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The Origins of Theoretical Population Genetics the Origins of Theoretical Population Genetics - Theoretical Population Genetics The Origins of Theoretical Population Genetics The Origins of Theoretical Population Genetics with a New Afterword William B. Provine The University of Chicago Press Chicago and London WituraM B. Provinzis the C. A. Alexander Professor of Biological Sciences at Cornell University. Heis the author of Sewall Wright and Evolutionary Biology and the editor of Evolution: Selected Papers by Sewall Wright, both published by the University of Chicago Press. The University of Chicago Press, Chicago 60637 The University of Chicago Press, Ltd., London © 1971 by The University of Chicago Afterword © 2001 by The University of Chicago All rights reserved. Printed in the United States of America IO 09 08 07 06 05 04 03 02 OI 12345 ISBN: 0-226-68463-6 (cloth) ISBN: 0-226-68464-4 (paper) Library of Congress Cataloging-in-Publication Data Provine, William B. Theorigins of theoretical population genetics / William B. Provine.—with a new afterword p- cm. Includes bibliographical references and index. ISBN 0-226-68463-6 (cloth : alk. paper)—ISBN 0-226- 68464-4 (pbk.: alk. paper) 1. Population genetics—History.I. Title. QH455.P77 2001 576.5'8'09—de21 2001027561 © 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. To Doris Marie Provine Contents INTRODUCTION 1x 1. Darwin’s THEory oF Natura SELECTION: THE ReE- ACTION DARWIN'S THEORY I THE REACTION 10 Thomas H. Huxley and “Natura non facit Sal- tum ad II Francis Galton, Regression, and Discontinuous Evolution 14 BACKGROUND TO THE CONFLICT BETWEEN MENDELIANS AND BIOMETRICIANS 25 KARL PEARSON: A SKETCH OF HIS EARLY LIFE 26 WELDON, PEARSON, AND BIOMETRY 29 WILLIAM BATESON AND DISCONTINUOUS EVOLUTION 35 THE PUBLIC CONTROVERSIES 45 The Cineraria Controversy 45 The Struggle over the Evolution Committee 48 THE CoNFLICT BETWEEN MENDELIANS AND BIOMETRI- CIANS 56 THE HOMOTYPOSIS CONTROVERSY 58 THE MUTATION THEORY 64 INHERITANCE IN PEAS 70 HEREDITY IN MICE 73 MENDELISM AND BIOMETRY 80 MEETING OF THE BRITISH ASSOCIATION, 1904 85 COAT COLOR IN HORSES 87 THE EFFECTS OF THE CONFLICT 8&9 DarwINIAN SELECTION: THE Controversy, 1900-1918 go THE ARGUMENT AGAINST DARWINIAN SELECTION QI Vi11 CONTENTS Wilhelm Johannsen’s Pure Line Theory g2 Criticism ofJohannsen’s Pure Line Theory 96 Herbert SpencerJennings and Pure Lines IOO Raymond Pearl and Pure Lines 104 Criticism ofthe Pure Line Theory 105 THE PROOF AND EXPLANATION OF THE EFFECTIVENESS OF SELECTION Ios William Ernest Castle and Selection Theory 109 The Multiple Factor Theory IT4 Thomas Hunt Morgan and Variationfor Evolution 120 Ocenothera Mutants 121 Pure Line Theory and Selection 122 Morgan's Theory ofEvolution 125 Castle and the Selection Problem 126 5. PopuLaTION GENETICS: [HE SYNTHESIS OF MENDELISM, DarwInIsM, AND BIOMETRY 130 EXPLORATION OF THE MATHEMATICAL CONSEQUENCES OF MENDELIAN HEREDITY BEFORE 1918 131 RONALD ALYMERFISHER 140 SEWALL WRIGHT ‘54 J. B. S. HALDANE 167 CONCLUSIONS 177 AppEnp1x: GALTON, PEARSON, AND THE LAw oF ANCESTRAL HEREDITY 179 BIBLIOGRAPHY 189 AFTERWORD 197 INDEX 207 Introduction Wehave cometogetherinto this hall from various distances, from various states and countries, to discuss the problems of our commoninterest concerning population genetics. It seems to me that the selection of the agenda for this Twentieth Symposium on Quantitative Biology has been most appropri- ate and timely, because the importance of population genetics, and its bearing on various other branches of biology, have now become recognized not only by investigators in these branches but also by men in the practical business of breeding and in manyotherfields of theory and application." WITH THESE WORDS THE CHAIRMAN OPENED THE COLD SPRING Harbor Symposium of 1955, that year devoted to population genetics. His statement is indicative of the wide recognition that population genetics has gained as an important field of biological research, with implications for other areas of biolos1- cal interest. Anthropologists, eugenicists, demographers, ecol- gists, breeders, and others have been muchinfluenced by the impinging ideas of population genetics. The term “twentieth-century Darwinism” has often been applied to modern population genetics. Although the term may be accurate as a description of the similarity between Darwin’s idea of evolution and that of most population geneticists regarding the role of natural selection in the origin of species, it is misleading because it suggests that population genetics developed from Darwin’s ideas. The development from Darwin’s ideas to population genetics was actually a tortuous one. The origin of population genetics is perhaps best under- stood as a product of the conflict between two views of evo- lution which were eventually synthesized. On one side was 1. Cold Spring Harbor Symposia on Quanttative Biology 20 (1955): v. 1x x INTRODUCTION Darwin’s belief in gradual evolution, produced by natural selection acting upon small continuous variations. On the other was Galton’s belief in discontinuous evolution, produced by natural selection acting upon large discontinuous varia- tions. Galton thought natural selection was ineffective acting upon the small variations Darwin envisioned. The conflict between these two views began with the publication of the first edition of The Origin of Species and did not end until population genetics provided a new,synthetic theory. In this account, I treat the historical development of the ideas which culminated in the laying of the theoretical foun- dations of population genetics. The theoretical foundations, sometimes termed “classical” population genetics, were laid between 1918 and 1932 by R. A. Fisher, J. B. S. Haldane, and Sewall Wright. Their work was stimulated in large part by the controversy over the continuity of evolution and the efficacy of naturalselection. I have not included the contributions of the Russian School —Chetverikov, Timofeev-Resovsky, Dubinin, and later Dobz- hansky—* to the study of natural populations because their work did not merge with “classical” population genetics until after the theoretical foundations were established. Chetverikov did publish in 1927 a very important paper in theoretical pop- ulation genetics, but by the time this paper was known in England and the United States, the theoretical construct erected by Fisher, Haldane, and Wright had progressed be- yondit. I am greatly indebted to Professors Allen G. Debus and Richard C. Lewontin of the University of Chicago for their careful criticisms and suggestions while I was undertaking this project. I owe special thanks to Professor Lewontin for taking many hours of his time to encourage my work in biology while I was a graduate studentin history. Doris Marie 2. For an account of the Russian School, see Mark B. Adams, “The Founding of Population Genetics: Contributions of the Chetverikov School, 1924-1934,” Journal of the History of Biology 1 (1968): 23—40. Introduction xt Provine carefully analyzed each chapter, removing deadwood and demandingclarification. I have incorporated so many of her suggestions that she is responsible for a substantial portion of whatever merit this work achieves. Sewall Wright kindly granted me two lengthy interviews which helped my understanding of his work. I also wish to thank Professor William Coleman of Johns Hopkins Uni- versity and Mr. Murphy Smith of the American Philosophical Society for their help in guiding me to and throughthe Bate- son papers. Finally, I am grateful to Mrs. Lina Hood and Mrs. Bernice White for typing services rendered. ] Darwin’s Theoryof Natural Selection: The Reaction Darwin’s THEORY When Charles Darwin boarded the Beagle in late 1831 for his famous voyage he took with him volumeone of Charles Lyell’s Principles of Geology, which had been published in 1830. Darwin’s teacher John Henslow had recommended the book to him with the admonition notto accept Lyell’s views. Like most geologists of the time, Henslow was a catastrophist. He believed the geological history of the earth was progres- sive, that is, showed significant changes, and was characterized by successive cataclysms with periods of little change in be- tween. Adam Sedgwick, president of the Geological Society in 1831 and Darwin’s other teacher of geology, was also a catastrophist. With two catastrophists as his teachers Darwin naturally adopted their general view of geological change. But he did not think catastrophism provided a complete ex- planation of geological change. Hesaid to friend,“it strikes me that all our knowledge about the structure of the earth is very much like what an old hen would know of a hundred- acre field, in a corner of whichsheis scratching.”* In the Principles Lyell challenged, as James Hutton had before him, the prevalent geological theories of catastrophism and progressionism. Lyell believed the geological history of the earth could be explained by the same agents that were operating at present, given enough time. This was the prin- ciple of uniformitarianism. He went further and rejected progressionism. He held that geological forces had made only minor changes in the earth’s surface, and not even a cumula- tion of geological events could cause a major change. The impact of volume one of Lyell’s Principles upon Dar- 1. Francis Darwin, ed., The Life and Letters of Charles Darwin, 3 vols. (London: John Murray, 1887), 2:348. 2 NATURAL SELECTION: THE REACTION win’s thought was quick and deep. Only twenty days after the start of the Beagle’s voyage, the first ten of which he was miserably seasick, Darwin landed at St. Iago in the Cape Verde Islands and was there convinced, as he later said, of “the infinite superiority of Lyell’s views over those advocated in any other work known to me.”* This was a remarkable transformationin such a short period of time. St. Iago was a perfect place to convince Darwin of Lyell’s belief in gradual geological change. At first glance, the island might have appeared to be a perfect example of catastrophism.
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