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LIVES OF A BIOLOGIST LIVES OF A BIOLOGIST A D V E N T U R E S I N A C E N T U R Y O F E X T R A O R D I N A R Y S C I E N C E JOHN TYLER BONNER HARVARD UNIVERSITY PRESS Cambridge, Massachusetts London, England 2002 Copyright © 2002 by the President and Fellows of Harvard College All rights reserved Printed in the United States of America Pages 18–20 were originally published as the foreword to On Growth and Form: Spatio-Temporal Pattern Formation in Biology, ed. M. A. J. Chaplain, G. D. Singh, and J. C. McLachlan (New York: John Wiley & Sons, 1999); reproduced with permission. Library of Congress Cataloging-in-Publication Data Bonner, John Tyler. Lives of a biologist : adventures in a century of extraordinary science / John Tyler Bonner. p. cm. ISBN 0-674-00763-8 (alk. paper) 1. Bonner, John Tyler. 2. Biologists—United States—Biography. 3. Biology. I. Title. QH31.B715 A3 2002 570'.92—dc21 2001051526 [B] CONTENTS Preface vii 1. The World of My Elders: 1900–1920 1 2. Becoming a Biologist: 1920–1940 23 3. Everything Peaks: 1940–1960 61 4. Revolution and Progress: 1960–1980 133 5. Coming Together: 1980–2000 169 Index 211 PREFACE My lovely old office in Guyot Hall in Princeton— which I have occupied for over fifty years—was built in 1910. It has thick oak tables, brown wood window frames, and four lights that hang from the high ceil- ing, each with a conical green glass shade. The only thing I have added are bookcases on three walls. It definitely has a beginning-of-the-century atmosphere. At this moment I am sitting in my study in our summer place in Cape Breton. This nineteenth-century house looks out onto the estuary of a beautiful river. The walls of the room are made of horizontal boards of uneven width that have been smoothed with a draw knife. As I look out of the window by my desk, I can see the occasional small boat on the water and patches of farmland on the distant hills. Right this minute all is enveloped in a morning mist. These two rooms, in some sort of symbolic way, have framed me for the century. Within that frame there have been enormous changes. Here I am with my laptop computer writing this, and periodically entering the Internet for refer- ences or for reading the latest in the New York Times. Biol- ogy, the world, and I have gone through many transmutations. I was born in 1920 and started my interest in biology in the early 1930s, which means that I have spanned a large portion of the last century—a century that, above all things, has been distinguished by an extraordinary series of advances in biology. If one compares the biology of 1900 with that of 2000, the change has been astounding. In the beginning there was the discovery of the genes on the chromosomes, followed vii by the discovery of embryonic induction; the interpretation of evolution in terms of changes in gene frequency in a popu- lation; the rise of an understanding of the biochemistry of the cell; and perhaps most important of all, the beginning of molecular genetics, which was followed by the molecular genetics of embryological development; the great, new insights into animal behavior; sociobiology; the simplifica- tion of ecological and evolutionary principles by means of mathematical models—and so many other discoveries. To one glancing at this history from a panoramic point of view, a number of things are evident. One is the pursuit of the gene and the extraordinary success of the totally reduc- tionist approach, where the molecules and their structure have been unmasked. Another is the application of that reductionist knowledge to the problems of evolution and developmental biology. A third is the idealization, the sim- plification, of the vast number of complexities that charac- terize practically all of what we now know in modern biology: for instance, the regulation of the molecular activi- ties within the cell, the complex structure of ecological communities, the functioning of the brain, the sequential steps of development, the patterns of animal behavior, and the intricacies of evolutionary change. I ask myself where I fit in with all this. Over the years I have tried to keep my eye on as big a picture as possible, and to see how it all comes together. Development is not divorced from evolution nor, for that matter, from behavior; genetics is intimately intertwined with every aspect of modern biology from the activities of the cell to those of the brain; the group- ing of organisms—either as social groups or ecosystems—are viii P R E F A C E intimately connected with considerations of development and evolution. One of the things that has happened during this hundred- year span is the enormous accumulation of new and fasci- nating facts—so much so that it is hard to see even the trees, let alone the forest; one could almost say that we are now focused on spots on the bark. A primeval urge inside me pushes me toward bucking this trend. However, I have not done it by simply dissolving into generalities, but have steered a quite specific path, a specific vision. As will be evident in the pages to come, that vision has been to consider all organisms as life cycles, an enormously unifying way of thinking of things for it automatically links development, evolution, behavior, genetics, physiology, ecology, even behavior; it is a thread that links all of biology. The concept of life cycles also lies at the foundation of my own evolution. I began with an interest in lower organisms, algae and fungi and related forms that confronted me with their trajec- tory from egg to adult, or spore to adult, in a way that could not be dismissed. Furthermore, my initial laboratory focus was on the development of the social amoebae (cellular slime molds) whose life cycle, as we shall see later, is so peculiar that it is impossible to avoid becoming conscious of life cycles. They have been for me the lens for all my work, both in the laboratory and in my writing: the life cycle is a great simplifying and unifying part of all living organisms. In particular it is the bond that connects development and evo- lution, subjects so central to all of biology and ones that have been receiving increasing current interest. And then I am conscious of my own life cycle. Here I am P R E F A C E ix

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