Flowers and Honeybees: A Study of Morality In Nature [Critical ed.] 9789004428539, 9789004428546, 9004428534

Citation preview

Flowers and Honeybees

Critical Plant Studies PHILOSOPHY, LITERATURE, CULTURE

Series Editor Michael Marder (IKERBASQUE/The University of the Basque Country, Vitoria)

VOLUME 6

The titles published in this series are listed at brill.com/cpst

Flowers and Honeybees A Study of Morality in Nature By

Christopher Ketcham

LEIDEN | BOSTON

Cover illustration: Flowers and Honeybees, by Scott Ketcham. Used with permission of the artist. Back cover illustration: Nexus, by Scott Ketcham. Used with permission of the artist. Library of Congress Cataloging-in-Publication Data Names: Ketcham, Christopher, author. Title: Flowers and honeybees : a study of morality in nature / by  Christopher Ketcham. Description: Leiden ; Boston : Brill Rodopi, 2020. | Series: Critical plant  studies, 2213-0659 ; 6 | Includes bibliographical references and index. Identifiers: LCCN 2020013324 (print) | LCCN 2020013325 (ebook) | ISBN  9789004428539 (hardback) | ISBN 9789004428546 (ebook) Subjects: LCSH: Philosophy of nature. | Nature—Moral and ethical aspects.  | Flowers. | Honeybee. Classification: LCC BD581 .K478 2020 (print) | LCC BD581 (ebook) | DDC  171/.7—dc23 LC record available at https://lccn.loc.gov/2020013324 LC ebook record available at https://lccn.loc.gov/2020013325

Typeface for the Latin, Greek, and Cyrillic scripts: “Brill”. See and download: brill.com/brill-typeface. ISSN 2213-0659 ISBN 978-90-04-42853-9 (hardback) ISBN 978-90-04-42854-6 (e-book) Copyright 2020 by Koninklijke Brill NV, Leiden, The Netherlands. Koninklijke Brill NV incorporates the imprints Brill, Brill Hes & De Graaf, Brill Nijhoff, Brill Rodopi, Brill Sense, Hotei Publishing, mentis Verlag, Verlag Ferdinand Schöningh and Wilhelm Fink Verlag. All rights reserved. No part of this publication may be reproduced, translated, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, without prior written permission from the publisher. Authorization to photocopy items for internal or personal use is granted by Koninklijke Brill NV provided that the appropriate fees are paid directly to The Copyright Clearance Center, 222 Rosewood Drive, Suite 910, Danvers, MA 01923, USA. Fees are subject to change. This book is printed on acid-free paper and produced in a sustainable manner.

Contents Acknowledgements ix Preface: Introducing the Meadow x Introduction 1 1 The Question This Study Explores 1 2 The Shape of This Study 7 Cited References 13 1 Optimization, MEP, and Mutualism 15 1 Introduction 15 2 Optimization 16 3 Maximum Entropy Production (MEP) 23 4 Mutualism 33 Cited References 39 2 Emergence of the Flower and Honeybee Mutualism and Flower and Honeybee Ontology and Morphology 44 1 Introduction 44 2 Evolution of the Flower Honeybee Mutualism 45 2.1 Emergence of Flowers and Their Pollinators 45 2.2 The Dawn of Angiosperms 45 2.3 Pollen Likely Attracted Insects First 48 3 Emergence 54 4 Angiosperm Morphology 56 5 Flower Morphology 59 6 Honeybee Eusociality and Morphology 61 6.1 Honeybee Eusociality 61 6.2 Honeybee Anatomy 63 6.3 The Honeybee Caste System 66 7 The Moral Honeybee 67 Cited References 69 3 Flower and Honeybee Epistemology and Behavior 72 1 Introduction 72 2 Angiosperm Epistemology and Behavior 74 2.1 Plant Knowing 74 2.2 Plants and Light 77

vi

Contents

3

4

5 6

2.3 Plants and Chemical ‘Smells’ 79 2.4 Plant Tactile Senses 81 2.5 Plants Don’t Feel Pain 81 2.6 Plants and Sound 82 2.7 Plants Can Sense Gravity 83 2.8 Plant Memory 84 2.9 Tree Interdependence? 86 Plant Intelligence—a Philosophical Discussion 88 3.1 Thinking and Being 88 3.2 The Flowering Plant as a Presence in the Present 91 3.3 Non-Duality in Plants 93 3.4 Plant Intelligence as Distributed Intelligence 95 3.5 Intelligence and the Flower and Honeybee Facultative Mutualism 99 Honeybee Epistemology and Behavior 102 4.1 Basic Honeybee Existence 102 4.2 Honeybee Foraging Behavior 103 4.3 The Honeybee Foraging Waggle, Tremble, and Bump Dances 106 4.4 Honeybee Personality 111 4.5 House Hunting Waggle Dance 114 Consciousness in Flowers and Honeybees 117 Moral Elegance 126 Cited References 127

4 Epigenetics 132 1 Epigenetics Defined 132 2 Promise of Epigenetics 134 3 Epigenetic Purposes 135 4 General Implications of Epigenetics 139 5 Implications of Epigenetics for Flowers and Honeybees 140 Cited References 142 5 The Good and the Emergence of Morality in the Flower and Honeybee Mutualism 144 1 Introduction 144 2 Asymmetricity 144 3 Responsibility 151 4 Reciprocal Responsibility 154 5 Up from Value 156

Contents

6 Hospitality 160 7 Pragmatic Naturalism 162 8 Altruism 164 8.1 Sociobiology 164 8.2 Three Forms of Altruism 165 8.3 Altruistic Punishment 167 9 Singer’s Requirements for Morality to Emerge Applied to Flowers and Honeybees 174 10 Epigenetic Rules 179 10.1 Animal Epigenetic Rules 179 10.2 Plant Epigenetic Rules 185 11 Naturalistic Fallacies and Naturalistic Facts 186 11.1 David Hume 186 11.2 G. E. Moore 188 11.3 An Anti-Naturalistic Fallacy 194 11.4 Optimization 196 12 Flower and Honeybee Oughts and Obligates 197 13 Morality in Nature 199 Cited References 201 6 Study Summary and a Critique of Maximization 206 1 Study Summary 206 1.1 Key Findings of This Study 214 2 A Brief and Preliminary Critique of Maximization 215 Cited References 223 Index 225

vii

Acknowledgements I would like to thank my wife Jan for giving me time to write this book and the encouragement to do so. Thank you also to the philosophy faculty at West Chester University of Pennsylvania, who graciously gave me time and a forum to present my findings to, students, faculty, and members of the public. Credit goes to my brother Scott W. Ketcham for his cover artwork that visually conveys what this project tries to accomplish. The illustrations in this study are by Mattias Lanas who penned detailed and crisp renditions of flower and honeybee morphology. Finally, thank you to all the flower and honeybee field and laboratory researchers who have and continue to unveil the stories of these fascinating and important creatures.

Preface: Introducing the Meadow The farmhouse is surrounded by meadow. The meadow has been fertilized for millennia by the weathering Berkshire hills and three streams that cut through the property. For most of the nineteenth and part of the twentieth centuries cows and horses fertilized the land with their droppings. That ended when the farmers retired. Once tilled fields of corn, beans, and hay have returned to meadow. Long before I was born, honeybees found a crack in the siding of the farmhouse. Between the clapboard outer and plaster inner walls they built their hive. Even after years of swarming to create new hives, the descendants of those first honeybees in the farmhouse thrived. In late July when Cicadas begin their siren calls, the scent of rotting honey settled in the second-floor bedrooms. Rotting honey smells of vinegar and birthday cake hot from the oven. Just after lunch when we are small and have our naps, we drift off to sleep in air steeped in the scent of honey. The first hard frost kills the flowers and the leaves fall and then the meadow lets go of its shades of green and puts on its blanket of brown. The snows come and its plants become dormant. Honeybees huddle in the hive between the farmhouse walls, vibrating against each other to create warmth. What honey has been made in summer is consumed during the winter months, hopefully enough to last until the first flowers bloom. Later, the snow melts and the ice turns black. Then one morning the water glimmers from the sun that clears the hilltops much earlier than in winter. Out in the meadow the honeybees forage the flowers that color the trees and the bushy wisteria. Out of gaps in the winter weathered straw, bulbous plants send up green shoots that emerge with the last of the snow melt. Then iris and tulips bloom in all colors during the first warm days. In late spring and early summer there are periods of bright blooms, some in the same places from year to year, some different. In the middle of sum­ mer when there is no rain, the meadow pauses and only the hardiest of flowers bloom. Late in summer as the heat begins to abate, the goldenrod that has been growing tall all summer pops open like a yellow carpet until the first hard frost. However, the hardy chrysanthemums bloom even after the first frost. Throughout the flowering season, the honeybees forage for nectar and wear pantaloons of yellow pollen on their hind legs. They flit from flower to flower, fertilizing them and assuring the continuity of flowers into the next season. In the hive, the hive workers busily turn floral nectar into honey for consumption by all in the hive including those growing in the nursery, and into storage honeycombs that can feed the honeybees over winter and when flowers do not

Preface: Introducing the Meadow

xi

bloom. Throughout the summer new honeybee workers emerge to first tend the hive and then, for most, to a short twenty-day life of foraging. The farmhouse is the home of the honeybee, but the meadow is her hunting ground. Once a year, half the hive consisting of honey-gorged worker bees swarm in a golden mass often at the upstairs window of the farmhouse. They buzz a din that can be heard throughout the bedrooms. Then suddenly they are gone with the old queen who has been denied food to slim her down so that she can fly to a new place where they will build a new hive. The new queen remains in the walls of the farmhouse to continue the effort to maintain the hive’s continuity in a place where honeybee history has long endured. We take for granted the work of the honeybee. We ignore their presence even as we enjoy the meadow as the afternoon sun angles down behind the hills. However, the declining sun shows how busy the air is above the meadow, punctuated by tornadoes of tiny bugs here and there. Dragonflies zip from one end of the field to the other, tilling the air like the farmer in a field. Veer they do like fighter planes and then resume their tilling. A Flycatcher lands on a tall stem, bending it near to the ground. Then she is off with a flit, to return sometime later to wait for the next bug to capture her attention. The honeybees fly straight from one flower to another, whatever flower may be available this day, this season, this moment in time. Then they slow and sometimes hover. At other times they simply slow and enter the flower straight away. Morning dew shows how many spider webs have been spun in the meadow. The spiders compete with the flowers for their pollinators. Even so, there are too many bugs in the meadow to count. At times there are sated spiders and honeybees with bright yellow baskets of pollen on their hind legs. At other times when few flowers bloom the spiders must wait and endure hunger. Meanwhile the honeybees have all fanned out far beyond the meadow looking for signs of flowers. The smell of honey in the farmhouse peaks at the height of summer and drifts slowly away as the cool of August-end settles in the valley. The days are shorter than they were in July. The fall flowers are beginning to bloom. The honeybees must continue to find them if they are to survive the winter. There are fewer flowers as the temperature drops and the spiders, tornado bugs, and honeybees must wait for the sun to warm the air before they can venture out. The dragon flies seem more hurried than in midsummer. The bounty now is less and the time brief for the chase. There is a median that runs through the year and this median bears its own sense of urgency. Urgency is absolute, but the response from creatures in the meadow is measured, not frantic. The rhythm of the meadow is a syncopation of weather and life. While the beat changes with the rhythm of the meadow,

xii

Preface: Introducing the Meadow

the tempo is always at the maximum sustainable pace or what this study calls optimal pace. The elegance of the meadow is that it is never more or never less than it can be. Even when humans have driven a plow through the meadow, the meadow eventually relaxes back into a rhythm that it can sustain. We can learn from the meadow.

Introduction 1

The Question This Study Explores

This study considers the question: “Is morality solely a human creation or can we discover evidence of or the antecedents of morality in nature?” I suggest that the antecedents of morality can be found in life’s existential propensity towards optimization, choosing the best among different options considering current conditions. Choosing the best option is ultimately towards preservation of the individual for purposes of genetic replication, whether direct as with a flower who can reproduce, or a sterile worker honeybee who cannot, but whose actions benefit the hive and the queen and males who can.1 There is an historic purpose for optimization. After life emerged in the early earth, it has survived many extinctions. It seems as if life’s overarching goal is to maintain the continuity of life. This continuity goal is not concerned with the success of any individual life form, but of life itself to survive and reproduce in perpetuity. Life experiments through mutation which is not always beneficial to the resulting life form. However, life also gives the living the possibility of epigenetically expressing genes (without mutation) in an alternative way that can be advantageous to the living life form. Life also gives many life-forms the ability to respond to stimulus, learn and, in some cases, pass this learning epigenetically or through teaching down to future generations. Life is not inherently cruel, because it provides creatures with ways towards optimizing their existence in their world. I suggest that it is not the notion that life seeks to renew itself in perpetuity that is the good, but that the process of optimization that many life forms employ towards this end is that which is towards the good. Think of it this way, if every creature always chose the decision to achieve the maximum result, then it likely would do whatever it took to achieve that objective without even considering risk to itself. Anything that stands in its way, even if not a threat to the individual, is likely to be attacked as interfering with its maximization effort. There would 1  The word ‘ultimate’ is used because each optimal decision is not necessarily made with self-preservation or even hive preservation in mind. As Nathaniel Barrett explains, “Self-preservation has the mark of motivation insofar as it seems to stem from an openended organizing principle of behavior rather than a set of instructions” Nathaniel Barrett, “On the Nature and Origins of Cognition as a Form of Motivated Activity,” Adaptive Behavior 2019, no. January (2019): 6. The honeybee’s motivation to forage is likely an organizing principle of day-to-day behavior, even though she ultimately produces the good for the hive by bringing home resources that can be turned into food.

© Koninklijke Brill NV, Leiden, 2020 | doi:10.1163/9789004428546_002

2

Introduction

be carnage and descent into Thomas Hobbes’ state of nature he maintained is, “[s]olitary, poor, nasty, brutish, and short.”2 Few might live to maturity as a result, and life would not be able to fill as many niches in the world as it does today. Therefore, optimization is not only contrary to Hobbes’ state of nature, but also contributes ultimately to the continuity of life itself. I suggest that optimization is fundamentally towards the good and primordial to the emergence of morality in nature itself. Given that reproduction of the genome is necessary for a species to thrive and evolve, optimization is an important process indeed. The process of optimization avoids unnecessary conflict and confrontation that is inherent in Hobbes’ state of nature. However, optimization is only towards the good. We must look at other aspects of existence for the emergence of morality in nature. Specifically, I suggest that studying the flower and honeybee facultative mutualism will provide an example where morality has emerged in nature.3 The theoretical underpinning of this assertion I take from Peter Singer’s three requirements for morality to emerge: the social group, restraint towards other members of the group, and judgment.4 This study maintains that the flower and honeybee facultative mutualism constitutes a social group, there is restraint between members of this social group, and that judgment is fundamental to the process of optimization that both species exhibit, both in their separate species’ decisions, and those involving the mutualism. The good comes from a condition of co-existence that emerges from the practice of optimization by both flowers and honeybees. The good arising in the facultative mutualism involves optimizing individual behavior in a way that does not intentionally cause undo harm either to self or others in the mutualism.5 This definition of the good is ideal because there will be exceptions where individuals may intentionally harm others. However, 2  Thomas Hobbes, Leviathan (London: Andrew Cooke, 1651), 78. 3  The honeybee family, or Apis Mellifera, consists of variants of an original species from Eurasia. When this study uses the term ‘honeybee’ that means any variant. Angiosperms are a family of hundreds of thousands of species. When this study flowers, in the generic sense in context of honeybees, that means only those flowers that honeybees forage. Since honeybees are found all over the world, this is likely thousands or even more potential species. Therefore both ‘flowers’ and ‘honeybees’ are useful generics that serve to simplify the concept that is their facultative mutualism. 4  Peter Singer, The Expanding Circle (Princeton, N.J.: Princeton University Press, 1981, 2011), 4, 5, 70. 5  Worker honeybees sting intruders and die. While this is ultimately a suicidal action, likely the worker is unaware of the consequences. There are many who believe that the hive is a superorganism and such an action of self-sacrifice represents only a small loss to the greater organism.

Introduction

3

in their facultative mutualism, flowers and honeybees do not usually cause undue harm to each other. Even though the flower and honeybee facultative mutualism exhibits goodness, and because there will also be genetic or even personality differences in individuals, there will be actors who make less than optimal choices that others would not have selected in the same circumstances. On balance, I suggest that the flower and honeybee facultative mutualism that emerges from generally optimal behavior by both species shows evidence of the moral notion of the good emerging in nature. Species in a facultative mutualism both exploit and benefit the other for at least one existential requirement—flowers exploit honeybee mobility to assist their reproductive needs; honeybees exploit flower pollen and nectar creation to enable the production of food for the hive. The flower and honeybee facultative mutualism is not exclusive because flowers have many insect or other pollinators and honeybees will forage any promising flower. Both have co-evolved to optimize their relationship and neither causes undue harm to the other.6 Optimization is also towards the avoidance of unnecessary risk. There­ fore, optimization inherently involves risk management. Choosing the best option based upon conditions requires avoiding conflict where possible, including not interfering with other life forms unnecessarily. Non-interference means that individual life forms tend to avoid participating in or precipitating conditions where unnecessary conflict could arise. The process of optimization is not only towards the teleological—passing down of genes—but is also towards optimal energy acquisition and use over time. Each life form must either make or acquire energy in order to survive. Life does so by taking the building blocks for energy production out of the environment or ecology. Through energy acquisition and use, life increases entropy, and likely increases entropy more than abiotic states.7 Research is discovering that the practice of optimization by life forms is by-and-large consistent with Newton’s second law of thermodynamics or entropy. Axel Kleidon and Ralph Lorenz explain earth’s state, “The Earth is a non-equilibrium system in a steady state.”8 The earth receives radiation from the sun and elsewhere and emits its 6  See: Judith L. Bronstein, “The Study of Mutualism,” in Mutualism, ed. Judith L. Bronstein (Oxford, UK: Oxford University Press, 2015), 11. 7  See: Joseph J. Vallino, “Ecosystem Biogeochemistry Considered as a Distributed Metabolic Network Ordered by Maximum Entropy Production,” Philosophical Transactions: Biological Sciences 365, no. 1545 (2010): 1424. 8   Axel Kleidon and Raph Lorenz, “Entropy Production by Earth System Processes,” in Non-Equilibrium Thermodynamics and the Production of Entropy: Life, Earth, and Beyond, ed. Axel Kleidon and Ralph Lorenz (Heidelberg, Germany: Springer 2005), 2.

4

Introduction

own radiation which produces a balance or steady state system. Life is an essential part of this balance today. Maximizing entropy is subject to environmental conditions that change over time, whether from natural forces like weather and volcanos, or the activities of life. Alternatively, in a closed bottle in space (a linear thermodynamic system), gas at one end will tend to dissipate until the gas is equally distributed in the bottle. This theoretical bottle receives no new input or output and therefore once maximum entropy is achieved, there is no further entropy, and there is no more information that can be derived from observing the system. However, since the earth continues to receive new solar radiation and expend its own, it is in continual flux as it tries to maintain its steady state, producing both more entropy and more information about the system. Life and other forces on earth tend towards maximum entropy like the bottle in space. However, earthly conditions restrain, for example, the equal distribution of entropic life forms e.g., desert versus jungle. Science is discovering that life tends to engage in processes that are towards Maximum Entropy Production (MEP) a theory that suggests when individual creatures optimize their behaviors, they tend to expend energy at a rate that is maximal according to current conditions.9 This is an important observation because it suggests that the propensity of life to behave towards optimizing energy use according to conditions, not only does not violate the second law of thermodynamics, but also serves to exemplify the operation of the second law. I suggest that optimization is a precondition for the emergence of moral­ i­ty. I am not suggesting that optimization is moral, but I am also not saying that it is not moral. As a precondition for morality, optimization helps facilitate the emergence of morality in circumstances where morality becomes part of the optimization process itself. This study considers the emergence of morality through the precondition of optimization in the facultative mutualism of flowers and honeybees. Honeybees (and pollinators in general) are required for the flower to consummate its sex act; and honeybees depend on flowers to produce energy resources for themselves and the hive. I suggest that optimization behaviors of both flowers and honeybees in their facultative mutualism are consistent with Singer’s requirements for morality to emerge. This study explores how the behaviors of both species show restraint for each 9  M EP is explored later in this study. For basic information see: Axel Kleidon, “Non­ equilibrium Thermodynamics and Maximum Entropy Production in the Earth System,” Naturwissenschaften 96, no. 6 (2009); “A Basic Introduction to the Thermodynamics of the Earth System Far from Equilibrium and Maximum Entropy Production,” Philosophical Transactions: Biological Sciences 365, no. 1545 (2010); Leonid M. Martyushev, “The Maximum Entropy Production Principle: Two Basic Questions,” Philosophical Transactions of the Royal Society B: Biological Sciences 365, no. 2010 (2010).

Introduction

5

other and how their individual judgments (optimal in the aggregate) serve both to sustain the mutualism as well as each species. Most social groups, including human, the eusocial honeybees and ants, and even the vast herds of the African savannah are generally comprised of a single species (though wildebeest and zebras often travel together). I maintain that the flower and honeybee facultative mutualism is a special kind of social group that involves dissimilar species who require each other in order to survive and reproduce. We can call the beehive a social group, just as we can call the wolf pack a social group, along with the prairie dog town, and the human city. In each of these, members of a single species constitute the group. There may also be rudimentary social groups in flowering plants. A recent study of tomato plants found that mycorrhizal networks on plant roots transmit information from pathogen infected tomato plants to other tomato plants.10 The flower and honeybee facultative mutual social group is different from the hive and the tomato plant relationship because it involves plant species and an insect species who are more unalike than alike. They are a social group for two very important reasons. First, they require each other. Second, they interact with each other with restraint during some part of their life cycle, but not the entire cycle. Human social groups are often not permanent or contain the same membership all the time. A person may enter and leave different social groups every day, beginning with family, then school or work, then club or organization, and back to family again. A person leaves the social group school upon graduation and another leaves a club by resigning. The graduate then may enter a new social group called work and the person who resigns from one club may join another. Flowering plants advertise their bloom; honeybees are attracted to the bloom and begin social interactions by entering the flower to secure nectar and pollen. Honeybees then distribute pollen to other flowering plants of the same species. When the flowering plant ceases to bloom, the honeybee no longer interacts with the plant. The foraging honeybee worker likely does not know she serves as pollinator for the flower. She knows that she secures what the hive requires from the flower. However, since each flower only produces a small part of what she and the hive need, she must fly from flower to flower and therefore ‘pays back’ the flower for her effort to produce the nectar.11 We can suggest that this relationship is also a form of commerce. The honeybee 10  See: Yuan Yuan Song et al., “Interplant Communication of Tomato Plants through Under­ ground Common Mycorrhizal Networks,” PloS one 5, no. 10 (2010): 2. 11  Most flowering plants that female honeybee foragers visit are bi-sexual. The feminine ‘she’ pronoun is used rather than ‘they’ which at times is awkward in a sentence. It is a female honeybee who transports the male gamete (pollen) from one flower to another where it can pollenate the ovule and produce seeds, or the young of the plant.

6

Introduction

gets paid in energy producing resources for her expense of energy distributing the pollen of the flower. The flower expends energy producing the nectar, but she is rewarded with genetic continuity. Note that like human commerce, the exchange is one commodity for a different commodity. For example, the honeybee is not food for the flower, but what the flower produces becomes food for the honeybee. Even the Venus Fly Trap who dines on flies who are attracted to the sweet smell in her vice-like leaves, produces a long stalk to her flowers so that pollinators do not become entrapped. Because the commodities exchanged are different (asymmetrical), there is no need for flowers and honeybees to compete. However, commerce cannot take place without rules of engagement. The facultative mutualism serves both species through its ontological co-evolved structure (supplier and buyer) and the acts of individuals who abide by Singer’s moral prerequisites of restraint and judgment. Therefore, it is possible to maintain that the flower and honeybee facultative mutualism is a social group that also has a mutually beneficial commerce function. The flower and honeybee mutualism is not exclusive. Many flowers have many different pollinators: bees, wasps, beetles, butterflies, hummingbirds. Honeybees will forage any flower that it considers promising. What emerges is a social group that has evolved over time using co-evolutionary processes that include genetic mutation towards mutual benefit and the active behaviors of individuals that maintain the mutualism. The flower and honeybee facultative mutualism has evolved over at least one million years from the emergence of honeybees. Honeybee antecedents (apoid wasps) began to evolve towards mutualism with flowers around one hundred million years ago, which means flower and pollinator mutualisms survived the dinosaur extinction. While the ontological and morphological requirements for the mutualism have coevolved for both flowers and honeybees over time, the mutualism is maintained by epistemological means—judgment and behavior—over the existential horizon of individuals of each species. Because individuals from both species use judgment to restrain from behaving adversely towards each other while they exploit the existential benefits that the other provides, together they form a social group that benefits and exploits each other. This study maintains that the flower and honeybee facultative mutualism is towards the good and meets the minimum requirements for conditions for the natural emergence of Singer’s morality. However, since this is a study of only one facultative mutualism in nature, I will not claim that all facultative mutualisms are structurally towards the good. Studies of other mutualisms are required to assess the emergence of morality (or not) for other than flowers and honeybees.

Introduction

7

Why study the flower and honeybee facultative mutualism for an origin of morality? Frans de Wall spent many years studying primates and our closest cousins on the evolutionary tree. He saw and then described behaviors that we would be hard-pressed to suggest do not have moral content.12 It is not so surprising that we might discover the antecedents of morality with those whom we share ninety percent of our DNA. We know that honeybees and ants, who are both insects of the Hymenoptera order, are eusocial and exhibit high degree of intra-colony cooperation. We know of many instances of mutualisms in nature. However, to study flowers and honeybees in context of the emergence of morality poses several challenges. First, there is an insect with a small but well-developed neurological system whose brain is structurally different from our own and other vertebrates but performs many similar functions. Second, no angiosperm has a central nervous system. The two species could not be more different and come from two separate branches of life. Yet they have spent a million years in a facultative mutualism that is both intimate and cooperative. The advantage for considering the emergence of ethics through flowers and honeybees is that both have been well studied by science. We know much about what they can do and are learning more and more about how they do it, including some of the electro-bio-chemical methods that underly their behaviors and processes. If we find even the antecedents of morality in the flower and honeybee facultative mutualism, it will become more difficult to give to humanity moral exclusivity. 2

The Shape of This Study

There is much to explore about flowers and honeybees towards discovering the good that arises in their facultative mutualism. The first four chapters of this study review the scientific literature in many areas of research associated with flowers and honeybees. Categories of scientific research explored in order in this part of the study include optimization, Maximum Entropy Production (MEP), mutualism, ontology and morphology, epistemology and behavior, and epigenetics.

12  For example, see: Frans de Wall, “Morality Evolved Primate Social Instincts, Human Morality, and the Rise and Fall of “Veneer Theory”,” in Primates and Philosophers: How Morality Evolved, ed. Stephen Macedo and Josiah Ober (Princeton, NJ: Princeton University Press, 2006); “The Tower of Morality.”

8

Introduction

This is a book written for both scientists and philosophers. The first four chapters are devoted to science associated with flowers and honeybees that are then related to the moral question this study explores. Some of the studies explored in these chapters are seminal and others remain controversial, but they show the effort of science to understand and explain flower and honeybee behavior and morphology that are then assessed through moral precepts. These chapters may be written in a style more academic than some readers would prefer. However, these chapters are important because they help to establish a scientific basis for the emergence of morality in nature. This study maintains that there is a scientific basis for the emergence of morality in nature. Following the considerable science associated with the fundamentals that underly this assertion are chapters that consider the emergence of moral concepts in the flower and honeybee facultative mutualism in context of the underlying science this study explores. Beyond informing the reader, a purpose for this study is to bring science and philosophy closer together in considering how morality is an emergent property in nature and not just some metaphysical notion without any scientific basis for understanding. Second, we ought to begin considering the purposes that morality might serve plants and animals. For example, the reasoning capabilities both plants and pollinators use that produce optimization, and the considerable responsibility and hospitality they show each other, likely has contributed to the explosion of the number of angiosperm species and pollinating insect species. Third, the flower and honeybee facultative mutualism has lasted a million years. Without restraint from harming the other, responsibility and hospitality towards the other, how long could such a social construct last? If both the scientist and philosopher begin to see the underpinning of morality in natural processes, they both can engage in a more meaningful discourse on morality and how it serves nature. However, this is a study only of the flower and honeybee facultative mutualism. There are many more mutualisms and other social constructs in nature that deserve similar treatment. One frustration the reader will encounter is that human notions of morality are not easily translated into the social construct of such diverse species as flowers and honeybees. Also, if human morality has its antecedents in nature, then we are right to engage in a critique of the anthropocentric view of morality while we search for other manifestations of morality in the world around us. Chapters one through four of this study are written in an academic style. Chapters five and six apply the scientific knowledge and theories to moral theory which has emerged from human-centric studies, but which are now applied to the flower and honeybee facultative mutualism. This study acknowledges the challenges of applying human notions of morality to non-neurological

Introduction

9

flowering plants and an insect species. Jeffrey T. Nealon explains why the fundamental study of life itself is important to the study of morality: Life is not something that’s owned by an organism, something hidden deep within it, to be protected against the outside at all costs; rather, life is the territory for the emergence of ‘interkingdoms,’ assemblages of heterogeneous processes. The animal territory for thematizing life, however important and apt it may be, tends to focus our attention on the biopolitical competition among individual organisms to the detriment of this robust sense of distributed, interconnected life.13 It is the ‘robustness of interconnected life’ that underpins the flower and honeybee facultative mutualism and therefore serves to extend Nealon’s argument for studying it. Chapter one reviews theories of optimization, MEP, and mutualism from the literature that are not exclusive to flowers and honeybees. This study explores how optimization, MEP, and mutualism are important to flower and honeybee behavior. This study later demonstrates how they are also important to the discussion of the emergence of morality in the flower and honeybee relationship. Chapter two first reviews the history of flower and honeybee evolution. Following is a discussion of flower and honeybee ontology and morphology. It is important to understand how both species’ morphology and associated processes have co-evolved to make their facultative mutualism possible. Ontology and morphology are discussed at a basic level, like what can be found in first texts on botany or Hymenoptera. However, cited references can provide the reader and researcher with access to more extensive information. The mutualism requires maintenance. Both species in some part of their existence interact with each other, and how they do so is discussed in the third chapter called epistemology and behavior. The epistemology and behav­ior chapter also considers flowers and honeybees in their existence outside of the mutualism. For honeybees, this includes activities associated with the hive; and for flowers, their activities in association with individual require­ments and interaction with other plant species and herbivores. The epis­te­mol­ogy and behavior chapter discusses the basics but also explores recent research in flowers and honeybees that better clarifies the scope of behaviors and capabilities both species demonstrate. As science begins to discover more about both species, their behaviors as individuals outside of the mutualism and behaviors within their facultative mutualism will come more clearly into 13  Jeffrey T. Nealon, Plant Theory (Stanford, Ca.: Stanford University Press, 2015), 119.

10

Introduction

view. For example, much has been learned how plants communicate availability to honeybees, and how honeybees announce their presence in the flower, both which improve flower and honeybee optimization towards their respective goals in the mutualism. Honeybee communication between hive members has long been studied, but recent research shows just how complicated and important this communication is to hive optimization. Studies of plants show that they can communicate the presence of herbivores and even other information to other plants both through the air, and in some cases, the roots. Chapter four introduces the relatively new science of epigenetics. Epi­ genetics (above genetics) is a process in nature that does not fit easily into ontological/morphological or epistemological/behavioral categories. Many genes can express themselves differently without mutating or changing the genetic sequence in chromosomes. Epigenetics explores how genes can modify their expression based upon stressors. Epigenetics, at least in some plants, can pass down what one generation has learned to future generations without any active teaching. In animals, stress during pregnancy in some studied species produce epigenetic changes in offspring to prepare them to address these stressors after they are born. Things are more complex with animals who rear their young because both nature (genes and epigenetics) and nurturing behavior may influence the capabilities of offspring to survive in their ecology. Epigenetics is generally towards optimization because it tries to prepare offspring for current conditions, serving as an additional existential process that will help the living creature in some cases, or its descendants in other cases, optimize behaviors according to present-day conditions. However, epigenetic processes are quite powerful and are also implicated in the emergence of cancer and other maladies. For this study, epigenetics provides additional data points on how flowers and honeybees use these processes to improve their chances of making optimal decisions both in their activities associated with their facultative mutualism and behaviors and activities outside of their mutualism. Epigenetics is another tool that plants and animals can use to better optimize their existence. Once the basis for and the basics of both flowers and honeybees and their facultative mutualism have been explored, the fifth chapter of this study considers how optimization helps the good emerge for these two species. As this is a study about morality, there are theoretical and practical aspects of morality that require exploration. Theories of naturalistic morality (moral naturalism) or ethics have not been widely accepted and their critique will be explored. This includes the admonition from David Hume that ought cannot be derived exclusively from is. Ought derived exclusively from is, is the first fallacy. Later,

Introduction

11

G. E. Moore suggests that a naturalistic fallacy is committed when one suggests that something is good because it exists in nature.14 Following the critique of naturalistic morality is a discussion from the point of view of proponents of naturalistic morality and what is required for a cogent theory of naturalistic morality to emerge that does not commit the aforementioned fallacies. After exploring both critique and support arguments, this study considers the flower and honeybee mutualism for evidence of naturalistic morality, principally through their behavior towards each other, and in their lives outside of the mutualism. This necessarily involves showing how the flower and honeybee mutualism overcomes many of the objections that critics of naturalistic ethics pose. This study also includes a discussion of how optimization facilitates the emergence of morality in nature. I submit from the outset, that the presence of morality in the flower and honeybee mutualism is in no way as sophisticated as morality explored and developed in the context of human activity. For example, we have no evidence that either honeybees or flowers have any conception of God or a higher being that has been fundamental to the development of much of human moral discourse. However, what this study maintains, is that while human morality may be ultimately more complex than morality found in the flower and honeybee mutualism, we cannot say that humanity has exclusive rights to claim moral theory and behavior as its own. This study maintains that there is, at least in flowers and honeybees, the emergence of the good that has evolved over time with their facultative mutualism construct that is and has been maintained for a million years by the optimal behaviors of individual actors. I suggest that this goodness does not commit the naturalistic fallacy because it is an emergent property of the relationship between flowers and honeybees. Because both flowers and honeybees are in an occasional and non-permanent relationship, and even though their relationship is required for the survival of both, we cannot also claim that the species that contribute to this facultative mutualism are inherently good. The sixth chapter of this study consists of two parts. The first summarizes the results of this study. The second part is a critique of humanity who, in our relationship with nature, tend to make decisions that maximize reward. Humans, rather than balance risk with reward, tend to eliminate risks and other exigencies in order to achieve maximum rewards. However, many human activities alter overall entropy because they involve eliminating life forces that may result in preventing the achievement of maximum entropy production 14  See: David Hume, Treatise of Human Nature (Gutenberg.org: Project Gutenberg, 2012). George Edward Moore, Principia Ethica (Cambridge UK: Cambridge University Press, 1903).

12

Introduction

that may be possible for a given ecology. The example used will be presentday monoculture crop farming practices. Scientists worry that the increase in carbon dioxide in the atmosphere is already producing adverse consequences, including global warming. I suggest that this is an important outcome that should be studied, but that we must also consider an additional source question that may contribute to global warming and that is whether the change of entropy in places where humans have altered ecologies on land and sea may also contribute to global warming and other contemporary problems. Even after experiencing four near-extinction events, life has re-emerged different morphological forms to produce more entropy than if the world was abiotic. Restoring entropic life production processes means reengaging the meadow to produce as much entropy that is possible based upon conditions. Most life produces MEP by making optimal decisions based upon given circumstances rather than through decisions that tend to maximize by eliminating risk or conditions that could give rise to risk. Eliminative processes tend to produce new consequences that, as we see in agricultural practices, likely produce less entropy than the natural meadow whose creatures tend to make optimal decisions based upon present conditions. There are two discourses that this study will not explore in any depth. The first is the very real problem of the disappearance of insects, and more specifically the honeybee through hive collapse syndrome, climate change, and other negative influences. This study presents the honeybee and how she has evolved to her current state of existence. It is for others to continue the good work on addressing these negative influences on insect and honeybee populations. As some angiosperm species are pollinated by means other than insects, mammals, or birds, these flowers would likely survive an insect extinction event. Defining a possible future without insects or a future where science actively seeks to preserve insect populations is for others to consider. This study also only lightly touches on how unfairly, or otherwise philosophy has treated animals and plants in relationship to humans. Michael Marder, Isabelle Stengers, Ilya Prigogine, and Jeffrey T. Nealon have made significant contributions to plant theory in philosophy.15 Even more work is being done

15  See: Ilya Prigogine and Isabelle Stengers, Order out of Chaos, Kindle ed. (London: Verso, 2017); Nealon, Plant Theory; Michael Marder, “Vegetal Anti-Metaphysics: Learning from Plants,” Continental Philosophy Review 44, no. 4 (2011); “Of Plants, and Other Secrets,” Societies 3, no. 1 (2013); Grafts (Minneapolis, Mn.: University of Minnesota Press, 2016); Plant-Thinking: A Philosophy of Vegetal Life, None (New York: Columbia University Press, 2013), Book; “Plant Intentionality and the Phenomenological Framework of Plant Intelligence,” Plant Signaling & Behavior 7, no. 11 (2012); “Plant Intelligence and Attention,”

Introduction

13

on animal ethics that also deserves attention.16 This study presents the flowering plant and the honeybee as co-equal partners in their social group. Both are presented as how they are in nature and not in relationship to any ordering of species in relationship to humans. The challenge of this study is to show these interesting creatures as how they present themselves to the world using the human-conceived notions of morality which, to this point, have resisted easy translation into species other than humans. This study now begins with a discussion of three scientific theories: Optimization, MEP, and mutualism. Cited References Barrett, Nathaniel. “On the Nature and Origins of Cognition as a Form of Moti­ vated Activity.” Adaptive Behavior 2019, no. January (2019): 1–15. doi:10.1177/ 1059712318824325. Bronstein, Judith L. “The Study of Mutualism.” In Mutualism, edited by Judith L. Bronstein, 3–19. Oxford, UK: Oxford University Press, 2015. Hobbes, Thomas. Leviathan. London: Andrew Cooke, 1651. Hume, David. Treatise of Human Nature. Gutenberg.org: Project Gutenberg, 2012. Kleidon, Axel. “A Basic Introduction to the Thermodynamics of the Earth System Far from Equilibrium and Maximum Entropy Production.” Philosophical Transactions: Biological Sciences 365, no. 1545 (May 12 2010): 1303–15. doi:10.1098/rstb.2009.0310. Kleidon, Axel. “Nonequilibrium Thermodynamics and Maximum Entropy Production in the Earth System.” Naturwissenschaften 96, no. 6 (2009/06/01 2009): 1–25. doi: 10.1007/s00114-009-0509-x. Kleidon, Axel, and Raph Lorenz. “Entropy Production by Earth System Processes.” Chap. 1 In Non-Equilibrium Thermodynamics and the Production of Entropy: Life, Earth, and Beyond, edited by Axel Kleidon and Ralph Lorenz, 1–20. Heidelberg, Germany: Springer 2005. Marder, Michael. Grafts. Minneapolis, Mn.: University of Minnesota Press, 2016. Marder, Michael. “Of Plants, and Other Secrets.” Societies 3, no. 1 (2013): 16–23. Marder, Michael. “Plant-Soul: The Elusive Meanings of Vegetative Life.” Journal of Environmental Philosophy 8, no. 1 (2011): 83–100. Plant Signaling & Behavior 8, no. 5 (2013); “Plant-Soul: The Elusive Meanings of Vegetative Life,” Journal of Environmental Philosophy 8, no. 1 (2011). 16  The literature on animal ethics is vast. One might begin with The Journal of Animal Ethics From the Oxford Centre for Animal Ethics: https://www.oxfordanimalethics.com/ what-we-do/publication/journal-of-animal-ethics/.

14

Introduction

Marder, Michael. Plant-Thinking: A Philosophy of Vegetal Life [in English]. None. New York: Columbia University Press, 2013. Book. Marder, Michael. “Plant Intelligence and Attention.” Plant Signaling & Behavior 8, no. 5 (2013): e23902-e02. doi:10.4161/psb.23902. Marder, Michael. “Plant Intentionality and the Phenomenological Framework of Plant Intelligence.” Plant Signaling & Behavior 7, no. 11 (2012): 1365–72. doi:10.4161/ psb.21954. Marder, Michael. “Vegetal Anti-Metaphysics: Learning from Plants.” Continental Philosophy Review 44, no. 4 (2011/11/01 2011): 469–89. doi:10.1007/s11007-011-9201-x. Martyushev, Leonid M. “The Maximum Entropy Production Principle: Two Basic Questions.” Philosophical Transactions of the Royal Society B: Biological Sciences 365, no. 2010 (May 12 2010): 1333–34. doi:10.1098/rstb.2009.0295. Moore, George Edward. Principia Ethica. Cambridge UK: Cambridge University Press, 1903. Nealon, Jeffrey T. Plant Theory. Stanford, Ca.: Stanford University Press, 2015. Prigogine, Ilya, and Isabelle Stengers. Order out of Chaos. Kindle ed. London: Verso, 2017. Singer, Peter. The Expanding Circle. Princeton, N.J.: Princeton University Press, 1981, 2011. Song, Yuan Yuan, Ren Sen Zeng, Jian Feng Xu, Jun Li, Xiang Shen, and Woldemariam Gebrehiwot Yihdego. “Interplant Communication of Tomato Plants through Underground Common Mycorrhizal Networks.” PloS one 5, no. 10 (2010): e13324, 1–11. doi:10.1371/journal.pone.0013324. Vallino, Joseph J. “Ecosystem Biogeochemistry Considered as a Distributed Metabolic Network Ordered by Maximum Entropy Production.” Philosophical Transactions: Biological Sciences 365, no. 1545 (2010): 1417–27. Wall, Frans de. “Morality Evolved Primate Social Instincts, Human Morality, and the Rise and Fall of “Veneer Theory”.” In Primates and Philosophers: How Morality Evolved, edited by Stephen Macedo and Josiah Ober, 1–82. Princeton, NJ: Princeton University Press, 2006. Wall, Frans de. “The Tower of Morality.” In Primates and Philosophers: How Morality Evolved, edited by Stephen Macedo and Josiah Ober, 161–81. Princeton, NJ: Princeton University Press, 2006.

Chapter 1

Optimization, MEP, and Mutualism 1 Introduction This chapter introduces three theories. The first, optimization, is something that is studied in many contexts, including biology, chemistry, economics, engineering, logistics, manufacturing, and computer science. The postal service generally establishes fixed regular letter delivery routes because most homes receive letter mail each day. For them, the fixed letter-carrier route is optimal. The parcel delivery service does not deliver daily to most homes. Each day requires the creation of an optimal route based upon many factors including delivery locations, timed deliveries, and energy efficiency including the weight of the load as it decreases throughout the day. In another example, internet search engines sometimes return millions of hits. They develop optimization routines to present the most relevant returns first to the searcher according to their research on what people generally are looking for with this term … of course, after placing relevant advertisements first. For this study, honeybees optimize their flight to known foraging locations and watch waggle dancers in the hive to discover new locations to which they can fly directly. Flowers have evolved to bloom when pollinators are available and perhaps when direct competition from other species are not blooming. These are all examples of optimal behavior. The second theory, Maximum Entropy Production, or MEP is the product of optimization by many species in the ecology. Newton’s second law of thermodynamics maintains that systems tend to increase in entropy over time. MEP is process theory that considers what kind of entropy is produced in non-linear thermodynamic systems like the meadow. These non-linear systems introduce constraints such as life that create entropy at different rates than linear systems. If living things are behaving optimally, considering the constraints that the ecology places in their way, then MEP theory suggests that maximum entropy likely is being produced. However, MEP is still a new theory, and what maximum entropy means in various contexts is still being studied. There is no consensus that MEP processes are universal in nature and there may be instances where MEP processes are not relevant in some life or ecological activities. The third theory is the mutualism. There are many mutualisms in nature where two species rely on each other for at least one life process or function.

© Koninklijke Brill NV, Leiden, 2020 | doi:10.1163/9789004428546_003

16

Chapter 1

Flowers and honeybees are a familiar mutualism to most. However, mammals and their gut bacteria are a mutualism. Gut bacteria breaks down food into chemicals that animals can use, and both species gain nourishment by doing so. Most plant roots are in a mutualism with mycorrhizal fungi that perform similar functions as mammalian gut bacteria, preparing essential nutrients like phosphorous and nitrogen for the roots to absorb, and in turn, the mycorrhizae feast on the sugary liquid in the plant’s roots. Lichen are an obligate mutualism of algae and fungus. Even our own eukaryotic cells (most animal cells) are believed to have incorporated their mitochondria from a bacterium that began a mutualism relationship with the eukaryote many millions of years ago. 2 Optimization Central to Singer’s requirements for morality to emerge is the agent’s capability to judge which requires reasoning. Optimization requires judgment and reasoning at several levels: the individual, both flowers and honeybees within their social groups, the extended society in the meadow, and the combined reason of billions of actors in a more global ecology. For this study, optimization is defined as choosing the ‘best’ decision among different options.1 ‘Best’ is contingent on the circumstances, and what is best can change from moment to moment. Optimization has been discovered not only in individual animal behaviors, but in evolutionary practices and larger ecological systems. While this study primarily considers behaviors of flowers and honeybees in their facultative mutualism and in extra-mutualism activities, this study reviews select studies of evolutionary and ecological optimization. First, is a discussion of optimization behaviors. Optimization in nature is not only subject to external constraints and contingencies, but also internal needs and goals of the individual. Personality, such as aggressiveness or risk adversity (both are associated with personal risk appetite) plays a role in determining optimal decisions. The individual also has affordances and constraints that come from being a member of a species. The honeybee has the option to fly from danger; the flower does not. The 1  “Optimization is central to any problem involving decision making, whether in engineering or in economics. The task of decision making entails choosing among various alternatives. This choice is governed by our desire to make the ‘best’ decision. The measure of goodness of the alternatives is described by an objective function or performance index. Optimization theory and methods deal with selecting the best alternative in the sense of the given objective function” Edwin K. P. Chong and Stanislaw H. Zak, An Introduction to Optimization (Hoboken, N. J.: John Wiley and Sons, Inc., 2013), xiii.

Optimization, MEP, and Mutualism

17

flower can release noxious chemicals to ward off herbivores; the honeybee can sting hive invaders. Each individual may also be genetically different from others of the species which may advance or retard its capabilities or reduce the number of options the individual can (or might) select from to make optimal decisions. For example, the less massive elephant seal male may not challenge the beach master for dominance, while a more massive male may. Therefore, optimization involves identifying and assessing important variables that can affect which decision is best. Even when categories of variables do not change, the value of each may impact which decision will be best. For example, the honeybee forager sees a promising new patch of flowers, but competition from other foragers is too great. Her best decision is likely to find a new patch or return to another one she remembers had less competition. Later in the day, competition has declined, so she decides to forage the promising patch. The flower seeking water often sends roots out in as many promising directions as possible. Once she finds water, she redirects water-seeking root growth towards the source rather than continue to send roots elsewhere. These are examples of making the best decision under the conditions and constraints posed by the environment in context of individual needs and goals. If, as this study assumes, that both flowers and honeybees can judge then they meet at least one requirement for Singer’s morality to emerge. Without the ability to judge, then making optimal decisions would be more-or-less a random act in a series of many more sub-optimal acts which would not likely serve the species or the mutualism or even the ecology over the long term. Studies of optimality may help explain some of the issues facing plants and animals and some of the processes that life has derived towards optimality. David Ackerly modeled tropical plant leaves in plants that produce leaves continually to determine what contributes to carbon gain decline which is thought to be a measure of whole-plant optimal performance.2 The question he studied was whether reduction in light (self-shading) or age of the leaf or both contribute to declining performance.3 While his model used only a limited number of samples, Ackerly suggests that self-shading contributed more to the drop in carbon gain decline than age.4 New leaf production costs energy, but maintaining leaves that are not productive wastes energy. Older but functional leaves might continue to be productive, but those (even younger) that are self- or canopy-shaded that are not productive should be shed optimally. 2  See: David Ackerly, “Self-Shading, Carbon Gain and Leaf Dynamics: A Test of Alternative Optimality Models,” Oecologia 119, no. 3 (1999): 300. 3  Ibid. 4  Ibid., 308.

18

Chapter 1

Paul Aigner considered the theory called the Most Effective Pollinator Principle (MEPP) that suggests that plants should evolve towards the most effective pollinators in their meadows. The models he developed propose that this is not always the case; that there may be other factors involved in plant-pollinator relationships other than effectiveness that produce evolutionary change. He concludes, “Plants may in fact often become specialized to their most effective pollinators, but there are as yet insufficient theoretical or empirical grounds to claim this as an organizing principle of floral ecology and evolution.”5 For example, humans have long kept honeybees in portable hives that they use to pollinate their crops. Honeybees also pollinate the other flowers they find valuable to forage. Studies are finding that the introduction of non-native or invasive species of bees and other pollinators may contribute to the decline in local pollinators, some who may be more effective pollinators for local plants.6 Locations where pollinators are not consistent year to year also present complications for MEPP theory. While there are exclusive flowerpollinator relationships, this is not the norm. Honeybees have been valuable for humans all over the world because they will forage many species of flowers. Others like bumblebees and butterflies may also forage many different species. In places where there are many different species of pollinators and where most available flowers are pollinated, it may not be optimal for many of these flowers to evolve towards any one pollinator. More fundamental to just pollination is whether outbreeding, the avoidance of self-pollination, has contributed to the long-term success of angiosperms. Antonio C. Allem reviewed many previous studies of angiosperm breeding practices. While most angiosperms are outbreeders, about a third are self-breeders, meaning they pollinate themselves without the assistance of pollinators.7 Allem provides an example where outbreeding became detrimental, “For example, strictly outbred maritime Danish races of Viola tricolor vanished in a few generations owing to inadequate pollinator service.”8 Today, 5  Paul A. Aigner, “Optimality Modeling and Fitness Trade-Offs: When Should Plants Become Pollinator Specialists?,” Oikos 95, no. 1 (2001): 183. 6  See discussions on the decline of different bee species: Mark J. F. Brown and Robert J. Paxton, “The Conservation of Bees: A Global Perspective,” Apidologie 40, no. 3 (2009); Carolina L. Morales et al., “Rapid Ecological Replacement of a Native Bumble Bee by Invasive Species,” Frontiers in Ecology and the Environment 11, no. 10 (2013); Jennifer C. Grixti et al., “Decline of Bumble Bees (Bombus) in the North American Midwest,” Biological Conservation 142, no. 1 (2009). 7  See: Antonio C. Allem, “Optimization Theory in Plant Evolution: An Overview of Long-Term Evolutionary Prospects in the Angiosperms,” Botanical Review 69, no. 3 (2003): 242. 8  Ibid.

Optimization, MEP, and Mutualism

19

insect species are vanishing at an alarming rate.9 Outbreeding may have been an optimal practice for early angiosperms when many pollinators evolved from earlier insect species to enter into mutualism relationships with angiosperms. As pollinator insect species disappear, it will be important to consider the impact on the efficacy of outbreeder species compared with self-breeders. What is optimal for angiosperms in the future will depend upon the conditions of the environment and the availability of pollinators. Canopy models in rainforests and elsewhere have long been studied in context of optimality. Niels P. R. Anten notes, “Canopy models have long been used to quantitatively relate the leaf area distribution and leaf photosynthetic characteristics of plants to their net photosynthetic carbon gain.”10 In a review of the literature on canopy modeling, Anten suggests that there are likely many factors that contribute to canopy optimization other than light. He notes that nitrogen availability is important and for many plants, nitrogen is absorbed through the roots. Some plants may be in nitrogen rich areas and others in nitrogen poor areas which will affect how the plant can grow. Plant height, leaf area, and leaf angle also affect optimality. He notes also that many studies of canopies are conducted at one specific moment in time rather than over a period of time where other things like infestation may impact canopy formation.11 For example, if one tree is cut down or falls down or loses limbs in a windstorm, there suddenly is more opportunity for surrounding trees to increase their branch and leaf capacity in the now vacant canopy. Anten also finds difficulty with canopy competition theories because in many settings there are many different kinds of competitors, “The problem with the competition theory is that, while there are maybe six to ten different resources that limit growth there can easily be more than 50 species of plants that coexist with each other.”12 Most competition models only use a small number of competitors. Therefore, increased complexity may engage oscillating competitive models that vary with the types of resources for which these species compete.13 Lars Chittka notes that, “[b]ee vision is indeed optimal for coding flower color … A preliminary result is that bee color vision is also optimal for 9  See: Scott Hoffman Black and Mace Vaughan, “Endangered Insects,” in Encyclopedia of Insects, ed. Vincent Resh and Ring T. Carde (Burlington, Ma.: Elsevier, 2009); Axel Hochkirch, “The Insect Crisis We Can’t Ignore,” Nature News 539, no. 141 (2016). 10  Niels P. R. Anten, “Optimal Photosynthetic Characteristics of Individual Plants in Vegetation Stands and Implications for Species Coexistence,” Annals of Botany 95, no. 3 (2005): 495. 11  Ibid., 503. 12  Ibid., 504. 13  Ibid.

20

Chapter 1

discrimination of leaf colors.”14 However, he does not suggest that the relationships that emerged between bees and angiosperms necessarily evolved the bee to see these colors. They may be the product of much earlier adaptations.15 Chittka does suggest that flowers may have adapted their color schemes to attract bees.16 However, he does not maintain that there is a one-to-one relationship to flower color and bee sight capabilities, and that these may be asymmetric due to floral limitations of producing the most optimal color schemes to attract bees.17 This suggests that species affordances or constraints may not always be easily adaptable and that optimization may include satisficing techniques that are quasi-optimal, but are still effective. These are but a few of the many optimization studies of natural processes that have been conducted in recent decades.18 Time and change represent other challenges to species’ optimization behaviors. The meadow changes from day to day. At certain times of the floral growing season there may be no flowers or a bounty. The honeybee hive uses various mean to accelerate the maturation and perhaps even retard the maturation 14  Lars Chittka, “Bee Color Vision Is Optimal for Coding Flower Color, but Flower Colors Are Not Optimal for Being Coded—Why?,” Israel Journal of Plant Sciences 45, no. 2–3 (1997): 126. 15  Ibid. 16  Ibid. There is even research that suggests that bees are attracted to the warmth of the color that they also can discern in their visual spectrum, see: Adrian G. Dyer et al., “Bees Associate Warmth with Floral Colour,” Nature 442, no. 3 (2006). 17  Chittka, “Bee Color Vision Is Optimal for Coding Flower Color, but Flower Colors Are Not Optimal for Being Coded—Why?,” 126. 18  Other relevant studies not summarized in this study include: Mathieu Cloutier et al., “A Systems Approach to Plant Bioprocess Optimization,” Plant Biotechnology Journal 7, no. 9 (2009); Roderick C. Dewar et al., “Optimal Function Explains Forest Responses to Global Change,” BioScience 59, no. 2 (2009); Cleiton B. Eller et al., “Modelling Tropical Forest Responses to Drought and El Niño with a Stomatal Optimization Model Based on Xylem Hydraulics,” Philosophical Transactions of the Royal Society B: Biological Sciences 373 (2018); Oskar Franklin, “Optimal Nitrogen Allocation Controls Tree Responses to Elevated Co2,” The New Phytologist 174, no. 4 (2007); Agren Goran I and Franklin Oskar, “Root: Shoot Ratios, Optimization and Nitrogen Productivity,” Annals of Botany 92, no. 6 (2003); Kouki Hikosaka, “Leaf Canopy as a Dynamic System: Ecophysiology and Optimality in Leaf Turnover,” ibid., 95, no. 3 (2005); Mathieu Lihoreau et al., “Bees Do Not Use Nearest-Neighbour Rules for Optimization of Multi-Location Routes,” Biology Letters 8, no. 1 (2012); William A. Mitchell and Thomas J. Valone, “The Optimization Research Program: Studying Adaptations by Their Function,” The Quarterly Review of Biology 65, no. 1 (1990); Miloš Nikolić and Dušan Teodorović, “Empirical Study of the Bee Colony Optimization (Bco) Algorithm,” Expert Systems with Applications 40, no. 11 (2013); Graham H. Pyke, “Optimal Foraging Theory: A Critical Review,” Annual review of ecology systemat­ ics 15, no. 1 (1984); Jan Scheirs, “Integrating Optimal Foraging and Optimal Oviposition Theory in Plant-Insect Research,” Oikos 96, no. 1 (2002).

Optimization, MEP, and Mutualism

21

of hive bees into foraging bees based upon the availability of flowers. Waggle dancers inform unemployed honeybee foragers where they can find floral patches. Communication saves energy and increases the number of successful foraging trips for the hive. The hive may increase or decreases its hive guarding diligence based upon how good the foraging has been. As hive guarding can be quite costly (stinging bees die; guards are not foraging) if the hive has built adequate supplies of honey, it may relax its defense against raiding from workers from other hives.19 Flowers need sunlight to produce energy, but they also need water. If there is a drought, the plant may retard growing new limbs or leaves to put energy into sending out more roots to find water. Once water is found, the plant can return to its strategy of growing branches and leaves to produce more energy. All of these are examples of how flowers and honeybees optimize their decisions and behaviors based on the circumstances of the environment over time in context of the affordances and constraints of the species. Species can only optimize to the extent they are capable. Consider that the flower cannot move to the other side of the meadow where a stream runs even in the deepest drought. However, the honeybee can fly to the stream to obtain water to cool the hive. Flower sessility is a constraint and honeybee mobility is an affordance when it comes to access to this stream during dry seasons. Capabilities and needs of species vary. Dormancy helps the perennial plant to survive the winter without consuming too much stored energy. Energy stored and conserved can be used at the beginning of the growing season to produce new stems, leaves, roots, and flowers. On the other hand, honeybee workers and their queen survive the winter huddled together in the hive eating down stores of honey for energy. Such a strategy gives honeybees an advantage because they can begin foraging in significant numbers once the floral season begins. Both flowers and honeybees have ontological/morphological and epistemological/behavioral means to select the best decision given the circumstances of the moment. As the studies in this chapter show, the world presents different variables in different combinations at different times, whether seasonal or otherwise. Both flowers and honeybees need to optimize their behaviors to maintain their optimal existence. Plants need an adequate number of leaves for photosynthesis. They have developed processes and light-sensing organs 19  “Our data therefore, strongly support the hypothesis that an adaptive shift in guard acceptance thresholds occurred,” Stephen G. Downs and Francis L. W. Ratnieks, “Adaptive Shifts in Honey Bee (Apis Mellifera L.) Guarding Behavior Support Predictions of the Acceptance Threshold Model” Behavioral Ecology 11, no. 3 (2000): 333.

22

Chapter 1

to move branches and leaves from shade to sun. They have developed other routines to maintain optimal leaf photosynthesis. For example, if an herbivore begins to chew on a plant’s leaf, many plants have the capability of producing noxious chemicals that may slow down predation or even force the herbivore to find a different plant to forage. We cannot say that optimization is the good. We can only say that optimization in the flower and honeybee facultative mutualism is towards the good, but not necessarily good for all parties. While it is true that flowers and honeybees show restraint towards each other, their considerable powers of reason and judgment and resulting behaviors do not always directly benefit the other. The honeybee avoids flowers beset by predators or competition. The flower ceases the production of nectar when it is pollinated. Avoiding predators is good for the honeybee. Ending the production of nectar is good for the plant because it preserves energy to mature fruits and seeds. However, both species show restraint towards the other as they go about their individual businesses. They do not cause undue harm to the other. There are no written rules or laws that flowers and honeybees follow. There also is no external arbiter or sovereign who establishes and enforces rules. We can say that their facultative mutualism has been evolutionarily driven. While this is true, the actions of millions of individuals from both species are necessary to draw both towards each other. It requires that both species optimally behave towards each other in ways that maintain and even strengthen their facultative mutualism. This is something both species have done on their own and together over time. The elegance of Singer’s three requirements for morality to emerge lies in simplicity. Morality can emerge from something as simple as the restraint shown by flowers and honeybees towards each other as each make optimal decisions for themselves and the mutualism. Their habituation towards each other has evolved optimal practices that have served each other well or their mutualism would not have survived. Both restraint and judgment have also served flowers and honeybees well. Their mutualism serves to preserve individuality, but not, ultimately, at the expense of the other. Judgment tempered by restraint maintains their facultative mutualism social group and the individual participants and species that are a part of the group. While they are co-dependent, they also can go about their individual lives without undue constraint. Contrast the flower and honeybee social group with human social groups. We are a one-species social group, we are co-dependent upon each other, but at the same time we lead independent lives where we make judgments in context of both the social group and individual needs. Honeybees and flowers exist in most parts of the world as do humans. One difference may be that flowers and

Optimization, MEP, and Mutualism

23

honeybees use the process of evolution to produce biotechnology (proboscis, nectaries) to enhance their co-dependence. Humans create external technologies (tools, cars) that require external rules and other systems for their use in a co-dependent society. These external rules require external bureaucracies to enforce them. Both human societies and the flower and honeybee facultative mutualism, even with their different technological and enforcement constructs, use judgment and restraint as tools for moral co-existence. Therefore, what underlies both human and flower and honeybee morality is not substantially different. Optimization is fundamental to the maintenance of the flower and honeybee facultative mutualism. If flowers and honeybees (and likely other species in their ecology) optimize their behaviors regularly and consistently, there is emerging evidence that this produces entropy in a way that is consistent with Newton’s second law of thermodynamics. 3

Maximum Entropy Production (MEP)

If we can assume that life is an emergent process, meaning that order is created out of what some might call chaos, then a major question that arises is whether life defies the laws of physics and chemistry. Within emergent theory is the problem of entropy. Newton’s second law of thermodynamics suggests that the universe and all its matter and energy tend towards decreasing order, not increasing order. Life increases order and thus poses a problem for emergent theories to define how life comports with the second law of thermodynamics. Nobel Laureate Ilya Prigogine and co-author Isabelle Stengers suggest how this increasing order might be possible: The famous law of increase of entropy describes the world as evolving from order to disorder; still, biological or social evolution shows us the complex emerging from the simple. How is this possible? How can structure arise from disorder? Great progress has been realized in this question. We now know that nonequilibrium, the flow of matter and energy, may be a source of order.20 If nonequilibrium is a source of order and such order emerged early life forms, then life’s capability of sustaining itself is not contrary to the laws of physics. 20  Ilya Prigogine and Isabelle Stengers, Order out of Chaos, Kindle ed. (London: Verso, 2017), xxix.

24

Chapter 1

What does this have to do with morality emerging in the flower and honeybee social group? The flower and honeybee facultative mutualism is a result of both biological and social evolution. The social group itself is an emergent process, meaning that it produces order. Life itself is in the business of creating order or complexity which, on the surface, seems diametrically opposed to the second law of thermodynamics. However, what science is learning is that life is a participant in the entropy process just as nuclear decay and black holes are. Life must consume or make energy to survive. In that respect, life contributes to entropy in the non-linear thermodynamic earth. Therefore, if we can say that life does not violate the second law of thermodynamics, then life’s emergence and the subsequent emergence of the flower and honeybee social group do not require us to enter a mystical or metaphysical explanation for how life is different from fundamental processes. Thus, the emergence of life and morality are natural processes that that are derived from the fundamental forces of nature. There does not have to be an intervening force whether metaphysical or otherwise. Morality therefore can be derived from the naturally occurring phenomenon called life. It is life’s unique contribution to entropy that this chapter explores. Axel Kleidon, et al., explain the second law of thermodynamics, “The second law states that for isolated systems that do not exchange energy or mass with their surroundings, the entropy of that system can only increase.”21 Kleidon, et al., define entropy in context of linear thermodynamic systems. This is the theoretical closed bottle in space that eventually distributes gas molecules evenly through the bottle. Earth, on the other hand, continually receives light from the sun, and life and other processes use and release that energy making the world a non-linear thermodynamic system but one that is a steady state. James Dyke and Axel Kleidon put forth a definition of Maximum Entropy Production to help explain the entropic process in complex systems: The proposed Maximum Entropy Production (MEP) principle states that sufficiently complex systems are characterized by a non-equilibrium thermodynamic state in which the rate of thermodynamic entropy production is maximized.22 21  Axel Kleidon, Yadvinder Malhi, and Peter M. Cox, “Maximum Entropy Production in Environmental and Ecological Systems,” Philosophical Transactions of the Royal Society B: Biological Sciences 365 (2010): 1297. 22   James Dyke and Axel Kleidon, “The Maximum Entropy Production Principle: Its Theoretical Foundations and Applications to the Earth System,” Entropy 12, no. 3 (2010): 613.

Optimization, MEP, and Mutualism

25

The question is whether the second law of thermodynamics holds in a nonlinear system where life is present. Life is a complex system, but life is also a self-organizing system. Filip J. R. Meysman and Stijn Bruers, note that Erwin Schrodinger wondered whether life defied the second law of thermodynamics that suggests that systems tend towards more entropy rather than less.23 Life seems to increase complexity and organization rather than dissipate it.24 While Schrodinger proposed some ideas to resolve this dilemma, Meysman and Bruers note that later researchers have come up with a stronger theory: Schrodinger’s idea was more generally reformulated as that biological systems can maintain a far-from-equilibrium state only through a continuous exchange of energy and matter with their environment, a process that is necessarily accompanied by entropy production.25 Though life maintains its organization, it must use energy to do so, and therefore contributes to the production of entropy in a manner that does not contradict the second law of thermodynamics.26 Meysman and Bruers suggest that the amount of entropy produced by life in the meadow likely exceeds the entropy that could be produced by the same meadow absent of any life.27 Life is a self-organizing system. Emergent life theories suggest that early life assembled from building blocks like amino acids and other chemicals that may have been surrounded by a lipid bag that eventually self-organized into what we might call single-cell animal life.28 Rod Swenson defines emergence as, “Spontaneous transformation of a set of components (generalized ‘atomisms’ 23  F ilip J. R. Meysman and Stijn Bruers, “Ecosystem Functioning and Maximum Entropy Production: A Quantitative Test of Hypotheses,” Philosophical Transactions: Biological Sciences 365, no. 1545 (2010): 1405. 24  Prigogine and Stengers offer this observation, “On the contrary, life seems to express in a specific way the very conditions in which our biosphere is embedded, incorporating the nonlinearities of chemical reactions and the far-from-equilibrium conditions imposed on the biosphere from solar radiation” Prigogine and Stengers, Order out of Chaos, Location 703. 25  Meysman and Bruers, “Ecosystem Functioning and Maximum Entropy Production: A Quantitative Test of Hypotheses,” 1405. 26  See: Axel Kleidon, “A Basic Introduction to the Thermodynamics of the Earth System Far from Equilibrium and Maximum Entropy Production,” ibid.: 1313. See also: E. D. Schneider and J. J. Kay, “Life as a Manifestation of the Second Law of Thermodynamics,” Mathemati­ cal and Computer Modelling 19, no. 6 (1994). 27  Meysman and Bruers, “Ecosystem Functioning and Maximum Entropy Production: A Quantitative Test of Hypotheses,” 1405. 28  See: Terrence W. Deacon, Incomplete Nature: How Mind Emerged from Matter (NY & London: W. W. Norton & Company, 2012), 288.

26

Chapter 1

or ‘particles’) from an incoherent state … to a coherent state.”29 Coherence is required for judgment to occur and judgment is required for optimization to occur. A direct line from emergent coherence of life through optimization can now be drawn. This is just more evidence that morality, as an emergent process in the coherent state that is life, is derived from more fundamental natural processes. This is helpful because it shows that self-organizing systems are not an anathema to nature or to the second law of thermodynamics. However, not all self-organizing processes in the universe produce life. Benard cells are an example of a non-life self-organizing process. In a Benard experiment, a viscous translucent fluid is heated from above and cooled from below. The once translucent fluid produces many opaque “Benard cells” that look like red blood cells or donuts with depressions in the middle but not holes.30 Once the heat and cool are removed, self-organization ceases. However, while life self-organizes and all individual life forms die, somehow the system called life maintains this process over much longer periods of time (billions of years) than a Benard experiment is likely to be conducted. The Benard cells do not mutate into other self-organizations; life does.31 The selforganization requires the expenditure of energy and the maintenance of that self-organization also requires expenditure of energy. If they don’t emerge into something more complex, amino acids, chemicals, and lipids will continue to contribute to entropy, but not at the same rate as if they self-organize into life. The difference between life and the Benard cell self-organization is the ability of life to sustain and replicate itself through the making of or capturing of environmental energy in such a way as to produce continuity. The Benard cell returns to the prior state once the narrow bands of externally supplied heat and cold are removed. Life is uniquely capable of searching for and acquiring energy to sustain itself. Second, life has developed a process to forestall the inevitable decay of organic materials and that is reproduction. Benard cells 29  Rod Swenson, “Emergent Attractors and the Law of Maximum Entropy Production: Foundations to a Theory of General Evolution,” Systems Research 6, no. 3 (1989): 188. 30  Ibid., 192. 31  Rod Swenson gives insight as to how life might be different from a Benard cell, “[t]hat the spontaneous ordering (or self-organization) that characterizes the visible world is a process of selection of which replicative ordering is a special case (the case where there is component replication)” Rod Swenson, “Selection Is Entailed by Self-Organization and Natural Selection Is a Special Case,” Biological Theory 5, no. 2 (2010): 177. Emphasis in original. Both the Benard Cell and life comport with MEP according to Swenson. Natural selection involves MEP but the difference is that there is component replication i.e. DNA reproduces itself. Even so, natural selection, mutation, and epigenetics, while, they may accord with MEP also serve to change the MEP produced even if in minute amounts when new changes take hold in the species.

Optimization, MEP, and Mutualism

27

cannot reproduce themselves; they can grow in number only when the conditions are right, and the amount of required chemicals are present. Life, on the other hand, can reproduce and this is where life is fundamentally different from other processes that comport with the laws of physics and chemistry. Reproduction, however, is not outside of either physics or chemistry law, nor is its continuity function a violation of the second law of thermodynamics. The continuity process, simply put, is consistent with self-organization which is an emergent process in nature. However, we are still uncertain how DNA and RNA emerged in nature. Unraveling that answer is the subject for other studies. Maximum Entropy Production is a “class of hypotheses” that try to explain just how life produces entropy and in what amounts.32 It is important to note that MEP is not a single theory. Rather, specific hypotheses emerge from investigations into different ecological and life functions. However, as Meysman and Bruers suggest, these hypotheses can be gathered under these two basic MEP notions: The MEP prediction is that the living system will not only have a higher entropy production than the non-living one, but also that the living system will self-organize itself so that its entropy production is maximized in some way.33 Life both increases entropy and organizes so that it will maximize entropy ­production. This study suggests that this maximization is achieved through the process of optimization through which life engages. Even with competition and predation, increases in entropy will require higher and/or more productive biomass. Therefore, optimization appears critical to MEP as well as to life itself. Meysman and Bruers say that MEP is a kind of, “[g]oal function, i.e. an extremal principle that is relevant for the development and operation of living systems.”34 The difficulty of ascribing a goal function to a living system is that it relies upon individual members of the living system to act in a manner that produces maximum entropy. Consider the notion of optimality in context with goals. If creatures were to always choose the greatest entropic alternative, likely the life system would devolve into carnage and death, ultimately leading to a less entropic abiotic system. Rather, life, somehow, seems 32  Meysman and Bruers, “Ecosystem Functioning and Maximum Entropy Production: A Quantitative Test of Hypotheses,” 1406. 33  Ibid. 34  Ibid.

28

Chapter 1

bent upon the maintenance of the life system in perpetuity. For want of a better term to explain this drive towards continuity, we can say it derives from the actions of ‘selfish genes.’35 Each optimal decision that each creature makes is ultimately towards this preservation of the life system. If the primordial goal of life is to sustain and reproduce its genetic code, then MEP, in part, will come from the collective optimal decisions of the creatures in the ecosystem that are towards reproduction. Therefore, an underlying goal that drives MEP in life systems is the propensity of life to orient towards continuity, derived from continuing genetic replication. Can we say that genetic mutation contributes to MEP? Likely it does, but in complex ways. Some mutations will not benefit the species, others will. The randomness of mutation seems contrary to the primordial goal of life towards continuity that is maintained by optimal decisions. Some mutations will advance creature capabilities, while others will not, or even cause existential problems. However, if we can maintain that a creature with advantageous genes and one with disadvantageous genes both make optimal choices, given what they were born with, then we can suggest that, yes, in the aggregate, the life system continues towards MEP even while the genetic code for some species may evolve to produce more or less entropy. The emerging science of epigenetics, which I will only mention briefly here, is discovering that knowledge of parents in some species can be handed down to offspring without education. For example, a study has shown that tobacco plants can pass down epigenetic defenses against pathogens to the next generation.36 Epigenetics does not change the order or composition of the chromosome, it does, however, change the expression of select genes. We can also offer that epigenetics is towards life’s goal of continuity. However, performance increases achieved through epigenetic processes in the tobacco plant likely reduce the entropy of the plant’s leaves being damaged by pathogens. On the other hand, they may also increase the entropy of the pathogen. Whether or not the new state that is reached is more or less entropic is not the critical issue. Based upon circumstances, e.g. epigenetic transfer of information on how to fight pathogens, change sets a new maximum entropic level in the ecology. Optimal behaviors, mutation, and epigenetics all serve to maintain life continuity in life systems. One of the reasons why researchers of MEP are hesitant to call it a single theory is that life is a complex organized system, and while 35  See: Richard Dawkins, The Selfish Gene (Oxford: Oxford University Press, 1989). 36  See: Alexander Boyko et al., “Transgenerational Changes in the Genome Stability and Methylation in Pathogen-Infected Plants: (Virus-Induced Plant Genome Instability),” Nucleic Acids Research 35, no. 5 (2007): 1714.

Optimization, MEP, and Mutualism

29

Meysman and Bruers suggest that life systems produce more entropy than abiotic systems, how they do so requires investigation into the different processes involved.37 Meysman and Bruers are not alone in suggesting that MEP should be thought of as a principal rather than a rule for the construction of different hypotheses of entropy production by life systems. Leonid Martyushev asks two important questions, “(i) can this principle claim to be the basis of all non-equilibrium physics? and (ii) is it possible to prove MEPP?”38 Based upon evidence from studies and theoretical work, Martyushev answers the first question with a qualified, “ ‘yes’ today.”39 While MEP studies to date have given credence to the underlying premise, there may be processes that do not conform to MEP. To the second he says: Note first that a principle like MEPP cannot be proved. Examples of its successful applications for description of observed phenomena just support this principle, while experimental results (if they appear) contradicting the principle will just point to the region of its actual applicability.40 Other theories from nature and physics have defied definitive proofs, so this is not unusual. Nor does the lack of proof negate the possibilities for the applicability of this principle. Tyler Volk and Olivier Pauluis have additional questions about the efficacy of MEP, including: [c]an MEP predict not just the overall state of entropy production of a system but also the details of the sub-systems of dissipaters within the system? Which fluxes of the system are those that are most likely to be maximized? How it is possible for MEP theory to be so domain-neutral that it can claim to apply equally to both purely physical-chemical systems and also systems governed by the ‘laws’ of biological evolution?41

37  Meysman and Bruers, “Ecosystem Functioning and Maximum Entropy Production: A Quantitative Test of Hypotheses,” 1415. 38   Leonid M. Martyushev, “The Maximum Entropy Production Principle: Two Basic Questions,” Philosophical Transactions of the Royal Society B: Biological Sciences 365, no. 2010 (2010): 1333. Note, MEPP is used but has the same meaning as MEP. 39  Ibid. 40  Ibid. 41  Tyler Volk and Olivier Pauluis, “It Is Not the Entropy You Produce, Rather, How You Produce It,” Philosophical Transactions: Biological Sciences 365, no. 1545 (2010): 1317.

30

Chapter 1

Volk and Pauluis suggest that in order to answer these questions we need to understand how entropy is produced, and ask, “Is it possible to determine the relative importance of specific processes for total entropy production?”42 Applied to biology, one of the problems they note is that sometimes in evolution entropy goes up and sometimes down even though, the overall entropy production rate is positive.43 Volk and Pauluis caution that changes in variables such as humidity or lack thereof in atmospheric convection models will produce different results.44 They note the considerable difficulty of applying MEP to biological systems: [M]EP theory has substantial problems when one attempts to apply it to biology. The problems come about because evolution will produce biological adaptations that might either increase or decrease the entropy production rate, so long as the entropy production rate is positive.45 I agree with this assertion, with the caveat that if life forms in general tend to make optimal decisions, as has been discussed, then even when biological adaptations are towards negative entropy, maximum entropy in these extant or even changing conditions continues to be produced. However, because life is a dynamic system, MEP becomes a dynamic process for which it will be difficult to predict the rate of MEP without more information. Volk and Pauluis put this problem into perspective, “Until MEP theory can make predictions about the details of the internal states of the dissipating systems, it will remain a heuristic.”46 Meaning, with life systems, MEP is more like a process of discover­ ing rather than a process of discovery. Joseph J. Vallino notes the difficulties of discovering MEP during major transient events: During major transient events, such as recovery from system collapse, MEP may not be a useful descriptor of system response. Rather, the community composition will likely play the dominant role in system dynamics as it reorganizes and attempts to return to MEP.47

42  Ibid. 43  Ibid., 1321. 44  Ibid. 45  Ibid. 46  Ibid. 47  Joseph J. Vallino, “Ecosystem Biogeochemistry Considered as a Distributed Metabolic Network Ordered by Maximum Entropy Production,” ibid.: 1424.

Optimization, MEP, and Mutualism

31

Vallino offers two such transient events, forest fires and invasive species: If a system organizes towards MEP over infinite time and space, then steepest descent routes must be inhibited, but they cannot be prevented from occurring. As already mentioned, a forest fire rapidly increases short-term entropy production, but at the expense of long-term entropy production. Invasive species can produce an identical phenomenon if the invading species propagates via oxidation of biological structure. However, if the invading species causes lower entropy production on longer time scales, then the new state will not be stable. On the contrary, if the invasive species increases averaged entropy production, then it should persist.48 Even after major transient events, I maintain that remaining creatures, or creatures that take advantage of a changing ecology, will continue to behave optimally. Likely MEP will increase over time as the place where the forest fire occurred recovers. Also, over time the place where the transient event occurs will reach a new equilibrium that may produce better predictability for MEP studies. Roderick C. Dewar considers plant optimization theory in context of MEP. He concludes: MEP can predict optimal plant behaviour that is reasonable from the perspective of natural selection. The different objective functions of these theories emerge as examples of entropy production on different spatiotemporal scales. Moreover, as a system-level thermodynamic principle, MEP extends the traditional optimization approach beyond individual plants to vegetation canopies and whole ecosystems. This suggests that MEP offers a unifying optimization principle for plant and ecosystem function, and that entropy production might be considered as a general objective function for biological systems.49 It is logical that time presents opportunities for living systems to regain equilibrium where MEP evolves to a different level. For example, even though an invasive species may not be contributing to the maximum entropy production that is possible for the meadow filled with a variety of life forms, both 48  Ibid. 49  Roderick C. Dewar, “Maximum Entropy Production and Plant Optimization Theories,” ibid.: 1434.

32

Chapter 1

the invasive species and the others who are evolving to cope with this new species and even compete with it, are all in the aggregate making optimal decisions towards their goals of continuity and reproduction. The forest fire is an example of where the land’s surface reverts to the originary near-abiotic state. However, even the burned-out land will not remain abiotic if there is life left on earth. We know from the many near life-extinction events, that life eventually returns to niches it once occupied even in completely different forms and combinations. Vallino offers, “However, because synthesis of biological structures cannot occur if entropy production is maximized instantaneously, we propose that information stored within the metagenome allows biological systems to maximize entropy production when averaged over time.”50 This suggests that the genome itself serves to regulate entropy through its capability for existing over a longer period of time. This Vallino suggest may be one reason why biotic systems and the passing of many generations of life forms produce more entropy than abiotic systems over time.51 In conclusion, I return to Martyushev, “Therefore, the possibility that all non-equilibrium thermodynamics and statistical physics can be constructed on the basis of the entropy production (actually the time derivative of the entropy) maximization appears to be very intriguing.”52 This study will proceed as Martyushev suggests, intrigued by the possibilities for the efficacy of MEP to fundamentally explain principles that underly biological processes such as the co-evolution and mutualism of flowers and honeybees but with an honest ‘reasonable doubt’ that it can yet be universally applied. Study examples are helpful in illustrating the possibilities for MEP analysis of various life processes. Chemotaxis is the process that bacteria use to move towards chemicals. In laboratory experiments on chemotaxis, Paško Županović et al., reported finding that the process is, “[r]ooted in the MEP principle.”53 Evidence of MEP can be found in laboratories using specific chemicals and specific bacteria, but what about the broader ecology? In a study conducted in the Amazon, Robert J. Holdaway, et al., report, “Our results indicated that forests had a higher rate of entropy production than pastures.”54 This is logical because the forest (especially a rainforest) has a 50  Vallino, “Ecosystem Biogeochemistry Considered as a Distributed Metabolic Network Ordered by Maximum Entropy Production,” 1417. 51  Ibid., 1417, 25. 52  Martyushev, “The Maximum Entropy Production Principle: Two Basic Questions,” 1334. 53  Paško Županović et al., “Bacterial Chemotaxis and Entropy Production,” Philosophical Transactions: Biological Sciences 365, no. 1545 (2010): 1401. 54  Robert J. Holdaway, Ashley D. Sparrow, and David A. Coomes, “Trends in Entropy Production During Ecosystem Development in the Amazon Basin,” ibid.: 1445.

Optimization, MEP, and Mutualism

33

greater biomass than the pasture. The rainforest may extend to tens of meters with life that exists from underground to the canopy. The pasture may extend only a meter high with some insect and bird activity above. Even though the niche called the pasture may have a complex life biome in its smaller footprint, it will logically produce less entropy than the more complex forest. It will, however, based upon the optimizing actions of the pasture species, produce MEP that is possible at this moment of the pasture’s existence. Stanislaus J. Schymanski, et al., used the Klausmeier model to study MEP, “It is based on the assumption that the existence of biomass permits increased infiltration and hence increased water use, while increased water use in return permits increased biomass growth. This leads to a positive feedback between biomass and transpiration.”55 The water-biomass feedback mechanism is just one force that contributes to biomass increasing entropy more than abiotic systems. Likely researchers will discover many more. While MEP will not directly help this study’s efforts to consider the emergence of morality in nature, MEP, along with the principle of optimality, shows that life and its activities over time do not violate fundamental principles of physics. This is important because this observation serves to critique notions that morality has been divinely given to humans exclusively. Morality is not constituted from outside of physics but is constituted within the physical laws. While soteriological revelation and the teaching thereof are important to human morality, I suggest they are not the source of morality in nature. Therefore, if optimality is consistent with MEP and MEP in general applies to life systems, and if optimality is primordial to the emergence of ethics, we can then say that morality is constructed from elements consistent with the physical laws of nature. If we can find antecedents of morality in nature, we can begin to look for evidence of morality in the flower and honeybee mutualism. First, however, we must understand what constitutes the flower and honeybee mutualism. 4 Mutualism Solitary species have emerged and thrived in the world. However, there are species who find it mutually beneficial to benefit from and, in some cases, exploit another species. Species have therefore discovered that it is optimal to engage with other species and even evolve their capabilities towards doing 55  S tanislaus J. Schymanski et al., “Maximum Entropy Production Allows a Simple Repre­ sentation of Heterogeneity in Semiarid Ecosystems,” ibid.: 1449.

34

Chapter 1

so. The mutualism of the flower and honeybee is co-evolutionary, meaning that both species needed to evolve in order to capitalize on their relationship. While this involves changing the underlying genetic structure of each e.g. the honeybee to drink nectar and the flower to make it, this doesn’t happen without the active participation of the individual species. They had to have used reason and judgment in order to more closely align their behaviors towards the other and likely this process also helped natural selection to favor mutations that benefited their mutualism. It is in the flower and honeybee mutualism that we begin to see the emergence of morality in nature. Their facultative mutualism is a social group, they do not intentionally harm each other, and they use judgment in their behaviors towards each other. As angiosperms have become the dominant plant species and so many insect species have evolved to both benefit from and exploit flowering plants, there must be very good reasons why this process is something that has lasted now an estimated one hundred million years. If life is an entropic process, then mutualism and the explosion of angiosperms and their pollinators, has served to increase MEP on earth. Mutualism is not the only successful process towards increasing entropy, but it is an important process that has found purchase in many different life forms. We cannot say that all mutualisms are towards the good. Each must be studied before making that determination. However, this study maintains that indeed the flower and honeybee facultative mutualism is towards the good because it comports with Singer’s requirements for the emergence of morality in nature. This chapter provides some basic information about the different forms of mutualism that will be helpful for others who wish to consider other mutualism constructs as candidates for contribution towards the good and other emergent instances of morality in nature. Judith L. Bronstein defines mutualism as, “Mutualism refers to all interspecific interactions, regardless of their specificity, intimacy, or evolutionary history.”56 As the definition implies, there are different manifestations of mutualism. For example, the obligate composite organism: the symbiote lichen has evolved different existential properties from its component life forms: fungus and algae. Bronstein also suggests that the warning call of the blue jay that many animals in the meadow recognize as such is an indirect mutualism that is cooperative rather than obligate.57 The flower and honeybee mutualism is between the extremes of obligate and cooperative. Bronstein gives the flower and honeybee mutualism these characteristics: “facultative”, “non56  Judith L. Bronstein, “The Study of Mutualism,” in Mutualism, ed. Judith L. Bronstein (Oxford, UK: Oxford University Press, 2015), 10. 57  Ibid., 11.

Optimization, MEP, and Mutualism

35

exclusive”, “co-evolved”, and “facilitative.”58 First, it is facultative, meaning that the mutualism is not associated with all life functions as is the lichen. The honeybee requires the flower’s production of nectar (carbohydrate) and pollen (protein) for nourishment; the flower requires the honeybee and other pollinators to spread pollen to other flowers of the same species so that seeds can be produced. All other life functions for either species are not directly associated with the mutualism. Neither the honeybee nor the flowers she forages are exclusive arrangements. While honeybees may not forage all species of flowers that emerge in the meadow, they will forage any flower that is suitable. The flowering plants that the honeybee forages are also pollinated by other species. Both honeybees and the flowers they forage are generalists. There are, however, specialist arrangements between flowers and pollinators where there is only one flower that the pollinator will forage, and the flower is shaped, colored, and scented to attract only one species of pollinator. Such an exclusive relationship might include an insect with a very long tongue to reach pollen because the bell of the flower is equally long. No other insect can reach the nectar of this flower and therefore the other insects will not forage this flower. Flowers and honeybees have co-evolved into their mutualism. The distant predecessor to the honeybee, the apoid wasp, was a carnivorous insect hunter. The fossil record shows some early angiosperms were insect pollinated, but there may have been others who were wind or water pollinated and therefore did not need insects or other pollinators.59 Current thinking is that the development of sticky pollen attracted early pollinators.60 Sticky pollen is just that, something that will attach itself to a carrier like an insect who can take it to other plants and pollinate the other plants. I speculate that the apoid wasp foraged from flower to flower looking for insects hidden inside or on the plant. In doing so it brushed the sticky pollen which stuck to various parts of its body. While flying from flower to flower looking for insects, it took pollen to other plants of the same species. The emergence of sticky pollen is not 58  Ibid. 59  Fossil record studies suggest that some early angiosperms may have been pollinated by insects. See: “We provide data to show that early fossil angiosperms were insect-pollinated” Shusheng Hu et al., “Early Steps of Angiosperm—Pollinator Coevolution,” Proceedings of the National Academy of Sciences 105, no. 1 (2008): 240. Also see: “We document three well-preserved examples of angiosperm herbivory, two assigned to modern insect genera, occurring early within the initial angiosperm radiation” C. C. Labandeira et al., “Ninety-Seven Million Years of Angiosperm-Insect Association: Paleobiological Insights into the Meaning of Coevolution,” Proceedings of the National Academy of Sciences of the United States of America 91, no. 25 (1994): 12280. 60  See: Hu et al., “Early Steps of Angiosperm—Pollinator Coevolution,” 240.

36

Chapter 1

co-evolutionary. The apoid wasp had not yet evolved. The apoid wasp may have realized during cleaning or other behaviors that the pollen satisfied its need for protein and may have learned to consume pollen or feed it to young as a dietary supplement. This still is not a co-evolutionary step. What happened to co-evolve the apoid wasp is uncertain. It could have been any number of things. It may have evolved, like the honeybee, pollen baskets on its hind legs, a better way of carrying pollen. Or, rather than hunt for insects, it evolved to be less aggressive, preferring pollen over the dangerous process of hunting. Plants photosynthesize light and carbon dioxide into sugars. Early angiosperms with sticky pollen may also have exuded some sugar product which the wasp or its successor found rewarding. The co-evolutionary race is now on. The apoid wasp evolved her mouth parts and gut to forage nectar and its hind legs to carry pollen and eventually became a honeybee and other members of extant bee or pollinating wasp species. The flower changed color, shape, and scent to attract the forager and evolved mechanisms to produce nectar in small quantities for the evolving wasp to forage. She also constructed herself so that the forager would have to pass by the sticky pollen on the way to the nectar treat. The evolutionary steps that led to present day flowers and honeybees have not been well preserved in the fossil record. Speculation is that both co-evolved asynchronously. E.g., the sticky pollen came first, then the wasp evolved to prefer sticky pollen over insects or perhaps it was a different co-evolutionary advance altogether.61 Today, the flower and honeybee mutualism is non-exclusive, but it is facultative, meaning that while both cannot do without the other for one major life function (food/reproduction), they both have robust lives outside of the mutualism. Consider that a plant species may bloom, for example, for two weeks out of a year. That is the only time that the flower and honeybee mutualism is active for that flower species. Finally, Bronstein notes that the flower and honeybee mutualism is facilitative, meaning that neither species intentionally harms the other.62 I use the word intentional because the actions of honeybees entering flowers may cause some wear and tear to the flower. Entering in and out of flowers likely causes some wear and tear to the honeybee. Consider also that neither competes with the other for common resources. Both have asymmetrical needs, and both have evolved to so require the other that to harm the other would not benefit the harmer. The honeybee receives only a small amount of nourishment from each flower. Flower blooms are often inconsistent during the growing season and the foraging honeybee needs to forage the entire growing 61  Ibid. 62  Bronstein, “The Study of Mutualism,” 11.

Optimization, MEP, and Mutualism

37

season to provide what the hive needs to survive year-round. The hive needs all the flowers she can find and forage. The honeybee forages many flower species and she will carry pollen from one species to different species. The flower that attracts honeybees must produce enough pollen to attract as many pollinators as possible because her pollinators are not specialists and mis-deliveries will be quite common. Therefore, both require the other and it would be folly to harm the other. Flowers grow only on plants. Honeybees are animals. They each come from completely different evolutionary branches of life. Yet, they both have co-evolved to require the other for existential needs. They have also evolved means for exchanging information. Flowers advertise color, shape, and scent to notify pollinators of their availability and suitability. When the flower is pollinated, she shuts down nectar production, some flowers change color to turn off advertising, and the flower withers and desiccates so that she no longer attracts pollinators. Flowers cannot see pollinators. However, when the honeybee forager enters the flower, she produces tactile sensations in the flower that can alert the flower to the presence of the pollinator.63 Different flower species have different responses to the tactile feel of the pollinator. One response is to loosen pollen so that it can stick to the pollinator. Others may release a drop of nectar. Still others do not respond to vibrations. Flower species have evolved to optimize their responses to the momentary event of the pollinator entering the flower. As the nectar reward is low for the pollinator, the pollinator has evolved to harvest as many flowers as it can in the time available to forage. For the honeybee forager this includes evolving a memory function to remember known flower locations. She has also evolved the capability of communication with other members of the hive to announce foraging locations (waggle dances), and the ability to translate foraging locations into flight patterns to help other workers fly more-or-less directly to the communicated site. Foraging honeybees can travel many kilometers in search of flower blooms.64 Honeybees needed to evolve not only to fly distances but also to be able to 63  E.g., “Bees that forage on poricidal flowers share a unique form of pollen harvesting. Upon alighting, female bees curl around the ‘anther cone,’ if present, or grasp clusters of stamens while rapidly shivering their large indirect flight muscles. This produces an audible buzzing sound and transmits strong vibrations to the floral androecium. In response to this vibration, the pollen is expelled (in 0.1–1.0 second) striking and sticking to the venter of the bee where it is later groomed and packed into the scopa” Stephen L. Buchmann, “Bees Use Vibration to Aid Pollen Collection from Non-Poricidal Flowers,” Journal of the Kansas Entomological Society 58, no. 3 (1985): 518. 64  Francis L. W. Ratnieks and Kyle Shackleton, “Does the Waggle Dance Help Honey Bees to Forage at Greater Distances Than Expected for Their Body Size?,” Frontiers in Ecology and Evolution 3, no. 31 (2015): 2.

38

Chapter 1

recognize major features of the landscape as markers to guide them back to distant flower patches.65 The flower and insect pollinator relationship has spanned nearly one hundred million years.66 For the last million years, flowers and honeybees have been a mutualism. Angiosperms comprise more than two hundred sixty thousand species and are today the predominant plant family.67 Some, as has been mentioned, are wind or water pollinated who do not require pollinators, therefore they have not co-evolved into a plant-pollinator mutualism relationship. Self-pollinators do not need pollinators either. Some self-pollinators may have first evolved into mutualisms with pollinators but may have evolved out of the mutualism for any number of reasons, including the lack of reliable pollinators. The future of plant pollinator relationships is not certain. While plant-pollinator mutualisms are predominant in angiosperms, the changing dynamic of the world, the growing rate of extinction of insect species, and the reduction of populations like honeybees due to hive collapse syndrome, may eventually require both flowers and pollinators to evolve to accommodate new existential and environmental exigencies. The flower and honeybee facultative mutualism, as this study suggests, meets Singer’s requirements for the emergence of morality: the social group, restraint towards other members of the group, and judgment. Co-evolution requires the participation of actors in the process to capitalize on and adjust to the morphological advances of the other. Awareness of the other is the first step in this process. After awareness, recognition that the other is good is required for the process to continue. This goodness must withstand the exigencies of both the environment and the vagaries of mutational evolution. The focus on the other must be covetous, but not like the Cheetah’s focus on 65  For more about the limits of honeybee vision see: Chittka, “Bee Color Vision Is Optimal for Coding Flower Color, but Flower Colors Are Not Optimal for Being Coded—Why?.”; Peter G. Kevan, Lars Chittka, and Adrian G. Dyer, “Limits to the Salience of Ultraviolet: Lessons from Colour Vision in Bees and Birds,” Journal of Experimental Biology 204, no. April (2001); Miriam Lehrer, “Spatial Vision in the Honeybee: The Use of Different Cues in Different Tasks,” Vision Research 34, no. 18 (1994); Adrian G. Dyer, Christa Neumeyer, and Lars Chittka, “Honeybee (Apis Mellifera) Vision Can Discriminate between and Recognise Images of Human Faces,” Journal of Experimental Biology 208, no. 24 (2005). 66  Pamela S. Soltis and Douglas E. Soltis, “The Origin and Diversification of Angiosperms,” Amercan Journal of Botany 91, no. 10 (2004): 1616. 67  According to: Labandeira et al., “Ninety-Seven Million Years of Angiosperm-Insect Association: Paleobiological Insights into the Meaning of Coevolution.” However, Douglas Soltis, et al., suggest the number is between three-hundred fifty and four hundred thousand: Douglas Soltis et al., Phylogeny and Evolution of the Angiosperms: Revised and Updated Edition (Chicago, Ill.: University of Chicago Press, 2018), 1.

Optimization, MEP, and Mutualism

39

the Gazelle. There must be a reciprocal coveting that acknowledges the good of the other or the mutualism will not progress. The goodness that the facultative mutualism develops, unlike the Cheetah and Gazelle, requires both the flower and the honeybee to benefit from and exploit the other. The Cheetah is dependent upon the Gazelle and both have co-evolved to run faster in their long association, but their social construct is antagonistic and, the Cheetah does not maintain restraint with her prey. Whether the autochthonous step towards the flower-pollinator revolution was sticky pollen or the wasp recognizing pollen to be nourishment, we may never know. What we do know that rather than become the predator-prey model like the Cheetah and Gazelle, the apoid wasp and the flower began to form a relationship based upon coequal benefit and exploitation of asynchronous resources. They began to form a social group, more casual at first and perhaps less cooperative than today, took care not to harm each other, and had to learn how to reason and judge for their own benefit and for the other. They eschewed competition for cooperation and mutually exploitational/beneficial commerce. The goodness comes not only from the morphological changes they have co-created to move more towards harmonizing with the other, but also through judgment and reason that both actors use to maintain this co-beneficial commerce relationship over many millions of years. Theirs is a society of equals. It is not a democracy where the majority rules. It is a social group where each strives over many millions of years to find ways of becoming a better servant to and beneficiary of what the other can provide. I cannot think of a better word for this social group than moral, a co-equal morality where there are no inter-species castes, no sovereigns, the ruled, or the subaltern. However, we are still at the beginning of the journey towards discovering morality in the flower and honeybee facultative mutualism. In the next chapters on ontology and morphology, epistemology and behavior, there is further discussion of restraint and judgment by both species. The nature of the flower and honeybee social group is considered in context of behaviors by both species. Cited References Ackerly, David. “Self-Shading, Carbon Gain and Leaf Dynamics: A Test of Alterna­ tive Optimality Models.” Oecologia 119, no. 3 (May 1999): 300–10. doi:10.1007/ s004420050790. Aigner, Paul A. “Optimality Modeling and Fitness Trade-Offs: When Should Plants Become Pollinator Specialists?”. Oikos 95, no. 1 (2001): 177–84.

40

Chapter 1

Allem, Antonio C. “Optimization Theory in Plant Evolution: An Overview of LongTerm Evolutionary Prospects in the Angiosperms.” Botanical Review 69, no. 3 (2003): 225–51. Anten, Niels P. R. “Optimal Photosynthetic Characteristics of Individual Plants in Vegetation Stands and Implications for Species Coexistence.” Annals of Botany 95, no. 3 (Feb 2005): 495–506. doi:10.1093/aob/mci048. Black, Scott Hoffman, and Mace Vaughan. “Endangered Insects.” In Encyclopedia of Insects, edited by Vincent Resh and Ring T. Carde, 320–24. Burlington, Ma.: Elsevier, 2009. Boyko, Alexander, Palak Kathiria, Franz J. Zemp, Youli Yao, Igor Pogribny, and Igor Kovalchuk. “Transgenerational Changes in the Genome Stability and Methylation in Pathogen-Infected Plants: (Virus-Induced Plant Genome Instability).” Nucleic Acids Research 35, no. 5 (2007): 1714–25. Bronstein, Judith L. “The Study of Mutualism.” In Mutualism, edited by Judith L. Bronstein, 3–19. Oxford, UK: Oxford University Press, 2015. Brown, Mark J. F., and Robert J. Paxton. “The Conservation of Bees: A Global Perspec­ tive.” Apidologie 40, no. 3 (2009): 410–16. Buchmann, Stephen L. “Bees Use Vibration to Aid Pollen Collection from Non-Poricidal Flowers.” Journal of the Kansas Entomological Society 58, no. 3 (1985): 517–25. Chittka, Lars. “Bee Color Vision Is Optimal for Coding Flower Color, but Flower Colors Are Not Optimal for Being Coded—Why?”. Israel Journal of Plant Sciences 45, no. 2–3 (1997): 115–27. Chong, Edwin K. P., and Stanislaw H. Zak. An Introduction to Optimization. Hoboken, N. J.: John Wiley and Sons, Inc., 2013. Cloutier, Mathieu, Chen Jingkui, Caroline De Dobbeleer, Michel Perrier, and Mario Jolicoeur. “A Systems Approach to Plant Bioprocess Optimization.” Plant Biotech­ nology Journal 7, no. 9 (Dec 2009): 939–51. doi:10.1111/j.1467-7652.2009.00455.x. Dawkins, Richard. The Selfish Gene. Oxford: Oxford University Press, 1989. Deacon, Terrence W. Incomplete Nature: How Mind Emerged from Matter. NY & London: W. W. Norton & Company, 2012. Dewar, Roderick C. “Maximum Entropy Production and Plant Optimization Theories.” Philosophical Transactions: Biological Sciences 365, no. 1545 (2010): 1429–35. doi: 10.1098/rstb.2009.0293. Dewar, Roderick C., Oskar Franklin, xe, kel, xe, Annikki, Ross E. McMurtrie, and Harry T. Valentine. “Optimal Function Explains Forest Responses to Global Change.” BioScience 59, no. 2 (2009): 127–39. doi:10.1525/bio.2009.59.2.6. Downs, Stephen G., and Francis L. W. Ratnieks. “Adaptive Shifts in Honey Bee (Apis Mellifera L.) Guarding Behavior Support Predictions of the Acceptance Threshold Model.” Behavioral Ecology 11, no. 3 (2000): 326–33.

Optimization, MEP, and Mutualism

41

Dyer, Adrian G., Christa Neumeyer, and Lars Chittka. “Honeybee (Apis Mellifera) Vision Can Discriminate between and Recognise Images of Human Faces.” Journal of Experimental Biology 208, no. 24 (2005): 4709–14. Dyer, Adrian G., Heather M. Whitney, Sarah E. J. Arnold, Beverley J. Glover, and Lars Chittka. “Bees Associate Warmth with Floral Colour.” Nature 442, no. 3 (08/02/online 2006): 525. doi:10.1038/442525a https://www.nature.com/articles/ 442525a#supplementary-information. Dyke, James, and Axel Kleidon. “The Maximum Entropy Production Principle: Its Theoretical Foundations and Applications to the Earth System.” Entropy 12, no. 3 (2010): 613–30. doi:10.3390/e12030613. Eller, Cleiton B., Lucy Rowland, Rafael S. Oliveira, Paulo R. L. Bittencourt, Fernanda V. Barros, Antonio C. L. da Costa, Patrick Meir, et al. “Modelling Tropical Forest Responses to Drought and El Niño with a Stomatal Optimization Model Based on Xylem Hydraulics.” Philosophical Transactions of the Royal Society B: Biological Sciences 373 (2018): 201–14. doi:10.1098/rstb.2017.0315. Franklin, Oskar. “Optimal Nitrogen Allocation Controls Tree Responses to Elevated Co2.” The New Phytologist 174, no. 4 (2007): 811–22. Goran I, Agren, and Franklin Oskar. “Root: Shoot Ratios, Optimization and Nitrogen Productivity.” Annals of Botany 92, no. 6 (Mar 2003): 795–800. doi:10.1177/ 007327538902700101. Grixti, Jennifer C., Lisa T. Wong, Sydney A. Cameron, and Colin Favret. “Decline of Bumble Bees (Bombus) in the North American Midwest.” Biological Conservation 142, no. 1 (2009): 75–84. Hikosaka, Kouki. “Leaf Canopy as a Dynamic System: Ecophysiology and Optimality in Leaf Turnover.” Annals of Botany 95, no. 3 (Feb 2005): 521–33. doi:10.1093/aob/ mci050. Hochkirch, Axel. “The Insect Crisis We Can’t Ignore.” Nature News 539, no. 141 (2016): Online. Holdaway, Robert J., Ashley D. Sparrow, and David A. Coomes. “Trends in Entropy Production During Ecosystem Development in the Amazon Basin.” Philosophical Transactions: Biological Sciences 365, no. 1545 (2010): 1437–47. Hu, Shusheng, David L. Dilcher, David M. Jarzen, and David Winship Taylor. “Early Steps of Angiosperm—Pollinator Coevolution.” Proceedings of the National Acad­ e­my of Sciences 105, no. 1 (2008): 240–45. Kevan, Peter G., Lars Chittka, and Adrian G. Dyer. “Limits to the Salience of Ultraviolet: Lessons from Colour Vision in Bees and Birds.” Journal of Experimental Biology 204, no. April (Jul 2001): 2571–80. Kleidon, Axel. “A Basic Introduction to the Thermodynamics of the Earth System Far from Equilibrium and Maximum Entropy Production.” Philosophical Transactions: Biological Sciences 365, no. 1545 (May 12 2010): 1303–15. doi:10.1098/rstb.2009.0310.

42

Chapter 1

Kleidon, Axel, Yadvinder Malhi, and Peter M. Cox. “Maximum Entropy Production in Environmental and Ecological Systems.” Philosophical Transactions of the Royal Society B: Biological Sciences 365 (2010): 1297–302. Labandeira, C. C., D. L. Dilcher, D. R. Davis, and D. L. Wagner. “Ninety-Seven Million Years of Angiosperm-Insect Association: Paleobiological Insights into the Meaning of Coevolution.” Proceedings of the National Academy of Sciences of the United States of America 91, no. 25 (1994): 12278–82. Lehrer, Miriam. “Spatial Vision in the Honeybee: The Use of Different Cues in Different Tasks.” Vision Research 34, no. 18 (1994): 2363–85. Lihoreau, Mathieu, Lars Chittka, Steven C. Le Comber, and Nigel E. Raine. “Bees Do Not Use Nearest-Neighbour Rules for Optimization of Multi-Location Routes.” Biology Letters 8, no. 1 (2012): 13–16. doi:10.1098/rsbl.2011.0661. Martyushev, Leonid M. “The Maximum Entropy Production Principle: Two Basic Questions.” Philosophical Transactions of the Royal Society B: Biological Sciences 365, no. 2010 (May 12 2010): 1333–34. doi:10.1098/rstb.2009.0295. Meysman, Filip J. R., and Stijn Bruers. “Ecosystem Functioning and Maximum Entropy Production: A Quantitative Test of Hypotheses.” Philosophical Transactions: Bio­ logical Sciences 365, no. 1545 (2010): 1405–16. Mitchell, William A., and Thomas J. Valone. “The Optimization Research Program: Studying Adaptations by Their Function.” The Quarterly Review of Biology 65, no. 1 (1990): 43–52. Morales, Carolina L., Marina P. Arbetman, Sydney A. Cameron, and Marcelo A. Aizen. “Rapid Ecological Replacement of a Native Bumble Bee by Invasive Species.” Frontiers in Ecology and the Environment 11, no. 10 (2013): 529–34. Nikolić, Miloš, and Dušan Teodorović. “Empirical Study of the Bee Colony Optimi­ zation (Bco) Algorithm.” Expert Systems with Applications 40, no. 11 (2013/09/01/ 2013): 4609–20. doi:https://doi.org/10.1016/j.eswa.2013.01.063. Prigogine, Ilya, and Isabelle Stengers. Order out of Chaos. Kindle ed. London: Verso, 2017. Pyke, Graham H. “Optimal Foraging Theory: A Critical Review.” Annual review of ecol­ ogy systematics 15, no. 1 (1984): 523–75. Ratnieks, Francis L. W., and Kyle Shackleton. “Does the Waggle Dance Help Honey Bees to Forage at Greater Distances Than Expected for Their Body Size?” [In English]. Frontiers in Ecology and Evolution 3, no. 31 (2015-April-01 2015): 1–7. doi:10.3389/ fevo.2015.00031. Scheirs, Jan. “Integrating Optimal Foraging and Optimal Oviposition Theory in Plant-Insect Research.” Oikos 96, no. 1 (2002): 187–91. Schneider, E. D., and J. J. Kay. “Life as a Manifestation of the Second Law of Thermo­ dynamics.” Mathematical and Computer Modelling 19, no. 6 (1994): 25–48. doi:https:// doi.org/10.1016/0895-7177(94)90188-0.

Optimization, MEP, and Mutualism

43

Schymanski, Stanislaus J., Axel Kleidon, Marc Stieglitz, and Jatin Narula. “Maximum Entropy Production Allows a Simple Representation of Heterogeneity in Semiarid Ecosystems.” Philosophical Transactions: Biological Sciences 365, no. 1545 (2010): 1449–55. Soltis, Douglas, Pamela Soltis, Peter Endress, Mark W. Chase, Steven Manchester, Walter Judd, Lucas Majure, and Evgeny Mavrodiev. Phylogeny and Evolution of the Angiosperms: Revised and Updated Edition. Chicago, Ill.: University of Chicago Press, 2018. Soltis, Pamela S., and Douglas E. Soltis. “The Origin and Diversification of Angio­ sperms.” Amercan Journal of Botany 91, no. 10 (Oct 2004): 1614–426. doi:10.3732/ ajb.91.10.1614. Swenson, Rod. “Emergent Attractors and the Law of Maximum Entropy Production: Foundations to a Theory of General Evolution.” Systems Research 6, no. 3 (1989/09/01 1989): 187–97. doi:10.1002/sres.3850060302. Swenson, Rod. “Selection Is Entailed by Self-Organization and Natural Selection Is a Special Case.” Biological Theory 5, no. 2 (2010): 167–81. Vallino, Joseph J. “Ecosystem Biogeochemistry Considered as a Distributed Metabolic Network Ordered by Maximum Entropy Production.” Philosophical Transactions: Biological Sciences 365, no. 1545 (2010): 1417–27. Volk, Tyler, and Olivier Pauluis. “It Is Not the Entropy You Produce, Rather, How You Produce It.” Philosophical Transactions: Biological Sciences 365, no. 1545 (2010): 1317–22. Županović, Paško, Milan Brumen, Marko Jagodič, and Davor Juretić. “Bacterial Chemotaxis and Entropy Production.” Philosophical Transactions: Biological Sciences 365, no. 1545 (2010): 1397–403.

Chapter 2

Emergence of the Flower and Honeybee Mutualism and Flower and Honeybee Ontology and Morphology 1 Introduction Flowers and honeybees could not be more different. Angiosperms (flowering plants) come from the plant kingdom and honeybees (Apis Mellifera) the animal kingdom. Where flowers take root is where they stay, while honeybee foragers can fly kilometers from their hive in search of food. Flowering plants make food from sunlight using photosynthesis and gather water and minerals predominantly through their roots. Honeybees gather carbohydrates (nectar) and protein (pollen) by foraging flowers of the flowering plants. Honeybees also search for water to cool the hive and tree gum for hive-building. However, each species has co-evolved to require and accommodate the other. The honeybee worker mouth parts have evolved to suck up nectar and store it in her crop where it is offloaded in the hive through the process of trophallaxis. She has hairs on the back of her hind legs where pollen sticks to form bright yellow pantaloons. The flowering plant has evolved her flower to attract honeybees through color, smell, and shape. The bell of the flower both accommodates and directs the honeybee forager down into the depths of the flower where she passes by the pollen stamens that release sticky pollen onto the honeybee on her way down to the nectar she can forage, sometimes deep inside. We can say, therefore, that the flower has adapted her shape to fit the honeybee body so that she can release sticky pollen onto the honeybee who will carry it to other flowers. The honeybee has adapted her body to fit the flower and her mouthparts to capitalize on the nectar treat that the flower provides. She has modified her hind legs to transport pollen to other flowers and to her hive. The honeybee serves as the vehicle to transport pollen of insect pollinated flowers to other flowers of the same species where some of the sticky pollen is released to pollenate the other flower. Plants become dormant in winter, and while honeybees remain in a ball deep within the hive, they do not hibernate like some mammalian species but remain active enough to vibrate their body to warm each other. The flowering plant stores in both her stems and the roots, nutrients she creates from sunlight, and minerals and water she forages through her roots. In the first part of this chapter is a history of the emergence of flowers and honeybees. © Koninklijke Brill NV, Leiden, 2020 | doi:10.1163/9789004428546_004

Emergence of the Flower and Honeybee Mutualism

2

45

Evolution of the Flower Honeybee Mutualism

2.1 Emergence of Flowers and Their Pollinators First a caveat. The fossil record has many holes and remains woefully inadequate in the number of examples of many species. However, new sources of fossil material, new species, and more examples of known species are being discovered every day. Therefore, what follows is subject to considerable future revision as we learn more about the past. The purpose of this chapter is not to establish firm boundaries for the emergence of angiosperms and pollinators, but to consider the process of emergence and how each species began an affinity towards each other that eventually led to the rather robust and widespread relationship between angiosperms and their pollinators that exists today. After the emergence of the angiosperm pollinator relationship began to take hold, both angiosperms and pollinators have continued to evolve many new species that retain the fundamental flower-pollinator relationship. It appears that evolutionary efforts of mutation to produce alternative processes have not yet evolved something better. Perhaps nature herself resists change to a process that has been so successful for the partnership between plant and animal species—the angiosperms and their pollinators. That said, there are many variations of angiosperm pollination that do not include insects or other pollinators, including self-pollination, and wind and water pollination. Some angiosperms are pollinated by a single species, others attract many different species of pollinators. We find pollinating bees, wasps, beetles, mammals, and birds among species who are attracted to nectar and serve flower species by spreading pollen from plant to plant. Note that honeybees receive all their carbohydrate and protein requirements from foraging opportunistically any suitable angiosperm species. Next, we explore the historical record for evidence of the emergence of angiosperms. 2.2 The Dawn of Angiosperms Xin Wang offers many possibilities in the fossil record for the predecessor to angiosperms, “Among living plants the Gnetales (Ephedra, Gnetum, and Welwitschia) are a group considered currently by many to be the most closely related to the angiosperms. Gnetum lives today in tropical forests, while Ephedra and Welwitschia are dry-climate or desert plants.”1 Angiosperms are vascular plants which Wang distinguishes from nonvascular plants, “Vascular plants are distinguished from non-vascular plants by their special and efficient water conducting system, the vascular bundle.”2 1  Xin Wang, The Dawn Angiosperms (Heidelberg, Germany: Springer, 2010), 5. 2  Ibid., 19.

46

Chapter 2

Vascular bundles are important because the flower requires a continuous water column from root to stem and leaf to supply the parts of the flower with sugar from photosynthesis, hormones it produces, and water and minerals from the roots. The properties of water are such that these tiny water columns can avoid cavitation or breaking for the many meters tall that some flowering trees grow. Plants other than angiosperms also have vascular bundles.3 Beyond vascular bundles, there has been an evolution in how angiosperms are defined. Wang explains, “Angiosperms were originally defined by having seeds that are enclosed since it is exactly what the word angiosperm means. A closed carpel provides angiosperms an added protection against predation and harsh environments including desiccation, as well as a self-incompatible system, and adds a pre-zygotic selection in addition to the post-zygotic one, which is common in other seed plants.”4 Closed carpel is not unique to angiosperms, “[s]ome of the conifers have demonstrated the same tendency to enclose and protect their seeds after pollination.”5 The closest Wang can come up with to distinguish angiosperms from other plants is, “[a] physically enclosed ovule at pollination appears to be an optimal and sufficient criterion.”6 Wang suggests three stages for angiosperm development. The first stage was in the Jurassic (-201.3 to -145 Mya—million years ago) or even earlier where, “Pioneering angiosperms during this stage are experimenting with various possibilities.”7 In the middle stage or the Cretaceous (-145 to -66 Mya), “This is the developing and radiating period for angiosperms.”8 It is in this second period that angiosperms began to dominate. Note, however, that the end of the Cretaceous is also the boundary line between the age of the dinosaurs and the beginning of the age of mammals. Angiosperms survived the dinosaur extinction and have continued to evolve and flourish, perhaps because many other-than-angiosperm competitors went extinct. In the late stage or Cenozoic (-66 Mya to today), angiosperms dominate, and, “Ecologically, angiosperms develop more coherent and mutually beneficial relationships with animals, especially insects, birds and mammals. The co-evolution between angiosperms and animals results in many specialized features of both.”9 Douglas Soltis, et al., provide important evolutionary and phylogenic facts about the angiosperms. Angiosperms are a seed plant. Other extant seed plant 3  Ibid. 4  Ibid. 5  Ibid. 6  Ibid., 26. Emphasis in original. 7  Ibid., 190. 8  Ibid. 9  Ibid.

Emergence of the Flower and Honeybee Mutualism

47

phyla include, “cycads, conifers, gnetophytes, and Ginkgo.”10 The oldest fossil pollen grains are from the Early Cretaceous one hundred thirty one million years ago which is why this date is generally accepted as the known beginning of angiosperms, even though plants had earlier experimented with angiospermy in the Jurassic.11 Some think that early angiosperms developed in woody environments; others think the earliest were aquatic.12 However, as Wang suggests, the “dominating doctrine” is that they have one common ancestor.13 Wang also notes, “There is no consensus on the origin place for angiosperms.”14 Wang also suggests that the contribution of pollinators to Angiosperm success and speciation is not yet understood, “How much animals contributed to the success of angiosperms is an open question, as there were few changes in insects and reptiles corresponding to the changes in angiosperms during the Cretaceous.”15 Wang notes that the features of flowering plants that differentiate them from other plants likely is not the only reason for their success, “It appears that angiospermy alone cannot account for the success of the angiosperms. It may have been the combination of many features as well as biotic and abiotic factors that have contributed to the success of angiosperms since the Middle Cretaceous.”16 Wang suggests some, but not all possible angiospermy features that may have contributed to angiosperm success: [p]olyploidy, gene duplication, vessels, low-carbon-cost-for-shoot physiology, reticulate leaf venation, more efficient light usage, rhizomatous and lianoid habits, extensive vegetative propagation, high photosynthetic rates, plant-insect relationships, plant-dinosaur interaction, unique ability to response to high CO2 levels, climate change, higher vein density, fast growth rate, weedy habit, plant-bacteria association, short reproductive cycle, high speciation rate and low extinction rate, chemical defense mechanism, occurrence of endosperm, landscape connectivity, environmental influence, and horizontal gene transfer.17 Delving into these possible advantages is beyond the scope of this study. 10  Douglas Soltis et al., Phylogeny and Evolution of the Angiosperms: Revised and Updated Edition (Chicago, Ill.: University of Chicago Press, 2018), 1. 11  Ibid., 25. 12  Ibid., 33. 13  Wang, The Dawn Angiosperms, 191. 14  (Wang, 2010, p. 191). 15  Wang, The Dawn Angiosperms, 192. 16  Ibid., 193. 17  Ibid.

48

Chapter 2

Flower origin requires a different discussion. Else Marie Friis, et al., note that there are two different theories for the origins of the angiosperm flower: the Euanthial and the Pseudanthial, “Under the Euanthial Theory the angiosperm flower is interpreted as a simple, bisexual, uniaxial system bearing spirally arranged lateral leaf-like appendages … Under the Pseudanthial Theory the angiosperm flower is interpreted as a compound, pluritaxal (multiaxial) structure potentially homologous to the cone of conifers or Gnetales.”18 There is as yet no comprehensive theory as to the origin of angiosperm flowers. Most angiosperms are bisexual, meaning they contain both male and female reproductive organs, but only some flowering plants self-pollinate.19 For those who do not self-pollinate there is generally a delay between “pollen production and stigma receptivity.”20 This time delay prevents self-pollination. Most angiosperms that do not self-pollinate (most angiosperms) rely on pollinators like insects, followed by wind and then water.21 Therefore, it is important to understand the basics of the mechanisms that flowers use to attract pollinators like honeybees. 2.3 Pollen Likely Attracted Insects First Flowering plants that attract insects like honeybees have sticky pollen. This means that the pollen remains on the stamens until it can be transferred to the insect that scrapes by the stamen as it seeks the nectar farther down in the flower. Some flowering plants respond to the vibratory presence of insects by loosening pollen so that it can more easily attach to the pollinator. At least one plant in the Andes, where both the climate and presence of pollinators is inconsistent from year to year, has learned how to extend stamens filled with pollen in a manner timed to when pollinators have previously visited in the recent past.22 In still other plants, pollinators like some bees (but not honeybees) buzz their wings which alerts the plant to loosen pollen.23 Insects like honeybees have hairs on their back legs that accumulate this pollen, some of which 18  Else Marie Friis, Peter R. Crane, and Kaj Raunsgaard Pedersen, Early Flowers and Angio­ sperm Evolution (Cambridge, UK: Cambridge University Press, 2011), 5. 19  Ibid., 11. 20  Ibid., 417. 21  Ibid. 22  “The findings at species level suggest that individual plants may be able to adjust the timing of their pollen presentation to the actual pollination scenario they experience” Moritz Mittelbach et al., “Flowers Anticipate Revisits of Pollinators by Learning from Previously Experienced Visitation Intervals,” Plant Signaling & Behavior e1595320, no. e1595320 (2019): 1. 23  Stephen L. Buchmann, “Bees Use Vibration to Aid Pollen Collection from Non-Poricidal Flowers,” Journal of the Kansas Entomological Society 58, no. 3 (1985): 517.

Emergence of the Flower and Honeybee Mutualism

49

is left behind at other flowers, some which is brought back to the hive where it becomes a rich protein source for the hive. The sticky pollen resists natural forces like wind and rain. Animals that brush by may dislodge some pollen but if the flower petals protect the pollen, the pollen may remain attached until a pollinator arrives. Douglas Soltis, et al., explain that pollinator attraction varies and that flowers use different substances to attract specific or generic pollinators, “Various means of pollinator attraction, in addition to pollen, have evolved in flowers, such as nectar, oil, resin, and perfume, which are needed for food, nest building, or attraction of mates. Whereas pollen and nectar are used by many pollinators, oil, resin, and perfume are used by some highly specialized bees.”24 Some flowers offer only pollen as a reward for pollinators.25 However, the sheer variety of flowering plants and the species that pollinate them suggest that there has been a long process of co-evolution that attracts diverse pollinators and has served to increase speciation of angiosperms. As flowering plants evolved, they ‘encouraged’ carnivorous apoid wasps to evolve mouth parts and other features to ingest and digest nectar and carry sticky pollen from one plant to another. Soltis, et al., explain: Much of floral diversity has been shaped by interactions of angiosperms with their pollinators. Floral organization and construction reflect broad pollinator syndromes e.g. so-called ‘hummingbird flowers’ vs. ‘bee flowers’, but additional features may also result from selection by pollinators and be crucial elements of floral diversity. Most prominent among these attributes—and the best studied—is color variation, particularly anthocyanins and their underlying genetic and biosynthetic pathways.26 Floral shape and color can attract or repel honeybees. Plants use flowers as colorful and shapely billboards to advertise to pollinators who have specific requirements. Pollinators will frequent flowers that advertise to their needs. Insects like honeybees will also investigate flowers they have not seen before to determine whether they can provide the required resources, meaning both quality of resource and accessibility (they can reach the nectar with their proboscis.) Once pollinated, some flowers change color, most stop producing nectar, and the flower itself desiccates by losing petals. Energy is then channeled from flower maintenance to seed and fruit production. There are countless ways that angiosperms and their pollinators interact that science has 24  Soltis et al., Phylogeny and Evolution of the Angiosperms: Revised and Updated Edition, 364. 25  Ibid. 26  Ibid., 376.

50

Chapter 2

discovered but are beyond the scope of this study that concentrates on flowers that are attractive to honeybees. ‘Flower’, as in flowering plant, is difficult to define.27 However there are different characteristics of angiosperms that are common. Xin Wang explains, “As reticulate leaf variation is a rarity in gymnosperms or ferns, but a common character in angiosperms, thus, unsurprisingly, it is a frequent identifier for angiosperms. This character, related to the efficiency of material transport within plant bodies, may have contributed to the success of angiosperms in their struggles to compete against their rivals.”28 As previously noted, the first angiosperms from the pollen fossil record can be dated to one-hundred-thirty million years ago in the early Creta­ceous period.29 There is no fossil evidence of true angiosperms before the Creta­ ceous period. Early flowering plants were generally small and most produced only one seed.30 Soltis, et al., suggest it was pollen that attracted insects to them.31 Pollen is rich in vitamins, lipids, proteins amino acids, starch, and sometimes oils.32 Between 100 and 93 million years ago the fossil record records a significant increase in angiosperm diversification and insects that complete the full metamorphic cycle (egg, larva, pupae, adult) at about the same time. Hymenoptera, the family which bees (and ants) come from, complete the full metamorphic cycle.33 What is not known for certain is whether the first angiosperms were woody or aquatic plants.34 Today angiosperms are woody trees, non-woody flowers like tulips, water plants like the water hyacinth, and so-called air plants (epiphytes), like orchids who capture humidity from their roots that do not burrow into the ground or the tree they attach to. Orchid roots, however, are highly absorbent to capture water in the form of mist, rain, or dew.

27  Wang, The Dawn Angiosperms, 17. 28  Ibid., 18. 29  Soltis et al., Phylogeny and Evolution of the Angiosperms: Revised and Updated Edition, 25. 30  Ibid., 27. 31  Ibid. 32  W. Scott Armbruster, “Evolution and Ecological Implications of ‘Specialized’ Pollinator Rewards,” in Evolution of Plant-Pollinator Relationships, ed. Sébastien Patiny (Cambridge, UK: Cambridge University Press, 2011), 46–47; Denis Michez, Maryse Vaderplanck, and Michael S. Engel, “Fossil Bees and Their Plant Associates,” in Evolution of PlantPollinator Relationships, ed. Sébastien Patiny (Cambridge, UK: Cambridge University Press, 2011), 134. 33  Soltis et al., Phylogeny and Evolution of the Angiosperms: Revised and Updated Edition, 29. 34  Ibid., 33.

Emergence of the Flower and Honeybee Mutualism

51

Today pollinators abound. There are more than 120,000 species of extant Hymenoptera (bees/wasps/ants) alone.35 Friis, et al., suggest that bees have coevolved the most with angiosperms: Bees are probably the group of pollinators that have co-evolved most closely with angiosperms and this relationship may account for much of the diversity of angiosperm flowers. Flowers that are pollinated by bees (melittophilous flowers) are often blue, yellow, or ultraviolet with nectar guides, and are open in the early morning. Frequently, flowers are zygomorphic, which forces the visiting bee into a position where it makes contact with stigma or pollen. Attraction is mainly visual and the reward is generally nectar and/or pollen.36 The diversity of angiosperms includes size, from 1 mm to over 100-meter eucalyptus trees, to single minute stamen flowers to giant blossoms more than 90 cm in diameter.37 Angiosperms are the only flowering plant. However there have been many ‘key innovations’ in angiosperms that have led to their considerable genetic radiation and diversity.38 These include the flowers and the evolution of organs into new and different functions e.g. the nectaries that produce nectar.39 For those flowers pollinated by animals (insects, birds, etc.) there are a number of variations. Variations can include “lipped flowers”, “tubular flowers of various lengths and widths”, and “specialization for a variety of pollinators”.40 There are also “revolve flowers”—that is, flowers that have several canals to reach the nectar, on which pollinators have to rotate around the center of the flower to gather all available nectar.41 Soltis et al., suggest, “This increased body contact with the anthers and stigma thus may enhance pollination success.”42 There are many more floral variations that are beyond the scope of this study. Our honeybee is a latecomer to the angiosperm relationship, evolving about a million years ago. While some bees are oligolectic, meaning that they forage in a narrow range of flower species, the honeybee is polylectic, meaning she will forage a wide range of flowering species. The honeybee is an opportunistic 35  Friis, Crane, and Pedersen, Early Flowers and Angiosperm Evolution, 421. 36  Ibid., 422. 37  Ibid., 1. 38  Soltis et al., Phylogeny and Evolution of the Angiosperms: Revised and Updated Edition, 343. 39  Ibid., 345. 40  Ibid., 363. 41  Ibid. 42  Ibid., 363–64.

52

Chapter 2

forager. The earliest bees are estimated to have evolved from Apoid (predatory) wasps about a hundred million years ago, that began to co-evolve with angiosperms thirty million years or so after angiosperm (pollen) known advent.43 As has been previously speculated, perhaps the Apoid wasps ate and learned to feed the pollen to their young as they would prey, providing a quite similar stew of nutrients for their young and themselves. The Insect consuming pollen rather than spreading pollen is an expensive problem for the plant. If the Apoid wasp or other pollen-eaters eat too much, the flower will be unable to replicate herself. This is especially devastating to annual plants that may flower only once. Flowers, therefore, needed to develop another method of attracting pollinators—nectar. Both the Apoid wasp and the angiosperm needed to evolve for the problem of excess pollen consumption to be resolved. These early Apoid wasps likely chewed the pollen. Their mandibles had not yet involved sucking systems that the later bees developed to acquire nectar.44 Michez, et al., suggest that likely wasp species began to evolve at this time and some became bees who no longer were predators but became full-time pollinators.45 The honeybee did evolve mouth parts to be able to siphon nectar, but also developed hairs on her hind legs to capture the sticky pollen that plants evolved that better attach to pollinators. The honeybee worker accumulates pollen on her hind legs but returns to the hive where hive workers offload this pollen which they use for food for the young and other purposes. As the honeybee flies from flower to flower, some of the pollen sticks to the new flower. Because the honeybee is polylectic, some pollen left behind at new flowers are not of the same species and therefore will not pollinate. Even with the advantages of insect pollination over wind or water pollination, abundant pollen must be produced because much is lost to mis-delivery and to honeybee consumption. The compromise worked out evolutionarily between the honeybee and the angiosperm includes the honeybee predecessor Apoid wasp’s emergent need for nutrients beyond protein and the need of the angiosperm to have a better chance to distribute pollen to other plants of its species. The key to making this process work was the evolution of nectar in flowers and nectar collecting systems in their pollinators. It appears from the literature and the fossil record that the explosion of angiosperm speciation corresponded with an explosion in insect pollinators. This gives credibility to the notion that the angiosperm had developed a very successful process for enticing and rewarding insects to assist in their reproductive process. As angiosperms began to rapidly fill new niches or those taken 43  Michez, Vaderplanck, and Engel, “Fossil Bees and Their Plant Associates,” 134. 44  Ibid. 45  Ibid.

Emergence of the Flower and Honeybee Mutualism

53

over from other plant species, they expanded not only in number of species but in number of individuals. As more species of angiosperms evolved there was a need for more insects to act as pollinators and some of these became specialists while others remained polylectic. Some researchers suggest that insect pollination evolved first and wind pollination later.46 As previously mentioned, others believe that wind or water pollination came first. If insect pollination came first, wind pollination could have been a mutation that took hold, or that also could have become necessary because of environmental pressure. One possibility is that where wind pollination evolved there were not enough pollinator insects and therefore wind pollination became an effective alternative to what had otherwise become a most effective process of insect pollination in other areas. It also could be possible that some angiosperms that did not develop sticky pollen that would cling to pollinators needed another process to distribute pollen. What is important to this study is not whether insect pollination came first or later, but that it continues to exist today, and robustly so. According to Michez, et al., the dinosaur extinction event sixty-five million years ago, that many believe was caused by or influenced by an asteroid or comet, had limited effects on bee speciation. However, a later global cooling event caused significant extinction in bee species, but this global cooling helped to evolve the modern bee.47 In addition to pollen that the Apoid wasps used to supplement predation, the sucrose, fructose, dextrose mixture in nectar offered additional energy benefits to later bee pollinators to supplement both the diet of their larvae and to supply energy for foragers. Honeybees have taken this energy production to the next step by producing honey that is not only rich in sugar but other nutrients (some from pollen) and serves the hive not only in young-rearing but also in wintering-over. Honey is also resistant to bacteria and other pathogens that might harm the young. The fossil record to date does not permit us to fill in all the blanks about the co-evolutionary process that led to both the explosion of angiosperm speciation and corresponding increase in pollinator insects. However, even with the aforementioned deviations, the mutualism between pollinator and angiosperm has resisted wholesale evolution towards a different approach, quite likely, because it has been such a successful process for both. The explosion of angiosperm species beginning in the later Cretaceous and into today suggests that even when flowers mutate into other species and other environmental 46   David Winship and Shusheng Hu, “Coevolution of Early Angiosperms and Their Pollinators: Evidence from Pollen,” Palaeontographica Abteilung B 4, no. 6 (2010): 103. 47  Michez, Vaderplanck, and Engel, “Fossil Bees and Their Plant Associates,” 140 & 43.

54

Chapter 2

niches, evolution resists changing the fundamental pollination process angiosperms and their pollination partners have developed. 3 Emergence We gain from the fossil record that both flowers and the apoid wasp evolved and radiated into many new species. This has become one of the most successful partnerships in nature. We can offer the theory that this is all about evolution and mutation. Mutation and time become an ontological engine that drives morphological changes that emerge mutualisms like the flower and honeybee. Co-evolutionary processes are likely even more powerful drivers of change because they require both species to concentrate time, energy, and intention on the other. However, there is more than just evolution involved in this process. Cooperation requires the rise, first of a recognition of the other, and then a reflective reciprocal gaze between the species in the mutualism. Attention to and intentionality towards the other becomes a focus of this reciprocal gaze. Nor is gaze the best word, because flowers cannot see the honeybee, though many can feel her presence. Gaze, therefore, is phenomenological and intentional, but not always visual. Because the relationship between flowers and honeybees is occasional and casual (non-competitive and non-violent) this reciprocal gaze must also be anticipatory and an expectation that the other will arrive at the flower or appear to honeybee. Both the flower and honeybee are embodied, meaning they cannot escape themselves. Yet they are never without each other even when they are not proximally located. Both are embedded in the world but embeddedness for each includes the other who is more than just the object of the other’s perceptivity. In other words, the flower and honeybee existence is more than the singularity of each being, or ipseity. Jean Luc Nancy provides insight, “Being cannot be anything but being-with-one-another, circulating in the with and as the with of this singularly plural coexistence.”48 Nancy is speaking about humans. This ‘being-singular-plural’ means that we become with others with whom we interact. While ontologically separate we are dependent upon each other not only for experience, but for our very becoming. However, we are a singular species. The flower and honeybee relationship is more than just being-singular-plural. Their reflective reciprocal gaze exceeds experience. They produce a kind of synthesis of existential horizons of reciprocal intentionality that engender the

48   Jean-Luc Nancy, Being Singular Plural (Stanford: Stanford University Press, 2000), 3. Emphasis in original.

Emergence of the Flower and Honeybee Mutualism

55

flower and honeybee singular-plurality.49 This synthesis of existential horizons requires anticipatory awareness in both species. The flower uses her considerable sensory skills (light, heat, vibration) to prepare for flowering, a critical existential requirement that leads to reproduction. Her anticipatory awareness at the time of flowering enables processes that have been dormant the rest of the year. She must be capable of responding as required when her pollinators arrive. She extends her horizon to the honeybee who extends hers to the flower. The synthesis of their horizons begins with the a priori nature of their respective gazes, but their horizons extend towards each other requiring priori response to the presence of this important other. Their horizons synthesize when the flower and honeybee come together in their intimate consummation of their mutualism. While both the flower and honeybee maintain their independent existences, their being-singular-plural is more than just experiential, it is existential. They are two separate existents whose synthesis of existential horizons is a requirement of continued existence. Therefore, there must exist between the two species a moral construct that is existentially driven and that also involves behavioral restraint and judgment that is towards the mutualism as well as towards horizonal others with whom they co-exist (e.g. hive mates; kin flowers). This moral construction likely began very early in the relationship of flowers and the apoid wasp, and while this moral construct encouraged ontological changes caused by genetic mutation, it also required intentional action of the actors in the developing mutualism. This emergent moral system could not have come solely from mutation and evolution. It requires the active participant and maintenance by extant actors who recognize the value of the other and what is required to preserve that value. In other words, while there is a considerable a priori underpinning to this moral construct, it requires priori action by the agents to sustain this relationship so that both can benefit from and exploit the mutualism. With the flower, it is interrelating processes and awareness that enable this sustaining. With the honeybee it is focused intentionality and perhaps even emotion, as will be discussed, that enable her to act to secure this moral relationship—the social group that is flowers and honeybees. We all know in general what a plant looks like, but it is important to understand basic morphology because morphology will inform the discussion of plant behavior and epistemology. 49  Adapted from Maurice Merleau-Ponty: “Therein it exceeds perceptual experience and the synthesis of horizons—as the notion of a universe, that is to say, a completed and explicit totality, in which the relationships are those of reciprocal determination, exceeds that of a world, or an open and indefinite multiplicity of relationships which are of reciprocal implication” Maurice Merleau-Ponty, Phenomenology of Perception, trans. Colin Smith (London: Roudedge and Kegan Paul, 1962), 71.

56 4

Chapter 2

Angiosperm Morphology

Even with variation, including size, color, and duration (e.g. annual versus perennial), the basic structure of the flowering plant generally contains these morphological elements.

Emergence of the Flower and Honeybee Mutualism

57

From the bottom and generally under soil or ground are the roots. The roots grow and branch through the soil in search of water and mineral nutrients (nitrogen, phosphorous, sulphur, and some metals) that the plant requires. Generally, the root system in most flowering plants develops a mutualistic relationship with mycorrhizae fungus that are found in the soil. The fungus obtains carbohydrates from the plant through its roots and the plant benefits because the fungi is more efficient in absorbing water and processing minerals into useable substances that both the fungus and the plant need. The fungi also are capable of harvesting and processing phosphates and iron from the soil, which the plant cannot. The fungi then can transfer these ions to the plant through the root system. The mutualism between plants and mycorrhizae began before the first flowering plants evolved because older plant families have mycorrhizae mutualisms. Also some mycorrhizae are compatible with more than one species of plant.50 Recent studies have found carbon transfer between different species of plants through their common mycorrhizae species mutualism as Suzzane W. Simard, et al., explain, “The amount of carbon exchanged between B. papyrifera and P. menziesii is indicative of a tightly linked plant—fungus—soil system.”51 The mycorrhizae attach to the root cells. Some species penetrate the root, others do not. More about the process of the exchange of nutrients, water, and minerals is beyond the scope of this project but can be found in most botany textbooks. For annual plants, while the roots may store nutrients until they are needed, long-term storage is unimportant as the plant only lives one growing season. Other flowering plants like tulips build large bulbous structures underground to store nutrients to maintain existence during the non-growing season and to provide nutrients to produce the flowers that tulips bloom in the early spring. Other plants like trees also have extensive root systems down which the sap flows in the fall when photosynthesis ends and rises to provide energy for buds to produce leaves and flowers for the next growing season. Maple sap is harvested in the spring when it begins to flow. The roots are attached to the stem. The tree trunk is simply a large stem. At intervals along the stem are nodes from which branches and/or leaves grow. Some tall flowering trees have few branches and then only at the canopy of the forest. Others like shrubs may have complex branches that have many twigs from many nodes. The stem acts like a highway or like vertebrate vascular systems that send nutrients, volatile chemicals, and electrical signals 50  Suzanne W. Simard et al., “Net Transfer of Carbon between Ectomycorrhizal Tree Species in the Field,” Nature 388, no. August (1997): 579. 51  Ibid., 581.

58

Chapter 2

from the roots to the ends of stems, flowers, and leaves and in some cases back again. The water column flows upward in summer when the leaves transpire or release some water to act as a coolant much as humans do when we sweat. Plants like cacti use other schemes to prevent excessive transpiration. The leaves and perhaps some branches and stems contain photosynthetic cells that take in carbon dioxide and sunlight and convert these into sugars and the waste product, oxygen. It is beyond the scope of this study to explain the photosynthetic process. However, the sugars produced by photosynthesis provide nourishment for the flowering plant, the mycorrhizal fungi on her roots, and her pollinating honeybees. The first life on earth was anaerobic because there was little or no oxygen in the air. The early photosynthetic organisms, the stromatolites and their descendants, produced so much oxygen that it reached toxic levels for the anaerobic life forms, touching off the first great life extinction event in earth’s history.52 We are indebted to plants for the oxygen we breathe. Flowers generally emerge from nodes on stems called axillary inflorescence, which may be quite short or long as in orchids which are often longer than the plant itself. Some plants like Tulips may produce only a single flower and other angiosperms may produce hundreds. Both roots and stems can grow for the entire life cycle of the flowering plant. Sometimes, as in trees, growth produces girth not only to support the tree vertically but to strengthen against wind or other natural phenomenon. Branches may grow or die or be damaged by wind, lightning, herbivores, pestilence, or other natural phenomenon. Roots may grow and die because they find no useful nutrients. Roots can also be cut by animals or humans who dig nearby. Many flowering plants live in temperate zones which means that they may only produce leaves and flowers during the growing season. Others in more tropical regions may produce flowers year-round and not shed leaves in the fall. However, even those plants that are more-or-less evergreen, will replace leaves and branches now and again as leaves begin to lose function or the light changes to where they no longer can produce photosynthesis. Some flowering plants may not flower until specific conditions are met. Desert plants often bloom only after a drenching rain which may not occur for many years. Other plants may have developed other schemes to flower under other circumstances. With so many species, the opportunity for specialization has arisen many times. 52  E.g., “The evolution of oxygenic photosynthesis in cyanobacteria is widely considered to have been the cause of this event” D. T. Flannery and M. R. Walter, “Archean Tufted Microbial Mats and the Great Oxidation Event: New Insights into an Ancient Problem,” Australian Journal of Earth Sciences 59, no. 1 (2012): 2.

Emergence of the Flower and Honeybee Mutualism

59

Buds form at the end of stems which then develop into new growth stems and leaves either later in the season or the next year. The buds in perennial plants in temperate areas winter over and begin the process of growth when spring conditions are conducive for growth and development. Both changes in temperature and the increase in the length of daylight can trigger flower and leaf production. Many non-flowering species like conifers have the same basic morphological characteristics as angiosperms. However, flowers are unique to angiosperms and require a separate morphological discussion. 5

Flower Morphology

There are too many types of flowers to explore each in this study. How­ ever, there are some basic characteristics that apply to most flowers. Adrian Bell explains the basic parts of the flower: The various components of a flower are attached in sequence along a usually very short and variously shaped central axis, the torus or floral receptacle at the end of the flower stalk (pedicel). Due to the shortness of the receptacle the most distal, female organs, collectively called the gynoecium, appear to be in the centre of the flower and to be surrounded by the more proximal male organs (androecium-stamens), which are in turn surrounded by perianth segments (such as petals and sepals).53

53  Adrian D. Bell, Plant Form: An Illustrated Guide to Flowering Plant Morphology (Portland, Oregon: Timber Press, 2008), 178.

60

Chapter 2

While there are many variations on this theme (specifically with how the ovary is positioned) this is the general shape of the flower. Pollen develops on the stamens and pollen is the male equivalent of sperm in mammals. Some flowers self-pollinate, others have developed mechanisms to prevent selfpollination. The gynoecium contains the ovary and from the ovary emerge the ovule which is the female equivalent of the mammalian egg that is produced by the ovary. When fertilized, the ovule becomes a seed. When pollen lands on a suitable ovule, it produces a pollen tube that penetrates the ovule and fertilizes the ovule. Angiosperms that are the subject of this study rely upon insect or other pollinators to carry the pollen from one flower to the ovules of other flowers of the same species. Insect species like honeybees are non-specific (polylectic) foragers, meaning that they will visit flowers that are visually attractive (color and shape) and with a bell into which they can both fit and reach the nectar which flower nectaries (nectar glands) produce. Honeybees are also attracted to the scent of flowers.54 Honeybees are curious creatures and will investigate new flowers that may have never bloomed in their territory before. A honeybee may return again and again to specific flower patches if she can obtain what the hive has informed her are nutrients that the hive requires at this time. She may not return to a patch of flowers that no longer produces nectar (or never has) or for other reasons such as the influx of competitors or predators. This has been a basic discussion of angiosperm ontology and morphology with some discussion of how plants and pollinators co-evolved over the one hundred thirty-million-year evolution of known angiosperms. Honeybees likely evolved from carnivorous apoid wasps and have only been a species for about a million years or so. As honeybees are animals, they require their own morphological discussion.

54  Studies have shown that honeybees, for example, can also sense the tactile feel of the flower to determine whether it is suitable to be foraged or avoided, “Previous work has described the remarkable discovery that honeybees can learn to identify flowers by touch when neither visual nor olfactory cues are available” Heather M. Whitney et al., “Conical Epidermal Cells Allow Bees to Grip Flowers and Increase Foraging Efficiency,” Current Biology 19, no. 11 (2009): 948.

Emergence of the Flower and Honeybee Mutualism

6

61

Honeybee Eusociality and Morphology

6.1 Honeybee Eusociality Bees in general (and ants) are believed to have evolved from wasps beginning one hundred million years ago.55 The honeybee emerged in Eurasia about one million years ago. Honeybees have become eusocial, meaning they share the rearing of offspring, have a caste system (workers, queen, drones or males), and overlapping maturation generations.56 The concept of overlapping generations does not perfectly correlate with human generations. For example, there are no living grandparents. Workers are all sisters or half-sisters. Rather than generations, there are different levels of worker maturation in the hive. Immature workers tend the hive and mature workers generally become foragers. Therefore, overlapping generations is not useful without the adjective maturation. Additionally, hive mates cooperate with each other and there is little strife in the hive. This does not mean that there are not challenges. New queens can emerge together and, generally, there is only one queen in the hive. Males die after they mate, but those that have not mated by the end of the season are pushed out of the hive where they will not survive for long. Hives also must remove sick and dead bees.57 Finally, honeybees can distinguish, likely by smell, raiding workers from other honeybee hives. Guard workers at the mouth of the hive are tasked with warding off these raiding bees. Some lose their lives

55  J ames L. Gould and Carol Grant Gould, The Honey Bee, ed. Carol Grant Gould (New York: Scientific American Library: Distributed by W. H. Freeman, 1988), 20. 56  This is the definition, with this study’s insertion of maturation, that Gadagkar suggests was popularized by C. D. Michener, Raghavendra Gadagkar, “Why the Definition of Eusociality Is Not Helpful to Understand Its Evolution and What Should We Do About It,” Oikos 70, no. 3 (1994): 485. This definition serves this study’s purposes for the honeybee, however Gadagkar believes overlapping generations should not be part of the definition, ibid., 486. Martin A. Nowak, et al., propose a different definition of eusociality to include these features, “(1) the formation of groups. (2) The occurrence of a minimum and necessary combination of pre-adaptive traits, causing the groups to be tightly formed. In animals at least, the combination includes a valuable and defensible nest. (3) The appearance of mutations that prescribe the persistence of the group, most likely by the silencing of dispersal behaviour” M. A. Nowak, C. E. Tarnita, and E. O. Wilson, “The Evolution of Eusociality,” Nature 466, no. 7310 (2010): 1062. 57  See: P. Kirk Visscher, “The Honey Bee Way of Death: Necrophoric Behaviour in Apis Mellifera Colonies,” Animal Behaviour 31, no. 4 (1983); M. Spivak et al., “Hygienic Behavior in the Honey Bee (Apis Mellifera L.) and the Modulatory Role of Octopamine,” Journal of Neurobiology 55, no. 3 (2003).

62

Chapter 2

in this effort either through injury or because they deploy their stingers which means certain death.58 Honeybees have evolved hive-making capabilities which provide shelter in which the existent members of the hive at the end of the flowering season can winter over in temperate climates by vibrating wing muscles to generate heat. While the natural range of honeybees extends through Eurasia, east Asia, and Africa, humans have exported the honeybee to most places in the globe where honeybees can exist. Eusociality meets Singer’s requirements for morality to emerge. The hive is a social group, restraint is shown other members of the group, and there is considerable judgment shown by the three castes in the hive as they go about their business. It is not inconsistent with other constructs of morality to suggest that the honeybee can be both a member of the eusocial group or the hive as well as the flower and honeybee social group. Humans navigate through different social groups (work, family, bureaucracy) every day. Unlike human caste systems that are prejudicial in nature, the honeybee caste system is both biological and based upon segregation of work responsibilities. No caste is superior to the other. All castes perform necessary functions for the hive and the hive could not exist without all castes. The queen and males are born into their roles as producers of new honeybees. They have no other job during their lifetime. Workers are sterile females. As they mature, they perform different services for the hive, from nursery tending, cleaning, foraging, scouting, and even guarding. Some workers may mature into foragers, others may not. As will be discussed later, personality may have a role in determining the job progression individual workers go through. Like the flower and honeybee facultative mutualism, there is no external arbiter or sovereign in the hive. However, in order to achieve optimal results, honeybees have evolved communication strategies that encourage or discourage specific behaviors of workers (which will be discussed in more detail in later chapters). At different times the hive may require pollen, and at other times nectar. Interestingly, much of the behavior altering strategy is negative. Workers bump dance other workers to stop them from dancing foraging locations that do not suit the hive. Delays in trophallaxis to returning foraging workers may inform the foraging bee that what she is bringing back now does not meet the hive’s current needs. These honeybee communication strategies contribute to optimal decision-making in the hive. Eusociality contributes 58  Africanized honeybees are known for their aggressiveness. When the hive is threatened, say by bear, large numbers of workers exit the hive and follow and sting the bear for much longer than the typical honeybee.

Emergence of the Flower and Honeybee Mutualism

63

to the flower and honeybee facultative mutualism by maintaining optimality in the hive. While the need for water to cool the hive is important right now and not of benefit to the flower, the long-term implications of cooling the hive means that new worker larvae will be kept cool enough to survive the heat wave and eventually become foragers. Meanwhile, the plant may have shut down the production of nectar in order to provide more energy to sustain root growth in search of water. The survivability actions of both will not only preserve the individual species, but in the long run benefit the facultative mutualism. The worker interfaces directly with the flower. The queen and males benefit from the mutualism (food) but do not interface directly with the flower. It is the worker who uses her considerable powers of judgment to forage the right flowers at the right time. The flower and her environ provide appropriate foraging clues: color, shape, scent, predators, and competitors. When she returns to the hive, the worker may dance good foraging locations to another worker in the hive. This represents a force multiplier for both the hive and the flower. A new flower emerges. A worker finds it and dances its location to an unemployed forager who visits the flower and dances to another. Soon a steady stream of hive workers is flying back and forth to the flower from the hive. Once the flower is pollinated, she shuts down the production of nectar and the procession of honeybees slows and stops as foragers learn that this location no longer produces. It is not difficult to suggest that eusociality, and particularly the communication capabilities of workers, evolved to help other workers judge and decide optimally. That eusociality also contributes to the good of the hive and to the good of the facultative mutualism is also now apparent. How honeybees do what they do requires insight into honeybee ontology. 6.2 Honeybee Anatomy Honeybees are insects, of the order Hymenoptera which includes wasps, bees, and ants. This is a large order of about one hundred twenty thousand species. As with other insects the honeybee body is tripartite: head, thorax, and abdomen. Like other insects the honeybee has eight legs. The head has compound eyes which can see ambient light and simple eyes to see the ultraviolet that helps honeybee workers see pollen and where to enter the flower. The honeybee’s eyesight is not strong, meaning the worker likely only sees shapes like trees without much detail. Honeybees can see more clearly when they are close to an object.59 They have mandibles that they use to manipulate honeycomb and a proboscis containing a tongue (like a straw) 59  Gould and Gould, The Honey Bee, 40–43.

64

Chapter 2

used not only to suck up nectar from flowers but also to taste. Workers have the longest proboscis. Because the males and the queen do not forage, their probosci are smaller. Finally, there are two antennae which serve as both a tactile and taste/smell organ. Inside the head is the brain of about one hundred thousand neurons (humans have eighty billion neurons). As we learn more and more about honeybee brain biochemistry and anatomy, we are discovering that many of the biochemical processes are like that of humans and other mammals. Brain structure is also similar, but instead of a prefrontal cortex that serves as the executive function of mammalian brains, honeybees and other Hymenoptera have mushroom bodies which are believed to serve a similar function.60 Behind the head is the thorax where the legs and the wings and 60  Z  hengzheng S. Liang et al., “Molecular Determinants of Scouting Behavior in Honey Bees,” Science 335, no. 1225 (2012): 1227. See also: Jean-Marc Devaud et al., “Neural Substrate for Higher-Order Learning in an Insect: Mushroom Bodies Are Necessary for Configural Discriminations,” Proceedings of the National Academy of Sciences 112, no. 43 (2015); Cordula Durst, Stefan Eichmüller, and Randolf Menzel, “Development and Experience Lead to Increased Volume of Subcompartments of the Honeybee Mushroom Body,” Behavioral and neural biology 62, no. 3 (1994); Birgit Ehmer and Wulfila Gronenberg, “Segregation of Visual Input to the Mushroom Bodies in the Honeybee (Apis Mellifera),” Journal of Comparative Neurology 451, no. 4 (2002); Syed Abid Hussaini and Randolf Menzel, “Mushroom Body Extrinsic Neurons in the Honeybee Brain Encode Cues and Contexts Differently,” Journal of Neuroscience 33, no. 17 (2013); HaDi

Emergence of the Flower and Honeybee Mutualism

65

most of the muscle mass of the honeybee is located. Honeybees have six jointed-legs, and like all insects they do not have an internal skeleton but an exoskeleton which on the honeybee is not as heavily armored as a ground beetle … the honeybee must conserve weight in order to fly. There are two large forward wings and two smaller aft wings. The larger wings serve to produce lift and the smaller wings help like rudders and stabilizers on an airplane to direct the flight of the honeybee. Unlike butterflies, the honeybee does not flap its wings, rather flight muscles, “[a]lternately compress the thorax vertically and horizontally.”61 The snap up and down motion produces a much higher wing beat per second than, for example, butterflies. Honeybees need to fly long distances (perhaps 10–12 km) quickly, so they need fast wing beats. Honeybees can hover like hummingbirds. The butterfly feeds for herself; the honeybee forager feeds both for herself but more importantly the hive. The forward pair of honeybee legs have notches to clean antennae. All legs assist in locomotion. The rear pair have what are called pollen baskets which consist of hairs to which sticky pollen from flowers adheres. All legs have taste receptors. Foraging workers often can be observed with what appear to be bright yellow bulbs on their upper hind legs which means that they have acquired much pollen on their journey. Some they leave behind when they forage other flowers, and some returns with the forager to the hive. The third chapter of the honeybee frame is the abdomen where there is the crop to store nectar for offloading in the hive, and digestive organs.62 The worker alone has at the very back of her abdomen a stinger that is used to defend the hive against honey raiders and other predators or invaders. The worker stings only once because when used, the stinger disembowels the worker and continues to pump venom into the target. The worker then dies. Whether she knows this will occur is not understood. MaBouDi et al., “Olfactory Learning without the Mushroom Bodies: Spiking Neural Network Models of the Honeybee Lateral Antennal Lobe Tract Reveal Its Capacities in Odour Memory Tasks of Varied Complexities,” PLoS Computational Biology 13, no. 6 (2017); Mary J. Palmer et al., “Cholinergic Pesticides Cause Mushroom Body Neuronal Inactivation in Honeybees,” Nature Communications 4, no. March (2013); Fei Peng and Lars Chittka, “A Simple Computational Model of the Bee Mushroom Body Can Explain Seemingly Complex Forms of Olfactory Learning and Memory,” Current Biology 27, no. 11 (2017); Christina Scholl et al., “Light Exposure Leads to Reorganization of Microglomeruli in the Mushroom Bodies and Influences Juvenile Hormone Levels in the Honeybee,” Developmental Neurobiology 74, no. 11 (2014); Nicholas J. Strausfeld, “Organization of the Honey Bee Mushroom Body: Representation of the Calyx within the Vertical and Gamma Lobes,” Journal of Comparative Neurology 450, no. 1 (2002). 61  Gould and Gould, The Honey Bee, 41. 62  Ibid.

66

Chapter 2

6.3 The Honeybee Caste System There are three castes in a honeybee hive, queen, male (drone), and worker. The queen is a fertile female who can live for five years or so and can lay up to two thousand eggs per day, each one deposited in its own cell. The egg metamorphoses into a larva, a pupa, and then emerges as an immature adult. How the larvae are fed and cared for by the nursery tenders determines into which caste it will be born. Nursery tenders coerce larva into their future caste mainly through nutrition allocation. Males have only one purpose and that is to mate with the queen on the mating flight. They have no other duties in the hive. Once they mate, the drones die. At the end of the mating season if there are males who have not successfully mated, they are pushed from the hive and they die from exposure. The worker, on the other hand, is an infertile female. Her ovaries are underdeveloped and only rarely do workers lay eggs, which are often destroyed by nursery tenders. The general maturation process takes the worker from cleaner, “nurse, food storer, and forager,” for the last twenty of her sixty to eighty or so adult days.63 This is a general trajectory because some workers do not become foragers and others may serve as hive guards. Foragers may at times serve as scout bees to find locations for foraging by other workers or as inveterate foragers. Forging takes a toll on workers. Inevitably they work themselves to death. Once a year, the hive swarms. The swarm is a tight ball of bees who leave the hive with the old queen. The swarm generally forms not far from the original hive. About a third to half of hive bees will swarm with the old queen. The swarm serves honeybees not only to increase genetic diversity, but also to increase the number of foragers (to the benefit of flowers), and to migrate honeybees to new territories. However, the swarm first must find a suitable new place to build a hive. In the swarm only about five percent of the swarm serve as scout bees to find suitable new hive locations. Others remain with the old queen. It is not yet understood how a worker chooses to become a scout bee. It is not known what decision process honeybee workers use to decide to stay with the new queen in the old hive or to swarm with the old queen to build a new hive, though studies suggest that there may be environmental cues that trigger behavioral changes in workers, and this may affect which workers leave and which workers stay.64 Swarms generally occur in the spring when the hive 63  T  homas D. Seeley, The Wisdom of the Hive (Cambridge, Ma.: Harvard University Press, 1995), 240. 64  There is emerging interest in how the swarm is generated. For more see: M. Woyciechowski and K. Kuszewska, “Swarming Generates Rebel Workers in Honeybees,” Current Biology 22, no. 8 (2012).

Emergence of the Flower and Honeybee Mutualism

67

births more workers than the hive can handle. During this time, the queen is put on a diet so that she can fly, and when the hive is ready for the new queen, the swarm occurs.65 7

The Moral Honeybee

The honeybee is embodied and embedded in her world. Honeybees of all castes exist together in a hive where there is little strife. Each individual has a job to do and, remarkably, fifty-thousand or so hive mates who require the same nourishment do not compete with each other for resources. Perhaps one reason that this is the case is that honeybees exist in the present, but their efforts are temporally towards the future e.g. the coming winter. In this they exemplify Maurice Merleau-Ponty’s assertion, “Time is not a line, but a network of intentionalities.”66 Honeybee intentionalities are always already towards the hive and the safety and security of the hive. As we learn more about the thickness of the experience of the honeybee in the world, we begin to understand with Merleau-Ponty: The solution of all problems of transcendence is to be sought in the thickness of the pre-objective present, in which we find our bodily being, our social being, and the pre-existence of the world, that is, the starting point of ‘explanations’, in so far as they are legitimate—and at the same time the basis of our freedom.67 Honeybees exist in a thick present, and while they anticipate the future in terms of what the hive needs, they do not dwell in suppositions about the future. They use the past to optimize the present through understanding each location in the meadow for whether there is good foraging or not. They can communicate what they know to others in the hive. They are mindful creatures who thicken the present through their attention to the phenomenological details they can discern, act upon, and communicate to others. The network of intentionalities for the honeybee worker is associated with finding resources in the world that she can return to the hive for its use, whether it be pollen, nectar, water, or tree gum. Outside the hive, her intentionality towards the world is principally oriented towards flowers 65  Gould and Gould, The Honey Bee, 27. 66   Merleau-Ponty, Phenomenology of Perception, 417. 67  Ibid., 433.

68

Chapter 2

which she does not unduly harm when she visits. Inside the hive she is oriented towards others and is receptive to their direction and guidance—all in service to the hive. This orientation towards the other rather than self is a feature of eusociality, but it also maintains the facultative mutualism, because when she enters the flower it is not for her but for her family—the hive. I speculate that the worker sees the flower as an extension of her hive because the flower is essential to the survival of the hive. She likely only records for orientation purposes other phenomenon like trees and rocks. If the flower is an extension of her hive, then there is every reason for her to treat the flower the same as the hive and its living members. This is not unlike the farmer who treats the milk cow as one of the family because she provides for the family. The morality of the hive extends into the world because the flower is an extension of the hive family which she strives to bring horizonally closer to her hive. Every action she takes, from remembering the locations of forageable flowers, to communicating their location to others from her strong memory are designed to bring these distal members of the hive closer together through optimal means. Both the hive and her flowers constitute her aggregate social group, all who are members of her family. She shows restraint towards all family members and she uses her considerable powers of reason and judgment to morally interact with all. It is important to study what is significant to the actors in the mutualism in order to explore how their social group is constructed and what this might mean for any moral constructs that result. For example, leafcutter ants are eusocial, but they cut down plant stems and leaves to bring to their hive and likely provide little in return to the plant. Once inside the ant nest, the leaves and stems are fed to special fungi. The ants cultivate consume the fungus, leaving enough to act as culture to produce more fungi. The flower represents both family and food for the honeybee but is otherwise left whole after the honeybee concludes her foraging. Contrast this with the fact that much of the fungi and significant amounts of plant matter that are fed to the fungi are sacrificed to feed the hungry leafcutter ants. There is a different moral construct at work in the leafcutter ant cultivation activities. For example, the leafcutter ant’s relationship with the fungus produces moral dilemmas associated with any process of farming (meat or plant) where the life form is consumed. Only resources dedicated to the pollinator are consumed by the pollinator in the flower and honeybee facultative mutualism. These are resources sacrificed to the mutualism, certainly at expense to the plant, but are not otherwise dedicated. Consider the milk cow. The cow produces milk for the farmer but is not otherwise harmed directly. However, her calf which started her lactation process likely was slaughtered as veal or was sent to another farm as dairy

Emergence of the Flower and Honeybee Mutualism

69

replacement. The fact that the flower and honeybee both benefit and exploit each other engenders a relationship and a moral construct that may be unique to facultative mutualisms and perhaps the flower and honeybee relationship. They are co-equal in their gain or loss without unduly sacrificing anything more than they are willing to give. Nor do the flower or the honeybee attempt to exploit more than the other is capable. Over millions of years, both species co-evolved to their present state. Each must have discovered something good in this co-evolutionary process beyond the fact that they became more capable of benefitting and exploiting each other physically. As they became more and more dependent upon the other, they developed a relationship and stronger moral reciprocity which, for the flower and honeybee facultative mutualism, has lasted a million years. This moral reciprocity that exploits and benefits both parties without undue harm to either produces a unique social and moral construct that is worth studying and comparing to other social constructs. More about honeybee hive, foraging, and swarm behavior will be considered in the next chapter on epistemology and behavior. Cited References Armbruster, W. Scott. “Evolution and Ecological Implications of ‘Specialized’ Pollinator Rewards.” Chap. 3 In Evolution of Plant-Pollinator Relationships, edited by Sébastien Patiny, 44–67. Cambridge, UK: Cambridge University Press, 2011. Bell, Adrian D. Plant Form: An Illustrated Guide to Flowering Plant Morphology. Portland, Oregon: Timber Press, 2008. Buchmann, Stephen L. “Bees Use Vibration to Aid Pollen Collection from Non-Poricidal Flowers.” Journal of the Kansas Entomological Society 58, no. 3 (1985): 517–25. Devaud, Jean-Marc, Thomas Papouin, Julie Carcaud, Jean-Christophe Sandoz, Bernd Grünewald, and Martin Giurfa. “Neural Substrate for Higher-Order Learning in an Insect: Mushroom Bodies Are Necessary for Configural Discriminations.” Proceedings of the National Academy of Sciences 112, no. 43 (2015): E5854–E62. doi:10.1073/pnas.1508422112. Durst, Cordula, Stefan Eichmüller, and Randolf Menzel. “Development and Experience Lead to Increased Volume of Subcompartments of the Honeybee Mushroom Body.” Behavioral and neural biology 62, no. 3 (1994): 259–63. Ehmer, Birgit, and Wulfila Gronenberg. “Segregation of Visual Input to the Mushroom Bodies in the Honeybee (Apis Mellifera).” Journal of Comparative Neurology 451, no. 4 (2002/09/30 2002): 362–73. doi:10.1002/cne.10355.

70

Chapter 2

Flannery, D. T., and M. R. Walter. “Archean Tufted Microbial Mats and the Great Oxidation Event: New Insights into an Ancient Problem.” Australian Journal of Earth Sciences 59, no. 1 (2012): 1–11. Friis, Else Marie, Peter R. Crane, and Kaj Raunsgaard Pedersen. Early Flowers and Angiosperm Evolution. Cambridge, UK: Cambridge University Press, 2011. Gadagkar, Raghavendra. “Why the Definition of Eusociality Is Not Helpful to Understand Its Evolution and What Should We Do About It.” Oikos 70, no. 3 (1994): 485–88. doi:10.2307/3545789. Gould, James L., and Carol Grant Gould. The Honey Bee. Edited by Carol Grant Gould New York: Scientific American Library: Distributed by W. H. Freeman, 1988. Hussaini, Syed Abid, and Randolf Menzel. “Mushroom Body Extrinsic Neurons in the Honeybee Brain Encode Cues and Contexts Differently.” Journal of Neuroscience 33, no. 17 (Apr 24 2013): 7154–64. doi:10.1523/JNEUROSCI.1331-12.2013. Liang, Zhengzheng S., Trang Nguyen, Heather R. Mattila, Sandra L. Rodriguez-Zas, Thomas D. Seeley, and Gene E. Robinson. “Molecular Determinants of Scouting Behavior in Honey Bees.” Science 335, no. 1225 (Mar 9 2012): 1225–28. doi:10.1126/ science.1213962. MaBouDi, HaDi, Hideaki Shimazaki, Martin Giurfa, and Lars Chittka. “Olfactory Learning without the Mushroom Bodies: Spiking Neural Network Models of the Honeybee Lateral Antennal Lobe Tract Reveal Its Capacities in Odour Memory Tasks of Varied Complexities.” PLoS Computational Biology 13, no. 6 (Jun 2017): 1–29. doi:10.1371/journal.pcbi.1005551. Merleau-Ponty, Maurice. Phenomenology of Perception. Translated by Colin Smith. London: Roudedge and Kegan Paul, 1962. Michez, Denis, Maryse Vaderplanck, and Michael S. Engel. “Fossil Bees and Their Plant Associates.” Chap. 5 In Evolution of Plant-Pollinator Relationships, edited by Sébastien Patiny, 103–64. Cambridge, UK: Cambridge University Press, 2011. Mittelbach, Moritz, Sandro Kolbaia, Maximilian Weigend, and Tilo Henning. “Flowers Anticipate Revisits of Pollinators by Learning from Previously Experienced Visitation Intervals.” Plant Signaling & Behavior e1595320, no. e1595320 (2019): 1–5. doi:10.1080/15592324.2019.1595320. Nancy, Jean-Luc. Being Singular Plural. Stanford: Stanford University Press, 2000. Nowak, M. A., C. E. Tarnita, and E. O. Wilson. “The Evolution of Eusociality.” Nature 466, no. 7310 (Aug 26 2010): 1057–62. doi:10.1038/nature09205. Palmer, Mary J., Christopher Moffat, Nastja Saranzewa, Jenni Harvey, Geraldine A. Wright, and Christopher N. Connolly. “Cholinergic Pesticides Cause Mushroom Body Neuronal Inactivation in Honeybees.” Nature Communications 4, no. March (2013): 1–8. doi:10.1038/ncomms2648. Peng, Fei, and Lars Chittka. “A Simple Computational Model of the Bee Mushroom Body Can Explain Seemingly Complex Forms of Olfactory Learning and Memory.”

Emergence of the Flower and Honeybee Mutualism

71

Current Biology 27, no. 11 (2017/06/05/ 2017): 1706. doi:https://doi.org/10.1016/ j.cub.2017.05.037. Scholl, Christina, Ying Wang, Markus Krischke, Martin J. Mueller, Gro V. Amdam, and Wolfgang Rössler. “Light Exposure Leads to Reorganization of Microglomeruli in the Mushroom Bodies and Influences Juvenile Hormone Levels in the Honeybee.” Developmental Neurobiology 74, no. 11 (2014/11/01 2014): 1141–53. doi:10.1002/ dneu.22195. Seeley, Thomas D. The Wisdom of the Hive. Cambridge, Ma.: Harvard University Press, 1995. Simard, Suzanne W., David A. Perry, Melanie D. Jones, David D. Myrold, Daniel M. Durall, and Randy Molina. “Net Transfer of Carbon between Ectomycorrhizal Tree Species in the Field.” Nature 388, no. August (1997): 579–82. Soltis, Douglas, Pamela Soltis, Peter Endress, Mark W. Chase, Steven Manchester, Walter Judd, Lucas Majure, and Evgeny Mavrodiev. Phylogeny and Evolution of the Angiosperms: Revised and Updated Edition. Chicago, Ill.: University of Chicago Press, 2018. Spivak, M., R. Masterman, R. Ross, and K. A. Mesce. “Hygienic Behavior in the Honey Bee (Apis Mellifera L.) and the Modulatory Role of Octopamine.” Journal of Neurobiology 55, no. 3 (Jun 2003): 341–54. doi:10.1002/neu.10219. Strausfeld, Nicholas J. “Organization of the Honey Bee Mushroom Body: Represen­ tation of the Calyx within the Vertical and Gamma Lobes.” Journal of Comparative Neurology 450, no. 1 (2002/08/12 2002): 4–33. doi:10.1002/cne.10285. Visscher, P. Kirk. “The Honey Bee Way of Death: Necrophoric Behaviour in Apis Mellifera Colonies.” Animal Behaviour 31, no. 4 (1983): 1070–76. Wang, Xin. The Dawn Angiosperms. Heidelberg, Germany: Springer, 2010. Whitney, Heather M., Lars Chittka, Toby J. A. Bruce, and Beverley J. Glover. “Conical Epidermal Cells Allow Bees to Grip Flowers and Increase Foraging Efficiency.” Current Biology 19, no. 11 (2009/06/09/ 2009): 948–53. doi:https://doi.org/10.1016/ j.cub.2009.04.051. Winship, David, and Shusheng Hu. “Coevolution of Early Angiosperms and Their Pollinators: Evidence from Pollen.” Palaeontographica Abteilung B 4, no. 6 (2010): 103–35. Woyciechowski, M., and K. Kuszewska. “Swarming Generates Rebel Workers in Honeybees.” Current Biology 22, no. 8 (Apr 24 2012): 707–11. doi:10.1016/j.cub .2012.02.063.

Chapter 3

Flower and Honeybee Epistemology and Behavior 1 Introduction It is important to heed the admonition of Martin Giurfa, that the non-scientific community creates misunderstandings attributing human qualities to animals, thus: “[t]he critical question is not, therefore, whether insects achieve ‘marvelous feats’, but how they achieve them.”1 Ontology and morphology gives us some guidance of how flowers and honeybees can do what they do. However, what they know, how they know what they know, and how they make decisions that ultimately require morphological processes to effect these decisions, are answers we much search for to discover how flowers and honeybees achieve marvelous feats. These are the subject of epistemological and behavioral inquiry. Much has been learned how flowers interact with their environment and what processes plants use to access necessary resources such as light, water, and minerals. While plants are rooted in place, they have developed ontological strategies of growth to reach towards light. Such efforts would be haphazard if the plant could not sense light. However, the tips of shoots have light-sensing cells that guide the movement or growth of existing stems or leaves towards the light. Light-sensitive cells are gravity-negative oriented, reaching upward and outward to find light. On the other hand, underground roots cannot use light to find water or minerals and must rely upon mycorrhizae fungus to aid in sensing and then finding sources of minerals which the mycorrhizae then process into substances both useful for the plant and the fungus. Roots are generally gravity-positive oriented, meaning they tend to grow downward and outward in the soil, rather than up (there are exceptions to this). Roots must also be able to sense dampness in order to move towards productive water sources. Herbivores like caterpillars abound. Plants have developed processes to identify when leaves are being munched on by insects and other herbivores and have evolved methods for releasing noxious chemicals to repel herbivores. Plants also use gaseous ethylene that alerts other plants of encroaching predation. Ethylene also is the chemical that encourages fruit to ripen. Plants use 1  Martin Giurfa, “Cognition with Few Neurons: Higher-Order Learning in Insects,” Trends in Neurosciences 36, no. 5 (2013): 285.

© Koninklijke Brill NV, Leiden, 2020 | doi:10.1163/9789004428546_005

Flower and Honeybee Epistemology and Behavior

73

many hormones and pheromones to regulate their processes and, in some cases, communicate predation or kinship with other plants and even flower availability (perfume) to attract pollinators. There are many differences between plants and animals. Plants produce their own carbohydrates (sugar) but absorb minerals and water through their roots; animals (generally) consume protein, carbohydrates, and minerals produced by other life forms (though many get some minerals directly from the soil). Some animals drink water; others get water from what they eat. Higher order animals have brains or central processing systems. While plants have no brains, they do have distributed processing systems that interact with each other in service to the entire plant. Plant processes interact with each other based upon internal and external stimuli and electro-chemical signals but do so without a brain. An animal brain receives input from various phenomenon and directs diverse processes, particularly its capabilities of locomotion, to serve the needs of the animal. The plant can grow, die back, or move certain organs like stems or roots towards nutrients, light, or away from competitors or predators. The flowering plant cannot move from its location as many animals can. Even the animal called a barnacle that becomes permanently attached to a rock, in its larval stage was mobile and used this mobility to find an appropriate place to attach itself. Honeybees are eusocial insects, meaning that they have overlapping maturation generations, share the care for young, and have a caste system consisting of the fertile egg-laying queen, a small number of fertile males whose job it is to mate with the queen, and workers, sterile females who through their lifetime can perform many jobs, from tasks in the nursery, to foraging outside of the hive, or as guards to prevent other hives from raiding honey stores or to fight-off predators. There is a significant body of literature on honeybee behavior and epistemology. Like other insects, honeybees have brains that perform tasks similar to those of vertebrates and mammals, and even use similar chemicals in these activities. There is little strife in the hive, though there are both chemical, and behavioral feedback mechanisms to coerce individuals to behave in ways that might better serve the hive. Honeybees can sense their world, process what they see, feel, hear, taste, and touch, and can judge what they believe is the optimal approach to what they intuit. They have developed elaborate behavioral strategies to communicate the presence of forageable flowers and to communicate suitable new locations to build hives during the process called the swarm. These so-called waggle dances are explored in this study to give insight into how honeybees interact with their environment and each other. However, it is also important to understand that honeybees are also adept at flying great distances from the hive and are able to find their way

74

Chapter 3

back which means they have the capability of identifying and remembering markers such as trees, meadows, or streams to follow back to the hive and to return again to the foraging site. They can also identify suitable foraging sites and avoid predators at these sites or in flight. Plants do not have brains, but recent studies have shown that plants, using their distributed existential systems, have considerable capabilities to make decisions and even communicate with other plants. 2

Angiosperm Epistemology and Behavior

2.1 Plant Knowing A key element of the emergence of morality is the ability to reason or judge. Judgment is also important in optimization with a further requirement that the creature chooses the best option considering the circumstances. The actions of thousands of creatures making optimal decisions produces MEP which is consistent with the second law of thermodynamics. The question that is critical to the emergence of morality in the flower and honeybee facultative mutualism is to what extent can flowering plants judge or reason. If they cannot, then the effort to develop a picture of morality in their relationship with honeybees is likely futile. Both Marder and Nealon have explored how philosophy has either ignored plant existentiality, dismissed it, or has relegated it to the lowest possible level of being in the hierarchy of life.2 Until recently, science has also largely ignored the being-becoming or existentiality of plants. There are recent studies of plant behavior that Daniel Chamovitz and others have reviewed that do show that plants judge based upon environmental conditions presented to them. A most difficult problem is that plants have no neurons, so what we think about being and becoming for plants must come without animal neurons and brains. Plants, like animals, have multiple systems that perform specific functions like growth, photosynthesis, absorption of water and minerals, searching for light and water, and protection against predation. Like animals, plants use both chemical and electrical means in these processes that help the plant make optimal judgments. What is difficult to wrap one’s head around is that they accomplish many of the same things that animals do without any central nervous system. They simply do what they do without 2  See, for example: Jeffrey T. Nealon, Plant Theory (Stanford, Ca.: Stanford University Press, 2015); Michael Marder, Plant-Thinking: A Philosophy of Vegetal Life, None (New York: Columbia University Press, 2013), Book; “Plant-Soul: The Elusive Meanings of Vegetative Life,” Journal of Environmental Philosophy 8, no. 1 (2011).

Flower and Honeybee Epistemology and Behavior

75

an executive function (brain). Their connected and interconnected processes, including vascular processes, serve specific functions and work together to make appropriate and optimal decisions for the plant and perhaps other plants. Therefore, if philosophy is to develop a way of thinking about plants, it needs to do come up with different ways of describing plant existentiality. Important to this study of the emergence of morality in nature, we must be able to describe how these brainless plants can develop a relationship with and form a social group with an insect that does have a central nervous system and a complex executive functioning brain. Through Daniel Chamovitz and others, this chapter reviews recent research on the existential capabilities of plants. Necessarily, the researchers in this study have tried to relate animal process terms like seeing, hearing, and feeling to plant processes and existentiality with some difficulty. While plants, for example, can sense and process what to do when light hits light-receptive cells, they cannot see like the honeybee. However, both species respond to light using the capabilities they have and therefore, while different, are adjusting behaviors related to what light they have sensed. Plants cannot talk, but they can communicate. The color of their flowers becomes an advertisement to honeybees either to forage or not. We can say that coloration, by in large, is genetically derived, but how did evolution discover the need to produce the attractive color? The activities of many generations of predecessor flowers, perhaps epigenetically or through active decision-making, must make choices that help the mutation take hold and maintain that mutation for as long as it attracts honeybees. Therefore, when we talk about plant existentiality, we must consider the plant under her terms. Unfortunately, most of the terms we can use to describe the flowering plant’s terms are animal terms that are inadequate to explain just what plants are experiencing. Given these limitations and caveats, this chapter must establish whether flowering plants can reason or judge for this effort of discovery of morality in nature to continue. Chamovitz explains that there are certain genes that helps a plant be aware, “[i]f it’s in the light or the dark” and these same genes are in humans and, “[a]lso regulate (among other developmental processes) response to light in both.”3 Photosynthesis began with the evolution of stromatolites 3.7 billion years ago and the first great extinction was caused by these creatures who introduced oxygen in large quantities into the atmosphere that killed off most

3  Daniel Chamovitz, What a Plant Knows: A Field Guide to the Senses, Kindle ed. (Oxford, UK: Scientific American/Farrar, Straus and Giroux, 2012), 3.

76

Chapter 3

early anaerobic life.4 One requirement for photosynthesis is that suitable sunlight must be available. Because plants are rooted in place, they developed strategies to bend or grow organs towards the light. The meadow is a dynamic place. In spring before the leaves emerge on the trees, the plants underneath the canopy may find suitable light for photosynthesis, but as trees unfurl their leaves and grow, lower plants must find sunlight to survive. Chamovitz uses the word know, in association with plants but acknowledges its use is unorthodox, “Plants don’t have a central nervous system; a plant doesn’t have a brain that coordinates information for its entire body. Yet different parts of a plant are intimately connected, and information regarding light, chemicals in the air, and temperature is constantly exchanged between roots and leaves, flowers and stems, to yield a plant that is optimized to its environment.”5 The plant’s environment is limited to the place where it has taken root. The plant therefore does not need capabilities that animals require to move quickly. However, a plant must be capable of sensing its environment, learn from that environment, and make appropriate and optimal choices on how to react to environmental stimuli. Because it is rooted in one place it must become a better knower of place than its more mobile counterparts, the animal. Without a mind, the plant is ever ‘mindful’ of its environment through consistent and changing cues that it senses continually. Some flowering plants, as will be discussed, use epigenetic processes to pass along some of what they have learned to offspring, not through teaching or mutation, but through changing the expression of specific genes that do not alter the chromosomal sequence. The plant must have capabilities to sense the environment she is in because she cannot escape the local ecology in which she sprouted. She needs light, water, and minerals to survive and the flowering plant in this study must produce flowers that will attract honeybee foragers for pollination. She therefore must be able to adjust herself to changes in light, water, and minerals. Unlike many animals, plants do not stop growing during their lifetime, whether it be to move stems towards the light, grow taller to reach the canopy and sun, or to grow roots to reach water and minerals, or all the above.

4  Prior to cyanobacteria the oceans were anoxic, “During stage 1, from 3.85 to 2.45 Ga, atmospheric O2 was almost certainly less than a few parts per million, except possibly during the period between 3.0 and 2.8 Ga. The oceans were almost certainly anoxic except perhaps in oxygen oases within the photic zone” Heinrich D. Holland, “The Oxygenation of the Atmosphere and Oceans,” Philosophical Transactions of the Royal Society B: Biological Sciences 361, no. May (2006): 912. 5  Chamovitz, What a Plant Knows: A Field Guide to the Senses, 5.

Flower and Honeybee Epistemology and Behavior

77

Higher order animals have central processing systems, not only to maintain the organism, but also to process phenomenological input. Plants distribute diverse maintenance and processing systems in different parts of the plant where they are connected by vascular systems. However, these systems must often interact in ways that support the plant’s existence. Animals, as mobile creatures, require quick response to phenomenon, whether food or predator. Plants that are imperiled also must react quickly. However, their processes for moving even hormones that react to danger are slower, measured not always in seconds or less, but sometimes in minutes or hours. In the meadow there are honeybees with brains and flowers with interactive processes. There are also the smallest single or multi-cell creatures in mud puddles that use flagella to move and do not have brains to facilitate movement towards or away from food or predation. Nature provides different affordances and constraints to existential beings based upon the ecological conditions in which the creature exists. 2.2 Plants and Light Plants discern the ultraviolet and infrared. Chamovitz explains, Plants can tell when there is very little light, like light from a candle, or when it’s the middle of the day, or when the sun is about to set into the horizon. Plants know if the light is coming from the left, the right, or from above. They know if another plant has grown over them, blocking the light, and they know how long the lights have been on.6 In general, plants respond phototropically only to blue light.7 Plants have developed methods not only to record the presence of particular wavelengths of light but also to retain that information over time. They do this without memory neurons. Plant processes use biochemistry to accomplish this task just as animals use biochemicals to produce memory and store it. While the processes may be different, they are towards the same end, the use of light to record time and temporality that can be used by the animal or plant to begin, sustain, or end processes towards the maintenance of the health of the creature. A classic experiment conducted by Charles Darwin discovered that phototropism or sensitivity to light occurred at the tip of the seedling and not in other places on the young plant, even though the seedling bent further down

6  Ibid., 9–10. 7  Ibid., 12–13.

78

Chapter 3

the stem.8 However, more recent experiments show that phototropism is found in the leaves of plants and if all the leaves are cut off, the plant will not respond phototropically to the light.9 Later experiments show that plants engage in photoperiodism, meaning, “[p]lants measure how much light they take in.”10 Scientists also have discovered that, “Plants were differentiating between colors: they were using blue light to know which direction to bend in and red light to measure the length of the night.”11 However, even a short burst of red light during the night can induce some plants to flower and others not to flower. Therefore it, “[i]s not the length of the day but the length of the continual period of darkness” that the plant measures.12 Plants are sensitive to different wavelengths of red light and react accordingly, and, “[w]e can say that the plant remembers the last color it saw.”13 Practically then, as Chamovitz explains, “In nature, the last light any plant sees at the end of the day is farred and this signifies to the plant that it should ‘turn off’. In the morning, it sees red light and wakes up. In this way a plant measures how long ago it last saw red light and adjusts its growth accordingly.”14 Some greenhouse operations, for example, use red light in early AM hours when the earth day is short to extend the growing period and perhaps induce flowering. Therefore, the use of light can be used to ‘fool’ greenhouse plants into thinking it is summer all the time, thus extending the growing period to year-round. Light is both a signal and food generator for the plant. Because they are rooted in one place, they must seek light in order to survive.15 So, motility, though it is restricted to one place, is important to the plant to seek the light it needs to survive. Chamovitz notes, “If a plant senses it is in the shade, it will start growing faster to get out. And plants need to survive, which means they need to know when to ‘hatch’ out of their seeds and when to reproduce.”16 Animals, on the other hand, have developed mobility to search for food sources and it is likely that animals ‘see’ differently from plants. Chamovitz explains, “Although plants see a much larger spectrum than we do, they don’t see in pictures. They don’t have a nervous system that translates light signals into pictures. Instead, they

8  Ibid., 14. 9  Ibid., 20. 10  Ibid., 17. 11  Ibid., 18. 12  Ibid., 20. 13  Ibid., 21. 14  Ibid., 22. 15  Ibid., 23. 16  Ibid., 26.

Flower and Honeybee Epistemology and Behavior

79

translate light signals into different cues for growth.”17 Despite millions of years of divergent evolution there are chemical receptors that humans and plants share such as cryptochromes which are blue-light receptors. Both humans and plants have circadian clocks to help each regulate the day-night cycle which, in humans, the cryptochromes and blue light reset our circadian clock.18 So, we can say that plants can sense or ‘see’ light but for purposes she needs like photosynthesis and growth. Animals use light to locate other creatures and food and process light differently from plants. However, both animals and plants use light for internal circadian regulation. 2.3 Plants and Chemical ‘Smells’ Chamovitz explores the different ways that plants can ‘smell’, “Plants know when their fruit is ripe, when their neighbor has been cut by a gardener’s sheers, or when their neighbor is being eaten by a ravenous bug; they smell it.”19 The hormone ethylene encourages ripening. Fruits emit ethylene and one rotting fruit can encourage all the others to ripen more quickly, “But ethylene is particularly important for plant aging as it is the major regulator of leaf senescence (the aging process that produces autumn foliage) and is produced in copious amounts in ripening fruit.”20 Colin Tudge records that prior to electricity, kerosene was burned in storage facilities where fruit ripened. Kerosene produces black oily soot but also ethylene that encourages ripening. When electric warming was introduced, the nasty soot disappeared, but the fruit did not ripen—the missing ingredient, ethylene.21 We know that animals are attracted to ripening and even rotting fruit. As the deer consumes the apples dropped in the meadow, the seeds pass through her digestive system and are deposited in places that may be far away from the parent tree. Therefore, ethylene serves the plant in her quest to produce viable offspring and to increase the species’ footprint on the land. Chamovitz notes that not all flowering plants use photosynthesis. Some vine plants lack chlorophyll and instead attach themselves to other plants and survive by eating their juices.22 Such vines appear to find their prey by smell.23 Trees and their leaves attacked by, for example caterpillars, emit a chemical signal that alerts other trees to produce noxious chemicals to ward 17  Ibid. 18  Ibid., 29. 19  Ibid., 33. 20  Ibid., 37–38. Emphasis in original. 21  Colin Tudge, The Tree (New York: Three Rivers Press, 2995), 272. 22  Chamovitz, What a Plant Knows: A Field Guide to the Senses, 39. 23  Ibid.

80

Chapter 3

off the caterpillars. The questions scientists have been asking is whether this chemical release is a form of intentionality on the part of the attacked plant or is just something that occurs with damage.24 More answers to this question are likely to emerge with future study. Lima beans respond to beetle infestations in two ways as Chamovitz notes, “The leaves that are being eaten by the insects release a mixture of volatile chemicals into the air, and the flowers (though not directly attacked by the beetles) produce a nectar that attracts beetle-eating arthropods.”25 Scientists are not convinced that plants ‘talk’ to each other, rather, perhaps, “[t]he neighboring plant must be practicing some form of olfactory eavesdropping on an internal signal actually intended for other leaves on the same plant.”26 In fact, experiments have shown that the chemical warnings emitted from an attacked leaf warn the other leaves on the same plant. Other plants that are close by and can ‘smell’ the same chemicals also benefit from the warning.27 Science does not yet know whether the release of chemicals that warn other plants is nothing more than a byproduct of the act to warn other leaves on the affected plant, or is part of a plant socialization structure we do not yet understand. Plants make salicylic acid to ward off infection, and this humans use to reduce fevers in the derivative of the chemical known as aspirin.28 Plants give off a variety of smells. “The smells induce different pollinators to visit the flower and seed spreaders to visit fruits,” and as Chamovitz records, the author Michael Pollan, “[i]nfers these aromas can even induce people to spread flowers all over the world.”29 Can plants smell? Chamovitz suggests, “[a]s of 2011 only one receptor for a volatile receptor, the ethylene receptor, has been identified in plants.”30 We await the results of future research, of which some is likely under way, for answers to this question. It is logical that the plant has developed sophisticated processes to find necessary light, water, and minerals in the location in which she germinates. She has no other choice but to resolve to thrive in the location where she becomes rooted. Naturally, as well, she must develop protection against predators, whether bacteriological or larger herbivores who will consume her photosynthetic leaves, attractive flowers, nutritious stems, or delectable roots which may be the targets of pigs and other animals who use considerable olfactory 24  Ibid., 37 & 39. 25  Ibid., 48. 26  Ibid., 51. 27  Ibid. 28  Ibid., 54. 29  Ibid., 57. 30  Ibid., 60.

Flower and Honeybee Epistemology and Behavior

81

powers to discover tubers and other edible roots. Flowering plants do all of this and perhaps even communicate with other plants without a single neural cell. 2.4 Plant Tactile Senses Plants do sense their surroundings and as Chamovitz suggests: Not only do they know when they’re being touched, but plants can differentiate between hot and cold, and know when their branches are swaying in the wind. Plants feel direct contact: some plants, like vines, immediately start rapid growth upon contact with an object like a fence they can wrap themselves around, and the Venus flytrap purposely snaps its jaws shut when an insect lands on its leaves. And plants seemingly don’t like to be touched too much, as simply touching or shaking a plant can lead to growth arrest.31 Plants use a form of hydraulics to control other movement. Cell walls are hard and can hold water, “By pumping water in and out of cells, the plant can control how much pressure is applied to the cell wall. The pulvinus cells are found at the base of each Mimosa leaflet and act as mini hydraulic pumps that move leaves. When the pulvinus cells are filled with water, they push the leaflets open; when they lose water, the pressure drops and the leaves fold into themselves.”32 Therefore plants can sense both the outside world and their internal processes and can react optimally to change they perceive. 2.5 Plants Don’t Feel Pain Do plants feel pain? Chamovitz suggests they do not, “While plants feel touch, they don’t feel pain. Their responses are also not subjective. Our perception of touch and pain is subjective, varying from person to person.”33 An experiment using a hot iron on tomato plant leaves generates an electrical signal that passes through the leaves and the stem, “The leaf did not feel pain. The tomato responded to the hot metal not by moving away from it but by warning other leaves of a potentially dangerous environment.”34 The advantage that the tomato plant has over the honeybee is if the honeybee organ (say a wing or a leg) is damaged, it cannot be replaced and the forager likely will perish. The tomato plant not only can warn other leaves and stems to move away from 31  Ibid., 61. 32  Ibid., 74. 33  Ibid., 82. 34  Ibid., 83.

82

Chapter 3

the heat, it can produce new organs (leaves, stems, and roots) to replace that which has been damaged. In places where there are frequent fires, some plants have evolved seeds with hard shells that pop open only when the fire arrives. Others have root systems that can produce new stems when those above the surface are burned off. While animals and plants ‘feel’ differently, Chamovitz notes similarities at deeper cellular levels: Despite the differences between the ways that plants and animals react to touch and other physical stimulations at the organismal level, at the cellular level the signals are hauntingly similar. Mechanical stimulation of a plant cell, like the mechanical stimulation of a nerve, initiates a cellular change in ionic conditions that results in an electrical signal. And just like in animals, this signal can propagate from cell to cell, and it involves the coordinated function of ion channels including potassium, calcium, calmodulin, and other plant components.35 2.6 Plants and Sound In 2012, Monica Gagliano, et al., studied chili plants and discovered that they could communicate with each other, but not necessarily through traditionally identified means, “Furthermore as seeds grow into seedlings, they are able to discriminate among neighboring species and modify their growth patterns accordingly, without necessarily relying on known determinants, such as volatile chemicals, direct physical contact or changes in infrared light wavelengths.”36 She speculated then that perhaps sound was a means of communication and called for additional plant acoustic studies.37 She noted previous studies where plants produce sound waves, but the question of how sounds are produced and what, if any, sounds plants can sense and react to is not understood.38 However, she does explain that a considerable number of flowering plants release pollen only after the vibrations of bee wings reach the appropriate frequency: “For example, some 20,000 species use buzz pollination where the pollen is released from flowers only when they are vibrated at

35  Ibid., 84. 36  Monica Gagliano et al., “Acoustic and Magnetic Communication in Plants: Is It Possible?,” Plant Signaling & Behavior 7, no. 10 (2012): 1347. 37  Monica Gagliano, “Green Symphonies: A Call for Studies on Acoustic Communication in Plants,” Behavioral Ecology 24, no. 4 (2012): 793–94. 38  “The Flowering of Plant Bioacoustics: How and Why,” Behavioral Ecology 24, no. 4 (2013): 800.

Flower and Honeybee Epistemology and Behavior

83

the correct sound frequency, a feat achieved by bees that have co-evolved to vibrate their flight muscles appropriately.”39 In subsequent studies Gagliano and fellow researchers began to see that plants do react to vibrations that produce what we call sounds. For example: We found that roots were able to locate a water source by sensing the vibrations generated by water moving inside pipes, even in the absence of substrate moisture. When both moisture and acoustic cues were available, roots preferentially used moisture in the soil over acoustic vibrations, suggesting that acoustic gradients enable roots to broadly detect a water source at a distance, while moisture gradients help them to reach their target more accurately.40 More work on plant bioacoustics must be done before we gain a clear picture on how plants ‘hear’ vibrations, what it means to them, and how they can make decisions on how to respond. However, for purposes of this study, it appears that many bees communicate their presence in the flower through wing beat vibrations which the flowers can then react to by releasing sticky pollen onto pollinators. Others like honeybees do not ‘buzz’ their wings but if Chamovitz is correct, the flowering plant can still feel the presence of the honeybee and likely can discern whether she is a predator or a pollinator. Therefore, the sophistication of flower and bee communication is more than just sight and scent, but also can involve acoustics associated with wing vibrations or the tactile vibration of feet on flowers. Consider also that plants are sessile creatures and must know more about the location into which they were born than animals who are more mobile. It stands to reason that flowering plants would evolve to improve their phenomenological sensing capabilities wherever possible. Whether all plants can ‘hear’ and respond to acoustic vibrations (other than buzz pollination) remains largely unanswered. 2.7 Plants Can Sense Gravity Plants when turned upside down slowly reorient leaves and roots.41 Is this gravity or something more? Chamovitz explains, “[t]he cells in the extreme ends of the root (in the region called the root cap) sense gravity and help a

39  Ibid. 40  Monica Gagliano et al., “Tuned In: Plant Roots Use Sound to Locate Water,” Oecologia 184, no. 1 (2017): 151. 41  Chamovitz, What a Plant Knows: A Field Guide to the Senses, 120.

84

Chapter 3

plant know where down is.”42 However, life is more complicated than sensing up from down as Chamovitz explains, “A plant can be pulled in many directions at once. Sunlight hitting a plant at an angle causes it to bend toward the rays, while the sinking statoliths within a plant’s bending branches tell it to straighten up. These often-conflicting signals enable a plant to situate itself in a position that is optimal for its environment.”43 2.8 Plant Memory Plants do not have memories like humans because they do not have brains, yet they can remember. Chamovitz notes that studies with pea plant tendrils provide evidence that with certain light or touch routines, that the tendrils, “[h]ad stored this information and recalled it once he [the researcher] placed it in the light.”44 Some plants need a period of cold weather to thrive or even flower. Cherry trees, for example, flower after winter when there is 12 hours of sunlight, but not in the fall when there is the same amount of sunlight. This has to do with a combination of gene triggers and other factors such as light and temperature.45 As has been mentioned and will be discussed in greater detail in a later chapter, plants can pass on knowledge to future generations through epigenetic processes. Chamovitz notes, “What is truly amazing about epigenetics is that it facilitates memory not only from season to season within a single organism but from generation to generation.”46 Recent studies have shown that there is transgenerational memory in some plants. Chamovitz explains, when these plants are stressed, they change expressions of their genes that are passed down to next generations. Even though the offspring did not experience the stressors their parents did, the offspring are given the epigenetic tools their parents learned to counteract these stressors.47 This occurs without genetic mutation (genes can express themselves in many ways), without changes in the chromosomal sequence, and without the parent ‘teaching’ the offspring. Chamovitz explains how this works, “[t]he parents formed the memory of the stress, retained it, and passed it on to their children,

42  Ibid., 121. 43  Ibid., 138. 44  Ibid., 143. Item in bracket added. 45  Ibid., 157. 46  Ibid., 160. FYI: “epigenetics is the study of heritable changes in gene expression or cellular phenotype caused by mechanisms other than changes in the underlying DNA sequence” Evgenya Popova and Colin J. Barnstable, “Epigenetics Rules,” Journal Of Ocular Biology, Diseases, And Informatics 4, no. 3 (2012): 93. Emphasis in original. 47  Chamovitz, What a Plant Knows: A Field Guide to the Senses, 129.

Flower and Honeybee Epistemology and Behavior

85

and the children recalled the information and reacted accordingly, in this case, with increased genetic changes.”48 Chamovitz notes that there are similarities in how plant and animal memory work, “Many of the mechanisms involved in plant memory are also involved in human memory, including epigenetics and electrochemical gradients. These gradients are the bread and butter of neural connections in our brains, the seat of memory as most of us understand it.”49 Plant memories, however, are procedural and not episodic as with animals.50 From Endel Tulving, Chamovitz understands that there are three levels of consciousness, procedural, sematic, and episodic.51 Plants maintain the lowest level of consciousness—procedural,“[t]he ability of organisms to sense and react to internal and external stimulation”, but not semantic or episodic.52 According to Tulving, procedural: “[i]s concerned with how things are done—with the acquisition, retention, and utilization of perceptual, cognitive, and motor skills.”53 Therefore, if we can agree with Tulving’s definition of consciousness, plants are minimally but only procedurally conscious. Consciousness, however, is a contentious issue that is subject to broad current debate in the literature. Whether Tulving is right that sensing and reacting to stimuli is a form of consciousness, is subject to considerable skepticism. However, we know that plants sense and react to stimuli, so whatever we call this, we understand that this is part of plant existence. We have seen that they can bend to light, grow roots towards water, some can react to vibrations of honeybees by loosening pollen, and can produce noxious chemicals when attacked by herbivores. Rather than suggest that plants are intelligent with all the controversy and debate over what intelligence means, Chamovitz says that plants are aware: Plants are acutely aware of the world around them. They are aware of their visual environment; they differentiate between red, blue, far-red, and UV lights and respond accordingly. They are aware of aromas surrounding them and respond to minute quantities of volatile compounds wafting in the air. Plants know when they are being touched and can distinguish different touches. They are aware of gravity: they can change 48  Ibid., 161. 49  Ibid., 162. 50  Ibid., 166. 51  Ibid. 52  Ibid. 53  Endel Tulving, “Memory and Consciousness,” Canadian Psychology/Psychologie Cana­ dienne 26, no. 1 (1985): 2.

86

Chapter 3

their shapes to ensure that shoots grow up and roots grow down. And plants are aware of their past; they remember past infections and the conditions they’ve weathered and then modify their current physiology based on these memories.54 This study will return to the subject of consciousness in both plants and honeybees later. For now, however, we can say with Chamovitz, that plants are aware, exhibit Tulving’s first level of consciousness, and can remember and use this memory to make optimal future decisions … This they do without any neurological memory cells. So far, studies have not found that plants generate emotions, nor do we have any information that they suffer.55 However, we are beginning to understand that some plant ecologies may interact more than what has been previously thought. 2.9 Tree Interdependence? Peter Wohlleben used his considerable powers of observation as a forester to consider what he has seen in the forest. Much of what he has observed he can link only tangentially to existing scholarship, but others are ripe for scientific investigation to determine why and how certain processes in the forest appear to occur. Likely there are electrical, chemical, genetic, or pheromonal causes to some of what he observes. There may also be symbiotic relationships in roots between plants, fungi, or microbes that have yet to be discovered or understood. For example, Wohlleben speculates that a forest full of trees is interdependent, that if one or more trees goes missing, gaps could increase sunlight on the forest floor that would dry it out and wind could enter the gaps causing even more treefall.56 While he notes that science has identified that trees of the same species in some forests have interconnected root systems, we do not have any good answers as to what benefits the trees derive from this interconnection. Is there nutrient exchange, electrical communication, pheromone communication, and, if so, what purposes do these exchanges serve? In a study (of non-angiosperm species) Kevin J. Beiler et al., found: In summary, we found that most trees in a multicohort old-growth forest were linked in a scale-free MN [mycorrhizal network], where large 54  Chamovitz, What a Plant Knows: A Field Guide to the Senses, 170. 55  Ibid., 172. 56  Peter Wohlleben, The Hidden Life of Trees: What They Feel, How They Communicate— Discoveries from a Secret World (Vancouver, BC Canada: Greystone Books, 2016), 4.

Flower and Honeybee Epistemology and Behavior

87

trees served as hubs, with implications for understorey regeneration and functional continuity in the stand. To ensure that old-growth Douglas-fir forests remain resilient and self-regenerative following disturbance, our findings support a management approach that conserves large trees or groups of trees and their mycorrhizal fungal associates.57 More on the composition of forests and the implications of such hub-like systems requires more investigation, but Beiler et al.’s, research suggests possibilities for ecological social group-type structures where old and larger trees might benefit proximal others. As has been noted, flowering plants do alert other flowering plants of herbivore predation. For example, Wohlleben explains that Acacia trees give off ethylene gas when giraffes nibble on their leaves. As we have seen elsewhere, ethylene is a powerful pheromone that has many uses. In this case, observers understand that ethylene encourages neighboring acacias to produce noxious chemicals that giraffes don’t like. Once out of range of the ethylene affected trees, the giraffes can browse for a short time without tasting the noxious chemicals.58 The tomato plant study, previously mentioned, suggests that more investigation needs to be done on root communication between same-species and even different species with identical or similar mycorrhizal networks.59 We have discovered that plants and the subject of this study, angiosperms, are quite remarkable creatures. They can sense and react to their environment, can communicate availability to pollinators, and even communicate predation through the air and, in some cases, through their roots. We speculated with Wohlleben that there might be more to the organization of the forest than just the random placement of trees. We have offered sensing and reaction as an elementary form of consciousness that is aware, even if this consciousness is not as rich as that of the human or the honeybee. The next major question we must ask is whether plants are intelligent beings. This study approaches plant intelligence through a philosophical lens, more to address questions associated with intelligence, not to develop empirical scientific answers to the question of plant intelligence.

57  K  evin J. Beiler et al., “Architecture of the Wood‐Wide Web: Rhizopogon Spp. Genets Link Multiple Douglas‐Fir Cohorts,” New Phytologist 185, no. 2 (2010): 551. 58  Wohlleben, The Hidden Life of Trees: What They Feel, How They Communicate—Discoveries from a Secret World, 7. 59  Yuan Yuan Song et al., “Interplant Communication of Tomato Plants through Under­ ground Common Mycorrhizal Networks,” PloS one 5, no. 10 (2010).

88

Chapter 3

3 Plant Intelligence—a Philosophical Discussion 3.1 Thinking and Being Martin Heidegger asks, “What is called thinking?”60 We must consider that question and ask, “Do plants think?” First, I will, with N. Katherine Hayles, suggest that there is a difference between cognition and thinking: To avoid confusion, I will reserve ‘thinking’ for what conscious entities such as humans (and some animals) do, and ‘cognition’ as a broader term that does not necessarily require consciousness but has the effect of performing complex modeling and other informational tasks. On this view, we can say that while all thinking is cognition, not all cognition is thinking.61 I suggest that the plant performs ‘complex modeling and other information tasks’ perhaps without consciousness as we have traditionally thought of consciousness (Perhaps even different from Tulving’s first level but with similar effects for the plant). I will have more to say about theories of plant consciousness in a later chapter. What I will maintain at this juncture is that plants cognize rather than think, leaving the term thinking to those beings like honeybees who have a separate neurocognitive processing system.62 I do not think that this distinction diminishes plant capabilities because the distinction between thinking and cognition when it comes to plant activities and 60  Martin Heidegger, What Is Called Thinking trans. Fred D. & J. Glen Gray Wieck (New York: Harper & Row Publishers, 1968). 61  N. Katherine Hayles, “Cognition Everywhere: The Rise of the Cognitive Nonconscious and the Costs of Consciousness,” New Literary History 45, no. 2 (2014): 201. 62  Whether minds that think have what is called a global workspace is a controversial idea. Michael Tye explains why he thinks that bees and humans are similar, “Finally, there is evidence that the neural architecture of the bee supports a global workspace. According to some scientists, this suffices for consciousness. No such claim is being made here. The point I wish to make is simply that there is an important functional similarity in the underlying architecture of bee and mammalian brains. This point, together with the points made above, undercuts attempts to defeat the use of behavioral similarities as grounds for experimental via an application of Newton’s rule and the same effect, same cause principle” Michael Tye, Tense Bees and Shell-Shocked Crabs (Okford, UK: Oxford University Press, 2017), 153. Flowers have no global workspace and their cognition likely is structured differently from that of humans and honeybees. Therefore, while we may eventually find that flowers somehow think, until we do, I suggest that we use the term cognition that implies similar functionality between plants and animals but without the intervening brain in plants.

Flower and Honeybee Epistemology and Behavior

89

Heidegger’s notion of what is presence in presence is no different in plants and animals. Heidegger answers ‘what is called thinking’ with: But in the meantime we have learned to see that the essential nature of thinking is determined by what there is to be thought about: the presence of what is present, the Being of beings. Thinking is thinking only when it recalls in thought the [what is presence in presence] That which this word indicates properly and truly, that is, unspoken, tacitly. And that is the duality of beings and Being.63 It is not controversial to suggest that flowering plants are beings if we can say that all life forms are beings. Let us assume, therefore, that all life forms are beings. We can also point to the flower and its plant as a what is present when we see it in the meadow. Is that all we can say that it is a being that is present? No, because we must ask what is presence in this presence? In other words, is there more to this being, the flowering plant, than just being and being present? We must consider what is the presence of this plant being that is present, and this is the more difficult task. Heidegger maintains that there is a two-fold nature to being. First there is the ontological presence—I see that there is a flower in the meadow. That gives us an object that we can locate phenomenologically. Yet this object we see we cannot discern much more than its relation to other objects, its dimensions, and colors and maybe its smell. If this plant is a being, to discern the second nature of being—the being of beings, we must explore what the presence of this presence is. Parmenides suggested in Fragment VIII of his proem about the being of beings: “That which exists, whatever it may be, is ungenerated and imperishable, homogeneous and continuous, unmovable and unchangeable and therefore the only thing there is.”64 This brings us to the question of what is the nature of this being of beings other than as a structured continuity that is unalterable? Is this being of beings like a rock stuffed into every being? What purpose would that serve? Would we not have to find quite different rocks to stuff into a dog, a flower, a honeybee, or a water buffalo? Intuitively we know that their existential and ontological existences are quite different. However, Parmenides suggests that this being of beings is undifferentiated, so a separate construct of being (rock) 63  Heidegger, What Is Called Thinking 244. Item in bracket added to replace Greek eov. 64  L. Tarán, Parmenides: A Text with Translation, Commentary, and Critical Essays (Princeton, NJ: Princeton University Press, 1965), 192. See also: John Palmer, Parmenides and Presocratic Philosophy (Oxford: Oxford University Press, 2009).

90

Chapter 3

is not possible for each being. Even so, Heidegger suggests that humans are the only beings who know they have being and, “[f]or which being is an issue for itself.”65 Can this being that understands it has being be fundamentally different from other beings who have being? Being must be something that engages the being in a way that is not only a presencing but also an opening to the world. This is the universality of being whether it applies to humans, flowers, or honeybees. Therefore, this being must be of the kind that can accept the attachment of being which means that not only the present being becomes a presencing, but also is open to the world. It is the openness to the world that is necessary for a being to be able to discern what is presence in presence. Is the flowering plant in the meadow a proper being? We have already said that this is the case, and therefore it must have been open to become attached by Parmenides’ being. This requires the plant to be open to the world. If such a being is not open to the world then the being of being would not serve any purpose and, as I maintain, could not be a being. Being requires an opening (for being to attach) in the world to be open to the world. What we have not yet explained for the plant is how it’s being that is open to the world can cognize, meaning how it can enable its presence in presence. This flowering plant in the meadow has no brain or nervous system. How can we suggest that it can cognize? The question can be further modified into what is required for cognition? First, there must be an opening to the world. We can say that the flowering plant is open to the world because it has multiple world sensing capabilities. However, what is required next is a means to communicate what the plant senses to appropriate organs in its body. Honeybees and humans have sensory nerves that report what the sense organs have discovered (the presence) to a brain that is tasked with discovering the presence of the presence. This involves not only discerning the presence of the presence, but also discerning the meaning of the presence of the presence to the creature itself. Then the creature must inform the various organs in the body of what to do about this presence for which now its presence of the presence has been assessed by the honeybee brain. Yet we know that the brain and the nervous system of honeybees and humans do not just direct the muscles to move, they report certain conditions to other organs that release chemicals into the vascular system that engage other organic processes such as increased heart rate or attention. The plant has no such second system for thinking, it only has a vascular system. Yet it must somehow cognize in order to direct systems and processes. We 65  Martin Heidegger, Being and Time trans. John MacQuarrie, & Edward Robinson (New York: HarperCollins Publishers, 1962), 182, H143.

Flower and Honeybee Epistemology and Behavior

91

know from science experiments that the plant is open to the world and can acquire information from the world from its different sensory capabilities from different organs. We know that certain plant stem cells can discern light and its direction, and roots can discern through their mutualism construct with mycorrhizae what nutrients have been discovered and are being processed by the mycorrhizae. So, the problem becomes evident straight away. How do plant organs communicate this information to other plant organs if there is no separate electrochemical highway to inform them that information, that the organ cannot directly sense but is important to that organ, is available? We can show how the vascular system connects to all organs of the plant. We know that this vascular system can move nutrients and pheromones created by one or more organs throughout the plant where they can be accessed where necessary and appropriate. We also know that plants release pheromones into the air which can alert the plant’s other leaves or its neighbors of attacking herbivores. We know that roots of some plants that are connected to other plants by mycorrhizae networks can communicate both pheromonal and other information to other plants. We know also that some plant roots when they encounter other roots discern whether the roots are from the same plant, same species, or other species. We also know that plants use not only chemical processes, but also bioelectric processes without a connective neural system. All of these are recognized plant processes. Without a central nervous system, it is tempting to suggest that the plant is nothing more than processes, perhaps sophisticated, and even interconnected and interactive processes, but nothing more. That this vascular system serves as a highway for these processes to operate, but there can be no cognition because the plant does not have neurons or a brain. However, it is perhaps also anthropocentric hubris to suggest that honeybees and humans are more than just processes without explaining how this could be. It is the province of philosophy to discern this ‘more than’, explain it, and demand its purposefulness. More about thinking and cognition as it relates to flowers and honeybees will be considered in the later discussion on consciousness. Now, however, it is important to better understand how a plant as present is also a presence. 3.2 The Flowering Plant as a Presence in the Present While Heidegger wants to discern the being of being human as the being who understands it has being, he refrains from dismissing the being of being from any other being. Even while he explains human thinking, he opens the discourse of thinking/cognition for other creatures such as the flowering plant in the meadow:

92

Chapter 3

It is enough if we dwell on what lies close and meditate on what is closest; upon that which concerns us, each one of us, here and now; here, on this patch of home ground; now, in the present hour of history.66 While he is speaking of human meditative thinking, he is not describing it as being ‘high-flown’, rather he is speaking in terms that can be translated into plant existentiality. The plant is ‘on this patch of home ground’ where it is fixed and has no choice but to ‘dwell on what lies close’. He notes also another problem with human thinking that it tends to wander from what is ‘in the present hour of history’ and what concerns us in this temporal moment. While the plant can retain experiential knowledge for use in the future, it has no process for being other than ‘on this patch of home ground, in the present hour of history’. Simply, the plant does not have the capacity to speculate on things other than those that confront the plant in this moment, e.g., where would be the optimal direction to grow a new stem, now that the canopy has shaded this branch. Recall that the Buddha associates speculative thinking with dukkha or suffering. The mind that wanders into speculation can ignore what the moment brings to cling to extraneous threads we might call wishful thinking. Wishfulness clings to that which is wished and for what purpose can that serve but to deny the present in favor of clinging to that which may not ever be possible to obtain? The Buddha also admonishes that even if the wish is granted, clinging, grasping, and craving even for the being of the being itself produces just more dukkha. The flowering plant in the meadow is always already in the present. Therefore, even though she can create memory through biochemical processes that may or may not be epigenetic (for herself or offspring), she will use those remembrances only when necessary as in what the moment requires. We know that honeybees and humans also create biochemical memories in neurons and neural systems. Therefore, we cannot rule out the idea that there can be similarities between plant and animal memory formation or the purpose for storing memories in order to facilitate appropriate and effective responses when confronted with the same conditions in the future. Flowering plant mobility is restricted to growth in place. She is in contact with the ground, the air, and the light from the sun. As she cannot move from where she roots, she needs only to be able to discern ‘that which is close’, ‘on this patch of home ground now’. Therefore, she assesses probabilities to produce the best decision based upon present circumstances and the information generated from these. In search of nutrients or the sun to make them, the flower can only extend or modify her reach into the surrounding dirt, air, or water for 66  Discourse on Thinking: A Translation of Gelassenheit (New York: Harper & Row, 1969), 47.

Flower and Honeybee Epistemology and Behavior

93

she cannot venture into another meadow.67 Roots grow into the soil. Promising root avenues receive more growth than those that do not. Non-promising roots die back in order to conserve energy. Vines extent tendrils that also seek probable purchase based upon what the vine has encountered in previous movement. The plant continues to make decisions using probability assessments of where light is or where nutrients have or could be found. Therefore, any speculation she makes as to where light will be found is based upon information she has at this moment. The flower does not muse about retirement and what it will take to retire comfortably; assuredly nor does the honeybee. Locality restriction does simplify both what the flower is capable of or needs to accomplish to exist. However, the plant is open to the world and is a going towards what it needs to survive, or a going away from that which does not or would otherwise harm the plant. This going towards or going away from in the plant does not take the form of locomotion but growth, extension, and retraction from a fixed location. 3.3 Non-Duality in Plants There are duality constructs we use to describe elements of nature: night day, sun shadow, predator prey, winner loser. However, this duality is illusory and not ideal. Day and night on the equator are equal each day, but the amount of sunlight depends upon whether the season is wet or dry. Therefore, all days and nights are not the same. Predators often do not capture prey and prey often escape predators. Both predators and prey exhibit scars of their past encounters. Sunlight and shade can vary with the position of the sun in the sky, and the shaded forest floor plant in the morning may be sunlit in the afternoon. The plant is decidedly non-dual because it is always already all middle. We can suggest that both the subject and object of the plant are non-dual. This is because the plant has no intervening organ that speculates. It can act only upon what is presented in the moment. Even so, the plant makes optimal decisions based upon information provided. Because the root may not have good information as to where nutrients can be found, it must grow numerous roots in different directions until the probability of nutrient resource finding can be improved. We can therefore say that the root system, not a separate mind, must use its capabilities of growth or retreat in order to discover that which it needs. The root may augment growth capabilities with sensing capabilities such as water vibration, dampness, and information that mycorrhizae transmit to the root that they are now processing valuable chemicals or minerals. 67  Yet her color, shape, and scent call in honeybees from many kilometers away.

94

Chapter 3

The problem is that it is not difficult to suggest that both honeybee and human minds are separate from the body in some form or the other. This was Rene Descartes’ pronouncement. That somehow mind supervenes the brain and the body and thus produces a gestalt that is greater than the sum of the brain and body’s ontological structure. One of the problems with this notion is that mind is associated with the distal brain and therefore the mind somehow is not located within the center of the being (the honeybee or human existent). While the leaf and root organs of the flowering plant in the meadow are distally located, the plant is proximally connected by its vascular system. Michael Marder is right, that the seed germinates in the middle, extending both the root into the ground and the stem into the sky from the middle.68 The flowering plant in the meadow has no mind to cause this illusion of duality, therefore the plant is always already in the middle of itself. What do I mean by middle? The plant eventually grows up, down, and in circumference. The middle, that began with the seed, therefore, becomes stretched. An imperfect analogy is the balloon. As we blow up the balloon, the surface area expands, but the middle of the balloon is still the middle. It has not moved. This analogy is imperfect because the plant, unlike the balloon, can increase its mass. In the fall, the tulip (bulb plant) compresses itself (like the balloon that is deflated) into its bulbous root where it remains until the spring where once again it begins to grow in all suitable directions. If the flowering plant in the meadow is always already on this patch of home ground in the present hour of history, it is, by definition, a ‘mindful’ creature (which we can describe as non-mindful mindfulness). How can this be if the plant does not have a mind? Buddhist mindfulness dissolves the notion of a separate mind and locates the being of being in the core of the being, the middle. The middle way of Buddhism is not simply the avoidance of extreme behavior but locating, orienting, towards the presence in presence. The plant has no intervening device (brain) that would cause the creature to distally locate or even relate its presence away from the presence in presence. As such the presence in presence is always centered in the whole of the plant. The plant cannot out-think herself because there is only that which is presence in presence and nothing more. Plant existentiality as coming from the middle suggests that the plant exists non-dually. Plant organs: roots, stems, leaves, flowers, are connected by a vascular system that brings nutrients from roots and sugars created by the leaves into the middle, always the existential middle of the plant because there are no ends to the plant—everything even at the extremities is still in the middle of 68  Marder, Plant-Thinking: A Philosophy of Vegetal Life, 63.

Flower and Honeybee Epistemology and Behavior

95

it all. Despite the location of these different organs that may be meters apart, there is always already a middle without ends. There is a reason for this nonduality because there is no mind-body problem for the plant. The plant has no distal mind that humans and insects do, with a neural system that operates in parallel to its vascular system. Thus, the plant can have no cognitive sense that it is something different from the body of the plant and its functionality. There is only plant. This is contrary to what Rene Descartes suggested that the human mind and the body are separate and a dualism. Arthur Schopenhauer refuted the notion of duality: So we are inseparably connected as necessary parts of one whole, which includes us both and exists through us both. Only a misunderstanding can set up the two of us as enemies in opposition to each other, and lead to the false conclusion that the one contests the existence of the other, with which its own existence stands and falls.69 Duality likely is illusory. There are other plant/animal similarities. The plant’s vascular system not only distributes nutrients to where they are needed, but it also sends chemical or electrochemical signals (like animals do) from where the need for them are generated to other parts of the plant that may have some use for them. While the processes of leaf photosynthesis and root osmosis are different, they are not independent from the plant which exists as it does without the dualist illusion of a separate mind that is the executive of it all. The plant has no executive will which it is why the plant exists in the middle and without ends. Schopenhauer suggested that all beings have will. Plant will is not distally formed but is associated with all its interrelated processes. Senses produce input that are assessed by processes and action is taken where necessary to respond to what has been sensed e.g. release noxious chemicals to all leaves when an herbivore is detected munching just one leaf. If the plant is all middle, and it can react to stimuli by engaging multiple organs to react, can we call any of what plants do intelligence? 3.4 Plant Intelligence as Distributed Intelligence Plant intelligence, of course, is not the same as animal intelligence. Plant intelligence is not routed through a central processing system that determines courses of action. Rather plant intelligence is distributed, meaning that each organ, though separate in functionality from others, is interconnected with all 69  Arthur Schopenhauer, The World as Will and Representation Ii, trans. R. B. & J. Kemp Haldane, vol. II (London: Kegan Paul, Trench, Trubner & Co. Ltd., 1910), 18.

96

Chapter 3

other organs through a proximal vascular system that serves the entirety of the plant. Plant will is distributed and ubiquitous as well and is more like the primordial will that Arthur Schopenhauer called the thing in itself. This whole, including both, is the world as representation, or the phenomenon. After this is taken away, there remains only the purely metaphysical, the thing-in-itself, which in the second book we shall recognize as the will.70 Schopenhauer maintains that all beings have will. Says Schopenhauer: But now, as the counterpoise to this truth, I have stressed that other truth that we are not merely the knowing subject, but that we ourselves are also among those realities or entities we require to know, that we ourselves are the thing-in-itself.71 Schopenhauer is not talking about plants, but about us. While we come from different branches of life, both plants and humans are attached by ubiquitous will. We, plants, honeybees, and humans have will and therefore, each of us, are the thing in itself, without ends, all middle. Nor are intelligence and will the same. Intelligence varies; will, according to Schopenhauer, is as ubiquitous as it is the thing in itself.72 My philosophy alone leads us out of this dilemma; in the first place it puts man’s real inner nature not in consciousness, but in the will. This will is not essentially united with consciousness, but is related to consciousness, in other words to knowledge, as substance to accident, as something illuminated to light, as the string to the sounding board; it comes into consciousness from within just as the corporeal world comes from without. Now we can grasp the indestructibility of this real kernel and true inner being that is ours, in spite of the obvious extinction of consciousness in death and its corresponding non-existence before birth. For the intellect is as fleeting and as perishable as is the brain, and is the brain’s product, or rather its activity. But the brain, like the whole

70  Ibid. 71  Ibid., 195. Emphasis in original. 72  Ibid., 291.

Flower and Honeybee Epistemology and Behavior

97

organism, is the product or phenomenon of, in short a secondary thing to, the will, and it is the will alone that is imperishable.73 We can debate whether plants are conscious or intelligent, but fundamentally, like all other creatures, plants have will. Intelligence is secondary and different from the will. Therefore, even if we believe that plants are not intelligent, are not conscious, and are bereft of other capabilities that animals appear to display, we are all willing creatures. The will is therefore a fundamental and imperishable function of existence. This means that will is always already available. Will does not perish with the dying creature because will is not incorporated into the creature. Will attaches itself to the creature and does not expire when the creature does. As Christopher Ketcham notes: We live and die but will continues. When a new being is born, will attaches itself to the individual. Death extinguishes consciousness but will continues. Schopenhauer mused that it is this will within us that gives us the experience of feeling that there is continuity beyond this impermanent life. It is will: not soul, not intellect, not consciousness that continues indefinitely like Parmenides’s being. However, Schopenhauer was, like Parmenides and being, at a loss to explain the origin of will beyond the fact that it enters consciousness.74 Foundationally, therefore, the plant is a willing creature as are all life forms. If we can have faith in this thought, we can then agree with Jean Paul Sartre that “existence comes before essence”75 Being and will are the necessity we must understand before we can consider essence which describes the existential condition of the plant which we do know is different from that of the honeybee and the human. However, as recent research is beginning to show, plants and animals have more similar capabilities and functions than once thought. Like the nervous system of animals, the plant vascular system transports information throughout the plant in the form of chemicals and electrical signals. Some information may be meaningful and valuable for the roots, but not for the leaves. Brains perform similar functions, directing the production of skin sweat when warm and adrenal gland adrenalin when threatened. 73  Ibid., 199–200. 74  Christopher Ketcham, “Schopenhauer and Buddhism: Soulless Continuity,” Journal of Animal Ethics 8, no. 1 (2018): 17. 75   Jean-Paul Sartre, “Existentialism Is a Humanism,” ed. Walter Kaufman, Existentialism from Dostoyevsky to Sartre (Whangarei, New Zealand: Pickle Partners Publishing, 2016). 210.

98

Chapter 3

Other information may be meaningful and valuable to all plant organs, such as the presence of herbivores on or near the plant. Some information is transmitted to other plants through the air or through the roots. The mutualism of some plant roots and mycorrhizae can even transmit this information through mycorrhizal networks that connect separate plants of the same or even different species. From Plotinus, Marder suggests that plant thoughts are “growth-thoughts.”76 Growth and its obverse, reduction, complete the whole of stationary motility that is central to the non-dualism of plant existentiality. The plant is both a growing and reducing thing at the same time. While the roots are searching for food, some die away because they find nothing. At the same time, the stem adds branches towards the sun and lets branches and leaves die when the sun no longer reaches them. The growth/reduction equilibrium process that encompasses the entire plant is always already towards optimality. Even the release of noxious chemicals to ward off attacking herbivores serves growththought by mitigating unwanted reduction in photosynthetic surface area. While there is an ontological difference between the distal intelligence of animals and the distributed intelligence of plants, both exhibit capabilities we can call intelligent. Fundamentally, I suggest that intelligence begins when a creature regularly produces the optimal response to stimulus over the existential temporality of the existent. We can suggest that human intelligence is more complex because it includes abstract thoughts, but fundamentally we too could not exist for long if we were not cognizant of our environment and could not make optimal decisions towards our own survival. Plant organs: stems, roots, leaves, flowers all have different functions as do animal organs. Plants can sense light, minerals, predation, and can even record whether roots they encounter are of the same species or other species as do animals when they encounter other animals. Plants are interconnected by a vascular system that permits the distribution of information both chemically and electronically like animals do. Plants can learn and remember, often this memory lasts for many years. So can animals. The vascular network in plants not only serves as a way of moving nutrients to appropriate organs for processing or storage but also for storing and distributing information. However, plants also use airborne pheromones to warn other leaves on the same plant that herbivores are attacking. Plants do not have any vascular operations that require a central processing system. Plant principal functions are organ driven, meaning that all decisions 76  Marder, Plant-Thinking: A Philosophy of Vegetal Life, 129; “What Is Plant-Thinking,” Klesis—Revue Philosophique 25 (2013).

Flower and Honeybee Epistemology and Behavior

99

are made at the organic level. The vascular system provides chemical and electrical information of what the other organs are doing or are experiencing. Therefore, plants have sensing/acting processes in each of the organs and each of these performs different services to the entire plant. These sensing/ acting processes are interconnected through the vascular system that serves, in part, as an information highway to inform other organs of decisions, threats, or other issues the organ is dealing with. I suggest that plants operate less like the central processing system of a computer and more like the internet which also has nodes (organs) that do different things and transmit that information to other nodes as is necessary and relevant. What plants do not have is an independent neurological system connected to each organ or to each organ’s sensing capabilities. It is both distributed (organs) and connected (vascular system). We might explain plants as interconnected distributed creatures without an independent nervous system. While we can suggest that the vagaries of evolution and the many millions, perhaps even billions of years of independent plant evolution from animals have evolved this differentiation, but there may be other reasons for why plants do not have a nervous system that is independent from a vascular system. Two differences become obvious: locomotion and time. Flowering plants move by growing or retarding organ growth (roots, stems, leaves, flowers, tendrils) from a fixed location. Thus, they have an anchored existence from the moment their seed sprouts. Plant movements are, in general, significantly slower than animals. Yet, they can release warning pheromones quickly within minutes or hours through the air or through the vascular transmission of information. However, we must understand that while a stem may take days to reorient itself towards the sun, the sensory cells that cause the stem to move receive and process the change in sunlight obtain information as quickly as we might when we squint from looking directly at the sun. Therefore, we cannot definitively maintain that plants necessarily ‘experience’ time any differently from animals, even though their process of movement, in most cases, involves organic change or growth that takes time. Both plants and animals have similarly functioning circadian clocks. We can therefore say that the circadian process of both plants and animals is temporally equivalent. The next question is whether the flower and honeybee faculty mutualism is intelligent beyond co-evolution that appears to have intelligently designed both to require, benefit from, and exploit each other. 3.5 Intelligence and the Flower and Honeybee Facultative Mutualism The question of whether plants are intelligent is important not only because plant and animal capabilities can be described differently, but also because we have two species, flowers and honeybees, that use different existential systems

100

Chapter 3

(vascular vs. vascular and neurological) that receive, assess, and process phenomenological input who have come together into a facultative mutualism that has lasted for at least a million years. It is important to understand that both evolution through mutation and the behavior of existents have created the existential construct called flower and honeybee facultative mutualism. The activities of both vascular and neurologically driven honeybees have co-evolved with vascular (only) driven angiosperms to both benefit from and exploit the other. We have many other examples of mutualisms in nature like the root-mycorrhizal mutualism, or the human-gut bacteria mutualism. The flower and honeybee facultative mutualism is important to study because we have so much information about both flowers and honeybees. However, what is significant as well, is that there are two distinctly different life forms who come from fundamentally different branches of life but who have found a way to exist in a community where difference is both an advantage and a challenge to maintain. The advantage to the flower and honeybee mutualism is that dependence on each other are for asymmetrical needs: food for honeybees; facilitation of the sex act for flowers. However, while their existential needs do not compete, they fulfill their respective requirements at the same location and that is the flower that is produced by the flowering plant. They both have had to develop means for informing the other of availability and presence. This they have accomplished through different means, including flower communication through advertisement of color, shape, smell, and even tactile feel. The honeybee informs the flower of her presence by landing and foraging, which in some instances, encourages the flower to loosen pollen or to move pollen carrying stamens closer to the pollinator. All this is done with a plant that has no brain and an insect who does. It is difficult not to see that a kind of intelligence emerges from the flower and honeybee mutualism that combines the talents of both but exceeds both in the gestalt of the mutualism. It combines the distributed intelligence of the flower with the distal intelligence of the honeybee, that while asymmetrical, produces a form of intelligence that is both an inward seeking and outward seeking unity that is always already permanent for the two species, but impermanent at the same time; because, like other forms of intelligence, it is always prepared for the exigencies and contingencies that the world will bring to the mutualism and the partners in the mutualism bring to it. Nor is this gestalt of intelligence something that we must confine to this particular mutualism—what about all the others who live in the meadow? Optimization occurs throughout the meadow and this contributes not only to the flower and honeybee facultative mutualism but to all existents who call the meadow home. This suggests that it is time for science to reconsider what it means to think or cognize in terms of what nature provides its creatures

Flower and Honeybee Epistemology and Behavior

101

in the form of capabilities and their abilities to react to or with creatures in the meadow that may be totally different than themselves. This study will not offer such a reconsideration, but does provide some evidence that will help others to think differently about how do diverse species of flowering plants and an animal create such a durable construct as a facultative mutualism that functions as a social group where each member is existentially irreplaceable to the other. The flower can sense whether an insect is an herbivore like a caterpillar or a honeybee who is a pollinator. The caterpillar it takes actions against, and at the same time encourages the honeybee to enter her flower to consummate their facultative mutualism relationship. Therefore, it is important to consider the flower under her terms without resorting to anthropomorphic prohibitions against such general notions of thinking/cognition, consciousness, intelligence, or even ethics as it can be considered in other species. What we require are better definitions of terms such as thinking/cognition, consciousness, intelligence, ethics, judgment, and the passions to consider how other creatures exist and explain how they perform similar existential functions using quite different ontological and epistemological means. However we move down the path towards gaining better understanding and definition to basic concepts like thinking/cognition, consciousness, intelligence, ethics, judgment, and passions, I remind the reader of the initial assertion in this study that most life that use optimal processes to make and act upon decisions appear to be oriented towards the good. Angiosperms, in general, make optimal decisions from stimuli they receive. They do not process this stimulus through a distal brain (or even moredistributed brain like the octopus) but through a vascular system that connects different processes in different organs that are tasked with controlling different existential requirements of the plant. Fundamentally, plants and animals move towards both existential and intergenerational goodness by reacting optimally to stimuli. Therefore, it is important for science to recognize that optimal decision-making is a shared trait of both flowers and honeybees and is also instrumental to the success and durability of their facultative mutualism. We have established that flowering plants can judge and reason. There­fore, we can continue to consider the flower and honeybee facultative mutualism to discover the origin of morality in nature and in a social group that is otherwise than human. We will have an easier task in explaining the honeybee’s capabilities to reason and judge because they are animals and have a central nervous system with a sophisticated brain. Even if this task is simpler than for the flowering plant, we still must consider how honeybees do what they do and later in this study must broach the inevitable paradox of how such totally

102

Chapter 3

different life forms have come together to form a social group that has lasted at least one million years. As the equal partner in the flower and honeybee facultative mutualism, and, the more mobile partner, it is important to assess how honeybees use their considerable mental and physical skills to not only facilitate their mutualism but also achieve intergenerational goodness through the facultative mutualism and outside of it. Research into honeybee waggle dances began in earnest in the nineteen twenties with research done by Karl von Frisch who earned a Nobel prize in biology for his discoveries that the tiny honeybee has created a kind of language to effectively communicate valuable information to others in the hive. His foundational research and that from many others, not only suggests that symbolic language is not exclusively a human capability, but also that it evolved separately and distinctly in the insect who has significantly fewer neurons than even the most primitive mammals. Language and other skills that science are uncovering from studies of honeybees provide us with solid evidence that honeybees spend a considerable amount of cognitive and behavioral energy towards optimizing their own decisions while they navigate ecological conditions that regularly change over time. Their commitment to intergenerational goodness also, like mammals, involves multiple generations as existence in the hive at any one time. Shared nursery duties, foraging, and other behaviors produce a hive society for which we can consider whether any of these actions can be theorized as ethical practices and to determine whether we can formulate moral theory for honeybee hive society. 4

Honeybee Epistemology and Behavior

4.1 Basic Honeybee Existence Honeybees live in hives, often built in hollowed out trees or branches that provide insulation from the heat of summer and cold of winter and to some extent protect against honey predators like bears. The entrance to the hive is small which provides a small surface area to protect against incursions from raiding foragers from other hives and predators like some wasps that prey on honeybees. The opening to the hive in our farmhouse wall (when I was a child was covered by aluminum siding) was about the size of a ten-penny nail head which is unusually small. The hive has a dance floor near the entrance of the hive where foragers communicate where flowers are, and where nearby hive bees can perform trophallaxis, the process hive bees use to offload from foraging bees both the nectar they have gathered in their crop and the pollen

Flower and Honeybee Epistemology and Behavior

103

that covers their hind legs. Deeper in the hive is the nursery and a chamber for the queen. Honeycomb is continually constructed by hive worker bees from resources the foragers bring to the hive. In addition to pollen and nectar, honeybees forage for tree gum which is used as a kind of glue to build hive structure. Foragers also bring water used to cool the hive in summer. Beating wings evaporate the water that cools the hive and the sensitive nursery. The queen may live five years. Spring and summer-born workers live sixty to eighty days, but those who are born in the fall live four to six months because they winter-over in the hive. The few males live until they can successfully mate with the queen on the mating flight and then they die. Those males who do not mate successfully at the end of the mating season will be ejected from the hive and will die. Workers are sterile females. As immature adults they first tend to a series of different hive duties and then most become foragers, and some become hive guards. Workers are the only caste in the hive to change jobs during their lifetime. (Much more about hives can be discovered in more comprehensive texts noted in this footnote.)77 4.2 Honeybee Foraging Behavior Ants and honeybees are both eusocial insects. However, for most species of ants, their foragers do not fly. Ant foragers must also travel distances (not as far as honeybees in most cases) to obtain necessary resources for the hive. As most ants are ground-bound (though many climb), they release pheromones to form trails to alert others to these resources. As more and more foraging ants travel the trail, the scent becomes stronger which encourages more foragers to follow the trail to obtain resources and return them to the nest. If you spill some sugar on the counter, the first ant that finds it will produce a pheromone trail back to the nest. In a short time, a whole procession of ants travels from sugar spill to the nest and back again. However, honeybees have evolved to fly to potential foraging resources. A pheromone trail would dissipate too quickly in the air and a ground pheromone trail would be impractical. They must use their memory and a version of language called the waggle dance to recall and 77  See: Thomas D. Seeley, The Wisdom of the Hive (Cambridge, Ma.: Harvard University Press, 1995); Thomas D. Seeley, Honeybee Democracy (Princeton, NJ: Princeton University Press, 2010); James L. Gould, “Honey Bee Cognition,” Cognition 37, no. 1–2 (1990); Randolf Menzel, “The Honeybee as a Model for Understanding the Basis of Cognition,” Nature Reviews Neuroscience 13, no. November (2012); Randolf Menzel and Alison Mercer, Neurobiology and Behavior of Honeybees (Berlin Germany, New York: Springer-Verlag, 1987); Thomas D. Seeley and Royce A. Levien, “A Colony of Mind,” Sciences 27, no. 4 (1987); James L. Gould and Carol Grant Gould, The Honey Bee, ed. Carol Grant Gould (New York: Scientific American Library: Distributed by W. H. Freeman, 1988).

104

Chapter 3

communicate the location of valuable resources: flowers. Randolph Menzel notes some of the challenges associated with honeybees and their dependency on flowers: Flowers are unreliable, widely distributed food sources, normally offering minute rewards. Flowers of the same kind tend to bloom in close proximity, because plants of the same species growing in patches often bloom simultaneously, or a single plant has many blossoms. Thus, a patch of flowers of the same kind has a location in space and exists for some time, perhaps longer than the lifespan of an insect pollinator. A typical habitat consists of several to many patches of flowers, some of the same species, some of others; pollinators must choose between them.78 The honeybee can fly ten kilometers or more from the hive to search for resources.79 Flowers may bloom in the meadow where the hive is in one week and in a far-off field the next week. The honeybee forager prefers certain colors of flowers they discern in both the color spectrum and ultraviolet light but will investigate species that the forager has not seen before. Rather than use pheromones as a guide to return to resources, the honeybee uses its memory to record distance from the hive, direction from the hive, and a triangulation to the position of the sun in the sky.80 The honeybee forager can remember features of the ecology such as meadows, streams, and trees to help guide it to and from the foraging site. Menzel explains: Insects’ learning capacity and richness of memories are usually underestimated, but studies of learning and memory in honeybees (under both natural and laboratory conditions) demonstrate that learning is fast, and comprises various levels of cognitive processing, such as generalization, categorization, concept formation, configuration, and contextdependency (Menzel & Giurfa 2001). Memory is rich, highly dynamic, and long-lasting.81 78  Randolph Menzel, “Behavioral and Neural Mechanisms of Learning and Memory as Determinants of Flower Constancy,” in Cognitive Ecology of Pollination: Animal Behaviour and Floral Evolution, ed. Lars Chittka and James D. Thomson (Cambridge, UK: Cambridge University Press, 2001), 21. 79  M. Beekman and F. L. W. Ratnieks, “Long‐Range Foraging by the Honey‐Bee, Apis Mellifera L.,” Functional Ecology 14, no. 4 (2000): 490. 80  Seeley, The Wisdom of the Hive, 37. 81  Menzel, “Behavioral and Neural Mechanisms of Learning and Memory as Determinants of Flower Constancy,” 21.

Flower and Honeybee Epistemology and Behavior

105

However, honeybee foragers need to learn and learn quickly. They forage only for the last twenty days or so of their lives. Menzel explains that new foragers perform a learning flight which produces “landscape memory” of the terrain flown with the hive as the geographic center.82 Foragers have a strong spatial memory that they retain for their foraging life. In addition to foraging in the discovered landscape, foragers retain complex information about resources in the landscape. Menzel explains: The locations of rewarding sites are characterized by their particular features and are memorized accordingly. Bees learn the local features (signals, localization relative to landmarks, reward conditions) of two to four feeding sites, and behave accordingly: they choose the correct color at the correct time and place (Menzel et al. 1999; Lehrer 1999) or the correct color pattern at the correct step in a sequence (Collett 1992); they choose the correct odor at a particular time (Koltermann 1971); they indicate the correct direction and distance to one of two feeding sites according to time of the day (von Frisch 1965, table 37); and, they match the frequency of their visits to the reward quantities of at least four feeding sites (Greggers & Menzel 1993).83 Therefore, foragers can assess the quality of foraging resources not only by location, but by color, smell, sugar content, and even predator or competitor concentration. Whether they return to these resources depends upon their quality and the contemporaneous need of the hive. As Menzel notes, “Furthermore, bees have the capacity to switch their motivation according to recent experience and activate remote memory according to the motivational change.”84 For example, the honeybee assesses that the rewarding patch is overrun by competitors or predators and does not return. Delays in trophallaxis in the hive can provide foragers negative feedback on either the quality of what they have brought or the need for that resource at this time.85 Flowers and honeybees have co-evolved to profit from and exploit each other. The flower cannot move from its location to better position itself for 82  Ibid., 23. 83  Ibid., 24. 84  Ibid. 85  See: Walter M. Farina and Josue A. Núñez, “Trophallaxis in the Honeybee, Apis Mellifera (L.) as Related to the Profitability of Food Sources,” Animal Behaviour 42, no. 3 (1991); Andrea A. Tezze and Walter M. Farina, “Trophallaxis in the Honeybee, Apis Mellifera: The Interaction between Viscosity and Sucrose Concentration of the Transferred Solution,” ibid., 57, no. 6 (1999).

106

Chapter 3

its preferred pollinators. Flowers cannot speak which means they cannot hail pollinators. However, they do advertise their availability by shape, color, and scent. If the honeybee sees and smells a desirable flower, she must also assess the quality of nectar and the presence of competitors or predators at the site. Some flowers do not replenish nectar which may discourage the forager from returning to the site. The presence of too many competitors may also stress nectar replenishment, making the flower undesirable. However, flowers with depleted nectar and those that only provide pollen and not nectar may entice the honeybee to return if the hive has signaled to the worker that pollen is needed now. Once the flower is pollinated it may continue to look desirable, but likely the flower has shut down its nectar production and will soon desiccate in order to complete the seed fertilization process. As Menzel says, the nectar reward for the honeybee forager from each flower is small and inadequate to satisfy the honeybee.86 Therefore, the honeybee hive will employ forty thousand or so foragers to find and return to promising foraging sites. If pheromones cannot be used to communicate to other bees promising foraging sites, an alternative means of communicating valuable resources is necessary. Honeybees have developed an elaborate waggle dance to communicate the location and quality of resources, including locations where honeybee predators may be present. The different tremble dance alerts hive bees that a forager requires trophallaxis. There is also a bump-dance that provides negative feedback to waggle dancers about what they are dancing. Finally, there is a house-hunting waggle dance performed by scout honeybees during the swarm. As scout honeybees fan out, they assess the quality of potential locations and return to the swarm to dance potential sites they have found. Much has been learned about what and how much information is communicated by honeybees in their dances. 4.3 The Honeybee Foraging Waggle, Tremble, and Bump Dances The origin of the honeybee was in Eurasia, but honeybees have migrated to or been exported all over the world. Different species of honeybee have distinctly different foraging dances. However, one recent study reared two different species, one European, and one Asian in one hive, and discovered that the two species can understand the waggle dance of the hive.87 The waggle dance is most likely a genetic adaptation but also has a learning 86  Menzel, “Behavioral and Neural Mechanisms of Learning and Memory as Determinants of Flower Constancy,” 21. 87  Songkun Su et al., “East Learns from West: Asiatic Honeybees Can Understand Dance Language of European Honeybees,” PLoS One 3, no. 6 (2008): 1.

Flower and Honeybee Epistemology and Behavior

107

component.88 This implies that the foraging waggle dance is both biological and a learned behavior. Foraging workers are mature non-reproductive females. Foragers leave the hive to search for resources required by the hive. As they flit from flower to flower, they observe and remember sugar content, location, density of competitor foragers, and danger, such as predators in the area. Foragers return to the hive laden with resources harvested during the journey. When they land in the hive some report to a dance floor and perform a waggle dance to another worker in the hive. The waggle dance communicates a triangulation vector with three points, the hive, the location of the sun, and the angle of deviation from both which is the direction to the food source.89 The dance also communicates distance through the number of times the dancer turns around in the dance. This suggests that honeybees can record and remember standard units of measure which they translate into distance.90 Surprisingly, this information transfer likely only requires 7.5 bits of information.91 Compact and efficient communication is necessary because honeybees only have a million neurons compared with the eighty billion or so of humans.92 In comparison, the navigation computer that directed the spacecraft that landed men on the moon and returned them to the earth had capabilities to store 4k or four thousand bits of information. Honeybees can navigate to new locations using far fewer bits of information. Brian R. Johnson and James C. Nieh explain the mechanics. There can be only one unemployed forager who watches the waggle dance which takes about five seconds to perform, “If the unemployed bee has already visited the source the waggle dancer is advertising, then it is reactivated and returns to it 88  Honeybee foragers are quick learners to be sure. They perform multiple jobs during their short lifetimes. Even with tremendous neuroplasticity, the limitation of one hundred thousand neurons suggests that learning a waggle dance without some innate ability to do would be a monumental task for a job that lasts twenty days. The chapter on epigenetic rules in this study may have bearing on how the foraging and other waggle dances emerged and became part of the behavioral repertoire of the worker. However, this idea has not been studied in any detail. I suggest that the Asian and European dance variants are not so different that the imported worker cannot learn how to interpret the other than innately constituted dance. For example, while Canadian French and the French spoken in France are different, members of each can grasp most of what the other is saying without significant new learning. 89  Seeley, The Wisdom of the Hive, 37. 90  Roger Schürch, Margaret J. Couvillon, and Madeleine Beekman, “Editorial: Ballroom Biology: Recent Insights into Honey Bee Waggle Dance Communications,” Frontiers in Ecology and Evolution 2016, no. February (2016): 4. 91  Ibid. 92  Giurfa, “Cognition with Few Neurons: Higher-Order Learning in Insects,” 285.

108

Chapter 3

(Seeley 1995). If the novice has not been to the source before, then it learns the location of the food source and attempts to locate it.”93 The foraging dance follower not only obtains the location of the resource, but also the smell of the resource on the dancer.94 The following forager can also sense (hear) the vibrations of the dancer’s wings. The higher the pitch and the more energetic the dance researchers believe may communicate higher quality resources in terms of sugar concentration, etc.95 The tremble dance of the honeybee is similar to the waggle dance but is used by a forager when the foraging bee perceives a long delay in unloading its nectar. The tremble dance’s purpose is to recruit receiver bees to offload what the forager has brought to the hive.96 She may also spread the scent of her resource load during the tremble dance which may help recruit receiver bees. An analogy to the tremble dance is the truck driver who grouses to the dispatcher at the warehouse that he is losing money because the delivery cannot be offloaded quickly enough because an inadequate number of warehouse workers showed up for work today. The dispatcher has had to optimize the truck unloading process to put perishable loads first and this poor truck driver’s load of nails last. Thus, the tremble dance likely helps the honeybee hive optimize its workload by performing trophallaxis quickly on higher quality resources, leaving lesser quality resources to be offloaded when receiver bees have time to do so. There could be multiple reasons for delays in trophallaxis. The first is that there may be inadequate numbers of receiver bees that can be corrected by maturing workers more quickly to participate in offloading.97 Second, the scent of the resources that the offloading bee releases may suggest to the receiver bees that the quality of the resource is low and that receivers 93  B  rian R. Johnson and James C. Nieh, “Modeling the Adaptive Role of Negative Signaling in Honey Bee Intraspecific Competition,” Journal of Insect Behavior 23, no. 6 (2010): 463. 94  Martin Hammer and Randolf Menzel, “Learning and Memory in the Honeybee,” Journal of Neuroscience 15, no. 3 (1995): 1617. 95  Seeley, The Wisdom of the Hive, 39. See also: Wolfgang H. Kirchner, “Vibrational Signals in the Tremble Dance of the Honeybee, Apis Mellifera,” Behavioral Ecology and Sociobiology 33, no. 3 (1993): 169. 96  Johnson and Nieh, “Modeling the Adaptive Role of Negative Signaling in Honey Bee Intraspecific Competition,” 463. 97  For the emerging science on pheromonal maturation processes and honeybee neuroplasticity see: Isabelle Leoncini et al., “Regulation of Behavioral Maturation by a Primer Pheromone Produced by Adult Worker Honey Bees,” Proceedings of the National Academy of Sciences of the United States of America 101, no. 50 (2004); Andrés Arenas et al., “Behavioral and Neural Plasticity Caused by Early Social Experiences: The Case of the Honeybee,” Frontiers In Physiology 4, no. Article 41 (2013).

Flower and Honeybee Epistemology and Behavior

109

would profit more from offloading foragers who have brought higher quality resources. Research is just beginning to suggest that foragers may learn from delays in trophallaxis that they should forage for higher quality nectar or pollen resources, or other resources like water or tree gum that the hive needs at this moment.98 Trophallaxis may also involve some pheromonal or hormonal exchange that provides more direct information to the foraging and offloading honeybee than intuition obtained from the delay itself.99 More research needs to be conducted before we can better understand the important information that trophallaxis provides both the offloading and receiving honeybee. There is also a bump or stop dance.100 If a following forager senses there is danger at the location that the dancing bee is dancing, the following forager can bump the dancer, and this encourages the dancer to stop dancing. Hive honeybees can also bump the dancer if the assessment of resource quality does not meet current hive needs. This negative feedback must be considered in context of the foraging waggle dance. James C. Nieh notes that there has been significant research on positive feedback in honeybee colonies such as the waggle dance, but not much on negative feedback.101 He says about feedback loops, “In superorganisms, individuals within a social group act as cooperative vehicles for gene propagation, and their actions often rely on a network of self-organizing behaviors, rather than centralized control.”102 Nieh’s experiments show that, “[c]onspecific attacks at food sources lead to the production of stop signals.”103 Nieh found that when workers experience aggressive food competition at a food source, they tended to bump dance more in the hive. If they smelled the same scent of the food source on the waggle dancer where 98  E.g., “Recruiting efficiency, i.e. the number of recruited foragers, depends on the profitability of the food source in terms of the concentration of the sugar solution (von Frisch 1965) as well as the flow rate at a constant concentration (Nfifiez 1971)” Farina and Núñez, “Trophallaxis in the Honeybee, Apis Mellifera (L.) as Related to the Profitability of Food Sources,” 391. 99  E.g., “Here, we report on the identification of a substance produced by adult forager honey bees, ethyl oleate, that acts as a chemical inhibitory factor to delay age at onset of foraging. Ethyl oleate is synthesized de novo and is present in highest concentrations in the bee’s crop. These results suggest that worker behavioral maturation is modulated via trophallaxis, a form of food exchange that also serves as a prominent communication channel in insect societies” Leoncini et al., “Regulation of Behavioral Maturation by a Primer Pheromone Produced by Adult Worker Honey Bees,” 17559. 100  Parry M. Kietzman and P. Kirk Visscher, “The Anti-Waggle Dance: Use of the Stop Signal as Negative Feedback,” Frontiers in Ecology and Evolution 3, no. 14 (2015): 1. 101  James C. Nieh, “A Negative Feedback Signal That Is Triggered by Peril Curbs Honey Bee Recruitment,” Current Biology 20, no. 4 (2010): 310. 102  Ibid. 103  Ibid.

110

Chapter 3

they experienced conspecific aggression they were more likely to bump dance that dancer.104 However, if the non-dancing bee had been the aggressor at the food source, she would not likely bump dance the waggle dancer who danced about the same location where she was the aggressor. Similarly, if the bee had been attacked by a biting ambush predator at the foraging site or had been exposed to a honeybee alarm pheromone left by other foragers, she would also more likely bump dance the waggle dancer.105 What Nieh observed was: The stop signal is a brief vibrational signal lasting 150 ms. at around 380 Hz. It is frequently delivered by a sender butting her head into a recipient, although the sender may also climb on top of the receiver … I will use the term “stop signal” because experiments show that this signal can cause waggle dancers to stop dancing and leave the nest. Playbacks of the stop signal (artificial vibrations of the comb) reduced waggle dance durations by 59% and recruitment by 60%.106 From other researcher observations, Nieh points out that crowded food sources or longer waits at such food sources will also elicit more stop signals.107 Nieh’s research maintains, “My preliminary observations suggested that conspecific fighting over rich food increased stop signal production. Such fighting could occur in the context of nest robbing but is probably not common for floral resources … Predator attacks may be a natural trigger for stop signals.”108 However, Nieh notes that negative feedback does not often work on the first try and that the feedback loop may require many stop signals before waggle dancers stop dancing the imperiled resource.109 Nieh surmises, “Receivers requiring multiple stop signals are, in effect, integrating negative feedback from multiple information sources, and the colony-wide effect of recruitment cessation thus arises as an emergent property of multiple, independent actors signaling and receiving information about food patch conditions.”110 Nieh found that pinching the dancer’s leg was more effective than the release of the danger pheromone, “Like a natural attack, pinching also sharply decreased 104  Ibid. 105  Ibid. 106  Ibid. 107  Ibid. See also: Johnson and Nieh, “Modeling the Adaptive Role of Negative Signaling in Honey Bee Intraspecific Competition.” 108   Nieh, “A Negative Feedback Signal That Is Triggered by Peril Curbs Honey Bee Recruitment,” 311. 109  Ibid., 314. 110  Ibid., 314–15.

Flower and Honeybee Epistemology and Behavior

111

the number of waggle dance circuits. Alarm pheromone did not affect waggle dancing production, although there was a 4-fold decrease in the average number of waggle circuits. Thus, more dangerous attack stimuli appear to elicit stronger responses (more stop signals, fewer waggle dance circuits) as compared to alarm pheromone alone, which involved no physical contact.”111 Nieh concludes: In summary, a forager’s experience at a patch and her foraging motivation influence her decision to recruit. For example, honey bees perform fewer waggle runs after returning from dangerous as compared to safe flowers. However, one individual’s decision to cease recruiting does not stop recruitment by other waggle dancers. By sending stop signals, she can inform foragers visiting the same location of adverse foraging conditions and provide negative feedback to counteract waggle dancing by others. Thus, collective actions of the superorganism arise from the positive and negative feedback of multiple actors, with negative feedback cycles providing greater precision and speed for labor reallocation.112 Conditions in the world are always changing. Flower patches mature and desiccate. Other honeybees or insects might over-forage a previously rich site, and during periods of drought or the approach of winter, the hive might need to accept lesser quality resources to make enough honey to help it survive the winter. Dancing and feedback to dancers are instrumental in the hive optimizing its foraging behavior. These are the basic dances that help both foragers and receiver bees optimize resource gathering and processing for the hive. What we have not yet explored is whether there are honeybee personality differences that also contribute to how honeybees choose which decision to make, e.g., forage where there has been predation or not. Are there, for example, more aggressive foragers or hive guardians who may make more risky decisions because of their personal preferences? 4.4 Honeybee Personality Honeybee workers are not all identical twins. For the most part, unless a new queen takes over, the hive has a common mother, but up to ten males have mated with her so there are many half-sisters. We see different personalities in other animals, some driven by genetics (larger antler racks in some deer), 111  Ibid., 314. 112  Ibid., 315.

112

Chapter 3

others through learned behavior. Honeybees from even the same hive show differences. We cannot say that honeybee foragers are all perfect foragers. They make mistakes. Keith D. Waddington suggests that individual workers, given the same conditions, can arrive at different decisions: Although general patterns of pollinator foraging behavior have been found, variation among individual foragers has not been well studied. Individuals observed in foraging experiments often differ in their behavior even when given the same problem (e.g., choosing among flowers in the same patch; Waddington & Holden 1979). Inter-individual variation might be due to differences in experience, such that each forager has a slightly different experience in the same patch and makes decisions based on different information (see Thomson & Chittka, this volume). Sampling error could produce important differences in experience among individuals, especially if the sample of previous flower visits used to make future decisions is small. However, genetic differences among individuals may also contribute to behavioral variation.113 More recent research suggests the possibility that individual honeybees have different personalities.114 Alexander Walton & Amy L. Toth studied individual worker behavior, “[o]ur data suggest some individuals may be more likely to be highly interactive with other workers (e.g., engaging in food sharing), while other individuals are consistently less interactive.”115 They offer that the possible purpose for individuation includes, “We suggest that individual-level personality differences have the potential to contribute to colony division of labor by creating variation in individual tendencies to perform different tasks.”116 This is not unexpected because it has long been understood that some workers become hive guardians while most do not. While all scout bees in the swarm are workers, only about five percent of workers become scout bees. Also, some hive bees do not mature into foraging bees. Whether personality difference is genetic, learned, or a combination of both is not understood. However, recent 113  K  eith D. Waddington, “Subjective Evaluation and Choice Behavior by Nectar- and Pollen-Collecting Bees” in Cognitive Ecology of Pollination: Animal Behaviour and Floral Evolution, ed. Lars Chittka and James D. Thomson (Cambridge, UK: Cambridge University Press, 2001), 41–42. 114  See: Alexander Walton and Amy L. Toth, “Variation in Individual Worker Honey Bee Behavior Shows Hallmarks of Personality,” Behavioral Ecology and Sociobiology 70, no. 7 (2016). 115  Ibid., 999. 116  Ibid.

Flower and Honeybee Epistemology and Behavior

113

research into the possibility that honeybees can experience emotion suggests that personality may be partially driven by experience. Therefore, consistent risk taking or avoidance behavior in different individuals may be influenced by experience. Melissa Bateson, et al., agitated honeybee workers. They suggest that based upon the responses of shaken bees compared to non-shaken bees, that the shaken bees produced the same kind of brain chemicals that humans do when agitated, “This finding reinforces the use of cognitive bias as a measure of negative emotional states across species and suggests that honeybees could be regarded as exhibiting emotions.117 While research in emotional states of honeybees is both new and controversial, we do know that honeybees have significant capabilities for reacting to different phenomenon.118 Yet again, even in a single hive, there can be significant genetic diversity. Robert E. Page, Jr. et al., hypothesize that polyandry increases genetic diversity and may result in individual specialization by workers in the hive and may increase the capabilities of the hive to deal with different environmental exigencies.119 Margaret K. Wray, et al., suggest that hive personality plays a role in hive sustainability: The colonies in our study showed consistent behavioural differences in traits such as defensive response, foraging activity and undertaking, and several of these traits were correlated as part of a behavioural syndrome. Furthermore, some of these traits were strongly tied to colony productivity and winter survival.120 However, the origins of such differences have not been studied to any great extent. Finally, we do know that hormones and other factors play a role in the maturation of workers into foragers. Isabelle Leoncini, et al., report that if older workers are present in the hive this retards the maturation of workers into foragers. They suspect that the process of trophallaxis and the release of hormones from the mature worker play roles in the process of regulating hive 117  Melissa Bateson et al., “Agitated Honeybees Exhibit Pessimistic Cognitive Biases,” Current Biology 21, no. 12 (2011): 1070. 118  For additional discussion on evidence for honeybee emotion see: Zbigniew Lipiñski, “The Emotional Nature of the Worker Honeybee (Apis Mellifera L.),” Journal of Apicultural Science 50, no. 1 (2006). 119  Robert E. Page et al., “Effects of Worker Genotypic Diversity on Honey Bee Colony Development and Behavior (Apis Mellifera L.),” Behavioral Ecology and Sociobiology 36, no. 6 (1995): 387. 120  Margaret K. Wray, Heather R. Mattila, and Thomas D. Seeley, “Collective Personalities in Honeybee Colonies Are Linked to Colony Fitness,” Animal Behaviour 81, no. 3 (2011): 559.

114

Chapter 3

bee maturation into foragers.121 They suspect also that the pheromone ethyl oleate is transmitted from the foraging bee to the hive bee during trophallaxis and this not only serves to facilitate food exchange from forager to hive and nest worker, but also serves as a communications process for which the pheromone can be used to retard hive bee maturation.122 Research suggests that the quality of sucrose that the returning forager has to exchange also modulates the rate of trophallaxis. Andrea A. Tezze and Walter M. Farina say: Moreover, irrespective of viscosity, donor foragers may provide information about the richness of the exploited sugar solution during unloading. Thus, in the social context of the hive, in addition to the traditional communication strategies already shown and studied in bees (e.g. dance behaviour, emission of chemical signals, vibrational and sound communication), trophallactic behaviour may be an effective process by which to exchange quantitative information about the profitability of the nectar sources exploited by foragers.123 As nectar and pollen are so critical to the survival of the honeybee hive, it is not surprising that the honeybee has evolved complex processes to find and assess the quality of floral resources and developed multiple capabilities like waggle, tremble, and bump dances, and trophallaxis to communicate not only where floral resources are so others can follow, but also the quality of those resources and any dangers associated with those resources. Just as critical to hive survival is the swarm’s need to find a new location to build a hive. 4.5 House Hunting Waggle Dance The house hunting waggle dance performers are worker scouts who dance on top of the swarm that develops when a new queen is born, and the old queen leaves the nest. Some workers leave the hive with the old queen; others stay in the old hive with the new queen. Who leaves and who stays is not 121  Leoncini et al., “Regulation of Behavioral Maturation by a Primer Pheromone Produced by Adult Worker Honey Bees,” 17559. For a theoretical perspective on how this might work see: Gro Vang Amdam and Stig W. Omholt, “The Hive Bee to Forager Transition in Honeybee Colonies: The Double Repressor Hypothesis,” Journal of Theoretical Biology 223, no. 4 (2003). 122  Leoncini et al., “Regulation of Behavioral Maturation by a Primer Pheromone Produced by Adult Worker Honey Bees,” 17563. 123  Farina and Núñez, “Trophallaxis in the Honeybee, Apis Mellifera (L.) as Related to the Profitability of Food Sources.”; Tezze and Farina, “Trophallaxis in the Honeybee, Apis Mellifera: The Interaction between Viscosity and Sucrose Concentration of the Transferred Solution,” 1325.

Flower and Honeybee Epistemology and Behavior

115

yet well understood.124 The house hunting waggle dance relies on what is believed to be a consensus approach.125 As scouts pan out, they find locations that are suitable and not. Scouts report back to the swarm and dance the quality of the site they have found. Researchers suggest that the more energetic the dance, the better the location, according to the reasoning of the dancing scout.126 Other scouts then fly to that location and return to dance their opinion about the location. It has been suggested that over time consensus builds because the better sites are danced enthusiastically, and others are not danced at all. Eventually there is agreement and the swarm flies in mass to the new location.127 What has been observed, but is not fully understood how it is triggered, that the swarm produces a piping signal which may inform the swarm that consensus has been reached. Whether this is related to an increase in the temperature of the swarm due to an increase in energetic behavior or some other reason is just now being studied.128 The house-hunting process is critical to the swarm’s survival. While the workers, and males who leave the hive have gorged on honey (even while they have starved the queen so that she is light enough to fly), they must not only build a new hive but also create new stores of honey for the winter. Most swarms occur early in the foraging season to give the swarm time to build a proper hive and to create adequate stores of honey to winter-over. The house-hunting process requires both searching scouts and swarming bees who do not leave the swarm. The scouts look for home sites that meet certain qualification as to size and protection from the elements. Rather than rely on a single scout to find the optimal location, the many scouts fan out initially and discover what they can find. Each discovered potential location may or may not be adequate for hive needs. This is why, when a site is danced, other scout observers may or may not visit the site based upon the information 124  See: Wayne M. Getz, Dorothea Brückner, and Thomas R. Parisian, “Kin Structure and the Swarming Behavior of the Honey Bee Apis Mellifera,” Behavioral Ecology and Sociobiology 10, no. 4 (1982); M. Woyciechowski and K. Kuszewska, “Swarming Generates Rebel Workers in Honeybees,” Current Biology 22, no. 8 (2012). 125  See: Nigel R. Franks et al., “Information Flow, Opinion Polling and Collective Intelligence in House-Hunting Social Insects,” Philosophical Transactions: Biological Sciences 357, no. 1427 (2002): 1569. 126  Ibid. 127  Ibid. 128  See: Thomas Schlegel, P. Kirk Visscher, and Thomas D. Seeley, “Beeping and Piping: Characterization of Two Mechano-Acoustic Signals Used by Honey Bees in Swarming,” Naturwissenschaften 99, no. 12 (2012); P. Kirk Visscher and Thomas D. Seeley, “Coordinating a Group Departure: Who Produces the Piping Signals on Honeybee Swarms?,” Behavioral Ecology & Sociobiology 61, no. 10 (2007).

116

Chapter 3

learned from the dancing forager, including the dancer’s enthusiasm for the site. As scouts return from visited sites, they will dance or not dance the visited site. Soon, fewer and fewer sites are danced. Both the scout and swarm bees appear to participate in the consensus decision to fly to a final chosen site to begin the construction of a new hive. The honeybee uses the combined capabilities of many individuals to provide adequate information to make the important decision of choosing a new hive location. The swarm appears (we do not yet know how) to arrive at a consensus. We do not know whether an intellectual, pheromonal, sensory, or behavioral trigger (or a combination) completes the consensus decision, but we do know that when consensus is reached, the swarm flies off in mass to the new location. As with the speculation that individual workers have individual personalities, recent research suggests that different hives and swarms exhibit different ‘personalities’.129 Margaret K. Wray, et al., found that while different swarms did not significantly vary the time it took to search for a new hive location, swarms did differ in the number of waggle dances performed.130 The implication is that differences in thoroughness and consensus building between swarms may play a role in long-term hive sustainability. However, this has not yet been tested. We can compare house-hunting with foraging waggle dance processes. The house-hunting waggle dance requires the participation of the entire swarm. The foraging waggle dance and bump dances are local, involving the dancer and an observer. The importance of the house-hunting process is critical to the entire swarm. Therefore, the optimal process of consensus building requires multiple and repeated inputs in order to make an optimal decision. On the other hand, there are thousands of foraging journeys in process every day and each one only represents a small contribution to the hive. Even so, the process is optimal because the foraging waggle dance provides valuable information to an unemployed bee and the dancing bee as to where to forage next. Also, research suggests that the tremble dance may inform the dancing worker that either hive needs have changed, or that the resources the dancing worker has foraged are inadequate to meet hive needs. The bump dance can inform that there is danger at the site being danced. 129  See: Wray, Mattila, and Seeley, “Collective Personalities in Honeybee Colonies Are Linked to Colony Fitness.”; Margaret K. Wray and Thomas D. Seeley, “Consistent Personality Differences in House-Hunting Behavior but Not Decision Speed in Swarms of Honey Bees (Apis Mellifera),” Behavioral Ecology and Sociobiology 65, no. June (2011). 130  “Consistent Personality Differences in House-Hunting Behavior but Not Decision Speed in Swarms of Honey Bees (Apis Mellifera),” 2061.

Flower and Honeybee Epistemology and Behavior

117

Flowers and honeybees have co-evolved ontologically into their facultative mutualism. Both use optimal decision-making processes not only for their own personal (and hive) existential purposes but also to sustain their mutuality. Both flower and honeybee epistemology and behavior show signs of consciousness and intelligence, and perhaps emotion plays a role in honeybee existence. Consciousness in both flowers and honeybees is explored next. 5

Consciousness in Flowers and Honeybees

The question of consciousness in either species associated with the flower and honeybee facultative mutualism requires additional discussion. Chamovitz makes a case for plant awareness and does so through Tulving’s first level of consciousness: procedural. According to Tulving, procedural: “[i]s concerned with how things are done—with the acquisition, retention, and utilization of perceptual, cognitive, and motor skills.”131 Plants certainly perceive, and when stimulus is received, processes (chemical, electrical, pheromonal) are triggered to react accordingly and optimally. Stimulus: leaf chomp; response: release noxious chemicals. Others likely would not call this consciousness, but as we will soon see, awareness might involve cognition (without neurons) that may not be called conscious but is pertinent to how plants navigate their situatedness in the world. The second Tulving level of consciousness, or semantic memory, “[h]as to do with the symbolically representable knowledge that organisms possess about the world.”132 We know that honeybees transmit abstract information during the waggle dance, including direction and distance from the hive. Honeybees can translate distance into units of measure and triangulate that with known locations: origin and destination. Therefore, we can say that honeybees exhibit the second level of Tulving’s consciousness or semantic memory. We know that flowers have evolved to attract honeybees through color, shape, and scent. While these are purely ontological manifestations, how did the plant, who cannot see the same way the honeybee can, produce an artifice that attracts honeybees who can? Certainly, the flower that is the best fit for the honeybee is the one that has a better chance of reproducing and thus increasing the number of chromosomes that produce more flower shapes, colors, and scents that honeybees prefer. We can stop there, but there is also the nagging question how extant plants might have contributed to this modification. It is 131  Tulving, “Memory and Consciousness,” 2. 132  Ibid.

118

Chapter 3

possible that some flowers have genes that can produce different hues. The Toadflax flower, for example, can produce an asymmetrical and a symmetrical variation through epigenetic processes.133 Plant learning and the passing down of this knowledge through epigenetics is certainly imaginable as a method to provide the best possible advertisement for pollinators. Finally, episodic memory, the third level of consciousness, according to Tulving is, “Episodic memory mediates the remembering of personally experienced events.”134 We know that honeybees can remember flower patches and discern whether they are valuable or not by the quality of their resource and whether the last time they visited there were competitors or pre­dators about. Through the bump dance they can relay problems with the site to workers who dance this location. Flowering plants record the duration and time of blue and red light which helps them regulate existential processes. It is also known that plants can remember certain events for many years and can use this knowledge to react accordingly when similar stimuli occur in the future. Recent Andean research this study has cited shows that some flowers in areas of infrequent and uncertain pollinator visitations remember the time of recent pollinator visitation and can extend fresh stamens at the same time the next day to optimize their possibility of releasing pollen to the (hopefully) returning pollinator. Can we not say that this is episodic memory? Both flowers and honeybees (and humans) use bio-electro-chemical processes to accomplish what they do. This includes the recording of consequences of events and how the individual acted during that event for their recall later to serve useful purposes in the future when similar events occur. The honeybee uses her memory to optimize her foraging capability. Plant studies have shown that plants can use memory of events in ways that can help combat infestation, and even display pollen to optimize pollination—but this is accomplished without any memory neurons. Therefore, we must confront the issue that flowers do not possess the central nervous system and neurological structure that underpins most consciousness research. I suggest that artificial intelligence research has had to confront the same issue and has begun to look at problems such as consciousness through a different lens. N. Katherine Hayles considers what she calls “cognitive nonconsciousness” that may help to explain just how plants can demonstrate Tulving’s different memory constructs without having a mind to

133  Adrian Bird, “Perceptions of Epigenetics,” Nature 477, no. 24 (2007): 396–97. 134  Tulving, “Memory and Consciousness,” 2. Emphasis in original.

Flower and Honeybee Epistemology and Behavior

119

do so.135 Through Stanislaw Lem, Hayles explores how complex problems can be solved without language: Reasoning that translating tasks into formal languages may be unnecessary for solving complex problems, Lem proposes a form of evolutionary computation programmed in natural media, an ‘information farm’ in which systems could successfully perform cognitive modeling functions without consciousness.136 Bees translate complex problems without verbal language but produce visual, sound, smell, and tactile representations in their dances (symbols) that serve as language other foragers can understand. Therefore, we can say that honeybees have translated some tasks into formal language to solve some of their foraging problems. Plants also communicate nonverbally to their pollinators through complex representations of availability including shape, color, and scent. However, the advertisement is similar to a static (non-electronic) roadside billboard—it says the same thing for a short period and generally the plant cannot modify the message when conditions change (though there are some flowers that change color when they become pollinated). Rather, frost will end the message as will pollination, but if insect predators should invade the flower, she cannot warn the pollinators of their presence. Plants also seem to solve complex problems without translating them into language. Let us assume for the moment that plants can do this without consciousness as defined by Tulving or anyone else. How could this work? Hayles suggests there are several requirements that are necessary to produce what she calls the artificial life mantra, “From simple rules to complex patterns or behaviors.”137 “First, these systems operate within evolutionary dynamics, that is, they are subjected to fitness criteria that select certain states out of the diverse range available.”138 The plant has evolved capabilities, so its range of states is limited to the affordances and constraints that have molded the plant over its evolutionary history. Plants are aware, meaning that they are always already in the world and ready to receive stimuli. When stimuli are received, there likely will be multiple options the plant will have to choose from, the simplest being do nothing or do something. For example, the wind 135  Hayles, “Cognition Everywhere: The Rise of the Cognitive Nonconscious and the Costs of Consciousness,” 200. 136  Ibid. 137  Ibid., 201. 138  Ibid.

120

Chapter 3

sways branches of trees to produce occasional shade on the flower’s leaves. Should she move towards where the sun appears to be, or wait to see whether this is an impermanent phenomenon? The fitness requirement is whether leaf and branch orientation are optimal now or would be in another configuration. Likely she will decide this based upon the effects such intermittent shading has on photosynthesis. Hayles’ second criterion is, “they are adaptive; they change their behaviors as a result of fitness challenges such as homeostasis for the cell.”139 The flowering plant produces noxious chemicals to ward off the challenge of chomping caterpillars. When her leaves are not under this stress, she does not release these noxious chemicals. Hayles’ third criterion is, “they are complex, composed of parts interacting with each other in multiple recursive feedback loops, or what Andy Clark calls ‘continuous reciprocal causation.’ Consequently, they exhibit emergence, results that cannot be predicted and that exceed the sum of their parts.”140 The plant is metaphysically all middle, but its processes are interconnected by the vascular system that produces multiple feedback loops from stimuli such as light and darkness, sunlight and shade, water near the roots and not, heat and transpiration, and root encroachment on others of the same species. I suggest predictability can be found in a linear system. The sun warms the rock to a certain temperature that can be predicted by the duration and intensity of sunlight. The flower will use transpiration and other means not to exceed certain temperatures even as she tries to optimize her use of solar energy. Optimization is not a linear process like sunlight on the rock because it requires a decider to assess options and then make decisions, something which linear processes cannot do. The gestalt of the plant is that the behaviors and memory of the plant resemble that of animals many believe are conscious, like the honeybee. Fourth and finally, Hayles requires that, “they are ‘constraint driven,’ which implies that the individual agent’s behaviors are guided by simple instincts or rules that constrain them to certain productive paths.”141 The plant orients itself towards the light and when turned upside down its anti-gravity sensors return the plant leaves towards the sun and its gravity sensory roots towards the earth. We can say that this is instinctual, but both the gravity and antigravity moves are the subject of simple rules that do not require language or even consciousness to perform. Other activities of plants such as the recording 139  Ibid., 200. 140  Ibid., 200–01. 141  Ibid.

Flower and Honeybee Epistemology and Behavior

121

of events for future use are rule driven as well, but can we call them instincts or is there something more? The something more Hayles suggests can be expressed as nonconscious cognition, “Nonconscious cognitive systems are distinct from the processes that generate them because they show an ‘intention toward’ not present in the underlying material processes as such.”142 Plants are intentioned towards the world centrifugally—they continually grow—and this involves more than just the sum of their processes. This study also suggests that the flower and honeybee facultative mutualism is more than just co-evolved ontological affordances and constraints that lead both species to perform activities towards each other autonomously and instinctually. What this ‘more than’ in flowers and in the flower and honeybee facultative mutualism deserves attention and definition. Optimal decision making from multiple options abound in the flower and honeybee relationship that cannot be explained by process driven theories alone. Hayles contrasts the glacier tumbling down a hill with honeybee hive workers collaboratively constructing their hexagonal cells in the hive.143 The glacier cannot adapt; it must flow down in a manner that gravity, water, and slope require. Therefore, the process of the glacier moving down the hill emerges nothing more from the processes that move water and earth. The honeybees who stand in a circle spit wax that eventually becomes the optimal hexagon, an emergent phenomenon. One worker working alone would likely not produce the same result. The hexagon that results occurs outside of the cognitive activities of the workers who participate in its construction. Hayles suggests that cognitive nonconscious processes like the collective action of hive bees, “demonstrate emergence, adaptation, or complexity.”144 The question is, how can we think beyond an aggregate of simple processes when trying to cognize the flower? Hayles suggests that we consider this: As a general concept, the term ‘cognitive nonconscious’ does not specify whether the cognition occurs inside the mental world of the participant, between participants, or within the system as a whole. It may operate wholly independently from consciousness, as in the cases of bees and termites, or it may be part of the larger system such as a human,

142  Ibid., 201. 143  Ibid. 144  Ibid.

122

Chapter 3

where it mediates between material processes and the emergence of consciousness/unconsciousness.145 However, cognitive nonconsciousness in artificial intelligence does not imply cognition in an organic creature. As previously mentioned, Hayles makes a distinction between cognition and thinking where, “[w]hile all thinking is cognition, not all cognition is thinking.”146 Hayles frees us up from the neurological ‘thinking’ to pursue cognition as not having to arise from a separate neurological pathway that controls other functions of the thinking creature. Rather, cognition permits us to think in terms of distributed processes that send information back and forth based upon stimulus in order to produce an optimal response to what has been sensed. The buzz pollinated flower reacts to the specific vibration and loosens pollen for the plant’s significant other. The flower can perform complex modeling, make optimal decisions, and produce the optimal response when required without requiring either thinking or traditional notions of animal consciousness. Optimization research looks at processes in nature such as ant and termite colonies, plant canopy structure, and many chemical processes to develop algorithms to use in artificial intelligence and logistics. What these optimization routines produce is a form of noncognitive cognition as Hayles has defined the term. The diverse orientation possibilities (individual, group, process) for cognitive nonconsciousness therefore can be more comfortably applied to the emergent properties of the flower, the honeybee, and their mutualism without requiring consciousness, individually or collectively. Therefore, rather than try to shoehorn animal consciousness processes and properties into the plant because we have no other explanation for their complexity that seems beyond connected processes, we can consider this complexity that solves complex problems without language through the lens of cognitive nonconsciousness. Artificial intelligence research may in the future be helpful in better cognizing how plants do what they do rather than trying to equate plant intelligence with animal intelligence, especially where consciousness is involved. What cognitive non-consciousness also does when it defines cognition differently from thinking, is remove the need for an anthropomorphic comparison. This does not mean that the earlier discussion of thinking in this study is obviated. Substituting cognition now for thinking in context of the flowering plant helps to differentiate the human who has a mind and the flower who does not. What Hayles has done is name thinking the thing that animals with central nervous 145  Ibid., 202. 146  Ibid., 201.

Flower and Honeybee Epistemology and Behavior

123

systems do. Plants and artificial intelligence systems who do not have a central nervous system cognize rather than think. Fundamentally both thinking and cognition are associated with producing optimal decisions. Cognition can be observed both in creatures who think and creatures, and artificial intelligence who do not. The difficulty is distinguishing between artificial and life-generated cognition without thinking. Could the notion of soul be the difference between artificial and life cognition? Marder includes in the construction of plant soul traditional philosophical notions of what constitutes soul and, “a certain manner and rhythm of movement” which in plants can be seen as growth and decay.147 Plants can replace lost or damaged limbs, leaves, roots, and sometimes flowers. He says, “The plant’s life is expressed in an incessant, wild proliferation, a becoming-spatial and a becoming-literal of intentionality.”148 The plant is a being who is always already a becoming as its continual growth and decay attest. Plants have a metaphysical right to be existents for their own sake and not ours. However, the flowering plants extend this invitation to the equality of existence to the other, the honeybee who has become an intimate in its existential struggle. Marder initiates this discussion with these questions: For, what if plant-soul and plant-thinking let the other pass through them without detracting from its alterity? What if they grow so as to play this role more effectively, to welcome the other better? And what if all this is accomplished thanks to the essential incompletion of linear growth that does not return to itself but is, from the very outset, other to itself? What if, finally, this inherent respect for alterity spelled out a key meaning of vegetative life?149 The flower reaches centrifugally out to the world through growth and advertisement to welcome the honeybee to her. She never stops growing and decaying which means that she cannot produce a teleological ontological structure while alive. She welcomes the alterior (different) honeybee to her who is not only ontologically alterior but her orientation to the flower is asymmetrical, centripetal. This asymmetry, as will be explained, is essential to the maintenance of the facultative mutualism. The flower, on the other hand, maintains her centrifugal focus without thinking but she does use cognition to help her

147  Marder, “Plant-Soul: The Elusive Meanings of Vegetative Life,” 85. 148  Ibid., 95. 149  Ibid., 98.

124

Chapter 3

accomplish her existential goals. I turn to an artificial intelligence example to help exemplify what the plant does. The smart speaker when on is always aware. It hears what the world produces in sounds but remains in that listening-only mode until it receives a particular auditory stimulus—its name being called. One smart speaker reacts to alert the vocalizer that she has received this particular auditory stimulus by turning on a light. This alerts the vocalizer to ask a question or command her to perform a certain action. After the vocalizer asks the question or makes the command, the smart speaker interprets the command and performs what the vocalizer has asked (if she understands). Once the command has been accomplished, she turns off her light and then returns to awareness mode. Consider the flower now in context of the actions the smart speaker takes. First, the flower is always in receptive mode, always aware. When an insect enters the flower, its tactile presence alerts the flower (like turning on the light) that an insect has arrived. The insect buzzes at the frequency the flower understands, and this command generates her response (for the smart speaker: the alarm is set for 10:30) which is to loosen pollen. If the frequency is not a frequency she recognizes as something she can respond to, she does not loosen pollen. When the insect leaves, the flower returns to awareness mode where she will remain until another stimulus begins. However, there is more to the flower than just smart speaker logic. After she is pollinated, the flower turns off not only her awareness of insect function, some change color, and all flowers desiccate eventually, because what had been the equivalent of smart-speaker technology converts itself into seed and fruit making technology. No smart speaker can do that today. Therefore, while cognitive nonconsciousness can explain some of what the plant does in terms that do not require thinking or consciousness, the complexity of plant processes is that they produce intricate emergences like the transition from flower production to seed production. As artificial intelligence becomes more sophisticated to where it can both learn and evolve processes to conform to environmental exigencies, we may be able to consider these capabilities through the lens of what plants can do. We also can consider cognitive nonconsciousness through the facultative mutualism construct of flowers and honeybees. The mutualism requires the participation of two cognitive beings much like the smart speaker and its vocalizer. We know that the smart speaker can learn from consistent vocalizations and through algorithms and behind the scenes assessment of humans, to react to new or slightly different stimuli. As such, both the smart speaker and the vocalizer benefit from their interaction and can evolve their interactions towards their ‘mutual’ benefit over time. Much the same happens with the flower and honeybee facultative mutualism (without the human intervention)

Flower and Honeybee Epistemology and Behavior

125

that serves to strengthen the relationship over time and co-evolve the mutualism. This living-creature mutualism is inter-dependent for at least some of their different life functions. The smart speaker and vocalizer do not achieve the level of dependency because the smart speaker has no existential life functions. However, this suggests that a fundamental difference between the smart speaker and vocalizer relationship and the flower and honeybee facultative mutualism is existential dependency on the alterior other. Therefore, the cognitive capabilities of the flower and the honeybee must be different from that of artificial intelligent routines. Artificial intelligence uses cognition, but this cognition cannot yet produce the same mutual interdependency that flowers and honeybees can. We can say that honeybees are thinking cognitive creatures, not just cognitive creatures (even if some of their cognitive work is unconscious). If flowers are cognitively nonconscious just like artificial intelligence, what does the ‘creature’ component provide the flower that artificial intelligence does not? I suggest that there is a gestalt to the flower and plants in general. Because they are all middle their processes are always already engaged with the world, not just in the form of limited awareness to the human voice as is the smart speaker to her world. Certainly, more complex smart speakers can be built to asses visual and other cues. However, life does not just develop routines to react to external and internal stimulus, it also maintains the continuity of life itself which it does through the change of its capabilities over time through genetic and epigenetic processes that adapt routines and processes towards changes in the ecology and environment. That optimization (while it is also the province of artificial intelligence) has become the means for plant and animal life to assess whether morphological change is becoming necessary or not. Once the options to the problem fall outside the realm of the artificial intelligence algorithm (even if minimally so), the non-learning capable algorithm can no longer perform (without human intervention). This can happen when life confronts environmental conditions that no longer permit its existence. However, if the change is not life threatening, the life form evolves itself to alter its routine. Evolution is both deliberative and the result of random mutation. Deliberation towards optimal decisions is the province of the existent. Some of what the creature learns can be passed down to future generations epigenetically (or through teaching). Other evolutionary requirements are achieved through random mutations that may produce beneficial or harmful results in offspring. Complicating this even further, the flower and honeybee mutualism uses co-evolutionary practices to bring the two participants into better alignment. Their asymmetrical orientation towards each other encourages this continual effort to better align the two species. Therefore, flower capabilities for cognition are enhanced by other processes.

126

Chapter 3

While I agree with Hayles that we cannot say that artificial intelligences think, I suggest that plant cognition in concert with these other processes produces a gestalt that artificial intelligence has yet to achieve. The problem is how to define this gestalt objectively. Whether this gestalt can emerge only through life processes remains to be seen. If advances in artificial intelligence begin to imitate existential processes including reproduction, random mutation, and co-evolution we may begin to gain greater insight as to what makes up this gestalt. 6

Moral Elegance

Plants are aware and they cognize, and honeybees think. They live separate lives but are co-dependent upon each other: honeybees for nourishment; flowers for reproduction. They both benefit from and exploit each other’s capabilities but do so with considerable restraint and they make optimal judgements for themselves and the social mutualism. Honeybees are eusocial and flowering plants may co-exist with other plants in an interdependent structure that is not yet well understood. While they became a mutualism through coevolutionary processes, they require a moral process of co-existence to sustain the mutualism. This moral process is governed by the optimal behaviors that both existents employ. Optimality ultimately produces MEP which is consistent with the second law of thermodynamics. From the science presented so far, we can begin to see that morality emerges from natural processes. We have also observed that Singer’s elegantly simple rules: social group, restraint, judgment are present in nature and are exemplified through the flower and honeybee facultative mutualism. Morality does not require written rules nor a sovereign as the flower and honeybee facultative mutualism shows. Also, moral systems like that of flowers and honeybees can endure, in this one instance for at least a million years. While both flowers and honeybees have evolved during their association, they have consistently adhered to Singer’s three requirements for the emergence of morality in nature. While processes like the honeybee waggle dances and even flower nectar production or coloration may have evolved over time, the principal driver for both species has been to optimize their existence which in turn maintains their social construct as fundamentally moral and likely more efficient as a result. This chapter has given us insight into what plants and honeybees do and some, even if limited information, for how they do it. There is, however, an existential condition beyond ontology/morphology and epistemology/­behavior

Flower and Honeybee Epistemology and Behavior

127

that must be considered and that is the nascent but rapidly developing science of epigenetics that proposes a third realm of engagement with the world and that is through changes in genetic expression that do not involve mutation but also influence individual and offspring viability. Cited References Amdam, Gro Vang, and Stig W. Omholt. “The Hive Bee to Forager Transition in Honey­ bee Colonies: The Double Repressor Hypothesis.” Journal of Theoretical Biology 223, no. 4 (2003/08/21/ 2003): 451–64. doi:https://doi.org/10.1016/S0022-5193(03)00121-8. Arenas, Andrés, Gabriela Ramírez, María Sol Balbuena, and Walter M. Farina. “Behavioral and Neural Plasticity Caused by Early Social Experiences: The Case of the Honeybee.” Frontiers In Physiology 4, no. Article 41 (2013): 2–11. doi:10.3389/ fphys.2013.00041. Bateson, Melissa, Suzanne Desire, Sarah E. Gartside, and Geraldine A. Wright. “Agitated Honeybees Exhibit Pessimistic Cognitive Biases.” Current Biology 21, no. 12 (2011): 1070–73. doi:https://doi.org/10.1016/j.cub.2011.05.017. Beekman, M., and F. L. W. Ratnieks. “Long‐Range Foraging by the Honey‐Bee, Apis Mellifera L.” Functional Ecology 14, no. 4 (2000): 490–96. Beiler, Kevin J., Daniel M. Durall, Suzanne W. Simard, Sheri A. Maxwell, and Annette M. Kretzer. “Architecture of the Wood‐Wide Web: Rhizopogon Spp. Genets Link Multiple Douglas‐Fir Cohorts.” New Phytologist 185, no. 2 (Jan 2010): 543–53. doi:10.1111/j.1469-8137.2009.03069.x. Bird, Adrian. “Perceptions of Epigenetics.” Nature 477, no. 24 (2007): 396–98. Chamovitz, Daniel. What a Plant Knows: A Field Guide to the Senses. Kindle ed. Oxford, UK: Scientific American/Farrar, Straus and Giroux, 2012. Dötterl, Stefan, Ulrike Glück, Andreas Jürgens, Joseph Woodring, and Gregor Aas. “Floral Reward, Advertisement and Attractiveness to Honey Bees in Dioecious Salix Caprea.” PloS one 9, no. 3 (2014): e93421, 1–11. Farina, Walter M., and Josue A. Núñez. “Trophallaxis in the Honeybee, Apis Mellifera (L.) as Related to the Profitability of Food Sources.” Animal Behaviour 42, no. 3 (1991/09/01/ 1991): 389–94. doi:https://doi.org/10.1016/S0003-3472(05)80037-5. Franks, Nigel R., Stephen C. Pratt, Eamonn B. Mallon, Nicholas F. Britton, and David J. T. Sumpter. “Information Flow, Opinion Polling and Collective Intelligence in House-Hunting Social Insects.” Philosophical Transactions: Biological Sciences 357, no. 1427 (Nov 29 2002): 1567–83. doi:10.1098/rstb.2002.1066. Gagliano, Monica. “The Flowering of Plant Bioacoustics: How and Why.” Behavioral Ecology 24, no. 4 (2013): 800–01. doi:10.1093/beheco/art021 %J Behavioral Ecology.

128

Chapter 3

Gagliano, Monica. “Green Symphonies: A Call for Studies on Acoustic Communication in Plants.” Behavioral Ecology 24, no. 4 (2012): 789–96. doi:10.1093/beheco/ars206 %J Behavioral Ecology. Gagliano, Monica, Mavra Grimonprez, Martial Depczynski, and Michael Renton. “Tuned In: Plant Roots Use Sound to Locate Water.” Oecologia 184, no. 1 (2017/05/01 2017): 151–60. doi:10.1007/s00442-017-3862-z. Gagliano, Monica, Michael Renton, Nili Duvdevani, Matthew Timmins, and Stefano Mancuso. “Acoustic and Magnetic Communication in Plants: Is It Possible?”. Plant Signaling & Behavior 7, no. 10 (2012): 1346–48. doi:10.4161/psb.21517. Getz, Wayne M., Dorothea Brückner, and Thomas R. Parisian. “Kin Structure and the Swarming Behavior of the Honey Bee Apis Mellifera.” Behavioral Ecology and Sociobiology 10, no. 4 (1982): 265–70. Giurfa, Martin. “Cognition with Few Neurons: Higher-Order Learning in Insects.” Trends in Neurosciences 36, no. 5 (May 2013): 285–94. doi:10.1016/j.tins.2012.12.011. Gould, James L. “Honey Bee Cognition.” Cognition 37, no. 1–2 (Nov 1990): 83–103. Gould, James L., and Carol Grant Gould. The Honey Bee. Edited by Carol Grant Gould New York: Scientific American Library: Distributed by W. H. Freeman, 1988. Hammer, Martin, and Randolf Menzel. “Learning and Memory in the Honeybee.” Journal of Neuroscience 15, no. 3 (Mar 1995): 1617–30. Hayles, N. Katherine. “Cognition Everywhere: The Rise of the Cognitive Nonconscious and the Costs of Consciousness.” New Literary History 45, no. 2 (2014): 199–220. doi:10.1353/nlh.2014.0011. Heidegger, Martin. Being and Time Translated by John MacQuarrie, & Edward Robinson. New York: HarperCollins Publishers, 1962. Heidegger, Martin. Discourse on Thinking: A Translation of Gelassenheit New York: Harper & Row, 1969. Heidegger, Martin. What Is Called Thinking Translated by Fred D. & J. Glen Gray Wieck. New York: Harper & Row Publishers, 1968. Holland, Heinrich D. “The Oxygenation of the Atmosphere and Oceans.” Philosophical Transactions of the Royal Society B: Biological Sciences 361, no. May (2006): 903–15. Johnson, Brian R., and James C. Nieh. “Modeling the Adaptive Role of Negative Signaling in Honey Bee Intraspecific Competition.” Journal of Insect Behavior 23, no. 6 (2010/11/01 2010): 459–71. doi:10.1007/s10905-010-9229-5. Ketcham, Christopher. “Schopenhauer and Buddhism: Soulless Continuity.” Journal of Animal Ethics 8, no. 1 (2018): 12–25. Kietzman, Parry M., and P. Kirk Visscher. “The Anti-Waggle Dance: Use of the Stop Signal as Negative Feedback.” [In English]. Frontiers in Ecology and Evolution 3, no. 14 (2015-February-17 2015): 1–5. doi:10.3389/fevo.2015.00014. Kirchner, Wolfgang H. “Vibrational Signals in the Tremble Dance of the Honeybee, Apis Mellifera.” Behavioral Ecology and Sociobiology 33, no. 3 (1993): 169–72.

Flower and Honeybee Epistemology and Behavior

129

Leoncini, Isabelle, Yves Le Conte, Guy Costagliola, Erika Plettner, Amy L. Toth, Mianwei Wang, Zachary Huang, et al. “Regulation of Behavioral Maturation by a Primer Pheromone Produced by Adult Worker Honey Bees.” Proceedings of the National Academy of Sciences of the United States of America 101, no. 50 (2004): 17559-64. doi:10.1073/pnas.0407652101. Lipiñski, Zbigniew. “The Emotional Nature of the Worker Honeybee (Apis Mellifera L.).” Journal of Apicultural Science 50, no. 1 (2006): 49–62. Marder, Michael. “Plant-Soul: The Elusive Meanings of Vegetative Life.” Journal of Environmental Philosophy 8, no. 1 (2011): 83–100. Marder, Michael. Plant-Thinking: A Philosophy of Vegetal Life [in English]. None. New York: Columbia University Press, 2013. Book. Marder, Michael. “What Is Plant-Thinking.” Klesis—Revue Philosophique 25 (2013): 124–43. Menzel, Randolf. “The Honeybee as a Model for Understanding the Basis of Cognition.” Nature Reviews Neuroscience 13, no. November (2012): 758–68. Menzel, Randolf, and Alison Mercer. Neurobiology and Behavior of Honeybees. Berlin Germany, New York: Springer-Verlag, 1987. Menzel, Randolph. “Behavioral and Neural Mechanisms of Learning and Memory as Determinants of Flower Constancy.” Chap. 2 In Cognitive Ecology of Pollination: Animal Behaviour and Floral Evolution, edited by Lars Chittka and James D. Thomson, 21–40. Cambridge, UK: Cambridge University Press, 2001. Nealon, Jeffrey T. Plant Theory. Stanford, Ca.: Stanford University Press, 2015. Nieh, James C. “A Negative Feedback Signal That Is Triggered by Peril Curbs Honey Bee Recruitment.” Current Biology 20, no. 4 (2010): 310–15. Page, Robert E., Gene E. Robinson, M. Kim Fondrk, and Medhat E. Nasr. “Effects of Worker Genotypic Diversity on Honey Bee Colony Development and Behavior (Apis Mellifera L.).” Behavioral Ecology and Sociobiology 36, no. 6 (1995): 387–96. Palmer, John. Parmenides and Presocratic Philosophy. Oxford: Oxford University Press, 2009. Popova, Evgenya, and Colin J. Barnstable. “Epigenetics Rules.” Journal Of Ocular Biology, Diseases, And Informatics 4, no. 3 (2012): 93–94. doi:10.1007/s12177-012-9088-8. Sartre, Jean-Paul. “Existentialism Is a Humanism.” In Existentialism from Dostoyevsky to Sartre, edited by Walter Kaufman Whangarei, New Zealand: Pickle Partners Publishing, 2016. Schlegel, Thomas, P. Kirk Visscher, and Thomas D. Seeley. “Beeping and Piping: Characterization of Two Mechano-Acoustic Signals Used by Honey Bees in Swarming.” Naturwissenschaften 99, no. 12 (2012/12/01 2012): 1067–71. doi:10.1007/ s00114-012-0990-5. Schopenhauer, Arthur. The World as Will and Representation Ii. Translated by R. B. & J. Kemp Haldane. Vol. II, London: Kegan Paul, Trench, Trubner & Co. Ltd., 1910.

130

Chapter 3

Schürch, Roger, Margaret J. Couvillon, and Madeleine Beekman. “Editorial: Ballroom Biology: Recent Insights into Honey Bee Waggle Dance Communications.” [In English]. Frontiers in Ecology and Evolution 2016, no. February (2016-January-07 2016): 4–6. doi:10.3389/fevo.2015.00147. Seeley, Thomas D. Honeybee Democracy. Princeton, NJ: Princeton University Press, 2010. Seeley, Thomas D. The Wisdom of the Hive. Cambridge, Ma.: Harvard University Press, 1995. Seeley, Thomas D., and Royce A. Levien. “A Colony of Mind.” Sciences 27, no. 4 (1987): 38–43. doi:10.1002/j.2326-1951.1987.tb02955.x. Song, Yuan Yuan, Ren Sen Zeng, Jian Feng Xu, Jun Li, Xiang Shen, and Woldemariam Gebrehiwot Yihdego. “Interplant Communication of Tomato Plants through Underground Common Mycorrhizal Networks.” PloS one 5, no. 10 (2010): e13324, 1–11. doi:10.1371/journal.pone.0013324. Su, Songkun, Fang Cai, Aung Si, Shaowu Zhang, Jürgen Tautz, and Shenglu Chen. “East Learns from West: Asiatic Honeybees Can Understand Dance Language of European Honeybees.” PLoS One 3, no. 6 (2008): e2365, 1–9. Tarán, L. Parmenides: A Text with Translation, Commentary, and Critical Essays. Princeton, NJ: Princeton University Press, 1965. Tezze, Andrea A., and Walter M. Farina. “Trophallaxis in the Honeybee, Apis Mellifera: The Interaction between Viscosity and Sucrose Concentration of the Transferred Solution.” Animal Behaviour 57, no. 6 (1999/06/01/ 1999): 1319–26. doi:https://doi .org/10.1006/anbe.1999.1110. Tudge, Colin. The Tree. New York: Three Rivers Press, 2995. Tulving, Endel. “Memory and Consciousness.” Canadian Psychology/Psychologie Canadienne 26, no. 1 (1985): 1–12. doi:10.1037/h0080017. Tye, Michael. Tense Bees and Shell-Shocked Crabs. Okford, UK: Oxford University Press, 2017. Visscher, P. Kirk, and Thomas D. Seeley. “Coordinating a Group Departure: Who Produces the Piping Signals on Honeybee Swarms?”. Behavioral Ecology & Sociobiology 61, no. 10 (Jan 2007): 1615–21. doi:10.1007/s00265-007-0393-3. Waddington, Keith D. “Subjective Evaluation and Choice Behavior by Nectar- and Pollen-Collecting Bees”. Chap. 3 In Cognitive Ecology of Pollination: Animal Behaviour and Floral Evolution, edited by Lars Chittka and James D. Thomson, 41–60. Cambridge, UK: Cambridge University Press, 2001. Walton, Alexander, and Amy L. Toth. “Variation in Individual Worker Honey Bee Behavior Shows Hallmarks of Personality.” Behavioral Ecology and Sociobiology 70, no. 7 (2016/07/01 2016): 999–1010. doi:10.1007/s00265-016-2084-4. Wohlleben, Peter. The Hidden Life of Trees: What They Feel, How They Communicate— Discoveries from a Secret World. Vancouver, BC Canada: Greystone Books, 2016.

Flower and Honeybee Epistemology and Behavior

131

Woyciechowski, M., and K. Kuszewska. “Swarming Generates Rebel Workers in Honeybees.” Current Biology 22, no. 8 (Apr 24 2012): 707–11. doi:10.1016/ j.cub.2012.02.063. Wray, Margaret K., Heather R. Mattila, and Thomas D. Seeley. “Collective Personalities in Honeybee Colonies Are Linked to Colony Fitness.” Animal Behaviour 81, no. 3 (2011/03/01 2011): 559–68. doi:10.1016/j.anbehav.2010.11.027. Wray, Margaret K., and Thomas D. Seeley. “Consistent Personality Differences in House-Hunting Behavior but Not Decision Speed in Swarms of Honey Bees (Apis Mellifera).” Behavioral Ecology and Sociobiology 65, no. June (2011/06/15 2011): 2061– 70. doi:10.1007/s00265-011-1215-1.

Chapter 4

Epigenetics 1

Epigenetics Defined

Darwin taught us that species mutate into new species. Genetics has taught us that mutations of specific genes are responsible for producing new species. New species come about because of environmental changes or other existential challenges that individuals must deal with during their lives. Mutation is a long and probabilistic process. It does little help to species endure acute drought or other temporary contingencies. As we learn more how DNA functions, science is learning that many genes are not always turned on. Some, like stem cells are responsible for growth and development and eventually turn off. Other genes may not ever turn on until environmental conditions warrant their use. Some may just be relics from our past. Epigenetics is the study of genetic expression of individual or combinations of genes during the lifetime of individual existents. Genetic expression can be on, off, or some other mode that is not mutation, but a variation on gene functioning that is a capability of the gene itself. There is emerging evidence in some life forms that the changes in genetic expression of parents, for example after dealing with trauma and disease, can be passed down to offspring and perhaps into many future generations. Each creature is born with capabilities to exist in its environment. Its genes provide these capabilities. When change makes it untenable for the creature to exist as is in its environment, and if given enough time to evolve, the process of mutation can help species evolve capabilities to cope with change. Sandwiched in between the quick individual response to current conditions and the very long process of mutation is epigenetics, or intermediate processes that are activated only by the individual and species when conditions warrant. What makes these epigenetic processes special is that they can, for example, help existents and offspring ward off an acute plague, but can be shut off when the plague no longer affects existents. For want of a better phrase, they provide ‘temporary mutation,’ not of the chromosome order but of the expression of the gene until that expression change is no longer required. To use a crude analogy, epigenetics is the blue tarp pulled from the basement to the roof after the hurricane. Before the hurricane, the blue tarp is not helpful and remains in storage. After the hurricane is gone and the roof is repaired, the blue tarp goes back into the basement where it can be used again should another hurricane form. Owners of the structure can sell the house with the blue tarp in

© Koninklijke Brill NV, Leiden, 2020 | doi:10.1163/9789004428546_006

Epigenetics

133

the basement so that successive owners will have the benefit of the temporary repair device. Science has discovered epigenetic processes in both plants and animals. Epigenetics is important to this study of the emergence of morality in nature because it is an additional tool that species can use to enhance personal capabilities. Species can use epigenetics to optimize their existence and perhaps that of their offspring. Epigenetics factors in the experience of the creature to make changes in genetic expression. In other words, epigenetics is activated by experience and the interface of the creature with her environment. Keeping creatures viable and functioning can help them maintain social structures like the flower and honeybee facultative mutualism, perhaps even serving to strengthening the relationship over time. Singer does not include durability of the social structure as a requirement for the emergence of ethics in nature. However, the fundamental relationship between flowers and honeybees has endured for at least a million years. While it is true that both flowers and honeybees have evolved during this relationship, their social group has also endured, and they continue to thrive as a group, making decisions towards the preservation of their social group, while using also restraint in their actions towards each other. If morality in a single social group can endure for a million years or more, this should give humanity pause to consider how important morality can be to species and their relationship to others in the greater ecology. In the last million years, both flowers and honeybees have had to endure much in the way of climate change including the last ice age that ended about ten thousand years ago. Still their facultative mutualism has endured and perhaps one reason for this is the process called epigenetics. Epigenetics does not fit easily into ontological/morphological or epistemological/behavioral categories. Arthur Riggs defines epigenetics as, “[t]he study of mitotically and/or meiotically heritable changes in gene function that cannot be explained by changes in DNA sequence.”1 To which Adrian Bird revises the definition to; “[t]he structural adaptation of chromosomal regions so as to register, signal or perpetuate altered activity states.”2 In both definitions, epigenetics is the study of changes in the expression of genes without changing the order of genes in chromosomes. Epigenetics is not mutation where one or more genes or the chromosome order are irretrievably changed during the 1  Adrian Bird, “Perceptions of Epigenetics,” Nature 477, no. 24 (2007): 396. Note: Mitosis is where a cell divides into two identical cells. Meiosis is where the cells divide into four cells, each with half the chromosomes of their parents and where daughter cells are genetically different. 2  Ibid., 398.

134

Chapter 4

reproductive process. Evgenya Popova and Colin J. Barnstable outline the six primary mechanisms of epigenetics: 1. 2. 3. 4. 5. 6.

DNA methylation Chromatin remodeling by chromatin-associated proteins Histone modifications Histone variants and their composition Non-coding regulatory RNAs Chromatin 3D structure including DNA looping and nuclear territories3

Explaining how each of these mechanisms work is not within the scope of this study. What is important is to understand that life has developed at least six ways to modify gene expression without changing the chromosome order. Popova and Barnstable note that, “Genes can be in one of several functional states: transcribed, poised for transcription, inactivated, and silenced.”4 Epigenetic mechanisms can switch genetic states of some genes to serve specific purposes. 2

Promise of Epigenetics

During the life of any individual creature, the creature experiences the exigencies in the world of its existence. Many higher life forms, both plant and animal, can learn from these exigencies. Epigenetics is a kind of learning function that can alter gene functionality or ‘activity’ states in order to help the organism cope with such changes. Adrian Bird notes the promises for study of epigenetics, “Also included is the exciting possibility that epigenetic processes are buffers of genetic variation, pending an epigenetic (or mutational) change of state that leads an identical combination of genes to produce a different developmental outcome.”5 Epigenetic processes convert experiences of the parent or the individual into genetic expressions that can be reversed or even changed in future generations. Peter Sarkies notes, “Nevertheless, so far the molecular studies discussed in this article suggest that transgenerational epigenetic changes in the absence of sequence change may be at best a minor 3  Evgenya Popova and Colin J. Barnstable, “Epigenetics Rules,” Journal Of Ocular Biology, Diseases, And Informatics 4, no. 3 (2012): 93. 4  Ibid. 5  Bird, “Perceptions of Epigenetics,” 398.

Epigenetics

135

contributor to short term adaptation and long-term evolution.”6 Rather than go the more permanent and experimental route of mutation, in epigenetics the gene can express itself differently depending upon the stressors the individual experiences. This process appears to have the hallmark of an intermediate step in evolutionary processes that is between individual learning and permanent species change. However, Sarkies also suggests, “An interesting possibility therefore is that organisms have evolved mechanisms in order to prevent epigenetic changes from becoming fixed in populations. The ability of epigenetic mechanisms to respond directly to the environment might be dangerous if their activities were to result in fixed gene expression differences for subsequent generations.”7 These are both important concerns because we know that some plants can pass along height and flowering time information to future generations.8 Often this can be passed along to many future generations, purportedly as long as this information is valuable to succeeding generations. If, for example, the temporal presence of pollinators should change, the plant can cease passing along outdated information about flowering time to offspring and replace it with new information. Plant stress situations such as predation may also be passed along. For example, Alexander Boyko discovered evidence of intergenerational epigenetics information transfer in tobacco plants that were subjected to biotic stressors.9 3

Epigenetic Purposes

Corné M. J. Pieterse suggest a purpose for epigenetics, “However, less frequent stress situations may only persist for only 1 or a few generations, which is often too short for genetic adaptations to establish in the population.”10 An interme6  Peter Sarkies, “Molecular Mechanisms of Epigenetic Inheritance: Possible Evolutionary Implications,” Seminars in Cell & Developmental Biology (2019, In Press): 8. 7  Ibid. 8  E.g., “Studies in the plant Arabidopsis thaliana showed that alterations in DNA methylation can be transmitted through several generations” Corina Nagy and Gustavo Turecki, “Transgenerational Epigenetic Inheritance: An Open Discussion,” Epigenomics 7.5 (2015): 781. Frank Johannes, et al., found heritability in plant height and flowering time: Frank Johannes et al., “Assessing the Impact of Transgenerational Epigenetic Variation on Complex Traits,” PLoS Genetics 5, no. 6 (2009): 1. 9  Alexander Boyko et al., “Transgenerational Changes in the Genome Stability and Methylation in Pathogen-Infected Plants: (Virus-Induced Plant Genome Instability),” Nucleic Acids Research 35, no. 5 (2007): 1714. 10  Corné M. J. Pieterse, “Prime Time for Transgenerational Defense,” Plant Physiology 158, no. 2 (2012): 545.

136

Chapter 4

diary step between temporary existential knowledge and permanent genetic change, as Bird intimates, may guide species over the long-term towards genetic evolution that is less random than has been previously thought. This may be how some species seem to adapt more quickly to environmental change, even towards new speciation over fewer generations than what a pure random mutation might offer. Bird suggests: An implicit feature of this proposed definition is that it portrays epigenetic marks as responsive, not proactive. In other words, epigenetic systems of this kind would not, under normal circumstances, initiate a change of state at a particular locus but would register a change already imposed by other events.11 This notion of epigenetic responsive assistance to species without mutation requires significant additional research. For example, Gideon Ney and Johannes Schul studied two isolated species of katydids. While they discovered evidence of epigenetics in both species, they did not find evidence that epigenetic processes contributed to “species isolation.”12 Therefore, there is an open question of whether epigenetics and the much longer-term and morepermanent process of mutation can be correlated or connected. Epigenetic processes that can prepare future generations for stressors can be thought of as boosting the possibility that the genes of the parents through their offspring will have a better chance of continuity than if there was no epigenetic process. Also, these stressors may change, and the parent can produce epigenetic changes to adapt to these new stressors. However, epigenetic studies have found that stressors on mammal mothers during pregnancy can epigenetically alter her offspring, perhaps not for the better.13 For example, 11  Bird, “Perceptions of Epigenetics,” 398. 12  Gideon Ney and Johannes Schul, “Epigenetic and Genetic Variation between Two Behaviorally Isolated Species of Neoconocephalus (Orthoptera: Tettigonioidea),” Journal of Orthoptera Research 28, no. 1 (2019): 17. 13  The literature on epigenetics is rich with study on epigenetic cancer and other disease triggers and negative implications of inadequate or injurious care on developing fetuses or infants that have long-term implications the maturing child that even may explain antisocial behavior in some adults. See this small selection for more information on the negative implications of epigenetics: Yang Wang et al., “Epigenetic Regulation and Risk Factors During the Development of Human Gametes and Early Embryos,” Annual Review of Genomics and Human Genetics 20, no. August (2019); Peter D. Fransquet et al., “The Epigenetic Clock as a Predictor of Disease and Mortality Risk: A Systematic Review and Meta-Analysis,” Clinical Epigenetics 11, no. 1 (2019); Laura Ramo-Fernández et al., “The Effects of Childhood Maltreatment on Epigenetic Regulation of Stress-Response

Epigenetics

137

Autumn J. Bernal and Randy L. Jirtle, found evidence of negative epigenetic intergenerational transfer in mice they suggest might have human implications: The sensitivity of the human epigenome to low levels of EDCs [endocrine disrupting compounds] will directly influence the health of current and future populations. If EDCs are shown to interfere with epigenetic programming at current exposure levels, researchers speculate that in addition to altering disease susceptibility, they could also be contributing to the documented increase in human infertility.14 The question of human trauma experienced by either the mother or father or both associated with epigenetics is an important area of research. During the U.S. Civil War, the Confederacy prisoner of war camps were notoriously overcrowded, with little food, medicine, or shelter. The conditions were so bad at Andersonville, that the camp commander was hanged after the war. A study of children of non-POW soldiers, POWs in the worst camp conditions, and POWs in the better camp conditions found mortality differences in POW male children, “The sons of ex-POWs imprisoned when camp conditions were at their worst were 1.11 times more likely to die than the sons of non-POWs and 1.09 times more likely to die than the sons of ex-POWs when camp conditions were better.”15 This effect was not found in daughters and was only found in sons born after the war. Even though Dora L. Costa, Noelle Yetter, and Heather DeSomer could not rule out psychological or cultural causes for such an effect, they maintain that the implications are the trauma of being in the camps had an epigenetic effect on male children born after their ordeal.16 On the other hand, Laura Ramo-Fernandez, et al., did not discover that childhood maltreated (CM) mothers passed along epigenetic markers on specific stressresponse-associated genes to their children:

Associated Genes: An Intergenerational Approach,” Scientific Reports 9, no. 1 (2019); Dan L. Longo and Andrew P. Feinberg, “The Key Role of Epigenetics in Human Disease Prevention and Mitigation,” The New England Journal of Medicine 378, no. 14 (2018). 14  Autumn J. Bernal and Randy L. Jirtle, “Epigenomic Disruption: The Effects of Early Developmental Exposures,” Birth Defects Research Part A: Clinical and Molecular Teratology 88, no. 10 (2010): 943. Item in brackets added. 15  Dora L. Costa, Noelle Yetter, and Heather DeSomer, “Intergenerational Transmission of Paternal Trauma among Us Civil War Ex-Pows,” Proceedings of the National Academy of Sciences 115, no. 44 (2018): 11215. 16  Ibid.

138

Chapter 4

Although future studies are needed to confirm the results e.g. in an epigenome-wide approach, our results have an important implication: from the point of view of DNA methylation, the offspring of CM-exposed mothers does not seem to necessarily display the same epigenetic patterns as their mothers in the targeted CpG sites. In this case, professionals should focus on psychosocial factors during the first years of life, which might prospectively buffer the potential transmission of CM-associated consequences.17 The civil war study did find what they classify as epigenetic changes after parental trauma, but the childhood maltreatment study did not, suggesting family environment may play a role in the intergenerational transfer of stressors and behavior associated with maltreatment. Linda Witek Janusek et al., saw maternal stress influencing maternal behavior that may influence epigenetic changes in pre-natal and early post-natal nurturing. Evidence primarily exists for maternal psychosocial experiences (i.e. mood and exposure to stress, adversity, or trauma) to associate with epigenetic modification to offspring genes involved in neurobehavioral pathways (i.e. glucocorticoid, oxytocin, and serotonin system genes) … Epigenetic transmission of adverse early life experiences to the offspring genome most often occurs during the prenatal and early postnatal periods, when developing systems are more sensitive to environmental signals.18 Mothers who are depressed may not be as attentive as other mothers and this may produce negative epigenetic development in the womb and in the newborn that in combination produce maladaptive behaviors, that can follow the child into adulthood. Janusek, et al., suggest, “Emerging work suggests interventions that foster positive maternal—infant interactions may attenuate the epigenetic impact of early life stress.”19

17   Ramo-Fernández et al., “The Effects of Childhood Maltreatment on Epigenetic Regulation of Stress-Response Associated Genes: An Intergenerational Approach,” 7. 18  Linda Witek Janusek, Dina Tell, and Herbert L. Mathews, “Epigenetic Perpetuation of the Impact of Early Life Stress on Behavior,” Current Opinion in Behavioral Sciences 28 (2019): 1. 19  Ibid.

Epigenetics

4

139

General Implications of Epigenetics

Three human trauma studies have found three different ways that children can be affected by parental stress. The civil war study suggests that stress may be passed down epigenetically and can influence the mortality of children. The second study of childhood maltreated mothers did not show epigenetic transfer, but that perhaps the reason why childhood maltreatment passes through generations is more psychological. The third study suggests that stress on the mother, for example due to maternal depression, may produce negative epigenetic results both for the child who is not yet born and soon after birth where the child is most sensitive to environmental conditions. The suggestion is that good genes are not always enough. Epigenetic processes may have some influence on the success of offspring, but that parental behavior, especially early in the child’s life is highly influential on the child’s future behavior. As mentioned previously, epigenetic studies are relatively new, and we are likely to discover much more about the effects of environmental and internal stressors on individuals and their offspring. However, there may be a difference in epigenetics between sessile plants who do not raise their young and mammals and other animals like honeybees who do. Evidence reported in this study suggests that some plants can pass along information to offspring on how to defend against herbivores and other information such as plant height and flowering times which likely are consistent with the ecology in which parents are born and which offspring will also likely take root. This may provide the next generation with some advantage that the plant parent cannot provide through direct parental nurturing. However, we also know that plants can release ethylene and other volatile compounds through the air to warn of such things as caterpillar infestation. Ethylene is a compound that most plants produce, so this information can be passed to many different species. A recent study of tomato plants found, “Here, we show that CMNs [common mycorrhizal networks] mediate plant-plant communication between healthy plants and pathogen-infected tomato plants.”20 The tomato plants of the same species have common mycorrhizal fungi associated with their root structures and these can interconnect different tomato plants to produce a communication structure that alerts one plant that another has been adversely affected by pathogens. These plants are likely all from the same generation, so we are probably not seeing living parental experience being passed down to offspring, though that is possible with 20  Yuan Yuan Song et al., “Interplant Communication of Tomato Plants through Underground Common Mycorrhizal Networks,” PloS one 5, no. 10 (2010): 1. Item in bracket added.

140

Chapter 4

perennial flower patches. However, this does suggest that there are more complex intra-species communications between living members of the same plant generation than has been previously thought.21 Nature versus nurture becomes blurred in epigenetics. Epigenetics, as we have seen, can pass along information learned or experienced by the parental generation to future generations. If future generations have similar experiences, it appears, at least with plants, that they can epigenetically pass some of this information along to their future generations. If their experience is different, they may also be able to alter the expression of genes epigenetically to offspring to reflect this new experience. However, in human studies, science has discovered possible epigenetic effects from stress that is passed along to children in some cases, but not in others. Mutation, parental epigenetics and parental nurturing (in some combination or in some instances) may either help or hinder the development and success of their children. 5

Implications of Epigenetics for Flowers and Honeybees

There has been limited research on plant epigenetics, and the literature on honeybee epigenetics is sparse. As more research into epigenetics is conducted, we likely will learn much more how epigenetics can both help and harm individuals and future generations. Epigenetics is a powerful genetic tool that, especially in plants, can help prepare current and future generations by providing learning without education. While some communication between plants has been confirmed (air and root), there is minimal evidence that plants nurture their young.22 There is evidence that some plants, at least, when they encounter the roots of others, will stop growing in that direction.23 We cannot, 21  In another study on beans, researchers discovered that mycorrhizae networks can transmit information about aphid infestation, “We present the first experimental evidence that herbivore-induced signalling molecules can be transferred from plants infested with aphids to uninfested neighbours via a common mycelial network” Zdenka Babikova et al., “Underground Signals Carried through Common Mycelial Networks Warn Neighbouring Plants of Aphid Attack,” Ecology Letters 16, no. 7 (2013): 840. 22  This is not to deprecate the instincts of Wohlleben who suggests that there may be more interconnectedness and interdependency in forests where there are both young and old from the same genetic line Peter Wohlleben, The Hidden Life of Trees: What They Feel, How They Communicate—Discoveries from a Secret World (Vancouver, BC Canada: Greystone Books, 2016). 23  See: Gordon G. McNickle, “Root Foraging Behaviour of Plants: New Theory, New Methods and New Ideas” (University of Alberta, 2011); Katja Schiffers et al., “Root Plasticity Buffers Competition among Plants: Theory Meets Experimental Data,” Ecology 92, no. 3 (2011).

Epigenetics

141

however, call this nurturing. As flowering plants likely have limited direct influence over the actions of their offspring, epigenetic transfer of information to future generations is a powerful tool, that in limited respects, replaces this nurturing function. As they mature, honeybee workers take on different jobs and have different interactions with each other. Feedback mechanisms (like the dances) produce a kind of nurturing to maturing honeybees in the honeybee hive. Maturation pheromone or hormone transfers from older workers to younger workers may initiate epigenetic processes to retard or accelerate maturation. Workers do not reproduce so they cannot pass down epigenetic changes to offspring. How and what they forage and what the nursery workers feed larvae may have an impact on the next generation. For example, different years may produce different qualities and quantities of food and other resources. Whether this can or does introduce epigenetic change in the developing worker is not known. In other animals that nurture their young, sometimes bad or ineffective parenting can produce adverse epigenetic changes that can affect offspring for life and may or may not be transmitted to future generations. While epigenetics has been shown to be a process that nature evolved to prepare the young and future generations to meet exigencies of the environment, it appears that this powerful genetic tool can trigger adverse epigenetic changes associated with bad parenting or other stressors experienced during maturation. The implications of this are that when optimization is compromised (maternal depression, bad offspring raising, etc.), less than optimal behavior may result. Adverse epigenetic changes may also affect the mortality or morbidity of the offspring. Stress to the parent alone, as theorized by the Civil War prisoner study, may produce intergenerational epigenetic change that adversely affects offspring. Epigenetics and mutation are powerful tools that nature has developed to produce both positive and negative changes in organisms, families, and even species. Mutations that do not benefit the species may not survive many generations. Reactions to stress that produce adverse epigenetic changes may also produce a similar result. The introduction of epigenetics to the discussion of optimization returns to the notion that individuals can optimize only to the extent they are capable. Epigenetics also exemplifies the notion that there are deviations from species norms that affect maximum entropy production alongside the effects produced by differences between individuals and their optimization capabilities. Stressors like war, pandemics, opioid addicted mothers, and other systemic processes may change the MEP equation in ways like that of the forest fire or the invasive species. When optimization parameters are reset, then MEP values will change as a result. This observation will become more important in the final chapter of this study

142

Chapter 4

that considers the problem of maximizing crop production versus optimizing crop production. However, before we get to the question of maximization versus optimization in human activities compared to flowers and honeybees, it is important to develop the idea of the good and the emergence of morality in the flower and honeybee mutualism. Cited References Babikova, Zdenka, Lucy Gilbert, Toby J. A. Bruce, Michael Birkett, John C. Caulfield, Christine Woodcock, John A. Pickett, and David Johnson. “Underground Signals Carried through Common Mycelial Networks Warn Neighbouring Plants of Aphid Attack.” Ecology Letters 16, no. 7 (2013/07/01 2013): 835–43. doi:10.1111/ele.12115. Bernal, Autumn J., and Randy L. Jirtle. “Epigenomic Disruption: The Effects of Early Developmental Exposures.” Birth Defects Research Part A: Clinical and Molecular Teratology 88, no. 10 (2010): 938–44. doi:10.1002/bdra.20685. Bird, Adrian. “Perceptions of Epigenetics.” Nature 477, no. 24 (2007): 396–98. Boyko, Alexander, Palak Kathiria, Franz J. Zemp, Youli Yao, Igor Pogribny, and Igor Kovalchuk. “Transgenerational Changes in the Genome Stability and Methylation in Pathogen-Infected Plants: (Virus-Induced Plant Genome Instability).” Nucleic Acids Research 35, no. 5 (2007): 1714–25. Costa, Dora L., Noelle Yetter, and Heather DeSomer. “Intergenerational Transmission of Paternal Trauma among Us Civil War Ex-Pows.” Proceedings of the National Academy of Sciences 115, no. 44 (2018): 11215–20. doi:10.1073/pnas.1803630115. Fransquet, Peter D., Jo Wrigglesworth, Robyn L. Woods, Michael E. Ernst, and Joanne Ryan. “The Epigenetic Clock as a Predictor of Disease and Mortality Risk: A Systematic Review and Meta-Analysis.” Clinical Epigenetics 11, no. 1 (April 11 2019): 1–17. doi:10.1186/s13148-019-0656-7. Janusek, Linda Witek, Dina Tell, and Herbert L. Mathews. “Epigenetic Perpetuation of the Impact of Early Life Stress on Behavior.” Current Opinion in Behavioral Sciences 28 (2019/08/01/ 2019): 1–7. doi:https://doi.org/10.1016/j.cobeha.2019.01.004. Johannes, Frank, Emmanuelle Porcher, Felipe K. Teixeira, Vera Saliba-Colombani, Matthieu Simon, Nicolas Agier, Agnès Bulski, et al. “Assessing the Impact of Transgenerational Epigenetic Variation on Complex Traits.” PLoS Genetics 5, no. 6 (2009): e1000530, 1–11. Longo, Dan L., and Andrew P. Feinberg. “The Key Role of Epigenetics in Human Disease Prevention and Mitigation.” The New England Journal of Medicine 378, no. 14 (2018): 1323–34.

Epigenetics

143

McNickle, Gordon G. “Root Foraging Behaviour of Plants: New Theory, New Methods and New Ideas.” University of Alberta, 2011. Nagy, Corina, and Gustavo Turecki. “Transgenerational Epigenetic Inheritance: An Open Discussion.” Epigenomics 7.5 (2015): 781–94. Ney, Gideon, and Johannes Schul. “Epigenetic and Genetic Variation between Two Behaviorally Isolated Species of Neoconocephalus (Orthoptera: Tettigonioidea).” Journal of Orthoptera Research 28, no. 1 (2019): 11–19. Pieterse, Corné M. J. “Prime Time for Transgenerational Defense.” Plant Physiology 158, no. 2 (2012): 545–45. Popova, Evgenya, and Colin J. Barnstable. “Epigenetics Rules.” Journal Of Ocular Biology, Diseases, And Informatics 4, no. 3 (2012): 93–94. doi:10.1007/s12177-012-9088-8. Ramo-Fernández, Laura, Christina Boeck, Alexandra M. Koenig, Katharina Schury, Elisabeth B. Binder, Harald Gündel, Jöerg M. Fegert, Alexander Karabatsiakis, and Iris-Tatjana Kolassa. “The Effects of Childhood Maltreatment on Epigenetic Regulation of Stress-Response Associated Genes: An Intergenerational Approach.” Scientific Reports 9, no. 1 (2019/04/18 2019): 1–12. doi:10.1038/s41598-018-36689-2. Sarkies, Peter. “Molecular Mechanisms of Epigenetic Inheritance: Possible Evolutionary Implications.” Seminars in Cell & Developmental Biology (2019/06/21/ 2019, In Press). doi:https://doi.org/10.1016/j.semcdb.2019.06.005. Schiffers, Katja, Katja Tielbörger, Britta Tietjen, and Florian Jeltsch. “Root Plasticity Buffers Competition among Plants: Theory Meets Experimental Data.” Ecology 92, no. 3 (2011): 610–20. Song, Yuan Yuan, Ren Sen Zeng, Jian Feng Xu, Jun Li, Xiang Shen, and Woldemariam Gebrehiwot Yihdego. “Interplant Communication of Tomato Plants through Underground Common Mycorrhizal Networks.” PloS one 5, no. 10 (2010): e13324, 1–11. doi:10.1371/journal.pone.0013324. Wang, Yang, Qiang Liu, Fuchou Tang, Liying Yan, and Jie Qiao. “Epigenetic Regulation and Risk Factors During the Development of Human Gametes and Early Embryos.” Annual Review of Genomics and Human Genetics 20, no. August (2019/08/30 2019): 21–40. doi:10.1146/annurev-genom-083118-015143. Wohlleben, Peter. The Hidden Life of Trees: What They Feel, How They Communicate— Discoveries from a Secret World. Vancouver, BC Canada: Greystone Books, 2016.

Chapter 5

The Good and the Emergence of Morality in the Flower and Honeybee Mutualism 1 Introduction The first four chapters of this study considered the botany/biology of plants and honeybees. While effort has been made to establish the context for the emergence of ethics through optimization and the flower and honeybee facultative mutualism, a working theory for how morality emerges has not been proposed. This question is approached systematically in this chapter, first through asymmetry of benefit, exploitation, and orientation both flowers and honeybees have towards each other and then through their reciprocal responsibility and hospitality to each other. I then assess these conditions of their facultative mutualism through Kitcher’s pragmatic naturalism which leads directly into a discussion of altruism, and how both flowers and honeybees exhibit altruism in its many forms. I conclude this portion of the discussion considering Singer’s requirements for morality to emerge followed by Michael Ruse and Edward O. Wilson’s origin of morality through the concept of epigenetic rules. The second part of this chapter addresses the naturalistic fallacies and whether or not the efforts of this study can avoid the fallacies of argument and definition both Hume and Moore argue against. Following the naturalistic fallacy discussion, I show how the effort of this chapter leads to a logical conclusion that morality can emerge in nature (without fallacies) as evidenced by the flower and honeybee facultative mutualism. I then summarize the moral ideas that have emerged through this discussion. We begin by considering the asymmetries of the flower and honeybee relationship. 2 Asymmetricity There are three asymmetries associated with the flower and honeybee facul­tative mutualism. The first two are: the benefits they receive and the exploitations they take from each other are both asymmetrical. The honeybee benefits from the food resources (nectar and pollen) that the flower produces for the honeybee. She exploit’s the flower’s energy to do this. The

© Koninklijke Brill NV, Leiden, 2020 | doi:10.1163/9789004428546_007

The Good and the Emergence of Morality

145

flower benefits from the honeybee’s facilitation of her sex act by carrying pollen to other members of her species. She exploits the flight and pollen carrying capabilities of the honeybee to do this. This means that neither competes for the same resources. This is important because it helps minimize the conflicts between the two species. This is also important because the responsibility that I suggest emerges in the flower and honeybee facultative mutualism is reciprocal. Reciprocity of responsibility that emerges in this mutualism does not have to deal with exigencies caused by the need to acquire a common resource. Rather, both are involved not only with advantaging themselves but advantaging the other through processes other than competition. Outside of their mutualism, flowers and honeybees compete with others for other resources, but the mutualism does not face the same challenges. What emerges is a form of commerce where the exchange for one resource is for a different resource. This is not unlike human commerce where money is exchanged for goods or services and vice versa, or grain is exchanged for a horse’s shoeing. The third asymmetry comes from orientation to the world. The flower is oriented centrifugally towards the world, meaning she reaches out to grasp the honeybee through her flower advertisement. The honeybee forager is oriented centripetally towards the earth and to the flower who is always already inextricably tied to where it is located.1 These asymmetric focuses keep the two oriented towards each other always, meaning that they reinforce the mutualism through the continuity of their respective reciprocal but asymmetrical gazes. The flower extends herself into the world through growth, but even farther than the tips of her roots or stems can reach through her advertisement to the honeybee that says, “visit me.” The flower is both centrifugally focused on the world that comes to her and the honeybee who comes to her. The honeybee is centripetally focused on the earth, the middle, from which the plant emerges. Asymmetry of orientation, while it produces a device from which morality can emerge, does not produce the morality of the flower and honeybee mutualism. The asymmetrical orientation strengthens the notion that the optimal actions of both species serve the social group formed by the mutualism. The difference in the flower and honeybee social group compared to many human social groups is that neither compete with the other for the same commodity. This means that any scarcity (not internally generated) is the result of forces individuals do not control (weather, fire, genetics, external competition, predation, etc.) and not the result of competition between flowers and honeybees. There is no need then for strife over scarcity of resource between the two 1  Centrifugal: away from center. Centripetal: towards the center.

146

Chapter 5

partners in this mutualism. The two simply obtain what they can get from the other that the other can give. Asking more from the other would not produce anything more. Since both require the other, it is not logical to intentionally harm the other in order to gain scarce resources the other cannot produce. The honeybee hive is both a dwelling and to dwell. As a dwelling it serves as a city to house all three castes of honeybees. It is therefore a structure. The hive is also existential, where it is to dwell but to dwell is without ends (like the flower) as the hive has no chief executive that oversees its function and viability. All honeybees are existentially tied to dwell in the dwelling that is the hive. They cease to exist without it. While there is much more that can be said about the notion of to dwell as it relates to hive existentiality, it is important to note that the interdependent but independently mobile honeybee foragers are always already oriented towards the middle by virtue of their existential need to dwell.2 Therefore, it comes as no surprise that the honeybee forager is oriented centripetally towards the flower (who is all middle) and the earth to which most flowers are inextricably tied. The flower, because it is all middle and is anchored to the earth, is part of the earth and therefore represents the centripetal center towards which the honeybee forager focuses. Even as the honeybee forager enters the flower, she never loses her centripetal orientation to the world of flowers, because she will leave one flower and seek the next. While she is an independent existent, she is always dependent upon the hive even as her attention is centripetally drawn to the world of the flower. The hive queen and males are centripetally focused on the hive and procreation. Even as the swarm leaves the hive to search for another place to build a hive, the hive soon returns to its centripetal focuses of the hive bees 2  Heidegger explains to dwell with the fourfold, “But ‘on the earth’ already means ‘under the sky.’ Both of these also mean ‘remaining before the divinities’ and include a ‘belonging to men’s being with one another’ ” “Building Dwelling Thinking” in Poetry, Language, Thought (New York: Harper & Row, 1971), 351. The fourfold of dwelling is earthbound, but under the sky and in the realm of heaven and is possible because people are a social species who need and desire to be with others. However, the dwelling is more: “Rather, dwelling itself is always a staying with things. Dwelling, as preserving, keeps the fourfold in that with which mortals stay: in things” ibid., 353. This means that the beehive or the dwelling is a place to stay in and to dwell by existents. Hive species like bees cannot exist long without a hive where young are born, honey is stored, and serves as a shelter from the elements and predators. For humans, to dwell is something learned. Says Heidegger, “The proper dwelling plight lies in this, that mortals ever search anew for the essence of dwelling, that they must ever learn to dwell” ibid., 363. Emphasis in original. Honeybees are born into castes that restrict some degrees of freedom to dwell—but favor others. Honeybees have certain biochemical capabilities that enable them to dwell within the degrees of freedom that the caste allows, but honeybees also can learn which means they can learn how to dwell, particularly when environmental change requires changes in to dwell.

The Good and the Emergence of Morality

147

towards the hive. For foraging workers, and because this study suggests that flowers are members of the honeybee’s family, both the hive and the flowers are always already in centripetal focus. The individual flower and honeybee may only associate with each other for a few days or weeks while the flower is blooming, and the flower is producing adequate nectar. The rest of the year they do not associate with each other. However, this reciprocal—centrifugal/centripetal—orientation towards each other does not change. When the perennial flower blooms the next year, honeybees who were not alive the previous year will forage her. The persistence of this reciprocal orientation I believe serves two functions. First, it maintains the reciprocal focus that the two have on each other even as exigencies of the world try to intrude. Second, that this reciprocal orientation has been instrumental in preserving the focus of the participants on the mutualism and each other. This is likely one reason why the flower and honeybee facultative mutualism has endured for a million years. Even so, both have co-evolved over their million-year association, meaning that behaviors and morphologies changed over time. Their reciprocal orientation towards each other has not changed and this is likely one reason why their mutualism remains strong. What also helps is that they continue to benefit and exploit each other asymmetrically and therefore have not introduced strife that would require a change in how flowers and honeybees relate and treat each other. However, the flower and honeybee are quite different species and it bears looking deeper into how both maintain their asymmetry of orientation, benefit, and exploitation. The plant, as has been explained, is all middle. It has a vascular system that extends from root to leaf tip but no neurological system which means there is no executive that guides the plant’s behavior. However, as this study has explored and Michael Marder reminds us, “As we know, plants do not have a central nervous system but this does not prevent them from sending complex bio-chemical messages, for instance, through their roots and altering their growth patterns as a result.”3 We cannot maintain that a central nervous system makes the sentient animal superior over the plant, because the plant performs similar functions without the benefit from (or burden of) a parallel processing system called the brain and its associated sensory system. The plant can sense, assess, and react to what it senses without having to process this through neurons. That said, there are metaphysical differences between the flower and the honeybee. The plant is all-in where it is. Each organ or system has a function, and communication between functions helps the whole plant deal with exigencies that 3  Michael Marder, Grafts (Minneapolis, Mn.: University of Minnesota Press, 2016), 53.

148

Chapter 5

the world presents to the plant. Plants are the embodiment of situatedness, an other-than-mindful mindfulness that is always already positioned to respond to phenomena as it comes to the plant. This is the nature of the being that is plant, a being in expectation that the world will bring itself to the plant. However, the plant is not in stasis nor is it inert like the rock that lets the world come to it but does nothing about being put into shade or pushed around by water. When the world comes to the plant, the plant uses centrifugal means to embrace the world. When the world brings shade to the plant she extends her branches to reach the sun. When the world refuses to bring water or minerals to the plant, the plant extends her roots deeper into the world to reach her bounty. Plants dominate the landscape where it is possible to germinate, root, and reproduce. Many plants use water or air routines to scatter pollen so that they can reproduce. It is only the angiosperms who have developed a partnership with insects, birds, and other animals to pollinate their flowers directly, rather than only through indirect means like wind or water. The flowering plant has learned how to extend her centrifugal reach into the sky far beyond where her branches touch. She beckons the world to come to her, and the world does in the form of pollinators. While we call plants sessile creatures, they require mobility (air, water, insect) for those who are not self-pollinators in order to reproduce. Bees and other pollinators heed the call of the flower and assist in the mobility she requires to reproduce. Flowers reach out into the world through growth, but they also reach out into the world beyond their morphological capabilities and bring it back to themselves through the assistance of mobile honeybees. Marder gives a metaphysical explanation of the plant and her middleness, “The middle pertains to a non-totalizable synthetic unity, such as the plant, spanning divergent milieus outside of it: the earth and the sky, darkness and light, the moisture of the soil and the dryness of crisp air.”4 This non-totalizable synthetic unity is beyond a whole because the flower creates a gestalt by extending herself into the world beyond her tactile reach, and because she includes within this gestalt the creatures we call pollinators as part of her existential but non-totalizable synthesis. Angiosperms, through color, shape, and scent reach out to pollinators to encourage them to come to the flower in the meadow or into the orchard or tall forest. The honeybee forager, on the other hand, in this reciprocal construct is centripetally oriented towards the flower. Even when the honeybee finds nectar in the tallest tree, she is not anywhere else but proximal to the earth in the form of the tree that is all middle. The honeybee and the flower become one 4  “Vegetal Anti-Metaphysics: Learning from Plants,” Continental Philosophy Review 44, no. 4 (2011): 475.

The Good and the Emergence of Morality

149

with the earth for the shortest moment that the honeybee is inside the flower foraging for her nectar treat. This consummation of process also produces a non-totalizable synthetic unity we call the mutualism. The honeybee forager may not know the metaphysics of communing directly with the earth as she flits from flower to flower, but she is drawn centripetally to the earth and the situatedness of the plants she so covets. Marder suggests a beginning for the discussion of morality in the plant honeybee facultative mutualism, “The ethics of plants, proceeding from their own standpoint, will perennially return to this middle place literally suspended between heaven and earth.”5 It is the flower who arranges the environment for the emergence of morality in her relationship with the honeybee. Her nonmindful mindfulness and attention to the world brings the world towards her as she extends herself into the world. She calls to the honeybee and the honeybee responds. It is this calling out that first brought the apoid wasp to her as the wasp searched for insects who lived among her flowers and leaves. The wasp likely learned that insects found refuge in or about the flowers of angiosperms. It is the flower who likely moved first to establish a relationship with the wasp by developing sticky pollen so that the wasp could help her spread her genes. Flowers and honeybees co-evolved, meaning that they both had to change to accommodate each other. This not only took yeoman genetic and epigenetic engineering, but sustained intentionality from generations of extant agents towards each other to forge this relationship. The relationship between flowers and honeybees has been carefully constructed from the cumulative actions of both towards each other. While the flower and honeybee facultative mutualism took considerable genetic engineering to evolve, it was the constant attention by individual existents to the emerging partnership that has guided genetics towards the mutualism, rather than away from it. Social group-building requires inward focus in order to create durable constructs. The flower honeybee social group has lasted a million years, and the plant pollinator relationship has endured one hundred million years of frequent contingency and existential crises, of which the most profound was likely the dinosaur extinction event sixty-five million years ago. A fundamental question is why, why the emergence of a partnership between a plant and an animal species? Prior to one hundred thirty million years ago plants self-pollinated or used wind or water to reproduce. There are many plant species like conifers and grasses who use some of the same pollination methods as the first vascular plants species that emerged a billion years or so ago on the earth. What changed with the angiosperms? Likely the world 5  Ibid., 475. Emphasis in original.

150

Chapter 5

came to a point where traditional methods of pollination had reached a plateau. Stasis is an anathema to life that continues not only to evolve when the world changes, but also to experiment to see whether a new mutation will provide advantage to the offspring of an extant species. It is perhaps serendipitous that the angiosperm flower emerged to begin a new process of pollination. We don’t know when insects first discovered flowers and flowers began to accommodate the insects, but it was likely early in the emergence of angiosperms. Using fossil evidence, S. C. Capellari, et al., suggest that bees emerged about the same time as the eudicots (pollen with three pores), now a prevalent flowering plant species.6 Which came first, the eudicots or bees, is not known. However, both bees and eudicots have evolved many species during their existential time together. There have long been mutualisms like the eukaryote cell and its mitochondria, the mammal-gut bacteria mutualism, the lichen, and many more. When flowers and honeybees commenced to build a new mutualism, this process was already well-understood in nature. Likely there were indirect mutualisms such as the warning calls of animals during the time of the dinosaurs. Plants and their mycorrhizal fungi were probably in a mutualism construct before the emergence of the first plant-insect mutualism. The honeybee (along with some other pollinating Hymenoptera) are eusocial. The honeybee hive, like the plant is all middle because it has no executive function. Each member of the caste performs its duties without a sovereign or a common distal brain. The queen and her males, the eggs, larvae, and pupae, and many of the immature workers live and work within the hive, existing centripetally, leaving only during the swarm to find another place to exist centripetally. The mature workers who are foragers also exist centripetally, where the locus of their attention is the hive, but the focus of their attention is the world, specifically the flower. The hive, even though its caste members have both minds and vascular systems, is always already all middle. Therefore, plant existentiality that is a colony of processes that produces a non-totalizable synthetic unity, is like the hive which also is a non-totalizable synthetic unity made from inter-dependent but independently mobile cognitive/thinking individuals. We can therefore say that the hive is made from divisible individuals who are indivisible from the hive. Honeybees do not survive long without their hive. The non-totalizable synthetic unities of the hive and the flowering plant are compatible because their dance of their mutuality is asymmetrical (e.g. lead and follow). The sessile flower calls the mobile honeybee towards her 6  S. C. Cappellari, H. Schaefer, and C. C. Davis, “Evolution: Pollen or Pollinators—Which Came First?,” Current Biology 23, no. 8 (2013): R316.

The Good and the Emergence of Morality

151

(shape, color, scent), and the honeybee receives the call and uses her locomotion capabilities to consummate the relationship by flying to the flower. Thus, both centrifugal (flower) and centripetal (honeybee) forces come together at the flower. This dance repeats itself in perpetuity for as long as flowers continue their centrifugal reaching out during the growing season, which, in more-tropical regions may be year-round. The flower and honeybee mutualism is not obligate like the lichen where new existential properties are created from two discrete species. Even so, the flower and honeybee facultative mutualism is intimate, meaning that they both come into direct contact with each other. They do so, however, for different purposes. This asymmetry of the flower and honeybee dance, as has been discussed, produces asymmetrical social commerce where one commodity (pollen and nectar) is exchanged for a completely different other (pollination assistance). In the mutualism, the flower maintains her centrifugal orientation to the honeybee at all times she is available for the honeybee. She maintains her call to other pollinators even as the honeybee forages within her flower. The honeybee continually maintains her centripetal orientation to the flower, because even as she emerges from the one flower, she is focused on finding the next one to enter. The forager’s satisfaction is achieved proximally with the flower while the flower is satisfied only when the distally carried pollen fertilizes another flower of the species. The cycle of flower calling and honeybee entering is maintained until the flower calls off the dance because she is pollinated, or the honeybee can find no more satisfaction by entering the flower. The dance is an apt metaphor, because after a traditional human dance concludes, the partners separate amicably even to move to opposite sides of the room. While the flower and honeybee orientation to each other and the benefits they receive and the exploitations they take from each other are asymmetrical, I suggest this produces an environment for responsibility to emerge in their facultative mutualism. 3 Responsibility Science has considered what flowers and honeybees do and has also spent considerable amount of time studying how they do it. This study has not endeavored to explore in detail the hows of the bio-electro-chemical underpinnings of these behaviors, but much of the research this study cites on flower and honeybee behavior considers the what through the how. The why is often difficult to discern in scientific study, which often leads to speculation, caveat,

152

Chapter 5

and calls for more research. This research needs to be done. However, the aggregate of research done on flower and honeybee behavior is at a point where we can begin to assess the cumulative effects of their behaviors on their respective existences and in their mutualism social group in order to better understand the form of morality that emerges. In one respect, this morality is more enduring than human morality. The flower and honeybee have spent a million years (and longer ancestrally) of evolutionary time and effort to co-evolve themselves into mutualistic partners. Humans have not had to co-evolve but have had only themselves to blame or celebrate for the morality they have emerged over two hundred thousand years modern humans have walked the earth. The morality of the flower and honeybee facultative mutualism may be less complex than human morality, first, because their relationship is limited in time and function. Even so, flowers and honeybees have an important but limited means of communicating with each other as do humans. Third, the co-evolved but asymmetrical interdependence and orientation mitigates competition and strife between the two species. Rather than develop complex rules of conduct against doing harm to the important but different other, the two species have evolved their morality through fewer rules of conduct, but these are rules that are towards advantaging the other as they also advantage themselves. The flower and honeybee facultative mutualism exemplifies Emmanuel Levinas’s ethical mandate that I am infinitely responsible for the other human with whom I interact—without exception. Levinas never tried to apply his theory of responsibility to animals or other life forms.7 To apply Levinasian moral theory of responsibility to the flower and honeybee facultative mutualism, we need to think what responsibility means to these two species. To begin with, Levinas’s notion of responsibility is asymmetrical, meaning I am responsible to the other, but it is the business of the other to decide whether to reciprocate. Says Levinas, “[I] am responsible for the Other without waiting for reciprocity, were I to die for it.”8 This puts the burden on me to be responsible to and for the other. I suggest that the asymmetry of benefit and exploitation in the flower and honeybee facultative mutualism eliminates the need for asymmetrical responsibility. In fact, both flowers and honeybees are responsible for and to each other for all their activities associated with their 7  Levinas saw the face of the other human as the location for human ethics. Levinas commented that he could not know whether the snake had a face and would need to study this idea further: Tamra Wright, Peter Hughes, and Alison Ainley, “The Paradox of Morality: An Interview with Emmanuel Levinas,” in The Provocation of Levinas: Rethinking the Other, ed. Robert Bernasconi, & Wood, David (London and New York: Routledge, 1988), 172. 8  Emmanuel Levinas and Philippe Nemo, Ethics and Infinity: Conversations with Philippe Nemo, trans. Richard Cohen (Pittsburgh, PA: Duquesne University Press, 1985), 98.

The Good and the Emergence of Morality

153

mutualism. Reciprocity is therefore more-or-less guaranteed. That there is reciprocity in the flower and honeybee mutualism eliminates Levinas’s additional requirements that I, the responsible person, must be more passive than passive and must also substitute myself for and to the other even to my own peril. The mutualism social group created by flowers and honeybees considers both the vagaries of nature as well as the metaphysical requirements of each to co-exist with the other in a responsible manner that respects the rights and existentiality of the other. It does not, for example, require the honeybee to enter a flower guarded by a dangerous predator to serve the flower first; nor does it require the flower to continue producing nectar for the honeybee even after she has been pollinated. Ultimately, neither of these actions benefit the mutualism because they have agreed through a million years of co-evolution and cooperative behavior to respect the needs of the other as they exploit the other for what they require. Levinas’s asymmetry, substitution, and passivity are not necessary for these two species who have used other means to mitigate conflict and detrimental behaviors towards the other. Non-exclusivity means that in most flower and pollinator (with notable exceptions) relationships, neither is solely dependent upon the individual other. This means that the failure of one pollinator to pollinate a flower, does not doom the plant’s reproduction. Should a flower cease producing nectar, the honeybee will search for another flower. Asymmetry of orientation, benefit, and exploitation have also produced fair rules of commerce for which there is limited cheating to gain advantage for either species at the expense of the other. Co-evolution means that while either the flower or the honeybee may gain temporary advantage from an asynchronous mutation, the other through behavior, mutation, and epigenetic means can eventually achieve parity with the other. While some flowers who advertise to honeybees do not produce nectar, they do produce pollen which the honeybee also requires. However, there is not much evidence that honeybees cheat with their flower partners, though within the hive there can be cheating. The groundwork for this reciprocal responsibility is laid by their asymmetrical orientation towards each other, the asymmetrical benefits they receive from the other, and asymmetrical exploitations they take from the other. This established foundation is important because it permits both species to behave optimally together in their mutualism. Optimality in this mutualism eliminates the choices of strife, war, and for the most part, cheating. The foundation of mutual cooperation without gaining undue advantage or doing undo harm makes optimal choice simpler, because participants no longer must deal with the consequences of advantage-gaining that can harm self or other. Optimal choices are therefore restricted to those preserving the self from the exigencies of nature that may be associated with the mutualism

154

Chapter 5

but arise outside of it (e.g., predators and competitors at flowers), and those associated with achieving the optimal exploitable result from the benefits the other provides. This study has shown that optimality considering present conditions, in general, is towards maximum entropy production. We also know that forest fires produce moments of extreme entropy production. We can now also suggest that human war and strife are like the forest fire in nature, they engender moments of extreme entropy production. However, what follows forest fires and war is less entropy production, which means that war and forest fires in many respects limit optimality and the resulting entropy that is possible to be produced. Levinas suggests that the peace that follows war is part of the war.9 We see that this is the case in the history of humanity through its conflicts and peace cycles that seem destined to continue for as long as humanity exists. The flower and honeybee facultative mutualism has no war-peace cycle. It has not invented war for its social group and appears to have gone out of its collective way to avoid strife in any form that may affect the other in the construct. What effect on aggregate entropy production has the flower and honeybee facultative mutualism had and how does it compare with humanity and its considerable deviations into war and strife? This question deserves further study. The difference between human activities and flower and honeybee activities towards producing entropy are considered in the final chapter of this study. In summary, while there are similarities in the moral conduct of the flower and honeybee facultative mutualism with that of humans, the basis for flower and honeybee morality is founded on responsibility without the specter of war, strife, undue advantage, or undue harm as significant possibilities. Next, it is important to show how this responsibility process works. 4

Reciprocal Responsibility

This study considers morality through two contexts: descriptive and normative. Descriptive are the optimal behaviors observed in the flower and honeybee facultative mutualism that both species accept, have maintained, and sustain each existentially. Consistency in behavior leads to a normative description of a code of conduct that emerges from the rational application of these optimally derived behaviors. This normative code of conduct I will call reciprocal responsibility. The optimal decisions made by participants in the flower and 9  Emmanuel Levinas, Totality and Infinity: An Essay on Exteriority, trans. Alphonso Lingis (New York, New York: Springer, 1969), 22.

The Good and the Emergence of Morality

155

honeybee facultative mutualism are both rational and optimal, not only for the mutualism construct, but also for the individual participants in the mutualism social group. It is difficult to object to the idea that there are biological and behavioral origins of this code of conduct where optimal rational decisions are regularly and consistently being made to support the mutualism and maintain it even as the world throws existential and evolutionary exigencies at both species. The durability of reciprocal responsibility is such that it has produced a code of conduct that evolves for sure, but likely has remained normatively reciprocally responsible for at least a million years. Humans seem incapable of maintaining reciprocal responsibility for any length of time whether in business, interpersonal, or inter/intra-societal activities. Likely this can be attributed in part to the scarcity of common resources that all humans require or believe they require. Humans, in general, continue to seek advantage, cheat, and kill to benefit self over others for these scarce resources. Flowers and honeybees have asymmetrical needs and exploit each other for asymmetrical benefits. Therefore, scarcity does not flow from the social group they both have created, but from the greater world. Even the flower with a weak nectar production capability, gives what it can. Flowers and honeybees, it seems, have little or no incentive to seek advantage at the expense of the other. The power of reciprocal responsibility, given the co-evolved asymmetrical orientation, advantage, and exploitation that underpins the emergence of this reciprocity, is such that the normative has been maintained even as both species have continued to co-evolve. A million years ago there would have been a different flower mix and honeybees may have both looked and behaved differently, but I suggest that their normative of reciprocal responsibility has not wavered. The success of the flower and honeybee mutualism has produced new species of flowers or migrated older species to new locations as the honeybee has found a home throughout the world. Reciprocal responsibility in the flower and honeybee facultative mutualism is all of these: cooperative, co-optive, and satisficing. Cooperative, meaning they behave to benefit each other. Co-optive, meaning they also exploit each other but not at undue or unexpected expense to the other. Satisficing, meaning that sometimes the best decision is not to engage behaviors towards the mutualism at this time—avoiding the predator inhabited flower; stop producing nectar after the flower is pollinated. Cooperative, co-optive, and satisficing behaviors associated with their mutualism are optimally decided. This optimality is always already contextually maintained because both species asymmetrically exploit and benefit from each other. The mutualism requires both exploitation and benefit which means that reciprocal responsibility becomes the most optimal outcome of most decisions, even when the decision is not to forage or

156

Chapter 5

to end nectar production. Reciprocal responsibility means that I look out for myself while looking out for you and vice versa. If your exploitation of me is of benefit to you, why would you wish to harm me in this effort? Conversely, my exploitation of you is of benefit to me. Neither benefit nor exploitation causes undue harm or excessive benefit to the other in this mutualism. While there can be scarcity locally in the flower and honeybee facultative mutualism (few pollinators; few flowers) there is no advantage to either social participant to try to exploit the other more than what is possible to exploit. Lack is not the result of the mutualism, but of inherited capabilities or outside forces that neither party can directly control. It is in the plant’s best interest to produce as many flowers as optimally possible to attract honeybees for reproduction, and it is in the honeybee’s best interest to produce as many foragers as optimally possible to increase its food production. The decision for both species in actions associated with the mutualism are always contextually the same: the rational decision is optimal if it is judged to be towards reciprocal responsibility to the other of the mutualism in context of what is also desirable for the individual and the species e.g., avoid predators. Levinas’s theory of responsibility requires something to generate the need for responsibility and for human understanding of whom I am to be responsible. Levinas maintains the locus for this origin of my understanding of this responsibility is in the human face. The question next considered, is whether flowers and honeybees have faces in the same way that Levinas envisioned the face. 5

Up from Value

The movement towards the flower and honeybee facultative mutualism first required that each discover value in the other. This necessitates that the other is recognized and understood by the other as an important and valuable other. This recognition must be more than something that brushes by the flower like the cow or the mass of forest that the honeybee cognizes from the ambient optical array she sees before her. Therefore, recognition of otherness is an important requirement for the flower and honeybee mutualism to begin. The flower recognizes the honeybee as an important other from her tactile presence; the honeybee recognizes the flower as an important other by her shape, color, and scent. However, recognition of otherness is only the first step. Establishing the value of the other beyond simple recognition must occur. The apoid wasp needed to discover the protein value of the pollen of the flower as she was

The Good and the Emergence of Morality

157

foraging for insects. Therefore, after recognition of otherness, discovery of value, in the case of flowers and honeybees, requires proximity to the other. Emmanuel Levinas suggests, for example, that humans recognize the face of the other before me, not as a face, but as the metaphysical locus of humanity.10 He even sees God speaking through the face, “There are these two strange things in the face: its extreme frailty—the fact of being without means and, on the other hand, there is authority. It is as if God spoke through the face.”11 Edith Wyschogrod elaborates, “In Levinas’s account, the passive, preoriginary self of ipseity is a living system, one for which not love but a preoriginary openness to the other who cannot be conceptualized is the condition of ethics.”12 This means that (through the recognition of the face) another human stands before me and therefore my obligation to the other is the same as it is to any other human. The other is infinitely other to me and always will be, but the face begins this relationship filled with otherness. We likely can say the same thing about the flower and the honeybee. Both are infinitely different or alterior to each other (even more so than humans to each other) and forever will be, but they need points of recognition to recognize that this is an other who is important to me. Plants needed to develop a ‘face’ that the honeybee forager would recognize as being an other who is valuable to her and, by extension, to her hive. The angiosperm already had the rudiments of the device that would become the recognizable face to the honeybee: the flower. She just needed to mold that face through genetic sculpting to produce the color, shape, and scent that the honeybee would recognize as being the face of this important other. The honeybee, as has also been discussed, can see colors, but her color (yellow, green, blue) and ultraviolet spectrum are limited. The face of the flower cannot exceed the honeybee’s spectral capabilities, or the honeybee will not recognize her, just as we do not recognize chimpanzee faces as human. This facial recognition was necessary to precipitate the centripetal urge of the honeybee to seek and enter the important other. The honeybee also needed to create a face for the flower who cannot see her, but surely the flower can feel the honeybee’s 10  This is one of many places where Levinas discusses the locus of humanity as the face, “By essence the prophetic word responds to the epiphany of the face, doubles all discourse not as a discourse about moral themes, but as an irreducible movement of a discourse which by essence is aroused by the epiphany of the face inasmuch as it attests the presence of the third party, the whole of humanity, in the eyes that look at me” ibid., 213. 11  Robert Bernasconi and David Wood, The Provocation of Levinas: Rethinking the Other (London and New York: Routledge, 1988), 169. 12  Edith Wyschogrod, Crossover Queries: Dwelling with Negatives, Embodying Philosophy’s Others (New York: Fordham University Press, 2006), 188.

158

Chapter 5

body which serve as a face of sorts, the source of recognition that the honeybee is nigh. The construction of the recognizable face that denotes important otherness was still not enough to establish a durable mutualism. The next steps towards constructing an effective mutualism likely took even more time to co-evolve. These included: sticky pollen, the apoid wasp developing a desire for nectar, the change of wasp mouthparts into that of a honeybee, the need for an alternative means to carry necessary pollen because mouthparts had changed to favor nectar, and the reconstruction of the flower to produce nectar and to attach pollen to the hairs that the honeybee evolved to replace mouth parts as a pollen transportation device. However, the flower’s centrifugal orientation and the honeybee’s centripetal orientation have never changed, enabling both to co-evolve in ways that have made their mutualism strong enough to last a million years. As I have said before, this persistency of consistent focus (centripetal and centrifugal) towards the other has helped not only to define and construct the mutualism but sustain it over time. I also suggest that consistency of focus in the mutualism has driven the individual actors towards making optimal decisions associated with the mutualism itself. The foraging honeybee in the meadow is focused centripetally on the flower, and optimal decisions she makes are through this centripetal orientation towards her goal of foraging for her hive. The flower maintains her centrifugal focus towards extending herself into the world to attract pollinators, and all her optimal existential and evolutionary decisions are outwards when the time comes to reproduce. We can say that even when the flowering plant is not in reproductive mode, her centrifugal efforts to grow roots towards nutrients and water, and stems and leaves towards the sun, ultimately is towards readying herself to produce flowers, seeds, and fruits. Because their orientation towards each other has remained in tight focus, both species have evolved to become responsible to and for the other for the asymmetrical existential requirements that are the subject of the mutualism. They first, as Levinas suggests, recognize the face of the other (those flowers that recognize the feel (shape) of the honeybee, and the foraging honeybee the color, shape, and scent of the flower). At the present state of the mutualism this is adequate for both to recognize that this is an important other to me and I must be responsible for this other even as I exploit the other. Unlike Levinas’s notion of responsibility, neither the flower nor honeybee become radically passive or substitute each other for the other.13 Rather, this responsi13  Levinas explains how the other makes me vulnerable, a host as hostage, who, to be responsible, must substitute myself for this other, “Vulnerability, exposure to outrage, to wounding, passivity more passive than all patience, passivity of the accusative form,

The Good and the Emergence of Morality

159

bility to the other is bi-directional, even as the exploitation and benefits to the other are asymmetrical. I maintain that the flower and honeybee mutualism produces responsibility to the other without the additional requirements that Levinas imposed: substitution for the other and radical passivity. Levinas made responsibility to the other unlimited and without exception. The flower stops producing nectar when it needs to; the honeybee stops visiting the flower when it needs to. Responsibility without exception is not necessary for the flower and honeybee facultative mutualism because such an absolute requirement would not lead to optimal decision making that is essential for the facultative mutualism to thrive and continue. Neither the flower or honeybee are passive and neither substitute themselves for the other, but reciprocal responsibility is achieved while conflict is avoided. This is not to suggest that Levinas’ responsibility ethic is wrongheaded, because he was thinking only about human interactions. There is the possibility for conflict between humans who may have similar responsibility needs and existential goals. The flower and honeybee facultative mutualism has evolved in such a way that conflict is avoided because each has asymmetrical needs. As a result, exploitations are also asymmetrical but are optimally derived in a way that no undue harm is done to the other. If responsibility for the other, as Levinas suggests, is an essential element of human morality, then we cannot deny that responsibility in the plant honeybee facultative mutualism is also an essential element of their relationship. If responsibility is essential to morality, then can we say that if flowers and honeybees are acting responsibly towards each other, that they are acting morally in their mutualism construct which this study maintains is a social group as is any human social group? I suggest with Michael Marder that we can.14 There is inter-species communication, rules of conduct, responsible behavior, durability, commerce, and the need of the other for the other which makes the flower and honeybee facultative mutualism a social group. Marder has consistently maintained that the history of philosophy is a history of excluding plants from the mainstream discussion to advantage trauma of accusation suffered by a hostage to the point of persecution, implicating the identity of the hostage who substitutes himself for the others: all this is the self, a defecting or defeat of the ego’s identity. And this, pushed to the limit, is sensibility, sensibility as the subjectivity of the subject. It is a substitution for another, one in the place of another, expiation” Emmanuel Levinas, Otherwise Than Being, trans. Alphonso Lingis (Pittsburg, Pa.: Duquesne University Press, 1974), 15. 14  Marder says, “Ethics as such is an offshoot of plant-thinking. If ethics, understood a la Levinas, is the relation to the other, then it must be rooted in the ontology of vegetal life, heteronomously defined by a striving to alterity” Michael Marder, Plant-Thinking: A Philosophy of Vegetal Life, None (New York: Columbia University Press, 2013), Book, 182.

160

Chapter 5

animals (particularly humans) in the philosophical discourse.15 The flower and honeybee facultative mutualism suggests that the traditional philosophical dis­course that ignores or deprecates the role of plants is simply not tenable. I main­tain that not only does the flower and honeybee facultative mutualism restore the place of plants in the philosophical discourse, responsibility has been fundamental to the establishment of the flower and honeybee facultative mutualism social group that has produced the morality derived from this mutualism. The centrifugal focus of flower intentionality complements the centripetal focus of honeybee foraging intentionality which has led to the development of a durable social group that requires the participation of both creatures to maintain the construct (and sustain each other) and even evolve it over time. 6 Hospitality While Singer maintains that restraint is necessary for the emergence of morality, we see hospitality in the actions of both flowers and honeybees. Both the flower and the honeybee serve as host and guest in their hospitality construct. As the flower and honeybee are reciprocally responsible to the other, they are also reciprocally hospitable to the other. Each is prepared ontologically to accommodate the other’s hospitality as well as exploit the other’s hospitality but through different means. The flower fits the honeybee and the honeybee fits the flower. The worker accommodates the flower’s pollen through hairs to which the pollen sticks when she carries them to the next flower. The flower is constructed to fit the honeybee and is long enough to accommodate her proboscis so that the nectar treat can be harvested. Jacques Derrida maintains that in asymmetrical human hospitality the host is hostage to the guest.16 Kevin D. O’Gorman notes through B. J. Malina that, “The stranger will rarely, if ever, reciprocate hospitality, thus they are forever indebted to the host.”17 In the flower and honeybee facultative mutualism the host is paid back not in kind but with an appropriate reward. The guest not only exploits the hospitality of the host, but she pays back this hospitality in a suitable manner. 15  Ibid., 2. Also see: Marder, “Vegetal Anti-Metaphysics: Learning from Plants.”; “Of Plants, and Other Secrets,” Societies 3, no. 1 (2013). 16  Jacques Derrida, Adieu to Emmanuel Levinas, trans. Pascale-Anne Brault, & Michael Nals (Stanford, CA: Stanford University Press, 1999), 55. 17  Kevin D. O’Gorman, The Origins of Hospitality and Tourism (Oxford, UK: Goodfellow Publishers Limited, 2010), 49.

The Good and the Emergence of Morality

161

The distinction between host and guest loses meaning in this relationship because both the flower and honeybee are guest and host simultaneously even if the reward (pollination or conversion of nectar into food) is asymmetrical in the hospitality event. In human society, the invited guest usually brings a gift to the host, but likely the gift is not commensurable to what the host provides the guest. In the flower and honeybee hospitality event, the value of what the host and guest give to the other cannot be value quantified because they are completely different, like adding apples and oranges. Suffice it to say that over time, the benefit to both is adequate to meet the needs of the individual who is both guest and host. Over time, the cost to both is optimal and within the existential requirements of both as well. In human hospitality there is a host to whom the guest travels. This may be Abraham’s tent in the desert, or the home in the suburb. We may be tempted to describe the fixed location flower as the host and her visitor the honeybee as guest. That would be true if both did not reciprocate hospitality in the event. This is more than hospitality as is typically envisioned in human terms. Rather this is a co-hospitality that is matrixial and may have no direct equivalent in human society. The flower owns its nectar until it gives it away to the foraging honeybee; the honeybee owns its energy until it gives it away by flying from flower to flower and pollenating each. Consider that both the flower and honeybee through their co-hospitality eventually produce acts of creation through food for the honeybee queen and males, and assistance in the sex act for the flower. As has been explored there is both asymmetry and reciprocity in the flower and honeybee relationship. There is asymmetry of ontology, and asymmetry in what each benefits from and exploits the other: food versus sex assist. Finally, their orientation to the world is asymmetrical, with flowers centrifugally and honeybees centripetally focused. This helps to preserve the social construct because neither compete with the other for the same resource. This means that competitive conflict can be avoided and does not even need to be addressed. At the same time, flowers and honeybees are both reciprocally responsible to each other and co-hospitable. Co-hospitality and co-responsibility align because flowers and honeybees benefit and exploit each other. This means that the act of exploitation from one produces an asymmetrical benefit to the other and vice versa. Therefore, each participant in this facultative mutualism must both be responsible and hospitable simultaneously. This approach to co-existence may only be possible for two completely different creatures like flowers and honeybees. While humans may avoid conflict with each other, their needs are symmetrical and inevitably conflict for scarce shared resources

162

Chapter 5

is bound to arise. Not having to deal with conflict caused by competition may be one reason why the flower and honeybee facultative mutualism has persisted for a million years. If we are to locate morality in nature, we must eventually confront the naturalistic fallacies of both Hume and Moore. Before that discussion, it is important to consider naturalism through a pragmatic lens and this requires a review of Kitcher’s pragmatic naturalism and the issue of altruism which is an important issue for morality deriving morality from nature. This will help us prepare arguments for why the flower and honeybee mutualism is towards morality and how this natural emergence of morality does not commit naturalistic fallacies. 7

Pragmatic Naturalism

Philip Kitcher explains that the origin of naturalism comes from Charles Darwin’s strategy, “[f]or explaining facets of the contemporary organic world in terms of the history of life. Pragmatic naturalism proposes that we adopt the same strategy in the case of ethics.”18 Pragmatic naturalism seeks to find the origin of ethics and morality by studying the history of life. If ethics can be associated with the history of life, then ethics and morality are subjects that are not exclusive to humanity. While humans may have been the first to cognitively articulate the concepts of ethics and morality, this does not mean that the antecedents of this abstract idea cannot have originated in other life forms. The effort of pragmatic naturalism is to discover contemporary ethical and moral origins that are relevant to or derived from the actions of otherthan-human life forms. Or, as Kitcher puts it, “([u]nderstand contemporary phenomena in terms of the processes that have given rise to them.)”19 Pragmatic naturalism cites its origin in Darwin’s investigation of the history of life. If we are to proceed down this path, we must consider whether we can discern any evolutionary origins of naturalistic morality. David Sloan Wilson et al., suggest, “Moral systems are a product of natural selection, as surely as immoral acts. Put another way, there is a temptation to act morally and to insist upon moral behavior in others, just as there is a temptation to act immorally.

18  Philip Kitcher, “Naturalistic Ethics without Fallacies,” in Evolution and Morality, ed. James E. Flemming and Sanford Levinson (New York: New York University Press, 2012), 5. Emphasis in original. 19  Ibid., 5.

The Good and the Emergence of Morality

163

Which temptation prevails is likely to be highly context-dependent.”20 Consider the context of human morality. First, humans are social beings. Wilson, et al., are correct to assert, “Human societies around the world are governed by moral systems that classify behaviors into ‘right’ and ‘wrong’ based largely on the criterion of common welfare.”21 Common welfare suggests that morality requires consideration of the group or social group. This means that while there may exist in humanity the temptation to act immorally or for-oneself-only, the social group has developed standards of conduct that consider the welfare of others and has developed coercive means to enforce those standards. Wilson et al., suggest that many have been too quick to dismiss the notion of group selection as part of the process of ethical development. Wilson et al., say, “As Darwin realized, it is hard to explain how moral individuals outcompete immoral individuals within a group, but easy to explain how groups of moral individuals outcompete groups of immoral individuals.”22 Turning to a concept called altruistic punishment through Bowles and Gintis, they explain the process of coercion that may have developed in human societies, “[a] plausible explanation for the evolutionary success of this strategy is that groups with a high fraction of altruistic punishers would have sustained cooperation more successfully than groups with fewer punishers, and so would have prevailed over them.”23 What they are suggesting is that negative feedback for behaviors is central to enforcement of and maintenance of common welfare. Before discussing altruism punishment in more detail, a preliminary discussion of altruism is required. Kitcher calls his naturalism, pragmatic naturalism, consistent with Darwin’s own view that looks at the history of life to discover ethical origins.24 While Kitcher provides a framework, the real work is analyzing flowers and honeybees, not only through their asymmetry, but also their behaviors which means revisiting optimization and mutualism through a behavioral lens. Before we can discuss optimization and mutualism pragmatically, Kitcher gives us some guidance in this regard with his separation of altruism into three categories: biological, psychological, and behavioral.25 These altruistic categories permit us to lift the veil of anthropomorphism that has covered the subject of morality 20  David Sloan Wilson, Eric Dietrich, and Anne B. Clark, “On the Inappropriate Use of the Naturalistic Fallacy in Evolutionary Psychology,” Biology and Philosophy 18, no. 5 (2003): 677. 21  Ibid. 22  Ibid., 679. 23  Ibid. 24  Kitcher, “Naturalistic Ethics without Fallacies,” 5. 25  Ibid., 7.

164

Chapter 5

for so many years to begin to consider it in context of other creatures of nature. Altruism is just what we see in the honeybee hive, in the relationship of some flowering plants towards each other, and in the mutualism itself. We also see what Ernst Fehr and Simon Gachter call altruistic punishment in the honeybee hive which gives us some guidance for how and why honeybees cooperate in their hive.26 8 Altruism 8.1 Sociobiology E. O. Wilson coined the neologism Sociobiology to become a system of study: “Sociobiology is defined as the systematic study of the biological basis of all forms of social behavior, including sexual and parental behavior, in all kinds of organisms, including man.”27 What Wilson wanted to know is how can or does altruism evolve in nature? Peter Singer quotes Wilson on altruism that, “[i]s the central theoretical problem of sociobiology.”28 However, sociobiology has become more than the quest to discover the origins of altruism alone. Kitcher offers a narrow sociobiology definition: Narrow sociobiology is more selective. Its questions are evolutionary questions. So, in posing the question why animals engage in the forms of behavior that they do, narrow sociobiology construes the quest as asking for a specification of the actual workings of evolution: how did the behavior originally evolve? How is it maintained?29 This study, in the narrow sociobiology tradition, has considered the coevolution of both flowers and honeybees that has led to their formation of a facultative mutualism social group. Also, the individual flower and honeybee behavior that has evolved in this mutualism has been considered through the metaphysical lens of asymmetry and the behavioral lens of optimization to consider how the reciprocal responsible and hospitable behaviors of both have been maintained over time. However, the subject of altruism has not yet been addressed. 26  See: Ernst Fehr and Simon Gächter, “Altruistic Punishment in Humans,” Nature 415, no. January (2002). 27  Edward O. Wilson, “What Is Sociobiology?,” Society 15, no. 6 (1978): 10. 28  Peter Singer, The Expanding Circle (Princeton, N.J.: Princeton University Press, 1981, 2011), 5. 29  Philip Kitcher, Vaulting Ambition (Cambridge, Ma.: MIT University Press, 1985), 115. Emphasis in original.

The Good and the Emergence of Morality

165

8.2 Three Forms of Altruism Singer suggests that the three requirements: social group, restraint, and reason are necessary conditions for the emergence of morality, but that altruism must be present to facilitate morality. Therefore, a discussion of altruism is necessary. Peter Singer says that the location of the ethical is in altruism.30 Altruism goes beyond just cooperation, for example, what we see when a flock of geese fly together and change positions, sometimes to lead, and sometimes to follow, that improves long-journey flight sustainability. Altruism requires definition if we are to proceed. Philip Kitcher sees three forms of altruism, biological, psychological, and behavioral. [a] biological altruist is an organism that acts to augment the reproductive success of an other organism at reproductive cost to itself. Psychological altruists, by contrast, are animals with a particular type of structure in their psychological lives: when they come to believe that their actions will have consequences to other animals, they adjust their preferences to align those preferences more closely with those they attribute to the others, and they do so without expectations that their subsequent actions will promote the wishes they previously had. [b]ehavioral altruists are animals who act like psychological altruists: they look as though they are aligning their wishes with those of their beneficiaries, although their reasons for doing so may result from a desire to promote their desired ends (they may be thoroughly Machiavellian).31 Angiosperms, by and large, are not biological altruists with their own and different plant species. Are angiosperms psychological altruists? From root and forest studies we are beginning to see tomato plants use their root systems to warn each other. Many plants release ethylene that alerts others of predation. This is most likely cooperation because neither the warning nor warned plant may adjust its behavior to conform to the other. However, Peter Wohlleben suggests it may be the case that there are some adjustments made by old-growth trees to accommodate newer growth, perhaps at expense to themselves.32 More study must be conducted before we can classify these behaviors as

30  Regarding sociobiology and the problem of altruism, Singer says: “Sociobiology bears on ethics indirectly, through what it says about the development of altruism, rather than by a direct study of ethics. Since it is difficult to decide when a chimpanzee or a gazelle is behaving ethically, this is a wise strategy” Singer, The Expanding Circle, 5. 31  Kitcher, “Naturalistic Ethics without Fallacies,” 7. 32  Peter Wohlleben, The Hidden Life of Trees: What They Feel, How They Communicate— Discoveries from a Secret World (Vancouver, BC Canada: Greystone Books, 2016), 4.

166

Chapter 5

psychologically altruistic. There may be some instances where plants act like behavioral altruists which I will leave to others to research. We know that honeybee workers are biological altruists, not because they have agreed to sterility, but because they are coerced into sterility through the nutritional adjustments made to developing siblings by sister nursery tenders. Honeybee eusocial structure somehow encourages cooperation with others, for others, with reproductive cost to themselves. There are genetic theories why workers agree to live and work eusocially that are beyond the scope of this study. The relationships workers have with other workers in the hive suggest that they regularly operate as psychological altruists. The honeybee that dances a flower patch is bumped by another worker because that worker believes the patch to be substandard. The dancing bee stops her dance to align her preference to the other and does not continue her dance and may even heed the warning of the bump dancer to look for another place to forage. Olav Rueppell, et al., report evidence that injured or sick honeybees can self-remove themselves from the hive.33 Honeybees generally are not behavioral altruists within their hive. However, when honey stores are robust, studies show that guard bees can let up their guard so that they do not attack raiding bees from other hives.34 This inaction appears on the surface to align with the wishes of the other, perhaps the more needy hive, but these guard bees simply are making a pragmatic decision that aggressive hive guarding is not worth the risk and therefore the benefit the lax guarding behavior is principally for the hive that is being raided. I believe that flowers and honeybees have co-evolved a para-psychological altruism in their facultative mutualism. The structure of their asymmetrical mutualism produces alignment by both to meet the needs of the other. Over millions of years, flowers have aligned their flower shapes, color, scent, and nectar capabilities to attract the honeybee and her predecessors who evolved to require pollen and nectar to survive. The honeybee has evolved herself to better perform the pollination process for the flower. Yet altruistic process 33  Their experiment: “We challenged honey bee foragers with prolonged CO2 narcosis or by feeding with the cytostatic drug hydroxyurea. Both treatments resulted in increased mortality but also caused the surviving foragers to abandon their social function and remove themselves from their colony, resulting in altruistic suicide” Olav Rueppell, M. K. Hayworth, and N. P. Ross, “Altruistic Self‐Removal of Health‐Compromised Honey Bee Workers from Their Hive,” Journal Of Evolutionary Biology 23, no. 7 (2010): 1538. 34  Stephen G. Downs and Francis L. W. Ratnieks, “Adaptive Shifts in Honey Bee (Apis Mellifera L.) Guarding Behavior Support Predictions of the Acceptance Threshold Model,” Behavioral Ecology 11, no. 3 (2000): 326.

The Good and the Emergence of Morality

167

is only half of the equation, because the flower must exploit the honeybee’s mobility in order to achieve its mutualism requirement of pollination and the honeybee exploits the nutritional producing capabilities of the flower. This results in para-psychological altruism because the flower and honeybee through their realignment also expect to receive something in return, e.g. Kitcher’s ‘the wishes they previously had.’ Therefore, the flower and honeybee mutualism is somewhere in-between psychological and behavioral altruism as defined because unlike the relaxed guarding in the hive, there is real alignment with the asymmetrical needs of the other. If the needs of flowers and honeybees were symmetrical (like that of the hive that is raided and that of the raiding bee) then we could call the efforts of the mutualism towards altruism behavioral. More study on mutualisms in general is required before we can say whether para-psychological altruism extends to other mutualism constructs. While we see evidence of altruism’s different forms in the flower and honeybee mutualism, there is evidence of altruistic punishment in the honeybee hive. 8.3 Altruistic Punishment Ernst Fehr and Simon Gachter explored why humans cooperate. They cite different theories. The first is, “The theory of kin selection focuses on cooperation among individuals that are genetically closely related.” The second is “[t]heories of direct reciprocity focus on the selfish incentives for cooperation in bilateral long-term interactions.” A third is, “The theories of indirect reciprocity and costly signaling show how cooperation in larger groups can emerge when the cooperators can build a reputation.”35 While each theory can explain some aspects of human cooperation, they do not explain why genetically diverse persons cooperate when actions of the group are not repeatable and generate little reputation as a result. In such circumstances, they ask, what is the mechanism of cooperation?36 They offer punishment as the solution to these gaps in understanding cooperation. They suggest, “If those who free ride on the cooperation of others are punished, cooperation may pay.”37 This may be true, but they ask, who will enact the punishment? They see a second order public-good for punishment: Everybody in the group will be better off if free riding is deterred, but nobody has an incentive to punish the free riders. Thus, the punishment of free riders constitutes a second-order public good. The problem of 35  Fehr and Gächter, “Altruistic Punishment in Humans,” 137. 36  Ibid. 37  Ibid.

168

Chapter 5

second-order public goods can be solved if enough humans have a tendency for altruistic punishment, that is, if they are motivated to punish free riders even though it is costly and yields no material benefits for the punishers.38 In the investment game they constructed, to avoid punishment and gain an optimal reward, members were instructed to act in concert with the group norm. What Fehr and Gachter wanted to discover is: who would punish and why would they punish? The second will be explored first: why actors would act to punish free riders? They surmise that the emotion of those who took it upon themselves to punish the free riders is relevant, “Free riding may cause strong negative emotions among the cooperators and these emotions, in turn, may trigger their willingness to punish the free riders.”39 Their study results suggest, “[t]hat free riding causes strong negative emotions and that most people expect these emotions.”40 Meaning, that free riders know that they generate negative emotions in others. From their study they discovered three things. First, the above average contributors did most of the punishment of free riders—this answers their first question—who the punishers are. Second, the greater the deviation from the norm, the more punishment increased. They say, “This is exactly what would be expected if negative emotions are the proximate cause of the punishment, because negative emotions became more intense as the free rider deviated further from the others’ average investment.”41 Next: Third, if negative emotions cause punishment, the punishment threat is rendered immediately credible because most people understand they trigger strong negative emotions when they free ride. Therefore, we should detect an immediate impact of the punishment opportunity on contributions at the switch points between the punishment and the nopunishment condition. This is what we observed.42 David Hume suggests that human behavior is a product of both passion and reason. The Fehr and Gachter’s study reports that negative emotion arises in those who act according to societal norms when they see that others are 38  Ibid. 39  Ibid., 139. 40  Ibid. 41  Ibid. 42  Ibid.

The Good and the Emergence of Morality

169

deviating from societal norms. Also, that those who deviate from the norm know that they will generate negative emotion in those who act within the norm of the group. We can see from this that negative emotion generation can stir those who experience it to act to bring others into line. Therefore, there is a connection between negative emotion and inflicting punishment. Negative emotion can also be associated with passion that is fundamentally towards some objective or goal or need of the individual and group (e.g. in this game). Negative emotion is not the passion but results from the inability to achieve the passion (goal), for example, because others are not playing by the agreedupon rules. Negative emotion becomes a process that produces punishment as a means for returning the game to where those who deviate are altruistically punished to reform to the norm. The reasoning aspect of Hume’s observation of human behavior likely is contained in the notion that following societal norms or rules is generally good not only for the group but for individuals in the group. What Fehr and Gachter do not explain is why the free riders would free ride in the first place if: 1) they know it will cause negative emotion in others, 2) that they know they may receive punishment as a result. There may be several reasons, including misunderstanding the rules, and the possibility for greater self-advantage of the free ride even with the cost of punishment. More study about the emotional and reason state of the free rider is necessary. Fehr and Gachter conclude: Thus, our evidence suggests that the evolutionary study of human cooperation in large groups of unrelated individuals should include a focus on explaining altruistic punishment. Moreover, because altruistic punishment occurs among genetically unrelated individuals and under conditions that rule out direct reciprocity and reputation formation, the above-mentioned theories do not readily account for altruistic punishment.43 Honeybee hives are made up from closely related individuals (not the unrelated individuals in Fehr and Gachter’s study). However, there is some research that honeybees can experience negative emotions (shaken honeybees).44 We 43  Ibid., 131. 44  For example, “Honeybee perception is highly emotional. This emotionality is manifested by characteristic subjective and objective symptoms. Due to this emotionality the primary form of bee consciousness may reflect different primordial affects type of anxious, fear, rage etc. The JH-3 appears as the modulator of all emotional reactions in honeybees. As result of this, the phenomenon emotional nature of bee perception strongly influences

170

Chapter 5

know also that there is punishment in the form of negative feedback in the hive for those who do not act within the norms required for the hive to function optimally. Negative feedback includes the bump dance, tremble dance, and delayed trophallaxis. In the house-hunting dance, negative feedback comes from the reduced number of visits by other scouts to the location a scout has found. This negative feedback can encourage the scout to search for another location or visit locations that are gaining attention and dancing by other scout bees. What we do not know is whether in the normal course of events where conforming honeybees give punishment in the form of negative feedback, whether the punisher experiences negative emotion when observing this deviation, and whether, like Fehr and Gachter’s study participants, the offending honeybee can expect this negative emotion to occur. Something, however, must trigger the punishing honeybee’s actions that provides negative feedback to the offending worker. That the offending bee quite often ceases the offensive action and moves on to more productive activities suggests perhaps that negative emotion does play a role in both the generation of negative feedback by the punisher and acquiescence to return to the norm by the offending bee. The honeybee hive is relatively strife free. If the free riding of others generates negative emotion in those who do not free ride, what is the emotional state in a social group or group where there is little or no free riding? I ask, does the emotional dial swing from negative emotion to positive emotion when there is no free riding, or is the emotional state of the group where there is no free riding a kind of neutral state, and if so, what is this emotional state? Humans do experience what we call happiness and even euphoria (not drug induced). Is this the result of compliance with societal norms or something different? More research is necessary to gain answers to these questions. As has been discovered, flowering plants can communicate by air released pheromones, and some communicate through mycorrhizal networks in their roots. These are positive warnings against negative results from herbivore its behaviour and behavioural development” Zbigniew Lipiñski, “The Emotional Nature of the Worker Honeybee (Apis Mellifera L.),” Journal of Apicultural Science 50, no. 1 (2006): 58. Also: “We show for the first time that agitated bees are more likely to classify ambiguous stimuli as predicting punishment. Shaken bees also have lower levels of hemolymph dopamine, octopamine, and serotonin. In demonstrating state-dependent modulation of categorization in bees, and thereby a cognitive component of emotion, we show that the bees’ response to a negatively valenced event has more in common with that of vertebrates than previously thought. This finding reinforces the use of cognitive bias as a measure of negative emotional states across species and suggests that honeybees could be regarded as exhibiting emotions” Melissa Bateson et al., “Agitated Honeybees Exhibit Pessimistic Cognitive Biases,” Current Biology 21, no. 12 (2011): 1070.

The Good and the Emergence of Morality

171

infestation or pathogens. Plants of the same species often stop growing roots that encroach on proximal others.45 While we can derive reason from the optimal actions of plants in moving towards water, sun, and other nutrients, can we attribute emotion to the plant? The plant is a middle without ends. It is a vascular system that deploys many of the bio-electrochemical processes that animals do but has no parallel executive neurological function. We may not be able to assign emotion to the plant. However, we can attribute passion to the flower through the efforts she makes to find sunlight, water, nutrients, and reach out to draw in the honeybee to her flower. While passion and emotion appear to be connected, the question that follows is whether emotion is required in all passionate circumstances? I cannot say whether the flowering plant experiences emotion, but I maintain that she is passionate about doing what is necessary to produce the good for herself and her mutualism through her optimal decision-making process. The passions therefore may be augmented by the emotions in creatures that can experience emotion, but the passions exist in creatures regardless of whether they can experience emotion. If there are negative emotions that produce altruistic punishment, what can we say about negative feedback that honeybees receive? The sensory system of the dancing honeybee receives the pinch on the leg from a following forager who does not want her to dance this location. The dancing bee’s brain receives the information that this is a pinch and orders other processes into motion, perhaps to release hormones to stop the sensation and to direct her muscles to stop dancing. It is perhaps possible then that honeybee passions are augmented by emotion caused by this altruistic punishment, but this requires further study. The flowering plant senses the pinch from chewing caterpillars and releases volatile chemicals to make her leaves distasteful. Perhaps she also releases a warning to other flowers through the air or through her roots. While both the pinch on the leg of the honeybee waggle dancer by the follower and the pinch of the caterpillar munching are ‘pinches’, they set into motion different responses. The pinch of the honeybee leg is negative feedback to an action she is performing. The pinch of the caterpillar notifies the plant of predation 45  See: Jeffrey A. Klemens, “Kin Recognition in Plants?,” Biology Letters 4, no. 1 (2008): 58; Meredith L. Biedrzycki et al., “Root Exudates Mediate Kin Recognition in Plants,” Commu­ nicative & Integrative Biology 3, no. 1 (2010); Meredith L. Biedrzycki and Harsh P. Bais, “Kin Recognition in Plants: A Mysterious Behaviour Unsolved,” Journal of Experimental Botany 61, no. 15 (2010); María A. Crepy and Jorge J. Casal, “Photoreceptor-Mediated Kin Recognition in Plants,” New Phytologist 205, no. 1 (2015); Susan A. Dudley, Guillermo P. Murphy, and Amanda L. File, “Kin Recognition and Competition in Plants,” Functional Ecology 27, no. 4 (2013).

172

Chapter 5

so that she can set into motion processes to protect her other leaves and perhaps warn others of encroaching predation. It would be fallacious to call both ‘pinches’ equivalent because both species are not alike and their response to each is quite different. However, both pinches set processes into motion to stop the pinch from recurring. The question is whether there is emotional content that is generated from these two similar stimulus and response situations? Preliminary research on honeybee emotion suggest that there is a possibility that there is. There is not the same research for plants. We must therefore ask what constitutes emotions? Are, for example, emotions feelings? If so, the flower feels the caterpillar, feels the sun or shade, feels water, and feels the honeybee vibrating in her flower. Does feeling require higher orders of consciousness than just stimulus-response which Tulving suggests is the lowest level of consciousness and one that flowers show evidence of? If there are levels of consciousness, then can there also be levels of emotion that are associated with the passions? Or, if plants are not conscious but cognitively nonconscious, can the plant exhibit emotion or is this only possible for those who are cognitively conscious? The question of emotion in plants and honeybees requires further research that is beyond the scope of this study. However, I suggest that the passions are fundamental to both plant and animal life and that even if plants cannot feel or express emotions, this does not hinder flowering plants from making optimal decisions towards the good in their flower and honeybee facultative mutualism. Hume maintains that both the passions and reason have a role in human behavior. Why should this not also be the case for the flowering plant who reacts reasonably and intentionally to the pinch of the chewing caterpillar? Fehr and Gachter suggest that negative emotion is generated when people deviate from a human social group’s norm. First, the person who plays by a social group’s rules experiences negative emotion when others do not play by the rules. Those who deviate know that this negative emotion will be generated. They also suggest that the greater the deviation, the greater will be the generation of negative emotion.46 Perhaps there is emotional value in punishing the free rider because it may dissipate or reduce the negative emotion in the punisher when the free rider comes into compliance. What Fehr and Gachter do not explain is what biological function produces the negative emotion. Their study taught the participants the rules of an investment game they were to follow. If they followed the strategy, they would gain the reward. It was in all participants’ best interests to follow the strategy. The game was not something that had been encoded in their genes nor was it an 46  Fehr and Gächter, “Altruistic Punishment in Humans,” 139.

The Good and the Emergence of Morality

173

evolutionary adaptation. However, what underlies this game is the notion that everyone benefits when everyone plays by the rules of the game. There may be an element of naturalism here—fair rules in a fair game should be followed if everyone is to benefit optimally. The question is whether this general rule ‘fair rules in a fair game’, for example, is learned or innate to the human condition? Free riders reduce optimality and generate negative emotion which can spur human individuals to exact punishment on the free riders in order to bring them into compliance. What this game also suggests is that while humans may have fundamental understandings such as fair rules in a fair game should be followed, there are no dictates as to what game must be played. What are the origins of such common understandings such as fair rules in a fair game should be followed? There are three categories we might consider. The first is a biological origin. The second is that these fair rules all are learned. Third, that the fair rules have been given to us by God or some other supernatural process. Kitcher rejects the notion of supernatural origin. Hume suggests, “Reason is, and ought only to be the slave of the passions, and can never pretend to any other office than to serve and obey them.”47 Therefore, as the product of human desires, moral values (reason) are something that may not be part of the biological condition of the human species but are produced by it. This may mean that moral values are learned. However, as we will soon discover, our cranial capacity may be too small for us to begin with a blank slate where we must learn all the moral constructs we consistently have to negotiate. G. E. Moore suggests that, “[e]thics aims at discovering what are those other properties belonging to all things which are good.”48 However, Moore admonishes us that we cannot conflate good with the good because good is undefinable. We cannot say that nature is inherently good. It is towards the discovering of the origin of those properties of things that produce the good that we will turn and that is towards considering whether there is an evolutionary and thus biological origin of morality for such notions as fair rules in a fair game should be followed or that the process of optimization can produce the good, but is not good in itself. However, we must enter the evolutionary and biological origin of morality with some trepidation because there has been so much push back by those who wield the naturalist fallacy and Hume’s notion that we cannot derive ought exclusively from is as arguments against such an origin. Before venturing into naturalistic fallacies, we will return to Singer’s

47  David Hume, Treatise of Human Nature (Gutenberg.org: Project Gutenberg, 2012), 219. 48  George Edward Moore, Principia Ethica (Cambridge UK: Cambridge University Press, 1903), 10.

174

Chapter 5

three requirements for morality in order to consider them specifically in context of the flower and honeybee facultative mutualism. 9

Singer’s Requirements for Morality to Emerge Applied to Flowers and Honeybees

Altruism may be central to the question of sociobiology and essential for morality. Peter Singer’s explanation of how morality emerges provides a foundation that comports with how both flowers and honeybees behave towards each other and how each species behaves towards its kin, or in the case of plants, also companion plant species that often co-exist together but may not be in their own separate (formal) mutualism. Much of moral discussion begins with human moral development and Singer is no exception. Singer first rejects Jean Jacques Rousseau’s origin of ethics, his theory that humans developed from solitary creatures.49 Singer also rejects the origin of ethics from a divine lawgiver.50 Consulting the fossil record, Singer notes that hominids and proto-humans that evolved into humans were social animals.51 Singer says that the first thing that is required for an ethics to emerge is the social group: Since ethics is a form of social behavior—more than that, no doubt, but that at least—ethics falls within the scope of sociobiology. One might, of course, raise questions about the extent to which ethics has a biological basis; but if the origins of ethics lie in a past which we share with many non-human animals, evolutionary theory and observations of nonhuman social animals should have some bearing on the nature of ethics.52 With this statement, Singer opens the door to the possibility that ethics and morality could emerge in other social animals. Honeybees are considered eusocial insects, meaning they collectively raise young, have intra-species castes, and multiple maturation generations are present in the hive. This suggests that any social norms produced by the hive can be maintained over time as new members of each caste are born and die. As has been previously discussed, evidence exists that when honeybees from one species are deposited 49  Singer, The Expanding Circle, 3. 50  Ibid., 58. 51  Ibid., 4. 52  Ibid., 5.

The Good and the Emergence of Morality

175

and raised in another honeybee species’ hive, they can understand the dance of the other which is different from their ancestral dance. Therefore, intergenerational learning and knowledge transfer is possible in the honeybee. Likely, therefore, there is both a biological and a learning component to honeybee social norms. Flowers may or may not exist in social groups with other flowers or other plants. More study is required to answer that question. This study maintains that the facultative mutualism is a social group for its flower and honeybee members, therefore flowers are members of at least one social group along with their mutualism companions, the honeybees. Singer’s second requirement is restraint towards other members of the group: A social grouping cannot stay together if its members make frequent and unrestrained attacks on one another. Just when a pattern of restraint toward other members of the group becomes a social ethic is hard to say; but ethics probably began in these pre-human patterns of behavior rather than in the deliberate choices of fully fledged, rational human beings.53 I suggest in addition to the group, and restraint, a form of communication is necessary to develop social conventions that can be passed on to others. The biological forms of communication can come from any sensory capability the species has. Honeybee pheromones trigger cognitive and even physical changes in others of the hive. Visual cues may warn or attract another to approach. The bee waggle dance, with its sun-aligned orientation, turns, wing beats, sounds, scents, and touch, is a rather sophisticated form of shared communication. We know that flowers communicate to each other through ethylene emissions and some root networks. While the bump dance means nothing to the flower, and ethylene likely has no meaning for the honeybee, they communicate in their facultative mutualism through a common language that involves vision and tactile recognition. We also have observed that flowers communicate their availability through the shape, color, and scent of their flowers, and honeybees communicate their presence to the flower through tactile means. If communications and sociality are required for the emergence of ethical thinking and behavior, we can heed the words of Ludwig Wittgenstein that there could be no such thing as a private language, “The individual words of this language are to refer to what can only be known to the person speaking; to his immediate private sensations. So another person cannot understand the 53  Ibid., 4.

176

Chapter 5

language.”54 The waggle dance communication is likely both biological and learned. Richard D. Alexander suggests that both genetics and social learning have a part to play in the development of moral rules for a species: [m]orality need not be contrary to natural selection or inconsistent with it but that, at least as practiced and perhaps also as imagined by most, it may instead be a logical outgrowth or extension of the practice of social reciprocity by a complexly social organism which changes as a result of both genetic evolution and cumulative social learning.55 There is relatively little strife in the honeybee hive, which indicates a pattern of restraint among hive members. Alexander Walton and Amy Toth have found that some individual bees are more interactive or engage in more food sharing behavior than others, which they attribute to personality difference.56 However, what is not known is whether this is due to genetic difference or learned behavior.57 The hive is genetically diverse because the queen has mated with up to ten males. Therefore, genetic differences likely have some role to play in personality differences. In some species, flowers of the same species show restraint towards each other by limiting root encroachment. The flower also shows restraint to the honeybee and vice-versa. Flowering plants are not eusocial species. Plants compete for sunlight with other plants, even those of the same species. Underground, roots also must compete for resources. Plants send out roots that can touch others of the same or different species. Root plasticity studies are beginning to suggest, “[t]here is evidence for a number of species that this plasticity allows individuals to avoid overlap of their rhizospheres, hereby buffering competition.”58 On the other hand, there is competition underground for resources, and one study by Liesje Mommer, suggests, at least for two plant species:

54  Ludwig Wittgenstein, Philosophical Investigations, trans. G. E. M. Anscombe (Oxford: Basil Blackwell, 1958), Chapter 243, p. 88e. 55  Richard D. Alexander, “A Biological Interpretation of Moral Systems,” Zygon 20, no. 1 (1985): 5. Emphasis in original. 56  Alexander Walton and Amy L. Toth, “Variation in Individual Worker Honey Bee Behavior Shows Hallmarks of Personality,” Behavioral Ecology and Sociobiology 70, no. 7 (2016): 999. 57  Ibid., 1009. 58  Katja Schiffers et al., “Root Plasticity Buffers Competition among Plants: Theory Meets Experimental Data,” Ecology 92, no. 3 (2011): 611.

The Good and the Emergence of Morality

177

[r]oot responses to nutrient distribution in a competitive environment depend on the competitive strength of the neighbouring species. The foraging response of the superior species (R. palustris) was hardly affected, but that of the inferior species (A. stolonifera) was greatly inhibited and even reversed by competition: instead of proliferating in the nutrient rich patch, it increased root growth and foraging activity in less favourable patches.59 Kevin J. Beiler, et al., cite emerging evidence that the ecology of the forest is somewhat hierarchical but also mutually beneficial, “[w]here large trees served as hubs, with implications for understorey regeneration and functional continuity in the stand.”60 This suggests that there may be a form of plant community in forests and perhaps even in meadows. Root studies suggest that animals and plants forage in similar ways.61 If this is the case, then we can consider foraging morality for both flowers and honeybees using similar terms. There is growing evidence that plants of the same species or even different species of plants that share compatible mycorrhizae may cooperate or at least not compete at the root level. For some researchers, the forest appears to have nodes of old-growth trees that facilitate the controlled growth of other neighboring trees.62 This has led to speculation that there is a form of cooperation at least in some ecologies between plants of the same and even different species. However, this study assumes only that flowering plants show restraint to honeybees. Singer’s third requirement is the capability of reason. We know that honeybees have considerable reasoning capabilities and therefore they meet all three requirements for the emergence of morality—at least at the hive level. Honeybees, however, protect the hive from other hive foragers bent on raiding honey stores in their hive. Guard workers can even fight these other hive-bees to the death. Guarding behavior varies depending upon whether foraging is good or poor. As has been mentioned previously, researchers have found that

59  Liesje Mommer et al., “Interactive Effects of Nutrient Heterogeneity and Competition: Implications for Root Foraging Theory?,” Functional Ecology 26, no. 1 (2012): 66. 60  Kevin J. Beiler et al., “Architecture of the Wood‐Wide Web: Rhizopogon Spp. Genets Link Multiple Douglas‐Fir Cohorts,” New Phytologist 185, no. 2 (2010): 551. 61  Gordon G. McNickle, “Root Foraging Behaviour of Plants: New Theory, New Methods and New Ideas” (University of Alberta, 2011), 2. 62  For example see: Wohlleben, The Hidden Life of Trees: What They Feel, How They Communicate—Discoveries from a Secret World; Colin Tudge, The Tree (New York: Three Rivers Press, 2995).

178

Chapter 5

when nectar resources are high, both raiding and guarding behavior decline.63 Guarding and raiding are risky endeavors and it makes logical sense that more guarding and raiding occur when the hive is most threatened by the lack of forageable material. Flowers use considerable reasoning power to bend towards the sun, to send roots to water and minerals, and to fend off herbivores and pathogens. They use light to set circadian clocks and use duration of sunlight and darkness to plan their photosynthetic processes. Therefore, while we are uncertain that flowers form social groups with other plants, they show restraint to some other plants and to honeybees and they do reason. That they are in a social group with honeybees suggests that plants do meet Singer’s requirements for ethics to emerge, at least in their facultative mutualism construct. However, it is important to understand what constitutes the social group for the flower and honeybee facultative mutualism. The flower and honeybee social group is similar to that of human societies. Flowers require their pollinators and the pollinators require flowers to exist. Humans require other humans. Like humans, flowers and honeybees are co-dependent upon each other. However, only a few flower and insect/bird species are in exclusive relationships. Like human social groups where one moves from one group (family-to-work-to-recreation, etc.) to another in the course of the day, the honeybee forages any flower species that can serve her needs and the non-exclusive flower receives any pollinator who has the capability of spreading her pollen. Second, there is considerable restraint between flowers and honeybees. The honeybee does not damage the flower beyond normal wear and tear, and, for most species, flowering plants do not injure or capture pollinators. Even the Venus Fly Trap has developed a very long stem from which flowers grow that keeps pollinators away from its trap-sprung leaves whose sweet smell attracts non-pollinating insects like flies.64 Third, both species in the mutualism use reason to make decisions even though their morphologies engage different processes that make decisions.

63  Downs and Ratnieks, “Adaptive Shifts in Honey Bee (Apis Mellifera L.) Guarding Behavior Support Predictions of the Acceptance Threshold Model,” 326. 64  “Remarkably, though the Venus Fly Trap’s cage-like leaves exude sweet substances that only a pollinator would love, the plant does not regularly consume its pollinators that consist of, “the sweat bee Augochlorella gratiosa (Halictidae) and the longhorned beetle Typocerus sinuatus (Cerambycidae) had the highest relative abundance, while A. gratiosa and the checkered beetle Trichodes apivorus (Cleridae) carried the largest pollen loads” Elsa Youngsteadt et al., “Venus Flytrap Rarely Traps Its Pollinators,” The American Naturalist 191, no. 4 (2018): 541.

The Good and the Emergence of Morality

179

Though the flower and honeybee facultative mutualism appears to meet Singer’s conditions for how morality can emerge in nature, there may be more than just behavioral reasons to think so. Michael Ruse and Edward O. Wilson suggest there is a biological origin for morality that is constructed from so-called epigenetic rules that are beyond learning and any notion of a supernatural origin for morality. 10 Epigenetic Rules 10.1 Animal Epigenetic Rules As Alex Walter notes, at least since Hume and his ought derived from is admonition, there has been resistance to the notion that there can be a biological or evolutionary origin for morality.65 On the other hand, Michael Ruse and Edward O. Wilson posit that we can avoid anti-naturalism fallacies if we consider morality through a biological and evolutionary lens. Ruse and Wilson suggest three possible origins for morality in humans.66 The first is, “[d]ivinely placed within the brain or else outside the brain awaiting revelation.”67 For this they find no evidence. Even the bible does not offer a divinely implanted ethics. This brings us to the next origin of ethics, the blank slate approach. We are born with no moral instincts or knowledge and must learn it all. For this Ruse and Wilson suggest, “In fact, the blank-slate brain might require a cranial space many times that actually possessed by human beings.”68 Even in the bible with its Sodom and Gomorrah and other conditions of amoral existence, there runs through the lessons evidence of knowledge of good and bad moral behavior. The teachings of the bible are more towards adherence to long-standing moral rules, rather than new invention. Ruse and Wilson explain, “The evidence from both genetic and cognitive studies demonstrates decisively that the human brain is not a tabula rasa.”69 If the blank slate would require much larger cranial capacity than humans possess, then there must be at least some other foundation for moral behavior. This, Ruse and Wilson suggest must be biological, “In 65  Alex Walter, “The Anti-Naturalistic Fallacy: Evolutionary Moral Psychology and the Insistence of Brute Facts,” Evolutionary Psychology 4, no. 1 (2006): 36. 66  Whether these are all the possible origins, I cannot say. However, these three origins cover a lot of proposed theoretical ground plowed by those who are interested in the origin of ethics. 67  Michael Ruse and Edward O. Wilson, “Moral Philosophy as Applied Science,” Philosophy 61, no. 236 (1986): 174. 68  Ibid., 180. 69  Ibid.

180

Chapter 5

short, there appears to be no escape from the biological foundation of mind.”70 If our brain is not a blank slate, what is it about the brain that can produce other than revealed or learned morality? Ruse and Wilson suggest that we as a species must have evolved “epigenetic rules” that are: [g]enetically based processes of development that predispose the individual to adopt one or a few forms of behaviours as opposed to others. The rules are rooted in the physiological processes leading from the genes to thought and action. The empirical heart of our discussion is that we think morally because we are subject to appropriate epigenetic rules. These predispose us to think that certain courses of action are right and certain courses of action are wrong. The rules certainly do not lock people blindly into certain behaviours. But because they give the illusion of objectivity to morality, they lift us above immediate wants to actions which (unknown to us) ultimately serve our genetic best interests.71 The underlying processes of the mind/brain are essentially pre-programmed for basic moral thinking that provide us, in the least, some initial advantage towards continuity that serves our personal best interest as well as that of the human genome. Conceptually this is consistent with other known epigenetic processes that, for example, pass along means for plant offspring to combat pestilence. Ruse and Wilson offer ideas for how these epigenetic rules might have evolved: [e]nsembles of genes have evolved through mutation and selection within an intensely social existence over tens of thousands of years; they prescribe epigenetic rules of mental development peculiar to the human species; under the influence of the rules certain choices are made from among those conceivable and available to the culture; and finally the choices are narrowed and hardened through contractual agreements and sanctification.72 As a social species we likely have evolved our thinking which has included changing the way we act. The Darwinian process of natural selection selects for those who follow the epigenetic rules, in the long run. Likely, this also involves genetic changes that can facilitate the evolving rule-book that humans have 70  Ibid. 71  Ibid. 72  Ibid., 180–81.

The Good and the Emergence of Morality

181

encoded in their brains. For example, we do not deliberate on whether it is right or wrong to avoid hitting a child who crosses in front of our car, we just do. Perhaps these epigenetic rules help us make good decisions quickly when there is not time to deliberate. Ruse and Wilson reviewed an experiment that asked speakers of multiple languages to put their color terms on a chart. Even with the different origins of language, the different speakers located their terms (by shade) with few exceptions in the columns that respond to the four basic colors.73 This does suggest that there are innate processes for identifying colors and shades in humans that are not subject to modification by other thought processes. The eye has developed over millions of years of evolution across the animal kingdom to discern the basic colors and hues (to various degrees of specificity depending upon the species). Therefore, it matters not what we call colors. The physical properties of light do not change and have not changed since the first sun emerged in the infant universe. However, the question is, how do we get to the notion of epigenetic rules for behaviors that are not associated with fundamental physics like light waves? Ruse and Wilson suggest that we have evidence of behavioral constraint in humans: More precisely, whenever development has been investigated with reference to choice under conditions as free as possible of purely experimental influence, subjects automatically favoured certain choices over others. Some of these epigenetic biases are moderate to very strong, as in the case of colour vocabulary. Others are relatively weak. But all are sufficiently marked to exert a detectable influence on cultural evolution.74 If this is the case and we do not have a blank-slate mind, then we do not also have unrestrained free will. Our epigenetic rules caution us from entering Hobbes’ unrestrained state of nature. Ruse and Wilson also suggest, “And the hypothesis of morality as a product of pure culture is refuted by the growing evidence of the co-evolution of genes and culture.”75 The epigenetic rules appear to act as both a foundation for action and as a nagging consciousness that moves one to act more towards what the social group believes is good rather than in another direction. We are at the nascent stage of understanding epigenetics in context of behaviors and we have only limited information about how these rules produce moral distinctions for us to follow. However, 73  Ibid., 182. 74  Ibid. 75  Ibid., 185.

182

Chapter 5

what Ruse and Wilson suggest is that we should stop denying the biological aspect of human behavior and instead increase our investigation of the origins of these epigenetic rules and consider how we apply them in a social context. Towards this end they make a rather strong assertion: We believe that implicit in the scientific interpretation of moral behaviour is a conclusion of central importance to philosophy, namely that there can be no genuinely objective external ethical premises. Everything that we know about the evolutionary process indicates that no such extrasomatic guides exist.76 Therefore, the blank slate and the deity implanted ethics are not plausible. While we evolve ethics through social activities, and there are variations of ethical application in different societies, fundamentally moral behavior emerges from biological processes. If this is the case, right and wrong are foundationally epigenetic rules for which to navigate existence for social species. These are going to vary by species because species have different existential needs as they suggest, “It follows from what we understand in the most general way about organic evolution that ethical premises are likely to differ from one intelligent species to another.”77 However, we must ask whether there can be that much difference in moral behavior that is genetically derived if there are fundamental rules of social existence that are generally attributable to most social life forms? We can only discover answers to these through discerning whether there are common moral practices in social species and what moral practices might differ. We can then compare these to human social practices or between other social species. Ruse and Wilson suggest that we define ethics in this manner, “Let us define ethics in the ordinary sense, as the area of thought and action governed by a sense of obligation—a feeling that there are certain standards one ought to live up to.”78 Therefore, the process of ethics discovery involves considering the origins of ‘a sense of obligation’ and the standards that are promulgated by this sense of obligation. We must also seek answers to these additional questions. How strong is this sense of obligation in a species? What are the standards and how flexible are these? What enforcement mechanisms are built into these epigenetic rules, sense of obligation, and standards (e.g. altruistic punishment)? We know that human and other social societies (like eusocial 76  Ibid., 186. 77  Ibid. 78  Ibid.

The Good and the Emergence of Morality

183

insects) evolve themselves and their behavior over time. While there may be epigenetic rules, a sense of obligation, and standards of behavior, we must also understand how these rules, obligations, and standards can be modified over time. Meaning, what of our own cognitive and social activities can contribute to change? While a blank slate may not be possible, what of ethical behavior is epigenetically derived, and what may come from experience, change, or decision making under uncertainty using such tools as optimization? Humans create new technology that makes old technology obsolete. For example, the automobile replaced the horse and buggy. While the rules of driving (side of road) and who has the right of way at interchapters did not change with the arrival of the car, other rules and practices had to be developed because the new technology required them. However, the fundamental notion that one should not run down the other driver of either the car or the buggy has not changed. Therefore, we may discover epigenetically derived rules, obligations, and standards that are not easily changed because they are fundamental to social existence, regardless of which social species we are speaking about. We also are likely to find that other rules, obligations, and standards are not so fixed, and that other social species may engage in activities like cannibalism that are against human sensibilities in most societies. The workers of the honeybee hive are all daughters of the queen (prior to the swarm) but there may be as many as ten fathers who mated with the queen. We know that workers appear capable of engaging in the waggle dance straight away. We also know that different honeybee species have different dances, but when one worker of a different dancing species is raised in a foreign hive, she can understand this new dance. Therefore, there likely is both a genetic and learning component to the honeybee dance behavior. There likely is a genetic origin for the waggle dance behavior which carries with it the moral responsibility to dance suitable locations and provide negative feedback through the bump dance where necessary. As the workers do not reproduce, and the queen and males do not dance, dancing is may or may not be something that is epigenetically derived in honeybees. However, and this is pure speculation, it may be possible that all honeybees have dancing capabilities but what workers are fed in the nursery epigenetically turns on these dancing genes. As was discussed in the chapter on epigenetics, external environmental and other conditions can produce epigenetic changes. The queen in any hive must be fed, kept warm but not hot, and she produces eggs to meet hive needs. Some years she may not be fed well because there is little nectar that is collected. We do not know whether this can produce epigenetic changes in future generations that might affect worker behavior or capabilities. Nor do we

184

Chapter 5

know whether stresses during development can produce epigenetic changes in workers that may affect their work. However, we can say that the honeybee worker appears born capable of dancing, and if dancing is essential to the hive, we can begin to see that dancing is towards the good and, in general, meets the requirement of Ruse and Wilson that honeybees, like humans, are not born tabula rasa. We know that some flowers can epigenetically pass along to offspring ways of combatting pathogens they have experienced. We know also that some plants of the same species recognize each other’s roots and limit encroachment. There is also evidence from canopy and other research that plant networks tend to optimize their growth patterns that limit competition to some extent. An argument can be made that even though the plant has no separate neurological process, its other processes and organs seem capable of what might be called moral actions. While they both benefit from and exploit each other, we also see that both flowers and honeybees in their facultative mutualism and interactions do not intentionally cause undue harm to the other species. The asymmetrical orientation that each has to the other, even when the honeybee enters the flower, likely mitigates much of the possibility for strife because both species do not compete for the same resources. Certainly, co-evolution has played a part in enabling both species to cooperate with each other. However, each individual plant and each honeybee worker must optimize her behaviors towards the other to achieve their asymmetrical objectives. This is something that individuals must do on their own even though they may bring to this moment a million years or more of co-evolutionary genetic change. Each, for the most part, makes decisions that are optimal for both the individual (and the honeybee worker for her hive) and the mutualism construct. This requires restraint which, in concert with Ruse and Wilson’s other-than-blank-slate approach, appears to be something that has become a natural construct of both species. For example, after pollination, some flowers change color to no longer be attractive to the honeybee. This not only protects the flower from the actions of pollinators who might damage the developing seed, but it also directly helps the honeybee because she will not make the fruitless trip to the flower that has stopped producing nectar. Therefore, even as the flower shuts down her system of attracting honeybees to her, she has evolved to do so in such a way as to advantage the important other to her, the honeybee. Alex Walter reminds us to consider natural facts on their own merit to determine whether they may be ethical facts. It is difficult to look at the flower and honeybee mutualism and not see the emergence of the good from natural facts. However, one must do so with the caveat that all aspects of the flower and honeybee mutualism do not

The Good and the Emergence of Morality

185

produce the good. There are flowering plants that do not produce nectar and the honeybee may take the pollen of one flower and not ever find another of the same species to visit. 10.2 Plant Epigenetic Rules We know that different species of flowers sometimes exist near each other.79 We do not know why they choose to exist near each other. We also know that when their roots encounter others of their own kind, some species tend not to invade the space of the other of their species.80 There may be epigenetic rules for such behaviors; yet again these behaviors may result only from genetic mutation. Whether we can call these behaviors moral is another question. Even if we cannot call these plant actions moral, we can suggest that plants, like animals, may have developed epigenetic rules for existing with other plants in various environs. Towards this end we do know that flowering plants appear to optimize activities such as canopy height and proximity to other plant species that may assist both in optimizing sunlight. Also, a forest without a lot of gaps can facilitate defense against wind.81 If animals have epigenetic rules, then, without further study, we cannot rule out that plants may also have epigenetic rules.82 We have spent considerable time making the argument for the emergence of morality in nature. However, we have not yet made a convincing case that our 79  Anthony Trewavas, Plant Behavior and Intelligence (Oxford, UK: Oxford University Press, 2014), 185. 80  See: Zdenka Babikova et al., “Underground Signals Carried through Common Mycelial Networks Warn Neighbouring Plants of Aphid Attack,” Ecology Letters 16, no. 7 (2013); Yuan Yuan Song et al., “Interplant Communication of Tomato Plants through Underground Common Mycorrhizal Networks,” PloS one 5, no. 10 (2010); Anthony Trewavas, “What Is Plant Behaviour?,” Plant, Cell & Environment 32, no. 6 (2009). 81  See: Cleiton B. Eller et al., “Modelling Tropical Forest Responses to Drought and El Niño with a Stomatal Optimization Model Based on Xylem Hydraulics,” Philosophical Transactions of the Royal Society B: Biological Sciences 373 (2018); Niels P. R. Anten and Heinjo J. During, “Is Analysing the Nitrogen Use at the Plant Canopy Level a Matter of Choosing the Right Optimization Criterion?,” Oecologia 167, no. 2 (2011); Agren Goran I and Franklin Oskar, “Root: Shoot Ratios, Optimization and Nitrogen Productivity,” Annals of Botany 92, no. 6 (2003); Kevin Liu and Tandy Warnow, “Treelength Optimization for Phylogeny Estimation,” PLoS ONE 7, no. 3 (2012); C. Rikard Unelius et al., “Non-Host Volatile Blend Optimization for Forest Protection against the European Spruce Bark Beetle, Ips Typographus,” ibid. 9, no. 1 (2014). 82  Finally, Ruse and Wilson suggest that evolution has prepared us to deal with short-term moral dilemmas, but we have been less than stellar in our longer-term thinking. To demonstrate this problem, we need only turn to the question of human involvement in climate change and our rather slow response to conditions that seem to be changing with some rapidity, Ruse and Wilson, “Moral Philosophy as Applied Science,” 192.

186

Chapter 5

efforts are nothing more than the fallacies that Hume and Moore warned us against committing. It is now appropriate to consider both Hume and Moore’s admonitions and see whether we have both avoided fallacious arguments, and more importantly, embarked upon a better path towards which the discussion of morality and goodness might take us. 11

Naturalistic Fallacies and Naturalistic Facts

11.1 David Hume David Hume maintains that we cannot derive ought exclusively from is. Hume explains this famous assertion: I cannot forbear adding to these reasonings an observation, which may, perhaps, be found of some importance. In every system of morality, which I have hitherto met with, I have always remarked, that the author proceeds for some time in the ordinary way of reasoning, and establishes the being of a God, or makes observations concerning human affairs; when of a sudden I am surprised to find, that instead of the usual copulations of propositions, is, and is not, I meet with no proposition that is not connected with an ought, or an ought not. This change is imperceptible; but is, however, of the last consequence. For as this ought, or ought not, expresses some new relation or affirmation, it is necessary that it should be observed and explained; and at the same time that a reason should be given, for what seems altogether inconceivable, how this new relation can be a deduction from others, which are entirely different from it. But as authors do not commonly use this precaution, I shall presume to recommend it to the readers; and am persuaded, that this small attention would subvert all the vulgar systems of morality, and let us see, that the distinction of vice and virtue is not founded merely on the relations of objects, nor is perceived by reason.83 Therefore, we cannot derive something from an object without it being a condition of that object. As David Sloan Wilson, et al., explain, “Hume claimed that ethical statements cannot be deduced exclusively from factual statements.”84 Exclusively is the equivocation that must be investigated. They explain, “The 83  Hume, Treatise of Human Nature, 248. 84  Wilson, Dietrich, and Clark, “On the Inappropriate Use of the Naturalistic Fallacy in Evolutionary Psychology,” 670. Emphasis in original.

The Good and the Emergence of Morality

187

word ‘exclusively’ is a crucial part of the naturalistic fallacy. If we remove it, the statement ‘ought cannot be derived from is’ implies that the facts of the world have no relevance to ethical conclusions.”85 The effort of this study is to consider how the facts of the world of flowers and honeybees are relevant to morality as we understand it. Oliver Curry suggests that Hume expressed the idea that, “[m]oral values are the product of certain natural human desires. Hume argued that human behaviour is a product of passion and reason.”86 It is the premise of this study that optimization, making the best choice among multiple options, can be studied morally. If we stay with Hume, any ethical conclusion we reach on optimization is not derived exclusively from the factual but requires an intervening premise that is a product of both passion and reason. Hume considers both passion and reason to be integral to human behavior. However, Hume also says, “Reason is, and ought only to be the slave of the passions and can never pretend to any other office than to serve and obey them.”87 We need some impetus in order to make rational decisions. Simon Blackburn calls these “appetitive states.”88 These appetitive states include, “[a]ppetites themselves, needs, desires, cares and concerns, attitudes, and emotions.”89 These appetitive states are juxtaposed against the doxastic states, “These are the states of persons representing the world to themselves, and forming or recognizing valid or probable relations of implication between their representations.”90 An urgent question is whether we can attribute both passion and reason to other-than-human behaviors, specifically those of flowers and honeybees. This will be discussed later in this chapter. First, however, can we develop a syllogism whereby ought is not entirely derived from is? Wilson et al., provide a persuasive example of a deductively invalid ethical statement and how it can be corrected. The deductively invalid statement first: – Torturing people for fun causes great suffering (factual premise). – Torturing people for fun is wrong (ethical conclusion).91

85  Ibid., 671. 86  Oliver Curry, “Who’s Afraid of the Naturalistic Fallacy?,” Journal of Evolutionary Psychology 4, no. 1 (2006): 234. 87  Hume, Treatise of Human Nature, 219. 88  Simon Blackburn, “Précis of Ruling Passions,” review of Ruling Passions, Simon Blackburn, Philosophy and Phenomenological Research 65, no. 1 (2002): 122. 89  Ibid. 90  Ibid. 91  Wilson, Dietrich, and Clark, “On the Inappropriate Use of the Naturalistic Fallacy in Evolutionary Psychology,” 670.

188

Chapter 5

This is Hume’s objection that we cannot derive the ethical conclusion from the factual without an intervening statement, the ethical premise. This revision makes the statement deductively valid: – Torturing people for fun causes great suffering (factual premise). – It is wrong to cause great suffering (ethical premise). – Torturing people for fun is wrong (ethical conclusion).92 We can apply this reasoning to optimization. First, the invalid deductive statement: – Optimization is choosing the best decision among many options (factual premise). – Choosing the best decision among many options is a good thing to do (ethical conclusion). Now the valid deductive statement where an ethical premise is added. – Optimization is choosing the best decision among many options (factual premise). – It is good to choose the best decision among many options (ethical premise). – Choosing the best decision among many options is a good thing to do (ethical conclusion). In both examples, the ethical premise is not a condition of the factual but is outside the factual. However, Hume only launches this discussion. 11.2 G. E. Moore G. E. Moore goes beyond Hume to formulate what he calls the ‘naturalistic fallacy’. Moore was not against the emergence of morality in nature per se, but he did not like the way that naturalism proponents argued for naturalism. Their argument, he says can be expressed as, “ ‘ You are to do this, because most people use a certain word to denote conduct such as this.’ ”93 Moore’s objection to this approach is that, for example, the word pleasure is defined by itself because there is no intervening definition of good that can show why the word

92  Ibid., 671. 93  Moore, Principia Ethica, 12.

The Good and the Emergence of Morality

189

pleasure is good.94 Moore says that, “But if he confuses ‘good’ which is not in the same sense a natural object, with a natural object whatever, then there is a reason for calling that a naturalistic fallacy.”95 The good of the decision is not contained within the process (object) called optimization. It is the result of the decision constructed from Humean passion and reason both which are outside the object (the decision). However Moore also states that, “I now am insisting that good is undefinable.”96 He clarifies this with, “I do not mean to say that the good, that which is good, is thus indefinable; if I did think so, I should not be writing on Ethics, for my main object is to help towards discovering that definition.”97 The good, is different from good he suggests, “As it is, I believe the good to be definable; and yet I still say that good itself is indefinable.”98 Therefore, the good of something or some action may be definable but the word good alone as an unmodified noun is not definable. Even so, the good does not belong to the object. The facts of the decision speak for themselves without modification. The honeybee flew towards the flower is a simple statement. We cannot ascribe goodness to this action without an intervening thought as to why. Moore suggests that there are two alternatives if his premise that good is not definable is wrong: In fact, if it is not the case that good denotes something simple and indefinable, only two alternatives are possible: either it is a complex, a given whole, about the correct analysis of which there may be disagreement; or else it means nothing at all, and there is no such subject as Ethics.99 The alternatives to his proposition are good is either all or nothing. The all can be taken to mean that all of nature is good even though we may disagree on the particulars. Or, if there is no subject as ethics then the good has no meaning. Moore finds neither of these alternatives acceptable. The good, as Moore suggests, if it is to apply to the object e.g. the decision, that decision must always be good. For example, I make the statement nectar is good for the honeybee. However, rotten nectar is never good. Nectar that contains pesticides is not good. Therefore, we cannot say that nectar is always good. We know that the same decision made in other circumstances may not 94  Ibid. 95  Ibid., 13. 96  Ibid., 9. 97  Ibid., 8–9. 98  Ibid., 9. Emphasis in original. 99  Ibid., 15.

190

Chapter 5

be good and may be deleterious to the decider. The honeybee who decides to fly towards the flower guarded by a predator may not ultimately produce the good for the honeybee. More fundamentally are the questions of cause and effect when something is declared to be good. Moore explains: Whenever we judge that a thing is ‘good as a means’, we are making a judgment with regard to its causal relations: we judge both that it will have a particular kind of effect, and that that effect will be good in itself. But to find causal judgments that are universally true is notoriously a matter of extreme difficulty.100 Cause and effect therefore also must be good and where can we find such universality either in nature or in ethics? We have considered the proposition that nectar is always good and found it lacking. Moore suggests: Hence we can never be entitled to more than a generalisation to a proposition of the form: ‘This result generally follows this kind of action’; and even this generalisation will only be true, if the circumstances under which the action occurs are generally the same.101 We have difficulty ascribing goodness to nectar in general because there are circumstances where nectar is not good. What Moore does not want us to do is to try to develop generalized definitions of the good but consider the individual judgements and actions in context of the circumstances where the action takes place to discover the good or not. This Moore states rather succinctly as: With regard then to ethical judgments which assert that a certain kind of action is good as a means to a certain kind of effect, none will be universally true; and many, though generally true at one period, will be generally false at others.102 The question we now must face is whether optimization, the choice of the best option in a given circumstance is good in itself, or does this assertion run afoul of the naturalistic fallacy? Moore provides guidance:

100  Ibid., 22. Emphasis in original. 101  Ibid. Emphasis in original. 102  Ibid. Emphasis in original.

The Good and the Emergence of Morality

191

In other words, to judge that an action is generally a means to good is to judge not only that it generally does some good, but that it generally does the greatest good of which the circumstances admit. In this respect ethical judgments about the effects of action involve a difficulty and a complication far greater than that involved in the establishment of scientific laws. For the latter we need only consider a single effect; for the former it is essential to consider not only this, but the effects of that effect, and so on as far as our view into the future can reach. It is, indeed, obvious that our view can never reach far enough for us to be certain that any action will produce the best possible effects. We must be content, if the greatest possible balance of good seems to be produced within a limited period.103 I suggest that Moore would not have a problem with us saying that optimization is a process for assessing the possibilities for the good in a given circumstance (even if this is limited only to a narrow period of time.) John Teehan and Chris DiCarlo offer such an affirmation: In order to resolve a problematic situation, to make a moral judgment, we need to have a clear grasp of the situation at hand and the possible consequences of various options. Whatever contributes to our understanding of the situation, contributes to our judgment of what we may construe as the good in that situation.104 However, if we go further and suggest that optimization is inherently good, we commit the naturalistic fallacy. Also, with Moore, we cannot project into the distant future whether what looks like was towards the good today, will, in the long run, produce the good. We know that even the best decision may not in the long run produce the good as an intelligent being may define it. Teehan and DiCarlo summarize the lesson of the naturalistic fallacy: The deeper lesson of the Naturalistic Fallacy is that ethics is not about identifying pre-existing moral definitions. It is, instead, an ongoing process of deliberation concerning what is right/good to do.105

103  Ibid., 23. Emphasis in original. 104  John Teehan and Christopher DiCarlo, “On the Naturalistic Fallacy: A Conceptual Basis for Evolutionary Ethics,” Evolutionary Psychology 2, no. 1 (2004): 43. 105  Ibid.

192

Chapter 5

Morality therefore is a deliberation. Optimization is inherently a deliberation over available options. Moore explains the aim of ethics in more detail: And it is a fact, that Ethics aims at discovering what are those other properties belonging to all things which are good. But far too many philosophers have thought that when they named those other properties they were actually defining good; that these properties, in fact, were simply not other, but absolutely and entirely the same with goodness. This view I propose to call the ‘naturalistic fallacy’ and of it I shall now endeavour to dispose.106 We cannot define good. Moore, however, suggests that rather than try to define good, we remain open to the good as it may unfold, “But if we recognise that, so far as the meaning of good goes, anything whatever may be good, we start with a much more open mind.”107 Therefore, we can consider the process of optimization in relation to the good for a specific situation, but we cannot call the process as being good in itself. The good is indeed a difficult subject because it requires knowing not only the circumstances in which the good may be thought, but the object that is a subject in these circumstances. Good is indefinable because it is subjective, but the good of a decision can be deliberated based upon the circumstances in which the decision was made. Therefore, our syllogism: – Optimization is choosing the best decision among many options (factual premise). – It is good to choose the best decision among many options (ethical premise) – Choosing the best decision among many options is a good thing to do (ethical conclusion). Is a valid deduction and it also does not commit Hume’s prohibition of exclusively relating the ought (ethical conclusion) with the is (the factual premise) because it has an intervening ethical premise. The question is whether this syllogism commits the naturalistic fallacy. Can we say that nature, natural selection, and evolution are good and therefore anything nature does which we might find inherently bad is good? Meaning, are we maintaining what ought to be rather than exactly what things are? Goodness is indefinable. 106  Moore, Principia Ethica, 10. 107  Ibid., 20.

The Good and the Emergence of Morality

193

What we need are examples from which we can derive the good. We can find the good in the choosing of the best, but we cannot say that the choice that is ultimately chosen through this process is always good, nor can we say that the process of optimization is always good. The best is relative, and we cannot give the quality of goodness to best because best is not only circumstantial but also best may ultimately not be good to the decision maker or others. The optimization process leading up to the choosing, based upon circumstances of the moment, however, contains qualities of goodness even though we cannot say that the process is or what is chosen is in fact good. However, after the decision has been made, we can say that the good derived from this decision is that it prevented the creature from suffering harm. That the decision is good for the creature existentially does not mean we can attribute the same quality of goodness to the object called the decision. What we have done in this syllogism, as Moore has admonished us not to do, is to describe how things ought to be rather than give an example that would show how the good is derived from the decision. Ultimately, I take this to mean that we cannot derive morality from evolution or biology directly without committing the naturalistic fallacy. We must use specific examples to derive things like the good, happiness, and other qualities. If we cannot derive the good from nature itself, we might turn either to the spiritual or suggest that humans and other beings begin with a blank slate and develop ethical constructs through experience alone. Both alternatives, as we have seen, have their own problems and produce fallacies of their own. What I suggest is that we need to better understand the processes of evolution, natural selection, and biology before rejecting naturalism. We need to consider naturalism, not through the lens where there is only good in nature, and that exigencies like diseases that kill infants are inherently good because they produce stronger people in the long run. We know, for example, that the optimization process may be towards the good, but ultimately may not produce the good. The process is not good but is in search of the good as it relates to the individual decision maker. I suggest that with flowers and honeybees, optimal decisions are in general made that are towards supporting the mutualism construct which means that ultimately, they may produce the good for both species—then again, they may produce good for one participant and not the other. The question that I propose we ask is: can we use science to discover just what in nature can be considered through the lens of the good without categorically ascribing nature or the process (e.g. optimization) as good in itself? I suggest that this study has uncovered aspects of behaviors towards the good in the flower and honeybee facultative mutualism that are difficult to deny as being towards the good or as being natural evidence of moral behavior that is

194

Chapter 5

towards the good. At the same time, with Moore, we should remain open to the possibilities of the good emerging from other aspects of nature without precondition or assumption of the answer. That said, there is a very real possibility that an anti-naturalism fallacy has emerged from the discourse on the naturalistic fallacy. 11.3 An Anti-Naturalistic Fallacy David Sloan Wilson et al., suggest, “[t]he facts of the world do have ethical implications, which the naturalistic fallacy was never intended to deny.”108 Alex Walter maintains that there is an anti-naturalistic fallacy that suggests that there are no natural facts that are of interest to the ethical discourse. His refutation of this fallacy begins with, “[w]e must recognize that while not all natural facts are relevant to ethical or moral discourse, all facts that are relevant to ethical and moral discourse will nonetheless be natural facts. To hold that values are non-natural facts is to commit the anti-naturalistic fallacy.”109 We have already seen that if we correct the ought derived from is fallacy using an intermediate statement of ethical value, we are no longer making a fallacious argument. That only corrects the argument. Whether natural facts are relevant to ethical discourse is the subject of Walter’s critique of the antinaturalistic fallacy. Walter cites many researchers who use the naturalistic fallacy and Hume’s ought/is dictum, “[f]or the purpose of drawing limits around the scope of scientific inquiry into ethics and morality.”110 Walter says, “Moore believes there is one metaphysical entity that does qualify as the ‘good.’ ”111 Walter explains: I believe that once evolutionists understand that proponents of the naturalistic fallacy are committed to ethical objectivism—which entails that values are supernatural facts, they will have no more fear of the ethical relevance of brute facts than they fear that creationists will successfully argue that the universe was created by divine providence.112 Walter maintains that rather than embrace the naturalistic fallacy, we should critique it. The question that Walter wants us to consider is whether values 108  Wilson, Dietrich, and Clark, “On the Inappropriate Use of the Naturalistic Fallacy in Evolutionary Psychology,” 670. 109  Walter, “The Anti-Naturalistic Fallacy: Evolutionary Moral Psychology and the Insistence of Brute Facts,” 34–35. 110  Ibid., 33. 111  Ibid., 34. 112  Ibid.

The Good and the Emergence of Morality

195

are supernatural facts. If good cannot be defined and is not inherent in, for example, the word nectar, then it must be something that is produced outside the subject—that Moore also wanted to emphasize. Walter suggests this leads towards a divine origin of morality. Rather than divine origin, this study suggests that there is growing evidence that the flower and honeybee natural facultative mutualism is capable of not only searching for the good but emerging the good out of both passion and reason that both employ in engaging their social group. I agree with Walter that we must begin with the recognition that all facts are natural but not all natural facts, “[a]re relevant to ethical or moral discourse.”113 What must be searched for are the facts that lead us towards what we might later call the good. I believe we can agree with Moore that neither flowers nor honeybees are inherently good. However, our task, as was Moore’s, is to discover the origin of the good in nature. In this study the way towards recognizing the good begins with the facultative mutualism that both species have co-evolved over at least a million years. This social group shows evidence of the good because there is restraint towards the other resulting from the need (Hume’s passions) to both benefit from and exploit the other. This passion, though asymmetrical, has been maintained over time and perhaps for the duration of the facultative mutualism. The evidence of this natural fact of restraint resulting from passions, is missing something and that is Humean reason. Reason, in context with flowers and honeybees is associated with optimal decision making by both species that is, in general, towards the benefits and exploitation passions associated with the social group or facultative mutualism. Optimal decisionmaking is a process that involves both the passion and reason of the optimizer. We know that, in general, flowers and honeybees demonstrate that they are reciprocally responsible and reciprocally hospitable to each other in association with their need to exploit and benefit the other. Good is not inherent in the flower and honeybee social group, restraint, reciprocal responsibility, or optimization. However, it is difficult to suggest that the good does not emerge from their combined efforts and effects. At some point, in order to find the good, we need to agree on certain natural facts that are associated with producing goodness. John Dewey offers guidance towards this effort. He says, “But morals based upon concerns with facts and deriving guidance from knowledge of them would at least locate the points of effective endeavor and would focus available resources upon them.”114 This is what this study has done without resorting to whether such facts produce eudaimonia, happiness, hedonism or 113  Ibid. 114  John Dewey, Human Nature and Conduct (New York: Henry Holt and Co., 1922), 12.

196

Chapter 5

some other traditional moral end. By focusing our efforts on ‘points of effective endeavor’ we can begin to suggest, as Dewey does next, that morality likely has its origin in nature, “A morals based on the study of human nature instead of upon disregard for it would find the facts of man continuous with those of the rest of nature and would thereby ally ethics with physics and biology.”115 This study offers the behaviors of flowers and honeybees in their facultative mutualism towards engaging the critique of the naturalistic fallacy that some maintain rejects the notion that morality can emerge in nature. We have seen that both flowers and honeybees are reciprocally responsible and hospitable to each other in their mutualism social group. This reciprocal responsibility is maintained by natural acts of both species that support the mutualism in this reciprocal responsibility/hospitality continuum. Reciprocal responsibility and hospitality are maintained by behavior, and as has also been discussed in this study, through the combined efforts of both species to make optimal decisions in this regard. A discussion of how optimization can be thought of as being toward the good without engaging the naturalistic fallacy will bring us towards a basis for establishing evidence of morality in nature. 11.4 Optimization The process of optimization can produce the good. However, we have clarified that statement with the caveat that optimization itself cannot be defined as good in itself—always good. Optimization therefore can be expressed as towards the good. The decisions that the optimization process make produce behaviors that can be analyzed for goodness. Creatures like flowers and honeybees ‘naturally’ optimize. Therefore, we must ask what is the origin of optimization if it is not through some natural bio-evolutionary process? From birth many creatures must be able to assess their environment and make decisions quickly in order to survive to reproduce. Even human infants are born with a startle response to loud noises or falling that is not learned. If there are processes of assessment and reaction that are bio-evolutionary, can we consider such processes like optimization that may or may not be learned through terms that are associated with bio-evolution without committing the naturalistic fallacy? I suggest that we can if we avoid the notion of generalizations such as optimization is good, and optimization always produces the good. The process of optimization considers circumstances based upon creature needs. The needs (Humean passions) drive reason, both which flowers and honeybees generally use towards deriving an optimal decision. What is important to 115  Ibid.

The Good and the Emergence of Morality

197

learn is what of this process emerges from bio-evolution processes and what is learned? Naturalism suggests that everything results from natural, not spiritual properties. How can we define the good in natural processes without also calling the processes good as Moore admonishes? Philip Kitcher proposes a pragmatic naturalism that we can consult the history of life to discover processes that have given rise to current practices such as moral actions and perhaps even the optimization process. I suggest that this is not at odds with Moore who made it clear that we be observers of the world first before judging any morality or lack thereof that emerges. 12

Flower and Honeybee Oughts and Obligates

Flowers grow when they need to and stop growing, even to discard older growth, based upon external stimuli such as sun, water, the presence of minerals, and the vibration of the honeybee. It seems that flowers are obeying certain oughts whenever presented with stimuli. Whether these oughts are genetic, epigenetic, or learned, they drive the flower towards making optimal decisions. The obligation to grow when circumstances warrant seems fundamental to the existential nature of the plant. The variable is where and how and that is left to the flower to decide based upon the internal and external circumstances the flower finds herself. What are the obligates of honeybees? While we cannot read the content of the dancing honeybee’s mind, we can investigate the changes this dance generates in the behavior of other foragers in the hive. Researchers have found that the dances (generally) generate specific behaviors for those foragers who observe and follow the dancer’s instructions.116 Therefore, we can translate honeybee worker communication into human language for comparative purposes. If we can also investigate whether there are 116  See: Parry M. Kietzman and P. Kirk Visscher, “The Anti-Waggle Dance: Use of the Stop Signal as Negative Feedback,” Ecology and Evolution 3 (2015); Wolfgang H. Kirchner, “Vibrational Signals in the Tremble Dance of the Honeybee, Apis Mellifera,” Behavioral Ecology and Sociobiology 33, no. 3 (1993); Fernando Wario et al., “Automatic Methods for Long-Term Tracking and the Detection and Decoding of Communication Dances in Honeybees,” Frontiers in Ecology and Evolution 3, no. 103 (2015); Francis L. W. Ratnieks and Kyle Shackleton, “Does the Waggle Dance Help Honey Bees to Forage at Greater Distances Than Expected for Their Body Size?,” ibid., no. 31; Madeleine Beekman et al., “Honeybee Linguistics—a Comparative Analysis of the Waggle Dance among Species of Apis,” ibid., no. 11; Roger Schürch, Magaret J. Couvillon, and Madeleine Beekman, “Ballroom Biology: Recent Insights into Honey Bee Waggle Dance Communications,” ibid. 2016, no. February (2016).

198

Chapter 5

consistent biochemical changes that occur in honeybees who make optimal decisions on whether to follow the dancing bee or not, we may discover that there is an emotional connection to such decision making in addition to the logical approach it is believed some foragers make which may be translated as, “Well she does sound like she found a promising foraging site. I will keep her advice in mind, but I have a better one. If when I get there it should be overrun by competitors or predators, I will follow her advice.” Or, for another forager, “I am not presently employed, and she sure has found a good foraging site. I will follow her advice because I really need to get back to work.” For both workers the nagging obligation is that they must both get back out and forage for resources for the hive—this is the right thing to do. However, what is right depends upon circumstances and personal knowledge of where the best resources are. Therefore, the obligate is “I should forage” and the variable is where. If we can extend Ruse and Wilson’s epigenetic rules paradigm to honeybees, there is something (emotion perhaps; passions for sure) that reveals to the forager the obligation to forage. This obligation only generates attention to the behavior she needs to perform to meet this obligation—forage. The obligation, however, does not contain content for how to go about fulfilling this obligation for the hive. The obligation says only: Forage. She must use other faculties of observation, communication, memory, and optimal choice to find the best way to fulfill the obligation through behaviors she determines are best. In other words, the ought to forage (the nagging obligation) is derived from the fundamental epigenetic ethical rules that say that the good usually comes from foraging. The ought (I ought to forage) emerges from the is (the biological and/or epigenetic rule that says I should forage). However, what is not derived from either the ought or is, is the method she will choose (logically and within her capabilities) to fulfill her obligation. We therefore must ask whether this biological or epigenetic origin is at odds with philosophical understanding that began with David Hume who suggests that, ‘we cannot exclusively derive ought from is.’ The later G. E. Moore further refines Hume’s maxim and calls the problem the naturalistic fallacy. If we want to go down the path of Ruse and Wilson, we must first consider whether the biological origin of morality violates Hume’s admonition. I suggest that it does not. My reasoning is simple. The key term in Hume’s maxim is exclusively: ‘One cannot exclusively derive ought from is’. The premise of Ruse and Wilson is that the ought for ethical behavior is socially constructed and then imprinted genetically. This becomes an epigenetic rule. For honeybee foragers, it is ‘I should forage.’ What good means, I have agreed with Moore, is undefinable. However, we cannot deny that the good of foraging is that it benefits the hive. Therefore, the obligate ‘I should forage’ serves as the ought in Hume’s admonition. What

The Good and the Emergence of Morality

199

Ruse and Wilson see in humanity is that emotions that derive obligation produce the ought directly from the epigenetic biological rule. However, the ought cannot be exclusively derived from the is because neither give us the answer how exactly to behave in the circumstance or the honeybee any direction for where to forage in the present-day meadow. I ought to forage is a given from the epigenetic rule: forage. How and where I should forage is not given in this ought from is. The act of foraging is both biological (proboscis action) and cognitive, (where to stick the proboscis in the flower). Where to forage is subject to both individual knowledge and knowledge that the forager derives from others. Therefore, because she reasons, her ought is not derived exclusively from is: she has a choice. She is not always already good (the naturalistic fallacy) because she can make mistakes. She tries to make the ‘best’ and ‘most optimal’ decision about where to forage based upon knowledge which she reasons into optimal behavior. However, what began this process of engaging in foraging behavior is the epigenetic rule that says I should forage. What we do not know is how the obligate to forage is generated in the honeybee. Is it through some emotional process we have not discovered, or do honeybees and perhaps other invertebrates employ something other than emotion that we have not discovered? I don’t propose to answer these questions now. They require further study. Where I agree with Ruse and Wilson is that it would be best to lay aside the naturalistic fallacy and pursue studies that investigate whether there is in fact an evolutionary biological origin for ethics. If after suitable study we can find no evolutionary biological origin for ethics, and the spiritual origin, and blank slate theories are implausible, then we must look in another direction. I do not think we will ultimately need to look for that other direction because we have discovered how the good can be generated from (the towards the good) process of optimal decision-making that underpin the flower and honeybee facultative mutualism that exhibits the emergence of goodness for all the reasons that this study suggests. However, this is a limited study of one mutualism and therefore studies of other social groups are required in order to make a stronger claim to the origins of morality in nature. It may be that there are other approaches to the emergence of ethics in life that we have not yet discovered. 13

Morality in Nature

This study began with the question: “Is morality solely a human creation or can we discover evidence of or the antecedents of morality in nature?” Specifi­ cally, this study considered the flower and honeybee facultative mutualism to

200

Chapter 5

determine whether both well-studied species provide evidence of morality emerging in nature. A major criticism of ethical naturalism purportedly comes from both Hume and Moore. I suggest that Hume, rather than saying that morality could not emerge in nature, more specifically was critiquing rather poorly constructed arguments made by proponents of naturalistic morality and moral naturalism. Moore suggested that his term ‘naturalistic’ was not as important as ‘fallacy’, “And I do not care about the name: what I do care about is the fallacy.”117 Moore’s critique was of the idea of good, which he could not define. He also wanted to end the practice of associating good with the concept or object, for example, yellow. To associate goodness with the object requires that the object always be good. Such generalizations lead to bad philosophy. Rather, he suggests that we begin with a blank slate, and consider how the good could arise without precondition or pre-conceived notions of the emergence of goodness. This is all that both Darwin and Kitcher have asked us to do, to look at the historical and contemporary record of life in order to discover naturalism without fallacies. This is what this study has endeavored to do through two devices, optimization, and mutualism, both associated with flowers and honeybees. The behavioral process called optimization begins with the notion that when presented with multiple options, many (if not most) life forms rationally choose the best option based upon environmental and existential conditions. This does not mean that optimization is good, because sometimes the best decisions may not produce the good for the deciding species or others. Even so, the emerging science of MEP suggests that optimization is central to how life produces entropy likely in greater amounts than if the world were in an abiotic state. Therefore, optimization assists life in complying with Newton’s second law of thermodynamics. Optimization, correspondingly, must be a natural fact. The question then asked was whether optimization is, in general, towards the good. I suggest that the qualified answer is ‘yes’, and perhaps this is generally the case in nature, but, heeding Moore’s admonition against generalizations, we must secure more evidence from the greater biotic world. I ask, is optimization generally towards the good in other than the flower and honeybee facultative mutualism? Even proponents of MEP give the same qualified answer, that while there is increasing evidence of MEP that emerges from conducted studies, a blanket declaration of MEP as a universal is not appropriate. In summary, the emergence of morality in nature has been developed through just one social group, the flower and honeybee facultative mutualism. The descriptive elements of morality are derived from the optimal actions 117  Moore, Principia Ethica, 14.

The Good and the Emergence of Morality

201

taken by both species. The normative comes from the structure of the social group, the facultative mutualism, which is asymmetrical regarding benefits and exploitations of each other and the asymmetrical but reciprocal focus of each species towards the other that is both persistent and durable. From the descriptive and normative elements of this ethical construct we have adapted Levinas’ theory of responsibility to a theory of reciprocal responsibility which is applicable, at this point, only to this social group. Both flowers and honeybees are also reciprocally hospitable to each other, serving in both roles as host and guest simultaneously. Reciprocal responsibility and reciprocal hospitality are the normative moral theories that emerge from the normative construct: the facultative mutualism. However, the implications of reciprocal responsibility and hospitality are significant. If no one in the society must compete for one or more mutually required resources, then the need to develop rules for sharing are not necessary. This means that conflict that can arise from cheating or other consequences of the need to share a common resource does not regularly arise in such a social construct. When we look at human social groups, it quickly becomes apparent that the quest for the same required resources is one cause for much conflict and inequality in the human societal condition. The flower and honeybee social group has no (or very limited) need to address such potentials for conflict and so the efforts of the participants, while optimal towards each individual’s needs, also become optimal to the mutualism. Because there is no need to compete, actions associated with the mutualism social group are either neutral or advantageous to the other, with very few instances where they are disadvantageous. Why? There is no rational reason for either species to harm the other in the pursuit of a resource because neither competes for a common resource and both depend upon the other for different resources. The final chapter of this study summarizes the whole study, and then considers an oblique obverse to optimization—maximization—in context of human activities on planet earth. Cited References Alexander, Richard D. “A Biological Interpretation of Moral Systems.” Zygon 20, no. 1 (1985): 3–20. Anten, Niels P. R., and Heinjo J. During. “Is Analysing the Nitrogen Use at the Plant Canopy Level a Matter of Choosing the Right Optimization Criterion?”. Oecologia 167, no. 2 (2011): 293–303.

202

Chapter 5

Babikova, Zdenka, Lucy Gilbert, Toby J. A. Bruce, Michael Birkett, John C. Caulfield, Christine Woodcock, John A. Pickett, and David Johnson. “Underground Signals Carried through Common Mycelial Networks Warn Neighbouring Plants of Aphid Attack.” Ecology Letters 16, no. 7 (2013/07/01 2013): 835–43. doi:10.1111/ele.12115. Bateson, Melissa, Suzanne Desire, Sarah E. Gartside, and Geraldine A. Wright. “Agitated Honeybees Exhibit Pessimistic Cognitive Biases.” Current Biology 21, no. 12 (2011): 1070–73. doi:https://doi.org/10.1016/j.cub.2011.05.017. Beekman, Madeleine, James C. Makinson, Margaret J. Couvillon, Kaitlyn Preece, and Timothy M. Schaerf. “Honeybee Linguistics—a Comparative Analysis of the Waggle Dance among Species of Apis.” [In English]. Frontiers in Ecology and Evolution 3, no. 11 (2015-February-17 2015): 1–11. doi:10.3389/fevo.2015.00011. Beiler, Kevin J., Daniel M. Durall, Suzanne W. Simard, Sheri A. Maxwell, and Annette M. Kretzer. “Architecture of the Wood‐Wide Web: Rhizopogon Spp. Genets Link Multiple Douglas‐Fir Cohorts.” New Phytologist 185, no. 2 (Jan 2010): 543–53. doi:10.1111/j.1469-8137.2009.03069.x. Bernasconi, Robert, and David Wood. The Provocation of Levinas: Rethinking the Other. London and New York: Routledge, 1988. Biedrzycki, Meredith L., and Harsh P. Bais. “Kin Recognition in Plants: A Mysterious Behaviour Unsolved.” Journal of Experimental Botany 61, no. 15 (2010): 4123–28. doi:10.1093/jxb/erq250. Biedrzycki, Meredith L., Tafari A. Jilany, Susan A. Dudley, and Harsh P. Bais. “Root Exudates Mediate Kin Recognition in Plants.” [In eng]. Communicative & Integrative Biology 3, no. 1 (Jan-Feb 2010): 28–35. doi:10.4161/cib.3.1.10118. Blackburn, Simon. “Précis of Ruling Passions.” Review of Ruling Passions, Simon Blackburn. Philosophy and Phenomenological Research 65, no. 1 (2002): 122–35. Cappellari, S. C., H. Schaefer, and C. C. Davis. “Evolution: Pollen or Pollinators—Which Came First?”. Current Biology 23, no. 8 (2013): R316–18. doi:10.1016/j.cub.2013.02.049. Crepy, María A., and Jorge J. Casal. “Photoreceptor-Mediated Kin Recognition in Plants.” New Phytologist 205, no. 1 (2015): 329–38. doi:10.1111/nph.13040. Curry, Oliver. “Who’s Afraid of the Naturalistic Fallacy?”. Journal of Evolutionary Psychology 4, no. 1 (2006): 234–47. doi:147470490600400120. Derrida, Jacques. Adieu to Emmanuel Levinas. Translated by Pascale-Anne Brault, & Michael Nals. Stanford, CA: Stanford University Press, 1999. Dewey, John. Human Nature and Conduct. New York: Henry Holt and Co., 1922. Downs, Stephen G., and Francis L. W. Ratnieks. “Adaptive Shifts in Honey Bee (Apis Mellifera L.) Guarding Behavior Support Predictions of the Acceptance Threshold Model.” Behavioral Ecology 11, no. 3 (2000): 326–33. Dudley, Susan A., Guillermo P. Murphy, and Amanda L. File. “Kin Recognition and Competition in Plants.” Functional Ecology 27, no. 4 (2013): 898–906. doi:10.1111/ 1365-2435.12121.

The Good and the Emergence of Morality

203

Eller, Cleiton B., Lucy Rowland, Rafael S. Oliveira, Paulo R. L. Bittencourt, Fernanda V. Barros, Antonio C. L. da Costa, Patrick Meir, et al. “Modelling Tropical Forest Responses to Drought and El Niño with a Stomatal Optimization Model Based on Xylem Hydraulics.” Philosophical Transactions of the Royal Society B: Biological Sciences 373 (2018): 201–14. doi:10.1098/rstb.2017.0315. Fehr, Ernst, and Simon Gächter. “Altruistic Punishment in Humans.” Nature 415, no. January (2002): 137–40. Gadagkar, Raghavendra. “Why the Definition of Eusociality Is Not Helpful to Understand Its Evolution and What Should We Do About It.” Oikos 70, no. 3 (1994): 485–88. doi:10.2307/3545789. Goran I, Agren, and Franklin Oskar. “Root: Shoot Ratios, Optimization and Nitrogen Productivity.” Annals of Botany 92, no. 6 (Mar 2003): 795–800. doi:10.1177/ 007327538902700101. Heidegger, Martin. “Building Dwelling Thinking” Translated by Albert Hofstadter. Chap. IV In Poetry, Language, Thought, 145–61. New York: Harper & Row, 1971. Hume, David. Treatise of Human Nature. Gutenberg.org: Project Gutenberg, 2012. Kietzman, Parry M., and P. Kirk Visscher. “The Anti-Waggle Dance: Use of the Stop Signal as Negative Feedback.” [In English]. Ecology and Evolution 3 (2015-February-17 2015): 1–5. doi:10.3389/fevo.2015.00014. Kirchner, Wolfgang H. “Vibrational Signals in the Tremble Dance of the Honeybee, Apis Mellifera.” Behavioral Ecology and Sociobiology 33, no. 3 (1993): 169–72. Kitcher, Philip. “Naturalistic Ethics without Fallacies.” In Evolution and Morality, edited by James E. Flemming and Sanford Levinson, 1–30. New York: New York University Press, 2012. Kitcher, Philip. Vaulting Ambition. Cambridge, Ma.: MIT University Press, 1985. Klemens, Jeffrey A. “Kin Recognition in Plants?” [In eng]. Biology Letters 4, no. 1 (2008): 67–70. doi:10.1098/rsbl.2007.0518. Levinas, Emmanuel. Otherwise Than Being. Translated by Alphonso Lingis. Pittsburg, Pa.: Duquesne University Press, 1974. Levinas, Emmanuel. Totality and Infinity: An Essay on Exteriority. Translated by Alphonso Lingis. New York, New York: Springer, 1969. Levinas, Emmanuel, and Philippe Nemo. Ethics and Infinity: Conversations with Philippe Nemo. Translated by Richard Cohen. Pittsburgh, PA: Duquesne University Press, 1985. Lipiñski, Zbigniew. “The Emotional Nature of the Worker Honeybee (Apis Mellifera L.).” Journal of Apicultural Science 50, no. 1 (2006): 49–62. Liu, Kevin, and Tandy Warnow. “Treelength Optimization for Phylogeny Estimation.” PLoS ONE 7, no. 3 (2012): 1–9. doi:10.1371/journal.pone.0033104. Marder, Michael. Grafts. Minneapolis, Mn.: University of Minnesota Press, 2016. Marder, Michael. “Of Plants, and Other Secrets.” Societies 3, no. 1 (2013): 16–23.

204

Chapter 5

Marder, Michael. Plant-Thinking: A Philosophy of Vegetal Life [in English]. None. New York: Columbia University Press, 2013. Book. Marder, Michael. “Vegetal Anti-Metaphysics: Learning from Plants.” Continental Philosophy Review 44, no. 4 (2011/11/01 2011): 469–89. doi:10.1007/s11007-011-9201-x. McNickle, Gordon G. “Root Foraging Behaviour of Plants: New Theory, New Methods and New Ideas.” University of Alberta, 2011. Mommer, Liesje, Jasper van Ruijven, Corien Jansen, Harry M. van de Steeg, and Hans de Kroon. “Interactive Effects of Nutrient Heterogeneity and Competition: Implications for Root Foraging Theory?”. Functional Ecology 26, no. 1 (2012): 66–73. doi:10.1111/j.1365-2435.2011.01916.x. Moore, George Edward. Principia Ethica. Cambridge UK: Cambridge University Press, 1903. O’Gorman, Kevin D. The Origins of Hospitality and Tourism. Oxford, UK: Goodfellow Publishers Limited, 2010. Ratnieks, Francis L. W., and Kyle Shackleton. “Does the Waggle Dance Help Honey Bees to Forage at Greater Distances Than Expected for Their Body Size?” [In English]. Frontiers in Ecology and Evolution 3, no. 31 (2015-April-01 2015): 1–7. doi:10.3389/ fevo.2015.00031. Rueppell, Olav, M. K. Hayworth, and N. P. Ross. “Altruistic Self‐Removal of Health‐ Compromised Honey Bee Workers from Their Hive.” Journal Of Evolutionary Biology 23, no. 7 (2010): 1538–46. Ruse, Michael, and Edward O. Wilson. “Moral Philosophy as Applied Science.” Philosophy 61, no. 236 (1986): 173–92. Schiffers, Katja, Katja Tielbörger, Britta Tietjen, and Florian Jeltsch. “Root Plasticity Buffers Competition among Plants: Theory Meets Experimental Data.” Ecology 92, no. 3 (2011): 610–20. Schürch, Roger, Magaret J. Couvillon, and Madeleine Beekman. “Ballroom Biology: Recent Insights into Honey Bee Waggle Dance Communications.” Frontiers in Ecology and Evolution 2016, no. February (2016): 4–6. Singer, Peter. The Expanding Circle. Princeton, N.J.: Princeton University Press, 1981, 2011. Song, Yuan Yuan, Ren Sen Zeng, Jian Feng Xu, Jun Li, Xiang Shen, and Woldemariam Gebrehiwot Yihdego. “Interplant Communication of Tomato Plants through Underground Common Mycorrhizal Networks.” PloS one 5, no. 10 (2010): e13324, 1–11. doi:10.1371/journal.pone.0013324. Teehan, John, and Christopher DiCarlo. “On the Naturalistic Fallacy: A Conceptual Basis for Evolutionary Ethics.” Evolutionary Psychology 2, no. 1 (2004): 32–46. Trewavas, Anthony. Plant Behavior and Intelligence. Oxford, UK: Oxford University Press, 2014.

The Good and the Emergence of Morality

205

Trewavas, Anthony. “What Is Plant Behaviour?”. Plant, Cell & Environment 32, no. 6 (2009/06/01 2009): 606–16. doi:10.1111/j.1365-3040.2009.01929.x. Tudge, Colin. The Tree. New York: Three Rivers Press, 2995. Unelius, C. Rikard, Christian Schiebe, Björn Bohman, Martin N. Andersson, and Fredrik Schlyter. “Non-Host Volatile Blend Optimization for Forest Protection against the European Spruce Bark Beetle, Ips Typographus.” PLoS ONE 9, no. 1 (2014): 1–10. doi:10.1371/journal.pone.0085381. Walter, Alex. “The Anti-Naturalistic Fallacy: Evolutionary Moral Psychology and the Insistence of Brute Facts.” Evolutionary Psychology 4, no. 1 (2006): 33–48. Walton, Alexander, and Amy L. Toth. “Variation in Individual Worker Honey Bee Behavior Shows Hallmarks of Personality.” Behavioral Ecology and Sociobiology 70, no. 7 (2016/07/01 2016): 999–1010. doi:10.1007/s00265-016-2084-4. Wario, Fernando, Benjamin Wild, Margaret J. Couvillon, Raúl Rojas, and Tim Landgraf. “Automatic Methods for Long-Term Tracking and the Detection and Decoding of Communication Dances in Honeybees.” [In English]. Frontiers in Ecology and Evolution 3, no. 103 (2015-September-25 2015): 1–14. doi:10.3389/fevo.2015.00103. Wilson, David Sloan, Eric Dietrich, and Anne B. Clark. “On the Inappropriate Use of the Naturalistic Fallacy in Evolutionary Psychology.” Biology and Philosophy 18, no. 5 (2003): 669–82. Wilson, Edward O. “What Is Sociobiology?”. Society 15, no. 6 (1978): 10–14. Wittgenstein, Ludwig. Philosophical Investigations. Translated by G. E. M. Anscombe. Oxford: Basil Blackwell, 1958. Wohlleben, Peter. The Hidden Life of Trees: What They Feel, How They Communicate— Discoveries from a Secret World. Vancouver, BC Canada: Greystone Books, 2016. Wright, Tamra, Peter Hughes, and Alison Ainley. “The Paradox of Morality: An Interview with Emmanuel Levinas.” Translated by Andrew Benjamin & Tamra Wright. Chap. 11 In The Provocation of Levinas: Rethinking the Other, edited by Robert Bernasconi, & Wood, David, 168–81. London and New York: Routledge, 1988. Wyschogrod, Edith. Crossover Queries: Dwelling with Negatives, Embodying Philosophy’s Others. New York: Fordham University Press, 2006. Youngsteadt, Elsa, Rebecca E. Irwin, Alison Fowler, Matthew A. Bertone, Sara June Giacomini, Michael Kunz, Dale Suiter, and Clyde E. Sorenson. “Venus Flytrap Rarely Traps Its Pollinators.” The American Naturalist 191, no. 4 (2018/04/01 2018): 539–46. doi:10.1086/696124.

Chapter 6

Study Summary and a Critique of Maximization 1

Study Summary

Not long after the earth became a planet, life emerged. Unlike other emergent properties in nature like Benard cells that return to prior states after external application of heat/cool, light, pressure, etc. end, life has maintained itself over billions of years on this planet. I suggest that one unique property of life is its quest for perpetuity. Even four near-extinction events in the history of life have not returned the earth to the state where it was prior to life’s first sustainable emergence. A process that life uses to sustain its emergence is DNA which may have evolved after the first proto life emerged and began to sustain itself. DNA has two major functions. First, it preserves the recipe for creating new life forms because no individual life is immortal. Second, it is capable of mutation which means that as circumstances of the world change, new life forms can be adapted to meet new challenges. Mutation has also helped life find new niches in the world in which to thrive. In fact, the first great extinction event was when stromatolites developed the process of photosynthesis where they could manufacture what was necessary from sunlight and carbon dioxide to sustain their emergence. The resulting oxygenation of the atmosphere poisoned anaerobic species. We might call what resulted from this extinction event the photosynthetic revolution. Sometime after, plants emerged and when some evolved vascular systems, this eventually led to the emergence of angiosperms, the flowering plants, the earliest of which in the fossil record (pollen) are recorded one-hundred-thirty-one million years ago. Oxygenation of the atmosphere was not long an impediment to life. Animal species evolved to use oxygen towards mobility and other processes, and along the way, insects soon appeared. Insects for many millions of years were carnivorous creatures either eating other insects, or like mosquitos and tics, supping from the bodily fluids of other creatures. Others like aphids drilled into plant vascular systems to access the sugary fluid that is a product of photosynthesis. When angiosperms first emerged, likely from one originary species, there most likely were no animal pollinators as there are today. Traditional means of getting pollen to stigma in vascular plants included wind and water which are subject to the vagaries of nature and require massive amounts of pollen production because of low probability of individual pollen spores finding another of the same species. This means much energy production of the wind or water

© Koninklijke Brill NV, Leiden, 2020 | doi:10.1163/9789004428546_008

Study Summary and a Critique of Maximization

207

pollinated plant goes towards producing pollen at the expense of other processes. What happened next is only now being pieced together. Two things must have occurred to bring insects and flowers together. First was pollen that could stick to the insect that could be carried to other flowers of the same species. Second, the insect needed to be attracted to the flower species for this process to be established. How this happened is uncertain. Perhaps the apoid wasp, the predecessor to the bees, found insects huddled among flowers and so began to include flowers in their hunting routines. The many hundreds of thousands of pollinating insect species and the many hundreds of thousands of angiosperm species now in existence in most of the world’s ecologies show how successful and important this process of pollination has become. This study began its evolutionary journey with the apoid wasp, the predecessor to the bee and the Hymenoptera distant ancestor of the honeybee. The apoid wasp evolved from carnivorous insect hunter to a species that preferred pollen protein over insect protein. At the same time the sticky pollen of the flower attached itself to the apoid wasp who carried it from flower to flower. It is likely that the apoid wasp had not yet developed a taste for carbohydrate nectar and perhaps the flower had not yet produced such a product. However, over time, the apoid wasp became a bee who no longer pursued insects for nourishment. The flower by now had developed means for producing nectar and the bee the means to consume and benefit from this nectar, all the while still carrying the pollen from flower to flower after it stuck to her. The co-evolution arms race, as it were, honed the honeybee into a flower specialist and at the same time the flower modified herself into species that would attract honeybees and encourage them not only to drink her nectar but to search for others of the same species for a corresponding reward. Along the way, the honeybee evolved two important existential advantages. First, it became eusocial, living in a caste society where there is little strife and much communal activity. Second, it developed a means of communicating where flower patches are located to other foraging members of the hive, the waggle dances. Ants have long used pheromone trails to direct foragers to food sources. Honeybees fly to their food sources which makes pheromone trails impractical. The creature with a brain of just one million neurons can communicate abstract direction and distance as well as the navigation system on the first human moon landing with far fewer bits of information than the now primitive Apollo navigation computer. For a million years honeybees and flowers have co-evolved into the facultative mutualism that is the subject of this study. This study asks specifically, “Is morality solely a human creation or can we discover evidence of or the antecedents of morality in nature?” The remarkability of nature and her evolutionary processes aside, the effort of this study

208

Chapter 6

has been to determine whether we can find evidence of morality in nature in other than humans and in other than human social groups. As we have seen there is quite a debate (the extent which was only briefly reviewed for the basic arguments) over whether there is such a thing as natural morality. Proponents of the naturalistic fallacy point to Hume and Moore to critique the derivation of goodness from natural facts. Others, like David Sloan Wilson suggest that natural facts are all that we have. Therefore, we must address natural facts even if we eventually find them inadequate for the task of ethics. Others like Kitcher argue that these natural facts must be considered through the lens of evolution and the history of life on earth for evidence of the emergence of morality which he calls pragmatic naturalism. Michael Ruse and E. O. Wilson take this a step further, declaring that our brains are too small to learn morality from scratch and therefore there must be an epigenetic origin of morality that is handed down from generation to generation. However, all this discourse is just that without evidence of the natural emergence of morality in other than humans. It has been the effort of this study to supply this evidence in a narrow frame, the flower and honeybee facultative mutualism. Much of this study has explored the individual capabilities and behaviors of flowers and honeybees both within their existence that is conducted outside the mutualism and for those behaviors directly associated with their mutualism. However, agreeing with the basic premise of Moore’s naturalistic fallacy, we cannot say that even activities or processes that appear to produce good moral results are good. We can, however, search for the good that may arise from processes that generate behaviors, assess the effects or results of these behaviors, and then consider what that means in the context of the emergence of morality in flower and honeybee activities. The evidence for the emergence of the good in nature through the flower and honeybee facultative mutualism begins with the notion of optimization. Optimization is choosing the best decision from multiple available options, through a process that rationally considers the impact of those options, and then acts upon the chosen optimal decision. Optimization, in general, com­ports with Newton’s second law of thermodynamics: entropy. From optimi­za­tion decision making in nature comes the idea that some observed processes in nature are generally towards Maximum Entropy Production or MEP. While MEP proponents do not posit that all natural processes are towards MEP, there are many processes that science has studied that are believed to meet theoretical MEP criteria. This is important because life as an emergent property of nature appears to produce more complexity and order than it generates entropy, and without further analysis would seem to defy Newton’s second law. However, research has discovered that life, with its consumption of

Study Summary and a Critique of Maximization

209

considerable amounts of energy to maintain itself and reproduce, likely generates more entropy than if the earth were abiotic. If optimization contributes to entropy and, more specifically to MEP, then optimization is a natural process and natural fact that complies with the laws of physics. Optimization is likely practiced by most if not all creatures in the circumstances and environment where each exists. While optimization may be primordial to the emergence of morality, we cannot call optimization good because that would be fallacious. We need something more. Singer says there are three pre-conditions for the emergence of morality—the social group, restraint towards other members of the group, and judgment. Correspondingly, according to Singer’s theory, solitary creatures by themselves do not engender the emergence of morality. We must find social groups in this study of flowers and honeybees if we are to consider whether morality has or can emerge. We know that the eusocial honeybee hive is a social group, there is little strife in the hive, and honeybees make optimal decisions in their many life activities which suggests that they judge. While there is some evidence that flowering plants may have some social contact with members of their own species and even others, it is difficult to say flowers, in general, form and maintain social groups with other flowers. We can, however, look at the flower and honeybee facultative mutualism to see whether it is in fact a social group because certainly Singer’s other requirements, 1) restraint towards other members, and; 2) judgment leading to behaviors that are optimal to both species and the mutualism itself can be observed from studying flower and honeybee behavior towards each other. The structure of the flower and honeybee mutualism is such that it is both beneficial and exploitative or facultative. It is non-exclusive because flowers that honeybees forage are also foraged by other insects and pollinators. Honeybees also are opportunistic foragers—any suitably displayed flower they will forage. Their mutualism is not obligate because they require each other for only one life function: honeybees need flower-produced food; flowers need honeybee mobility to facilitate their sex act. Beyond that they are not dependent upon each other. However, both have co-evolved to require each other and to exploit each other in their mutualism. Finally, in concert with Singer’s second requirement, they show considerable restraint in their activities with each other which means that neither intentionally harms the other. From satisfying these requirements the stage is set for morality to emerge if and only if this mutualism is a social group. Consider these features of their facultative mutualism: there is durability, flowers and honeybees have co-existed and coevolved for at least a million years, they depend upon each other and even cannot exist without the other, they have developed rules of conduct towards

210

Chapter 6

each other (through optimal decision making: rationality), and have maintained this from generation to generation.1 Therefore, I maintain that the flower and honeybee mutualism is a social group made from the efforts of very different species. From the understanding that this facultative mutualism is a social group that meets Singer’s three requirements for morality to emerge, we can begin to search for where morality might have or possibly may emerge. The question is what kind of morality has/or could emerge from this facultative mutualism? It is important to delve into the structure of the mutualism, which for both species, benefits from the other and exploits the other. First, both the benefit and exploitation are asymmetrical. The honeybee gains food but the flower exploits forager mobility and energy to do so for her pollination requirements. The honeybee gains food but exploits the flower’s ability and energy to produce food for the benefit of the honeybee. Neither species competes with the other for the same resource; nor does either exploit the other the same way. Asymmetricity produces restraint in the facultative mutualism because neither competes with the other for any existential requirement gained from or exploited by the other. The third asymmetrical feature of the flower and honeybee facultative mutualism is that they are asymmetrically oriented towards the world and each other. The flower is all middle and is centrifugally oriented towards the world, reaching out to the world through her honeybee advertisement, the flower. Through the flower she calls the honeybee to her and the honeybee heeds this call. This is because the foraging honeybee is centripetally oriented earthward towards the flower in the meadow, orchard, jungle, or forest. Both become intimate when the honeybee enters the flower but retain their separate orientation towards the world. For even as the honeybee enters the flower, she will soon think about visiting other flowers, and the flower continues advertising for other pollinators and other bees to visit her. While their asymmetrical orientations are maintained continually, the individual flower and honeybee may have only a short relationship with each other lasting days or perhaps weeks. Through the part of the year where flowers do not bloom, neither the flower nor honeybee engage with each other. This does not suggest that their rather limited engagement is not a social group construct. People join clubs which may only be open for specific functions or only during specific times 1  Singer does not give us the definition of social group other than one is necessary for ethics to emerge. I suggest that rather than try to develop a formal definition of a social group, the features of the flower and honeybee facultative mutualism outlined here suggest both sociality and group behavior that we can intuitively suggest is a social group.

Study Summary and a Critique of Maximization

211

of the year. However, these clubs can be defined as social groups because of regular members, activities, and rules of conduct, and there is some measure of restraint towards other members of the club. This study considers asymmetricity as foundational to the emergence of morality in the flower and honeybee facultative mutualism, but what needs to be analyzed are the behaviors of these asymmetrically oriented beings. Through Levinas’s ethics of responsibility this study suggests that indeed, flowers and honeybees are responsible to each other. Like Levinas who locates morality in the existential human face of the other and where we recognize the other and know that we must therefore be responsible to the other, the flower creates its flower ‘face’ for the honeybee and the honeybee’s tactile presence when she enters the flower alerts the flower to her presence (an pseudo-face). Responsibility for the other, however, in Levinas’s human ethics is asymmetrical. I am responsible to the other, but it is the other’s business to reciprocate. I am also responsible to the other even to the detriment of my own existence. Humans often compete for resources. Levinas’s notion of asymmetrical responsibility, passivity before the other, and even substitution for the other are likely necessary to bring into focus the notion that human responsibility must somehow deal with competition for scarce common resources. This is not the case for the flower and honeybee mutualism. Any scarcity of resource is generally generated from outside the mutualism, e.g. genetics, lack of sun, water, or adverse weather. As the flower and honeybee do not compete for the same resource, they can be reciprocally responsible for each other because of their asymmetrical needs. This means they will not try to gain more from the other than what the other can (or wishes to) provide as might be the case when two humans fight over the last piece of bread or one sells another an inferior product at a superior price. Flowers and honeybees have maintained their reciprocal responsibility to each other for more than a million years, likely altering responsible behaviors as both species evolved. However, what likely has never changed is the reciprocal nature of their responsibility even as behaviors towards each other have evolved. From this reciprocal responsibility emerges the good associated with the flower and honeybee facultative mutualism. I maintain that we have indeed learned a lot from the meadow. Levinas tries to develop conditions for the emergence of responsibility in human interface, but he asks quite a lot of me who am responsible for this other whose face I cognize as being human. He suggests that I must be passive even to the point where I substitute myself for the other to the detriment of myself. Why? It is the business of the other to reciprocate. This asymmetry of responsibility likely is caused by the need for both of us to compete

212

Chapter 6

for the same existential resources. If there is only one piece of bread, and the other is hungry, Levinas’ responsibility ethic maintains I must let the other take it all if the other wants. War and human conflict suggest that humans are not always so inclined. As both species are reciprocally responsible, they are both reciprocally hospitable. They are both simultaneously host and guest and welcoming when they encounter the other. Both give hospitality and receive it. This reciprocity of hospitality dovetails with reciprocal responsibility that occurs because both benefit from and exploit the other asymmetrically. The flower beckons the honeybee and the honeybee responds to this hailing by entering the flower. Both have co-evolved processes and morphology that can capitalize on cohospitably from each other. They both give, as would the host, and receive as would the guest. The world is filled with illusory dualities like predator prey, competitive winner and loser, and conflict and avoidance. Even without strict dualities, the world is a contentious place. What is refreshing about the flower and honeybee mutualism is that both species are elementarily non-dual and without ends. The plant is all middle, growing and retreating from the center, which is ultimately attached to the great middle, the earth. The honeybee hive is all middle because there is no central authority who commands. Nor does the flower access a central authority because it has no brain. David Loy quotes Plotinus, “There were not two; beholder was one with beheld; it was not a vision compassed but a unity apprehended.”2 Unity is what we see with the flower and honeybee reciprocal gaze. Both are seers and are always seen which means that duality never emerges through their respective and continual gazes. While this may be Plotinus’s apprehended unity it is also a unity unbounded in its continuity over time: this moment to the next; this year to the next; intergenerational, with a million-year history. This continuity of gaze represents a timelessness that is outside of time as we might conceive of it. Consider that both flowers and honeybees are in the moment. While both can record existential events and refer to them when similar circumstances arise, they are always already in the moment. Jacob Boehm describes this timelessness, “He to whom time is the same as eternity and eternity the same as time is free from all contention.”3 I do not submit that contention is void in the existential lives of flowers and honeybees, but the flower and honeybee facultative mutualism

2  David Loy, Nonduality: A Study in Comparative Philosophy (Amherst, NY: Prometheus Books, 2012), 2. Kindle Edition. 3  B. Harding, The Doctrines of Jacob Boehm (New York: Macoy Publishing, 1919), 33.

Study Summary and a Critique of Maximization

213

is free from most contention and therefore may just be the timelessness that Loy and Boehm envision. Even in Levinas’s responsibility ethic there is a difference between you and me (beholder and beheld) which requires my absolute responsibility without any expectation of reciprocity from you. Flower and honeybee reciprocal responsibility does not require that the flower treats the honeybee a certain way and the honeybee responds identically in kind. There is no need because they are so different and their needs are so different, that reciprocity is given in return but not necessarily in kind. It is remarkable that such a non-dual social group could emerge from an existential world that has been called survival of the fittest. Even so, all around the periphery of this non-dual social group are contentions. Flowers fight for light and nutrients and against herbivores and pathogens. Honeybees fight competitors for access to flowers and escape predators who would eat them. At the hive, guard honeybees assail robber honeybees from other hives. The non-dual facultative mutualism serves as a respite from such contentions for both species. This mutuality is neither good nor evil. It avoids dualities and therefore duality is not a subject for the flower and honeybee facultative mutualism. The emptying of dualities is of some importance to the Buddhist notion of nirvana. Because insects and plants can achieve this level of emptiness from duality in their social group suggests that perhaps the Mahayana branch of Buddhism has a useful thought that all sentient creatures can become enlightened, which they call Buddha nature. Whether one is oriented towards historically Asian, African, or European philosophical thought, this social group is empty from duality which means that this is an important social group to study. This is why I believe we can all learn from the meadow where the flower and honeybee mutualism is conducted and consummated. The specter of competition for scarce synchronous resources between humans may be a condition we may not ever be able to overcome. Within notion of the emptiness from duality, the flower and honeybee reciprocal responsibility and reciprocal hospitality serve as a critique of the human condition. Specifically, how can people structure their relationships to where reciprocal responsibility and hospitality can be engaged naturally like the flower and honeybee? How can we construct social groups empty of duality? What amount of asymmetricity of benefit or exploitation can be accomplished in human social groups to where reciprocal responsibility and hospitality become common conditions of co-existence? Answers to these questions are for others to consider. However, I suggest that humans can learn from this flower and honeybee study’s exploration of what helps construct the reciprocal responsibility/hospitality and that is the consistent optimization we see in the behaviors of both, in their separate and mutualism endeavors, all of which

214

Chapter 6

likely produce more entropy than behaviors that tend towards maximizing benefit or exploitation of the other or of the environment. I believe this study has produced evidence that morality can emerge in nature, specifically in the flower and honeybee facultative mutualism. In this mutualism, optimization produces the conditions for reciprocal responsibility and hospitality that produces much of the good for the social group called flower and honeybee facultative mutualism. If optimization is towards the good, then optimization should be considered in context of human endeavors. Humans have the capability of altering natural conditions so that we can produce greater reward than what would be possible under the previous natural conditions. Flowers and honeybees have limited capabilities of altering natural conditions. Humans often maximize reward by removing conditions that would in nature require a different outcome to be optimal. This study shows that morality (and the good) can emerge in nature and, and in this study, does so through optimization processes that engender reciprocal responsibility. Next is a list of key findings of this study. 1.1 Key Findings of This Study – The flower and honeybee facultative mutualism is a social group. – While both flowers and honeybees co-evolve genetically, they require extant beings to behave in ways that are both towards and maintains their facultative mutualism. – Flowers and honeybees make optimal decisions towards each other which generally benefit themselves, the other, and their facultative mutualism. – Flowers and honeybees, by-and-large make optimal decisions that require judgment. – Optimal decision-making by flowers and honeybees is consistent with Maximum Entropy Production (MEP) theories that maintain that, in general, life produces entropy consistent with the second law of thermodynamics. – Optimization is a natural fact. – Optimization is a process that is antecedent to but towards morality. – Flowers and honeybees have an asymmetrical relationship to each other. They benefit and exploit each other for asymmetrical needs: nutrition versus procreation assistance. Therefore, they do not compete for scarce resources. – Both flowers and honeybees exhibit restraint towards each other. – Flowers and honeybees are asymmetrically but reciprocally focused on each other. Flowers: centrifugal. Honeybees: centripetal. This consistent reciprocal gaze has helped them maintain their relationship for a million years.

Study Summary and a Critique of Maximization

215

– Flowers and honeybees communicate with each other through visual and tactile means. – Flowers and honeybees have Levinasian ‘faces’ that announce the presence and importance of the alterior other. – Flowers and honeybees are reciprocally responsible to each other and are also reciprocally hospitable towards each other. – Because flowers and honeybees are reciprocally responsible and hospitable towards each other in their facultative mutualism they exhibit a modified form of Emmanuel Levinas’ responsibility ethic. – The flower and honeybee facultative mutualism characteristics meet Peter Singer’s requirements for morality to emerge. – Flowers and honeybees engage in what is being called para-psychological altruism with each other. – Naturalistic fallacies in this study were avoided. Neither optimization nor the facultative mutualism, nor flowers and honeybees were declared good. Rather, considering the responsible nature of their behaviors and their optimal decision-making, the construct called the flower and honeybee facultative mutualism is towards the good. – If the flower and honeybee mutualism is towards the good and both flowers and honeybees are reciprocally responsible and hospitable to each other, which are moral concepts, their mutualism provides one example where morality has emerged in nature outside of the purview of humanity. 2

A Brief and Preliminary Critique of Maximization

The question that this study now poses, is if we humans believe that maximization also produces the good, why do we not see much evidence of decision maximizing in species other than humans? “We can learn from the meadow,” deserves repeating. Recall that Robert J. Holdaway, et al., reported in their study of the Amazon, “Our results indicated that forests had a higher rate of entropy production than pastures.”4 It stands to reason that a meadow with its complex mix of plants and animals likely has a higher rate of entropy than the carefully crafted farm field that grows one species only. Earth with life likely produces more entropy than the world if it 4  Robert J. Holdaway, Ashley D. Sparrow, and David A. Coomes, “Trends in Entropy Production During Ecosystem Development in the Amazon Basin,” Philosophical Transactions: Biological Sciences 365, no. 1545 (2010): 1445.

216

Chapter 6

were abiotic. Holdaway et al., demonstrate that entropy production depends upon the concentration and perhaps the mix of life itself. Maximum entropy is achieved by the optimal actions of species in a natural setting in context of the conditions the (non-linear thermodynamic) world presents when the decision is made. Axel Kleidon and Ralph Lorenz note that studies suggest that, “[v]­egitated surfaces exhibit cooler surface temperatures and a lower surface albedo, which results in higher rates of entropy production.”5 Rather than radiate the heat from the sun, vegetated surfaces absorb it. This suggests that that vegetated surfaces channel some energy away from being radiated back into the atmosphere that would result in higher air temperatures. This is one reason why cities’ air temperatures are hotter than nearby meadows. Another is that leaves transpire which produces cooling water evaporation. Kleidon and Lorenz note that the higher entropy of vegetative states can be linked to both producers of energy (plants) and consumers (honeybees).6 Therefore, we can say that both flowers and honeybees participate in producing the higher entropy of a vegetative state. Furthermore, say Kleidon and Lorenz, through Alfred Lotka, “[t]hat assemblages of organisms can be viewed as ‘armies of energy transformers’ and the ‘[evolutionary]’ advantage must go to those organisms whose energy-capturing devices are most efficient in directing available energy into channels favorable to the preservation of the species.”7 In other words, the long-term advantage goes to the energy optimizers. Perhaps hundreds of species, both plants and animals, are making optimal decisions at any moment during the day in their meadow. Each is looking optimize which means finding a balance between the passion to maximize reward with reason that passionately wants to minimizing risk in every decision the individual makes. Risk and reward are adjudicated through the risk appetite of the individual. Honeybees avoid flowers where predators abound. A more aggressive honeybee might decide to enter a flower where competitors are present, while another from the same hive may not. Optimization requires making the best choice based upon the existential goals of the individual, considering both the risk and reward from multiple decision options in context of creature capabilities that help formulate basic risk appetite. While individual actions serve to shape the resulting environ to some extent, the entire meadow environ must continue to deal with exigencies that cannot be overcome by 5  Axel Kleidon and Ralph D. Lorenz, “Entropy Production by Earth System Processes,” in Non-Equilibrium Thermodynamics and the Production of Entropy: Life, Earth, and Beyond, ed. Axel Kleidon and Ralph D. Lorenz (New York: Springer Science & Business Media, 2005), 15. 6  Ibid. 7  Ibid.

Study Summary and a Critique of Maximization

217

the aggregate of individual decisions alone. These include weather, climate change, and humans. I suggest that humans are fundamentally maximizers. We want what we want and determine to obtain what we want, often regardless of the consequences or circumstances. For example, is it much more efficient and more economical for the farmer to plow under the meadow’s vegetation and add herbicides and pesticides to create a (near) aseptic environment to grow perhaps just one species of crop? A recent study by Claire E. LaCanne and Jonathan G. Lundgren note that, “Most cropland in the United States is characterized by large monocultures, whose productivity is maintained through a strong reliance on costly tillage, external fertilizers, and pesticides.”8 Cropland is de-naturalized to prepare it for the monoculture, for example: soybeans, corn, or wheat. Quite often orchards are cleared of underbrush and ‘undesirable’ plant and insect life e.g. worms. In this study LaCanne and Lundgren compared conventional monoculture farming with regenerative farming acreage to consider the effects on insect population, crop productivity, and overall profitability of the product. They outline the goals of regenerative farming: The goal of regenerative farming systems (Rodale, 1983) is to increase soil quality and biodiversity in farmland while producing nourishing farm products profitably. Unifying principles consistent across regenerative farming systems include (1) abandoning tillage (or actively rebuilding soil communities following a tillage event), (2) eliminating spatio-temporal events of bare soil, (3) fostering plant diversity on the farm, and (4) integrating livestock and cropping operations on the land.9 The idea of regenerative farming is to use the power of soil and the creatures that inhabit healthy soil to regenerate the soil naturally without the addition of pesticides, herbicides, and fertilizer. Cows, pigs, horses, and other farm animals provide natural fertilizer when left to roam in the fields. Such efforts may also reduce soil erosion and may require less water because the soil is rich with plant and other material that can hold water. As you might expect, the regenerative fields studied had many more insects than those of the conventionally tilled and treated farm fields. Crop yields were twenty-nine percent lower in regenerative fields. However, the regenerative farmers earned seventy-eight

8  Claire E. LaCanne and Jonathan G. Lundgren, “Regenerative Agriculture: Merging Farming and Natural Resource Conservation Profitably,” PeerJ, no. 2018 (2018): 1. 9  Ibid.

218

Chapter 6

percent more in net profit.10 As businesspersons, farmers understand that the amount of net profit they earn is more important than gross revenue. If they can increase their profit margins they will listen to those who have proven strategies to do so. More research is required, but regenerative farming strategies have potential to change crop farming strategy. The caveat with this approach is that the same acreage under regenerative farming produces lower yields per acre. Therefore, if the entire US were to change to regenerative farming methods, there would be less product to feed the burgeoning world population. More acreage would need to be farmed to make up for the shortfall. However, there likely will be other benefits that this one study has not contemplated, including reduced water consumption and the fact that as fields mature, they might develop insect populations that hold pestilence to cultivated crops in check. Perhaps, regenerative farming may help reclaim land that was once farmed or that could be farmed given the right conditions. For example, the ancient farming practice called Zai is being used to reclaim land that had been lost to the encroaching Sahara Desert.11 In Burkina Faso there is a short rainy season followed by drought. Modern farming methods do not work. How does Zai work? The farmer digs lots of pits, puts manure and other organic material inside each. This attracts termites and other insects who dig tunnels. When it rains, water is stored in these slick tunnels that hold water longer than bare soil or sand can. The farmer then plants crops. The plant roots grow into the tunnels and have access to water long after the rain. Waste material attracts insects who thrive on it, and in a partnership with the plant, give it time to grow and produce more waste material that the termites can then use themselves. Other insects and birds will find this place an oasis and may even improve the success of bird migration in the area. Other plants like trees protect the reclaimed land from the Sirocco winds. Such practices can help prevent famine and hold back the Sahara from increasing in size.12

10  Ibid. 11  See: M. N. Danjuma and S. Mohammed, “Zai Pits System: A Catalyst for Restoration in the Dry Lands,” Journal of Agriculture and Veterinary Science 8, no. 2 (2015). See also: Boubacar M. Moussa et al., “Combined Traditional Water Harvesting (Zaï) and Mulching Techniques Increase Available Soil Phosphorus Content and Millet Yield,” Journal of Agricultural Science 8, no. 4 (2016). 12  See also for additional techniques: Robert Zougmoré, Abdulai Jalloh, and Andre Tioro, “Climate-Smart Soil Water and Nutrient Management Options in Semiarid West Africa: A Review of Evidence and Analysis of Stone Bunds and Zaï Techniques,” Agriculture & Food Security 3, no. 1 (2014).

Study Summary and a Critique of Maximization

219

Recall that Holdaway’s research suggests that richer life environments produce more entropy than those with fewer species and/or biomass.13 A question that needs to be asked is to what extent has life as an optimizing force contributed to the earth maintaining a life-viable environment for billions of years, despite four near-extinction events and countless hot, cold, and other cycles? This is the fundamental question that regenerative farming science asks but does so at the local level and does so, not just for purposes of restoring environmental balance, but to improve the net income of farmers who deploy the practice. The benefits of such practices are two-fold: first to the ecology which is difficult to measure in dollars or even quality of life standards, and second, to the farmers who see realized improvements in quantifiable profits as a result. Life, in general, makes optimal choices based upon the environment presented to it at this moment even when the timeline of its goals is longer. For example, the honeybee must stockpile adequate reserves of honey to survive the winter but can only produce as much honey as the environment gives it tools to do so. The hive can, when necessary, alter the number of workers and allocate them to tasks in order to optimize this effort. Humans have not been successful in regulating their population. As population increases without restraint there are two measures of carrying capacity that we will confront. The first is the capability of the earth to produce enough food to feed the people. Second is quality of life. To support higher populations, populations who have excess food and resources will be confronted with increasing costs to maintain this imbalance and perhaps even go to war (or build walls) to resist allocating these excesses to others that, if reallocated, will reduce quality of living for those who have come to depend on more. One consequence of the need to feed increasing populations we see in human activity today, is that arable land is being repurposed towards other than agriculture e.g. roads, cities, homes. The remaining arable land that is tilled is made more and more aseptic to prepare for crops that are bred or engineered to produce the maximum amount of human consumables per acre as possible. While these hybrids may produce more entropy per plant than naturally occurring variants (e.g. larger, or more fruit), they are planted in earth that has been first made nearly aseptic to eliminate herbivores, pests, and pathogens. At the same time, species of plants that would have normally grown in the field have been tilled under or killed with herbicides. The field then must be prepared with fertilizers that are designed to facilitate the maximal growth of the planted crop. Leaving the meadow’s root systems intact would have retained water 13  Holdaway, Sparrow, and Coomes, “Trends in Entropy Production During Ecosystem Development in the Amazon Basin,” 1445.

220

Chapter 6

and reduced runoff. The tilled field in many environs now must be irrigated both to replace lost water and to achieve the maximum growth possible for the crop per acre. In some areas, aquifers are not being replenished adequately to offset the irrigation load. Crops are often rotated in order to replenish some of the soil nutrients that are lost by single-crop agriculture. However, even this is inadequate, and fertilizers, herbicides, and pesticides must be applied each year. The economics of farming requires this continual increase in yield per year not only for economic reasons, but to satisfy demand. New hybrid seeds are developed to fine tune the crop to local environmental, soil, and other conditions like pest resistance. However, hybridization quite often also reduces the genetic variety which means that opportunistic herbivores, pests, and pathogens frequently find ways to exploit the lack of genetic diversity. Pathogens and pests also become resistant to pesticides and ever stronger pesticides must be produced. Pesticide and herbicide residue may also be harmful to beneficial insects like honeybees and other pollinators, and leached fertilizers can harm streams and aquifers. As the arms race to produce super-crops increases, there is ever greater need to control the environment into which these seeds are planted. The risk to other than maximum crop yield must be forcefully eliminated as much as possible. This is what I mean by maximization. The flower and honeybee only have tools to optimize their risk and reward equation according to their risk appetite. They make decisions that seek to maximize their reward but within risk levels that they or the species can tolerate and only within circumstances they are capable of controlling. Obviously, humans have the technology to control much more of the environment than flowers and honeybees, but even then, humans cannot control the weather. What humanity does in present-day agriculture is to create an environment where natural and other risks are eliminated or reduced as much as possible. This produces higher yields per acre to be sure, but with what effect on entropy? The fundamental question I ask now and for which I do not have a good answer, is if life has so long existed making optimal decisions subject to ecological and other circumstances (with limited capabilities of altering those circumstances) and as a result this likely has produced and does produce more entropy than a configuration of the earth without life, what effects do increasing human maximization practices have on the earth’s entropic production? This includes the mostly non-vegetated areas like cities and roads, and the controlled vegetated areas like single-crop planting and suburban (single species) lawns. We suspect that human activity is increasing carbon dioxide in the atmosphere and see that the earth is warming at the same time. Science suggests

Study Summary and a Critique of Maximization

221

that these two outcomes are connected. I ask what amount of both outcomes is the product of human activity caused by fossil fuel use, and what also may be the result of the reduction of space where life can make optimal decisions in order to produce MEP? Even as I ask this question, it is with the understanding that there may be entropy processes at work that we do not yet understand or have not identified an adequate number of variables to make predictions. For example, Charles H. Lineweaver notes that the sun in earth’s earlier history was dimmer than it is today and thus could not warm the earth as it does today. He wonders whether early life’s contribution to methanogenesis, rather than cool the earth as entropy is more wont to do, actually served to warm the early earth and thereby reduced its entropy production.14 That may then have set the stage for even more life, leading towards a steady-state earth filled with life that produces MEP. From this speculation comes the inevitable question of what effect do the reduction in biomass and vegetated surface and the increase in sun’s radiation today (through its aging process) produce—global warming, or do we not yet know? Consider that abiotic surfaces of varying reflectivity are going to absorb or reflect light according to a formula based upon how much radiation each receives and its inherent reflective capability. This was true a billion years ago and today. Life, however, makes optimal decisions on how to produce and consume energy, based upon several factors, including and not limited to, the amount of direct radiation received. Plants can refrain from photosynthesis (producing energy) when searching for water which consumes energy through root growth. Honeybees may consume as much energy per individual in drought years when there are fewer flowers but produce less honey to winter over. The hive during the drought likely will reduce the number of foragers so that they do not waste energy on fruitless trips. The black rock that absorbs energy makes no decision; rock physics makes the decisions. Life makes optimal decisions based upon the circumstances of the environment (the same sun that hits the rock) but in context of its needs and its capabilities. Therefore, life presents a different dynamic for entropy than a pure rocky surface. We do not yet have the models that show just how optimization affects the entropy of biomass compared with abiotic or limited biotic areas of the globe. One question worth asking is whether increases in the cooler and more entropic meadow land mass can counteract the increases in carbon dioxide that serve to increase the temperature of the atmosphere. Obviously, an 14  C  harles H. Lineweaver, “Cosmology and Biological Reproducibility: Limits on the Maximum Entropy Production Principle,” in Non-Equilibrium Thermodynamics and the Production of Entropy: Life, Earth, and Beyond, ed. Axel Kleidon and Ralph D. Lorenz (New York: Springer Science & Business Media, 2004), 75.

222

Chapter 6

increase in carbon dioxide consuming plants will reduce carbon dioxide in the atmosphere, but how much … and, would both increases in biomass entropy and the reduction in atmospheric carbon through photosynthesis help to reduce climate change purportedly caused by carbon dioxide? Is increase in optimizing ‘meadow’ biomass even possible to any significant extent if human populations continue to grow? I am suggesting that a better understanding of the physics of optimization would help us better understand the role of life in the regulation of entropy on earth. A fundamental objection that will be raised is what I am recommending is a return to the human as hunter-gatherer. Rather, what I am asking is that we look at the role of entropy production in climate change. If we determine that nature as a maximum entropy producer has contributed to the delicate balance of earth’s steady state, we should consider how our activities affect entropy, not just how humans increase the amount of carbon dioxide in the atmosphere. Hans Jonas considered the history of technology. Early in our history, technology’s impact was local and, in many cases, individual. The blacksmith or cobbler made the shoe for one horse or one person. The dagger was an interpersonal weapon. Today factories make shoes and we have impersonal nuclear weapons. Therefore, until recently, technological impact was limited to local activities. Jonas also suggests that because technology was local and interpersonal, “advanced planning” was not an issue.15 Today, technology can affect generations far past our own. The question of intergenerational justice looms in the matter of climate change. We must begin to consider the longer-term implications of decisions we make today. If our decisions prove in the longer term to not meet expectations, we must then alter these practices, always with an eye to the longer term rather than the immediate reward. This study has shown how important optimization is to flowers and honeybees in their facultative mutualism. By extension we have also looked at research that suggests that MEP can be generated in other environments with other species of life, all who are making optimal decisions. Human development and our tendency to maximize by clearing away obstacles to gain reward, rather than work within the parameters those obstacles dictate, may produce different amounts of entropy compared to a life-filled world without humanity. Whether the net entropy produced by a life-filled world with humanity (today) is more or less than one without humanity, I am not able to offer an opinion. However, I suspect that there likely is a difference and that difference means 15  Hans Jonas, “Technology and Responsibility: Reflections on the New Tasks of Ethics,” Social Research 40, no. 1 (1973): 35.

Study Summary and a Critique of Maximization

223

something to the capability of the world to sustain life over the long-term. I am not naive enough to suggest that the species that wants ever more—will want willingly what the flower and honeybee can accept through their optimal decision-making process. However, there may be things we can do, do more of, or do less of to produce MEP through gaining more assistance from the world community of life that has kept the earth verdant for many years, even as the earth and the universe have tried to make life extinct many times. Cited References Danjuma, M. N., and S. Mohammed. “Zai Pits System: A Catalyst for Restoration in the Dry Lands.” Journal of Agriculture and Veterinary Science 8, no. 2 (2015): 1–4. Harding, B. The Doctrines of Jacob Boehm. New York: Macoy Publishing, 1919. Holdaway, Robert J., Ashley D. Sparrow, and David A. Coomes. “Trends in Entropy Production During Ecosystem Development in the Amazon Basin.” Philosophical Transactions: Biological Sciences 365, no. 1545 (2010): 1437–47. Jonas, Hans. “Technology and Responsibility: Reflections on the New Tasks of Ethics.” Social Research 40, no. 1 (1973): 31–54. Kleidon, Axel, and Ralph D. Lorenz. “Entropy Production by Earth System Processes.” Chap. 1 In Non-Equilibrium Thermodynamics and the Production of Entropy: Life, Earth, and Beyond, edited by Axel Kleidon and Ralph D. Lorenz, 1–20. New York: Springer Science & Business Media, 2005. LaCanne, Claire E., and Jonathan G. Lundgren. “Regenerative Agriculture: Merging Farming and Natural Resource Conservation Profitably.” PeerJ, no. 2018 (2018/02/26 2018): 1–12. doi:10.7717/peerj.4428. Lineweaver, Charles H. “Cosmology and Biological Reproducibility: Limits on the Maximum Entropy Production Principle.” In Non-Equilibrium Thermodynamics and the Production of Entropy: Life, Earth, and Beyond, edited by Axel Kleidon and Ralph D. Lorenz. New York: Springer Science & Business Media, 2004. Loy, David. Nonduality: A Study in Comparative Philosophy. Amherst, NY: Prometheus Books, 2012. Moussa, Boubacar M., Abdoulaye Diouf, Salamatou I. Abdourahamane, Jørgen Aagaard Axelsen, Karimou J. M. Ambouta, and Ali Mahamane. “Combined Traditional Water Harvesting (Zaï) and Mulching Techniques Increase Available Soil Phosphorus Content and Millet Yield.” Journal of Agricultural Science 8, no. 4 (2016): 126–39. Zougmoré, Robert, Abdulai Jalloh, and Andre Tioro. “Climate-Smart Soil Water and Nutrient Management Options in Semiarid West Africa: A Review of Evidence and Analysis of Stone Bunds and Zaï Techniques.” Agriculture & Food Security 3, no. 1 (2014/11/03 2014): 1–8. doi:10.1186/2048-7010-3-16.

Index Abdomen (Honeybee) 63, 65 Ackerly, David, carbon gain, self-shading 17 Affordances and Constraints 16, 20, 21 Aigner, Paul, Most Effective Pollinator Principle 18 Alexander, Richard D., morality and natural selection (see also Morality) 176 Allem, Antonio C. outbreeders and self-breeders 18 Altruism 163–174 Altruistic punishment 163, 167–174 Negative emotion 169 Behavioral 163, 165–167 Biological 163, 165 Para-Psychological 166–167 Psychological 163, 165, 166–167 Sociobiology 164 Angiosperms (see also Wang, Zin, Chamovitz, Daniel) 44 Awareness 85  Consciousness (see also Endel Tulving)  85–86, 117 Gravity 83–84 Memory 84–85  Pain 81 Plant knowing 76 Plants and light 77–79 Circadian rhythms 79 Cryptochromes 79 Light and photosynthesis 78 Phototropism and blue light 77 Ultraviolet and infrared 77 Smell 80 Sound (see Gagliano, Monica) Tactility 81 Behavior Growth thoughts 98 Intelligence, Distributed Intelligence  95–99  Processes 73, 75 Salicylic acid 80 Flower morphology 59–60 Bell, Adrian 59 Defined 50

Gynoecium 60 Light-sensitive cells, gravity negative  72 Nectaries 60 Ovary 60 Ovule 60 Pollen tube 60 Roots 72 Stamens 60 Flower Origin Euanthial 48 Pseudanthial 38 Fossil record 45 Cretaceous Origin 50 One common ancestor 47 Stages for angiosperm development  46 Plant Morphology 56–59 Axillary inflorescence 58 Buds 59 Mycorrhizae 57, 177 Roots 57 Stem 57 Vascular plants defined 45 Closed carpel 46 Important contributors to angiosperm success 47 Seed plant 46 Vascular bundles 46 Antem, Niels P. R. forest canopies, canopy models, competition theory 19 Antennae 64 Anthropocentric, Anthropomorphic 8, 101 Antinaturalism fallacy (see also Moore, G. E., Naturalism) 179 Appetitive states 187 Apis Mellifera (see also Honeybees) 44 Apoid Wasp 6, 35–36, 49, 52, 149, 158 Asymmetrical, asymmetry (see also Hospitality, Mutualism, Responsibility)  6, 36, 100, 123, 144–151, 145, 147, 150, 152, 153, 155, 161, 164, 184, 195 Asynchronous 36, 39, 153 Axillary inflorescence 58

226 Bateson, Melissa, honeybee emotion, shaken bees (see also Honeybee behavior)  113, 169 Bee (see Honeybee) Beiler, Kevin J., tree hubs 86–87, 177 Benard cells 26 Bernal, Autumn, J. and Randy L. Jirtle, negative epigenetics (see also Epigenetics)  136–137 Bird, Adrian (see also Epigenetics) Epigenetics defined 133, 136 Promise of epigenetics 134 Blackburn, Simon, appetitive states 187 Boehm, Jacob, timelessness 212 Boyko, Alexander, tobacco plant epigenetics  135 Bronstein, Judith L., mutualism (see also Mutualism) 34, 36 Buddha, Buddhism Buddha nature 213 Dukkha 92 Buds 59 Bump dance (see Honeybee) Buzz pollination 48, 83 Canopies, canopy models, competition theory 19 Capellari, S. C., bee emergence with eudicots  150 Carpel (see Angiosperms) Castes (see Honeybees) Centrifugal 123, 145, 147–148, 150, 158, 160 Centripetal 123, 145–146, 150–151, 158, 160 Chamovitz, Daniel (see also Angiosperms) Awareness 85  Consciousness (see also Endel Tulving)  85–86, 117 Gravity 83–84 Memory 84–85  Pain 81 Plant knowing 76 Plants and light 77–79 Circadian rhythms 79 Cryptochromes 79 Light and photosynthesis 78 Phototropism and blue light 77 Ultraviolet and infrared 77

Index Plants and sound (see Gagliano, Monica) Smell 80 Tactility 81 Chemotaxis 32 Chittka, Lars, bee vision (see also Honeybee)  19–20, 20 Circadian rhythms 79 Civil War Study of epigenetics (see also Epigenetics, Costa, Dora L., Noelle Yetter, and Heather DeSomer) 137 Coevolution 34, 35–36, 38, 52, 54, 69, 125, 153, 155, 184 Cognitive nonconsciousness (see also Hayles, N. Catherine) 118–123 Commerce 8, 145, 153 Consciousness (see Tulving, Endel, Chamovitz, Daniel) Continuity 1, 28 Costa, Dora L., Noelle Yetter, and Heather DeSomer, Civil War POWs and epigenetics 137 Cretaceous 50 Crop (Honeybee) 65 Cryptochromes 79 Curry, Oliver, moral values 187 Darwin, Charles Mutation 132 Phototropism 77 Derrida, Jacques, hospitality (see also Hospitality) 160 de Wall, Frans 7 Dewar, Roderick C., maximum entropy production, (see also Maximum entropy production) 31 Dewey, John, morality (see also Morality)  195–196 DNA 7 Dualism, Non-Dualism Descartes, Rene 95 Schopenhauer, Arthur 95 Embedded 54 Embodied 54 Emergence 23, 25 Benard cells (see also Benard cells) 26 Definition 25

Index Entropy Maximum entropy 4 Maximum Entropy Production (MEP) (see also Maximum entropy production) 4, 11, 23–33 Epigenetic, epigenetics 1, 132–143 (see also, Popova, Evgenya and Barnstable, Colin J.; Bird, Adrian; Civil War Study of Epigenetic; Janusek, Linda Witek; Ney, Gideon; Pieterse, Corné M. J.; Sarkies, Peter) Above genetics 10 Defenses 28 Defined 28, 132–133 Epigenetic Rules (See also Ruse, Michael and E. O. Wilson) 144, 179–186, 198–199 Mechanisms 134 Negative aspects 136–138 Promise of 134–135 Purposes 134–137 Ethics (see Morality) Ethylene 72, 79, 139 Euanthial 48 Eukaryote cell 150 Eusocial (see also Honeybee) 61–63, 150 Honeybee castes 61–62, 66–67 Males (drones) 62, 66, 103 Queen 62, 66, 103 Workers 62, 66–67, 103 Overlapping maturation generations 61 Existence before essence 97 Facilitative (mutualism) 35–36 Facultative (mutualism) 2–3, 5, 34, 125 Fehr, Ernst and Simon Gachter, altruistic punishment (see also Altruism)  164, 167–174 Flower (see also Angiosperm) Definition 50  Focused intentionality 55 Friis, Elsa Marie (See also Angiosperms, Honeybee) Bee evolution 51 Flower origin Euanthial 48 Pseudanthial 48

227 Gagliano, Monica, Plants and sound (see also Angiosperms) 82–83 Genetics and Mutation 28 Gestalt 100, 125 Giurfa, Martin, how insects do what they do  72 Good, The Good 1, 11, 22, 101, 171, 173, 188–194 Gynoecium 60 Hayles, N. Catherine Cognition 88 Cognitive nonconsciousness 118–123 Thinking 88 Head (Honeybee) 63 Heidegger, Martin  Being 89–91 Thinking 89 Hive, Hive society, Hive personality (see Honeybee) Hobbes, Thomas 2 Holdaway, Robert J., entropy, forests v. pastures 32, 215–216 Honeybee Anatomy Abdomen 63, 65 Antennae 64 Bee vision (see also Chittka, Lars)  19–20, 20 Compound eyes 63 Ultraviolet 63 Crop 65 Digestive organs 65 Exoskeleton 65 Head 63 Legs 64–65 Mandibles 63 Mushroom bodies 64 Pollen Baskets 65 Proboscis 63 Stinger 65 Thorax 63–64 Wings 64–65 Behavior Dances Bump dance 62, 106, 109–111 Dance floor 102

228 Honeybee, Behavior, Dances (cont.) House hunting dance 106, 114–117 Consensus 115–116 Tremble dance 106, 108 Waggle dance 106–108 Distal intelligence 100 Emotion (see also Bateson, Melissa)  113 Hive guarding 21 Hive society 102 Hive personality 113 Landscape memory 105 Learning flight 105 Maturation 20 Negative feedback 109, 170 Personality 111–113, 176 Scout bee (swarm) 66 Swarm 66 Trophallaxis 62, 102, 105, 108–109, 113–114 Castes 61–62, 66–67 Males (drones) 62, 66, 103 Queen 62, 66, 103 Workers 62, 66–67, 103 Evolution 51 Origin 106–107  Hospitality 160–162 Co-hospitality 161 Reciprocal hospitality 195, 212 House-hunting dance (see Honeybee) Hume, David (see also Naturalistic fallacy, Moore, G. E.) Ought exclusively from is 10, 173, 186–188, 198 Passion and reason 168, 172–173, 187, 195 Hymenoptera 7, 9, 50–51, 63 Intelligence, Distributed Intelligence  95–99 Ipseity 54, 157 Janusek, Linda Witek, negative epigenetic effects (see also epigenetics) 138 Johnson, Brian R. and James C. Nieh, waggle dance (see also Honeybee) 107 Jonas, Hans, technology 222

Index Ketcham, Christopher, will (see also Schopenhauer, Arthur) 97 Kitcher, Philip (see also Naturalism) Narrow sociobiology 164 Pragmatic naturalism 162–165, 197 Klausmeier model of maximum entropy production (see also Maximum entropy production) 33 Kleidon, Axel, Maximum Entropy Production (see also Maximum entropy production) 24, 216 With James Dyke 24 With Ralph Lorenz 3 LaCanne, Claire E. and Jonathan G. Lundrgen, regenerative farming 217–218 Language (see Angiosperm, Honeybee, Wittgenstein) Legs (honeybee) 64–65 Leoncini, Isabelle, retardation of maturation of honeybees (see also Honeybee) 113 Levinas, Emmanuel (see also Responsibility) Face 157–158 Passivity 153 Peace and war 154 Reciprocity 152–153, 211 Responsibility 152, 154, 156, 158, 211, 213 Substitution 153, 159 Light (see Angiosperms) Linear thermodynamic system (see also Non-equilibrium system) 4 Lineweaver, Charles H., sun’s history 221 Mandibles (Honeybee) 63 Marder, Michael, plant existentiality  12, 74, 94, 98, 123, 147, 149, 159 Martyushev, Leonid, maximum entropy production (see also Maximum entropy production) 29, 32 Maximization 1, 11, 215, 217, 220 Maximum Entropy Production (MEP)  23–33, 74, 154 (see also Dewar, Roderick C., Holdaway, Robert J., Kleidon, Axel, Martyushev, Leonid, Robert J., Meysman, Philip J., Vallino, Joseph J., Volk, Tyler) Menzel, Randolph, flower reliability  104, 106

229

Index Merleau-Ponty, Maurice Intentionalities 55, 66–67 Time 67 Meysman, Filip J. R. and Stijn Bruers (see also Maximum entropy production) Maximum Entropy Production (see also Maximum entropy production)  25, 27, 29 Schrodinger and the second law of thermodynamics 25 Michez, Vaderplanck, and Engel, wasp evolution (see also Apoid wasp) 52–53 Middle, plant 94 Mommer, Liesje, root response to competition 176 Moore, G. E. Good, the good (see also Good, the good)  173, 189–194 Morality and ethics 189, 192–193 Naturalistic fallacy 11, 144, 173, 188, 191, 193 Morality Descriptive and normative 154, 200–201 Co-equal 39 Elegance 126–127 Emergent property 8, 149, 174, 214 Levinas 152, 159  Moral actions 184 Mutualism 145 Naturalistic 10, 162 Obligation 182 Reciprocity 69, 152 Most Effective Pollinator Principle 18 Mushroom bodies (Honeybee) 64 Mutation 54 Mutualism 33–39 (see also Bronstein, Judith) Coevolution 34, 35–36, 38, 52, 54, 69, 125, 153, 155, 184 Definition 34 Facilitative 35–36 Facultative 2–3, 5, 34, 125 Asymmetrical needs 100, 144 Benefit and exploit 39, 144 Intelligence of 99–100 Indirect 34 In nature 33

Non-exclusive 3, 6, 35–36 Obligate 34 Mycorrhizae (see also Angiosperms)  5, 91, 177 Nancy, Jean Luc, being singular plural  54–55 Naturalism Anti-naturalism fallacy (see also Walton, Alexander) 194–196 Natural Facts 184, 197 Pragmatic Naturalism (see also Kitcher, Philip) 162–165, 197 Naturalistic fallacy (see also Moore, G. E.)  11, 144, 173, 188, 191, 193 Nealon, Jeffrey T., plant theory 9, 74 Nectaries (see also Angiosperms) 51 Newton, Isaac Second law of thermodynamics 3, 23, 24 Prigogine, Ilya and Isabelle Stengers, nonequilibrium 23 Ney, Gideon and Johannes Schul, epigenetics in katydids 136 Nieh, James C., negative feedback waggle dance (see also honeybee) 109–111 Non-Duality, Duality 93–95 Non-equilibrium system (see also Linear thermodynamic system) 3, 24 Non-interference 3 Non-mindful mindfulness 149 Obligate (mutualism) 34 Obligation (moral) 182 O’Gorman, Kevin D., hospitality (see also Hospitality) 160 Oligolectic 51 Optimization, optimize, optimal, optimality  1–2, 4, 16–23, 27, 101, 153–154, 192, 195, 220–222 Best decision 17, 158, 164, 184 Definition 16 Identifying and assessing variables  17, 200 Towards the good 196 Otherness, the other 156 Oughts and Obligates for flowers and honeybees 197–199 Outbreeders and self-breeders 18

230 Ovary 60 Ovule 60 Page, Robert E., polyandry 113 Parmenides, being (see also Heidegger, Martin) 89 Passivity 157 Perpetuity 206 Personality (see Honeybee) Photosynthesis 78 Pieterse, Corne M. J., epigenetic purposes (see also Epigenetics) 135 Plant (for complete detail see Angiosperms) Plant theory (see also epigenetics) 12 Marder, Michael 12 Nealon, Jeffrey T. 12 Prigogine, Ilya 12 Stengers, Isabelle 12 Pollan, Michael, human spread of plants  80 Pollen (see Angiosperms) Pollen Baskets (Honeybee) 65 Pollen tube (Angiosperms) 60 Pollen value to pollinators 50 Pollination (see also Angiosperms) Self-pollinate 38 Water 38 Wind 38 Polyandry 113 Polylectic 51 Popova, Evgenya and Colin J. Barnstable, mechanisms of epigenetics (see also Epigenetics) 134 Proboscis (Honeybee) 63 Pseudanthial 38 Ramo-Fernandez, Laura, negative aspects of epigenetics (see also Epigenetics) 137 Reason and judgment (see Singer, Peter) Reciprocal (responsibility) 145, 154–156, 159–160–161, 195, 213 Reciprocal Gaze 54, 145 David Loy 212 Reciprocity 69, 152 Regenerative farming 217–219 Reliability, flower 104, 106 Responsibility (see also Levinas, Emmanuel)  151–154

Index Commerce 145 Common resource 145 Mutualism 160 Reciprocal 145, 154–156, 159–160–161, 195, 213 Restraint (see Singer, Peter) Riggs, Arthur, epigenetics defined  (see also epigenetics) 133 Risk 2 Appetite 216, 220 Roots 72 Rueppell, Olav, honeybee self-removal 166 Ruse, Michael and Edward O. Wilson (see also Epigenetics) Define ethics 182 Epigenetic rules 144, 179–186, 198–199 Sarkies, Peter, epigenetics purposes (see also Epigenetics) 134–35 Sartre, Jean Paul, existence before essence  97 Scarcity 155, 213 Schopenhauer, Arthur, thing in itself 96 Schymanski, Stanislaus, J. Klausmeier model of maximum entropy production (see also Maximum entropy production) 33 Scout (Honeybee) 66 Second law of thermodynamics 3, 23, 24 Selfish genes 28 Sight (see Honeybee) Simard, Suzzane W., mycorrhizae (see also Angiosperms) 57 Singer, Peter (see also Altruism) 174–179 Altruism 174 Three requirements for morality 2, 22, 62 Social Group 34, 38–39, 160, 163, 165, 174, 178 Reason and judgment 16, 22, 34, 38–39, 55, 165, 177–178 Restraint 2, 22, 34, 38, 55, 160, 165, 175 Smell (see Angiosperms, Honeybee) Social group (see also Peter Singer) 5 Sociobiology (See also Wilson, E. O.) 164 Soltis, Douglas (see also Angiosperms) Evolutionary and phylogenic facts about angiosperms 46–47

231

Index Pollinator attraction methods of flowers  49–50 Woody or aquatic origin 50 Sound (see Angiosperms, Honeybee) Stamen (see also Angiosperms) 44 Stem (plant) 57 Sticky pollen (see also Angiosperms)  35, 44, 48–49, 149, 158 Stinger (Honeybee) 65 Stromatolites 58, 75 Swarm (Honeybee) 66 Swenson, Rod Emergence 25 Coherence 26 Synthetic unity 148–149, 150 Tactility 81 Technology 222 Teehan. John and Chris DiCarlo, moral judgment and clear grasp of situation  191 Tezze, Andrea A. and Walter M. Farina, trophallaxis (see also Honeybee) 114 The good (see Good, the good) Thorax (Honeybee) 63–64 Triangulation, honeybee direction finding (see also Honeybee) 104 Trophallaxis (see Honeybee) Tudge, Colin, ethylene and ripening (see also Angiosperms) 79 Tulving, Endel, consciousness (see also Chamovitz, Daniel) Episodic 85, 118 Procedural 85, 117 Semantic 85, 117 Vallino, Joseph J., maximum entropy production (see also Maximum entropy production) 30–32 Vascular plants defined 45 Volk, Tyler and Olivier Pauluis, maximum entropy production (see also Maximum entropy production) 29, 30 von Frisch, Karl, honeybee language (see also Honeybee) 102

Waddington, Keith D., honeybee personality (see also Honeybee) 112 Waggle dance (see also Honeybee) 37 Walter, Alexander, antinaturalism fallacy (see also Moore, G. E., Naturalism)  179, 194–196 Walton, Alexander and Amy L. Toth, honeybee personality (see also Honeybee) 112, 176 Wang, Xin (See also Angiosperm) Flower Defined 50 Fossil record angiosperms 45 Cretaceous Origin 50 One common ancestor 47 Stages for angiosperm development  46 Vascular plants defined 45 Closed carpel 46 Important contributors to angiosperm success 47 Seed plant 46 Vascular bundles 46 Wasp (see Apoid wasp) Will (see also Schopenhauer, Arthur) 97 Wilson, David Sloan, Anti-naturalistic fallacy (see also Moore, G. E.) 194–196 Moral systems 162–163 Ought from is (see also Hume, David)  186 Wilson, Dietrich, and Clark, inappropriate use of naturalistic fallacy (See also Moore, G. E., Naturalism) 187–188 Wilson, E. O., Sociobiology 164 Wings (Honeybee) 64–65 Wittgenstein, private language 175 Wohlleben, Peter, tree interdependence  86–87, 165 Wray, Margaret K., hive personality (see also Honeybee) 113, 116 Wyschogrod, Edith, passivity and ipseity (see also Levinas, Emmanuel) 157 Zai 218 Županović, Paško, chemotaxis 32