THE COMMONWEALTH AND INTERNATIONAL LIBRARY Joint Chairmen of the Honorary Editorial Advisory Board SIR ROBERT ROBINSON, O.M., F.R.S., LONDON DEAN ATHELSTAN SPILHAUS, MINNESOTA Publisher: ROBERT MAXWELL, M.C3 M.P. ZOOLOGY DIVISION General Editors: j. M. DODD AND SIR FRANCIS KNOWLES THE FOUNDATIONS OF GENETICS The forecourt of the Augustinian monastery of St. Thomas. On the left, the white marble statue of Mendel by the Viennese sculptor Charlemont, a tribute from "the friends of science to the investigator Gregor Mendel". It was unveiled in 1910 when it stood in the Klosterplatz outside the monastery wall. On the right, the sandstone monument erected by the monastery in 1922 on the occasion of the centenary celebrations of Mendel's birth. It stands in the garden plot (120 ft X 20 ft) that was used by Mendel in his pea experiments. The windows on the first floor above the monument and facing the street are those of the apartment occupied by Mendel during these years. On the right of this plot, on the ground-floor, is the room that was the refectory of the monastery and that now is the Mendel Memorial Hall. THE FOUNDATIONS OF GENETICS by F. A. E. CREW, F.R.S. Formerly Professor of Animal Genetics in the University of Edinburgh PERGAMON PRESS OXFORD - LONDON · EDINBURGH · NEW YORK TORONTO · PARIS · FRANKFURT Pergamon Press Ltd., Headington Hill Hall, Oxford 4 & 5 Fitzroy Square, London W.l. Pergamon Press (Scotland) Ltd., 2 & 3 Teviot Place, Edinburgh 1 Pergamon Press Inc., 44-01 21st Street, Long Island City, New York 11101 Pergamon of Canada, Ltd., Adelaide Street East, Toronto, Ontario Pergamon Press S.A.R.L., 24 rue des Écoles, Paris 5e Pergamon Press GmbH, Kaiserstrasse 75, Frankfurt-am-Main Copyright © 1966 Pergamon Press Ltd. First edition 1966 Library of Congress Catalog Card No. 65-27371 Printed in Great Britain by Billing & Sons Limited^ Guildford and London This book is sold subject to the condition that it shall not, by way of trade, be lent, re-sold, hired out, or otherwise disposed of without the publisher's consent, in any form of binding or cover other than that in which it is published. (2312/66) A great man lays upon posterity the duty of understanding him JOHN BUCHAN, Oliver Cromwell LIST OF ILLUSTRATIONS PLATES The forecourt of the Augustinian monastery of St. Thomas Frontispiece I. The Augustinian House of St. Thomas, Brno Facing p. 18 II. J. G. Kölreuter, Carl F. von Gärtner, Charles Naudin, August Weismann III. Francis Galton, E. B. Wilson, Pierre Louis François Levegue de Vilmorin, Johann Gregor Mendel IV. Carl Correns, Hugo de Vries, Erich von 50 Tschermak-Seysenegg, Th. Boveri V. William Bateson, R. C. Punnett, W. Johanssen, W. E. Castle VI. Charles B. Davenport, Thomas Hunt Morgan, Calvin B. Bridges, H. J. Muller VII. Giant chromosomes of the salivary glands of D. melanogaster 98 FIGURES PAGE 1. A dioecious plant 6 2. A monoecious plant 7 3. Fertilization in a flowering plant 8 4. The unfixable hétérozygote: the Blue Andalusian fowl 33 5. Mitosis 58 6. Factor interaction: Peacomb XRosecomb in the fowl 77 7a. Sex-linkage in Abraxas 81 ix X LIST OF ILLUSTRATIONS 7b. Sex-linkage in Abraxas: the reciprocal cross 82 8. A population compounded out of five pure lines of beans 91 9. The chromosome complex of the normal human male ; the karyotype of the normal human male 93 10. Meiosis 95 11. Pollen-grain formation 96 12. Embryo-sac formation 97 13. Cross-section of a grain of maize 98 14. Drosophila melanogaster: male and female 99 15. Conventional diagram of the chromosome complex of Drosophila melanogaster 100 16. Sex-linkage in Drosophila melanogaster 104 17. Sex-linkage in Drosophila melanogaster: the reciprocal cross 105 18. Sex-linkage in Abraxas: the explanation 107 19. Primary non-disjunction 116 20. Secondary non-disjunction 117 21. Bar-eye character in Drosophila melanogaster 118 22. Map of the chromosomes of Drosophila melanogaster 120 23. The giant chromosomes of the salivary gland ofDrosophila melanogaster: diagrammatic 151 PREFACE GENETICS has become the core science in the biological field; its basic tenets have profoundly influenced the rate and the direction of the development of all the biological and sociological sciences and have found their applications in many different spheres of scientific activity. The major interest in genetics has been passing from the bio logical to the molecular level where the basic nature of the living organism is being studied in almost incredible detail by means of the techniques and experimental methods of physics, chemistry and mathematics. For an understanding of the newer molecular genetics an adequate knowledge of what is now called classical genetics is essential. This book attempts to trace the historical development of genetics, throwing into prominence those contributions to advancing genetical knowledge which can surely claim an exceptional and enduring importance.* It also draws attention to the many and varied spheres of scientific activity in which genetics, in application, has been found to have its fruitful uses. F. A. E. C. * These are the scientific papers cited in the footnotes. XI ACKNOWLEDGEMENTS As THE approach in the writing of this book has been the historical, it follows that I have drawn very largely upon the writing of others. I hope that the inclusion of any book or scientific paper in the list of references at the end of this book will be regarded as an acknow ledgement of my indebtedness and as a gesture of gratitude to both author and publisher. I am very grateful to the late Dr. J. Krizenecky, of the section of the Moravian Museum in Brno which is concerned with the collec tion and conservation of Mendeliana, for the photographs of Gregor Mendel, of his statue and of the monastery in Brno; to Dr. B. M. Slizynski of the Institute of Animal Genetics, Edinburgh University, for the photograph of the giant chromosomes of Drosophila mela- nogaster; to Dr. Marcus Rhoades of the University of Indiana for the photographs of some of the founders of genetics which appeared as frontispieces in the journal Genetics and for his permission to re produce them; to Professor R. C. Punnett and to Professor H. J. Müller of Indiana University for photographs of themselves and to Professor Edward Castle of Harvard University for one of his father, for inclusion in the company of the founders and to Messrs. Oliver & Boyd of Edinburgh for permission to reproduce a number of illustrations in a book written by me and published by them many years ago. F. A. E. C. xiii CHAPTER 1 PLANT HYBRIDIZATION BEFORE MENDEL WHEN, long ago in man's eventful history, the hunter of wild animals and the gatherer of wild plants gave place to the herdsman and to the scatterer of grain, his dependence upon domesticated animals and cultivated plants began. It was inevitable that sooner or later he would seek ways and means of "improving" these animals and plants so that they might more fully satisfy his expanding needs. The accumulated experience of countless generations of husband men gradually became fashioned into a traditional craft and out of the breeders' methods of trial and error the art of breeding developed, to be used in the creation of new breeds and varieties attuned to the purposes for which they were being deliberately bred and to the conditions of the environment in which they were to be raised. When man had gained the mastery over his physical environment and the struggle for existence had become less fierce, the breeder ceased to look upon his animals and plants merely as objects of use and began to look to them for aesthetic satisfaction. He became a "fancier" who exercised the art of breeding in the production of animals and plants remarkable for their beauty or for their quaint- ness. The great variety of animals and plants of economic importance and their high quality and the great diversity of types of exhibition beasts and birds and flowers and fruits that are to be encountered in the agricultural, poultry, bird and flower shows that figure so large in the life of most countries today bear witness to the astonishing skill that the breeder and the fancier wield. These are the creations of the artist, not of the scientist, for no satisfactory explanation of the successes that these men achieved and of the 1