Evolution in Four Dimensions Life and Mind: Philosophical Issues in Biology and Psychology Kim Sterelny and Robert A. Wilson, editors Cycles of Contingency: Developmental Systems and Evolution Susan Oyama, Paul E. Griffiths, and Russell D. Gray, editors, 2000 Coherence in Thought and Action Paul Thagard, 2000 Evolution and Learning: The Baldwin Effect Reconsidered Bruce H. Weber and David J. Depew, 2003 Seeing and Visualizing: It’s Not What You Think Zenon Pylyshyn, 2003 Organisms and Artifacts: Design in Nature and Elsewhere Tim Lewens, 2004 Molecular Models of Life: Philosophical Papers on Molecular Biology Sahotra Sarkar, 2004 Evolution in Four Dimensions: Genetic, Epigenetic, Behavioral, and Symbolic Variation in the History of Life Eva Jablonka and Marion J. Lamb, 2005 Evolution in Four Dimensions Genetic, Epigenetic, Behavioral, and Symbolic Variation in the History of Life Eva Jablonka and Marion J. Lamb with illustrations by Anna Zeligowski A Bradford Book The MIT Press Cambridge, Massachusetts London, England © 2005 Massachusetts Institute of Technology All rights reserved. No part of this book may be reproduced in any form by any elec- tronic or mechanical means (including photocopying, recording, or information storage and retrieval) without permission in writing from the publisher. MIT Press books may be purchased at special quantity discounts for business or sales promotional use. For information, please email [email protected] or write to Special Sales Department, The MIT Press, 5 Cambridge Center, Cambridge, MA 02142. This book was set in Stone Sans and Stone Serif by SNP Best-set Typesetter Ltd., Hong Kong. Printed and bound in the United States of America. Library of Congress Cataloging-in-Publication Data Jablonka, Eva. Evolution in four dimensions : genetic, epigenetic, behavioral, and symbolic variation in the history of life / by Eva Jablonka and Marion J. Lamb with illustrations by Anna Zeligowski. p. cm.—(Life and mind) Includes bibliographical references (p. ). ISBN 0-262-10107-6 (alk. paper) 1. Evolution (Biology) I. Lamb, Marion J. II. Title. III. Series. QH366.2.J322 2005 576.8—dc22 2004058193 10 9 8 7 6 5 4 3 2 1 To our genetic, epigenetic, and cultural parents and offspring Contents Acknowledgments ix Prologue 1 I The First Dimension 5 1 The Transformations of Darwinism 9 2 From Genes to Characters 47 3 Genetic Variation: Blind, Directed, Interpretive? 79 II Three More Dimensions 109 4 The Epigenetic Inheritance Systems 113 5 The Behavioral Inheritance Systems 155 6 The Symbolic Inheritance System 193 Between the Acts: An Interim Summary 233 III Putting Humpty Dumpty Together Again 239 7 Interacting Dimensions—Genes and Epigenetic Systems 245 viii Contents 8 Genes and Behavior, Genes and Language 285 9 Lamarckism Evolving: The Evolution of the Educated Guess 319 10 A Last Dialogue 355 Notes 385 Bibliography 417 Index 447 Acknowledgments This book would not have been written without the encouragement and help of our friends, families, students and colleagues. We are grateful to all of them. Part of the book was written while E.J. was a visitor at the Museum of Vertebrate Zoology, Berkeley, University of California, and we would like to thank David and Marvalee Wake and their colleagues for the good working environment they provided,and Martha Breed and the WW group for their company and the wonderful nature trips. We also want to thank everyone working at the Cohn Institute for the History and Philosophy of Science and Ideas at Tel Aviv University for their help and support. Our debt to the students in the Cohn Institute and the participants in the “Networks in Evolution” seminar at the European Forum Alpbach 2002 is a big one. Their comments and criticisms made us clarify many of our ideas and arguments, abandon some of them, and think deeply about how we should present the material in this book. We hope that they will enjoy the final product. We have benefited from information and advice from many people, but our special thanks must go to those who have read and commented on various drafts of the book. Eytan Avital, Daniel Dor, Fanny Doljanski, Yehuda Elkana, Yehudit Elkana, Evelyn Fox Keller, James Griesemer, Revital Katznelson, Jawed Iqbal, Lia Nirgad, Christine Queitsch, Richard Strohman, Iddo Tavory, and Alan Templeton each read sections or chap- ters, and pointed out some of the errors and ambiguities in what we had written. Our long-suffering friends Lia Ettinger, Simona Ginsburg, and Joy Hoffman read drafts of the whole book, and their comments, criticism, and many valuable suggestions have made it a far better book than it would otherwise have been. Tom Stone and his colleagues at The MIT Press were helpful and encouraging throughout, and we thank them for their guid- ance and excellent editorial work.