Sex and Death: An Introduction to Philosophy of Biology

The University of Chicago Press, Chicago 60637

The University of Chicago Press, Ltd., London

© 1999 by The University of Chicago

All rights reserved. Published 1999

14 13 12 11 10        5 6 7 8

ISBN: 0-226-77303-5 (cloth)

ISBN: 0-226-77304-3 (paper)

Figure 6.3 is from Karen Arms and Pamela S. Camp, Biology, 3d ed. © 1987 by Saunders College Publishing. Reproduced by permission of the publisher. Figure 9.4 is from W. Hennig, Phylogenetic Systematics, © 1979 by the Board of Trustees of the University of Illinois. Reproduced by permission of the University of Illinois Press.

Library of Congress Cataloging-in-Publication Data

Sterelny, Kim.

    Sex and death : an introduction to philosophy of biology / Kim Sterelny and Paul E. Griffiths.

        p.      cm. — (Science and its conceptual foundations)

    Includes bibliographical references (p.      ) and index.

    ISBN 0-226-77303-5 (cloth : alk.paper). — ISBN 0-226-77304-3 (pbk.: alk. paper)

    1. Biology—Philosophy    I. Griffiths, Paul E.    II. Title.    III. Series.

QH331.S82    1999

570'.1—dc21

98-47555

CIP

The paper used in this publication meets the minimum requirements of the American National Standard for Information Sciences—Permanence of Paper for Printed Library Materials, ANSI Z39.48-1992.

ISBN: 978-0-226-17865-3 (e-book)

Sex and Death

An Introduction to Philosophy of Biology

Kim Sterelny and Paul E. Griffiths

The University of Chicago Press

Chicago and London

Science and Its Conceptual Foundations

A series edited by David L. Hull

to Melanie and Kate

Contents

Preface

Part I. Introduction

1. Theory Really Matters: Philosophy of Biology and Social Issues

1.1. The Science of Life Itself

1.2. Is There an Essential Human Nature?

1.3. Is Genuine Altruism Possible?

1.4. Are Human Beings Programmed by Their Genes?

1.5. Biology and the Pre-emption of Social Science

1.6. What Should Conservationists Conserve?

2. The Received View of Evolution

2.1. The Diversity of Life

2.2. Evolution and Natural Selection

2.3. The Received View and Its Challenges

Part II. Genes, Molecules, and Organisms

3. The Gene’s Eye View of Evolution

3.1. Replicators and Interactors

3.2. The Special Status of Replicators

3.3. The Bookkeeping Argument

3.4. The Extended Phenotype

4. The Organism Strikes Back

4.1. What Is a Gene?

4.2. Genes Are Active Germ Line Replicators

4.3. Genes Are Difference Makers

5. The Developmental Systems Alternative

5.1. Gene Selectionism and Development

5.2. Epigenetic Inheritance and Beyond

5.3. The Interactionist Consensus

5.4. Information in Development

5.5. Other Grounds for Privileging Genes

5.6. Developmental Systems and Extended Replicators

5.7. One True Story?

6. Mendel and Molecules

6.1. How Theories Relate: Displacement, Incorporation, and Integration

6.2. What Is Mendelian Genetics?

6.3. Molecular Genetics: Transcription and Translation

6.4. Gene Regulation

6.5. Are Genes Protein Makers?

7. Reduction: For and Against

7.1. The Antireductionist Consensus

7.2. Reduction by Degrees?

7.3. Are Genes DNA Sequences Plus Contexts?

7.4. The Reductionist Anticonsensus

Part III. Organisms, Groups, and Species

8. Organisms, Groups, and Superorganisms

8.1. Interactors

8.2. The Challenge of Altruism

8.3. Group Selection: Take 1

8.4. Group Selection: Take 2

8.5. Population-Structured Evolution

8.6. Organisms and Superorganisms

9. Species

9.1. Are Species Real?

9.2. The Nature of Species

9.3. The One True Tree of Life

9.4. Species Selection

Part IV. Evolutionary Explanations

10. Adaptation, Perfection, Function

10.1. Adaptation

10.2. Function

10.3. The Attack on Adaptationism

10.4. What Is Adaptationism?

10.5. Structuralism and the Bauplan

10.6. Optimality and Falsifiability

10.7. Adaptation and the Comparative Method

11. Adaptation, Ecology, and the Environment

11.1. The Received View in Ecology

11.2. History and Theory in Ecology

11.3. The Balance of Nature

11.4. Niches and Organisms

11.5. Reconstructing Niches

11.6. Unfinished Business

12. Life on Earth: The Big Picture

12.1. The Arrow of Time and the Ladder of Progress

12.2. Gould’s Challenge

12.3. What Is Disparity?

12.4. Contingency and Its Consequences

12.5. Mass Extinction and the History of Life

12.6. Conclusions

Part V. Evolution and Human Nature

13. From Sociobiology to Evolutionary Psychology

13.1. 1975 and All That

13.2. The Wilson Program

13.3. From Darwinian Behaviorism to Darwinian Psychology

13.4. Evolutionary Psychology and Its Promise

13.5. Evolutionary Psychology and Its Problems

13.6. Memes and Cultural Evolution

14. A Case Study: Evolutionary Theories of Emotion

14.1. Darwin on the Emotions

14.2. Sociobiology and Evolutionary Psychology on the Emotions

14.3. The Modular Emotions

14.4. Beyond the Modular Emotions

14.5. Emotion, Evolution, and Evolved Psychology

Part VI. Concluding Thoughts

15. What Is Life?

15.1. Defining Life

15.2. Universal Biology

15.3. Simulation and Emergence

Final Thoughts

Glossary

References

Index

Preface

This book began long ago and far away, in Chicago in 1993 when one of us (Sterelny) tried out the basic idea on David Hull and Susan Abrams, both of whom were supportive. On Sterelny’s return to the Antipodes, he continued to think about the project, and decided that a collaborative project would be more fun to do and would result in a better book. So he talked the idea over with Griffiths and with David Braddon Mitchell, an Australian philosopher of science with interests in both philosophy of biology and Australian botany. Thus the basic body plan of the book was laid down; a body plan that, in contrast to some others, has not remained impervious to developmental and other perturbations. After a couple of years of talking, we seriously got down to writing in 1995. Since 1996 this book has probably been the main project of the two survivors, Braddon Mitchell having been submerged by other plans. He did, however, have a major input into chapter 12.

There are, of course, many different ways to write an introduction to philosophy of biology. One option would be to use biological examples to stalk general issues in philosophy of science—the nature of theory and theory change, causation, explanation, and prediction. There is much to be said for such a book, for philosophy of science, in our view, has been too dominated by exemplars from theoretical physics. That matters: for example, the historical explanations central to, say, geology and evolutionary biology seem importantly different from those of physics. Still, that is definitely not the book we have written. This book is very much focused on the conceptual and theoretical problems generated by the agenda of biology, rather than pursuing a philosophy of science agenda through biological examples.

We have also chosen not to approach philosophy of biology by tracking the conceptual and theoretical development of evolutionary ideas, as David J. Depew and Bruce H. Weber have done in their Darwinism Evolving. There is an occasional nod to the history of the disciplines concerned, but the organizational spinal cord of our book is the conception of evolutionary biology that was developed in the classic works of Mayr, Dobzhansky, Simpson, and Stebbins in the 1940s. That conception, the modern synthesis, dominated evolutionary thinking at least into the late 1960s. The current problems of evolutionary theory have been largely, though not wholly, the result of pressures to rethink that conception. We have chosen to call this core conception the received view rather than the synthesis view because we represent it in a rather schematic and ideal form. The real synthesis was never wholly uniform, of course, and for the most part the variation within it has not been our concern.

We have called our book Sex and Death: An Introduction to Philosophy of Biology. First, the subtitle: The reader may have noticed that while it speaks of biology, in the preface we have written of evolutionary biology. Indeed, in the text we have focused on evolutionary biology. That focus is not exclusive: chapter 5 explores connections between evolutionary and developmental biology, and chapters 6 and 7 push this exploration further. Those chapters take up the relationship between the role of genes in evolutionary theory and the molecular biology of the gene. Moreover, chapter 11 is devoted to exploring the interplay between evolutionary biology and ecology. But it is true that we have discussed other areas of biology mainly as they relate to evolution. So evolutionary theory and evolutionary theorists loom large over this work. (So we too can say that if we have seen too little, it is because giants have been standing on our shoulders.) This emphasis is partly, we think, a reflection of the genuinely conceptually challenging nature of evolutionary theory. As we show (we hope) in the text, evolutionary theory really poses a striking compound of conceptual and empirical problems. But it is partly a historical accident, too. We have no doubt that there are similar problems in ecology, developmental biology, and molecular biology (at least), and we hope to have done at least a little to extend the reach of philosophy of biology into those areas.

Second, the title: We chose the title because it was fun. And philosophy of biology is fun. The living world is splendid and bizarre—far more bizarre than we, at least, could have imagined—and the conceptual problems posed in understanding it are wonderfully intriguing (and important, as we argue in chapter 1). We hope this book shows that. We nurture the illusion that it will both manifest our relish in the subject and perhaps infect others with the same disease.

The structure of the book is, we hope, evident from the analytic table of contents. Part 1 sets out the scope of the project. Parts 2–4 work through the core debates, as we see them, in evolutionary theory and associated branches of biology. In part 2 genes are at center stage: we discuss both the idea that evolutionary history is really, fundamentally the history of gene lineages and the relationship between the evolutionary and molecular understanding of genes. In part 3 the focus changes to organisms, groups, and species. An important connecting thread is the question of whether groups and species play a role in evolution importantly like that played by organisms. Natural selection is central to part 4, for that part is on evolutionary explanation, and the key controversy about evolutionary explanation is the role of selection. So in a sense parts 2–4 are the heart of the book. Part 5 takes up human evolution, and more particularly, the sociobiological debates and their relatives. Apart from the intrinsic interest of this subject, many of the issues about the nature of evolution and natural selection are nicely exemplified through their application to humans. Part 6 winds up the show. We here attempt to put the central debates about the nature of evolutionary processes and patterns in a broader context by asking whether the characteristic patterns and processes of life on earth are likely to be features of any living world.

We have tried to write a book for three audiences. We wanted a book that would be accessible both to biology students with little or no philosophy and philosophy students with little or no biology. So we have used as little technical jargon as possible. When we have used specialist terminology from either discipline we have explained it in the text (usually immediately after the term’s introduction) and, often, included it in the glossary. We have also made fairly liberal use of boxes in the text to discuss and explain more technical material. We have, however, tried to write the text so that no box is essential to following the flow of the argument. So readers should be able to skip the boxes if they like without losing the thread of the ideas. Many of the issues discussed in the book interconnect, one with the other. We have tried to help the reader follow these connections with parenthetic guides; for instance, (5.3) would indicate that the issue in play will be, or was, discussed further in section 5.3. Finally, we have provided Further Reading sections at the end of chapters 2–15 to introduce and orient newcomers to the literature.

So we hope this book is accessible to both philosophers and biologists without previous experience of the other area. Our third intended audience is, of course, our peers. This book is not a view from nowhere. It’s an introduction to philosophy of biology from our own perspective on the discipline. So it contains our own assessment of what matters and what does not—of what is central and what is peripheral. That perspective is not widely shared, for we are products of a hybrid zone (displaying, we hope, hybrid vigor rather than hybrid sterility). Our take on evolution integrates important elements of the adaptationist, gene-centric conception of evolution associated with the likes of Maynard Smith, Williams, and Dawkins with elements of the pluralist, hierarchical conception associated with Gould, Lewontin, and Eldredge. At the very least, we would like to convince others in the field that the space of viable options is larger than they might have supposed.

This book took a lot of writing, and we got a lot of help with that writing. First we would like to thank David Hull and Susan Abrams for their initial enthusiasm and continued support for the project. Second, we owe a lot to the opportunity to talk biology and philosophy of biology, over many years and on many occasions, with the following: Russell Gray, Peter Godfrey-Smith, Susan Oyama, Geoff Chambers, David Hull, Karen Neander, Michael Hannah, David Braddon Mitchell, and Lenny Moss. They helped to provide the intellectual matrix from which this book has grown. Third, Peter Godfrey-Smith, Geoff Chambers, David Hull, Elliot Sober, Richard Francis, and two University of Chicago Press reviewers read and commented extensively on the semi-final manuscript. To them we owe much: thanks (not enough; if you’re lucky you might get a beer as well). We thank Dan McShea, Susan Oyama, David Sloan Wilson, Alan Musgrave, James Maclaurin, Mike Dickison, Karola Stotz, Werner Callebaut, and Annemarie Jonson for reading and commenting on sizeable chunks of that same draft. We have three more specific thanks to give. First, we shamelessly borrowed, though with permission, the title of chapter 1 from R. D. Gray and J. L. Craig, Theory really matters: Hidden assumptions in the concept of habitat requirements (1991). Second, chapter 4 owes a lot to Griffiths’s collaborators on other publications, Robin D. Knight and Eva M. Neumann-Held. Third, chapter 12 owes much to David Braddon Mitchell.

There are some equally valuable nonintellectual inputs to acknowledge. Griffiths’s thanks go to his former home, the University of Otago, for the outstandingly supportive research environment in the Department of Philosophy and for Richard Briscoe’s valuable services as research assistant. Also to his present home, the University of Sydney, where he taught two courses based around this book and employed another indefatigable research assistant, Ross West, who prepared the illustrations.

Sterelny inflicted numerous extraordinarily rough drafts of various chunks of this book on students at Victoria University of Wellington in 1995 and 1996 and at the California Institute of Technology, also in 1996. He thanks them for suffering so patiently. He also thanks the Philosophy Department, Monash University for hospitality and support in 1994; the Philosophy and Law Program, RSSS at the Australian National University for similar hospitality and support in 1995, and Caltech for providing a home in 1996. His base institution, Victoria University of Wellington, has supported the project in many ways. It provided a grant for research assistance in 1997, which enabled him to employ James Mansell, who worked with great intelligence and enthusiasm in finding and tracking down references (thanks, James). It granted leave to visit the ANU in 1995 and, for a more extended period, Caltech in 1996. Most importantly, it remains a civilized and supportive environment in which to work. Finally, he would also like to thank Melanie Nolan for her (mostly) tolerant attitude to his various preoccupations with biology, preoccupations especially marked in the final burst of writing and rewriting this work.

Kim Sterelny, Wellington, New Zealand

Paul Griffiths, Sydney, Australia

Part I

Introduction

Chapter 1

Theory Really Matters: Philosophy of Biology and Social Issues

1.1   The Science of Life Itself

The results of the biological sciences are of obvious interest to philosophers because they seem to tell us what we are, how we came to be, and how we relate to the rest of the natural world. The media often report that scientists have discovered the original purpose of some common human trait—morning sickness during pregnancy is designed to prevent malformed fetuses (Profet 1992). Or a traditional but controversial claim about society is found to be a biological fact—boys are more prone to violence and in greater need of formal social training than girls. And the gene for this difference has been localized—the genes for good social adjustment are on the paternally derived X chromosome, which only girls receive (Skuse et al. 1997). In all these cases biology seems to yield clear factual answers to questions of enormous moral and social significance.

In the late nineteenth and early twentieth century many philosophers looked to biology for answers to basic questions of ethics and metaphysics. Herbert Spencer’s evolutionary synthetic philosophy was the most influential philosophical system of its time. Friedrich Nietzsche, hero of today’s post-modernists, believed that Darwin’s theory could demolish traditional views of humanity’s significance in the overall scheme of things. In America, the pragmatist Charles Saunders Peirce investigated the implications of evolution for the nature and limits of human knowledge. But mainstream philosophy in the universities of the English-speaking world took a very different view. At the opening of the twentieth century Bertrand Russell declared that the theory of evolution had no major philosophical implications. The sciences that had something to teach philosophy were mathematics (particularly mathematical logic) and physics. Physics was to serve as a role model for the other sciences, and for the next fifty years philosophers nagged biology for its failure to live up to its example. The well-known philosopher of science and mind J. J. C. Smart compared the biologist to a radio engineer. Biologists study the workings of a group of physical systems that happen to have been produced on one planet. Smart thought that such a parochial discipline was unlikely to add to our stock of fundamental laws of nature (Smart 1963).