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Francis Galton Pioneer of Heredity and Biometry Michael Bulmer

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Francis Galton Pioneer of Heredity and Biometry Michael Bulmer Francis Galton Pioneer of Heredity and Biometry Michael Bulmer 2003 THE JOHNS HOPKINS UNIVERSITY PRESS Baltimore and London Contents Acknowledgments xi Chronology xiii Introduction xv 1A Victorian Life 1 Family Background and Education 1 Travels 6 Eastern Europe, 1840 6 The Near East, 1845–46 7 South West Africa, 1850–52 11 Vacation Tours 18 Scientific Career 21 The Royal Geographical Society 22 Exploration in Central Africa 23 The British Association 27 Inventions 28 Meteorology 30 Heredity and Evolution 32 Psychology 32 Photography 34 Fingerprints 35 Characterization 36 vi Contents 2 Hereditary Ability 42 “Hereditary Talent and Character” (1865) 44 Hereditary Genius (1869) 46 English Judges 48 Comparison of Results for All Professions 50 Transmission through Male and Female Lines 54 The Reception of Hereditary Genius 57 Nature and Nurture 60 English Men of Science: Their Nature and Nurture (1874) 60 “The History of Twins” (1875) 64 Galton’s Hereditarianism 67 Epilogue 71 Appendix: Number of Kinsfolk 74 3 Eugenics 79 Galtonian Eugenics 79 Later History of Eugenics 84 Britain 84 America 87 Germany 92 The Rationale of Eugenics 98 4The Mechanism of Heredity 102 Galton’s Knowledge of Heredity in 1865 103 Biparental Inheritance 103 The Non-Inheritance of Acquired Characters 105 The Law of Reversion 107 Darwin’s Provisional Hypothesis of Pangenesis 108 Reversion 110 The Inheritance of Acquired Characters 112 Xenia and Telegony 113 Galton’s Reaction to Pangenesis 114 Galton’s Political Metaphor of Pangenesis 114 An Experimental Test of Pangenesis 116 Contents vii Galton’s Theory of Heredity in the 1870s 119 Similarities between Relatives 123 Galton’s Ideas on Heredity in 1889 127 Discussion 131 Weismann and the Continuity of the Germ-Plasm 132 De Vries’s Theory of Intracellular Pangenesis 133 Segregation 136 Blending Inheritance 138 Fleeming Jenkin and the Problem of Swamping 141 5 Four Evolutionary Problems 147 The Domestication of Animals 147 The Evolution of Gregariousness 150 The Fertility of Heiresses 153 The Extinction of Surnames 156 The Evolution of Sex 160 “A Theory of Heredity” (1875) 161 Three Unpublished Essays 163 6 The Charms of Statistics 168 Quetelet and the Average Man 169 Galton and the Normal Distribution 173 Hereditary Genius (1869) 173 Natural Inheritance (1889) 175 The Importance of the Normal Distribution to Galton 180 Galton’s Quincunx 182 Regression and the Bivariate Normal Distribution 184 Correlation 191 Two Concepts of Probability 196 The Development of Statistics 202 Appendix: Regression Theory 206 7 Statistical Theory of Heredity 209 A Theory Based on Pangenesis 210 viii Contents “Typical Laws of Heredity” (1877) 211 An Experiment with Sweet Peas 212 Solution of the Problem 215 Johannsen’s Experiments with Beans 218 The Inheritance of Human Height 224 The Advantages of Height 225 The Regression of Offspring on Mid-Parent 229 Kinship 231 Fraternal Regression 233 Variability in Fraternities and Co-Fraternities 235 8 The Law of Ancestral Heredity 238 Galton’s Formulation of the Ancestral Law 239 Galton’s Derivation of the Law in 1885 241 Derivation of the Law in 1897 244 Galton’s Law As It Should Have Been 247 Karl Pearson’s Interpretation of the Ancestral Law 250 The Ancestral Law and Mendelism 257 Weldon and Mendelism 259 Pearson and Mendelism 261 Yule’s Reconciliation of the Law with Mendelism 266 Appendix: The Regression on Mid-Ancestral Values 272 9 Discontinuity in Evolution 275 Galton’s Theory of Discontinuous Evolution 276 Stability of Type 277 Perpetual Regression 281 Selection Experiments 284 The Fallacy of Perpetual Regression 285 “Discontinuity in Evolution” (1894) 288 Speciation and Saltation 292 De Vries and The Mutation Theory 294 Punctuated Equilibria 297 Contents ix 10 Biometry 299 The Demonstration of Natural Selection 300 The Career of W. F. R. Weldon 301 The Common Shrimp 302 The Shore Crab 303 Stabilizing Selection in Snails 308 Bumpus’s Sparrows 309 Multivariate Selection 312 Quantitative Genetics 315 The Multiple Factor Hypothesis 316 The Hardy-Weinberg Law 318 Mendelian Theory of Quantitative Genetics 321 The Response to Selection 324 Coda 327 Appendix: Multivariate Selection Theory 329 Selection Differentials and Selection Gradients 329 The Response to Selection 331 References 333 Index 351 Acknowledgments I am grateful to University College London Library for access to the Galton and Pearson papers, and for permission to use extracts from them; to the Biometrika Trust and to Oxford University Press for permis- sion to quote extracts and to reproduce three figures from The Life, Letters and Labours of Francis Galton by Karl Pearson; to the librarians of the Radcliffe Science Library and the Bodleian Library at Oxford University for their help in locating references; to Professor Anthony Edwards of Gonville and Caius College, Cambridge and to an anonymous referee for valuable comments on the manuscript; to David Burbridge for helpful correspondence about Galton’s theory of heredity; to Gavan Tredoux for use of his excellent website www.galton.org; to my copy editor Alice Calaprice of Princeton, New Jersey; to Trevor Lipscombe, Julie McCarthy, and Martha Sewall at the Johns Hopkins University Press, for their help in improving the manuscript and in preparing it for publica- tion; and to Mark Boardman for making the scanned images. Chronology 1822 Francis Galton born, 16 February 1828–38 At school 1838–40 Medical education in Birmingham and London 1840 Journey to Eastern Europe 1840–44 Trinity College, Cambridge, reading mathematics 1844 His father died, and he abandoned medicine 1845–46 Journey to Near East 1846–49 Hunting and shooting in England and Scotland 1850–52 Journey to South West Africa 1853 Married Louisa Butler Tropical South Africa 1855 The Art of Travel 1857 Bought 42 Rutland Gate, London 1864 Burton-Speke dispute at British Association, Bath 1869 Hereditary Genius 1872 Stanley’s talk at British Association, Brighton 1874 English Men of Science: Their Nature and Nurture 1883 Inquiries into Human Faculty and Its Development 1889 Natural Inheritance 1892 Finger Prints 1897 Louisa Galton died 1908 Memories of My Life 1909 Essays in Eugenics 1911 Francis Galton died, 17 January xiv Chronology Honors and awards 1853 Gold Medal, Royal Geographical Society 1855 Member of Athenaeum Club by special election 1860 Fellow, Royal Society 1886 Royal Medal, Royal Society 1894 Hon. D.C.L., Oxford University 1895 Hon. D.Sc., Cambridge University 1901 Huxley Medal, Anthropological Institute 1902 Hon. Fellow, Trinity College, Cambridge Darwin Medal, Royal Society 1908 Darwin-Wallace Medal, Linnean Society 1909 Knighted 1910 Copley Medal, Royal Society Introduction Francis Galton made important contributions in many areas of science in the nineteenth century. He explored South West Africa, discovered and named the anticyclone, and wrote a book on fingerprints, in addition to his work on anthropometry, psychology, and photography. But most important were his pioneering studies of heredity, in the course of which he invented the statistical tools of regression and correlation, earning him the title of the father of biometry, the application of statistical meth- ods to evolutionary biology. Galton’s work on heredity was closely linked to the evolutionary the- ory of his half-cousin Charles Darwin. Darwin proposed that species evolve through descent with modification, and that modification is pre- dominantly due to the operation of natural selection on small inherited variations. The first part of Darwin’s thesis was quickly accepted by con- temporary scientists, but his theory of the mechanism of evolution by natural selection was not, because he did not have a convincing theory of heredity to explain how selection acted and because he had no direct evidence of natural selection. Thus the explanation of variability and heredity and their relationship to natural selection have been key ques- tions in the history of Darwinism from the publication of The Origin of Species in 1859 until today. The work of Galton and his followers played a central role in the early history of Darwinism, which has been increas- ingly recognized by historians of science (Provine 1971, Olby 1985, Gayon 1998). Their work in the application of statistical methods to the biological sciences was equally important (Porter 1986, Stigler 1986 and 1999). This book is an attempt to clarify and explain the development of Galton’s ideas on heredity and biometry, and to place them in their scien- tific context. xvi Introduction Ernst Mayr divides scientists into romantics and classics: “The roman- tic is bubbling over with ideas that have to be dealt with quickly to make room for the next one. Some of these ideas are superbly innovative; others are invalid if not silly.... The classic, by contrast, concentrates on the perfection of something that already exists. He tends to work over a subject exhaustively. He also tends to defend the status quo” (1982, 831). Galton was an archetypal romantic, an innovator with the gift of seeing problems in statistical terms, but lacking the mathematical ability and the inclination to push his ideas to their logical conclusion. Relying on intuition rather than mathematical expertise, he made mistakes and was sometimes confused or self-contradictory. He was also rather careless arithmetically; obvious numerical errors have been corrected in what follows without comment. Thus, Galton was a pioneer who left it to others to perfect his ideas. It has been a challenge to present a clear account of these sometimes unclear ideas without distorting them. Apart from the first, biographical chapter, this is an internalist account of the development of scientific ideas, based on the premise that scientific knowledge is not just a subjec- tive social construct (Ruse 1999), though scientists are fallible men and women who are constrained by their peculiar abilities, personalities, and social environment. Social constructionist historians, coming from a background in the social sciences, tend to stress the social context of science history and have little sympathy with “the old internalists who wrenched science from its social context and wrote ghostly histories of disembodied ideas” (Desmond 1989, 21).
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