “A Sociobiological Expansion of Evolution and Ethics,” in T.  H. Huxley’s Evolution and Ethics, with New Essays on its Victorian and Sociobiological Context, edited by James G.  Paradis and George C. Williams (Princeton, NJ: Princeton University Press, 1989), 179–214, 212. 2. George C.  Williams, “Huxley’s Evolution and Ethics in Sociobiological Perspective,” Zygon 23, no. 4 (December 1988): 383–438, 438. 3. Williams, “Sociobiological Expansion,” 437. 4. Williams’s 1988 essay in Zygon is an abridged version of “A Sociobiological Expansion of Evolution and Ethics” in his and Paradis’s 1989 edition of T. H. Huxley’s Evolution and Ethics (see full citations above in notes 1 and 2). Williams’s essay “Mother Nature is a Wicked Old Witch” appeared in Evolutionary Ethics, edited by Matthew H. Nitecki and Doris V. Nitecki (Albany: SUNY Press, 1993), 217–31. It is an abridged version of Williams’s article, “Gaia, Nature Worship and Biocentric Fallacies,” the Quarterly Review of Biology 67, no. 4 (1992): 479–86. The essay in Zygon is also reprinted in Issues in Evolutionary Ethics, edited by Paul Thompson (Albany: SUNY Press, 1995), 317–49. 5. Williams uses this opening sentence in “Sociobiological Expansion,” “Gaia,” and “Mother Nature.” 6. He cites, for instance, Stephen Jay Gould’s “Darwinism and the Expansion of Evolutionary Theory,” Science 216, no. 4544 (1982): 380–87. 7. George C. Williams, Adaptation and Natural Selection (Princeton, NJ: Princeton University Press, 1966), 255. 8. T. H. Huxley’s Evolution and Ethics, with New Essays on its Victorian and Sociobiological Context, edited by James G. Paradis and George C. Williams (Princeton, NJ: Princeton University Press, 1989). All references to Huxley’s text will refer to the pagination of the facsimile of Evolution and Ethics in this edition. 9. Williams, Adaptation, 6–7; Williams, “A Defense of Reductionism in Evolutionary Biology,” in Oxford Surveys in Evolutionary Biology, vol. 2, edited by Richard Dawkins and Mark Ridley, 1–27 (Oxford: Oxford University Press, 1985); Williams, “Gaia,” 485–6. 10. See Williams’s review of Robert J. Richards, Darwin and the Emergence of Evolutionary Theories of Mind and Behavior (Chicago: University of Chicago, 1987) in Journal of Evolutionary Biology 2 (1989): 385–87. Indeed, the chapter with which Williams disagrees, included as Appendix 2 in this book, was previously published as Robert Richards, “A Defense of Evolutionary Ethics,” Biology and Philosophy 1, no. 3 (1986): 265–93.

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11. A good anthology focused on evolutionary ethics is Michael Ruse, ed., Philosophy After Darwin: Classic and Contemporary Readings (Princeton, NJ: Princeton University Press, 2009). 12. E. O. Wilson, Sociobiology (Cambridge, MA: Belknap/Harvard University Press, 1975), 27. 13. Kitcher’s essay appeared originally in Evolution und Ethik, edited by K. Bayertz (Stuttgart: Reclam, 1993). It has been reprinted variously. I cite Philip Kitcher, “Four Ways of ‘Biologicizing’ Ethics,” in In Mendel’s Mirror: Philosophical Reflections on Biology (New York: Oxford University Press, 2003), 321–32, 330. 14. Michael Ridge, “Moral Non-Naturalism,” The Stanford Encyclopedia of Philosophy (Spring 2010 Edition), edited by Edward N.  Zalta, http:// plato.stanford.edu/archives/spr2010/entries/moral-­non-­naturalism/. 15. George E.  Moore, Principia Ethica (New York: Cambridge University Press, 1903), 46. Quoted in Richards, Darwin, 323. 16. Kitcher, “Four Ways,” 330. 17. Stephen Jay Gould and Richard C.  Lewontin, “The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme,” Proceedings of the Royal Society of London, Series B, 205, no. 1161 (1979): 581–98, 584–85. 18. Gregg Mitman, The State of Nature: Ecology, Community, and American Social Thought, 1900–1950 (Chicago: University of Chicago Press, 1992), 7. This is an era well after the one discussed by Peter J. Bowler in The Eclipse of Darwinism: Anti-Darwinian Evolution Theories in the Decades around 1900 (Baltimore: Johns Hopkins University Press, 1983). 19. Mitman, State of Nature, 212. 20. David Lack, “Of Birds and Men” New Scientist (January 16, 1969): 21–22, 22. 21. Gregg Mitman, “Where Ecology, Nature, and Politics Meet: Reclaiming The Death of Nature,” Isis 97 (2006): 496–504, 500. See also Kevin Dann and Gregg Mitman, “Exploring the Borders of Environmental History and the History of Ecology,” Journal of the History of Biology 30 (1997): 291–302. 22. See B. Andrew Lustig, Baruch A. Brody, and Gerald P. McKenny, Altering Nature: Concepts of ‘Nature’ and ‘the Natural’ in Biotechnology Debates, vol. 1 (Springer, 2008). 23. George C.  Williams and Doris C.  Williams, “Natural Selection of Individually Harmful Social Adaptations among Sibs with Special Reference to Social Insects,” Evolution 11, no. 1 (March 1957): 32–39. George C. Williams, “Measurement of Consociation among Fishes and Comments on the Evolution of Schooling,” Michigan State University Museum Publications, Biology Series 2 (1964): 351–83.

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24. Carl Zimmer, “Stretching the Limits of Evolutionary Biology,” Science 304, no. 5675 (2004): 1235–36. 25. Williams, Adaptation, 83. 26. Margaret Drabble, The Sea Lady: A Late Romance (Orlando: Harcourt, 2006), 151. 27. Harmon Holcomb and Jason Byron, “Sociobiology,” The Stanford Encyclopedia of Philosophy (Fall 2010 Edition), edited by Edward N. Zalta, http://plato.stanford.edu/archives/fall2010/entries/sociobiology/. 28. E. O. Wilson, On Human Nature (Cambridge, MA: Harvard University Press, 1978), x. 29. Richards, Darwin, 540; Williams, Adaptation, 94. 30. Williams, “Review,” Journal of Evolutionary Biology 2 (1987): 385–87. 31. Richards, Darwin, 540 n. 108. 32. Williams, “Review,” 386. 33. Huxley, Evolution and Ethics, 79–80. 34. Mary Midgley, “Sociobiology,” Journal of Medical Ethics 10 (1984): 158–60, 159. 35. George C.  Williams, Review of Arthur L.  Caplan, ed., The Sociobiology Debate (New York: Harper & Row, 1978) in the Quarterly Review of Biology 54, no. 2 (1979): 212. 36. Huxley, Evolution and Ethics, 16. 37. Paradis, Evolution and Ethics, 34. 38. Huxley, Evolution and Ethics, 19, 81, 10, 13, 19, 33. 39. Huxley, Evolution and Ethics, 43, 85. See Richard Hofstadter, Social Darwinism in American Thought, rev. ed. (Boston: Beacon Press, 1955): 95–6, 139. Also see Richards, Darwin, 540. 40. James Paradis, T. H. Huxley: Man’s Place in Nature (Lincoln: University of Nebraska Press, 1978), 86. 41. Paul Lawrence Farber, The Temptations of Evolutionary Ethics (Berkeley: University of California Press, 1998), 62, 65, 66, 69. 42. Williams, “Sociobiological Expansion,” 214, 180, 181. 43. Williams, “Sociobiological Expansion,” 184. 44. Williams, “Sociobiological Expansion,” 191–92, 193. 45. See Peter Singer, The Expanding Circle: Ethics and Sociobiology (New York: Farrar, Straus and Giroux, 1981). 46. See Donald T.  Campbell, “Social Morality Norms,” in Morality as a Biological Phenomenon: The Presuppositions of Sociobiological Research, edited by Gunther Siegmund Stent (Berkeley: University of California Press, 1981), 70. See also Donald T.  Campbell, “Comments on the Sociobiology of Ethics and Moralizing,” Behavioral Science 24, no. 1 (Jan 1979): 37–45. Campbell refers to an article that E. O. Wilson published in the New York Times, “Human Decency is Animal” (NYT Magazine,

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October 12, 1975): 38–50; the article is drawn directly from Sociobiology: The New Synthesis (1975). 47. Williams, “Sociobiological Expansion,” 191. 48. Williams, Adaptation, 256. 49. Williams, “Sociobiological Expansion,” 197, 212, 202. 50. Williams, “Sociobiological Expansion,” 207–8. 51. Williams, “Sociobiological Expansion,” 209; Huxley, Evolution and Ethics, 59. 52. Williams, “Sociobiological Expansion,” 212, 213. 53. Williams, “Sociobiological Expansion,” 214. 54. Williams, “Sociobiological Expansion,” 214; Williams, “Huxley’s Evolution and Ethics,” 403. 55. Richard Dawkins, A Devil’s Chaplain: Reflections on Hope, Lies, Science, and Love (Boston: Houghton Mifflin, 2004), 10. 56. William F.  Harms, “Adaptation and Moral Realism,” Biology and Philosophy 15, no. 5 (2000): 699–712. 57. Williams, “Sociobiological Expansion,” 212; Huxley, Evolution and Ethics, 82. 58. Williams, “Sociobiological Expansion,” 209, 210. 59. Henry David Thoreau, Walden and Resistance to Government: Authoritative Texts, Thoreau’s Journal, Reviews and Essays in Criticism, 2nd Edition, edited by William Rossi (New York: Norton, 1992): 147–48. 60. Comments on Williams’s essay follow his text in Zygon 23, no. 4 (December 1988). They include: Sarah Blaffer Hrdy, “Comments on George Williams’s Essay on Morality and Nature” (409–11); Michael Ruse, “Response to Williams: Selfishness is not Enough” (413–16); Ralph Wendell Burhoe, “On ‘Huxley’s Evolution and Ethics in Sociobiological Perspective’ by George C.  Williams” (417–30); John B.  Cobb, Jr., “Befriending an Amoral Nature” (431–36), and George C. Williams, “Reply to Comments on ‘Huxley’s Evolution and Ethics in Sociobiological Perspective’” (437–38). 61. Hrdy, “Comments,” 410. 62. Williams, “Huxley’s Evolution,” 385. 63. Ruse, “Response,” 415. 64. Williams, “Reply,” 438. 65. Williams, “Sociobiological Expansion,” 181–2. 66. Ruse, “Response,” 416. For Ruse’s views, see Michael Ruse, Evolutionary Naturalism: Selected Essays (London: Routledge, 1995). 67. Michael Ruse, “From Belief to Unbelief—And Halfway Back,” Zygon 29, no. 1 (1994): 25–35. George C.  Williams, “Ruminations on Ruse and Religion,” Zygon 29, no. 1 (1994): 37–43. 68. Williams, “Reply,” 437.

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69. George C.  Williams, “Gaia, Nature Worship and Biocentric Fallacies,” the Quarterly Review of Biology 67 (1992): 479–86. George C. Williams, “Mother Nature is a Wicked Old Witch,” in Evolutionary Ethics, edited by Matthew H. Nitecki and Doris V. Nitecki, (Albany: State University of New York, 1993), 217–31. 70. James W.  Kirchner, “The Gaia Hypothesis: Fact, Theory, and Wishful Thinking,” Climatic Change 52, no. 4 (2002): 391–408. See also Kirchner, “The Gaia Hypothesis: Conjectures and Refutations,” Climatic Change, 58 (2003): 21–45. 71. Northrop Frye, Anatomy of Criticism: Four Essays (Princeton, NJ: Princeton University Press, 1957), 223, 224, 231, 233. 72. Williams, “Gaia,” 479. 73. Richard A.  Kerr, “No Longer Willful, Gaia Becomes Respectable,” Science 22 (April 1988): 393–95; Charles Mann, “Lynn Margulis: Science’s Unruly Earth Mother,” Science (April 19, 1991): 378–81. 74. Williams, “Gaia,” 480, 481. 75. Williams, “Gaia,” 484. 76. Richard C. Lewontin, The Triple Helix: Gene, Organism, and Environment (Cambridge, MA: Harvard University Press, 2001), 109. 77. Williams, Adaptation, 247, 255. 78. Williams, “Gaia,” 481, 482, 484. 79. Williams, “Gaia,” 482, 483. 80. Williams, “Gaia,” 484, 485. 81. Jessica C.  Flack and Frans B.  M. de Waal, “‘Any Animal Whatever’: Darwinian Building Blocks of Morality in Monkeys and Apes,” Journal of Consciousness Studies 7, no. 1–2 (2000): 1–29, 4. 82. Williams, “Gaia,” 486. 83. Williams, “Mother Nature,” 229–30. 84. Sarah Blaffer Hrdy, Mother Nature: A History of Mothers, Infants, and Natural Selection (New York: Pantheon Books, 1999): xviii–xix. 85. Richard D.  Alexander, “The Challenge of Human Social Behavior,” Evolutionary Psychology 4 (2006): 1–32, 5. www.epjournal.net/wp-­ content/uploads/ep04132.pdf. 86. George C. Williams and Randolph M. Nesse, Why We Get Sick: The New Science of Darwinian Medicine (New York: Times Books, 1995), 10–11. 87. Richard C.  Lewontin, Biology as Ideology: The Doctrine of DNA (New York: Harper Perennial, 1993). 88. Alexander, “Challenge,” 5. See also M.  Flinn and R.  D. Alexander, “Culture Theory: The Developing Synthesis from Biology,” Human Ecology 10, no. 3 (1982): 383–400. 89. Frans de Waal, Good Natured: The Origins of Right and Wrong in Humans and Other Animals (Cambridge, MA: Harvard University Press, 2006).

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For de Waal’s condemnation of the “veneer theory,” see especially Frans B. M. de Waal, “Morality and the Social Instincts: Continuity with the Other Primates,” The Tanner Lectures on Human Values Delivered at Princeton University (November 19–20, 2003). 90. Charles Darwin, The Descent of Man (1871, reprint, Princeton, NJ: Princeton University Press, 1981), 88–89. 91. Paul Shepard, The Others: How Animals Made Us Human. Washington, DC: Island Press/Shearwater Books, 1996. 92. Philip Kitcher, “Ethics and Evolution” in Primates and Philosophers: How Morality Evolved, edited by Frans B. M. de Waal (Princeton, NJ: Princeton University Press, 2006), 120–39, 139. 93. Marek Kohn, A Reason for Everything: Natural Selection and the English Imagination (London: Faber, 2004), 325. 94. Herman Melville, “The Maldive Shark,” in John Marr and Other Sailors (1888). 95. Williams, Adaptation, 188–9. 96. Williams, Adaptation, 189. 97. Robert Stone, “Absence of Mercy,” in Bear and His Daughter (Boston: Houghton Mifflin, 1997), 28. 98. Williams, “Reply,” 438. 99. Williams, interview in John Brockman’s The Third Culture (New York: Touchstone, 1995), 141. 100. Marek Kohn, review of Survivors by Richard Fortey, The Independent (September 9, 2011). 101. Charles Darwin, Letter to J. D. Hooker, July 13, 1856. 102. Edward O.  Wilson, Biophilia (Cambridge, MA: Harvard University Press, 1984). Wilson, Naturalist (Washington, DC: Island Press, 2006). 103. Barry Commoner, The Closing Circle: Nature, Man, and Technology (New York : Knopf, 1971), 16–23. 104. Frans Roes’s Speak, Darwinists! site: http://www.froes.dds.nl/. 105. William Hamilton, Narrow Roads of Gene Land: The Collected Papers of W. D. Hamilton, vol. 1 (New York: W. H. Freeman, 1996), 15. Hamilton almost certainly rephrases Daniel Dennett’s overarching metaphor of Darwinism as a “universal acid” that “eats through just about every traditional concept, and leaves in its wake a revolutionized world-view, with most of the old landmarks still recognizable, but transformed in fundamental ways.” Daniel C. Dennett, Darwin’s Dangerous Idea: Evolution and the Meaning of Life (New York: Simon & Schuster, 1995), 63. 106. An interview with William D.  Hamilton, Human Ethology Bulletin 12, no. 4 (1997): 3–7, 6. 107. Frans Roes, “A Conversation with George C. Williams,” Natural History 107 (May 1998): 10–13.

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108. Galen Strawson, in “Letters: What can be Proved about God,” New York Review of Books 59, no. 19 (December 6, 2012): 65. See also the original: Galen Strawson, “Religion is a Sin,” a review of Mark Johnston, Saving God: Religion after Idolatry, and Surviving Death, London Review of Books 33, no. 11 (2011): 26–28. http://www.lrb.co.uk/v33/n11/ galen-­strawson/religion-­is-­a-­sin.

CHAPTER 8

History, Contingency, Constraint: A Book of Nature

Natural Selection: Domains, Levels, and Challenges (1992) is simultaneously Williams’s most technical and most personal book. For my purposes, the intricate technical problems it explores are less relevant than its literary aspects. The text consists of ten chapters, and as the title indicates, focuses first on domains. Williams defines two mutually exclusive domains of selection, “one that deals with material entities and another that deals with information and might be termed the codical domain.”1 Richard Dawkins credits Williams with coining the term codical domain.2 The material domain comprises atoms in the DNA of the gene. The codical domain comprises information coded in the gene. This distinguishes message from medium. As for the message, on the first page of Natural Selection, Williams echoes and expands an argument he presented in “Defense of Reductionism” (1985). “Successful biological research in this century has three doctrinal bases: mechanism (as opposed to vitalism), natural selection (trial and error as opposed to rational plan), and historicity.” When he includes historicity, he proceeds further here than in 1985, perhaps because he attempts to reconcile his views of multiple levels of selection with those of other biologists, especially Stephen Jay Gould and Richard Lewontin. Not only does he cite Gould’s Wonderful Life (1989), but he also emphasizes one of Gould’s favorite conceptions, contingency. He cites Gould, Lewontin, and Niles Eldredge scores of times. When Williams © The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 M. P. Cohen, George C. Williams and Evolutionary Literacy, Literatures, Cultures, and the Environment, https://doi.org/10.1007/978-3-031-11650-6_8

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confronts “the role of historical contingency in determining properties of the Earth’s biota,” he concedes at the onset that “the formation of the Earth was a unique event, and unique events have been altering evolutionary processes ever since, always keeping futures unpredictable from presents.”3 These contingent processes have determined the properties of the earth’s biota and become a key to understanding certain features of organisms. Indeed, “many features of living organisms are functionally arbitrary or even maladaptive.” In addition to historicity and contingency, he speaks of “related evolutionary constraints (phylogenetic, developmental, genetic) on the action of natural selection.”4 As a consequence, Natural Selection is thematically focused on historicity, contingency, and constraint. Stylistically, it reveals more explicitly than any of Williams’s other books the contingent and constrained history of his own thinking. He illustrates the constraints that impinge upon his writing from a tradition of literature dealing with natural selection—not just from Darwinian texts, but also from more recently published work. The resulting style reveals a kind of restraint that one might almost find akin to the modernist notion of extinction of personality, as in T. S. Eliot’s famous essay, “Tradition and the Individual Talent.” Williams edits a little anthology for the end of his book that includes excerpts from William Paley on the human eye and Galen on the human hand. He pays his debt to historical texts. These, he writes, are “worth close attention by all biologists. Both illustrate how adaptation can be explored by seeking conformities to a priori design specifications.” These texts are normally taken to be supportive of Williams’s own interest in design. Certainly he includes them to illustrate methods he admires. He recommends but does not have room to include Hermann Joseph Muller, whose Evidence of the Precision of Genetic Adaptation (1948) presents a “tightly reasoned argument” that has “a surprisingly modern ring after 44  years.” As Williams says of Muller, “his discussion, like so many of Fisher’s, Haldane’s, and Wright’s between 1930 and 1960, shows that theories are often understood and used effectively long before they are formalized.”5 Under the circumstances, it is hard not to see these texts also as codices he inherits. They reveal the genetic shape of Williams’s thought. Any book of only 150 pages entitled Natural Selection entails maximum compression and perhaps drives the plain style into a corner. The narrator is intimate and candid about his education and the development of his views, leading some readers to notice that the book almost seems to

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consist of soliloquys, vignettes, and short essays. Williams speaks a great deal about ideas that conform with or are contrary to his own as he tries to distinguish his views from those of others, often by gradation. His aim is consensus. He treats all reasonable views with respect, but is sharp, as always, about intellectual laziness, and his key aim is to eradicate what he calls “obfuscation.” At the same time, this book is more technical and wide-ranging than any other by Williams. It alludes to Darwin more than any other. It engages with philosophers like Elliot Sober, with questions about success and succession. The narrator reveals himself by his ambivalent tone, by statements that include double doubts like the following: “I have no reason to doubt that various new adaptations evolved in certain lineages have facilitated their subsequent evolutionary proliferation. I seriously doubt that there is any way of establishing this proposition, because there is no way of knowing how any such group would have fared without its supposedly special advantage.”6 As a contemporary review in the journal Evolution argues, “the centerpiece of the book is a resurrection and nearly complete overhaul of ideas he first presented in Adaptation and Natural Selection (1966).” More accurately, Natural Selection is part of a continuing reassessment written during sabbaticals in 1981–82 and 1988–89. The book covers a wide gamut of issues, and as the two reviewers also note, “seems more preoccupied with identifying the symptoms of a whole host of difficult problems in evolution, and less concerned with diagnosing the causes and offering remedies.”7 Another review, by the ornithologist Joel Cracraft in Bioscience, is quite negative, saying that, “like others before him, Williams wants a unified view of selection,” noting “surprisingly little critical analysis of the literature on levels of selection.”8 From Cracraft’s perspective—which corresponds to those of many other readers—the overall structure of the book is also puzzling. He believes that Williams “paints a much broader canvas than the earlier book, covering ideas of selection at different hierarchical levels, macroevolution, optimization, phylogenetic and developmental constraint, diversity, species concepts, morphological stasis, and numerous other topics.”9 I will argue that Williams may be mounting a defense and broadening the applicability of his previous methodologies. Despite the proliferation of relatively negative reviews, and whether Williams is right or wrong in his scientific theorizing, I think that something more operates in this book because it is an attempt to justify writing as an avenue to understanding. This emphasis may explain his inclusion, at the bottom

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end of his frame, of a set of literary models for discussing genetic texts. It is also my hypothesis, judging from reviews of his book by biologists, that this part of his design or intention may have been entirely lost on his scientific readership. Each domain of natural selection can be understood to have its own units. A codex is the basic unit in the codical domain. As usual for Williams, discussion starts at the level of the gene, but it does not end there. Williams believes that the language for higher-level processes is inadequate. The phrase “species selection,” for instance, has obfuscated discussions of higher levels of selection because it implies “something special, for natural selection, about the species level of classification”; he is no happier with the expression “taxon selection.” As he decides, “If I am right that all levels of genealogical inclusiveness from local gene pools to classes and phyla may be subject to selection, the term clade selection used by Stearns (1986) is a great improvement.”10 A clade is a group of organisms believed to have evolved from a common ancestor. As Cracraft says, “Williams uses clade— as in his clade selection—in a way entirely at variance from the common use of that term.” Cracraft simply does not think “a uniform view of selection can be applied across the different hierarchical levels.”11 For instance, a recent review argues that “clade selection is unpopular with philosophers who otherwise accept multilevel selection theory. Clades cannot reproduce, and reproduction is widely thought necessary for evolution by natural selection, especially of complex adaptations.”12 There is no reason to believe that Williams is right on this subject.13 Nor is Gould, probably, when he argues for species selection. But the argument of Natural Selection is interesting in its own right because it reveals how Williams attempts to expand gene selectionism into macroevolution. Natural selection, he argues, selects from codices, and there is literary resonance in the terms code and codex. When Williams concludes his book by articulating a set of challenges, “challenges” has multiple meanings, implying not only challenges for biological workers, but also challenges to the thinking of biologists.

The Domain of the Codex In Natural Selection, Williams develops an elaborately linked set of analogies closely related to the traditional conceit of the “Book of Nature.” Organisms, Williams believes, can be conceived as documents, and this approach is useful for investigating unique evolutionary histories. “The

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doctrine of historicity holds that the properties of the organic world, from the total biota to minute parts of animals and plants, are the results of unique historical contingencies, some of which are documented in features of the organisms themselves.”14 This analogy presents the whole biotic kingdom as a library. A library, Williams argues, is not the material on shelves and in drawers, but the information in codices. A catastrophic act of arson in the seventh century was a physical event at exactly the same time as the destruction of information in the library at Alexandria. Information can exist only as a material pattern, but the same information can be recorded by a variety of patterns in many different kinds of material. A message is always coded in some medium, but the medium is really not the message.15

Yet this argument, as he knows, will not quite work when one asks what exactly was or is endangered. To speak of a codical domain constitutes both an interesting innovation and an allusion to traditional and historical modes for representing information. In my own dictionary, a codex is a book, usually one produced before the advent of the printing press; codex is a name for a manuscript book. The term usually refers to ancient or medieval books—to scriptures, classics, ancient annals. Etymologically, codex refers to a systematic compilation of laws, a book of laws, the word literally indicating a tree trunk apparently sometimes used for immobilizing prisoners. A codex is written by hand on a wooden tablet. For Williams, “a unit of selection in the codical domain would be a codex (codices is the only plural in my dictionary).”16 Williams implies an analogy here. Partaking partially of the immortality of the gene, the codex of Williams is ancient. Our recent use of the term code to indicate a cipher or secret code reminds one that any codex calls out a desire to decipher the historical text. As Williams says, “In discussing the codical domain we use such terms as bits, redundancy, fidelity, and meaning.” In his own sly and recursive way, Williams uses the following example of a code as a set of laws within a text: “Machiavelli’s The Prince is a codex that might be described as a set of directions for taking unfair advantage of subordinates. I might reasonably point to a copy and term it an immoral book. If so, I would be generally understood to be talking about the message, not the medium.”17

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So, in this sense, when one faces the codex as literary structure, one is within a literary work by Williams. Stephen Stearns is a source for many of the ideas expressed in Natural Selection and, as one of Williams’s closest and most sympathetic readers, thinks of this text as “a collection of essays in which he reacted to the huge literature that had appeared since his earlier books and advanced some new ways of thinking about enduring issues.”18 I would say that the analogy of the codex gives Natural Selection more coherence than a mere collection. Unless one wants to assume a piecemeal association of the materials or chapters, there is reason to look at the book’s frame. His first chapter is perhaps unique in a technical monograph because Williams entitles it “A Philosophical Position” and elucidates his interest in the historicity and constraints that impinge upon natural selection. This is followed by a second chapter that explores how a biologist might read and write about these codices as if they were historical texts, past and present—or, one might say, engage in writing an elucidation of codes and texts. By Chap. 6 he expands into macroevolutionary theory, but he returns to codes and texts in an appendix that explores some of the literary tradition that has attempted to explain organic design through prose. His book is an experiment whose structure is meant to mirror the way flaws and omissions of design are produced by natural selection and uncover the way these flaws enter the perspectives of its observers.

Macroevolution, Dendrograms, and the Language of Clades Natural Selection announces Williams’s desire to enter discussions about macroevolution, to interest himself in change that occurs at what are sometimes conceived as “higher levels.” As he says, these discussions have failed to clarify whether gene pools or living organisms are at issue. “My use of gene and clade as strictly codical concepts demands other terms for their physical manifestations.”19 Macroevolution is evolution “on a grand scale,” and it is imagined to operate at or above the level of species, in contrast with microevolution, which focuses on evolutionary changes within a species or population. Those who study macroevolution are especially interested in the investigation of extinctions. In this sense, Natural Selection aims critique toward current macroevolutionary theory, perhaps in the same way that Adaptation and Natural

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Selection aimed at microevolutionary theory. It is fair to say that Williams finds much current macroevolutionary discourse to be crude and inaccurate. To extend his gene’s eye view to macroevolution’s levels of selection calls for considerable manipulation of his theoretical concerns and the language in which they are embodied, or as this concern is called by one review, “terminological haggling.”20 But the manipulation is more than terminological. Consider the clade. The term’s etymological root comes from the word for branch. In diagrammatic terms, one can think of a clade as a branch of the tree of life. The term clade was first used by Julian Huxley in 1958. To think in terms of clades is to think in terms of trees, since a clade is etymologically a branch and a dendrogram a “tree drawing.” Mark Ridley defines cladism as an evolutionary methodology “rigorously based upon the phylogenetic principle as a means of biological classification” where “a species is a lineage of populations between two phylogenetic branch points (or speciation events).” Cladism “recognizes species by branch points, independently of how much change occurs between them.”21 For Williams, only clade selection operates above gene selection, and he depends for authority upon the work of two scientists he knows well, Stephen C. Stearns and Leigh Van Valen. The book of Natural Selection has trees in it because it must. As Williams explains, a gene pool and all its descendant gene pools constitute a clade, and such entities are normally represented by diagrams. So he illustrates clades with dendrograms because “the history of gene pools in a clade and of genes in a gene pool can both be realistically modeled by dendrograms.” He thinks of a clade as an informational entity. As applied to “a general model of selection in the codical domain,” Williams begins, “information can proliferate and be edited by natural selection only if the selection affects the information at a greater rate than competing processes such as mutation and drift.” He manipulates several dendrograms to illustrate this process of proliferation and editing, because “to model success and failure in genic selection implies a definition of the term gene. A gene is that which reliably survives the process of meiosis intact,” or, as one can also say, “a gene is that which obeys Mendel’s law of independent assortment. Independent means independent of other genes in its transmission through generations” (see Fig. 8.1).22 As Stephen C. Stearns understands Williams’s point, “codical units of selection persist longer than their material carriers and must be able to persist and proliferate faster than they change. If this is the case, their

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Fig. 8.1  These dendrograms illustrate “natural selection of codices,” B being an enlargement of A. Thickness indicates numerical abundance. C and D illustrate a pattern of imagined punctuated equilibria, and lack indication of abundance. This figure appears in George C.  Williams, Natural Selection: Domains, Levels, and Challenges (New York: Oxford University Press, 1992), 14. Reproduced with permission from Oxford University Press

proliferation can be represented by a genealogy or dendrogram, and George claimed that ‘a good test of the susceptibility of an entity to natural selection is whether its history can be modeled successfully by a dendrogram.’”23 Williams also argues that because whole genotypes do not replicate themselves in sexual reproduction—and cannot be modeled by dendrograms—they cannot be units of selection. In a sense, he reminds the reader what reduction to the gene’s eye view requires. Genes may not be functionally independent because genes of a genotype may and do interact with both their genetic and ecological environments in complex ways that determine their fitness. Consequently, “when dendrograms are used to illustrate selection in the codical domain, continuous lines should be used only for information continuously present.”24

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To think clearly with dendrograms, the observer must keep track of both the codical and the material domains, the abstract and the concrete. It is also possible to assign a time scale to these diagrams. A diagram (Fig. 8.2) attempts to demonstrate “relationships among the concepts of gene, genotype, gene pool, and individual soma.” Bodies are concrete objects but other entities exist as information in the codical domain. “The differential success of individual genotypes in contributing genes to a common gene pool, as shown here, is the only legitimate meaning for the term

Fig. 8.2  Dendrograms generated by random branching and character changes. As Williams explains, “A shows a completely random phylogeny while B shows random changes opposed by normalizing clade selection.” This figure appears in George C.  Williams, Natural Selection: Domains, Levels, and Challenges (New York: Oxford University Press, 1992), 133. Reproduced with permission from Oxford University Press

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individual selection.” Kin selection complicates a dendrogram: “If individuals regularly interact with relatives, selection at the level of the gene will depend in part on effects on the relatives’ reproductive success.” Williams suggests this as a way to consider the sociobiology of plants. Think of a nut on a nut tree: it may appear to be part of the tree, but it is an individual. Further, “closer attention shows it to be a composite of several genetically distinct individuals of varying kinship, each of which must be selected to optimize its interactions with each of the others.”25 As an alternative, Williams suggests one might think of chicks in a nest. These situations require some assessment of kin selection or inclusive fitness. Well, kinship itself is not easily perceived, so “selection must make use of indirect evidence, often something obvious, like propinquity at a certain stage of the life cycle.” Propinquity is an interesting term. It implies kinship, or that the participants have some sense of nearness revealed through concrete signs. He believes that nest mates, for instance, “may make use of many kinds of comparisons of close associates with each other and with themselves” to perceive the closeness of their relations with others in the nest, and they “modify their behavior on the basis of this information.” Such “mechanisms of kinship assessment” play a role in what he calls “the behavioral ecology of natural populations.”26 The response to this thinking was not particularly enthusiastic. Biologists asked whether genes are actors, causal agents in evolutionary processes, or are merely keeping track of them and thus have only a bookkeeping role. In 1994, Gould argued that Williams’s codical domain cannot be “an alternative realm of causality,” and called it “just a fancy name for the necessary bookkeeping function of evolutionary calculation.”27 Gould and Okasha believe that, because codices are not material entities “‘out there’ in the Darwinian struggle,” “bookkeeping cannot explain causality.” Indeed, Okasha finds the charge of “confusing bookkeeping with causality” a key objection to the gene’s eye view—one related to the charge that such a view ignores the fact that genes’ effects on phenotypes are dependent on context.28

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Vitalism and Mechanism, Reduction and Holism: “A Philosophical Position” When Williams speaks of his philosophical position, he seems less concerned with establishing whether he is right or wrong about any single argument, but instead wishes to provide a panoramic and inclusive compendium of his perspective. He is at the age when many people retire. At one time he did not expect to have an influential voice, but by 1992 he does, and so the question becomes how he will use it. He also takes pains to show the history of his views by reflecting on his own education. Vitalism, a perspective out of his past, “maintains that the observable machinery has but limited autonomy and is controlled by a purposive entity peculiar to living organisms.” He remembers “Authors of biology textbooks that I read in the 1940s often presented vitalism as an idea with some adherents, but implied that mechanism was nearly universal among practicing professionals.”29 Williams believes that old school textbooks included obsolete ideas out of intellectual laziness. In a common pattern, mid-century textbooks included a summary treatment of evolution where neo-Darwinian processes were given equal space with other, clearly obsolete views. “Thereupon any axiomatic use of the theory was abandoned, and it was merely assumed that natural selection always promoted what was in some way good.” He portrays this myth as dished up like a kind of children’s tale where “predator-vulnerable animals resemble their backgrounds,” “adults unselfishly devote themselves to reproduction so that their species might persist,” and there was a chain of being from amphibians to reptiles to mammals. “Perhaps,” such treatments implied, evolution “even assured the emergence of modern civilization.”30 Among “commendable exceptions to the general intellectual laziness,” he mentions, surprisingly, Allee, Emerson, Park, Park, and Schmidt’s Principles of Animal Ecology (1948).31 These are the very proponents of group selection with whom he took exception in 1966 and again in the 1980s. Only later in his education, as he remembers, did he read Fisher, Haldane, and Lack. He also reminisces that “in those days few people read such mathematical discussions as Haldane’s and Fisher’s. We got our theory in words from Dobzhansky, Mayr, and Simpson, who were deeply concerned with natural selection, but gave little attention to its levels of operation.”32 In other words, Williams portrays himself as a reader of texts—often faulty texts that have taken a lifetime to explore.

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As one who has gotten his theory through words, Williams himself provides theory through words. He does not imagine that he can read “beneath” texts to the motives of the writers. “I know how I feel when I claim ‘Cogito ergo sum,’ and by intuition and analogy I assume that Descartes felt the same way. I doubt that I will ever have any more direct insight into Descartes’ or anyone else’s thinking.” No wonder public discourse and demonstration matter most to him, as they always have. As he had written before, “I have no inclination to deny the mental realm or belittle its philosophical importance. I am inclined merely to delete it from biological explanation, because it is an entirely private phenomenon, and biology must deal with the publicly demonstrable.” Philosophically, he retains his faith in the adaptationist program, claims that it has been in use for centuries, that its use is increasingly common, and that it has become “more explicitly based on natural selection during the last few decades.” Yet he describes its practitioners at a distance, in the third person: “They assume that an organism is already close to some maximum achievable level of adaptation, and on this basis they attempt to predict features not yet known.”33 Williams brings order to his philosophical position by distinguishing “mechanistic biologists,” who imagine an “organism-as-crystal,” and adaptationists, who conceive an “organism-as-artifact.” He believes that “biologists interested mainly in unique evolutionary histories” should adopt an “organism-as-document approach,” taking his textual approach further and further. This approach emerges from the “doctrine of historicity”—namely, that all properties of the organic world, “from the total biota to minute parts of animals and plants,” must be “results of unique historical contingencies.” Contingency and constraint expand his philosophical position and allow him to recognize the functionally arbitrary or even maladaptive features of living organisms. He marshals examples: the seven common vertebrae found in necks of giraffe, human, and mouse constitute a “functionally inexplicable uniformity.” He observes the way in which the digestive and respiratory tracts cross in the throats of all vertebrates, making them capable of choking on food. While it is comprehensible as a historical legacy, “this evolutionary short-sightedness has never been correctable.”34 Williams notes that science fiction texts frequently find evidence of the species Homo sapiens on other planets, as if evolution would always lead to the same organisms universally, but how could this be? Yet the popularity of science fiction is a phenomenon suggesting to him a certain silliness.

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“There is little grasp of natural selection or historicity outside biology. Within biology they are well understood only by those especially interested in evolution.” Historical ignorance dwells inside and outside biology. Although he is quite sure that mechanism, natural selection, and historicity, “the same three principles, and no others, will be found to apply everywhere in the universe where life has arisen,” this ubiquity seems to cause him to pause. Even while he endorses “Dawkins’ (1983) idea of universal Darwinism,” he also writes, “of course any of my three basic principles might be wrong. I suggested as much about mechanism when I discussed the possibility of an immaterial but physically active principle of mentality.”35 This may be tongue in cheek. We know he does not believe in immaterial biological principles, and does not believe that vitalism will arise from its ashes. There are, to conclude, “many other philosophical issues on which a biologist must necessarily have a position,” and they cluster around questions of causality and the nature of explanation. “Biologists (like engineers) are endlessly confronted by historical causation and by positive and negative feedback loops not normally considered by physicists, or by philosophers for whom physics is the exemplar of science.” Williams claims to be baffled, as D. S. Wilson has also been, by the terms holist and reductionist as used by various writers who assume that the former have specimen jars and aquariums and the latter petri dishes and test tubes. He jokes about the distinction. “In most discussions I have heard, the distinction depends mainly on the sizes of glassware one uses in research.” So as he indicates, “I work more with jars and aquariums but prefer to ally myself with the reductionist camp, mainly because holism is too often a seemingly respectable label for mysticism and pedantic obfuscation.”36 Quite simply, mechanism is to vitalism as reductionism is to holism. The latter terms or perspectives are useless.

Hand and Eye, Thought Experiments, Evolutionary Texts, and Contexts Williams’s method explores evolutionary theory with a set of thought experiments, which are expressed as narratives. There is, for him, no other way to answer “why?” questions than through this kind of writing as a mode of thought. The efficacy of thought experiments continues to be discussed by philosophers of science, but one thing is certain. As has been

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well documented by examining Darwin’s revisions to On the Origin of Species, thought experiments are not simply transcribed in language; they are conceived, revised, and perfected in and through the use of language. Thought experiments are not simply portrayed through narratives, but their origin also lies in carefully edited narratives.37 In a parallel, Williams argues later, “I suppose that evolutionists do claim that natural selection creates genetic information, but I think the intended meaning is that it edits information. The original texts were chemically generated patterns, information only of conditions prevalent at their points of origin.”38 When I claim that Williams’s thought experiments are interesting literary narratives, worthy of close examination for their genesis, style, and structure, and when I also suggest that reading them must be a kind of thought experiment at large, there is no secret about this. Williams announces early and often that his thought experiments employ a methodology that is a priori rather than empirical because these narratives cannot proceed by observation or physical experiment—though they can sometimes be tested by those means. Though carried out as written narratives, his own thought experiments, like Darwin’s, rely on material causes for phenomena. In this original sense of method, he is truly Darwinian. He is a Darwinian writer. Like Darwin’s, Williams’s thought experiments undergo transformations over the years as he asks slightly different questions and reveals different possible or tentative solutions. One must be attentive to these transformations to acquire a nuanced sense of his changing perspective. Critique is always central to the interest of Williams’s thought experiments. He tests various modes of explanation, integrating relevant empirical evidence as thoroughly as possible. This becomes more onerous and burdensome over the years as the empirical literature swells. Ultimately, his thought experiments are designed to test the limits of evolutionary explanation. In this sense they do not mean to be “original.” By testing and editing a variety of inherited modes of explanation, he opens up (or unpacks) questions key to various modes of answering. Thought experiments often explore what cannot be explained—yet—and offer challenges to future theorists. In this sense, Natural Selection (1992) is a culmination of his thinking, an open-handed review of many possible modes of Darwinian explanation. Not least of the challenges facing biologists, as Williams suggests, is that they find themselves doubly in medias res, wondering about natural selection eons after the origin of the natural process while writing within a

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tradition that spans centuries. Consequently, he frames the book with a few classic pre-Darwinian texts that discuss adaptation in what he calls admirable ways. Galen, Paley, and Hermann Joseph Muller captivate Williams because they “illustrate how adaptation can be explored by seeking conformities to a priori design specifications.”39 All three of these classic writings investigate perfection of design in organisms. Paley, who finds the watch and the designed eye, and Galen, who wonders about the design of the hand, would be familiar to most readers. These texts might easily be models of close observation for poets too, like Ezra Pound in ABC of Reading.40 Muller, a well-known geneticist, was awarded the Nobel Prize in medicine in 1946 for his discovery that x-rays can induce genetic mutations. The Muller text uses this tool of inspection to elucidate a different interest. Muller argues that “long persistent morphological, physiological or biochemical features, characteristic of large populations, should in general be assumed to have, or to have had, adaptive value, under the conditions in which these groups evolved, and that the degree to which they are normally developed is also of adaptive significance.” He goes so far as to justify applying what he calls “a ‘teleological’ viewpoint here, which is in no wise mystical.” As Muller puts it, if researchers should fail to find adaptive “significance, or function,” that failure would be “an indication of our ignorance, and of the need for a deeper understanding of their workings.” Muller’s conviction about adaptation applies especially to medicine, where he advises “caution in attempting the alteration of standard bodily reactions, or the extirpation or alteration of structures normally present, until the functions of these parts and processes, and of their normal degree of development, have been well determined.” Yet he also concludes that “selection itself has been subject to grave limitations,” so “the organism is not perfect in any absolute sense, otherwise there would be no use in medicine, or in any form of artificial ‘interference’ with nature.”41 These views correspond very closely to those of Williams.

The Material and Codical: The Gene as Text and Unit of Selection In Adaptation and Natural Selection (1966), Williams writes, “In this book I use the term gene to mean ‘that which segregates and recombines with appreciable frequency.’” He posits that “such genes are potentially

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immortal,” since they may reproduce faster than external agents can destroy them, and “also have a high degree of qualitative stability.” As we remember, the material gene is a sequence of nucleotides coding for a protein or part of a protein; it may exist in alternative forms, called alleles, and most do not change except in the rare event of mutation; genes have existed unaltered for millions of years. As a unit of heredity, Williams continues in 1966, “the genotype is a coded message that is interpreted in some way by the soma. A gene is one of a multitude of meiotically dissociable units that make up the genotypic message.”42 In the second chapter of Natural Selection, Williams engages in an elaborate revision and restatement of problems associated with genes, codes, and genotypes. In essence, he rebuilds theory from genes up. This becomes clear when he begins to speak of the codex, the codical domain, and “the conceptual complication that genes are parts of genotypes.” He admits, “while I think it important to keep separate the codical and material domains, I would not insist that the codex concept completely replace the idea of a replicator.”43 Williams introduces separate domains because one of his purposes is to provide a “gene’s-eye view” of what he calls “clade selection.” Yet what also strikes a reader is an extended analogy between the process of writing about design and the processes of natural selection. The links between the two are most apparent in his second chapter, where he reminds the reader that he has always regarded the gene as a cybernetic entity, as he first wrote in 1966: “In its ultimate essence the theory of natural selection deals with a cybernetic abstraction, the gene, and a statistical abstraction, mean phenotypic fitness.”44 In discussing the codical domain, “we use such terms as bits, redundancy, fidelity, and meaning,” while the material domain is discussed in terms of color, charge, density, and volume. “The only descriptor they have in common is time, and events in one domain can be established as before, after, or simultaneous with events in the other.” He presents both material and the codical domains as real domains. Nevertheless, the codical is his focus, and he insists that “the idea that the gene is a package of information, not an object, is neither new nor idiosyncratic.”45 The material of the gene is, of course, DNA. As a chemical constituent of life, DNA is part of a phenotype. But the codical concept of the gene is a tool to think with, which does not mean that genes are not real entities. To understand the differences between what he calls the material and codical domains, one must imagine sets of genes as texts, as information,

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and so he posits a distinction between medium and message in biology. Williams uses Don Quixote for an analogy. This familiar text begins, Idle reader: thou mayest believe me without any oath that I would this book, as it is the child of my brain, were the fairest, gayest, and cleverest that could be imagined. But I could not counteract Nature’s law that everything shall beget its like; and what, then, could this sterile, illtilled wit of mine beget but the story of a dry, shrivelled, whimsical offspring, full of thoughts of all sorts and such as never came into any other imagination.46

It is impossible not to see this opening passage—one that Williams pointedly does not extract or cite—as a shadow text, another sly allusion by Williams to his own writing, to nature’s laws, and perhaps even to tilting at windmills. The medium, however, is not the message. According to Williams, Don Quixote can be conceived of as coded information that is only sometimes transmitted as ink on paper. It is also an example of an evolving cultural text because “it is often transmitted visually and stored ephemerally in human brains.” The text can be transmitted on magnetic tapes or disks, yet “in any medium, Don Quixote can form an archive from which copies can be made to any other medium, but no matter what the medium, it is always the same book.” He asks his readers to “think of genes as analogous to books in this respect,” where DNA is the medium but not the message.47 Twenty-first-century literary critics may not approve Williams’s definition of a text. Critics who taught his generation of undergraduates— the New Critics—would be more amenable to Williams’s view. To say that Don Quixote is a text, as Williams understands it, is to say that its message is independent of its medium. The study of culture fundamentally requires continuity of information, so that “there is more intellectual interest in the survival of Don Quixote as information than in survival of particular ink patterns in paper books or of magnetic orientations on tapes.” Understanding the codical domain is a matter of understanding the way texts survive, proliferate, and can be transformed. The message may be thought of as an abstraction, a “package of information,” but to be passed on accurately, information must proliferate faster than it changes, so as to produce a genealogy recognizable by some diagnostic effects. When Williams includes a figure of a dendrogram (Fig. 8.1), it is drawn in an unusual and interesting way, “line thickness indicating numerical abundance” within the population. (That the explanation for this

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illustration uses more space than the illustration reveals how uncomfortable Williams is with graphic representations.) Endorsing Dawkins and Hull’s distinction between replicators (genes) and interactors (phenotypes), Williams writes, “For natural selection to occur and be a factor in evolution, replicators must manifest themselves in interactors, the concrete realities that confront a biologist. The truth and usefulness of biological theory must be evaluated on the basis of its success in explaining and predicting material phenomena.”48

Terminology, Determinations, and Individuals What kind of language is appropriate here? Throughout his entire career Williams makes a sharp distinction, like Tinbergen, between the mental, a realm he considers unknowable by biology, and behavior, which can be observed and studied. Natural selection cannot see the mental. Gene and clade remain key terms “as strictly codical concepts.” As for their physical manifestations, he speaks of a parallel process that he calls phylad selection, a phylad being a group of closely related species of common origin, as opposed to a phylum, which is a taxonomic distinction for closely related organisms. As he moves up the ladder to higher levels of selection, he finds the phrase species selection obfuscating because it “implies that there is something special, for natural selection, about the species level of classification,” and he even accuses himself of having coined an “inappropriate term,” taxon selection, in 1985, because it “implies that a clade must be taxonomically distinguished to be relevant to selection.”49 Just as natural selection cannot see the mental, so too it can see neither a species nor any taxonomic category: these are only human classifications. Williams reminds his reader in 1992 that the term kin selection was introduced by Maynard Smith in 1964, but “the individual property selected for when relatives regularly interact is what Hamilton (1964) called inclusive fitness.” He does not have any qualms about continuing to use Dawkins’s famous reference to “selfish genes”—an anthropomorphic concept borrowed from Williams. Williams makes his own distinction between medium and message partly to justify Dawkins’s characterization: “He might well recognize it [the gene] as the bearer of a selfish message: Exploit your environment, including your friends and relatives, in ways that maximize my proliferation.” Within an organism, it appears that selfishness is controlled. Williams quotes with approval an extended anthropomorphic metaphor by E. G. Leigh on the internal affairs between cells:

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“It is as if we had to do with a parliament of genes, which so regulated itself as to prevent ‘cabals of a few’ from conspiring for their own ‘selfish profit’ at the expense of the ‘commonwealth.’”50 Systems do interact, but rather than thinking of this in terms of selfishness, Williams says “natural selection is a system of corrective feedback that favors those individuals that most closely approximate some best available organization for their ecological niche.” So what is an individual? “The term individual in abstract discussions in population genetics usually refers to a genotype, produced by a sampling of genomes from a gene pool.” Botany “refers to physiologically defined individuals as ramets and the whole collection of ramets of the same genotype as a genet,” and so he extends this botanical terminology to animal clones.51 Individuals are determined by differences. Late in his book, Williams presents what he calls a complete list of “causes contributing to differences between two individuals.” These differences may occur in different ways: (1) Ontogenetic stage. Consider the tadpole and the frog as “a classic kind of taxonomic challenge (e.g. which tadpole goes with which frog?).” (2) Age. “They may be of different age, a common reason for being different in stage, but they could be of the same age but different stages. If so, the one in the more advanced stage probably has higher phenotypic fitness.” (3) Frequency-dependent strategy. Frequency-dependent selection occurs when the fitness of a genotype or phenotype in a population is related to its frequency in the population. This can be positive when the more common variant is fitter, and negative when the less common variant is fitter. “They may be employing different life-­ history strategies maintained by frequency-dependent selection,” not just regarding sex differences, but within a sex there may be more than one reproductive strategy, and seemingly similar individuals may behave differently, because some polymorphisms are triggered by environmental cues. (4) Optimized response. “They may show individual adjustments to environmental factors other than the strategies employed by associates.” (5) Genetic load. “One or both individuals may suffer from one or more kinds of inherited defect.” There may be “locally maladaptive

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genes” that allow sexual reproduction to generate “maladaptive genotypes from adaptive genes.” ( 6) Epigenetic load. “They may differ in effects of environmentally inflicted trauma or deficiency (relative to some ideal that may be seldom experienced): poorly healed or unhealed injury, parasite load, inadequate nutrition, trace-nutrient scarcity, culturally inherited low social status, deception or manipulation by a competitor or exploiter.”52 At large, individuals are defined by these possible differences in fitness. His list reveals, specifically, how an individual appears from the point of view of natural selection.

Myths About Species, Niches, Optimization, and Ecology The expression clade selection helps clarify two fundamental ecological concepts. The “law of competitive exclusion” says that two species that compete for the exact same resources cannot stably coexist. Ecological succession refers to predictable and sometimes orderly changes in the composition or structure of an ecological community. Clade selection gives ecologists “a concept a bit broader than competitive exclusion or ecological succession,” and “it does not require that selection act among ecologically competing entities.” Williams says the concept of clade selection “differs in practice in its emphasis on the geological rather than the ecological time scale, and perhaps on a broader spatial perspective.” Although “studies of ecological succession do not normally allow for major changes in the ecological niches of the populations that succeed each other,” such a practice “would be routine in considering clade selection. The main difference may be simply in the attitude and purposes of users of the concepts.”53 Although he speaks frequently of internal systems—reproductive, digestive, nervous—within organisms, Williams is quite skeptical about the concept of an orderly external ecological system for the same reason that he considers the phenomenon of group selection rare. Any goal for evolution, if one can speak in this teleological way, is chemical, mechanically “selfish,” and repetitive. The process is adaptive only at the same time

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as it is immensely inefficient because adaptation itself is ad hoc and directed only toward survival, which is to say reproduction. For this reason, there is a great deal of teleological trouble when, in discussions of natural selection, one attempts to speak of the strategy and tactics of a species. Williams distinguishes between designs and their parameters in an attempt to circumvent such teleological language. He praises one researcher, Mart R. Gross, who insists that “the term strategy be used only for characters or character complexes maintained by frequency-­dependent selection,” generally defined as when the fitness of a genotype depends on its frequency, positive or negative, in a population. “Each strategy is composed of tactics, and any quantitative aspect of a tactic, in Gross’ usage, could correspond to one of my strategy parameters subject to optimization.”54 There is a point to his tortured syntax. Generally speaking, as the foregoing suggests, Williams shuns any terminology that turns relative measures into absolute states of being. When biologists speak of natural selection achieving an optimum, they mean that “natural selection acting on variation in a quantitative character (parameter of a design or strategy) can often be expected to fix the mean value near the functional optimum and to minimize the variation about that mean.” As Williams notes, “use of this idea is often called optimality modeling. I prefer the term optimization, which implies a corrective tendency, rather than a state attained. Likewise I think that calling a character optimized is always more realistic than calling it optimal.”55 This is not nit-picking because optima are always relative, just as the concept of a species is fluid.

Avatar One definition of avatar is a visible manifestation or embodiment of an abstract concept, as in the physical form taken by the god Vishnu during his visits on earth. Williams uses the term to mean a concrete and local population of the virtual entity called a species. Niles Eldredge, the paleontologist, defines an avatar similarly while attempting to preserve the idea of a species. He begins with the economic premise that organisms interact locally with others of the same species, so that “such local populations of organisms of the same species in this explicitly economic sense are termed avatars, meaning local representatives of a particular species.”56 Each avatar plays a role and has its own “niche” within that local ecosystem.

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But for Williams, we must remember, an individual is nothing but the gene’s avatar. According to John Damuth, from whom Williams acquires the term, when selection is made from among populations within any community, these populations are avatars of species. Avatars, he argues, are close analogues to individual organisms in traditional selection theory. “Just as populations evolve by organismic selection, communities evolve by avatar selection, and more inclusive units, the higher-level analogues of the species, evolve as their component communities do.” Damuth argues that higher-level selection is congruent with processes at the lower, organism-­based level.57 A recent article on the concept of “niche” explains that a shoal of fish in a lake or a herd of buffalo can be conceived as avatars because they are “causally integrated” and “more or less reproductively isolated sub-populations.” The main point is that “avatars, not whole species, experience selective pressures at the level of the group.”58 Since a species is not a god, and since only populations participate in the processes of the material world, the term avatar represents something more concrete, actual, and realistic. For these reasons and others, Williams finds the very term species useless. “The species,” he writes, may be “a key taxonomic concept, but that is all it is.”59 He is so troubled by the way other biologists use Mayr’s “species concept” that he lists problems associated with using the term species.60 As Williams argues, “to endow the species concept with theoretical significance beyond that of a widely recognizable level in the taxonomic hierarchy” is a frequent practice and “a source of much mischief in biological discussions.” Not only does the concept carry “a lingering Aristotelian essentialism,” but “ecologists often use the term species to mean ‘that which occupies a niche in the trophic structure [food chain],’ and this assumption is violated by the great majority of the world’s species.” Insofar as anyone might say that a species inhabits a niche, the notion of a separate, unique niche is itself problematic. Indeed, discourse of species and niches tends to be tautological. To speak as if a species were a unique individual or inhabits a unique place in a system results in a variety of “erroneous implications.” This is particularly apparent when paleontologists use the term species for their specimens; consequently, “the species rank has been accorded unwarranted significance in clade selection” by a set of paleontologists and biologists.61 These concepts are fictive, as Williams argues in a section titled “Fallacy of species niche occupancy,” where the discussion of terminology reaches its highest pitch and comes to a crisis. If one is to understand how the

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concept of species is used in community ecology, one must also recognize, as has been done generally, “that the niche of a single species may vary widely over its geographic range.” Confusion reigns because “effects of ontogeny on trophic relations can be extreme and have led some workers to speak of ecologically different developmental stages of a single population as separate trophic or ecological species.” In his example, “tuna of 1 g, 100 g, and 10 kg may be of the same population of the same species, but can hardly play the same roles in community ecology.” As far as he is concerned, “this whole difficulty would disappear if there were an acceptable substitute for species for community ecologists,” and this is why he is tempted to use Damuth’s term “avatar for the collection of individuals of a single species found in a particular community.” Biologists would “still need terms for ecologically diverse developmental stages and morphs of the same avatar.”62 The idea of avatars draws the reader into the realm of literary representation, where, as I was taught, a literary symbol partakes of the object it represents. A famous passage from Moby Dick reveals exactly this process. Ishmael contemplates the dignified brow of the sperm whale, so immensely amplified, that gazing on it, in that full front view, you feel the Deity and the dread powers more forcibly than in beholding any other object in living nature. For you see no one point precisely; not one distinct feature is revealed; no nose, eyes, ears, or mouth; no face; he has none, proper; nothing but that one broad firmament of a forehead, pleated with riddles; dumbly lowering with the doom of boats, and ships, and men.63

And so too with the creatures Williams introduces to readers, avatars all of them, whose descriptions partake of the objects they represent. If the foregoing examples illustrate why Williams believes the language and thinking of macroevolutionary theory is jumbled, when he endorses the term avatar, he seeks a discourse that focuses on real biological entities while he tries to avoid suggesting essentialism, permanence, or fixation—suggestions that often blind biologists to important changes in populations and environments—yet he cannot succeed entirely. That is why the concept of species is fictive, as Williams makes even clearer when he speaks of the “fallacy of species individuality.” By this point in the book, the reader should imagine that a clade, though it represents a lineage, functions as an individual in the codical domain, and a phylad as an individual in the material domain, only, as he insists, “in a

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particular sense.” “Every such entity has a more-or-less recognizable beginning, at least a presumed end, and a unique history and set of attributes in between.” Yet the beginning of a new population or gene pool “may be poorly defined.” Consequently, the origin of a species “must be far more nebulous than population origins. Given that there are now two populations where there was only one, how much and for how long must they diverge before we can say that there are two species where there was only one?” The only possible answer is a guess, preferably by a neo-­ Darwinian taxonomist, that “the two forms will never again freely interbreed even if sympatric,” that is, dwelling in the same geographic locale.64 The origins of species are ordinarily less decisive than the origins of populations, as is apparent from his illustration drawn from the study of the human lineage. In a subversive and understated way, Williams indicates that the very expression “origin of species” is deceptive. Consider humans as avatars. The best work Williams can find suggests that the three lineages Pan, Gorilla, and Homo separated from a common ancestor nearly simultaneously. The researchers he cites speculate that newly diverged chimpanzee, gorilla, and human lineages were only subspecifically differentiated members of the same species. As Williams then supposes, “this must have been followed, in our own lineage, by a time when Australopithecus and Homo were only subspecifically distinct, and then by a time when H. sapiens had diverged only slightly from H. erectus.” He finds that “the compass of human behavioral adaptations cannot be infinite, but it is surprisingly broad.”65 As I imagine, he is saying that human behavioral adaptations are not original, but broadly sourced from and often included in multiple previous lineages. Those who speak of species as distinct entities imply that “clades of greater or lesser inclusiveness are not individuals in the sense claimed for species,” but as Williams demonstrates, populations may undergo both fission and fusion. Examples of fusion include two species of Mesoamerican towhee that may have fused in historical time or the interbreeding of blue goose and snow goose where they come into contact on the west coast of Hudson Bay. “It is now widely realized that both extrinsic and intrinsic isolation between populations may be temporary,” so the concept of “species individuality can only foster confusion.” This has practical implications that Williams underlines: “Concern for endangered species can be counterproductive when what is really needed is concern for evolutionarily significant units.”66 If the goal of evolutionary science is to understand

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change, then a desire to preserve the taxonomy of the present may be fruitless. Individual species are not as important as avatars.

How Important Is Clade Selection? Williams spells out his analogical reasoning for the reader. After all, “the ultimate prize for which all clades are in competition is representation in the biota.” Given the obvious difference in fitness among clades, one clade must have a set of genes more likely than another’s to lead to long-term clade survival. This means that the ratio of probabilities of cladogenesis and extinction is different in the two clades… . The microevolutionary analogue of the extinction of a clade is the extinction of a gene, not the death of an individual.

Williams is not alone in this kind of reasoning. He finds his view of clade selection conforms to that of Van Valen and is contrary to those of others. One must look for any group-level adaptations, such as a thoroughly cooperative social system, “as a product of clade selection and evidence of its operation in the past,” but the reasoning cannot be turned on its head. “To make it a prior condition for the occurrence of clade selection is like starting with a singing bird in an evolutionary explanation for bird song.”67 When a biologist looks at a viable population, what does he see? Williams reiterates, “In my opinion the recognized microevolutionary processes that form the heart of the neoDarwinian synthesis are an adequate description of the evolution taking place in any Mendelian population.” Clade selection must be a sum of microevolutionary processes working on individuals. Natural selection of alternative alleles acts largely independently at each locus. Therefore, natural selection must be the only force tending to maintain or improve adaptations shown by the ephemeral organisms formed by the ephemeral genotypes… . If one could look back through the evolution of our own or any other sexually reproducing species, back to well before the Cambrian, no other fitness-enhancing process of any importance would be found.

This is the hard line of gene selectionism. But it will not quite do. Having taken that position, I must also take another. The microevolutionary process that adequately describes evolution in a population is an utterly

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inadequate account of the evolution of the Earth’s biota. It is inadequate because the evolution of the biota is more than the mutational origin and subsequent survival or extinction of genes in gene pools. Biotic evolution is also the cladogenetic origin and subsequent survival and extinction of gene pools in the biota.

Nevertheless, “This [latter] position is not equivalent to a belief in the efficacy of clade selection.”68 What a remarkably ambivalent statement about a concept that he himself advocates! As one might ask, “Why is he trying to have it both ways? He can’t live with clade selection. And he can’t live without it. Or maybe clades can’t live without it?”

Historicity and Constraint In his sixth chapter, “Historicity and Constraint,” Williams discusses the doctrine of historicity “with related evolutionary constraints (phylogenetic, developmental, genetic) on the action of natural selection.” But he begins the chapter by reminding the reader that his book is about natural selection “with all discussion constrained by acceptance of the two other doctrinal bases of modern biology, mechanism and historicity.”69 Surely speaking about history and evolutionary constraint while also admitting that one’s own argument is historically constrained constitutes a kind of meta-argument. In 1995, Kurt Schwenk published “A Utilitarian Approach to Evolutionary Constraint,” in which Williams is not discussed. However, Schwenk discusses the neo-Darwinian focus on “positive or constructive view of evolutionary change” for which the adaptationist program has been criticized.70 Many believe that Williams exemplifies and fully endorses this program. Certainly he inspired it and defends it in several books. Yet “constraint” is a word Williams uses only once in Adaptation and Natural Selection (1966), and when it becomes a major theme in Natural Selection, one ought to pause. Williams’s new emphasis on constraint constitutes what I believe to be a kind of paradigm shift in his thinking. Williams’s most explicit appeal to Darwin’s texts comes in this context. He speaks of the final chapters of Darwin’s On the Origin of Species as a model for his own. “With hindsight we can see that Darwin’s chapters successfully surmounted most of the difficulties and answered most of the objections. I expect no comparable success, but hope that I can at least convince some others that evolutionary biology today, no less than in

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Darwin’s time, faces some serious anomalies and conceptual challenges.” Just as Williams chooses Galen and Paley for his appendices because their “main arguments are thoroughly mechanistic, and their occasional errors arise from the primitive state of physical science in their times, and from their total ignorance of natural selection and historicity,”71 so too he chooses Darwin as a methodological model residing within the history of evolutionary thought. Natural Selection, more than any of Williams’s other books, not only depends on Darwin’s scientific authority but also takes literary authority from him. Williams claims he is using Chaps. 6 and 7 of the 1859 edition as his model, but as the titles for Darwin’s Chap. 7 make clear, he is using the sixth edition of The Origin. There Darwin lays out the “Difficulties of the Theory” as a set of questions. It is also the chapter in which he discusses the eye. Darwin’s Chap. 7 begins by confronting an objection: “the weakest part of my theory is, that I consider all organic beings as imperfect.” He reminds the reader “what I have really said is, that all are not as perfect as they might have been in relation to their conditions.”72 Darwin’s chapters take the form of refutation. Williams’s concluding chapters take the form of anomalies and challenges frequently set out as questions that have not been satisfactorily answered. Some are particularly engaging, but they read like a laundry list. I can find no clear structure to them. So I shall pass over Chaps. 8 through 10 of Natural Selection to focus on Williams’s appendices.

Paley’s Eye, Darwin’s Eye, Williams’s Eye “In crossing a heath,” Paley wrote in 1836, “suppose I had found a watch upon the ground, and it should be inquired how the watch happened to be in that place; I should hardly think of the answer which I had before given, that, for any thing I knew, the watch might have always been there.”73 For the precision of its design, Paley likens it to the design of the human eye, and famously so. Williams ceaselessly returns to Paley’s eye, and to his argument from design, in order to dramatize how recursive the process of examining and conceiving its design must be. Williams’s condensed version of this literary tradition, which includes Paley on the eye, Darwin on the eye, and Williams on the eye, contains a clear paradigm shift. In a very real sense, the Williams eye and the Paley eye are literary objects, symbols of perspective, found embedded in literary history. Paley’s description itself depends on a historical legacy, for as

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Williams also observes, “Galen and Paley built heavily on the works of Aristotle.”74 Aristotle, in turn, was a pioneer of medicine. Darwin intervenes. Moderns are taught, as Mayr writes, that “after 1859 it was no longer necessary to invoke design, a supernatural agency, to explain the adaptation of organisms to their environment.”75 Yet biologists continue to invoke design, though in different ways. What similarities remain? Williams allows the reader to recognize similarities and differences in these perspectives on the design of the eye itself, and more importantly, on the process that has brought it about. Tim Lewens has recently said of explanatory adaptationism that its “salience is derived from Anglo-American biology’s roots in natural theory. This conclusion is ironic: far from expressing the enmity between modern biology and natural theology, explanatory adaptationism is testimony to the fellowship between the two traditions.”76 Indeed, Williams confronts his own kinship with natural theology directly and exploits it. Rather than arguing against Paley’s natural theology, Williams consistently develops it in the light of modern knowledge. As Paley wrote, “I know no better method of introducing so large a subject, than that of comparing a single thing with a single thing; an eye, for example, with a telescope. As far as the examination of the instrument goes, there is precisely the same proof that the eye was made for vision, as there is that the telescope was made for assisting it.”77 In “How to Carry Out the Adaptationist Program,” Ernst Mayr reminds his reader of Darwin’s comments on the eye.78 And since Darwin’s comments on the eye are part of The Origin’s Chap. 6, they are part of Williams’s model. Under the subtitle “Organs of extreme perfection and complication,” Darwin admits, “To suppose that the eye, with all its inimitable contrivances for adjusting the focus to different distances, for admitting different amounts of light, and for the correction of spherical and chromatic aberration, could have been formed by natural selection, seems, I freely confess, absurd in the highest possible degree.” Yet he bends to reason and decides, “the difficulty of believing that a perfect and complex eye could be formed by natural selection, though insuperable by our imagination, can hardly be considered real.”79 Williams’s eye is not the “organ of extreme perfection” that Darwin’s is. “For the same reason that it was once effective in the theological ‘argument from design,’” he writes in 1966, “the structure of the vertebrate eye can be used as a dramatic illustration of biological adaptation and the necessity for believing that natural selection for effective vision must have

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operated throughout the history of the group.”80 In principle, he notes, “any other organ could be used for illustration,” but the optics of the eye are particularly convincing. There is more to it than optics. Paley creates an analogy between the found watch and the eye; consequently he yokes time, space, and a history of design. As Williams contemplates time, space, and design through his entire career, he continues to transform Paley’s text, and by implication Darwin’s too. Thirty years after Adaptation and Natural Selection, when he returns to what he calls his “chosen classic,” Paley on the vertebrate eye, Williams does so not only because it was used as a “particularly forceful part of his theological argument from design,” but also because the eye, though “surely a superbly fashioned optical instrument,” is also “a superb example of maladaptive historical legacy.”81 The argument that Paley abstracted from—or imposed upon—the eye includes close observations and careful inference. Williams admires its lucidity. As Paley writes, “the inference, we think, is inevitable, that the watch must have had a maker; that there must have existed, at some time, and at some place or other, an artificer or artificers, who formed it for the purpose which we find it actually to answer; who comprehended its construction and designed its use.”82 Williams insists that such passages are “worth close attention by all biologists.”83 First of all, as he wrote in 1966, there is “no better illustration of this process of recognizing functional design from careful observation.”84 And because Paley’s watch is linked to Paley’s eye, Williams must transform it while following the tradition: “Adaptation is demonstrated by observed conformity to a priori design specifications.”85 Paley’s “superbly fashioned optical instrument” transforms itself under modern scrutiny to become “something else, a superb example of maladaptive historical legacy.” This is an argument Williams will use to address the medical profession soon. Transformation implies difference, so, as he puts it, “the fact of maladaptive design, however minimal in effect, spoils Paley’s argument that the eye shows intelligent prior planning,” and close investigation reveals “real and routinely demonstrable” defects in human vision.86 In Paley and ever after, watch becomes eye. Unlike some stone on the ground anyone might encounter, neither the watch nor the eye has been there forever (and nor in fact has the stone). Between two symbols, space transforms itself into time and vice-versa: Williams’s eye comes to exemplify “maladaptive historical legacy.” Simple spatial analysis reveals that the retina is upside down and light reaches the rods and cones of the retina

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only after passing through nerves and blood vessels. The human eye is, to this extent, a distorting lens. Williams historicizes this: “There would be no blind spot if the vertebrate eye were really intelligently designed. In fact it is stupidly designed, because it embodies many functionally arbitrary or maladaptive features, of which the inversion of the retina is merely one example. These features are there, not for functional but for purely historical reasons.”87 This, in turn, allows Williams to conclude Natural Selection with what I will call the “Allegory of an Eye Blink”: “A human eye blink takes about 50 milliseconds. This means that we are blind about 5% of the time when we are using our eyes normally.” Blindness is not a good thing. One can imagine many ensuing problems, events of importance in warfare and other arenas. Why, then, Williams asks, would natural selection not replace 95 percent visual attentiveness with 100 percent by designing eyes that blink alternately or independently? The answer can be imagined as “some sort of trade-off balance”—if nobody will be able to see a design that has never happened. He knows the standard argument: a “blink mechanism for both eyes at once may be much simpler and cheaper,” or the gain might not be worth the cost.88 But if independent blinking had ever appeared, Williams cannot see how synchronous blinking would have any advantage over independent blinking, certainly not enough to make it disappear. So, what of Williams’s idea of adaptive alternation or independent timing? “Am I assigning too much importance to 50 milliseconds of blindness every second?” he asks, which might be code for “Am I belaboring a trivial point?” Well, the eye is an elaborate and essential mechanism, and there must be a reason for the flaw in design, the flaw being a clue to the way design comes about. One must imagine many changes necessary to alter the machinery or mechanism. Could asymmetrical blinking be produced developmentally, would favorable mutations for it have other effects, and how would selection act? We would also like to know why certain kinds of animals, not just certain kinds of eyes, do not exist. The implications of Williams’s “Allegory of an Eye Blink” reach quite beyond Paley’s eye. We are told that the natural theologian moves from the observable mechanism or organ to the unobservable creator and the authority of the creator. Americans do not lack for modernist literary transformations of Paley’s argument from design. At the very least, a literate person would recall Robert Frost’s poem “Design” and Wallace Stevens’s poem “Anecdote of the Jar,” the first discovering a tiny design

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residing outside the human world, and the second discovering the way design is created within the mind. Williams, by breaking Paley’s argument, severs this implication of an unobservable creator and also erases final authority and finalized answers in several ways. Not only is there no designer of perfect organs, but there are also no perfect organs. On a more subtle level, in a post-Paley world— or even a post-Darwinian world, at least with regard to eyes—there can be no perfect or a priori explanations for organs or their adaptive value, and less explanation for the ones that we do not see. Yet good biological method calls for asking about what is not to be found. And Williams grounds his method on a priori thinking. Observations, Williams has argued, must be teleonomical, but explanation must also be historical, constrained by knowledge of the general process involved, including its contingencies. That explains in part why provisional stories cannot predict what evolution will do, but only what experiments can be conceived. The prediction of results is a tentative procedure. Consequently, every book by Williams has a provisional conclusion. The eye is functionally designed, one must say, but not intelligently designed; it is a contrivance without a contriver. So it is that Williams is able to argue about contemporary theories of “intelligent design”—a nearly obligatory task for all Darwinians— without mentioning it by name. This matter of tentative explanation also has a great deal to do with how one reads evolutionary literature. Most readers of nonfiction continue to seek final answers. Williams tells the reader how to think, but not what the answer will be. Of course the eye is for seeing, but seeing how well? And why not three eyes? It is perhaps emblematic that Williams also turns to R. A. Fisher in the conclusion of his last book specifically addressing biologists. Fisher believed that “the contemplation in natural science of a wider domain than the actual leads to a far better understanding of the actual,” and notes, in that context, that one ought to wonder what the consequences might be of three or more sexes. Unrealized possibilities, as Williams believed, are often as important as those that are readily apparent. What nature does not do ought to interest us for scientific and ethical reasons. So, as a parting shot, Williams expresses rosy confidence in a future with “many fine studies, theoretical and empirical, of adaptations shown by organisms,” but cautions, “I hope they also show increased attention to the adaptations not shown by organisms.”89

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Reconsiderations Regarding Adaptation, Genes, and Machines Bodies are ridiculous arrangements, like books. They are akin to ad hoc, made up fictions that barely suffice under the circumstances. But do suffice. Perhaps one has to be as old as Williams is, nearing what is normally taken as the age of retirement, to write a final book about evolutionary theory while watching theory itself become more complicated and various. It is hard to miss a tension between telling the story of his changing perspectives and telling the story of natural selection. When he integrates the “doctrinal base” of history into his doctrinal revision in the “Defense of Reduction,” he becomes much more inclusive in the perspectives he admits into his discussions, but he does not yet become a “pluralist”—to use Gould’s term. Gould argued, somewhat later, that pluralists “accept natural selection as a paramount principle (truly primus inter pares), but then argue that a set of additional laws, as well as a large role for history’s unpredictable contingencies, must also be invoked to explain the basic patterns and regularities of the evolutionary pathways of life.”90 Perhaps Williams increasingly conceives his role as a reader of his colleagues’ work, who can speak for their tradition. A few years later, when he is asked to comment on Gould’s work for John Brockman’s The Third Culture, he writes that “I have trouble understanding Gould’s persistent efforts to minimize the importance of natural selection, the adaptive changes it produces, and the other things it does.” Nevertheless, “It’s obviously true that there’s a lot of chance in evolution, at any level,” and particularly at the higher levels on the pure basis of small populations, because “the survival of one entity and the extinction of another is much more likely to be a chance event.” In this context, Williams admits, “Steve has done a great job of explaining the role of chance in macroevolution and its dependence on historical legacies. There may be a few scientists out there who are as good as Steve Gould, but there are just damn few who are good as he is at writing for a great range of readers.”91 Williams’s admiration for Gould’s writing reveals something about his own strategy. At large, one could assess his strategies over the decades, a strategy being the series of actions that define the behavior of a player.92 In this sense, Williams is a player in a game that has stakes, rules, and outcomes. (1) He becomes a player with Adaptation and Natural Selection in 1966; (2) he changes his focus to Sex and Evolution in 1975; (3) he reiterates the reductionist strategy in 1985; (4) in the late 1980s and early

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1990s, he makes several moves within evolutionary theory and evolutionary ethics; (5) in 1991, he begins to pursue Darwinian medicine with Nesse and publishes a book on the subject in 1995.93 Still engaged in a defense of reason, Williams’s selection of reasons has changed, or perhaps evolved. Gould expresses admiration for this book— or at least Chaps. 2 and 4—asserting that in it, Williams “may still be talking the talk of gene selectionism, but he is no longer walking the walk.” Gould remembers Williams sharing his views at a conference at Ohio State University in 1988, presenting “the substance of his 1992 book.” Afterward, in an interchange that followed, the philosopher Marjorie Grene said, “You’ve changed a lot.” According to Gould, “George Williams, one of the calmest and most laconic of men, replied: ‘It’s been a long time.’”94 As Williams would later comment wryly, “Adaptationist stories are not so much about adaptation as its opposite.”

A Big Question from the Big Book of Nature In September 1986, the Smithsonian’s Directorate of International Activities and the National Academy of Sciences co-sponsored a “National Forum on Bio-Diversity” held in Washington, DC. It included addresses and lectures by scientists such as E. O. Wilson, Paul Ehrlich, and Stephen Jay Gould. The sixth extinction, as it came to be called, constitutes a crisis of epic proportions. What Williams has to say about this is: The loss of the Stellar sea cow and the Adam-and-Eve orchid were the same kind of loss to historical scholarship as the burning of the library at Alexandria. The current wholesale extinction of organisms is especially tragic and ironic because we are only now learning to read history in molecular structure, where the writing may well prove clearer and more detailed than in morphology and other phenotypic end states. To the aesthetic and economic arguments for the conservation of taxa, we should add the argument that the white rhinoceros and the blue whale must be immensely informative books that we have not yet had the skill or the time to read.95

It is hard not to think of Williams as a fatalist. The library is burning, with its valuable codices, their information irretrievable; it will not return in any other form. Natural selection may be wasteful, but so are humans. This is biology noir, not the safe evolutionary theory that the modern synthesis provides. Williams’s light and way leads to very dark lessons

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about “man’s place in nature” and about “man’s nature.” Under these circumstances, for his next project he turns away from what one cannot remedy, to face what one might be able to resist. With such knowledge, how might one improve the lives of humans?

Notes 1. George C.  Williams, Natural Selection: Domains, Levels, and Challenges (New York: Oxford University Press, 1992), 11. 2. Richard Dawkins, “An Ecology of Replicators,” Ludus Vitalis 12, no. 21 (2004): 43–52. 3. Williams, Natural Selection, 3. 4. Williams, Natural Selection, 8, 72. 5. Williams, Natural Selection, 190, 191. 6. Williams, Natural Selection, 35. 7. R. K. Grosberg and T. M. Caro, “The Challenges of Natural Selection,” a review of Natural Selection: Domains, Levels and Challenges, Evolution 47, no. 6 (Dec. 1993): 1893–95, 1893. 8. Joel Cracraft, review of Natural Selection: Domains, Levels and Challenges, BioScience 43, no. 9 (Oct. 1993): 641. 9. Cracraft, review, 641. 10. Williams, Natural Selection, 24. 11. Cracraft, review, 641. 12. W.  Ford Doolittle, “Making the Most of Clade Selection,” Philosophy of Science 84, no. 2 (2017): 275–95. 13. Samir Okasha, “Does the Concept of ‘Clade Selection’ Make Sense?” Philosophy of Science 70, no. 4 (2003): 739–51. 14. Williams, Natural Selection, 6. 15. Williams, Natural Selection, 10. 16. Williams, Natural Selection, 11. 17. Williams, Natural Selection, 10, 15. 18. Stephen C.  Stearns, “George Christopher Williams 1926–2010: A Biographical Memoir,” National Academy of Sciences (Washington, DC, 2011), 13. http://www.nasonline.org/publications/biographical-­ memoirs/memoir-­pdfs/williams-­george.pdf. 19. Williams, Natural Selection, 23. 20. Grosberg and Caro, “Challenges,” 1893. 21. Ridley, “Cladism,” http://www.blackwellpublishing.com/ridley/a-­z/ Cladism.asp. See also http://www.blackwellpublishing.com/ridley/a-­z/ Cladistic_species_concept.asp. 22. Williams, Natural Selection, 23, 14, 27, 13, 18; see Fig. 2.1C. 23. Stearns, “George Christopher Williams,” 14; Williams, Natural Selection, 13.

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24. Williams, Natural Selection, 16, 18, 14. 25. Williams, Natural Selection, 17, 19, 20. 26. Williams, Natural Selection, 21. 27. Stephen J.  Gould, “Tempo and Mode in the Macroevolutionary Reconstruction of Darwinism,” Proceedings of the National Academy of Sciences 91 (July 1994): 6764–71, 6770. 28. Samir Okasha, Evolution and the Levels of Selection (New York: Oxford University Press, 2006), 211–17. Okasha demonstrates the weakness of this conception. See also Steven A. Frank “Natural selection. III. Selection versus transmission and the levels of selection,” Journal of Evolutionary Biology. 25 (2012): 227–43, 234. Frank uses Williams’s ideas for another assessment of clade selection. See also Stephen Jay Gould, The Structure of Evolutionary Theory (Cambridge MA: Harvard University Press, 2002), 1315, 1316, 1323. 29. Williams, Natural Selection, 3. 30. Williams, Natural Selection, 46–47. 31. W.  C. Allee, et. al., Principles Of Animal Ecology (Philadelphia: Saunders, 1948). 32. Williams, Natural Selection, 47. 33. Williams, Natural Selection, 4, 5. 34. Williams, Natural Selection, 6, 7. 35. Williams, Natural Selection, 7, 8. 36. Williams, Natural Selection, 8–9, 9. 37. James Robert Brown and Yiftach Fehige, “Thought Experiments,” The Stanford Encyclopedia of Philosophy (Fall 2011 Edition), edited by Edward N. Zalta, http://plato.stanford.edu/archives/fall2011/entries/thought-­ experiment/. See also Peter Swirski, Of Literature and Knowledge: Explorations in Narrative Thought Experiments, Evolution and Game Theory (New York: Routledge, 2007): 126–7. J.  Lennox, “Darwinian Thought Experiments: A Function for Just-so Stories,” in Thought Experiments in Science and Philosophy, edited by T. Horowitz and G. Massey (Savage, MD: Rowman and Littlefield, 1991): 223–46. 38. George C. Williams, “Reply to Johnson,” Biology and Philosophy 11, no. 4 (1996): 541. 39. Williams, Natural Selection, 190. See H.  J. Muller, “Evidence of the Precision of Genetic Adaptation,” Harvey Lectures 43 (1950): 165–229. 40. See Chapter 1 of Ezra Pound, ABC of Reading (New York: New Directions, 1934), 17–18. 41. Muller, “Evidence,” 229. 42. Williams, Adaptation and Natural Selection (Princeton, NJ: Princeton University Press, 1966), 24, 56. 43. Williams, Natural Selection, 10–16, 12.

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44. Williams, Adaptation, 33. Williams probably acquires the meaning of the term as “control and communication in animals, men, and machines,” from Norbert Wiener, Cybernetics, or Control and Communication in the Animal and the Machine (Cambridge, MA: The Technology Press, 1948). 45. Williams, Natural Selection, 10, 12. 46. Miguel de Cervantes, Don Quixote, translated by John Ormsby (Project Gutenberg). 47. Williams, Natural Selection, 11. 48. Williams, Natural Selection, 13, 14, 13. 49. Williams, Natural Selection, 23, 24. 50. Williams, Natural Selection, 19, 15, 43. 51. Williams, Natural Selection, 43. 52. Williams, Natural Selection, 91–93. 53. Williams, Natural Selection, 53. 54. Williams, Natural Selection, 60. 55. Williams, Natural Selection, 61. 56. Niles Eldredge, The Pattern of Evolution (New York: W.  H. Freeman, 1999), 163. 57. John Damuth, “Selection among ‘Species’: A Formulation in Terms of Natural Functional Units,” Evolution 39, no. 5 (1985): 1132–46. 58. Barry Smith and Achille C.  Varzi, “The nich,” Noûs 33, no. 2 (1999): 214–38. 59. Williams, Natural Selection, 119. 60. Ernst Mayr, Animal Species and Evolution (Cambridge, MA: Harvard University Press, 1963). 61. Williams, Natural Selection, 120. 62. Williams, Natural Selection, 120. 63. Herman Melville, Moby Dick, Chapter 79: “The Prairie.” 64. Williams, Natural Selection, 121, 122. 65. Williams, Natural Selection, 122, 98. 66. Williams, Natural Selection, 122, 123. 67. Williams, Natural Selection, 25, 26, 28. 68. Williams, Natural Selection, 31. 69. Williams, Natural Selection, 72. 70. Kurt Schwenk, “A Utilitarian Approach to Evolutionary Constraint,” Zoology 98 (1995): 251–62, 261. 71. Williams, Natural Selection, 106, 191. 72. I am using The Complete Work of Charles Darwin Online, edited by John van Wyhe (2002–), http://darwin-­online.org.uk/content/frameset?item ID=F391&viewtype=text&pageseq=1. 73. Paley quoted in Williams, Natural Selection, 195. 74. Williams, Natural Selection, 190.

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75. Ernst Mayr, “How to Carry Out the Adaptationist Program,” American Naturalist (1983): 324–34, 333. 76. Tim Lewens, “Adaptation” in The Cambridge Companion to the Philosophy of Biology, edited by Michael Ruse and David Hull, 1–21 (Cambridge, UK: Cambridge University Press, 2007), 21. Lewens argues this more elaborately in chapter 2 of Organisms and Artifacts: Design in Nature and Elsewhere (Cambridge, MA: MIT Press, 2004), 21–37. 77. Paley quoted in Williams, Natural Selection, 196. 78. Mayr, “How to Carry Out the Adaptationist Program,” 324. 79. Darwin, Origin (1859), 186, 187. 80. Williams, Adaptation, 6. 81. Williams, Natural Selection, 73. 82. Paley quoted in Williams, Natural Selection, 196. 83. Williams, Natural Selection, 190. 84. Williams, Adaptation, 258–9. 85. Williams, Natural Selection, 40. 86. Williams, Natural Selection, 72, 73. 87. Williams, Natural Selection, 72, 73. 88. Williams, Natural Selection, 152, 153. 89. Fisher quoted in Williams, Natural Selection, 153; Williams, Natural Selection, 153. 90. Stephen Jay Gould, “The Pleasures of Pluralism,” New York Review of Books (June 26, 1997), 47–52. 91. George C. Williams, “A Package of Information,” in The Third Culture: Beyond Scientific Revolution, edited by John Brockman (New York: Touchstone, 1995), 70–71. 92. See Herbert Gintis, Game Theory Evolving (Princeton: Princeton University Press, 2000), 5. 93. Game theory appears in Sex and Evolution (1975) and in Natural Selection (1992), 58. It is often tied to questions about engineering. See especially section 5.1 of Natural Selection for game theory and ESS. 94. Stephen Jay Gould, The Structure of Evolutionary Theory (Cambridge, MA: Harvard University Press, 2002), 644. 95. Williams, Natural Selection, 76. Bobbi Low calls Williams a guerrilla environmentalist, as I discuss later. See Bobbi S.  Low, “George Williams, Theoretician and Guerilla Environmentalist,” the Quarterly Review of Biology 80, no. 1 (2005): 72–83, 73.

CHAPTER 9

Repairing Human Natures: Why We Get Sick

Why We Get Sick: The New Science of Darwinian Medicine (1995), the book George C. Williams coauthored with Randolph M. Nesse, intends to inculcate evolutionary literacy in the medical community, integrate evolutionary thinking into the education of physicians, and educate patients in evolutionary explanations for medical problems. Speaking of the book’s origin and composition, Nesse remembers writing “in an engaging style, hoping that people would read it just for fun. That has worked out well.”1 As history has shown, the notion that humans are perfect has not worked out so well. We try to pretend. Prince Hamlet says: What a piece of work is a man! How noble in Reason! how infinite in faculties! in form and moving how express and admirable! In action how like an Angel! in apprehension how like a god! the beauty of the world! the paragon of animals! and yet …2

It is the “yet” that puzzles us all. A popular focus represents a logical professional step for Nesse, who chafed against the purely practical orientation of medical practice. Whether it was a logical step for Williams is more complicated. In a certain sense, it constitutes a continuation of the ethical turn he made with his essays on evolution and ethics, but it is also a kind of introverted turn toward human © The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 M. P. Cohen, George C. Williams and Evolutionary Literacy, Literatures, Cultures, and the Environment, https://doi.org/10.1007/978-3-031-11650-6_9

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biology, maybe even toward humanistic issues. Williams turned away from his focus on how biological processes function in the world at large—perhaps because medicine was one way to oppose those processes, or perhaps because, as Alexander Pope once wrote, “The proper study of mankind is man.” Why We Get Sick is, in that sense, a meditation on the biology of “human nature” that all humans have in common. Conceptions of “human nature” are not identical to ideas about the “human condition.” The cultural critic Raymond Williams, for instance, claims that there can be no static, interior, universal, or general human condition.3 In a different sense, when the philosopher Hannah Arendt writes The Human Condition, though she finds it rooted in the biological labor of reproduction, she also frames it as a cultural construct, an ideological structure with a history that is performed through human work and action in the public sphere.4 Because it merges with questions about the “human condition,” the question of “human nature” is one more contested territory. Nesse and Williams argue that both emerge from the human genome, from “biologically inherited conditions.”5 Consequently, the boundaries of nature and culture blur in this book—perhaps in a productive way. Here the interactions between nature and culture, genome and environment, are most interesting. What, for instance, is a “condition”? The term appears frequently in Why We Get Sick, in two different senses. The first is in the sense of a patient’s state of heath, or having a condition, and the second refers to the conditions under which humans have evolved, where they have been situated by necessity, by internal and external factors and circumstances. An endorsement of the book by E. O. Wilson slides between these two: “By bringing the evolutionary vision systematically into one of the last unconquered provinces, Nesse and Williams have devised not only means for the improvement of medicine but fundamental new insights into the human condition.”6 As Nesse and Williams argue extensively, humans live according to a biological nature that was designed to meet the challenges of past environments. Human biology reveals a historical legacy that constrains human resources for meeting changing environments in the present. Paradoxically, though medicine is, in many senses, a human’s best strategy in opposing biology, that is not entirely the case, because not all biology should be thwarted. As a basic principle, Nesse and Williams advocate remedying what can be remedied in human health and welfare, while not attempting to change what has been “adaptive” about human physical and mental capacities.

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As Randolph Nesse remembers, he “started looking for an evolutionary biologist who might want to collaborate on this project.” He met Williams, who had been looking for a physician collaborator. They talked and exchanged notes, seeking evolutionary explanations for cancer, schizophrenia, and Alzheimer’s disease. “Finally, in a simple but profound reframing, we recognized that diseases do not have evolutionary explanations. Instead, we turned our focus to aspects of the body that leave us vulnerable to disease. Then we made progress.” They created rules for composition as they sent drafts of chapters back and forth on computer disks. In his now no longer available blog, “The Skeptical Adaptationist,” he reminisced: One of us would write a first draft, the other would completely rewrite it. We agreed on a simple method for staying friends. The revising author could make whatever changes seemed sensible, but was not allowed to look back at his earlier draft. After half a dozen iterations for each chapter, we forgot about who came up with what. We argued at length about many issues, allergy, defenses, menopause, senescence, and morality. It was real back and forth, the kind of very productive careful listening and hard arguing that carves away vagueness.7

As the preface to Why We Get Sick indicates, Williams’s interest originated when he read a 1980 article by Paul Ewald, “Evolutionary Biology and the Treatment of the Signs and Symptoms of Infectious Disease.”8 Nesse and Williams acknowledge Ewald’s work and borrow his scheme for classifying infectious diseases.9 Ewald continues to work in the field and is now the editor of the Journal of Evolutionary Medicine, an open-access, peer-reviewed journal. Meanwhile, Randolph Nesse continues to edit The Evolution & Medicine Review. This is a burgeoning field and various authors have documented its emergence and growth.10

Originality and Argument Nesse and Williams begin by asking a Hamlet-like question, “Why, in a body of such exquisite design, are there a thousand flaws and frailties that make us vulnerable to disease?” This vulnerability to disorder applies not simply to physical but also to mental systems. They ask—though this is a rhetorical question—“are mental disorders diseases?” Sometimes, the text is prone to re-telling old jokes like “Doctors still say … ‘Take two aspirin and call me in the morning.’”11

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It is hard to know whether the blithe style of Why We Get Sick is engaging or disturbing, or whether such a tone befits the seriousness of the subject. The name of the subject is also debatable. Williams preferred the term “Darwinian medicine,” and Nesse “evolutionary medicine.” Nor is it clear that Darwinian or evolutionary medicine is a new science. In his historical survey, Joe Alcock sharpens this distinction, explaining that “Williams advocated for the eponymous ‘Darwinian’ label because of its connection to natural selection,” but “‘Darwinian medicine’ might be seen as the subset of evolutionary medicine that deals primarily with adaptation and natural selection,” while “‘evolutionary medicine’ allows for a more comprehensive array of evolutionary explanations in biomedicine.”12 The book is generally taken to apply an adaptationist perspective to both the physiological and psychological dysfunctions or vulnerabilities of humans. In this sense, it follows Williams’s established principles, including his principle of critique. Though the recommendations by Nesse and Williams are sufficiently new to be fluid, the methodologies come from traditional sources. Physicians normally look for “proximate explanations” to answer “what?” and “how?” questions, as Nesse and Williams note, while evolutionary explanations ask and attempt to answer “why?” questions about ultimate origins and functions. As Nesse defines the two kinds of explanations, the proximate “explains how organisms work by describing their structures and mechanisms and their ontogeny,” while the evolutionary “explains why organisms are the way they are by describing how selection shaped current forms and their phylogeny.”13 Their distinction between the proximate and the ultimate comes originally from Tinbergen and Mayr. By the 1990s, it had become a staple of adaptationists—almost a slogan: Look for the ultimate not the proximate cause!14 Nesse and Williams establish that knowledge of evolution can help physicians predict and consequently diagnose human maladies, disorders, or diseases. They repeatedly call attention to why questions. “Most medical research seeks proximate explanations about how some part of the body works or how a disease disrupts this function. The other half of biology, the half that tries to explain what things are for and how they got there, has been neglected in medicine.”15 But the adaptationist flavor of the book is itself a descendent of ways of thinking created by Williams. Their book expands on their opening salvo in an often-cited 1991 article entitled “The Dawn of Darwinian Medicine,” published in the Quarterly Review of Biology.16 The title is a nice touch, in view of Williams’s

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“the light and the way.” As Joe Alcock points out, scientists take this article as the landmark paper for evolutionary medicine, and it is indexed by the National Library of Medicine as having genetic selection as its major topic.17 The American title of the book is Why We Get Sick, in contrast with the British title, Evolution and Healing. Both use the same subtitle, The New Science of Darwinian Medicine. The cover of the American edition identifies Nesse as an MD and Williams as PhD. No other book by Williams includes his degree. In the acknowledgments, the authors state, “We are grateful to our agent, John Brockman, for convincing us that we could present serious new science in a book for a general audience.” It seems more likely that Nesse sought to publish a popular book, but negotiations with the infamous agent Brockman were somewhat strained. According to Nesse, Brockman thought Williams was rather strange, since he came to a meeting wearing untied tennis shoes and a threadbare sports coat.18 Brockman remembers the situation somewhat differently: “Meeting him for the first time when with Randy Nesse, who did all the talking (i.e. never stopped talking) about their planned book on Darwinian Medicine. Finally, I interrupted Nesse and asked the taciturn Dr. Williams, ‘Professor, what can I tell publishers about you?’ ‘Well,’ he replied, ‘I once wrote a little book for a university press, but it was thirty years ago. It probably won’t be of interest to them.’”19 By contrast, Nesse’s promotional energy is apparent in many publishing strategies that follow the book, not only in disseminating articles derived from it, but also in his catchy language and breezy style. Readers found this book lively but also quirky. Nevertheless it succeeds in continuing to catch the attention of doctors and patients. Nesse and Williams’s second premise is that the adaptationist program is an appropriate evolutionary method for investigating human attributes: “By suggesting the functional significance of some known aspect of human biology, you may logically be able to predict some other, unknown aspects. An appropriate investigation can then confirm that these characteristics are either there or not. If they are there, they may be of medical significance.”20 Their article, “The Dawn of Darwinian Medicine,” carefully qualifies this premise, as a “disciplined application of what has been called the adaptationist program.” Predicting, as Williams had previously argued, entails “predicting the existence of phenomena otherwise unsuspected” through carefully conceived experiments. Biologists have been busy publishing predictions. A “table” that takes up a full page of “Dawn” lists twenty-six articles published in Evolution during the year 1988. Of those, twenty are marked as establishing confirmation of prediction, only five refute their

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prediction, and one has mixed results. The point is that “the adaptionist program has been enormously fruitful in the fields of ecology and animal behavior, and in the study of life cycles.”21 Why not in medicine?

Lists and Audiences Why We Get Sick begins with a list of six major causes of disease or disorder, and this list deconstructs the ideal human body. Most causes are rooted in incomplete or unsatisfactory adaptation of human bodies, either within themselves or with respect to their environments. Some causes relate to the hostile environments that humans have faced in the past and face in the present. These causes of disease include (1) defenses, like coughs, that work or sometimes do not work; (2) infection, as an arms race between parasites and bodies; (3) novel or modern environments that make life difficult for bodies adapted to the world of “Stone Age” humans; (4) genes, and genetic quirks; (5) design compromises; and (6) evolutionary legacies. “The Dawn of Darwinian Medicine” lists in its abstract only four causes of “medical disorders”: infection, injuries and toxins, genetic factors, and abnormal environments. In 2008, Nesse and Stearns recite a similar list of categories in an attempt to explain why natural selection has left the body vulnerable to disease: “mismatch with the modern environment, pathogens coevolving with hosts, constraints on what selection can do, unavoidable trade-offs, reproduction at the expense of health, and defenses such as pain and fever that are useful despite causing suffering and complications.”22 These categories overlap in complex ways and one might dismiss the classification system on the basis of the incompleteness or incomplete separation of these classes. But the exposition of Why We Get Sick is after all an early exploration. It is structured thematically, moving from infectious disease and injury to genetic problems including aging, then to allergy, cancer, and sexual reproduction, and finally to mental problems. In other words, it ascends a kind of ladder from safe ground to less certain and much more sensitive ground, particularly when it calls upon evolutionary psychology in its penultimate chapter.

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Speaking of Infection Infection is an “endlessly escalating arms race,” as Paul Ewald argues, and Nesse and Williams’s chapter on infection is named “An Arms Race Without End.” They use a “functional classification of infectious diseases” that Ewald recommends.23 One can classify the observable acts or symptoms of hosts and pathogens using this system. Like any other arms race, infection can be spoken of in terms of guarding and attacking; engagement with a pathogen is immensely wasteful and very costly, especially for the host. Modes of defense include hygiene, the skin itself, pain and malaise that can lead to escape and avoidance, expulsion, and of course immunological response. Meanwhile, pathogens secure resources by stealth, by limiting their own damage, by evasion of defenses, and so on. According to Van Valen’s “Red Queen Principle,” “predator and prey coevolve in an escalating cycle of complexity.”24 So too with hosts and parasites. “We are in a relentless all-out struggle with our pathogens, and no agreeable accommodation can ever be reached.”25 Nesse and Williams’s book frames the discovery of antibiotics as a great medical advance that has led to a golden age, but at the same time predicts that these benefits may be short-lived because bacteria continue to evolve defenses against antibiotics. This is now a familiar argument. “The list of threats we face from antibiotic-resistant bacteria is long and frightening.” The evolution of virulence is “widely misunderstood,” and an “adequate theory” of it is needed. Parasites use the defenses of hosts, evolve rapidly, compete with each other within hosts, and often become more virulent when they have opportunities for transmission. They spread by personal contact, by what Ewald has called “cultural vectors,” in water, and in hospitals.26 The human immune system operates according to tradeoffs of costs and benefits. Nevertheless it can miscalculate, make mistakes, fail to respond when it should, or over-respond. Pathogens may escalate their own deceptions in meeting this antagonist and escape attack from the immune system. Pathogens can become much more virulent in novel environments, leading to epidemics. This is the language of immunological arms race.

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Contradictions of Bodies But that, perhaps, is to get ahead of the story. In order to alter or decenter medicine’s proximate perspective, Nesse and Williams must first deconstruct the traditional ideal of the human body. They do this most dramatically in the language of an article they published in Scientific American in 1998. The language there reveals, not the genesis of Nesse and Williams’s argument, but a clear shape in which it is couched for a popular medium. It begins with the traditional Williams paradox, but expressed in a more cheerful tone: “Thoughtful contemplation of the human body elicits awe—in equal measure with perplexity.” Using Williams’s favorite example, the eye, the argument proceeds to its turn: “Admiration of such apparent perfection soon gives way, however, to consternation.” In other words: The body is a bundle of such jarring contradictions. For each exquisite heart valve, we have a wisdom tooth. Strands of DNA direct the development of the 10 trillion cells that make up a human adult but then permit his or her steady deterioration and eventual death. Our immune system can identify and destroy a million kinds of foreign matter, yet many bacteria can still kill us. These contradictions make it appear as if the body was designed by a team of superb engineers with occasional interventions by Rube Goldberg.27

In Why We Get Sick, this “we perspective” is also striking. We choke; our arteries get clogged; we become nearsighted. We have allergies. We get autoimmune diseases. “Dozens of other bodily designs seem equally inept. Each may be considered a medical mystery.” A language of awe and admiration is undercut by a language of contradiction and consternation. This language is partially resolved by an argument that the design of the human body is predictably flawed, partly because many of its functions are bundled, rather than integrated, and “the body is a bundle of careful compromises.”28 It is hard to know what design or careful mean in this context. Can Darwinian medicine redesign the human body? No. Can it overcome flaws in design? Maybe, in some cases, but because human wellbeing—mental and physical—has been designed as a result of certain tradeoffs and historical legacies, some poor designs are irreversible. Can one “game the system” created by natural selection—by which I mean use the rules and procedures that protect that system in order, instead, to manipulate it for a desired outcome? A more successful strategy might be

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to redesign human behavior so that it takes cognizance of human weaknesses, flaws, and problems. But, as the recent pandemic of the years 2020–22 has shown, human behavior is most difficult to redesign. As a kind of counterproposal or even antithesis to the idea of maladaptation, Nesse posits a mechanism he calls the “smoke-detector principle.” We all know about the problems of the ones installed on the walls of our homes. “Yet the annoying false alarms, and the costs of the smoke detector and its occasional battery change, are well justified by the protection they provide against a major fire.”29 Nesse applies this principle not only to the function of allergies, but also to the efficacy of anxiety and other psychic maladies. “The defenses appear overresponsive because they are ‘inexpensive’ compared to the harms they protect against and because errors of too little defense are more costly than errors of too much defense.”30 This is Nesse’s argument, but it is based on Williams-style methodical calculations of costs and benefits.

Order and Disorder, Causes and Cures, Means and Motives Why We Get Sick begins with a set of disclaimers. The authors ask “people to think about their illnesses in a different way, to ask questions of their doctors, perhaps even argue with them, but certainly not to ignore their instructions.” This is, then, an intervention that seeks to alter the dynamic of the professional relationship between doctors and patients. They claim that their book “does not arise from a disapproval of current medical research or practice in Western industrialized nations,” and they insist that they “are urging not an alternative to modern medical practice but rather an additional perspective.”31 It also begins with a refutation. More than a few prominent evolutionary biologists, including Ronald A. Fisher and William Hamilton, espoused eugenic doctrines. They believed that it was possible to breed better humans and improve the human gene pool—or at least to forestall what they imagined might be the deterioration of the human genome as a result of modern medicine—by sterilizing and euthanizing “unfit” individuals.32 William Hamilton is quite explicit in the second volume of his collected papers. As he states, “I predict that in two generations the damage being done to the human genome by the ante- and post-natal life-saving efforts of modern medicine will be obvious to all.”33

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Nesse and Williams’s de-idealized perspective toward the human body does not mean they advocate such a project to improve its genetic makeup. They are careful to distance themselves from views like those of Hamilton. They “try to forestall several potentially dangerous misunderstandings.” As they put it in a section titled “What We Are Not Saying,” First of all, our enterprise has nothing to do with eugenics or Social Darwinism. We are not interested here in whether the human gene pool is getting better or worse, and we are emphatically not advocating actions to improve the species. We are not even particularly interested in most genetic differences between people, but much more in the genetic material that we all have in common.34

This point should be stressed. They demonstrate medical value for an evolutionary understanding of problems humans have in common. As they visualize these problems they see in individuals, they also recognize their prevalence in the human population. They sometimes exhibit a sense of humor. Huck Finn’s Pap appears in the introduction to chapter 5, when he trips over a tub of salt pork, barks his shins, and then kicks the tub to the detriment of his toes. We can laugh at him, but we are all falling over that same tub of salt and fat. “Today most of us can afford to eat more fat, sugar, and salt than is biologically adaptive.” What’s more, as a behavioral legacy from our nutritionally challenged ancestors, we are likely to eat all we can get. But the situation is still more complicated. Given a nutritionally rich environment, humans undergo faster growth that results in larger bodies that were rare in the past. “The blood-pressure-regulating mechanism, pushed to adjust the system outside the range for which it was designed, often overshoots, causing high blood pressure.”35 Nesse and Williams also caution: “A question about function can have more than one right answer. For instance, the tongue is important both for chewing and for speech, the eyebrows both for keeping the sweat out of the eyes and for communication. Second, the evolutionary history of a species or a disease is like any other kind of history.” There are no experiments that can reach into the past. “History can be investigated only by examining the records it has left.”36 In a desire to produce a lively text, the book uses a wide variety of rhetorical tools. For instance, at the beginning of chapter 7, an anonymous professor of medicine—a character, one might say, loosely based on Williams—tells his students, “In fact, our bodies don’t work very well …

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we are bundles of genetic flaws. The body is a fragile, jury-rigged device. Our job as physicians is to fix Mother Nature’s oversights.” The job is not to change the body, but repair or at least reduce the effects of these historical oversights. Repair is possible precisely because evolution builds bodies by bundling functions. Contradictory and loosely integrated functions can be separated into units or modules. Nerves themselves gather into bundles. Bodily responses can be perceived as bundled into a kind of psychic “emergency kit.” “Like all products of organic evolution, [the human body] is a bundle of compromises, each of which offers an advantage, but often at the price of susceptibility to disease.”37 Maladies come from many sources, such as rickets from vitamin D deficiency, but what of cabin fever, jet lag, or other problems caused by novel and modern environments? Sometimes maladies come from “maladaptive human design.” “It has seemed a bit odd until now even to ask how something maladaptive like disease might have been shaped by natural selection.” For example, a design where the tube that carries food to the stomach crosses the tube that carries air to the lungs means that “the human capacity for choking represents an ancient maladaptive legacy aggravated by a much later compromise.” Yet all bodily responses are not maladaptive. And whether there is a “prevalence of maladaptive human design features” is at issue. “Cancer is a collective term for maladaptive and uncontrolled tissue growths of all kinds.” Cancer is and is not of us. More to the point, “many design features, while not maladaptive, are functionally arbitrary and explicable only as historical legacies.”38 A body has no orderly state; it is a result of a set of historical events. Under these circumstances, to speak of disorder in an imperfect design makes little sense. However, “maladaptive extremes of anxiety, sadness, and other emotions make more sense when we understand their evolutionary origins and normal, adaptive functions.” A great deal more of the human can be understood as adaptations than by any other means; even “our emotions are adaptations shaped by natural selection.”39 But our flawed design includes a disordered mind. “Are Mental Disorders Diseases?” asks the last discursive chapter, which Nesse and Williams also contributed as the lead for Simon Baron-Cohen’s anthology, The Maladapted Mind (1997).40 The answer is yes and no. The language of bundling is striking in this argument and can be taken as part of the language of evolutionary psychology, especially that of Leda Cosmides of the University of California,

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Santa Barbara, which compares the structure of the mind to a Swiss army knife made of separable and independent modules.41

Author and Authority: Doctors and Patients With all coauthored texts, it is nearly impossible to disentangle the authors and determine who contributed which language, or which writer is the true authoring agent or authority. The content and ideas of their various coauthored articles on Darwinian medicine include, and are based on, various arguments made by George C. Williams. Yet by and large, they do not exhibit or exemplify his language, his emphasis, or the tentativeness of agenda that is typical of Williams. In that sense, they are filtered through the more confident language of Randolph Nesse, who is himself highly articulate but writes from a different background. Being a psychiatrist, he often develops his own favorite themes. Presumably Nesse had more autonomy in the sections of the book dealing with psychiatric questions. Further, there is the matter of audience. “The Dawn of Darwinian Medicine” (1991), their first in what became an ongoing series of coauthored publications, led later to “Evolutionary Biology in the Medical Curriculum: What Every Physician Should Know” (1997), which was clearly directed toward physicians and the medical community.42 Why We Get Sick (1995) and “Evolution and the Origins of Disease” (1998) address a more general audience. The two coauthors have to define the roles of at least two kinds of readers of their book, the doctors and the patients, and intervene between them. After all, to be a patient, in more than an etymological sense, is to suffer, and to be a doctor is to teach. The use of the term patient predates the use of doctor for a physician or healer; Nesse and Williams use the latter terms, physician and healer, interchangeably. As Michael F.  Antolin explains, reaching out to medical professionals from evolutionary studies calls for crossing a longstanding cultural divide: “it is perhaps surprising how little the biomedical field explicitly uses the language of evolutionary science, even in areas like the evolution of antibiotic resistance in microbes.”43 More proximately pertinent might be the questionable efficacy of lecturing to doctors on human nature or the human condition. This longstanding cultural divide manifests itself in practical questions about the curriculum followed by doctors, the culture in which they practice, and the tasks on which they focus attention. Language reveals this. The work of doctors is almost guaranteed to be focused on proximate

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causes, if for no other reason than that their patients come to see them only when they are already diseased or perceive a malady in themselves. Consequently, the title’s rhetorical emphasis on the “We” of sickness may be a weak gesture. Consider dialogue at a medical clinic. Nesse and Williams offer the reader a chance to “listen in” as a pre-Darwinian and a post-Darwinian physician talk to various patients. The first patient has a very painful toe, and asks his pre-Darwinian doctor about his gout, saying, “So why isn’t the body designed better? You would think there would be some system to keep uric acid levels lower.” The doctor answers, “Well, we can’t expect the body to be perfect, now, can we?” This pre-Darwinian physician is portrayed as giving up on science, dodging the patient’s questions, and “implying that such ‘why’ questions need not be taken seriously.”44 Such doctors probably do not recognize the distinction between proximate and evolutionary explanations, nor do they recognize the importance and legitimacy of evolutionary explanations for disease. A post-Darwinian physician gives an answer “closer to what the patient wanted and was entitled to.” Not only does he explain that “human uric acid levels are much higher than those of other primates,” and higher uric acid levels correlate with a longer lifespan, but he says: “It seems that uric acid protects our cells against damage from oxidation, one of the causes of aging. So natural selection probably selected for higher uric acid levels in our ancestors, even though some people end up getting gout, because those higher levels are especially useful in a species that lives as long as we do.”45 Then the doctor proceeds to traditional therapies for lowering uric acid levels. With a patient who has strep throat, the post-Darwinian physician offers a lecture on the coevolution of streptococcus with humans. What’s worse, he tells the patient, “when we make antibodies that attack the strep bacteria, those antibodies are prone to attack our own tissues as well. We are in a contest with the strep organism, but we can’t win because the strep evolves much faster than we do.” So, with an encouraging word he explains, “thank goodness we can still kill it with antibiotics, although this may be a temporary blessing.” Then he lectures about taking all your pills as a responsible citizen—“do yourself and the rest of the world a favor”— because “those antibiotic-resistant organisms make life difficult for us all.”46 A patient with a heart problem learns that his genes make him prone to high cholesterol. “Well, those genes aren’t harmful in the normal

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environment we evolved in. If you spent six or eight hours walking around each day to find food, and if most of your food was complex starches and very lean meat from wild game, you wouldn’t get heart disease.” When the patient wants to know why he craves foods that he shouldn’t eat, he is told, “I’m afraid we were wired to seek out certain things that were essential in small amounts but scarce on the African savannah.” The post-­ Darwinian physician continues, “You are right, it is a kind of a cruel joke—we do want exactly those things that are bad for us. Eating a healthy diet does not come naturally in the modern environment. We have to use our brains and our willpower to compensate for our primitive urges.”47 How useful these miniature Darwinian explanations or sermons might be to a patient’s physical and mental state is debatable.

Maladapted Mind: The Stone Age and Evolutionary Psychology Early in his career, Williams writes, “Despite the arguments that have been advanced (e.g., Dobzhansky and Montague, 1947; Singer, 1962), I cannot readily accept the idea that advanced mental capabilities have ever been directly favored by selection. There is no reason for believing that a genius has ever been likely to leave more children.” But human brains did grow, and Williams wonders whether “selection for acquiring verbal facility as early as possible might have produced, as an allometric effect on cerebral development, populations in which an occasional Leonardo might arise.” He notices that “this interpretation is supported by the apparent diversity of adult human mentality and by the close similarity of the sexes in intellectual endowment.” Yet he is also capable of saying that “a character selected for a specific adult-male function, such as the political leadership of a primitive society, would reach a high development in adult males only, and would be well standardized in this group.” This leads him to a phrase that evolutionary psychologists have been fond of quoting: “Is it not reasonable to anticipate that our understanding of the human mind would be aided greatly by knowing the purpose for which it was designed?”48 As I have noted, this piece of discourse becomes a rallying cry for evolutionary psychology. More than a few critics of evolutionary psychology have noticed three of its premises: (1) human beings are preprogrammed with specific knowledge, not born as blank slates; (2) most human behaviors and social

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structures have been selected for the ultimate cause of reproduction, the spread of genes; (3) many if not most personal, medical, and social problems come about because humans became adapted to a Stone Age environment, but must live in a modern world. These add up to suggest that human nature is fixed, static, and perhaps timeless. “Could it be a trait that was adaptive in the Stone Age but that now causes disease?” Nesse and Williams often ask in Why We Get Sick, and the answer is predominantly yes. Indeed, the text arrays assorted facts about “Stone Age” humans, including their longevity, mortality rate, reproductive behavior, the fat content of their diet, and their social and natural environment. The expression “Stone Age” appears over forty times in the text, signaling an acceptance of a single, completed, unchanged human nature that has remained unaltered by natural selection since then. As the coauthors put it, “we are specifically adapted to Stone Age conditions,” and our “bodies were designed for the world of the Stone Age hunter-­ gatherer.”49 There are indeed a number of sections in chapters nine and ten that detail conditions during the so-called Stone Age. The Stone Age premise includes what evolutionary psychologists sometimes name the “environment of evolutionary adaptedness” (EEA), a concept foundational for the discussions of behavioral disorders in Why We Get Sick. This raises the question of whether, strictly speaking, an evolutionary psychologist’s version might well have been better entitled Why We Are Sick. Evolutionary psychology of the 1990s offered a remarkably static view of human evolution, with a list of modules or instincts that came about “back then” and have changed little since. A eugenicist like Bill Hamilton felt that human evolution occurred faster than that, which is why he and his most important influence, Fisher, were worried. What troubles me most is this. Whatever allows Williams to introduce history as the third doctrinal base for successful biological research in 1985 and 1992 seems paradoxically at odds with Why We Get Sick’s way of thinking, which lacks sufficient historical dynamic. Evolutionary psychology argues, in the words of Cosmides and Tooby, that “our modern skulls house a stone age mind.”50 But as David J. Buller has argued, history cannot occur only “back then,” and the adaptation of mind is a continuing process.51 When one asks why about human nature, one enters that process, though constraints certainly channel biological possibilities. It is not my purpose to quote Williams against himself, but one does need to remember that adaptation is a “a special and onerous concept that should

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not be used unnecessarily.” The expression “Stone Age,” as used in Why We Get Sick, leads to vagueness, not precision.

Depression and the Dawkins Effect Therefore, I am skeptical about the possibility of pursuing a discussion of mental disorders with “disciplined adaptationism.” Such discussions call for scrutiny on several grounds. Part of physicians’ attention to proximate causes is dictated by a pragmatic need to alleviate suffering in a humane manner. It does no good to say that depression may ultimately have been an adaptation, as Randolph Nesse writes in several articles beginning in the 1990s and continuing into the twenty-first century. As he said to the New York Times, he sees depression potentially “as a useful response to situations in which a desired goal is unattainable,” or, as he has put it, “when one of life’s paths peters out into the woods.”52 As a psychiatrist interested in treating depression, Nesse focuses on the degree to which depression is shaped by unreachable economic, romantic, or professional goals that must be seen objectively and abandoned; as Nesse says, humans must often loosen themselves from goals gone sour, even long sought goals.53 But it will not do to tell a depressed patient that “if I had to put my position in a nutshell … I’d say that mood exists to regulate investment strategies, so that we spend more time on things that work, and less time on things that don’t.”54 Such statements constitute a form of blaming the patient. Though Nesse and Williams cannot plumb the origins of modern striving for goals, evolutionary explanations can certainly document their existence: “Watching these successful people on television arouses envy. Envy probably was useful to motivate our ancestors to strive for what others could obtain.” The book has its own perhaps unobtainable goal—a desire to change public policy—which the authors defend on the premise that “facts can, however, help us to achieve whatever goals we decide to strive for.” The language of human goals—as in the goal of medicine—and the idea that natural selection lacks goals are put into tension by the rhetorical plan of Why We Get Sick. One can see this in the description of an adaptation that putatively causes depression. “Natural selection involves no plan, no goal, and no direction,” as we know, but it also may prohibit most shared goals. In the case of pregnancy, what “would seem to be the ultimate in shared goals—a refuge from conflict, perfect unity of purpose between mother and fetus,” reveals something very different to

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researchers—a conflict between mother and offspring from the onset of pregnancy onwards.55 For all that, progress frames this book’s blithe tone, whose final chapter 15 is optimistically entitled “The Evolution of Medicine,” implying, precisely, progress toward a goal. The preface promises the “potential contribution of evolutionary biology to medical progress” and the discovery of “new ways in which Darwinism can aid the progress of medicine.”56 With regard to cancer, for instance, the book assures the reader of rapid progress, with more progress to come. The notion that human medical knowledge progresses by some evolutionary process is somewhat surprising, given Williams’s view that evolution by natural selection proceeds by attrition, sheer waste, and many deaths. Williams always attempts to speak for consensus—intelligent consensus—but this becomes more difficult as evolutionary studies proliferate. The title of the prospectus, “Dawn of Darwinian Medicine,” suggests an interesting aspect of this proliferation. Dawn itself is a code of four letters and can be expanded to Darwin by adding one consonant and one vowel. The code can be transformed again, with Darwin becoming a code for evolution, as the book’s index suggests: “Darwinian medicine. See evolutionary medicine.”57 A typical Williams title does not reach out for such codes to establish authority. While Williams or Nesse counts the articles in one volume of Evolution in order to establish a kind of consensus, somehow in Why We Get Sick this use of consensus is transformed into something more promotional, into a kind of “universal Darwinism”—a term probably coined by Richard Dawkins in 1983 and expanded into a theory broad enough to be applicable to human cultural change.58 I doubt that Williams ever believed in a “universal Darwinism,” or that one could speak of change in social institutions like medicine as “evolutionary.” The best part of Williams’s thinking has always been rooted in a desire to open discussion, not close it. Opening conversation has never been the strategy of Richard Dawkins, who often attempts to close discussion by winning it. Consider Dawkins’s recent plight. He could not entitle a book “the greatest story ever told,” so he titled it The Greatest Show on Earth, thus suggesting a circus, and possibly P.  T. Barnum’s quip that there is a sucker born every minute. There is a subtle but grave difference between writing a book to clarify thinking and writing a book to promote ideas—or to promote yourself. Perhaps this distinction is more difficult to see now in an age of blogs. However, the challenges Williams offers are meant to open the way, as I

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have argued, to start discussion rather than give definite answers. It is hard to know whether Why We Get Sick succeeds in opening discussion, though it is certainly a critique of modern medical practice’s single focus on the proximate reasons for maladies.

Clinching an Argument: “Many” and “May” Constructions Any success for the kind of argument Nesse and Williams make about proximate and ultimate causes of disease requires a clear link to pragmatic applications for Darwinian explanations. As they argue consistently, “many aspects of the body seem amazingly crude,” “many of us” develop problems, and “evolutionary ideas might well have significance for many medical problems.” They offer “many examples,” produced by “many scientists,” that may be of use to “many medical professionals.” There are also “many misconceptions” about fitness.59 The adjective many indicates a large but indefinite number, and when used too liberally, produces vagueness. Granted, the book is meant to be a “preliminary guide” and a “challenge.” As indicated, many physicians have asked evolutionary questions “apologetically, treating their own ideas not as serious hypotheses but as mere speculations undeserving of serious inquiry.” Possibilities for success in transforming these to “serious inquiries” are undercut by the particular way in which Why We Get Sick discusses evolutionary explanations. I will call the book’s predominant grammatical construction the “may construction,” where the auxiliary verb is used primarily to express possibility. It begins with certain kinds of examples. For instance, given certain “inept” bodily designs, “each may be considered a medical mystery.” Or, “since disease is often assumed to be necessarily abnormal, the study of its evolution may seem preposterous.”60 The argument asserts, in general, that “evolutionary hypotheses … predict what to expect in proximate mechanisms.” Spelled out, prediction is often expressed by weak stylistic construction like this: “if we hypothesize that the low iron levels associated with infection are not a cause of the infection but a part of the body’s defenses, we can predict that giving a patient iron may worsen the infection—as indeed it can.” What may appear in the patient’s symptoms is repeatedly discussed using “may constructions” that slip into evolutionary explanations, as in “individuals may be

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viewed as vessels created by genes for the replication of genes.” Certain genes may give benefits early in life. Consequently “by suggesting the functional significance of some known aspect of human biology, you may logically be able to predict some other, unknown aspects,” and “if they are there, they may be of medical significance.” If fever is a defense mechanism, “using drugs to suppress fever may sometimes make people sicker.” But “a given expression of a defense need not be adaptive, and that even when it is, it may not be essential.”61 Nevertheless, when the “may construction” is used to posit various evolutionary explanations for human behaviors, the very repetition over time undercuts itself, especially when doubled, as in a discussion of avoiding dangers by practicing hygiene: “Our tendencies to avoid contact with people who may be infectious may have the same significance.” So too, “our tendency to defecate away from others may prevent the infection of close associates, and social pressures to conform to such practices may protect us from infection by others.” Sometimes the “double may” construction is implied, as in: “The frequency of babies ‘spitting up’ may be another instance in which the baby manipulates the mother, in this case to make more milk than is in her interests.”62 Symptoms may be misinterpreted, of course: “Some sneezing may possibly be an adaptation that viruses use to disperse themselves.” “Diarrhea may represent a defense mechanism.” All interventions have costs and benefits. In many cases, Nesse and Williams’s evolutionary view mitigates against excessive intervention. But what about adding antioxidants to the diet? Taking vitamin C, vitamin E, or beta-carotene is not without some risk, but “adding extra antioxidants to the diet may have benefits that exceed the costs.” Streptococcus can be killed with antibiotics, “although this may be a temporary blessing.”63 This signals that many explanations are essentially speculative—as in “looking for the functions of sleep may well provide the knowledge we need to adapt it better to our present needs.” There are clues to suggest that “plantar fasciitis may result from environmental novelty.” “Menopause may protect the interests of our genes in existing children.” Sometimes these speculations are clichés: “At present we have reason to believe that many problems adults have in getting along with other people may have their origins in problems with the first attachment.” Sometimes they seem contrived, as in the idea that “low mood may be a variant of a hibernation response in some remote ancestor.” Some suggestions seem wildly speculative: for instance, “studies suggest that the serotonin system may

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function, in part, to mediate status hierarchies and that some low mood may be a normal part of status competitions.” It is hard not to laugh when one reads, “Some have suggested that the tendency to suspiciousness itself may compensate somewhat for the disadvantages of schizophrenia.”64 As Nesse told me in conversation, he and Williams went through the manuscript inserting the “may construction” because they did not wish, in many cases, to claim absolute causality or make unqualified diagnoses. One understands the need for due caution and fair-mindedness that leads to the “may” construction, but when it appears literally hundreds of times, it becomes an empty mannerism. After all, part of the question is whether this argument will convince doctors. As the authors admit, “medical scientists may also hesitate to consider functional hypotheses because of their indoctrination in the experimental method.”65 An excess of speculation is not likely to encourage such scientists. Williams uses a “may” construction frequently in Adaptation and Natural Selection, but in entirely different ways and to different effects, primarily to express contingency. For instance, he writes, “Sometimes the important effect of an adaptation from man’s point of view may not be its function.” He uses the construction to express skepticism, point out the limitations of certain ways of thinking, or suggest an abstract possibility rather than to expand on speculative ideas. He sometimes uses the construction to indicate a purely hypothetical dimension in his own thinking, as in “I am inclined to concede that mammalian tissues may be physiologically somewhat more specialized than those of a fish.” In a typical example, he uses it to underscore his tentativeness: “I will present some points of view that may facilitate the search for connections between the natural selection of alternative alleles and the phenomena of ecology and morphogenesis.” Or, “The distinction between somatic and genetic may be arbitrary at times.” Or, “We may be familiar enough with the physiology and ecology of the organism to state an opinion on how fit it is, relative to other organisms in the same population.” There is often a certain wit to his use, as in “necessity may have been the mother of invention, but she was not the inventor.”66 Sometimes Williams alternates between may and might as a way of developing a scenario: “Somewhere in man’s ancestry there may have been a protoamphibian that occasionally came out on land and pushed itself along with its fin lobes. Such animals might have responded by a minor thickening of the part of the skin that came in contact with the ground, just as our skins thicken wherever they are subjected to friction.”

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Or he uses the may and but construction: “Sex may act to increase population size and it may act to confer evolutionary plasticity, but these are effects, not functions.”67 In Why We Get Sick, the construction is used less to posit abstractions or ways of thinking and more to accumulate a set of speculations and suggest, by that accumulation, a possible set of actual diagnoses for specific maladies. To be charitable, this follows what is apocryphally known as the “Pauling principle,” named after Nobel Prize-winning chemist and peacemaker Linus Pauling: the way to get good ideas is to produce many ideas and throw the bad ones away. So one must be willing to generate questionable ideas in order to generate many ideas. This dynamic has some parallels in evolutionary theory, and might even be a theory of progress. The problem is that Williams’s collaboration with Nesse entails a capitulation—a serious surrender and breach of some important fundamentals of his literary style, most of all of critique. We can speculate that Williams may have been advocating the “may construction” out of fidelity to scientific caution—a fidelity that this book seems to have stretched to or past its breaking point.

Ambitious Projects and Implacable Realities Because Williams wrote primarily of evolutionary processes and consequent products, his reasoning must apply to humans as well as other organisms. Though reason must apply to brains as organs, the ability of reason to apply itself to the behaviors of a brain is limited. He himself indicated that limitation. As he quips, “it is undoubtedly due to something in the neural and psychic activities of its brain that a spider monkey spends most of its time in the trees.”68 At the beginning of chapter 13 of Why We Get Sick, one finds the following claim: “While our main focus in this book is on how evolutionary ideas can help to explain and prevent or cure specific medical diseases, here and in the next chapter we broaden our view somewhat to encompass emotional and behavioral problems that may or may not be considered medical disorders.” Nesse and Williams make a distinction between “problems associated with reproduction, such as diabetes during pregnancy or sudden infant death syndrome,” which are clearly diseases, and others, such as jealousy, child abuse, and sexual problems, [that] involve behavior and emotions. However we categorize them, they cause much

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s­uffering and make more sense in the light of evolution. Help from Darwinism does not end at the boundary between the medical and the social or educational. Darwinism is relevant to all aspects of human life, not just medicine.69

It is hard not to imagine this kind of statement as overreaching. Fortunately, most of their book is not devoted to adaptationist explanations of mental dysfunctions. But the bottom line is whether Darwinian medicine can provide ultimate causal analysis as a diagnostic tool, as a guide to medical practice, as a satisfactory explanation for patients, and as an improvement in human life, as promised.70 There are limits. The poles of Williams’s themes include the inexorable and ultimate linkages between reproduction and death, from which there is no escape. As Nesse has noted, these links are created by adaptation and maladaptation. When the poles of Williams’s themes also set clear thinking against illusion about the process of natural selection, they especially underline the limitations on the success of natural selection’s processes and on that of any project to repair or remedy them. To expand slightly, Williams sees a necessary and dynamic tension between benefits and costs, between optimization and constraint, between youth and aging, between reproduction and incapacity. None of these linkages are escapable because they are parts of a process. He observes the proliferation of various kinds of illusions, perhaps inevitable illusions, that humans can outwit disease or death. The source of such illusions or errors may be religious or aesthetic. Insofar as the discussion of Why We Get Sick remains within the constraints of the Williams framework, it is successful; when it reaches beyond, it risks becoming a parody of itself, or worse. Something more fundamentally problematic occurs when Williams abandons painstakingly built principles and methods. If the “onerousness” of adaptation as a concept is replaced by easy loose speculation, the ground that Williams built throughout his whole body of work, via critique, is undercut. The Williams voice, as William Hamilton found, is always associated with a tightly reasoned argument about a limited theme; it maintains “an almost ruthless objectivity” and reveals a “characteristic knack for seeing a problem from a totally fresh point of view.”71 But imagine what it might be like to be an author of a book that lacks one’s own voice. Finding Williams in Why We Get Sick is a somewhat hit-or-miss proposition.

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The Human Condition Once again, Williams returns to Paley’s eye in Why We Get Sick, and the anecdote undergoes several strange metamorphoses. Whether they are felicitous remains to be seen. The anecdote of the watch is transformed from first to second person and the watch is transformed into an image representing the aging human body. Once more then, “you are crossing a heath,” a heath being, etymologically, untilled, uncultivated land or even wasteland. But this time you find a reminder of your own body. The watch, after all, is a traditional gift at retirement. This watch is run down, in both senses. “So perfect in many respects, in others makeshift at best. How could the creator of such a masterpiece have allowed so many obvious flaws?” So Paley’s watch slips into another literary trope. It is indeed inscribed, like those given upon the occasion of retirement, though the text is visible only with a magnifying glass: “Model 3,500,001,859 is designed to give accurate time. It will self-repair most problems, except for those arising from manufacturing defects, novel environments, or design compromises. In the ideal environment, its mean life span is 85  years. Care for it well, and enjoy it while you can. Lifetime Watch Co.”72 Any human body is itself a mechanism, like this decaying watch that once seemed perfect. Now its flaws are obvious, its chain broken, its spring rusted, its crystal “laced with slight distortions.” So too with our bodies. Despite the “exquisite design,” we find “crude flaws”; despite “multiple defenses,” we suffer a “thousand vulnerabilities”; despite “capacities for repairs, our bodies inevitably deteriorate.” But physicians need not simply wonder at the incongruity of it all, as they did before Darwin.73 Heed it well, physicians! As an expression, the “human condition” falls somewhere between human nature and human action, yet is contained by neither. Clearly, social and environmental conditions play a role. The expression is used by a wide variety of writers from a seemingly disparate set of perspectives, and remarkably few define its domain. What people mean by human nature includes what we are and how we deal with it. From an evolutionary perspective, the human condition, though constrained by the past, cannot be static. It is part of a process; that process is disorderly, and it includes decay and dissolution. The human condition is disorderly and limited. The word “condition” is itself rooted in social state or status and comes from a Latin expression suggesting conversation. The term has evolved to

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mean situation or mode of being. Many schemes have been invented to improve the human condition. One scheme focuses on medicalizing it, particularly through psychiatry. As one critic argues, “To medicalize the human condition is to apply a diagnostic label to various unpleasant or undesirable feelings or behaviors that are not distinctly abnormal but that fall within a gray area not readily distinguishable from the range of experiences that are often inescapable aspects of the fate of being human.”74 People are predisposed to think of the human condition as general and static, powerful though flawed, unique yet predictable, and inescapable. Near the end of Consilience: The Unity of Knowledge, in which E. O. Wilson argues that evolutionary science can uncover the essential structure of the human condition, “proceeding from the deep history of genetic evolution to modern culture,” he predicts that “science for its part will test relentlessly every assumption about the human condition.”75 In this sense, Why We Get Sick is indeed a text that focuses on but cannot test the human condition. E. O. Wilson has spoken of the human condition as a singularity, because of its unlikeliness. “Homo sapiens is the only species of large mammal—thus large enough to evolve a human-sized brain—to have made every one of the required lucky turns in the evolutionary maze.” This seems somewhat silly and leads commentators to remember that questions about the universality of the human condition inhabit a gray borderland between philosophy and empirical science.76 Arguably, Williams has been going toward this question since 1957, when he constructed his argument about “Pleiotropy, Natural Selection, and the Evolution of Senescence.” He does not consider the human condition a singularity, so his approach to this sociobiological question is more moderate and much less imperialistic than that of E.  O. Wilson. What Williams thought and wrote of the design of humans, where his own voice is more clearly discernable, appears in “The Tithonus Error,” as we shall see.

Notes 1. Nesse’s reminiscences are to be found on “The Skeptical Adaptationist,” Randolph Nesse’s no-longer-live blog about the challenge of applying evolutionary principles in medicine and elsewhere,” http://skepticaladaptationist.com/2010/09/george-­williams-­1926-­2010.html. Also, correspondence with the author, April 1, 2012.

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2. William Shakespeare, The Tragedy of Hamlet, Prince of Denmark, Act II, Scene ii, lines 293–97. 3. Raymond Williams, The Long Revolution (London: Chatto and Windus, 1961), 57, 98, 352. 4. Hannah Arendt, The Human Condition (Chicago: University of Chicago Press, 1958). 5. Randolph M. Nesse and George C. Williams, Why We Get Sick: The New Science of Darwinian Medicine (New York: Times Books, 1995), 106. Also published in Britain as Evolution and Healing: The New Science of Darwinian Medicine (London: Weidenfeld and Nicolson, 1994). I use the American edition. 6. Nesse and Williams, Why We Get Sick, back cover. 7. Randolph Nesse, “George Williams, 1926–2010,” post on personal blog, “The Skeptical Adaptationist” (September 2010). 8. Paul Ewald, “Evolutionary Biology and the Treatment of Signs and Symptoms of Infectious Disease,” Journal of Theoretical Biology 86 (1980): 169–76. 9. Nesse and Williams, Why We Get Sick, ix, 32. 10. Joe Alcock, “Emergence of Evolutionary Medicine: Publication Trends from 1991–2010,” Journal of Evolutionary Medicine 1 (December 2012): 1–12, Article ID 235572, http://www.ashdin.com/journals/published. aspx?jid=jem. 11. Nesse and Williams, Why We Get Sick, 3, 27. 12. Alcock, “Emergence,” 9. 13. Randolph M. Nesse, “Tinbergen’s Four Questions Organized.” This onepage scheme can be downloaded as “Biology’s Four Questions” at http:// www-­personal.umich.edu/~nesse/Nesse-­Tinbergen4Q.PDF. 14. Nesse credits the following sources for this distinction: Niko Tinbergen, “On the Aims and Methods of Ethology,” Zeitschrift fur Tierpsychologie 20 (1963): 410–63; Ernst Mayr, The Growth of Biological Thought: Diversity, Evolution, and Inheritance (Cambridge, MA: The Belknap Press of Harvard University Press, 1982). 15. Nesse and Williams, Why We Get Sick, 7. 16. George C.  Williams and Randolph M.  Nesse, “The Dawn of Darwinian Medicine,” Quarterly Review of Biology 66 (1991): 1–22. 17. Alcock, “Emergence,” 4. 18. Conversation with Nesse, January 25, 2021. 19. https://www.edge.org/memberbio/george_c_williams. 20. Nesse and Williams, Why We Get Sick, 21. 21. Williams and Nesse, “Dawn,” 4, 3.

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22. Randolph M.  Nesse and Steven C.  Stearns, “The Great Opportunity: Evolutionary Applications in Medicine,” Evolutionary Applications 1, no. 1 (2008): 28–48, 38. 23. Williams and Nesse, “Dawn,” Table 3.1, 31, 47–8, 59–60. 24. See Leigh Van Valen, “A New Evolutionary Law,” Evolutionary Theory 1 (1973): 1–30, 17. 25. Nesse and Williams, Why We Get Sick, 49. 26. Nesse and Williams, Why We Get Sick, 54, 58, 60. 27. Randolph M. Nesse and George C. Williams, “Evolution and the Origins of Disease,” Scientific American 29, no. 5 (1998): 86–93, 86. 28. Nesse and Williams, Why We Get Sick, 5, 4. 29. Nesse and Williams, Why We Get Sick, 159. 30. Randolph M.  Nesse, “The Smoke Detector Principle: Natural Selection and the Regulation of Defenses,” Annals of the New  York Academy of Sciences 935 (2001): 75–85, 83. 31. Nesse and Williams, Why We Get Sick, x, xi. 32. See Gert Kordhoff, “William Hamilton’s Worries about the Future of the Human Genome,” http://wasdarwinwrong.com/korthof97.htm. 33. W.  D. Hamilton, Narrow Roads of Gene Land: The Collected Papers of W. D. Hamilton, Volume 2: Evolution of Sex (New York: Oxford University Press, 2002), xlvii. 34. Nesse and Williams, Why We Get Sick, 11. 35. Nesse and Williams, Why We Get Sick, 158, 154. 36. Nesse and Williams, Why We Get Sick, 24, 12. 37. Nesse and Williams, Why We Get Sick, 91, 213, 236. 38. Nesse and Williams, Why We Get Sick, 155, 137, 133, 31, 131, 241, 127, 179. 39. Nesse and Williams, Why We Get Sick, 209, 210. 40. Randolph M.  Nesse and George Williams, “Are Mental Disorders Diseases?” in The Maladapted Mind: Classic Reading in Psychopathology, edited by Simon Baron-Cohen (East Sussex, UK: Psychology Press, 1997): 1–22. 41. Jerome H.  Barkow, Leda Cosmides, and John Tooby, eds. The Adapted Mind: Evolutionary Psychology and the Generation of Culture (New York: Oxford University Press, 1995). 42. Randolph M. Nesse and George C. Williams, “Evolutionary Biology in the Medical Curriculum: What Every Physician Should Know,” Bioscience 47 (1997): 664–66. 43. Michael F.  Antolin, “Evolution, Medicine, and the Darwinian Family,” Evolution: Education and Outreach 4, no. 4 (2011): 613–23, 614. 44. Nesse and Williams, Why We Get Sick, 245. 45. Nesse and Williams, Why We Get Sick, 245, 246.

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46. Nesse and Williams, Why We Get Sick, 246. 47. Nesse and Williams, Why We Get Sick, 247. 48. George C.  Williams, Adaptation and Natural Selection (Princeton University Press, 1966), 14, 16. 49. Nesse and Williams, Why We Get Sick, 4, 134. 50. “Evolutionary Psychology: A Primer”: http://www.cep.ucsb.edu/ primer.html. 51. David J. Buller, “Four Fallacies of Pop Evolutionary Psychology,” Scientific American 300, no. 1 (2009): 74–81. For Buller’s complete critique, see David J. Buller, Adapting Minds: Evolutionary Psychology and the Persistent Quest for Human Nature (Cambridge, MA: MIT Press, 2005). 52. See Erica Goode, “Viewing Depression as a Tool for Survival,” New York Times (February 1, 2000), http://www.nytimes.com/2000/02/01/ health/viewing-­depression-­as-­tool-­for-­survival.html. See also a reply by Michael M. Gindi, “Depression Debunking,” New York Times (February 8, 2000), http://www.nytimes.com/2000/02/08/science/l-­depression-­ debunking-­723703.html. 53. Nesse and Williams, Why We Get Sick, 217, 232. 54. Nesse quoted in Goode, “Viewing Depression.” 55. Nesse and Williams, Why We Get Sick, 220–21, 248, 10, 12, 14, 197. 56. Nesse and Williams, Why We Get Sick, x. 57. Nesse and Williams, Why We Get Sick, 279. 58. See Susan Blackmore’s Universal Darwinism website, http://www.universaldarwinism.com/Blackmore%20Susan.htm. 59. Nesse and Williams, Why We Get Sick, 5, ix, 12, 14. 60. Nesse and Williams, Why We Get Sick, xi, 5, 7. According to Nesse, this construction was quite deliberate. He and Williams actively sought to tone down statements that were too deterministic if they could not be substantiated. Conversation with Nesse, January 25, 2021. 61. Nesse and Williams, Why We Get Sick, 7, 15, 202, 21, 27. 62. Nesse and Williams, Why We Get Sick, 33, 3, 204. 63. Nesse and Williams, Why We Get Sick, 36, 38, 239, 246. 64. Nesse and Williams, Why We Get Sick, 228, 238, 249, 220, 225. 65. Nesse and Williams, Why We Get Sick, 242. 66. Williams, Adaptation, 12, 47, 56, 61, 101, 122. 67. Williams, Adaptation, 79, 132. 68. Williams, Adaptation, 69. 69. Nesse and Williams, Why We Get Sick, 182–83. 70. Nesse and Williams, Why We Get Sick, 245–48. 71. Hamilton, “Gamblers,” in Narrow Roads of Gene Land, vol. 1 (1996), 363. 72. Nesse and Williams, Why We Get Sick, 235. 73. Nesse and Williams, Why We Get Sick, 235.

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74. Paul Chodoff, “The Medicalization of the Human Condition,” Psychiatric Services 53, no. 5 (2002): 627–28. 75. Edward O. Wilson, Consilience: The Unity of Knowledge (New York: Knopf, 1998), 13, 265. 76. Edward O.  Wilson, The Social Conquest of Earth (New York: Liveright, 2012), 45. See this response in a Scientific American blog post, Cadell Last, “Universality of Preadaptation for the Human Condition,” Scientific American blog (January 16, 2013), http://blogs.scientificamerican.com/ g u e s t -­b l o g / 2 0 1 3 / 0 1 / 1 6 / u n i v e r s a l i t y -­o f -­p r e a d a p t a t i o n -­f o r­the-­human-­condition/.

CHAPTER 10

The Dark Side of Biology: Plan and Purpose in Nature

Plan and Purpose in Nature was meant to be a responsible popularization of science for general audiences.1 It is part of the Science Masters series published by Weidenfeld and Nicolson, a series promoted and advertised by John Brockman, consisting of short books on exciting aspects of science written for the general reader by prominent scientists. “These short, easy-to-read, attractive books present cutting-edge ideas in a format that will enable a broad audience to attain scientific literacy.”2 Surely, one might expect limits to the subtlety of literary allusions in such books. But Williams leaves so many signs of literary games that it is fair for a critic to say that he plays with his reader, and also with Brockman’s project and its aims. These games have to do with the limits to asking what is remarkable and what is wonderful about the natural world. By the 1980s, games, or more precisely game theory, had become a well-entrenched aspect of evolutionary theory. The book was variously titled, and all the titles are relevant. The subtitle for the original British edition is The Limits of Darwinian Evolution. It is worth remembering that evolution was the topic for more than one popular book during this era. Characteristic of this is the bestseller Wonderful Life: The Burgess Shale and the Nature of History (1989) by Stephen Jay Gould. In the United States, Williams’s book appeared as The Pony Fish’s Glow: And Other Clues to Plan and Purpose in Nature, the title of the American edition emphasizing the allegorical story of the pony fish and © The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 M. P. Cohen, George C. Williams and Evolutionary Literacy, Literatures, Cultures, and the Environment, https://doi.org/10.1007/978-3-031-11650-6_10

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the importance of clues. But limits are also central to the argument of the book. To speak of “the limits of Darwinian evolution,” and what natural selection cannot do, is to indicate biological limits humans must understand. According to Richard Dawkins, Williams was unhappy when a publisher altered his original title, Plan and Purpose in Nature: The Limits of Darwinian Evolution, for the American edition.3 Why would Williams prefer one title over another? First and most obviously, because the original title is meant to be directly ironic, as even a review of it on Amazon. com points out. From Williams’s perspective, there is neither plan nor purpose, and all clues point to limitations on the power of natural selection. Williams argues principally that the evolutionary process shows “how far it is possible to go with trial and error alone”: “All the way to the human eye and hand and immune system and all the other well-­engineered machinery by which we, and all other organisms, solve the problems of life. At least that is the orthodox view in modern biology.” To say the process works by trial and error is for Williams to say it works stupidly, as we have seen, and accounts for both its power and its limitations, including the gross deficiencies that result.4 If one understands a process, even one like natural selection, one ought to be able to predict outcomes, or to put it another way, tell a story. So the first chapter of this book is entitled “Adaptationist Storytelling.” The link between prediction and story remains a key issue, as it has always been for Williams. The “adaptationist program” begins with questions of how and why. Answers to these questions “sometimes enable us to predict and make important discoveries.” As he repeats from the argument in “Defense of Reduction,” “the idea that theories about prior history can be predictive is often casually dismissed. This is because people think of prediction in terms of future history, when the important use of theory is to predict the outcome of investigations.”5 One tells a story and then tries to find out whether it is a true story. The predictability of an investigation can be tested. Williams dwells on this method of inquiry. Consequently, there is a difference between a “just-so story” and Williams’s introductory adaptationist story of how the pony fish got its glow. Scientists tell stories to each other in a process that leads to newer and better stories. Entailed is the question of finding and using “clues.” Williams uses a scientific analogy to explain this method, noting that the discovery of Neptune came about as the result of observed anomalies in the orbit of Uranus. He uses another, more complicated analogy to

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historical storytelling as illustrated by Schliemann’s excavation of the cities in Troy. Schliemann began with a myth and a guess. “So Schliemann’s theory, a narrative history with a specific geography, led to the discovery of great interest. This is routinely true of theoretical stories told by evolutionary biologists.”6 Williams’s use of Schliemann for a heroic figure in his introduction is very strange, not only because Schliemann was so entrepreneurial (he was a publicist and exploited the popular press), but also because he was an amateur, though literate in ancient languages. As most cultural historians notice, only fourteen years after Darwin’s story of evolution shook the human sense of identity, Schliemann offered a different narrative that seemed to validate a romantic, mythological past. Further, there are many reasons to believe that Schliemann’s discoveries were deceitful and fraudulent, as discussed in multiple writings by David A. Traill, whose biography of Schliemann was published in 1995.7 As he frequently does, Williams returns to arguments about design, and the story is more elaborate than in earlier versions. Here “mundane processes taking place throughout living nature can produce contrivances without contrivers, and these processes produce not only functionally elegant features but also as a kind of cumulative historical burden, the arbitrary and dysfunctional features of organisms.” The “idea of functional design,” nevertheless, “often takes this form: an organism is observed to have a certain feature, and the observer wonders what good it might be.”8 This story is about light and eyes. The biologist wonders, what good does it do for the pony fish to glow?

An Adaptationist Story of the Pony Fish Perch-like, with an interesting structure of jaws, the pony fish can extend its mouth into a downward-pointing tube and gets its name from this strange mouth, which looks like a pony’s nose when extended. There are many species of pony fish, and some are known as slimys or slipmouths. The common pony fish matures at about twenty centimeters long, ranges widely in river mouths and estuaries throughout the Indo-West Pacific, in the Red Sea, the Persian Gulf, and East Africa—Reunion, Comoros, Seychelles, Madagascar, and Mauritius—to Fiji, north to the Ryukyu Islands, and south to Australia. Pony fish are an important food in the tropics. But the point is, they glow.9 The story of the pony fish’s glow is not based upon Williams’s own research, though it is certainly informed by his knowledge. His narrative

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constitutes a re-telling of work done by John Woodland Hastings, an eminent investigator of bioluminescence. Williams mentions in passing the title of Hastings’s article, “Light to Hide By.”10 When Hastings chose his delicious title he suggested more than passing resemblance to devices of plot in Edgar Allen Poe’s detective story “The Purloined Letter.” Permutations of Hastings’s complete title, “Light to Hide By: Ventral Luminescence to Camouflage the Silhouette,” are interesting to entertain, not only because they emphasize observations about hiding in plain sight, but also because light, silhouette, and camouflage are emphasized by the Francophone diction of his terms. Most important, implications or intuitions about this subject are intimately connected to the shape of storytelling. How the pony fish glows is apparent. It has a light-producing organ, or photophore, deep inside its body, equipped with a reflector to shine its light downwards. “What good is the light?” Williams asks. “Can it really be adaptive for a fish to illuminate its own innards?” He produces a precise and detailed description of the colors of fish and their visibility, especially from below, from a predator’s perspective during daylight hours. He tells a story of war and camouflage, of course, but also a story about investigating these. One is tempted to say that Williams follows what thirty years earlier he called “the light and the way.” The pony fish reveals its interior, but that is not the function or purpose of its light. What is the purpose of its light? The world elicits productive questions and unproductive ones. “What is the purpose of the sounds we make when gnawing bones or biting celery?” Williams asks, “The answer: none.”11 Only good questions provide answers that have implications and can be checked. As any ichthyologist or fisherman knows, most fish are darker above than below and most have extremely white bellies. This is called “adaptive countershading.” But as seen from below, the fish will still appear as a shadow. However, in the pony fish’s habitat “there is no place to hide.” It stays in dark water, but if it is seen by enemies it will be seen from below as a silhouette—“unless, of course, the potential prey can extinguish the silhouette by making its belly glow in a way that matches the light coming from above.”12 Hastings tested this story in a dark room, where he could adjust the light above an aquarium. According to Heraclitus, “Nature loves to hide” (and so indeed does culture), and in plain sight. If you are a fish, you are likely to have a dark side and a light side. The light side illuminates and thereby hides you from predators below. The dark side hides you from predators above. Hiding is essential. Yet the pony fish

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illuminates its innards in order to hide itself to avoid becoming part of some other fish’s innards. It hides by illumination, by revealing itself. The phenomenon of hiding therefore depends on the perspective of the most important observer, the predator. And even predators take advantage of what is more generally described as countershading. People like to hide too, but for different reasons. I, for instance, go off in the woods and the Sierra Nevada, the “Range of Light,” as John Muir called it, and when I am in the mountains, like Muir I often wear gray clothing to blend in with the granite. The pony fish has no such options. Hiding, one might say, is obligatory in a place where big fish eat little fish—and so the metaphor grows. Williams’s Nature is, perhaps, a charnel house, a scene or place of great physical suffering and loss of life. When creatures hide to become invisible in such a scene, to disappear, they are solving a problem. This point of Williams’s story comes from an extract he introduces from Karl Popper, that “we can describe life, if we like, and living organisms as the only problem solving complexes in the universe.” The extract comes from an appendix to Popper’s The Open Universe, entitled “Scientific Reduction and the Essential Incompleteness of all Science.”13 Ernst Mayr had argued that “every advance in physiology began with the question, ‘What is the function of a given structure or organ?’” Just so, Hastings asks the same question, and as a result learns about the anatomical position, spectral emission, timing of illumination, and finally the role of the pony fish’s light in protecting it against predation. “So adaptationist storytelling is an effective device to make important scientific discoveries, but isn’t there a more aesthetic sort of triumph here?” Williams asks. He wonders about the pleasures in finding nature full of such examples of “beautifully effective machinery” and the satisfaction “that human ingenuity can explore and understand these marvels.”14 But Williams does not dwell on these pleasures and marvels for long, because there are constraints for this kind of storytelling. The political scientist Bradley A. Thayer points out that Williams uses Hastings’s story as a reply to Gould.15 As Williams forecasts in his preface, Plan and Purpose in Nature might have been entitled “The Adaptationist Program,” and the subject gets full development especially in chapter 2, “Adaptationist Storytelling.” Though the theme is introduced in the “Defense of Reductionism” (1985), especially in sections entitled “Just So Stories” and “Prediction,” he continues to marshal new examples to explore modes of storytelling by biologists in past and future tenses. He responds, as usual, to Stephen Jay Gould and Richard C. Lewontin’s article “The

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Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme” (1979). Thayer demonstrates how a Williams story and a “just-so story” differ in significant ways. Williams explains about his: “Right from the start, mine was consistent with the known facts about the pony fish and its habitat, and I was careful to make it consistent with what might be called the Darwinian constraints.” His constraints “refer only to well-established material processes in formulating the story. This means avoiding any supernatural factors and always including a way in which natural selection … could maintain the proposed adaptation.”16 Indeed, Williams tells the story about the pony fish in order to explore the value of adaptationist stories. “There may be an analogy between adaptationist explanations and Kipling’s ‘Just-So’ stories,” Williams writes, “but there is a closer one with detective stories.”17 As Poe does, Williams engages in a complex form of doubling when he introduces Sherlock Holmes, not as he appears in an original Arthur Conon Doyle story, but as portrayed in the United States, by Mark Twain in “A Double-Barreled Detective Story.” In Twain’s “tall tale,” Sherlock Holmes proves to be wrong in his rationalization of various clues and in his prediction about a murderer’s identity. But Williams’s point is that in Twain’s version, a Holmes story elicits a “contrary story” that is more accurate than that of the great British detective. Williams is, after all, an American writer, even if he published this book first in Britain. On the one hand, Williams re-tells part of Twain’s Holmes story—just as he has retold Hastings’s story—to demonstrate not only how “Holmes’s theory was superseded by another theory,” but also that “the reason for scientific theories being abandoned is always that a better theory has come along.” There are conventions for this kind of replacement. “In science as in sleuthing, acceptability also depends on adherence to accepted rules for storytelling.” Any story about Sherlock Holmes, for instance, is about “predictions of what various investigations would reveal.”18 Yet choosing Twain’s story is strange on several grounds: (1) Twain’s story is a satire of mystery novels; (2) not only does Twain’s Holmes pursue ratiocination to an absurd degree but also with the wrong result; (3) this story is a late production of Twain (1902) and generally considered inferior, perhaps the worst Holmes parody in the canon. And these are old men’s stories; Twain is sixty-seven years old and Williams seventy-one when they publish them. Beyond a certain point, one never knows about the ironies either might intend, and Williams’s playful ironies often create ambiguity and breed implication. Not a few readers have noticed that the subtitle “Clues

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to Plan and Purpose in Nature” is ironic, but they must also notice that, if is there is no plan or purpose, in such a world clues may deceive those who have false expectations. Williams returns to Hastings’s explanation for the pony fish’s glow. It must not only explain “when the light would be turned on, and what kind of light it would emit, but also why the pony fish keeps its photophore,” because “adaptationist stories are not about evolution so much as about its absence.” Thus, Williams concludes that “Gould’s criticism notwithstanding, adaptationist storytelling continues to be a powerful method for the discovery of important facts about living organisms.”19 This story of pony fish reveals not how they get their mechanisms, but why they keep them. Williams marks a limit for historical inquiry into their light, and also maps the limits for how far adaptationist stories can go. Illuminating limits is a habit for Williams, who often offers an exemplary story and then says it “goes too far.” Frequently the root of the problem— going too far—can be found in the process of analogizing, as we remember from Adaptation and Natural Selection: “Analogies between biological phenomena and human affairs can also be of value at the level of groups of individuals, but there is no simple and reliable guide to tell us where to stop.”20 As one might say about Twain’s tale, unchecked stories are likely to go too far. The teller cannot know where to stop until the story is tested or checked. Consequently, a reader can apply to Williams the same method he uses to test storytelling. Ask questions, he suggests. How does an author tell a story? Why does he tell a story? What are the rules, and where are the limits? What empirical reason do we have to believe the story? What is its function and design? He only seems to undercut these questions when he claims that adaptationist stories are “not about evolution so much as about its absence.” Absences, after all, are empirical clues to be found by the detective or investigator, private or public, professional or amateur. Yet there is something else, and I will return to that. Poe’s investigator, Dupin, is justly famous not only for his creativity or intellect, but also for placing himself in the mindset of the criminal. Dupin discovers a letter hidden in plain sight, whose contents have not been revealed. What analogues one wants to make of these acts within investigations remains to be seen, in light of many ironies.

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Games Natural Selection Plays: Seal Ethics and Mob Behavior It is a tall task to make complicated conceptions into “easy-to-read” stories. This is both a tonal and conceptual issue. One example of the problem is illustrated when Williams shows how game theory can be applied to biological problems. To depict natural selection as a game is to reduce the concept of nature in many ways. Indeed, the arrival of game theory as a tool of evolutionary biologists is a direct consequence of reductionist ideas that Williams, Hamilton, and Maynard Smith embraced in the 1960s, especially when they asserted that altruistic behavior is self-interest in disguise.21 Game theory, above all, abstracts strategies that exercise self-interest, without regard to history. Williams recognizes Maynard Smith as the pioneer and doyen of game theory in evolutionary studies.22 As Maynard Smith writes, “paradoxically, it has turned out that game theory is more readily applied to biology than to the field of economic behaviour for which it was originally designed.” This is true, to a great extent, because though “there are grounds for doubting whether human beings always behave rationally,” when “the concept of human rationality is replaced by evolutionary stability,” all outcomes can be measured on the single one-­ dimensional scale of Darwinian fitness, and without recourse to motive.23 One marvels at the coldness of this proposition, but more at its assertion that natural selection, because it lacks motive, acts more rationally than humans. The Crafoord Prize Committee of 1999 succinctly states that Maynard Smith introduced game theory analysis to biology and coined the term evolutionarily stable strategies (ESS): “Game theory is a mathematical method that may be used to analyse how an actor—‘player’—should behave where he cannot have complete information about what another actor—‘opponent’—is going to do.” Interestingly, Maynard Smith used game theory to show why “it is uncommon for members of the same species to kill one another in combat.” As the Crafoord Committee puts it, “Since no combatant can have complete information on the others’ capacities, the optimal course for both parties is to refrain from escalating the combat. The result—‘the evolutionarily stable strategy’—becomes a ritualised struggle (as among black grouse or ruffs) without bloodshed.”24 Game theory can be used to predict a great many other so-called strategies, including the stable sex ratio of a species.

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Williams worries a great deal about whether and when game theory models reality.25 In Natural Selection, he spells out a list of these reservations, given his own “special concept of adaptation as conformity to a priori design specification.” These reservations about game theory, many based on his view that historicity matters, are worth extracting in detail. The discussion depends upon the phenomenon of “frequency-dependent selection,” which occurs when the fitness of a genotype depends on its frequency in a population—that is, where a genotype has a different fitness when common than when rare. Williams presents his concerns about game theory as a list: (1) “The theory in no way accounts for the evolutionary origins of the alternatives it considers. It accepts them as historical legacies and deals with their relative numbers.” (2) “It is irrelevant to the ontogeny of the strategies considered. [For instance] It may tell us why manhood and womanhood or other sexual phenotypes have certain frequencies, but it leaves untouched the question of whether sex is environmentally or genetically determined, or even whether there are men and women, rather than a hermaphrodite population in which each individual partakes of both manhood and womanhood.” (3) “More than one strategy or ratio of strategies may be a stable equilibrium. The one established by selection may depend on the starting ratio, another kind of legacy from the past.” (4) “Game theory was designed to enable engineers and economists to arrive at the one best solution to a problem. Evolution seeks only the most easily attainable stable solution, which need not be best, nor even moderately good, for either population or individual.” (5) “An ESS payoff matrix by itself says nothing about evolutionary changes in descriptive details of strategies considered, such as maleness and femaleness. It can tell us about relative numbers of men and women, but nothing about specific male and female adaptations.” (6) “As noted above, a frequency-dependent equilibrium is not an optimum, for either an individual or its population, nor is it the solution to any kind of problem faced by an organism. It simply abolishes the problem.”26

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In 1992, Williams decides that the prevalence of mutually exclusive designs of maleness and femaleness resulting from frequency-dependent selection seems to be obviously an example of an ESS. Does he approve of this kind of storytelling five years later? In a section of Plan and Purpose titled “For the Harm of the Species” (which, significantly, follows “The Scarcity of Individual Subordination to Group Interests”), he sets out an argument about the failure of functional organization in natural selection. He works his way from a simple game toward a more complex transaction, using the prisoner’s dilemma with devastating effect. Game theory is a relevant mode of analysis here since the “absence of a functional organization for populations or species actually has worse consequences than might be imagined, because natural selection within the groups can produce results that not only fail to help the group as a whole but may be harmful, or at least systematically wasteful.”27 Williams uses, as an example, the effect of selection on the sex ratio of a population, asking, “What then determines what fraction of the total is male and what fraction female?” He finds the history of thought on this question “rather curious.” “Darwin worried about it a little, but shrugged it off,” and the idea was resurrected by Ronald A. Fisher, who in 1930 invoked what is known as frequency-dependent selection, which Williams calls “a rather elementary idea in traditional economic reasoning.” Imagine you are equally skilled as a smith and a carpenter, but have necessary resources to set up shop for only one trade. “You know that the village already has a smith. Should that influence your decision?” So too with sexual strategy, as Williams recounts a remarkably 1950s version of procreation and sexual strategies: sex ratio provides the simplest possible example for analysis of frequency-dependent selection in biology, using game theory as a payoff matrix. As he illustrates, “the individual labeled player has a choice of being a male or a female. The next individual it encounters, labeled opponent, will be one or the other. Whichever our player chooses to be, if the opponent turns out to be of the opposite sex, it wins in its game of reproduction. The winning is represented by the score of 1 in the matrix. If the opponent is of the same sex, there is no reproduction, no winning, score 0” (see Fig. 10.1). Under the circumstances, you would advise a player to flip a coin, unless he or she knows there is disparity in the sex ratio. “But suppose you know that the males are slightly in the majority in the population. Now you can offer sound advice: be a female; or, if it is too late for that, have a daughter, not a son.”28 This principle, Williams quips, explains how men choose their favorite singles bars.

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opponent male female player

male

0

1

female

1

0

Fig. 10.1  “Payoff matrix for the sex-ratio game.” Figure in George C. Williams, The Pony Fish’s Glow: And Other Clues to Plan and Purpose in Nature (New York: Basic Books, 1997)

In the long term, such game-playing means that “the minority sex stops being in the minority and the sex ratio stabilizes at equal numbers.” We know this because there are approximately equal numbers of men and women in the world, and they will produce nearly equal numbers of offspring: “Neither sex has an advantage, and selection on sex ratio disappears. It will reappear and act to reestablish the nearly equal numbers if ever this equilibrium is disturbed.” The question is complicated when one considers the life history of a population. Consequently, Williams asks about the age where there are equal numbers of males and females, the effect sons and daughters have on mortality rates or costs to the parents. When do sexes mature? Why would there be a slight preponderance of boys at birth? While these are questions that biologists discuss at great length, “their answers imply quite minor quantitative modifications of the 50:50 ratio expected from simple frequency-dependent selection.”29 Williams makes a turn when he introduces a deliberately grotesque example of elephant seals that Richard Dawkins explores in River Out of Eden (1995) to show how “the effect can be decidedly negative.” In breeding season, adult elephant seals crowd beaches. In these crowds of a small number of large males and a large number of small females, called harems, most males are unsuccessful and don’t make it to the beach. They remain celibate bachelors. As Williams writes, “They represent a waste of resources, because only a small fraction of them will reproduce. Yet because of frequency-dependent selection, the population goes on, generation after generation, producing about the same number of males as females.” He continues, “actually it is worse than would be inferred from just the equality of numbers,” because only the largest and strongest males win, while “frequency-dependent selection keeps the population producing a wastefully large number of males, and sexual selection goes on making

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each male wastefully large.” Large males are large consumers, yet females bear “the entire physiological burden of reproduction for both her own and her mate’s genes.” As Williams also writes, humans are like this too, though on a minor scale. Consequently, “the recent feminist slogan ‘men are not cost-effective’ is entirely correct biologically: there are too many of them; they are too big; they accomplish less than women per unit of resources consumed.”30 This situation allows Williams to take up the game of the prisoner’s dilemma, which he prefers to call the “trader’s dilemma.” The game “shows another way in which natural selection may have negative effects at the group level.” If, for instance, in any transaction both a buyer and a seller look out only for self-interest, in any trade both will choose to be dishonest. “The net expectation from all this rational decision making,” he comments, is that honesty is not the best policy. “The cheater’s winnings are greater, and some honest player is penalized. What happens if all players cheat? No one wins.” Yet everyone perceives dishonesty as the best policy: “No matter what your opponent does, you do better by cheating.”31 Unfortunately, when natural selection functions by this rule, the result is that everyone loses. Williams then applies the “trader’s dilemma” to a different kind of biological example, a school of fish in a pond, where optimum size should be determined by two factors, predation and nutrition: “When a predator attacks, it is likely that no more than one fish will die, because, once one is caught, the others can get away.” Under these circumstances, it is safest to join the largest school available, but “the bigger the school the greater the competition for food and the less there will be for each fish. The optimum school size will be that with the greatest excess of benefit, from predator avoidance, over cost, from decreased nutrition.” Thus the following dilemma: “Suppose ten is the optimum number, and a fish finds itself in a school of twenty. What should it do, purely from the standpoint of self-­ interest? If it stays, it and all the others will suffer from a food shortage. If it leaves, it would help all the others by increasing their nutrition, but it would expose itself to much greater risk of death from predation.” The math suggests that, following the rules of the game, “the twenty fish will continue to swim together, even though, from every individual’s perspective, it would be better to break up into two schools of ten.”32 A human faced with an “optimum trade-off between predation hazard and competition for food” might sort out the problem rationally. “Unfortunately, the only decision making the fish can manage is a simple

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‘I should stay in this bad situation’ versus ‘I should move away into a worse situation.’” This dilemma, he observes, inevitably results in schools that are too big; this happens regularly. “An enormous number of other examples could be described that would illustrate the principle that, although some groups, such as honeybee colonies, are functionally organized, most animal groupings are not. They are just mobs of self-seeking individuals.”33 We have heard this story before, and though one may wonder, one might not be calling it wonderful.

How Humans Experience Sex and Death These are ugly games, these stories about behavior in the world of no plan or purpose. Those who invented game theory for Cold War strategizing think like John McCarry’s fictional character Barney Wolkowicz: “His approach to human nature was so primitive that most sophisticated men could scarcely understand it.”34 But Barney is mostly right, like Bill Hamilton, who, according to some sources, was a “thing person” rather than a “people person,” for whom all social behavior operated according to beautifully clear and brutally simple rules that could be reduced to mathematical expressions.35 There is a foolishness of sympathy and a foolishness of distance. When Williams entertains game theory in Plan and Purpose, he does so to test various kinds of limits—and tests a reader’s patience too. When humans and their experience of sex and death become dominant subjects for his investigation, about a third of the way through his book, one can guess the resultant tone. As we remember, Williams’s theory of senescence treats it “as an evolved characteristic of the soma” to be found “wherever a soma has been evolved, but not elsewhere.”36 To put this in striking terms, to have a body is to age and die. The same evolutionary process first favors youth and later compels death. One also remembers that Alfred E. Emerson’s notions of “beneficial death,” published in 1960 and 1961, goaded Williams to study senescence because this was a process that could be studied.37 His writing always interests itself less in death than in processes associated with it, just as his earlier works focus less on sex than on sexual reproduction. Sterelny and Griffiths named their introduction to the philosophy of biology Sex and Death, they claimed, “because it was fun. And philosophy of biology is fun.”38 One wonders what Williams thought of their title, or their idea of fun. From the beginning, his work focused not simply on sex

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and death, but on the causal and fatal relationship between them. The relationship doesn’t seem like much fun, and it just keeps going on forever. Nor is there much fun in contemplating it. However, he entitles consecutive chapters of Plan and Purpose “The Human Experience of Sex and Reproduction” and “Old Age and Other Curses.” Focusing on the human experience of sex and death constitutes a change in his writing perspective but not in the substance of his views. Williams thus enters into an ethical realm he has been constructing for over a decade.

Family Values By re-telling stories normally devoted to dramatizing the sanctity of the family, Williams also means to contradict many religious assumptions. As he recapitulates his argument for the concluding chapter of Plan and Purpose in Nature, “I attempted to show that the eighteenth- and nineteenth-­century God-is-smart tradition is fallacious. Now I plan an attack on what I’ll call the God-is-good tradition.”39 Let us make sure we understand. For Williams, there is no God, and the processes He devised are evil. If there had been one, ditto. But Williams does not entertain readers with distracting thoughts of gods except as a rhetorical strategy. In an interview associated with the publication of Plan and Purpose, Frans Roes notes a reference to Buddhism in Adaptation and Natural Selection and asks Williams whether he has a special liking for Buddhism. Williams replies, My reading on this sort of thing is extremely limited, but as a doctrine I think Buddhism is more compatible with the spirit of scientific inquiry than what you get in the Old Testament. I think it is because of the explicit recognition in Buddhism that things are not naturally good. There is a lot of pain and suffering in the world, and that is because that’s just the way the world is. And the way to overcome this is to live a certain way and to be resigned to the inevitable imperfections of life.40

Within this context, Williams reverses the order of sex and death in Plan and Purpose, developing two examples of religious attitudes that make no sense in light of modern biological thought. First is the “holy-corpse fallacy.” “When people die, their relatives and friends behave as if there were some moral significance in the dead body. They ignore the fact that the ‘last remains’ are just that, material that happened, at the time of death, to

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provide the medium of expression for a human life.” As he points out, “The holy-corpse fallacy once had support from the biological concept of protoplasm, the special living matter of an organism,” that some imagined as a “stable entity” regulating the material flux of the body. “Protoplasm was often discussed in the biology courses I took in the 1940s. It is a term almost never heard today.”41 Though other species perform apparent rituals associated with dead bodies, their motives are obscure. A second, “related fallacy” Williams calls “the moment-of-conception fallacy.” When a human egg and sperm join to define a new and unique human genotype, this process “does not produce any human hopes and fears and memories or anything else of moral importance implied by the term human. The newly fertilized egg may have the potential for a fully human existence, but that potential was there even before fertilization. The same can be said of all the fertilizations that might have been.” Indeed, fertilization also means “an early death for millions of competing sperm.” Williams argues that, “besides being philosophically unacceptable, the moment-of-conception fallacy is biologically naive. It implies that fertilization is a simple process with never a doubt as to whether it has or has not happened. In reality, the ‘moment’ is a matter of some hours of complex activity” that includes “elaborate biochemical interactions” so that any “strictly biological definition of humanity would have to specify some point in this elaborate program at which the egg and sperm have suddenly been endowed with a single human life.” Other difficulties ensue. “If that single human life develops for a while and then divides to produce identical twins or triplets, are they to be considered one human being?” As he writes, “This would be contrary to almost everyone’s moral sensibilities.” Williams concludes that “the only realistic view is that a human life arises gradually.” Consequently, if one wants rules, “the recognition of the full humanity in a full-term newborn would be one such simple rule. I am not inclined to argue that it is the best possible, but it makes more sense than any recognition of fetal rights.”42 Such an argument articulates a desire to see from inside natural processes and use this perspective to change culture. To be against nature may mean many things, but most of all it also includes the professional risk of being a detective who has seen what is inside nature—at a lower level than most wish to observe—and who reports the results. Need the process of human bio-history, shaped by reproduction, be hidden by the ignorance of religious doctrines like these, he asks, or can an advancement in scientific thinking change a culture? I believe that Williams is pessimistic. As in

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the concluding sentence of The Great Gatsby, “So we beat on, boats against the current, borne back ceaselessly into the past.” How can the human experience not be a victim of its historical past? After such reasoning—or knowledge—what forgiveness? Williams approves “recognizing that an unremitting effort is required to expand the circle of sympathy for others. This effort is in opposition to much of human nature and is surely the struggle proposed by Thomas Huxley and seconded by me.”43 For Holmes Rolston III, a philosopher best known for his work in environmental ethics, Williams’s perspective is appalling. As Rolston puts it, “Williams and Dawkins, having from somewhere learned already to ‘condemn’ evolution, will welcome genetic engineering in their ‘unremitting effort to expand the circle of sympathy for others,’ ‘trying to teach generosity and altruism to others because we are born selfish.’”44 But this is not what Williams has written in his articles and his book about Darwinian medicine. Biology demonstrates, as Williams does say, that people live by illusions and that those illusions are damaging. Many temptations hide along the way to evolutionary ethics.45 And scientists are often brash. Because ethics must be about human behavior, sociobiology is inescapably tangled in discussions of ethics.

Biology Noir This is surely biology noir—dark, popular, hardboiled pulp writing. The literary term hardboiled has its own interesting history, suggesting careful and conservative behavior, learned through experience perhaps, as well as hardheadedness, shrewdness, and bluntness of expression. The hardest thing to beat, as an old expression goes, is a hardboiled egg. The hardboiled style has been particularly male, with implications about body language, gender, and power. Noir fiction is always about detection, though not always by a professional detective. Be he investigator or victim, the detective’s object is to uncover, expose, discover, reveal, disclose what people would prefer not to know or see, to take the cover off the pot and look in—and maybe even stir things around. Noir looks at the dark side of behavior that seeks power and money and wants to hide sexual desire, infidelity, and violence. Noir focuses on primitive urges, as in the work of American naturalist writers like Stephen Crane. More recently, Raymond Chandler (1888–1959) concludes in “The Simple Art of Murder” (1950), “It is not a very fragrant world, but it is the world you live in, and certain writers with tough minds and a cool spirit of detachment can make very

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interesting and even amusing patterns out of it.”46 Chandler’s midcentury America believes the world is not nice, or kind, or fair. This is not the best of all possible worlds. So, to focus on sex and power, infidelity and selfishness, wealth and poverty, is to seek the dark patterns of human behavior. How can these patterns not be considered maladaptive? It is the purpose of science to see the world, or certain aspects of it, with a kind of detachment—the world as it actually functions. This is particularly true for evolutionary biology, as it uncovers the processes that life undertakes. It is also true that, historically, evolutionary biology at large is a critique of certain ideologies and illusions. The detective in noir literature, interestingly enough, is not just a dispassionate observer, but also sometimes a romantic. This is less true of Williams, who behaves a bit more like Dashiell Hammett’s Continental Op.47 The Op avoids attracting attention and keeps his opinions and impressions to himself, sometimes sharing them with his confidential reader. He always starts his detective work cold, with as few presuppositions as he can. Nevertheless, he follows a regularized order and procedures, or at least he claims to do so. The Op asks a large number of questions and attempts doggedly to get them answered. He often turns these questions upside down or inside out. He proceeds at an even and sometimes slow pace despite pressure to get the case solved. The Op tries a few simple tricks, until—perhaps out of desperation—he stirs things up, and then looks into the pot he has stirred. Unlike Raymond Chandler’s Marlowe, the Op is unlikely to believe that he can redeem the world. In all art, Chandler writes, there is the quality of redemption.48 So too, perhaps, with scientists like E. O. Wilson, who believe that following natural laws will save us.

The Ethics of Sociobiology Williams concludes Plan and Purpose by scrutinizing “The Wilson Manifesto” in Sociobiology (1975). He examines a big chunk of what he considers to be that manifesto—though his extract is not the one usually chosen by commentators. Just as Williams enters questions of human behavior carefully, so too he frames his critique of Wilson carefully. As he writes, “Wilson’s book is a magnificent achievement and probably the most philosophically important biological work of the century, but perfection is an elusive quality.”49

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The phrase “perfection is an elusive quality” might equally apply to Williams’s view of adaptation. Even if human behaviors might be “designed,” Williams will not go as far as E. O Wilson did when he published “Human Decency is Animal” in the New York Times in 1975. Yet in some ways Williams wishes to go farther. The Times article comprises a manifesto drawn from Sociobiology, but not the one that Williams cites and extracts. Here is what Wilson argues in the Times: In hunter-gatherer societies, men hunt and women stay at home. This strong bias persists in most agricultural and industrial societies and, on that ground alone, appears to have a genetic origin…. My own guess is that the genetic bias is intense enough to cause a substantial division of labor even in the most free and most egalitarian of future societies…. Even with identical education and equal access to all professions, men are likely to continue to play a disproportionate role in political life, business and science.50

Reading this passage now, one realizes the consequences for scientists who do not see their own social and cultural illusions. There may also be consequences for not recognizing the constraints inherent in natural processes, and errors that come from refusing to recognize weaknesses in the design of natural entities; some do not wish to see—or would rather not think about—such limitations. But when Wilson transports biological predispositions—if that is what they are—directly into a cultural description of present behavior, and when he seems to insist that biological history prescribes current social roles, he appears repulsive to Williams and to me. As we remember, according to Wilson, “sociobiology is defined as the systematic study of the biological basis of all social behavior,” and he believes that ethical behavior is subsumed under social behavior, so that ethics is a subject for scientists. When Wilson proposes that “ethics … be removed temporarily from the hands of the philosophers and biologicized,” he enrages plenty of humanists, and the passage above reveals why.51 Yet, in 1997, Williams extracts the first paragraph of the first page of chapter 1 in Wilson’s Sociobiology, “The Morality of the Gene.” I provide the complete passage below and bracket the section of it that Williams quotes, which contains only Wilson’s description of the perspective of the modern biologist. Camus said that the only serious philosophical question is suicide. That is wrong even in the strict sense intended. The biologist, who is concerned

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with questions of physiology and evolutionary history, [realizes that self-­ knowledge is constrained and shaped by the emotional control centers in the hypothalamus and limbic system of the brain. These centers flood our consciousness with all the emotions—hate, love, guilt, fear, and others— that are consulted by ethical philosophers who wish to intuit the standards of good and evil. What, we are then compelled to ask, made the hypothalamus and limbic system? They evolved by natural selection. That simple biological statement must be pursued to explain ethics and ethical philosophers, if not epistemology and epistemologists, at all depths.] Self-existence, or the suicide that terminates it, is not the central question of philosophy. The hypothalamic-limbic complex automatically denies such logical reductions by countering it with feelings of guilt and altruism. In this one way the philosopher’s own emotional control centers are wiser than his solipsist consciousness, ‘knowing’ in evolutionary time the individual organism counts for almost nothing.52

As one can see, and as Williams reveals, E. O. Wilson’s project of biologizing ethics is not an afterthought, and though this paragraph frames the argument of his 1975 book, it is, in turn, framed by a discussion of suicide. This passage dramatizes that the Wilson project is not, as some have argued, merely an addendum to his chapter 27, nor is his discussion of suicide casual. Williams chooses E.  O. Wilson’s premise rather than his conclusion and insists, first, that his manifesto fails to follow the lead of his premise, and second that the premise itself is unduly restricted. Williams ignores Wilson’s discussion of suicide for other reasons. “Why ‘self-­ knowledge’ and not knowledge in general?” Williams asks, “And why just the parts of the brain most concerned with emotions?” Indeed, “Why not the whole system of neural control and cognition. Why just ethics and epistemology?”53 An interesting tactic, considering his own argument in the “Sociobiological Expansion,” where Williams writes that it would be unwise to base any argument about human ethics on natural processes— though he does so in reverse, by basing his position on what he has learned about what natural selection cannot do. (This is congruent with his move to focus increasingly over the years on maladaptations.) Put another way, Williams does not refuse to ground morality in evolutionary theory, but implies this move nevertheless by dialectical means, when he creates an antithesis and grounds morality in resistance to natural processes. He says that humans must learn how to be unnatural—and, insofar as they are ethical beings, have probably been doing just that all along.

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One must remember that Williams writes an evolutionary ethic, not an environmental ethic, and the differences in obligation between the two aims are huge. What, one might ask, are evolutionary ethical responsibilities directed toward? And to what end? To speak analogously, what is important and what is threatened? The answers are not so obvious as Wilson’s because they are humanistic. However, Williams’s discussion of Wilson’s manifesto is not about ethics, but about knowledge. He allows his critique of Wilson to conclude his book because he approves “the use of the theory of natural selection to understand human nature and the current human condition,” a project that he calls “an immensely challenging task that no one can hope to bring to anything like a finish. We can only try, and must,” because humans and all creatures are “utterly dependent” on natural selection. Perhaps this is most obvious in environmental and medical matters, as he notes, “but there is no aspect of human life for which an understanding of evolution is not a vital necessity.”54

What Sociobiologists Should Reveal Williams has no faith in a planner, because there is no plan. He finds it difficult to imagine that humans can find the roots of any adult sense of ethical value in theology or biology. Unlike E. O. Wilson, he suspects that human decency is not animal. Humans are alone with their own devices. Consequently, Williams differs from Wilson in his assessment of what sociobiology is supposed to study and what sociobiology is supposed to reveal. Williams dramatizes a sharp distinction between cultural and biological domains through his use of the word altruism. Although Plan and Purpose includes altruism in the index, the author never uses the term. Because, as he writes in an entry for the 1999 MIT Encyclopedia for Cognitive Sciences, “In biology, altruism has a purely descriptive economic meaning: the active donation of resources to one or more individuals at cost to the donor. Moral values or conscious motivations are not implied, and the ideas are as applicable to plants as to animals.”55 Kin selection, reciprocity, and manipulation explain such donations. When he nods to the argument made in 1994 by D. S. Wilson and Elliott Sober, that cooperative human groups historically “must often have prevailed in conflicts with groups of more consistently self-seeking individuals,” he also remarks, “The

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resulting greater prevalence of human altruism would be more likely to result from culturally transmitted than genetic differences.” For these reasons, Williams is, quite frankly, befuddled by Wilson’s search for the roots of valuable emotions and self-knowledge in biological evolution. Instead, Williams documents “the inherent limitations of human thought processes” for understanding time, space, and other relations, despite sometimes successful intuitive perceptions by humans.56 Intuition has been “useful in achieving genetic success,” but it is unreasonable to expect more of it. However, he recommends a journalistic account of the origin of virtue, Matt Ridley’s The Origins of Virtue, and Robert Wright’s bestseller on evolutionary psychology, The Moral Animal: Why We Are the Way We Are.57 He speaks of these as “excellent detailed discussions of altruism in the animal kingdom and in human evolution, and of the history of thought on these topics.” Williams’s own pronouncements are issued as challenges, and they are answered, to a great extent, by sociobiologists like Sarah Blaffer Hrdy and by developmental biologists like Mary Jane West Eberhardt; the work of both he warmly encouraged. This is his calling, the role he set for himself and others—a role he follows with integrity, as his letters to aspiring scientists and his unremitting editorial work for the Quarterly Review of Biology indicate.

Technical Matters It is dangerous for a biologist to enter the realm of philosophy, just as it is dangerous for a philosopher to comment on evolutionary theory. To judge by his allusions, Williams is knowledgeable about the philosophical literature surrounding evolutionary theory. He introduces his chapter on design with an epigraph from “Parts of Animals,” where Aristotle proposes that parts of the body must exist for the sake of some purpose or action, and “it is evident that the body as a whole must exist for the sake of some complex action.” Williams considers that the modern biological interpretation of “complex action” to which all adaptations contribute must be reproductive success, though he thinks Aristotle may have missed it.58 As he did in the Huxley essays, Williams invokes “Hume’s law” about the distinction between “is and ought.” He approves the “distinguished historian of science” Robert J. Richards’s discussion of “a lubricous slide from an is to an ought” and assents to the premise that “moral directives

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cannot be deduced from descriptive premises.”59 He then explores the lubricious slide as the result of “a more general kind of fallacy that might be called a domain mix.”60 One domain, as Williams explains, allows for material description like length, mass, density. This material domain can be distinguished from moral, mental, and codical domains. As Hume’s law indicates, “We cannot derive consequences from one domain using descriptors peculiar to another (that is, a domain mix).” Most obviously “we cannot deduce moral conclusions from physical premises.” Biological discussion—which is to say, writing that is abundant in libraries—he finds often confused in mixing material and codical concepts. “A triple domain mix, mixing material, codical, and mental, has been rampant in recent years” when discussions slip from physical aspects of nerve cells into the codical realm of the information carried by nerve cells—thus mistaking medium and message—and, from there, slide into analysis of pleasure and anxiety “and other concepts proper to the mental domain.” “From such flights of unreason the authors claim to have provided a physical explanation for mental phenomena.”61 Williams criticizes exactly this slippage in Wilson’s work, “for what I felt to be the inadequacies in his use of the theory of natural selection to understand human nature and the current human condition.”62 Almost by definition, E. O. Wilson has demonstrated that he is not so literate in philosophy. Though Williams mentioned Wilson years earlier in his “Sociobiological Expansion,” he refrained for over twenty years from commenting on “The Wilson Manifesto.” One wonders why.

Huxley, Price, Wilson, Suicide I return to the part of the Wilson Manifesto that Williams does not quote. In that vein, E. O. Wilson has written, “I’m reminded of what Bertrand Russell once said about people’s unwillingness to think about population growth. He said people would rather commit suicide than learn arithmetic.”63 Why indeed do scientists despair? Thomas Henry Huxley despaired famously and dramatically, and in that depth he wrote, “You see a meadow rich in flower & foliage and your memory rests upon it as an image of peaceful beauty. It is a delusion … Not a bird twitters but is either slayer or slain and … not a moment passes in that holocaust, in every hedge & every copse battle, murder & sudden death are the order of the day.”64 Huxley’s despair has been attributed to

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his grief over the death of his daughter. So too, George Levine has written eloquently about Darwin’s grief over the death of a daughter. One can attribute Williams’s views to no such tragedy. What if he simply articulates the way things are? Sometimes despairing people commit suicide. Evolutionary biologists are likely to remember George Price, who was strongly influenced by Williams’s work. As one distinguished scientist puts it, “Price was the first to see that ritualized behavior, game theory and evolutionary genetics belonged together in a coherent vision of behavioral evolution. Price’s vision started Maynard Smith’s work on evolution and the theory of games, which is among the most influential developments of the past few decades.”65 George Price killed himself in the same year that E.  O. Wilson published his manifesto in Sociobiology. E. O. Wilson’s father also committed suicide. Consequently, there is an unmistakable aspect to E. O. Wilson’s repeated discussion of philosophy and suicide. When he refers to Wilson’s manifesto, as I have noted, Williams trims the bracketing or framing passages about suicide from Wilson’s text, and the trimming makes one think about presence—and its absence. Wilson dismisses Camus in his manifesto in the same year Price commits suicide. There is some terrible mixing of categories here, of seeing and not seeing. The implication of Camus’s fable or allegory or myth is relevant to the story of George Price and much more. As Camus wrote at the beginning of The Myth of Sisyphus and Other Essays, “it is legitimate and necessary to wonder whether life has a meaning.” There is a danger in seeking normative values in the wrong place, as Camus argued, and one must choose to live without warrant from some power beyond, the myth of Sisyphus being “a lucid invitation to live and to create in the very midst of the desert.”66 It is embarrassing that E. O. Wilson does not understand Camus’s fable in 1975. You would have missed the 1960s if you didn’t know of Camus’s Sisyphus, or that “his rock is his thing.” Wilson missed what Williams does not: Camus’s image of the stone and the mountain, of “futile and hopeless labor,” in resisting natural force, in a “universe without master,” with the reward, however brief, of contemplation.67 According to William Hamilton, in Narrow Roads of Gene Land (1996), when George Price “convinced himself that something at least close to what I claimed was true, he became very depressed.” Sometime later, even after Price believed he had discovered what he thought of as his miraculous covariance formula, he remained depressed, cut his carotid artery

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with a scissors, and bled to death. Bill Hamilton was invited by the police to collect his effects and tells a most intimate and distressing version of Price’s suicide. He remembers Price as “a man as deeply shocked by kin selection as the Victorian lady had been by evolution itself”; he remembers walking across Price’s room as “his dried blood crackled on the linoleum,” and he remembers collecting the papers. “That is how his life became dreamlike for me and also how his colourful thread in my science and my life ran out.”68 Some twenty years later, Williams writes: With what other than condemnation is a person with any moral sense supposed to respond to a system in which the ultimate purpose in life is to be better than your neighbor at getting genes into the future generations, in which those successful genes provide the message that instructs the development of the next generation, in which that message is always ‘exploit your environment, including your friends and relatives, so as to maximize our (genes’) success,’ a system in which the closest thing to a golden rule is ‘don’t cheat, unless it is likely to provide a net benefit?’69

Raymond Williams argues that references made to nature are usually selective, according to the speaker’s general purpose, and that arguments about nature are often really arguments about humanity.70 Perhaps we always confuse what we think is first nature, defined as the natural process uninfluenced by humans, for what is in reality second nature—nature to which human labor, culture, and action is added. Humans construct a second nature because they can. There may be selfless persons, but there are no selfless genes. Selflessness, writes Williams, is a human invention, except under certain rare conditions. As Williams reminds us over and over again, natural selection, being statistical, reveals what happens predominantly—not what is conceivable, but what is likely to happen. Humans do not possess their own individual genomes, or perhaps even their selves, but they can contemplate, change, and resist forces that alter them. Well, that is badly put. But there are limits to human inventions.

Distractions, Wonder, Allegory Science aims to see clearly, to see without unintended and unexamined preconceptions, to see what truly is and is becoming, not through the lens of desire, hope, or culture. But to see aesthetically is always to see with

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preconception. It is onerous to see without aesthetic preconception; perhaps it is impossible. The effort, however, is worthy. As to writing: how is it possible to write well without concreteness? And since writing concretely, if only by the process of selection, implies an aesthetic, how can writing be concrete without revealing an aesthetic? Williams argues early that the ultimate essence of the theory of natural selection deals with abstraction on the level of gene and on the level of population.71 Aesthetic abstraction can distract an observer from lower levels; it may indeed blind the observer to the relations between lower levels and constituent parts. Williams believes that looking at the lowest possible level helps to eliminate ideological notions of control by processes from above, control by a god or any other planning force. Indeed, he suspects that seeking explanation at higher levels constitutes the inherently religious activity of seeking plan and purpose. One wonders whether abstraction might be inherently religious, but also whether seeing at a lower level can lead to a certain kind of belief. Consider wonder and awe. “If a child is to keep alive his inborn sense of wonder,” wrote Rachel Carson in A Sense of Wonder (1964), “he needs the companionship of at least one adult who can share it, rediscovering with him the joy, excitement and mystery of the world we live in.”72 Surely, though, an adult—and particularly a biologist—means something different by “I wonder.” Helping children to enjoy life is all well and good, but when language of wonder and awe is totally absent from the writings of George C. Williams, I take this to mean that his perspective is for adults. Wonder is valuable—and perhaps gives one reasons to live. What Williams provides gives one a perspective on those reasons, and especially the degree to which they are fictions. He is not writing children’s books. The danger of seeking lower levels is a longstanding theme in American literature. When Pip falls overboard in Moby Dick, The sea had jeeringly kept his finite body up, but drowned the infinite of his soul… [Carried] down alive to wondrous depths, where strange shapes of the unwarped primal world glided to and fro before his passive eyes … Pip saw the multitudinous, God-omnipresent, coral insects, that out of the firmament of waters heaved the colossal orbs. He saw God’s foot upon the treadle of the loom, and spoke it; and therefore his shipmates called him mad.

Melville’s Ahab is particularly interested in “a little lower layer,” and Ishmael wonders,

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Is it that by its indefiniteness it shadows forth the heartless voids and immensities of the universe, and thus stabs us from behind with the thought of annihilation, when beholding the white depths of the milky way? Or is it, that as in essence whiteness is not so much a color as the visible absence of color; and at the same time the concrete of all colors; is it for these reasons that there is such a dumb blankness, full of meaning, in a wide landscape of snows—a colorless, all-color of atheism from which we shrink.73

And so, Melville wrote to Hawthorne, “I have written a wicked book, and feel spotless as the lamb.”74 The life-sustaining patterns humans create are just that—patterns humans create. Williams decides that it is fruitless to seek these patterns as if they are inherent in natural phenomena. The higher light humans seek must be created. Below—below the pony fish—is darkness and predation. Humans create light from within and do so to hide from something. This means that Williams self-consciously writes allegories of natural selection and expects some awareness from his readers. After all, any written text becomes interpretable, through the process of simile, as an allegory or allegorical narrative. Sometimes such interpretability is, in retrospect, humorous, and humor, though not encouraged in a scientific writer, ought to be factored in. Williams looks down into the depths. Darwin looks up into his much-storied tree, which he first introduces as a simile: “The affinities of all the beings of the same class have sometimes been represented by a great tree. I believe this simile largely speaks the truth. The green and budding twigs may represent existing species; and those produced during former years may represent the long succession of extinct species.”75 Darwin’s narrative assumes a unity of life that now seems dubious to biology.76 However, Darwin’s representation of life as a great tree, even if no longer useful to biologists, has been inscribed within the biological literature as an allegory.77 Sometimes developing an allegory is a lifelong process, carrying along its humorous aspects, even its ridiculousness, as the writer casts and recasts the narrative. The narrative becomes an allegory by the process of abstraction and, eventually, its own reduction to absurdity. The allegory remains, even when its basic premises are no longer considered valid, even as absurdity shadows it. More to the point, there is a certain quaintness or silliness when Darwin expands his simile, and a certain humor in his working it out in detail.

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This may be true for Williams as well. This set of processes does not auger a good thing or a bad thing, but simply reveals how language and narrative work upon attentive readers, leave their traces in thought. Because contemplation of any narrative, as representation, offers an opportunity to turn it into a simile or an allegory, no narrative suffers complete demise. So I will claim with the Allegory of the Pony Fish, which I take to be Williams’s version of the classic Allegory of the Cave.

Notes 1. Plan and Purpose in Nature: The Limits of Darwinian Evolution (London: Weidenfeld & Nicolson, 1996) was published in the United States as The Pony Fish’s Glow: And Other Clues to Plan and Purpose in Nature (New York: Basic Books, 1997). I use the American edition, but refer to the book by the always-included portion of its title. 2. See http://www.williamcalvin.com/bookshelf/ScienceMasters.htm; http://www.brockman.com/press/1994.10.02.sundaytimes.html. 3. Richard Dawkins, A Devil’s Chaplain: Reflections on Hope, Lies, Science, and Love (Boston, MA: Houghton Mifflin Company, 2003), 207. 4. Williams, Plan, 13, vii, viii. This citation and all others for the book are to the 1997 American edition, The Pony Fish’s Glow: And Other Clues to Plan and Purpose in Nature. 5. Williams, Plan, 3. 6. Williams, Plan, 3. 7. David A. Traill, Schliemann of Troy: Treasure and Deceit (London: John Murray, 1995). 8. Williams, Plan, 8, 10, 14. 9. The light organ system of the pony fish is sexually dimorphic. See Prosanta Chakrabarty, et al., “Evolution of the Light Organ System in Ponyfishes (Teleostei: Leiognathidae),” Journal of Morphology 272, no. 6 (2011): 704–21. 10. J. W. Hastings, “Light to Hide By: Ventral Luminescence to Camouflage the Silhouette,” Science 173 (1971): 1016–17. 11. Williams, Plan, 14, 2. 12. Williams, Plan, 15. 13. Williams, Plan, 16, 150. Williams probably acquired this Popper extract as reprinted in Studies in the Philosophy of Biology: Reduction and Related Problems, edited by Francisco Jose Ayala and Theodosius Dobzhansky (Berkeley : University of California Press, 1974): 259–84. 14. Williams, Plan, 16.

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15. Bradley A.  Thayer, “Evolution and the American Social Sciences: An Evolutionary Social Scientist’s View,” Politics and the Life Sciences 23, no. 1 (December 7, 2004): 2–11, 8. 16. Williams, Plan, vii, 17. 17. Williams, Plan, 17. 18. Williams, Plan, 18, 19. 19. Williams, Plan, 19. 20. Williams, Adaptation and Natural Selection (Princeton, NJ: Princeton University Press, 1966), 253. 21.  See Herbert Gintis, Game Theory Evolving: A Problem-Centered Introduction to Modeling Strategic Interaction (Princeton, NJ: Princeton University Press, 2000), 155. 22. Williams, Natural Selection: Domains, Levels, and Challenges (New York: Oxford University Press, 1992), 56. 23. John Maynard Smith, Evolution and the Theory of Games (Cambridge, UK: Cambridge University Press, 1982), vii. 24. “The Crafoord Prize 1999,” The Crafoord Prize website (February 11, 1999), https://www.crafoordprize.se/press_release/the­crafoord-­prize-­1999. 25.  George C.  Williams, “When Does Game Theory Model Reality?” Behavioral and Brain Sciences 7, no 1 (1984): 117. 26. Williams, Natural Selection, 58. 27. Williams, Plan, 52. 28. Williams, Plan, 52–3, 53. 29. Williams, Plan, 54. 30. Williams, Plan, 54, 55. 31. Williams, Plan, 55, 56–7. Williams’s emphasis. 32. Williams, Plan, 57. 33. Williams, Plan, 57–8. 34. Charles McCarry, The Last Supper: A Novel (New York: Dutton, 1983), 119. 35. Ullica Segerstråle, Nature’s Oracle: The Life and Work of W. D. Hamilton (New York: Oxford University Press, 2013), 289. 36.  Williams, “Pleiotropy, Natural Selection, and the Evolution of Senescence,” Evolution 11 (1957): 398–411, 403. 37.  Williams, “Preface (1996),” in Adaptation and Natural Selection, Princeton Science Library Edition (Princeton, NJ: Princeton University Press, 1996), ix. 38. Kim Sterelny and Paul E.  Griffiths, Sex and Death: An Introduction to Philosophy of Biology (Chicago: University of Chicago Press, 1999), xii. 39. Williams, Plan, 152.

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40. Frans Roes, “A Conversation With George C. Williams,” Natural History 107 (May 1998): 10–13. 41. Williams, Plan, 158. 42. Williams, Plan, 159–60. 43. Williams, “Reply To Comments On ‘Huxley’s Evolution And Ethics In Sociobiological Perspective,’” Zygon 23, no. 4 (December 1988): 437. 44. Holmes Rolston III, Genes, Genesis and God: Values and their Origin in Natural and Human History (Cambridge, UK: Cambridge University Press, 1999), 264–72, 271. 45.  See Paul Lawrence Farber, The Temptations of Evolutionary Ethics (Berkley: University of California Press, 1994). 46. Raymond Chandler, “The Simple Art of Murder,” in The Simple Art of Murder (Boston: Houghton Mifflin, 1950). 47. Dashiell Hammett, The Continental Op (New York: Vintage Crime/ Black Lizard, 1975). Dashiell Hammett’s anonymous, nameless, faceless Continental Op pretends to be unimaginative and avoids attracting attention to himself. The Op is said (in literary terms) to be a “realistic” detective, a working man who quietly completes one task before going on to the next. When people burst out with stories when he interviews them, he doesn’t ignore these self-­ justifications, but listens passively, keeps his doubts to himself, and seeks what was omitted from the stories. 48. Chandler, “Simple Art.” 49. Williams, Plan, 161. 50. E. O. Wilson, “Human Decency Is Animal,” New York Times Magazine (October 12, 1975): 48–50. 51. E. O. Wilson, Sociobiology: The New Synthesis (Cambridge, MA: Harvard University Press, 1975), 4, 562. 52. Wilson, Sociobiology, 3. The bracketed portion of this passage is the part Williams quotes in Plan, 161. 53. Williams, Plan, 161. 54. Williams, Plan, 167. 55. George C. Williams, “Altruism,” in The MIT Encyclopedia of the Cognitive Sciences, edited by Robert A. Wilson and Frank C. Keil (Cambridge, MA: MIT Press, 1999): 12–14, http://ai.ato.ms/MITECS/Entry/williamsg.html. 56. Williams, Plan, 163. 57. Matt Ridley, The Origins of Virtue (New York: Viking Press, 1996). Robert Wright, The Moral Animal: Why We Are the Way We Are (New York: Vintage Books, 1994). 58. Williams, Plan, 39, 42, 51.

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59. Robert J. Richards, Darwin and the Emergence of Evolutionary Theories of Mind and Behavior (Chicago: University of Chicago Press, 1987), 73. 60. Williams, Plan, 164. 61. Williams, Plan, 164–66. 62. Williams, Plan, 167. 63. E.  O. Wilson, interview, “Divisive Ideas on ‘Unification,’” Los Angeles Times (July 9, 1998). 64. T.  H. Huxley, “Draft of Manchester Address,” (1887) is taken from Adrian Desmond, Huxley: From Devil’s Disciple to Evolution’s High Priest (Reading, MA: Helix Books, 1999), 558. 65.  Steven A.  Frank, “George Price’s Contributions to Evolutionary Genetics,” Journal of Theoretical Biology 175 (1995): S373–388, S 387. See also Oren Harman, The Price of Altruism: George Price and the Search for the Origins of Kindness (New York: Norton, 2010). 66. Albert Camus, The Myth of Sisyphus and Other Essays, translated by Justin O’Brien (New York: Vintage, 1955), v. 67. Camus, Myth of Sisyphus, 88, 91. 68. William Hamilton, Narrow Roads of Gene Land: The Collected Papers of W. D. Hamilton, vol. 1 (New York: W. H. Freeman, 1996), 320–23, 174. 69. Williams, Plan, 154. 70. Raymond Williams, “Ideas of Nature,” in Problems in Materialism and Culture (London: Verso, 1980), 67–85. 71. Williams, Adaptation, 34. 72. Rachel Carson, A Sense of Wonder (New York: Harper & Row, 1984), 45. 73. Herman Melville, Chapter 93, and Chapter 42, “The Whiteness of the Whale,” of Moby-Dick; or The Whale (1851). 74. Herman Melville, letter to Nathaniel Hawthorne, November 17, 1851. 75. Charles Darwin, On the Origin of Species, 6th edition (1872), 104–5. 76. W.  Ford Doolittle, “The Practice of Classification and the Theory of Evolution, and What the Demise of Charles Darwin’s Tree of Life Hypothesis Means for Both of Them,” Philosophical Transactions of the Royal Society B 364, no. 1527 (August 12, 2009): 2221–28. 77. The philosopher Joel Velasco has explored this problem vividly and extensively. See, for instance, Joel D. Velasco, “Species, Genes, and the Tree of Life,” The British Journal for the Philosophy of Science 61, no. 3 (2010): 599–619. See also Joel D. Velasco, “The Tree of Life,” in The Cambridge Encyclopedia of Darwin and Evolutionary Thought, edited by Michael Ruse (Cambridge, UK: Cambridge University Press, 2013).

CHAPTER 11

A Retrospective: How Shall a Human Face the End?

A Sociobiology of Writing Writing is at least as complicated as any other human behavior. So Williams’s themes, his ways of exploring them, his changes in perspective, and his rhetorical turns are part of a complex set of behaviors. Though I do not claim to have accomplished a definitive reading of his texts, surely this attempt demonstrates a task worth undertaking, not just for Williams, but also for other evolutionary theorists. I might say, echoing him, that many readers may have already concluded that I do not understand these texts, in themselves or in their context, but “at least I can claim on the basis of conflicting views in the recent literature, the consolation of abundant company.”1 Commentaries on Williams’s contributions to sociobiology reveal a constellation of understandings and misunderstandings. Yet sociobiology has always seemed to be in theoretical disarray, primarily because diverse people write it. As early as 1981, Sarah Blaffer Hrdy argued—by citing a critique made by Mildred Dickermann in 1979—that though there has been a tendency to identify the field with Wilson’s single monumental book, “there is no monolithic body of knowledge identical with the field of sociobiology. Rather, there is a wide divergence of opinion on scores of issues among those who would call themselves sociobiologists.”2

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 M. P. Cohen, George C. Williams and Evolutionary Literacy, Literatures, Cultures, and the Environment, https://doi.org/10.1007/978-3-031-11650-6_11

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Many of those who chose to study sociobiology were guided by Williams, who himself never sought a grand synthesis. In 1980, he wrote, “sociobiology can be defined as the study of social behavior, a set of observable phenomena. Thus its scope would be determined by the phenomena dealt with, not by principles used in ordering these phenomena.”3 He continued to argue that it is more useful for an investigator to understand the grounds for investigation than to construct a new synthesis. Anyone interested in scrutinizing what E. O. Wilson calls the “methods developed in the natural sciences” could find how Williams tested their strengths and discovered their limitations, how he focused especially on conflicting views and paradoxes presented by the problems of evolutionary theory. His integrity is itself a model; his books are useful because he dissects theory, but not in order to kill it. He teases ideas apart, but also shows a better way to pull them together. Readers sometimes lose their patience with this approach, but they learn from it. What he contributed can be seen in the archives of the Quarterly Review of Biology, to which he devoted decades. It can be seen in the citations and acknowledgments in books by scholars like Sarah Blaffer Hrdy and many others. His influence is most obviously evident in a set of papers by students and scholars he influenced. Williams risked saying witty things like “I predict that no extraterrestrial biota will ever be found in which the universal principles of mechanism, natural selection, and historicity will not prevail.”4 For, as he argued, “the truth and usefulness of biological theory must be evaluated on the basis of its success in explaining and predicting material phenomena,” making important discoveries, and predicting the outcome of investigations. As applied to humans, prediction followed this path: “By suggesting the functional significance of some known aspect of human biology, you may logically be able to predict some other, unknown aspects.”5 Of course there are bad predictions and good ones. Sociobiology is grounded in the evolution of behavior, and one must make a distinction between kinds. I find it reasonable to take Melvin Konner’s distinction that sociobiology does not study all kinds behavior, but is “the application of natural selection theory to the explanation of reproduction, especially the behavioral aspects of reproduction.” As Konner goes on to note, “anyone who investigates or writes about behavioral biology without recognizing the potential for grave misuse of it, proven many times in the last two centuries, is either a dangerous charlatan or a dangerous fool.”6 No doubt, there has been a bit of bad behavior

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during the era of Williams, some among scientists and more among popularizers. A great many people have made use of the writings of Williams, variously interpreting his texts to establish authority for their own work, depending on their own needs and desires, sometimes suiting themselves and justifying their views. So one has to be careful. This is what happens when a man gains influence, even if that fame is hidden in the halls and laboratories of universities, but it might also be related to the Brockman effect—in an age of “science for the easily bored.”7

Williams Symposium 2004 On April 4, 2004, a symposium was held at Stony Brook University to celebrate Williams’s contributions to evolutionary biology, ecology, and medicine. Essays based on the talks given were published in the Quarterly Review of Biology, where he had been an editor for over thirty years.8 Williams invited colleagues, students, collaborators, and friends, including Frank C. Erk, Steven P. Ferraro, Helena Cronin, George W. Barlow, Mart R.  Gross, Mary Jane West-Eberhard, Martin Daly, Margo Wilson, Randolph M. Nesse, and Bobbi S. Low. David Haig and Sarah Blaffer Hrdy were also present but did not publish papers. Some of these people, like West-Eberhard, engage in straight evolutionary biology, and others, like Bobbi Low, do what has come to be called applied evolutionary biology, or evolutionary application, for short. This community created by Williams crosses disciplines and includes biologists, ichthyologists, psychologists, a psychiatrist, behavioral ecologists, resource managers, and at least one philosopher. As with any celebration, mostly the speakers praise Williams. For instance, Helena Cronin, who has largely been engaged in defending the gene-centered view, sets a certain tone by announcing, “George has given us not only a gene-­ centered understanding of adaptations but also a gene-centered theory of environment.” She connects the gene-centered view, adaptations, and “the richer and more complex environments they rely on,” though I am not entirely sure I know what that means.9 Others complicate this picture. Some speakers who were very close to Williams for a long time describe the long and elaborate institutional growth of Stony Brook, engage in personal reminiscence, remember Williams as a generous teacher who humorously exposed his own failures to a student, critiqued his own experimental methods, which he often considered sloppy, and deferred from including himself as a coauthor of academic papers by his students.

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They remember thought experiments learned, so that, as one said, “my brain was rewired to make me think more like him.”10 As one researcher put it, “Williams’s simple model exposed departures, which is the mission of models.”11 One describes “Williams’s Principle” as stating “that reproduction has not only a benefit but also a cost to lifetime fitness,” and so only by understanding biological causes can “human society adequately address such issues as parenthood, birth control, abortion, adoption, postpartum depression, child abuse, and child neglect.”12

West-Eberhard and a Transition from Williams’s Principles There was also serious critique interlaced with personal response. “One of the fascinating observations about scientists is the way personality, or personal character, affects the research one does.” So Mary Jane West-­ Eberhard begins her paper “The Maintenance of Sex as a Developmental Trap Due to Sexual Selection.” As for Williams, “I suspect that he is suspicious of everything he reads, especially if people are starting to believe it. He will lurk there, cultivating his doubts until he finds the pivotal word, and then he will spring it on us.” She sees “a rugged individualist if ever there was one,” “a quiet, even a sneaky, revolutionary,” and a “soft iconoclast,” saying that “he would rather resist than follow a tide, but he does not shout about it.”13 This pleasant and humorous tone allows her to pursue what she calls a stalemate between their divergent theories of the origins of sex.14 As one who does not appreciate fads and distrusts expressions like “cutting-edge” when used for science, she values Williams’s work because it provides challenges, not answers.15 She prefers speaking of insects, not humans, and discussing science with scientists. Williams welcomed professional challenges to his work. Indeed, the manuscript for Natural Selection (1992) received close reading by West-­ Eberhard and by David Sloan Wilson. While West-Eberhard disputes the notion that social behavior can be understood only as “genetical,” as in the title of Hamilton’s 1964 articles, she also believes that Hamilton’s formulations were being used as a model for her field, developmental biology, and that Williams’s explication of them made him a foundational evolutionary theorist.16 Williams in turn wrote a glowing recommendation for the back cover of her book Developmental Plasticity and Evolution (2003),

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saying it is “full of novel and convincing arguments,” and that it “will be a classic that people will be quoting decades from now.” She argues at the symposium that, along with recombination as a cause, “sexual reproduction is maintained as an adaptive legacy of sexually selected and other male manipulations that have produced a reproductive dependence of females on males.”17 In the early twenty-first century, and as early as 1989, Mary Jane West-­ Eberhard began to show how “plasticity” and “development” are related terms and concepts that “have passed through a period of neglect.” She defines phenotypic plasticity as “the ability of a single genotype to produce more than one alternative form of morphology, physiological state, and/ or behavior in response to environmental conditions.”18 As she has argued more recently, “If the twentieth century was the age of the gene, the twenty-first century promises to be the age of the environment, and of gene expression (all of which is environment dependent).”19 Her vigorous work helped bring about that promise and, by 2003, she outlines a new synthesis in over 600 pages of text, beginning with “Gaps and Inconsistencies in Modern Evolutionary Thought.”20 In Developmental Plasticity and Evolution, West-Eberhard finds inadequate Williams’s and Dawkins’s arguments “that selection acts on genes or that genotypic differences are required for selection to occur. This has led to confusion regarding cause and effect in evolution.” As she asks, “If recurrent phenotypes are as much a product of recurrent circumstances as they are of replicated genes, how can we accept a theory of organic evolution that deals primarily with genes?” Indeed, her book is meant to replace the “unimodal adaptation concept,” as she calls the gene-centered view, whose prevalence “shows the power of an internally consistent set of ideas to reinforce a single line of thought, even when this means overlooking a large number of contradictory facts.” Yet, in the book she acknowledges “George Williams, whose approach to adaptive evolution I thought would be more different from mine than it turned out (happily) to be, and who went out of his way to travel to places where we could discuss the issues that arose.”21 This may be an overly optimistic view of their concord, considering how close he came to speaking of all discussions of plasticity as nonsense in 1992 when he suggested “that plasticity be abandoned entirely.”22 West-Eberhard is frustrated especially by a language of biology that leads to confusions about nature and nurture, and a failure to appreciate the complex process of relations between genotype and phenotype:

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Among the consequences of neglect of mechanisms in modern evolutionary biology are the problems that arise when the black box of mechanism is filled with imaginary devices. By imagining that there is a one-to-one relationship between genes and phenotype, for example, biologists lose sight of the role of the environment and polygenic influence in development and evolution.

As she notes, while Hamilton’s work uncovered “the genetical evolution of social behavior,” by writing a phenotype-centric view, she stands the genotype-phenotype distinction on its head, where “genes are followers, not leaders, in evolution.”23 As a counterpoint to Williams’s attempt to rid biology’s language of certain excesses, West-Eberhard emphasizes, “The conceptual vacuum caused by lack of an adequate theory of development and evolution has been filled by metaphors, such as genetic programming, blueprints for organisms, and gene-environment interactions.” For instance, “for Dawkins (1982), the genome is a recipe; the phenotype, a cake. A less lifelike image for the dynamic activity of genes as they participate in development can hardly be imagined.” This kind of writing and thinking, she believes, has led to incompleteness and inadequacy of sociobiological explanations.24 Her approach has been infectious for sociobiologists who focus on primates, particularly in the writing of Sarah Blaffer Hrdy.25 In Mother Nature (1999), Blaffer Hrdy recalls a lecture in which West-Eberhard said, “Nothing is genetically determined in the sense of determined by genes alone. No gene is expressed except under particular circumstances… . It’s a kind of biological illiteracy to talk about a gene for anything other than a particular protein molecule.” Hrdy had been convinced, and consequently her views coincided: “No gene or set of genes, or even any one mechanism influencing people to favor kin, has been identified. We do not know even a fraction of the ways that kin selection works.”26 The point for Blaffer Hrdy is precisely that childbearing and child rearing, for instance, must be understood as a social process that is also a developmental process. West-Eberhard’s views also broaden the questions that ought to be asked by those interested in Darwinian medicine so they include developmental issues, and she casts considerable doubt on what she calls the strong distinction between proximate and ultimate causes so prominent in discussions like those of Nesse and Williams. Resistance to ideas of development can be traced to this distinction that values ultimate questions

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over proximate ones, but as West-Eberhard puts it, “proximate mechanisms are excluded from ultimate (evolutionary) explanations, yet a proximate mechanism (development) produces the variation that is screened by selection.” Proximate causes matter.27 Last but not least, the conclusion of West-Eberhard’s book, entitled “One Final Word: Sex,” examines the maintenance of sex by framing the problem first in exactly the terms Williams uses in 1975. In so doing, she demonstrates his foundational contribution to all discussions of sex because he clarified the cost-benefit or tradeoff terms in which biological problems like the maintenance of sex can be most profitably pursued. What she wishes to add, looking backward and forward, is an argument Darwin made, that “there are reproductively essential social factors that can exaggerate and maintain extravagant traits like sex.”28 The social can and does alter the biological, which is why her lecture for the Williams symposium in 2004, based on this final chapter, was originally titled “What George Williams Said about Sex but Didn’t Really Believe.”29 As she puts it, it was not really the maintenance of sex that concerned him most deeply, but the primary function of sex, what sex is designed by natural selection to do.

Practical Applications Martin Daly and Margo Wilson follow Williams more closely. They focus on Williams’s concept of “future discounting,” seeing that “expenditure of reproductive effort constitutes a sort of gamble” and asking, “When is that gamble worth taking?” As they summarize, the “barely justified cost” of any act must be “(1) directly proportional to its expected positive impact on current reproduction (‘the magnitude of the threat or opportunity’); (2) directly proportional to the proportion of reproductive value that is immediately at stake; and (3) inversely proportional to the RRV [residual reproductive value] that will remain if the gamble is refused.”30 Bobbi Low prefaced her book Why Sex Matters (2000)—the manuscript of which Williams read—with an epigraph from William James in which a crab would not think of itself as a crustacean, but would only say, if it could speak, “I am MYSELF, MYSELF alone.”31 She uses the concepts of individualism and maladaptation when she engages in evolutionary applications of Williams. Low’s title for the symposium, “George Williams, Theoretician and Guerilla Environmentalist,” reveals a strategy that uses both his critique of Gaia theory and his more general perspective

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on adaptation. So when Low praises Williams’s critique of the “Gaia Hypothesis,” she adds her own of the related contemporary version of a complicated and historically interesting concept of the “Noble Savage.”32 Low considers the “Noble Savage” a myth or “paleofantasy,” though she does not use this term coined by anthropologist Leslie Aiello; it is borrowed by Marlene Zuk, who makes a similar argument.33 Like behavioral ecologist Zuk, who worked with William Hamilton at the University of Michigan, Low is willing to go another step and critique ecofeminism as a third, purely mythical concept, a kind of “noble womanhood.”34 In this sense, Williams authorizes Low to say that “both the Gaia and Noble Savage hypotheses are romantic perceptions that are unlikely to help us in our efforts to manage resources so that we do not deplete them quickly.”35 Evolution has not produced any perfectly adapted species, least of all humans. There is something in this that touches on a premise of Why We Get Sick, of a “Stone Age Man,” evolved to fit an “environment of evolutionary adaptedness.” Nostalgia for any Edenic condition, as Adrienne Rich once wrote, “is only amnesia turned around.”36 Low preaches, “We do not need to become Noble Savages—no one ever has been. Instead, we need to understand conflicts and confluences of interest, the evolution of cooperation, and future discounting. Noble Savages were neither more ethical than we are today nor better at managing for the far future.”37 Low describes her work as falling into behavioral ecology and life history theory. Because, as Marlene Zuk explains, sociobiology became “sufficiently politically laden,” scientists began to speak of the evolutionary basis of behavior in animals as “behavioral ecology,” and the human counterpart as “evolutionary psychology.”38 Low practices a little of both. Like David Lack, she often focuses on the relationships between reproduction and food (or natural resources) and is less engaged by the links between sex and death. It is a reasonable step from Williams’s view of adaptation to her argument that “we evolved to be efficient short-term environmental managers, not long-term global conservationists.” As for imagining that hunters and gatherers lived in a harmonious and balanced state, “to infer a conservation ethic from a low impact lifestyle is a logically inappropriate reverse reasoning of the sort I suspect would drive George Williams to distraction.” “In contrast, whenever we can find low impact in conditions of high population density, highly developed technology, and high profit to exploitative enterprise, we can confidently infer a strong conservation ethic.” She concludes that Williams “laid out the groundwork” in

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“recognizing the inherent selfishness of all living things (no Gaia), the amoral nature of natural selection—but at the same time, the selective importance of cooperative behavior.”39 Some of the followers of Williams fall prey to an unfortunate, perhaps even fatal conflation of thinking about biology and human reproduction and human values. Nowhere is this better illustrated than when Bobbi Low tells this story: A great philosopher never married, and, in a possibly apocryphal story I read as a child, on his deathbed he called for all of his works to be set upon his lap. When the works had been brought, he sighed, “All of this is less than the weight of one grandchild.” (And, to complete the story tidily, he promptly died.) His insight is an important one: What are resources for, anyway, if not to build our families?40

The problem, beyond doubt, is nailed by Colin McGinn, who writes, “Low is interpreting her biological perspective as a theory of conscious human motives, trying to reduce them all to the motive of reproduction.”41 I do not believe that Williams falls prey to such reductions, but some of his followers do, as can be noticed in the frequency with which they refer to the number of children and grandchildren he produced, after noting the irony of his fatal malady, Alzheimer’s, but that gets ahead of the story.

D. S. Wilson’s Critique Williams’s legacy cannot be adequately summarized by following the speakers at a symposium in his honor, and it is neither possible nor advisable to present all responses to Williams’s writings here. But David Sloan Wilson was sufficiently close to Williams, and encouraged by him, that his views bear noticing. He believes that Williams, as critic of “naïve group selection,” became guilty of “naïve gene selectionism.” As he has recently written with Elliott Sober, “In Adaptation and Natural Selection, Williams is a robust individualist (a position he tempers in 1992). He denies that group adaptations exist,” and as they continue, “Williams’ Principle, as we will call it, says that adaptation at a level requires selection at that level.”42 For them, “individualist” is not a term of respect for a theory, or perhaps for a man.

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D. S. Wilson has gone over the salient points repeatedly. He writes that “Williams didn’t reject group selection because only genes are replicators, but because (according to his assessment) within-group selection invariably trumps between-group selection.” As he continues, “I want to acknowledge that if George’s empirical claim did turn out to be correct, it would be a very strong argument for individualism, which is the claim that adaptations exist only at the individual level; that individuals are the only units that can be called self-interested; that a group cannot be described as an organism writ large.”43 It is clear that more is at stake than levels of selection. According to Elizabeth Lloyd, this is a matter of belief. She argues that Williams advanced an engineering definition of adaptation and “believed that it was possible to have evolutionary change result from direct selection favoring a trait without having to consider that changed trait as an adaptation,” and thus drove “a wedge between the notion of a trait that is a direct product of a selection process and a trait that fits his stronger engineering definition of an adaptation.”44 As Lewontin wrote more dispassionately in “The Units of Selection” (1970): “Although Williams demonstrated that we do not need population selection to explain evolution and although I have demonstrated that in most instances it would be an inefficient process, there is still the possibility that population selection does occur.”45 Possibility is not the same as probability, or importance. The controversy over levels of selection continues. In 2007, David Sloan Wilson and E. O. Wilson coauthored an article whose purpose was to restore clarity and unity to sociobiological theory. Their premise is that current sociobiology is in theoretical disarray, with a diversity of frameworks that are poorly related to each other. Part of the problem 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.

They blame the effectiveness of Williams’s Adaptation and Natural Selection for this predicament, and “the tenor of the times, which seemed to make the rejection of group selection a pivotal event in the history of evolutionary thought.”46 As they indicate, neither biology nor sociobiology has ever truly recovered from Williams’s critique, which preceded Wilson’s Sociobiology by some nine years.

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Indeed E. O. Wilson has defended and refined his concept of sociobiology over the years and attempted once more to create a grand synthesis in Consilience: The Unity of Knowledge (1998). His premise is that “the greatest enterprise of the mind has always been and always will be the attempted linkage of the sciences and humanities.” Consilience is a manifesto for that enterprise. The book’s purpose is to argue that “there is intrinsically only one class of explanation,” and “the only way to either establish or refute consilience is by methods developed in the natural sciences.”47 Williams saw no such synthesis coming.

Ecology Adaptation and Natural Selection impinged on both ecology and evolution, “tightening,” as the philosopher Gregory J. Cooper puts it, the relationship between the two when it dismissed loose talk of higher-level processes.48 The Williams critique called for considerable reassessment of the language and concepts of ecology, such as struggle, community, and balance, and cast some doubt on the accuracy of terms like ecosystem and community.49 When Williams used these terms in 1966 and 1992, he was careful to make sure that the reader understood them as terms for mechanism and especially cost-benefit mechanisms enacted by individuals. His work called for soul-searching by ecologists and triggered the creation of a new evolutionary ecology. In “Reflections on Reflections: Ecology and Evolutionary Biology” (1986), Douglas J. Futuyma traces the historical roots for the “new evolutionary ecology,” roots that come not only from the legacy of the “evolutionary synthesis” where “natural selection is not only real, it is powerful.” Futuyma names R. A. Fisher, Sewall Wright, and J. B. S. Haldane, who “had rescued natural selection in theory,” and Ernst Mayr, George Gaylord Simpson, Julian Huxley, and G. Ledyard Stebbins, among others, who “had found it compatible with the observations of systematists.” Theodosius Dobzhansky figures in this change, but “the British school of ecological genetics … helped to convince a generation of biologists that selection shapes all ends.”50 As Futuyma reminds the reader, when “debate over the nature of population regulation had become one of the central issues in ecology,” it found on one side David Lack’s “individual-selectionist interpretation of fecundity and territorial behavior,” and on the other the group selectionist interpretation of Wynne-Edwards. As Futuyma says, it was no contest. A

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generation had learned the essence of neo-Darwinism, especially from George C.  Williams, William Hamilton, and John Maynard Smith.51 As Stearns summarizes the Williams position, “All the processes observed in ecosystems are the aggregate consequences of features, such as photosynthesis and predation, that are advantageous to individual organisms.”52 Futuyma underlines the influence of Williams, whose “appeal lay in his clear exposition of ecological issues in a genetic framework.” Between 1966 and 1992, Williams began to emphasize historical constraints for individuals and ecological systems. Yet in practice, Futuyma found in 1986, “evolutionary ecologists tend to be almost as ahistorical as ecologists who concern themselves not at all with history but only with ‘things as they are.’”53 It is not surprising that ecology has been polarized over levels of selection. Some of the complexity of positions in ecology can be seen by perusing the recipients of the Eminent Ecologist Award, which the Ecological Society of America grants annually to a senior ecologist for an outstanding contribution to the science of ecology. This list includes Henry Allen Gleason, who endorsed an individualist view and cast doubt on traditional notions of associated species, so that “the phenomena of vegetation depend completely upon the phenomena of the individual” (1959 and 1953), and Charles Elton, who investigated the functional structure of a niche and the disruption of ecosystems by invasion (1961). Elton was famous for writing, “When an ecologist says ‘there goes a badger,’ he should include in his thoughts some definite idea of the animal’s place in the community to which it belongs, just as if he had said, ‘there goes the vicar.’”54 Robert MacArthur (1973) preceded Eugene Odum (1974), George C. Williams (1989), and more recent awards went to E.O. Wilson and Paul Ehrlich.55 One notices in this list proponents of dueling paradigms of ecology. For instance, Odum’s primary premise about ecosystem development was group selection, and he wrote his standard textbooks well after Williams wrote Adaptation and Natural Selection. Nevertheless, Williams’s ideas, which were taken up by younger evolutionary ecologists, helped undermine classical ecosystem theory. Older ecologists were not the only ones unhappy with Williams’s influence on the theory of ecosystems. Some environmental historians, like Donald Worster, became very unhappy with the disorderly and chaotic, disruptive and discontinuous dynamics of systems, an upshot of Williams’s thinking.56

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How Biologists React to the Williams Style When ecologist Laurence B.  Slobodkin, Williams’s colleague at Stony Brook University, wrote a review of Adaptation and Natural Selection for the Quarterly Review of Biology, he found it “an exciting, significant and important work,” but also confessed that “analysis of Williams’ position is made somewhat more difficult by his style.” Unlike other scientists, for whom “words are used as a kind of casing for data tabulations, equations, or pictures,” Slobodkin observed, “Williams, however, believes in words.” When Williams writes that “the most urgent requisite for the success of any science is that it have a name,” Slobodkin concludes, “this may be meant as a joke, but it is perhaps of interest that, given the concern with words that characterizes the rest of the book, I can’t really tell whether it is meant to be funny or not.”57 Some scientific colleagues thought that Williams had a cold and piercing style, though views of his style have softened since his death. According to his own statements, his purpose was not, at least originally, to please or to generate expectations, but to teach, critique, and especially to warn. How his style became a model in itself—of clarity and simplicity, embellished with a characteristic corrosive wit—can be seen in the opinions of his colleagues, who tended to conflate the style with the man. It is probably fair to say that they speak primarily of a style he established with Adaptation and Natural Selection. Robert Trivers wrote, “He was a beautiful man, with a very simple and clear style of thinking, in a warm and humble personality.” Richard Dawkins called him “universally respected, … a tall, shy, diffident scholar with legendary wisdom.” Daniel Dennett noted “that someone so incisive, such a corrosive corrector of sloppy thinking, could be so gentle and generous is something of a marvel. He was a lover of truth, not a lover of his own perspective and reputation.”58 My point is that they noticed his style. For his converts and followers, Williams seemed nearly godlike. Steve Pinker wrote, “In my mind, George Williams is one of the most brilliant writers in the history of science. His 1966 book Adaptation and Natural Selection was way ahead of its time.” Even Stephen J. Gould showered him with praise: “He’s always been at the forefront of theoretical clarity in the field.”59 For Niles Eldredge, “George really is the most important thinker in evolutionary biology in the United States since the 1959 Darwin centennial. It’s astonishing that he hasn’t gotten more credit and acclaim. He’s a shy guy, but a very nice

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guy, and a very deep and a very careful thinker. I admire him tremendously, even though we’ve been arguing back and forth for years now.” Lynn Margulis, though ambivalent about his influence, said, “People didn’t connect evolutionary thinking and mammalian behavior. Williams is credited for recognizing the importance of reproduction in mammalian behavior. He’s communicating a scientific truth in a resistant cultural milieu. The fact that he has few articulate predecessors enhances the importance of his work.”60 After noting that Williams developed “the theoretical groundwork for Dawkins’s vision,” Peter A.  Corning calls Adaptation and Natural Selection a “classic volume” and “a therapeutic cold-bath that had the salutary effect, overall, of purging evolutionary biology of some fuzzy thinking.” Yet Corning was troubled by Williams’s “extreme reductionist view” and his attacks on other views as fantasies or products of “romantic imagination.” What’s worse, “Williams burnished his curmudgeon image by heartily endorsing Huxley’s ‘tooth and claw’ vision of evolution.”61 From Trivers, Dawkins, Dennett, or Pinker, one can see how admiration of Williams’s style seems nearly congruent with approval of his perspective, while those who disagreed with Williams describe his style of writing in more ambivalent terms. For many, like Slobodkin, the style was too slippery. Though it was fairly clear that “group selection” was “demolished,” scientific readers who found the tone puzzling did and did not know exactly what position Williams took, if indeed he took a position. This is probably a universal problem for those who write critique, even in academic circles.

Self-Suspicion and Altruism As Thomas Shepard wrote in The Sincere Convert, “Suspect thyself much, and when, in this shipwreck of souls, thou seest so many thousands sink, cry out, and conclude, It is a wonder of wonders, and a thousand and a thousand to one, if ever thou comest safe to shore.”62 Nobody comes safe to that shore. If Thomas Shepard offers a Puritan and Calvinist theology of self-suspicion, modern theories of naturalistic epistemology also point toward the fragility and limitation of human understanding.63 To some extent, appreciating Williams’s skills requires understanding some things about cognition. Just as he suspected perfection and was interested in the limitations of organic design produced by natural selection, so one must also be interested in limitations to human thinking that

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come from constraints on the capabilities of human brains, perhaps from negotiation between hemispheres and a brain’s capacity to make mistakes. The brain is, Williams might say, maladapted for clear critical thought.64 Lists of cognitive biases have run into the hundreds, and there is really no limit to them. It is enough to say that Williams has been concerned with particular cognitive biases tied to imagining certain kinds of higher-order control in evolutionary processes—biases that ignore the limitations of design in evolutionary processes. But there are more general reasons for reading Williams. Most important, to learn what a critical intellect is, and how it works. Frans de Waal has argued long and hard that morality is “natural” and that those who disagree are Calvinists who believe in innate depravity. In Good Natured: The Origins of Right and Wrong in Humans and Other Animals, he asks, “So why should we let the ruthlessness of natural selection distract from the wonders it has produced?”65 I mention this because, in his desire to attribute morality to other animals, de Waal picks an argument with Williams in particular, though among the usual company of Bill Hamilton and Richard Dawkins. For over a decade now, de Waal has strenuously opposed Huxley’s view, which Williams endorsed and made more radical—a view de Waal names “veneer theory.” As he says, for Williams morality must be seen as a very thin “cultural innovation achieved by our species alone” by which humans supposedly “became moral by choice,” though we are not always so. Yet saying this sort of thing, de Waal insists, “assumes that deep down we are not truly moral. It views morality as a cultural overlay, a thin veneer hiding an otherwise selfish and brutish nature.”66 More than a few of de Waal’s critics have responded, and Philip Kitcher’s response is perhaps most useful. At large, however, de Waal joins Elliott Sober, David S. Wilson, George Price, and others in the quest for natural goodness.67 One can ransack natural processes, seeking the roots of human altruism, but to do so, says Williams, would be looking for human virtue in the wrong place. Humans will have to look for social and ethical order in our own culture; we will have to create sturdier social virtues because they often must work in opposition to the processes of biology. Once again, Williams would notice the way aesthetic distraction—looking for wonders—becomes dangerous because much of the beauty that humans perceive in the natural world hides violence. No wonder he uses satire for such a distracted perspective. Also he considers the matter of proportion: in terms of the overwhelmingly monumental total mass of

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life, how many moral animals are there? Surely the process of evolution cannot be deduced from similarities or differences in the behaviors of primates, or even of mammals. Humans are and are not ruled by their biological origins. But most creatures are entirely ruled by their origins and adaptations. Williams was full of self-suspicion, set high standards for himself, and spoke deprecatingly of his own work, sometimes with displeasure, and sometimes he simply spoke of its transience. In recalling an early essay, “Natural Selection of Individually Harmful Social Adaptations among Sibs with Special Reference to Social Insects,” as compared to Hamilton’s two essays, “The Genetical Evolution of Social Behavior” I and II (1964), he wrote, “my wife and I had published a clumsy treatment of a related topic, natural selection among nuclear families. It was a relief to have our ideas replaced by Bill’s simple proposal.”68 When Williams spoke of Adaptation and Natural Selection, he could only say, “I once wrote a little book for a university press, but it was thirty years ago.”69 When Williams reviewed Maynard Smith’s book, The Evolution of Sex, he wrote that it made his own work nearly obsolete—but he also argued at large that Maynard Smith’s proposals would also fail the test of time. What counted, he asked, or mattered? “Its lasting value will lie not so much in the validity of answers suggested as in questions formulated and confronted.”70 He could have said the same thing of his own work. So perhaps one can understand why, in Plan and Purpose, he speaks of Why We Get Sick as discussing the implications of modern Darwinism to medicine in “grossly inadequate” detail.71 However, to claim that he was overly modest is not to take him at his word. His word is Critique.

Age and Youth Williams’s last major essay is “The Tithonus Error in Modern Gerontology,” his Crafoord Prize lecture of 1999.72 He stood by John Maynard Smith and Ernst Mayr, between two older men given the prize. The lecture has been interpreted as recommending a program of research to prolong human life or “slow down the clock.” That is a rather optimistic version of Williams’s message. The abstract for “The Tithonus Error” reminds a reader that “Tithonus asked Aurora for eternal life, when he meant eternal youth.” Once more, Williams chooses an epigraph from the Victorian Era, from Tennyson’s

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poem, composed first in 1833, but achieving final form and publication in that magic and frightening year, 1859, the year of The Origin. The epigraph from Tennyson begins with Tithonus remembering that he asked Aurora, the goddess of dawn, for immortality, who “Then didst thou grant mine asking with a smile.” The epigraph ends on a sadder note: Of happy men that have the power to die, And grassy barrows of the happier dead. Release me, and restore me to the ground.

What we desire—and the questions we ask—determines what we can receive, and errors in mistaken desire and asking the wrong questions are legion. Apparently, the error of Tithonus is a common error. T. S. Eliot provides another version of this error in the epigraph for The Waste Land (1922). Eliot took his text from the Satyricon by Petronius, of the Sibyl at Cumae, and though Eliot does not do so, the passage can be translated into English as, “For with my own eyes I saw the Sibyl hanging in a bottle, and when the young boys asked her, ‘Sibyl, what do you want?’, she replied, ‘I want to die.’”73 Like the story of Tithonus, this too is a tale of love, life, and gifts gone ironically awry. According to legend, the Sibyl offered her virginity to Apollo for years of life totaling as many grains of sand as she could hold in her hand. But, when she refused his advances, he gave her immortality but not eternal youth. She forgot to ask for eternal youth. Petronius’s story of the Sibyl seems the same as that of Tithonus as told by Tennyson, but it is not. Both figures, one mythological and one perhaps partly historical, err in desiring eternal life when they should also have asked for youth—if not eternal youth, then a longer lasting youth. The difference between the stories is not that Tithonus does not ask to die. They both outlive anything that makes life worth it—they outlive the minds that made life meaningful and the body that gave life joy. They are of different genders, both prisoners but in different ways. Both wish to end the agony. So too, the tone in which their stories are told differs. She is victimized and hangs, shrunken, in a bottle. He does not. He is a heroic figure, and more of a talker than she. She asked immortality for herself, and Aurora asked it for him. For the Sibyl there is an added irony, because she was a prophet and able to tell the future. Tithonus sees a future in which all things die while dawn arises each day, ever new. He sees the woods decay

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and fall. “And after many a summer dies the swan.” He is consumed, as he says, by his own immortality. Like the Sibyl, he asks for release and to be restored to the ground. It is doubtful that even the ancients believed that wishes for immortality could be granted. It is certain that no modern human enjoys the deterioration brought by aging in our era, where longevity is artificially extended. But these stories of the desire for immortality or eternal youth reveal what humans desire from their lives, though for how long, and to what effect? Why learn so much, one might ask, acquire such capabilities, when one can use these for such a short span of time, and then the capabilities decay. The body and mind decay. What a human desires is determined by contexts. Context determines when life ceases to be worth living. Williams shows more moderation than Eliot in selecting Tennyson’s rather than Petronius’s imagery. He is also moderate in his questions. He wants to see if it is possible to retard but not postpone senescence. He continues his abstract, “Modern gerontological research makes the same mistake in its preoccupation with death, as if it were a programmed event in an organism’s life history.” He also ridicules Biblical language, writing that “for gerontologists, the second part of the ‘a time to be born, and a time to die’ idea ought to die.”74 In “The Tithonus Error,” Williams offers a philosophical intervention, not only a technical discussion. Williams says, why not intervene into human biological nature before the process of deterioration begins? Nature, after all does not know best. Biology doesn’t care about the individual after procreation is completed. Humans can care, though, and take precautions. He denies that death is a programmed event in life history. He does not discourage postponing events such as death, but makes the important distinction that “continuous processes such as senescence can be retarded, not postponed.” He finds a second fallacy in “the wear-and-­ tear concept from analogies with manufactured machinery.”75 Processes can be studied and understood. Williams offers a program. “Gerontology ought instead to investigate senescence, the decreasing effectiveness of mechanisms by which adult organisms avoid death or loss of fitness.”76 An organism, as he notes, is a place where interesting processes occur. So with his typical candor, Williams announces at the end of his abstract, “The death of a studied organism must necessarily end its usefulness in providing valuable data. It is of little scientific significance.” What, then, is of significance? First of all, the arguments he has made about evolutionary ethics apply. Surely the program Williams recommends

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in “The Tithonus Error” constitutes exactly a practical way of acting upon Huxley’s understanding “that the ethical progress of society depends, not on imitating the cosmic process, still less in running away from it, but in combatting it.”

A Eulogy for Bill Hamilton If there is an irony in the Tithonus essay it is that the second section is entitled “The Moulding of Senescence by Natural Selection,” named after the title of William D. Hamilton’s paper of 1966. With his usual modesty, Williams comments that Hamilton “provided the beginnings of rigorous thoughts on the evolution of senescence,” and he only provides a “brief verbal outline” of Hamilton’s argument. According to Hamilton’s view, as expressed by Williams, the optimal genotype “will be that which programs an optimal distribution of its finite fitness supply over the life cycle.” Natural selection will produce and maintain adaptation at any age proportional to the product of survivorship and reproductive value. Though an individual human’s genetic success might require survival through its thirtieth year, “Avoidance of death in the ninetieth year? Worthless.” This means that modern human longevity constitutes what Williams calls “‘manufactured time,’ with the implication that people in their sixties or older are artificial.”77 Hamilton wrote in a commentary on his own paper in 1996, that Williams “seemed to have understood the matter right through” in 1957.78 That Williams anticipated his own thoughts is a consistent theme in Hamilton’s memoirs, because he thought of Williams as like a lost older brother. Hamilton had received the Crafoord Award six years earlier, in 1993, and he died March 7, 2000, three months after the Tithonus essay appeared in print, probably as a complication of malaria. According to the biographical memoir by Alan Grafen, “Bill contracted malaria in the Congo, collapsed with hemorrhaging soon after returning to London, and died without fully recovering consciousness a few weeks later, at the age of only 63 years.”79 What Williams thought of Hamilton is revealed by an almost laconic one-page memorial, “Some Thoughts on William D.  Hamilton (1936–2000),” published in July 2000 in Trends in Ecology and Evolution. He writes, “One thing I can honestly brag about is my early recognition of Hamilton’s immense importance.” He remembers, “that the youthful Hamilton could be so far ahead of almost all living biologists on such

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important issues was mighty encouraging.” Williams notes that he particularly admired Hamilton’s “Geometry for the Selfish Herd” (1971), but he puts Hamilton’s thesis in strange terms: “the main reason for gregarious behavior in animals is to maximize competition for the bad things in life.” This is because, for instance, faced with a predator, “each will have companions to compete for the wolf’s attention and, therefore, each will be a less likely victim, even without such benefits as alarm calls or collective alertness.”80 The bad things in life. Like malaria, whose etymology is “bad air.” “There are other bad things in life that gregariousness can mitigate,” as Williams continues. If, for instance, there is a mosquito in the room bearing West Nile virus, the largest crowd would provide the safest environment. The good thing was to have Hamilton around, and not simply to take the sting. Perhaps the strangest and most touching part of his thoughts on Hamilton is that they conclude with conflict, with Williams wishing he could continue to argue with Hamilton about the evolution of sex—as he had been doing only a year ago. Ideas do not grow or reproduce, except by exchange and discussion. “I dislike one aspect of Bill Hamilton’s career—its brevity. That he will no longer be solving basic biological problems is sad news for biology.” Sad news: bad news. An internal rhyme. “Some Thoughts” is a startling piece of writing, almost a tone poem. Like Tennyson’s “In Memoriam,” it is an elegy constructed of a short swallow-flight of song that dips its wings in tears and skims away. It is beautiful in its awkwardness, strange and unassuming, with its almost clichéd decorous phrases, like “I took time,” “enjoy his hospitality,” or “most pleasant.” As with Tennyson, the narrator seems “all over the place,” yet by juxtaposition of short humble sentences, he suggests a pain he will only label as “the bad.”

A Digression on the Death of George C. Williams Whether Williams has any sense of humor about his given middle name, Christopher, literally “Christ-bearing,” or about his relationship to Hamlet’s soliloquy on human nature, or about his alliance with Huxley, one cannot tell from his writings. He becomes more contemplative and more literary as he gets older, as the essay on “The Tithonus Error” suggests. Did he transform himself into a literary character like Kurtz, who

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judged, according to Marlowe in Heart of Darkness, and spoke “The horror!”? Maybe. Who did he belong to? In “Tithonus,” Williams focuses on a small and universal horror, “that evolution is biased in favor of youth over age. A gene that increases youthful capacities even at a price to the survivors later in life, may often be favored.” He says finally that death is not as important as his own final entreaty, to seize the day. He knows that most people are resigned to the inevitable decline of several decades, so he asks, why not start earlier, to “direct more research into finding ways to prevent or at least retard the kinds of functional deterioration that we normally experience in our forties and fifties?” “Adding years to people’s lives is of course a worthy goal, but perhaps adding life to their years is even worthier,” as he concludes the essay.81 The end is inevitable. “Thus, though we cannot make our sun / Stand still, yet we will make him run.”82 What should humans fear, and what should they wish for? As is decorous, those who wrote remembrances of Williams after his death dwelled upon his family, his wife Doris and his four children, but in a real sense, his kin were those scientists with whom he thought. They were stunned by his senescence. Some, like Hamilton, taught him and learned to think with him about death, and sometimes even to think like him about this process, beyond inclusive fitness. As Williams thought, they were kin. It is hard to distinguish between reciprocal altruism and kin selection. As one commentator defines the problem, reciprocal altruism (RA) typically occurs between non-relatives, as distinct from kin selection (KS). “But because similar traits expressed by conspecifics are usually due in part to identical genetic determinants, individuals that engage in RA are likely to be doing so because of shared genes. This is true so long as there is any genetic determination of RA.”83 Samir Okasha also reminds us that “human behaviour is obviously influenced by culture to a far greater extent than that of other animals,” and one can, in theory, distinguish biological altruism from cultural altruism, but this too is a tough job.84 Yet one depends on one’s colleagues to test one’s ideas. As Stephen C. Stearns remembers, he first realized that George’s “cognitive function was starting to decline” when he was asked to write a paper for a symposium. “The document he submitted was so far below his previous standards that I thought it would damage his reputation to publish it, and I asked him to withdraw it. He did so, but I knew he was bothered.” This would have happened in about 1997, and as Stearns writes, was the beginning of the end. In 2002, “George told us that he had Alzheimer’s.” And

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then, after seven years, “When I returned to give the Darwin day talk at Stony Brook in 2009, George was not able to attend, and his colleagues told me that they had not seen him for some time.” Stearns concludes his touching biographical memoir by noting “It is ironic that it was a person who understood aging so well who suffered from an illness that so dramatically demonstrates how vulnerable our aging bodies are.”85 When obituaries speak of “irony,” perhaps they mean “unfair,” though surely an evolutionary biologist is not supposed to think this way. There are plenty of ironies. The first and most obvious one here is the temptation to conflate two meanings of “life history.” (This is not to fault Stearns, as almost any human in such a situation would do the same.) Williams himself pioneered the study of what can be called “the evolution of life-history patterns,” a process that reflects, in his view, tradeoffs that occur in an organism’s origin, growth, reproduction, longevity, and senescence.86 Stearns later wrote a definitive book on the subject, The Evolution of Life Histories (1992). Yet in another sense of the phrase, a life history is the series of events that make up a human individual’s life. How an individual’s “life history” might be written by an evolutionary biologist depends, perhaps, on a larger theory of life history. George C. Williams, a man of acute intellect and a pioneer in the understanding of the evolution of life history, was also like any other man. He would eventually write below his previous standards, and even experience the frustration of knowing it. That such a man in the end would be unable, in the most horrible sense, to attend, or that he would become simply unseen by his colleagues, may be ironic. For Williams was, in the highest or even noblest sense of the term, a collaborator. Yet how could such a remarkable man’s intellect be described by a close colleague as “cognitive function”? Randolph M. Nesse, another collaborator with Williams, has observed, “It was Alzheimer’s disease that took him. The irony is nearly unbearable.” In one sense, the concept of pleiotropy that Williams explored in 1957, when he was 31 years old, was an investigation of the multiple and sometimes competing effects of genetic traits on the phenotype, since some genes that enable increased fitness for young and fertile organisms may contribute to decreased fitness, disease, or senescence later in life.87 So there is some irony in that. Williams might not have considered it ironic at all that he would die from the same forces that eventually take all organisms, sooner or later. For Williams, there was no irony in the processes of life, but many ironies in the ways humans rationalized them. He

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shared this view with Ernst Mayr, who, when he told Lynn Margulis that he was dying of cancer at the age of 100 years joked, “‘Well, Lynn,’ he said cheerfully, ‘I will have to die of something.’”88 The worst rationalization, Williams thought, was in imagining that nature would save us, an illusion he satirizes in “Mother Nature is a Wicked Old Witch” (1993). Irony, of course, is the mode of tragedy, and in a certain sense, tragedy is latent in everything that Williams wrote, though he chooses to ironize often in satire. Williams engaged with the literary side of life histories. One article he coauthored in 1983 with Sarah Blaffer Hrdy—though she wrote it, and he signed on only so it might get noticed by other professionals—is “Behavioral Biology and the Double Standard,” and begins with an epigraph from George Eliot, “We don’t ask what a woman does—we ask whom she belongs to.”89 But of course we all belong to something we cannot control, and senescence is one part of a process to which we belong that we may be able to retard, though we cannot postpone it for long.

Darwinism, Comic and Tragic In “Coda: The Comic Darwin,” George Levine makes an argument about Darwin, the Writer (2011) that could never be made about George C. Williams. Levine uses Darwin’s conclusion to The Origin to emphasize how he speaks of “the accumulation of innumerable slight variations, each good for the possessor” and argues that “Darwin’s ‘Conclusion’ is surely comic in the largest sense of the term.” By this Levine means that, “although all good biologists know that ‘perfection,’ in an absolute sense, is simply impossible in Darwin’s world,” Darwin continues to use the word “perfected” in his conclusion. As Levine puts it, “There is the experience of perfection, even if part of what we learn is that it is not perfection at all.” So Levine dwells, almost against his will, on “the extraordinary diversity and richness of all that highly designed life with all that history” that is “quite wonderful again.”90 Would it were so! Would one could believe it. Levine’s wonder seems almost certainly a leftover piece of romantic sentiment that Darwin cannot help but express, and to which Levine also clings. As for Williams, his writing on the subject is stark and distinctly dark in tone, and so perhaps is the truth.91 Williams might not be right, but he is rigorous, and that is what one reads him for. A literary critic rarely asks about a text, “Is the author

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right?” Nor do I have the authority to ask whether Williams is right, whether he writes the truth. “What is truth? said jesting Pilate, and would not stay for an answer.” So begins Francis Bacon’s essay entitled “Of Truth.”92 Call them adaptations, these innumerable slight variations Williams suspected never lead to perfection, but only to a state, a condition, a fitness better than that of some others. They are individualistic yet not good for us as individuals; they only guarantee the continuance of life. What is good for continuing life when we are young does us in later. Perhaps such a view upends questions of belief. One is reminded of a dialogue from James Joyce’s Ulysses, a different dialogue from a more difficult book than the one that Bill Hamilton chose for an epigraph to his classic essay on senescence.93 In Ulysses, Haines asks Stephen about a song named not, as he supposes, “Joseph the Joiner” but “The Ballad of Joking Jesus,” that Stephen has had to endure from Buck Mulligan: –– Three times a day, after meals, Stephen said drily. –– You’re not a believer, are you? Haines asked. I mean, a believer in the narrow sense of the word. Creation from nothing and miracles and a personal God. –– There’s only one sense of the word, it seems to me, Stephen said.94

Notes 1. George C.  Williams, Sex and Evolution (Princeton, NJ: Princeton University Press, 1975), 169. 2. Sarah Blaffer Hrdy, The Woman Who Never Evolved (Cambridge, MA: Harvard University Press, 1981, 1999), 204, n. 32. 3. George C.  Williams, “Kin Selection and the Paradox of Sexuality,” in Sociobiology: Beyond Nature / Nurture?, edited by George W. Barlow and James Silverberg, American Association for the Advancement of Science Selected Symposium 35 (Boulder, CO: Westview Press, 1980): 371–84, 371. 4. George C.  Williams, Natural Selection (New York: Oxford University Press, 1992), 9. 5. Williams, Natural Selection, 13; Williams, The Pony Fish’s Glow: And Other Clues to Plan and Purpose in Nature (New York: Basic Books, 1997), 3; Randolph M. Nesse and George C. Williams, Why We Get Sick:

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The New Science of Darwinian Medicine (New York: Times Books, 1995), 21. 6. Melvin Konner, The Tangled Wing: Biological Constraints on the Human Spirit (New York: Holt, Rinehart and Winston, 1982), 15, 446. 7. See “Turning Mind into Money,” Focus, Nr. 41 (October 8, 2001), translated and reprinted in Edge: http://www.edge.org/documents/press/ focus.e.html. 8. The Quarterly Review of Biology 80, no. 1 (March 2005). The essays included are: Frank C Erk, “George C.  Williams: Personal Reminiscences” (7–11); Steven P. Ferraro “Red-Letter Days” (13–17); Helena Cronin, “Adaptation: ‘A Critique of Some Current Evolutionary Thought’” (19–26); George W.  Barlow, “How Do We Decide That a Species Is Sex-­Role Reversed?” (28–35); Mart R. Gross, “The Evolution of Parental Care” (37–45); Mary Jane West-Eberhard, “The Maintenance of Sex as a Developmental Trap Due to Sexual Selection” (47–53); Martin Daly and Margo Wilson, “Carpe Diem: Adaptation and Devaluing the Future” (55–60); Randolph M. Nesse, “Maladaptation and Natural Selection” (62–70); Bobbi S. Low, “George Williams, Theoretician and Guerilla Environmentalist”. (72–83)

9. Cronin, “Adaptation,” 22. 10. Ferraro, “Red-Letter Days,” 16. 11. Barlow, “How Do We Decide,” 33. 12. Gross, “Evolution of Parental Care,” 37, 43–44. 13. West-Eberhard, “Maintenance of Sex,” 47. 14. Mary Jane West-Eberhard, Developmental Plasticity and Evolution (New York: Oxford University Press, 2003). See especially chapter 31. 15.  See Marlene Zuk, “2003 Sewall Wright Award: Mary Jane WestEberhard,” American Naturalist 163 (2004): 1. 16. West-Eberhard, Developmental Plasticity, 5, 7. 17. West-Eberhard, “Maintenance of Sex,” 52. 18.  Mary Jane West-Eberhard, “Phenotypic Plasticity and the Origins of Diversity,” The Annual Review of Ecology, Evolution, and Systematics 20 (1989): 249–78, 249. 19. “BIO: Mary Jane West-Eberhard,” Evolution & Development 11, no. 1 (2009): 8–10, 10. 20. For a favorable review of this important book, see C. D. Rollo, “Life= epigenetics, ecology, and evolution (L = E 3): A review of Developmental Plasticity and Evolution, by Mary Jane West-Eberhard,” Evolution & Development 6 (2004): 58–62. 21. West-Eberhard, Developmental Plasticity, 16, 4, 7, ix. 22. Williams, Natural Selection, 91.

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23. West-Eberhard, Developmental Plasticity, 11, 5, 20. 24. West-Eberhard, Developmental Plasticity, 13, 15, 40. 25. Hrdy, who was there, writes, “Unfortunately it was not yet (and still is not really) true that Mary Jane West Eberhard’s ‘approach has been infectious for sociobiologists….’ I often wonder what George Williams would have thought about the growing influence of MJ’s concept of ‘social selection’ esp. as interpreted by Randy Nesse to explain the emergence of human morality and conscience. Superficially these views seem totally at odds with his darker vision re moral ideas. However, I think it possible, even likely, that George would have come round to MJ and Randy’s position. But maybe not. As far as I know, he never altered his opposition to Kristen Hawkes’ version of the Grandmother Hypothesis” (Personal communication, 2012). 26. Sarah Blaffer Hrdy, Mother Nature: A History of Mothers, Infants, and Natural Selection (New York: Random House, 1999), 56, 63. 27. West-Eberhard, Developmental Plasticity, 3. See also Peter D. Gluckman and Mark A. Hanson, “Living with the Past: Evolution, Development, and Patterns of Disease,” Science 305, no. 5691 (2004): 1733–36. 28. West-Eberhard, Developmental Plasticity, 637. 29. West-Eberhard, “Maintenance of Sex,” 47. 30. Daly and Wilson, “Carpe Diem,” 59. 31. Bobbi S.  Low, Why Sex Matters (Princeton, NJ: Princeton University Press, 2000), 3. 32. See Bobbi S. Low, “Behavioral Ecology of Conservation in Traditional Societies,” Human Nature 7, no. 4 (1996): 353–79. See also Ter Ellingson, The Myth of the Noble Savage (Berkeley: University of California Press, 2001). 33. Marlene Zuk, Paleofantasy: What Evolution Really Tells Us about Sex, Diet, and How We Live (New York: Norton, 2013), 6. 34. Low, Why Sex Matters, 255, 256; Marlene Zuk, Sexual Selections: What We Can and Can’t Learn about Sex from Animals (Berkeley: University of California Press, 2002), 34–46. 35. Low, “George Williams,” 76. 36. Adrienne Rich, “Turning the Wheel,” in A Wild Patience Has Taken Me This Far, Poems 1978–1981 (New York: Norton, 1981), 59. 37. Low, “George Williams,” 76. 38. Zuk, Sexual Selections, 1. 39. Low, “George Williams,” 74, 75, 81. 40. Low, Why Sex Matters, 92. 41. Colin McGinn, “Some Guys Have All the Luck: How the Dominant Males Rule the Reproductive Roost,” New York Times (January 9, 2000), http://www.nytimes.com/books/first/l/low-­sex.html.

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42. Elliott Sober and David Sloan Wilson, “Adaptation and Natural Selection Revisited,” Journal of Evolutionary Biology 24 (2011): 462–68, 462, 467. See also Elliott Sober and David Sloan Wilson, “Re-Introducing Group Selection to the Human Behavioral Sciences,” Behavioral and Brain Sciences, 17, no. 4 (1994): 585–654; Elliott Sober and David Sloan Wilson, Unto Others: The Evolution and Psychology of Unselfish Behavior (Cambridge, MA: Harvard University Press, 1998). 43. David Sloane Wilson, “Truth and Reconciliation for Group Selection X: Naïve Group Selectionism,” The Huffington Post (May 20, 2009), https://www.huffpost.com/entry/truth-­and-­reconciliation_b_188707. 44.  Elisabeth Lloyd, “Units and Levels of Selection,” The Stanford Encyclopedia of Philosophy (Fall 2005 Edition), edited by Edward N. Zalta, http://plato.stanford.edu/archives/fall2005/entries/selection-­units/. 45. Richard Lewontin, “The Units of Selection,” Annual Review of Ecology and Systematics, 1 (1970): 1–18, 14. 46.  David Sloan Wilson and E.  O. Wilson, “Rethinking the Theoretical Foundation of Sociobiology,” the Quarterly Review of Biology 82, no. 4 (December 2007): 327–48, 327, 328. 47. E.  O. Wilson, Consilience: The Unity of Knowledge (New York: Knopf, 1998), 8, 266, 9. 48. Gregory J.  Cooper, The Science of the Struggle for Existence: On the Foundations of Ecology (Cambridge, UK: Cambridge University Press, 2003), 56, 58. 49. Williams, Adaptation, 247, 265. 50.  Douglas J.  Futuyma, “Reflections on Reflections: Ecology and Evolutionary Biology,” Journal of the History of Biology 19, no. 2 (Summer 1986): 303–312, 307, 309. 51. Futuyma, “Reflections,” 308. 52.  Stephen C.  Stearns, “George Christopher Williams 1926–2010, A Biographical Memoir,” National Academy of Sciences (2011), 12. http:// www.nasonline.org/publications/biographical-­memoirs/memoir-­pdfs/ williams-­george.pdf. 53. Futuyma, “Reflections,” 308, 312. 54. Charles Elton, Animal Ecology (Chicago: University of Chicago Press, 1927), 64. 55.  A standard textbook of evolutionary ecology is Eric E.  Pianka, Evolutionary Ecology, fifth edition (New York: Harper Collins, 1994). See also Frank B. Golley, A History of the Ecosystem Concept in Ecology: More than the Sum of the Parts (New Haven, CT: Yale University Press, 1996); David R. Keller and Frank B. Golley, eds. The Philosophy of Ecology: from Science to Synthesis (Athens: University of Georgia Press, 2000).

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56. A. J. Zellmer, T. F. H. Allen, K. Kesseboehmer, “The Nature of Ecological Complexity: A Protocol for Building the Narrative,” Ecological Complexity 3 (2006): 171–82. Donald Worster, “The Ecology of Order and Chaos,” Environmental History Review (1990): 1–18. 57. L.  B. Slobodkin, “The Light and the Way in Evolution,” a review of Adaptation and Natural Selection. A Critique of Some Current Evolutionary Thought, by George C.  Williams, the Quarterly Review of Biology 41, no. 2 (June 1966): 191–94, 191. Citing Williams, Adaptation, 258. 58.  These appreciations appear on John Brockman’s Edge: “George C. Williams (1926–2010), A Remembrance with Robert Trivers, Richard Dawkins, Daniel C.  Dennett, Niles Eldredge, Randolph Nesse,” Edge, September 9, 2011, https://www.edge.org/conversation/robert_ trivers-­r ichard_dawkins-­d aniel_c_dennett-­n iles_eldredge-­r andolph_ nesse-­george-­c. 59. “George C. Williams (1926–2010), A Remembrance,” Edge. 60. “George C. Williams (1926–2010), A Remembrance,” Edge. 61. Peter J.  Corning, “Evolution and Ethics: An Idea Whose Time Has Come? (Part I),” Journal Of Social And Evolutionary Systems 19, no. 3 (1996): 277–85. 62. Thomas Shepard, The sincere convert discovering the paucity of true beleevers and the great difficulty of saving conversion (London: Printed by T. P. and M. S. for Humphrey Blunden, 1641). 63. Frans de Waal speaks of Williams and Dawkins as makers of “Calvinist sociobiology.” Both writers are, as he says, Huxleyans. See The Ape and Sushi Master: Cultural Reflections of a Primatologist (New York: Basic Books, 2001), 346. 64. See, for instance, Massimo Piattelli-­Palmarini, Inevitable Illusions: How Mistakes of Reason Rule Our Minds (New York: Wiley, 1994). 65. Frans de Waal, Good Natured: The Origins of Right and Wrong in Humans and Other Animals (Cambridge, MA: Harvard University Press, 1996), 16. 66.  Frans de Waal, “Morally Evolved: Primate Social Instincts, Human Morality, and the Rise and Fall of ‘Veneer Theory,’” in Primates and Philosophers: How Morality Evolved, edited by Josiah Ober and Stephen Macedo, with commentary by Robert Wright, Christine M. Korsgaard, Philip Kitcher, and Peter Singer (Princeton, NJ: Princeton University Press, 2006), 1–80, 6. 67. A popular account is Lee Alan Dugatkin, The Altruism Equation: Seven Scientists Search for the Origins of Goodness (Princeton, NJ: Princeton University Press, 2006). 68.  Williams, “Some Thoughts on William D.  Hamilton (1936–2000),” Trends in Ecology and Evolution 15, no. 7 (July 2000): 302.

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69. “George C.  Williams (1926–2010), A Remembrance,” Edge, https:// www.edge.org/conversation/robert_trivers-­richard_dawkins-­daniel_ c_dennett-­niles_eldredge-­randolph_nesse-­george-­c. 70. George C. Williams, “Mysteries of Sex and Recombination,” a review of The Evolution of Sex by John Maynard Smith, the Quarterly Review of Biology 53, no. 3 (September 1978): 287–89. 71. Williams, Plan and Purpose, 177. 72. George C. Williams, “The Tithonus Error in Modern Gerontology,” the Quarterly Review of Biology 74, no. 4 (December 1999): 405–15. 73. From Petronius, Satyricon: “Nam Sibyllam quidem Cumis ego ipse oculis meis vidi in ampulle pendere, et cum illi pueri dicerent: Σίβυλλα τί θέλεις. respondebat illa: άποθανεΐν θέλω.” 74. Williams, “Tithonus Error,” 405, quoting Ecclesiastes 3:2. 75. Williams, “Tithonus Error,” 405, 406. Fisher used death and actuarial tables for his work. These tables tell someone about beginning and end points of a process. They measure spans of time. They do not reveal the process itself. These tables distort human thinking about lifespans, and these distortions are very much with us today. The medical business measures this way when it commands that people live longer. 76. Williams, “Tithonus Error,” 405. 77. Williams, “Tithonus Error,” 406, 407. 78. W. D. Hamilton, Narrow Roads of Gene Land, vol. 1: Evolution of Social Behaviour (Oxford: Oxford University Press, 1996), 89. 79.  Alan Grafen, “William Donald Hamilton, 1 August 1936–7 March 2000,” Biographical Memoirs of Fellows of the Royal Society 50 (2004): 109–132, 126. 80. George C. Williams, “Some Thoughts on W. D. Hamilton (1936–2000),” Trends in Ecology and Evolution 15, no. 7 (July 2000): 302. 81. Williams, “Tithonus Error,” 407, 414. 82. Andrew Marvel, “To His Coy Mistress,” lines 45, 46. 83. Stephen I.  Rothstein, “Reciprocal Altruism and Kin Selection Are Not Clearly Separable Phenomena.” Journal of Theoretical Biology 87 (1980): 255–61. 84.  Samir Okasha, “Biological Altruism,” The Stanford Encyclopedia of Philosophy (Winter 2009 Edition), edited by Edward N.  Zalta, http:// plato.stanford.edu/archives/win2009/entries/altruism-­biological/. 85. Stearns, “George Christopher Williams,” 19. 86. So-called “life-history strategies” reveal a set of reproductive rates and survival probabilities experienced over a lifespan, the interaction between reproduction and survival. Some variables include age or size at first reproduction, offspring size and number, parental care, and senescence. 87. George C. Williams, “Pleiotropy, Natural Selection, and the Evolution of Senescence,” Evolution 11 (1957): 398–411.

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88.  L ynn Margulis, “Ernst Mayr, Biologist Extraordinaire,” American Scientist 93, no. 3 (2005): 200–1, 201. 89. Sarah Blaffer Hrdy and George C. Williams, “Behavioral Biology and the Double Standard,” in Social Behavior of Female Vertebrates, edited by Samuel K.  Wasser (New York: Academic Press, 1983), 3–17. BlafferHrdy uses another epigraph from George Eliot for Mother Nature (1999): “Mother Nature—who by the bye is an old lady with some bad habits … .” 90. George Levine, “Coda: The Comic Darwin,” in Darwin, the Writer, 211–22 (New York: Oxford University Press, 2011), 212, 213, 220. 91. Consider the title of the definitive history of sociobiology controversies: Ullica Christina Olofsdotter Segerstråle, Defenders of the Truth: The Sociobiology Debate (Oxford: Oxford University Press, 2001). 92. Francis Bacon, “Of Truth,” The Essays (1601). 93. Hamilton used the following epigraph from Portrait of an Artist as a Young Man, for “The Moulding of Senescence by Natural Selection” (1966): “—The most profound sentence ever written, Temple said with enthusiasm, is the sentence at the end of the zoology. Reproduction is the beginning of death. He touched Stephen timidly at the elbow and said eagerly: —Do you feel how profound that is because you are a poet?” 94. James Joyce, “Telemachus,” Ulysses.

Index1

A Abstraction, 73, 86, 92, 115, 124, 126, 152, 153, 242, 243, 285, 317, 318 Adaptation, 1, 30, 72, 99, 133, 166, 191, 228, 268, 298, 325 Adaptationist, 9, 36, 89, 90, 179, 238, 259, 268, 286, 294–299 Adaptationist program, 32, 123, 166, 167, 174, 176, 177, 192, 238, 252, 269, 294 Adaptationist storytelling, vs. just-so story, 297 Adaptive countershading, 296 Advanced organisms, 100, 128 Aesthetic distractions, 99–128, 337 Alcock, John, 32, 268, 269 Alexander, Richard, 211 Allegory, 81, 125–126, 183, 315–319 Allegory of the Eye Blink, 256

Alleles, 54–57, 105, 119, 153, 242, 251, 284 Allometry, 278 Alter, Robert, 103 Altruism, 12, 33, 40, 60–64, 136, 140, 167, 168, 171, 191, 195, 196, 200, 204, 308, 311–313, 336–338, 343 American Naturalist, 11, 119, 308 Analogies, 10, 13, 15, 17, 19, 42, 43, 55, 57, 59, 72–75, 77, 81, 83, 84, 91, 116, 117, 123, 126, 137, 141, 142, 146, 150, 156, 166, 173, 174, 176, 199, 211, 230–232, 238, 242, 243, 255, 294, 298, 299, 340 of codex, 230–232 gross immorality vs. moral indifference of nature, 199 Anthropomorphism, 41, 74, 171

 Note: Page numbers followed by ‘n’ refer to notes.

1

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 M. P. Cohen, George C. Williams and Evolutionary Literacy, Literatures, Cultures, and the Environment, https://doi.org/10.1007/978-3-031-11650-6

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354 

INDEX

Antolin, Michael F., 276 Apples, 83, 123, 124 Arendt, Hannah, 266 Austen, Jane, 63 Autoimmune disease, 272 Avatar, 247–251 Avatars, humans as, 250 Axelrod, Robert, 64, 136 B Bach, Johann Sebastian, 16, 99 Baron-Cohen, Simon, 275 Bean-bag genetics, 52 Beauty, 99–128, 149, 172, 201, 207, 213, 265, 314, 337 Behavior, 2, 30, 75, 99, 133, 168, 190, 236, 270, 300, 323 Beneficial death, 23, 78, 305 Betzig, Laura, 134, 135, 158 Bible, 102, 103 Biodiversity and extinction, 157 Biological theory, 4, 175, 212, 244, 324 explaining and predicting, 244, 324 Biologist, 1, 3–5, 7, 10–12, 16, 17, 22, 24, 30, 32–35, 37–41, 44–47, 50, 53, 55, 57–59, 72–74, 78, 82, 85, 86, 88–90, 92, 95n20, 99, 100, 102–106, 113–117, 119, 123, 124, 126, 127, 136, 138, 152–155, 165, 167, 176–178, 193–195, 199–201, 205, 216, 217, 227, 228, 230, 232, 236, 239, 240, 244, 247–249, 251, 254, 255, 257, 267, 269, 273, 295, 297, 300, 303, 310, 313, 315, 317, 318, 325, 328, 333, 335–336, 341, 344, 345 Biotic adaptations, 84, 85, 88, 100, 105–107, 120, 141, 214 Blaming the patient, 280

Bloom, Harold, 103 Bohr, Neils, 2, 24n1 Bookkeeping and causality, 236 Bookkeeping, process, 173 Book of Job, 202, 218 Book of Nature, 174, 182–183, 227–260 Brockman, John, 11, 12, 181, 258, 269, 293, 325 Brower, David, 13 Brownmiller, Susan, 149 Buller, David J., 279 Burhoe, Ralph Wendell, 203 C Calvino, Italo, 41 Campbell, D. T., 200 Camus, Albert, 310, 315 Carrol, Lewis, 123 Carson, Rachel, 49, 317 Chandler, Raymond, 308, 309 Cheating, 136, 139, 140, 149, 168–170, 200, 304 Chicago school of ecology, 193 Chomsky, Noam, 86 Clades, 230, 232–236, 244, 249–252 Clade selection, 230, 233, 235, 242, 246, 248, 251–252 Classical oration, 15 Clever Hans, the horse, 181 Close reading, 71, 174, 326 Cobb, John B., 203, 206 Codex, codices, 231 Coleridge, Samuel Taylor, 100, 183 Commoner, Barry, 217 Constraint, 12, 50, 119, 121, 128, 182, 227–260, 270, 279, 286, 297, 298, 310, 334, 337 Contingency, 17, 19, 30, 35, 89, 90, 174, 175, 177, 180, 183, 211, 227–260, 284 Cooper, Gregory J., 333

 INDEX 

Cosmic process, 189, 197, 198, 201, 210, 211, 341 Cosmides, Leda, 275, 279 Cost of meiosis, 137, 139, 140, 146, 169 Cost-benefit analysis, 42, 118, 136, 139, 140 Coyne, Jerry, 37 Cracraft, Joel, 229, 230 Crafoord Prize in Biological Science, 11 Crane, Stephen, 308 Crick, Francis, 45, 49–51, 67n58, 172, 173 Critical thinking, 23 Critique, 3, 7, 9, 13–15, 22–24, 29–64, 71, 77–79, 82, 88, 101, 123, 135, 155, 167, 170, 171, 177, 192, 193, 206, 207, 232, 240, 268, 282, 285, 286, 309, 312, 323, 326, 329–333, 335, 336, 338 Cronin, Helena, 325 Cues (Tinbergen), 75 Cybernetics, 73, 92, 152, 153, 242, 262n44 D Dagg, Joachim, 139, 167 Dalton, John, 13, 166 Daly, Martin, 325, 329 Damuth, John, 248, 249 Darwin, Charles, 17, 19, 24, 36, 45, 72, 79–81, 86, 94, 102, 104, 116, 117, 124, 125, 145, 149, 179, 180, 195, 196, 199, 211, 213, 216, 229, 240, 252–257, 281, 302, 315, 318, 329, 335, 344, 345 as authority, 104, 281, 346 on group selection, 195, 196 Darwinian constraints, 298

355

Darwinian medicine, 259, 268, 269, 272, 276, 281, 286, 308, 328 Dawkins, Richard, 12, 13, 21, 30, 31, 33, 39, 40, 56, 57, 71, 102, 123, 135, 165, 166, 172, 175, 181, 203, 211, 227, 239, 244, 280–282, 294, 303, 308, 327, 328, 335–337 de Sousa, Ronald, 32 De Waal, Frans, 210, 212, 214, 337 Death, 12, 23, 29, 43, 56, 60, 61, 74, 77–79, 82, 85, 88, 133–159, 172, 182, 215, 216, 251, 272, 281, 285, 286, 304–307, 314–316, 330, 335, 340–345, 351n75, 352n93 DeGregori, Thomas, 172 Demography, 78, 88, 107 Dendrograms, 126, 232–236, 243 Dennett, Daniel C., 9, 19, 33, 180, 186n47, 211, 224n105, 335, 336 Depersonalization, 20 Depuit, Jules, 136 Design, 18–20, 22, 34, 35, 37, 43–44, 49, 50, 59, 71–94, 101, 104–105, 107, 114–116, 124, 134, 142, 152, 178, 179, 192, 205, 211, 228, 230, 232, 241, 242, 247, 253–257, 267, 270, 272, 275, 287, 288, 295, 299, 301, 302, 310, 313, 336, 337 Design, of eye, hand, 241 Determinism, genetic, 212 Dickermann, Mildred, 323 Dillard, Annie, 201 DNA, 4, 45, 50, 51, 105, 172, 173, 176, 181, 227, 242, 243, 272 Dobzhansky, Theodosius, 5, 11–13, 20, 21, 25n16, 33, 58, 93, 144, 153, 156, 199, 237, 278, 333 Doctrine, 20, 30, 32, 35–38, 83, 84, 88, 96n38, 127, 179, 216, 231, 238, 252, 273, 306, 307

356 

INDEX

Domain mix, 314 Domains, codical vs. material, 227, 235, 242, 249, 314 Don Quixote, 243 Drabble, Margaret, 194 Dugatkin, Lee Alan, 38, 61 E Ecocriticism, 3 Ecological Society of America, 10, 334 Ecology, 2, 4, 16, 46, 47, 87, 140, 193, 209, 210, 217, 236, 246–247, 249, 270, 284, 325, 330, 333–334 Ehrlich, Paul, 78, 259, 334 Eiseley, Loren, 116 Eldredge, Niles, 175, 227, 247, 335 Eliot, T. S., 41, 145, 228, 339, 340 Emerson, Alfred E., 23, 78, 193, 237, 305 Emerson, Ralph Waldo, 3, 11 Eminent Ecologist Award, Ecological Society of America, 334 Environmental humanities, 6, 13 Environment of evolutionary adaptedness, 279, 330 Environmental uncertainty, 142, 156, 157 Erk, Frank C., 44, 325 Ethics, evolutionary, 10, 16, 34, 191–195, 198, 259, 308, 312, 340 Ethology, 16, 22, 32, 33, 46, 75 Eugenics, 273, 274 Evolution, 1, 8–13, 17, 20, 22, 30–33, 35, 38, 40, 43, 46, 47, 49–52, 55, 58–63, 72, 73, 83, 87, 92–94, 96n29, 100, 101, 104–107, 111, 114, 116, 118, 121, 123, 124, 126, 127, 133, 139, 142, 145, 150, 154–158, 170, 172–175,

177, 180, 182, 186n48, 189–218, 229, 230, 232, 237–239, 244, 246, 251, 252, 257, 258, 265, 268, 269, 271, 275, 276, 279, 281, 282, 286, 288, 293–295, 299, 301, 308, 312, 313, 315, 316, 324, 327, 328, 330, 332, 333, 336, 338, 341–344 Evolutionary ethics, 10, 16, 34, 190–195, 198, 259, 308, 312, 340 Evolutionary medicine, 10, 42, 268, 269, 281 Evolutionary psychology, 16, 32–36, 40, 149, 270, 275, 278–280, 313, 330 Evolutionary stable strategy (ESS), 155, 300–302 Evolutionist, 205, 240 Ewald, Paul, 267, 271 Extinction, 20, 74, 85, 105, 117, 118, 123, 155–157, 172, 175, 216, 228, 232, 251, 252, 258, 259 Eye, 11, 19, 46, 54, 83, 86, 89, 91, 92, 102, 113–116, 139, 228, 233, 234, 236, 239–241, 249, 253–257, 272, 274, 287, 294, 295, 317, 339 F Farber, Paul Lawrence, 198 Fecundity, 42, 77, 101, 108, 119, 120, 136, 144–151, 333 Fertilization, 107, 109, 134, 139, 148–150, 168, 169, 307 Fischer, R. S., 31, 104 Fisher, R. A., 11, 13, 16, 18, 25n16, 42, 82, 95n20, 107, 119, 141, 148, 152, 153, 228, 237, 257, 273, 279, 302, 333, 351n75 Fisher’s Fundamental Theorem, 82

 INDEX 

Fitness, 32, 33, 40, 54–57, 59, 62–64, 73, 82, 85, 92, 104, 107, 109, 120–122, 124, 134–137, 142, 144–146, 149, 150, 152, 153, 157, 167, 168, 170, 173, 178, 181, 182, 191, 199, 200, 234, 236, 242, 245–247, 251, 282, 300, 301, 326, 340, 341, 343, 344, 346 Fitness, vernacular, 82, 122 Flying fish, 79, 81 Foucault, Michel, 135 Frank, Steven A., 154 Frequency-dependent selection, 245, 247, 301–303 Frost, Robert, 256 Frye, Northrop, 207 Function, 10, 18, 19, 22, 30, 32, 36, 53, 59, 62, 76, 77, 82–89, 91, 103, 106–112, 114, 115, 122, 124, 125, 138, 142, 148, 152, 169, 192, 198, 207, 236, 241, 249, 266, 268, 272–275, 278, 283–285, 296, 297, 299, 304, 309, 329, 343 Functional design, 19, 85, 89, 114, 115, 205, 255 Futuyma, Douglas J., 15, 333, 334 G Gaia hypothesis, 179, 206–210, 330 Galen, 19, 93, 177, 196, 218, 228, 241, 253, 254 Galton, Sir Francis, 54 Game theory, 95n20, 134, 136, 140, 142, 155, 179, 293, 300–302, 305, 315 Gene, 13, 35, 73, 101, 133, 166, 194, 227, 270, 304, 329 Gene reductionism, 173, 177 Gene selectionism, 32, 172–174, 204, 230, 251, 259

357

Gene, selfish, 12, 13, 33, 40, 41, 166, 178, 181, 194, 202, 203, 215, 244 Gene’s eye view, 12, 33, 139, 233, 234, 236, 242 Genetic determinism, 212 Genetic drift, 101 Genetic information, 49, 50, 72, 86, 240 Genetic load, 245 Genotype, 18, 35, 43, 51, 52, 54–56, 105, 143–147, 156, 157, 173, 176, 234, 235, 242, 245–247, 251, 301, 307, 327, 341 Genre, 3, 5, 6, 93, 207 Gerontology, 11, 340 God-is-good tradition, 306 Gould, Stephen Jay, 5, 13, 15, 32, 39, 123, 126, 167, 175, 176, 179, 181, 192, 213, 227, 230, 236, 258, 259, 293, 297, 299, 335 Grafen, Alan, 153, 154, 341 Grandmother effect, 60 The Great Gatsby, 308 Grene, Marjorie, 157, 259 Griffin, Donald R., 175, 180 Griffiths, Paul, 12, 305 Gross, Mart R., 121, 123, 247, 325 Group selection, 10, 32, 33, 40, 64, 74, 78, 81, 82, 87, 88, 90, 91, 134, 138, 157, 167, 177, 193, 195, 196, 209, 237, 246, 332–334, 336 Gymnosperms, 74, 157 H Haeckel, Ernst, 39 Haldane, J. B. S., 11, 13, 18, 61, 62, 104, 111, 113, 152, 153, 158, 173, 228, 237, 333 Hamilton, William, 12, 13, 21, 31, 40, 41, 45, 64, 84, 104, 134–136, 273, 274, 286, 315, 330, 334

358 

INDEX

Hamilton’s Rule, 199 Hamlet, 265, 342 Hammett, Dashiell, 309, 321n47 Hardboiled style, 308 Hastings, Jon Woodland, 296–299 Hauser, Mark, 75 Heterozygosity, 106 Higher order, 99, 100, 337 Historicity, 35, 174, 227, 228, 231, 232, 239, 252–253, 301, 324 Holism, 208, 237–239 Holy corpse fallacy, 306 Homing, 41, 44, 46–48 Hostility of the universe, 210 Hrdy, Sarah Blaffer, 31, 91, 121–123, 201, 203, 204, 206, 210, 313, 323–325, 328, 345, 348n25 Hull, David L., 172, 175, 244 Human condition, 199, 266, 276, 287–288, 312, 314 Human nature, 40, 170, 202, 204, 206, 217, 265–288, 305, 308, 312, 314, 342 Hume, David, 191, 192, 196, 313, 314 Huxley, Aldous, 171, 190, 196, 199, 202, 203, 210, 211, 339 Huxley, Julian, 11, 22, 127, 203, 233, 335 Huxley, Thomas Henry, 20, 34, 93, 181, 189, 190, 192, 194–199, 203, 204, 206, 212, 215, 308, 313, 314, 338, 343, 344 Hybridization, 49, 106

Infanticide, 31, 101, 123, 201 Infection, 270, 271, 282, 283 Instinct, 53, 76, 83–84, 86, 111, 115, 194, 279 Intelligent design, 115, 257 Interactor, replicator, 244 Intervention, 77, 144, 272, 273, 283, 340 Is-ought problem, 191, 192

I Imagination, 1–24, 41, 86, 102, 112, 183, 243, 254 Inclusive fitness, 32, 33, 40, 62–64, 84, 85, 134, 135, 153, 167, 168, 170, 181, 200, 236, 244, 343

L Lack, David Lambert, 21, 47, 71, 79, 104, 108, 118–121, 148, 193, 237, 330, 333 Lanham, Richard, 94 Leopold, Aldo, 13

J James, William, 195, 329 Jones, Steve, 116 Joyce, James, 158, 183, 346 Judson, Horace Freeland, 51 Just-so stories, 32, 101, 179–181, 294, 298 K Kerouac, Jack, 44 Keywords, 3 Kin, 12, 85, 330, 345 Kin Selection (KS), 33, 40, 41, 64, 150, 167–169, 172, 181, 236, 244, 312, 316, 328, 343 Kirchner, James, 206 Kitcher, Philip, 191, 192, 213, 337 Kohn, Marek, 213, 215 Konner, Melvin, 324 Krieger, Murray, 183 K-selection, 42

 INDEX 

Levels of selection, 30, 90–92, 100, 135, 174, 175, 227, 229, 230, 233, 244, 332, 334 Levine, George, 315, 345 Lewens, Tim, 254 Lewontin, Richard C., 123, 167, 176, 177, 179, 181, 186n48, 192, 208, 211, 227, 297, 332 Life cycle, 59, 60, 86, 109, 141, 144, 147, 182, 236, 270, 341 Life history, 12, 16, 29, 41–43, 59, 60, 72, 118–121, 142, 144, 172, 181–183, 202, 245, 303, 340, 344, 345, 351n86 Life history theory, 42, 182, 330 Literacy, evolutionary, 1, 14–16, 30, 35–37, 170, 265 Lloyd, Elizabeth, 332 Loose style, 103 Lottery, of survival, 141 Love, 49, 172, 210, 213, 217, 296, 311, 339 Lovelock, James, 206–209 Low, Bobbi, 121, 145, 148–150, 158, 325, 329–331 M MacArthur, Robert, 182, 334 Macroevolution, 175, 177, 229, 230, 232–236, 258 Maladaptation, 58, 90, 146, 273, 286, 311, 329 Maladaptationist, 90 Maladies, 268, 273, 275, 277, 282, 285, 331 Malthus, Thomas Robert, 104 Margulis, Lynn, 139, 206, 207, 215, 336, 345 Mathematical expression, 305 Mathematical reasoning, 152, 154 Mathematical rhetoric, 152 May, Robert, 39

359

“May” syntactical construction, 282–285 Maynard Smith, John, 11, 21, 23, 33, 38, 40, 51, 61, 104, 134–138, 140, 152–155, 179, 244, 300, 315, 334, 338 Mayr, Ernst, 5, 11, 37, 47, 52, 61, 72, 83, 87, 93, 113, 114, 118, 126, 153, 156, 237, 248, 254, 268, 297, 333, 338, 345 McCarry, John, 305 McGinn, Colin, 331 Mechanism, 10, 13, 29, 30, 35, 47–49, 53, 76, 79, 82–89, 100, 105, 106, 110, 111, 114, 115, 118, 119, 123, 124, 133, 136, 147, 154, 156, 167, 171, 180, 204, 211, 227, 237–239, 252, 256, 268, 273, 274, 282, 283, 287, 299, 324, 328, 329, 333, 340 Medawar, Peter, 57, 107, 153 Medium and message, 243, 244, 314 Meiosis, 34, 105, 137, 139, 140, 146, 147, 165, 168, 169, 172, 233 Melville, Herman, 201, 213, 214, 317, 318 Mendelian population, 105, 251 Mental disorders, 267, 280 Mentalism, 22, 175, 181 Meyer, Axel, 60 Microevolution, 232 Midgley, Mary, 13, 197 Military metaphors, 199 Mistakes, 9, 85, 103, 111, 271, 337, 340 Mitman, Gregg, 192, 193 Mob vs. group, 90, 112 Models of reproduction aphid-rotifer, 142–143 cod-starfish, 144 elm-oyster, 142–143 strawberry-coral, 142–143

360 

INDEX

Modern synthesis, 5, 11, 12, 92, 93, 127, 152, 211, 259 Moment-of-conception fallacy, 307 Monod, Jacques, 49, 67n58 Moore, George E., 192 Morality, as veneer, 211–214, 337 Morgan, Lloyd, 81 Mortality, 42, 77, 101, 108, 109, 119, 120, 138, 144, 150, 151, 182, 279, 303 Mothering, 210 Mother Nature, 12, 190, 206, 210, 275, 345 Muller, Hermann Joseph, 228, 241 N Nagel, Thomas, 45 Narratives, 8, 14, 15, 17, 30, 41–43, 73, 80–82, 93, 102, 117, 125, 126, 140, 142, 144, 147, 152, 169, 170, 175, 178, 179, 182, 213, 217, 239, 240, 295, 318, 319 National Academy of Sciences, 10, 59, 259 Naturalism, 44, 45, 66n37, 198, 216–218 Naturalist, 38–41, 44, 45, 147, 201, 216–218, 308 Naturalistic fallacy, 192 Natural selection, 1, 31, 72, 99, 144, 166, 190, 228, 268, 294, 324 Nature, 1, 35, 73, 99, 140, 166, 189, 239, 266, 295, 329 Nature writing, 1, 5, 6, 40 Nesse, Randolph, 11, 19, 34, 40, 87, 89, 90, 259, 265–269, 276, 280–286, 288n1, 291n60, 325, 328, 344 New Critics, 36, 174, 243 Niche, 29, 45, 47, 157, 245–249, 334

Noir, 259, 308–309 Normative judgments from nature, 20, 199 O Oates, Joyce Carol, 5 Obfuscation, 181, 229, 239 Occam’s Razor, 80–82, 105 Odum, Eugene, 177, 334 O’Hara, Robert J., 126 Okasha, Samir, 152, 166, 236, 343 Old Testament, 103, 306 One-locus model, 173 Onerous, 14, 22, 23, 30, 37, 43, 73, 77, 83, 116, 179, 240, 279, 317 Optimization, 90, 122, 123, 134, 178, 179, 195, 229, 246–247, 286 Optimize, 121, 125, 149, 178, 236, 247 Optimized response, 245 Organic adaptations, 84, 85, 105, 107 Organism as artifact, 57, 238 as crystal, 238 as document, 230, 238 P Paley, Grace, 59 Paley, William, 18, 19, 88, 93, 114, 115, 196, 228, 241, 253–257, 287 Palmer, Craig T., 158 Panglossian paradigm, 192 Paradis, James, 190, 197, 198 Paradox, 17, 20, 52, 57, 134, 137, 138, 141–142, 167, 199, 203, 272, 324 Parental investment, 32, 42, 119, 136, 148, 182

 INDEX 

Parsimony, 37, 78–81, 95n29, 100, 123 Parthenogenesis, 134, 142, 146, 147, 151 Pathogens, 270, 271 Pauling, Linus, 285 Pauling principle, 285 Payoffs, 136, 140, 301–303 costs and benefits, 135 Phenotype, genotype, 43, 51, 54–56, 245, 327, 328 Phylad, 244, 249 Phylad selection, 244 Pinker, Steven, 19, 39, 86, 121, 211, 335, 336 Piracy, 139, 169, 170 Pittendrigh, Colin, 71, 113–115 Plain style, 5, 14, 38, 94, 99, 101, 102, 113, 228 Plasticity, 79, 149, 285, 327 Pleiotropy, 56–60, 78, 101, 288, 344 Pleistocene, 59, 145, 175 Poe, Edgar Allen, 296, 298, 299 Polar bears, 117 Pony fish, 293–299, 318 Pope, Alexander, 18, 266 Popper, Karl, 179, 297 Popularization, 293 Population genetics, 5, 52, 54, 55, 152, 245 Populations, 23, 31, 34, 42, 47, 48, 53, 55, 57, 60–63, 72, 74, 76–78, 81, 82, 84, 85, 88, 100, 101, 105–108, 115, 117, 119, 134, 137, 139, 141, 142, 144–148, 153, 155–157, 167, 168, 173, 176, 177, 180, 195, 197, 232, 233, 236, 241, 243, 245–251, 258, 274, 278, 284, 285, 301–303, 314, 317, 330, 332, 333 Population thinking, 47, 93, 126

361

Pound, Ezra, 241 Price, George C., 23, 314–316, 337 Prisoner’s Dilemma, 136 Progress, 3, 8, 33, 49, 73, 87, 88, 100, 104, 117, 121, 122, 126–128, 165, 166, 189, 198, 267, 281, 285, 341 Prophesy in science, 134 Proximate vs. ultimate explanations, 84–87, 268, 282, 329 Q Quarterly Review of Biology, 11, 190, 268, 313, 324, 325, 335 R Rationalism, 5 Reason, 1, 19, 20, 24, 29, 40, 41, 53, 61, 64, 71, 77, 79, 89, 102–104, 108, 109, 112, 113, 116, 117, 122, 138, 141, 145, 147, 149, 152, 156, 167, 171, 172, 178, 180, 192, 194–196, 199, 201, 203–205, 207, 208, 213, 229, 230, 232, 245–248, 254, 256, 257, 259, 265, 277, 278, 282, 283, 285, 295, 297–299, 311, 313, 317, 318, 337, 342 Reciprocal altruism (RA), 134–136, 168, 210, 343 Red Queen hypothesis, 123, 143 Reduction, 16, 17, 41, 42, 80, 87, 126, 139, 148, 165, 166, 169, 171–176, 182–183, 234, 237–239, 311, 318, 331 Reductionism, 11, 17, 165–183, 208, 239 Religion, 66n37, 74, 218 Replicator, 175, 181, 203, 242, 244, 332

362 

INDEX

Reproduction asexual, 57, 87, 137, 138, 141, 142, 145–147, 162n60 sexual, 1, 11, 30, 34, 57, 78, 87, 105–107, 113, 118, 133, 134, 137–139, 141–143, 146, 147, 152, 153, 158, 162n60, 167, 169, 170, 172, 176, 234, 246, 270, 305, 327 Reproductive effort, 109, 120, 122, 329 Reproductive value, 120, 149, 329, 341 Rhetorical emphasis, 91, 277 Rhetorical questions, 103, 267 Rich, Adrienne, 330 Richards, Robert J., 39, 195–197, 207, 313 Ricklefs, Robert E., 120 Ridley, Mark, 15, 55, 58, 92, 165, 233 Roes, Frans, 217, 218, 306 Roes, Frans, “Speak Darwinists,” 217 Rolston III, Holmes, 308 Romanticized view of nature, 99 Roulette, 137, 141 R-selection, 42 Ruse, Michael, 203, 205, 206 S Schiller, Friedrich, 38 Schliemann, Heinrich, 295 Schools, of fish, 52, 53, 77, 91, 99, 193, 304 Schwenk, Kurt, 252 Science Masters Series, 293 Seals, parable of, 213–214 Self-deception, 93, 215 Senescence, 11, 23, 29, 42, 55–60, 88, 135, 153, 182, 267, 305, 340, 341, 343–346, 351n86 Sexes, battle of, 110, 122

Sex ratio, 95n20, 106, 107, 140, 300, 302, 303 Sexual behavior, 30, 104, 110, 133, 140, 142, 155, 158 Sexual reproduction, 1, 11, 30, 34, 57, 78, 105–107, 113, 118, 133, 134, 137–139, 141–143, 146, 147, 152, 153, 156, 158, 167, 169, 170, 172, 176, 234, 246, 270, 305, 327 Shepard, Paul, 213 Shepard, Thomas, 336 Sherlock Holmes, 298 Simile, 116, 125, 126, 201, 318, 319 Simpson, George Gaylord, 5, 11, 93, 127, 153, 175, 237, 333 Sinervo, Barry, 118, 121 Singer, Peter, 33, 121, 200, 203, 278 Sisyphean genotype, 18, 121, 144, 146 Sisyphus, 18, 144, 146, 315 Skepticism, 140, 284 Slobodkin, Lawrence B., 335, 336 Sober, Eliott, 47, 81, 82, 96n29, 177, 229, 312, 331, 337 Social adaptations, 61, 110, 168, 193 Social Darwinism, 192, 196, 197, 274 Social donors, 41, 62, 84 Sociobiological Expansion, 190, 191, 199, 201, 203, 206, 311, 314 Sociobiology, 16, 29, 32–36, 72, 88, 99–128, 133–159, 167–170, 189–218, 236, 308–312, 315, 323–325, 330, 332, 333 Socrates, 43 Soma, 50, 52, 55–57, 119, 171–173, 193, 242, 305 Spandrel, 101 Species, benefit to or of, 87, 90, 100, 208 Species, essentialism, 248 Spencer, Herbert, 192, 197

 INDEX 

Stearns, Stephen C., 29, 35, 59, 60, 99, 119, 146, 182, 230, 232, 233, 334, 343, 344 Stebbins, G. Ledyard, 5, 11, 12, 25n16, 93, 333 Sterelny, Kim, 12, 305 Stimulus, 84, 86, 92–94 Stone Age Humans, 270, 279 Stone, Robert, 214 Stony Brook University, 2, 21, 335 Stony Brook University, Symposium 2004, 325 Storytelling, 45, 89, 170, 171, 179, 295–299, 302 Strawson, Galen, 218 Stupidity, of evolutionary process, 30, 202 Style, 4, 6, 7, 11, 12, 15–18, 23, 30, 37, 39, 52, 71, 78–81, 92–94, 101–104, 116, 123, 127, 171, 183, 206, 228, 240, 265, 268, 269, 285, 308, 335–336 Style, plain, 5, 14, 38, 94, 99, 101, 102, 113, 228 Suicide, 310, 311, 314–316 Survival, 19, 22, 35, 42, 48, 55, 60, 73, 76, 77, 82, 84, 85, 87, 89, 103, 105–108, 111, 118, 119, 122, 133, 136, 145, 156, 157, 178, 182, 198, 202, 216, 243, 247, 251, 252, 258, 341, 351n86 Suspicion, 189, 190, 200, 202 Syntax, 10, 76, 93, 114, 206, 247 T Teilhard de Chardin, Pierre, 20 Teleology, 60, 104, 105, 109, 113, 114, 116, 127 Teleonomy, 113–116 Tennyson, Alfred Lord, 199, 215, 338–340, 342

363

Testing stories, 170–171 Thayer, Bradley A., 297, 298 Thoreau, Henry David, 3, 89, 204 Thornhill, Randy, 86, 158 Thought experiments, 17, 71, 169, 201, 239–241, 326 Tinbergen, Niko, 16, 22, 32, 33, 46, 49, 74–76, 83, 86, 87, 101, 115, 180, 194, 244, 268 Tithonus Error, 288, 338, 340–342 Tooby, John, 279 Tradeoffs, 42, 59, 118, 121, 182, 256, 270–272, 304, 329, 344 Traill, David A., 295 Trees, evolutionary, 126, 233 Tree thinking, 126 Trivers, Robert, 13, 21, 31, 33, 42, 64, 104, 121, 134–136, 139, 140, 149, 150, 152, 168, 200, 215, 216, 335, 336 Twain, Mark, 18, 298, 299 U UCLA, 2, 44 Universal Darwinism, 239, 281 University of California, Berkeley, 16 V Van Valen, Leigh, 123, 233, 251, 271 Velasco, Joel C., 125 Vertebrate eye, 89, 254–256 Vitalism, 22, 35, 83, 96n38, 227, 237–239 Viviparity, 109 W Waddington, Conrad Hal, 180 Wallace Stevens, 256 Weismann, August, 23, 49, 50, 193

364 

INDEX

Welles, Sam, 12 West Eberhardt, Mary Jane, 313 Williams, Doris, 7, 8, 44, 62, 63, 84, 92, 123, 193, 343 Williams, George C., 1, 2, 4–9, 13–16, 21, 29, 30, 32, 33, 40, 44, 58, 61–63, 71, 84, 93, 94, 102, 118, 135, 158, 170, 171, 203, 218, 234, 235, 259, 265, 276, 303, 317, 327, 329, 330, 332, 334, 335, 342–345, 348n25 as adaptationist, 9, 32, 36, 89, 90, 123, 166, 176, 177, 179, 238, 252, 259, 268, 269, 297–299 on Buddhism, 218, 306 double doubts, 229 as maladaptationist, 90 as reader of texts, 237 reminisces, 102, 154, 237 restraint, 45, 90, 101, 104, 116, 228 retrospectives, 87 satire, 127, 206, 207, 298, 337, 345 self-assessment, 20

soliloquys, 229, 342 style, 4, 5, 7, 17, 23, 30, 38, 52, 71, 80, 99, 102, 103, 113, 116, 206, 335–336 writing strategies, 216 Williams, Raymond, 3, 44, 266, 316 Williams Principle, 121, 331 Wilson, David Sloan, 31, 64, 239, 312, 326, 331–333, 337 Wilson, E. O., 10, 21, 31–34, 39, 62, 135, 167, 168, 171, 182, 191, 192, 194, 195, 197, 200, 211, 216–218, 259, 266, 288, 309–315, 323, 324, 332–334 Wilson, Margo, 325, 329 Wimsatt, William, 177 Wright, Sewall, 11, 13, 18, 104, 152, 153, 228, 333 Wynne-Edwards, V. C., 78, 333 Z Zimmer, Carl, 194 Zygon magazine, 189, 195, 196, 203, 204, 206