DARWIN'S BIOLOGICAL WORK
Solanum rostratuni Dun. Two flowers of the racemose inflorescence showing the asymmetry of the reproductive parts. The style eurves downwards and always lies between the lower enlarged stamen and the main axis of the infloreseenee so that adjacent flowers of the inflorescence arc mirror images of each other. See note on p. ix,
DARWIN'S BIOLOGleAL WORK SOME ASPECTS RECONSIDERED BY P. R. BELL J. CHALLINOR J. B. S. HALDANE P. MARLER H. L. K. WHITEHOUSE J. S. WILKIE EDITED BY P. R. BELL CAMBRIDGE AT THE UNIVERSITY PRESS 1959
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CONTENTS List ofplates Preface page ix xi 1 THE MOVEMENT OF PLANTS IN RESPONSE TO LIGHT, by P. R. BELL 1 The Recognition of Light as a Cause of Plant Movement 2 The Originality of Darwin's Work 7 The Development of the Study of Phototropism since Darwin 12 The Chemical Nature of the Growth-promoting Substances 29 The Part Played by Auxin in Phototropism 34 Conclusion 47 2 PALAEONTOLOGY AND EVOLUTION by J. CHALLINOR 50 The General Showing of Palaeontology 50 The Imperfection of the Geological Record of the History of Life 54 The Kinds of Evidence in Evolutional Palaeontology 60 Stratigraphical Palaeontology 67 Evolution in the Groups 78 Evolutionary Series 89 Appendix: The Geological Table 100 8 NATURAL SELECTION, by J. B. S. HALDANE 101 A Sketch of Darwin's Opinions 101 A Sketch of Genetics 104 Definition and Measurement of Selection: Effects of Genotypic Selection III Centripetal Selection 117 v
Directed Selection Sexual Selection Contents page 124 181 Speciation by Allopolyploidy 182 The Rate of Evolution, and the Argument from Artificial to Natural Selection 184 Evolution and Geography 141 Instinct 148 Genetic Assimilation 145 Other Difficulties 147 4 DEVELOPMENTS IN THE STUDY OF ANIMAL COMMUNICATION, by P. MARLER 150 Communication by Taste and Smell 151 Visual Communication 157 Auditory Communication 172 The Evolution of Bird Song 181 The Origins of Communication 197 Animal and Human Language 202 5 CROSS- AND SELF-FERTILIZATION IN PLANTS, by H. L. K. WHITEHOUSE 207 Introduction 207 The Development of Ideas on the Cause of Inherited Variation 215 The Development of Ideas on Why it is Beneficial to the Progeny to Cross Individuals which Differ 226 Developments in Knowledge of Mechanisms for Favouring Cross-fertilization 284 The Development of Ideas on How the Various Mechanisms for Achieving Cross-fertilization have Evolved 251 The Outlook 258 vi
Contents 6 BUFFON, LAMARCK AND DARWIN: THE ORIGINALITY OF DARWIN'S THEORY OF EVOLUTION, by J. S. WILKIE page 262 Evolution and Theology in the Eighteenth Century 266 The Theory of Evolution in the Works of Buffon 270 Buffon and Lamarck 288 Lamarck's Theory of Evolution Conclusion 289 802 Notes and References 811 vii
Solanum rostratum Dun LIST OF PLATES frontispiece A letter written by Darwin nine days before his death, probably his last scientific inquiry, was a request to Professor J. E. Todd of Iowa for more information and seeds of Solanum rostratum, a Mexican plant becoming naturalized in Texas. Professor Todd had suggested that the two forms of the flower of this plant provided a mechanism for cross-fertilization and as soon as this came to Darwin's notice he was anxious to experiment with them. 'But', he added, 'if you intend to experiment on them, ofcourse you will not send me the seeds, as I shd, be very unwilling to interfere in any way with your work.' In fact, no further attention seems to have been given to this curious plant until the present day. I A Jurassic (Corallian) ammonite facing page 80 II RistonbetuZaria L.-normal and melanic (f. carbonaria) forms 128 III Ancient and modern forms of heterozygotes of B. betu- Zaria f. carbonaria 129 IV Oscillograms from magnetic tape recordings of Ceylon cicadas 176 V Heterostyled plants, dioecism, gynodioecism, cleistogamy 240 ix
PREFACE On 24 May 1859, reviewing the papers which had been read before the Linnean Society of London in the preceding session, the President, Thomas Bell, is recorded in the history of the Societyas having regretted that the session had passed without being marked' by any of those striking discoveries which at once revolutionize, so to speak, the department of science on which they bear', or which 'shall produce a marked and permanent impress on the character of any branch of knowledge'. Nevertheless, one contribution was certainly 'to produce a marked and permanent impress', for, on 1 July 1858, the communications of Charles Darwin and Alfred Wallace On the Tendency ofspecies to form Varieties had been read to the Society. Few judgments can have proved so erroneous as that of Thomas Bell. The story of the publication in the following year of The Origin ofspecies and of the controversy which ensued has often been told. It is less well known that feeling was intense even within the Linnean Society and several Fellows withdrew from it because Darwin was not summarily expelled. Yet, within ten years, the Council of the Linnean Society was more rigorous in its interpretation of 'Darwinism' than Darwin himself. This humble man, who, in diffidently sending specimen chapters of The Origin ofspecies to his publisher, requested that after perusal they be sent to a 'lady being excellent judge of style [who] is going to look out for errors for me', was indeed beginning to effect a revolution. Yet we should not think of Darwin solely in relation to evolution and natural selection. He was in fact one of the foremost geologists of his day and his early papers made noteworthy contributions to that science. His work came to encompass the whole field of biology, but was never superficial. Several of his publications, for example The 17arious Contrivances by which Orchids are fertilized by Insects, have not yet been superseded. It is true, of course, that evolution came to be a unifying principle in all Darwin's scientific work, but such unifying principles are a great stimulus to biological progress. xi
Preface The contributions of Darwin to experimental biology have received less attention than his descriptive and theoretical work, but theywere byno meansunimportant. Forexample, he recognized the importance of plant physiology and pressed for the establishment of laboratories for its study. Starved though he was of equipment, his penetrating observations have been the origin of some of the most active branches of contemporary biology. For a century and more biologists have drawn freely and profitably upon Darwin's work and his flashes of originality continue to stimulate his successors. It is for these reasons, not because of the endless and often ignorant controversies which have become popularly associated with his name, that Darwin occupies a unique place in the history of biology. It is to put this point of view that Darwin's Biological Work is being published at this time when the name of Darwin is again before the public. All of the contributors are actively engaged in biological research (and research into the development of ideas in biology is considered to be an essential part of the science). Our hopes are the same as Darwin's when he submitted the manuscript of The Origin ofspecies: 'the book ought to be popular with a large bag of scientific and semi-scientific readers '. xii
ACKNOWLEDGEMENTS Acknowledgements are gratefully made to the following: to Mr G. Atkinson, for the frontispiece; to the Controller, H.M. Stationery Office, for figs. 2 and 3 in the article by J. Challinor (fig. 12 from British Regional Geology, The Welsh Borderland, and fig. 25 from The Hampshire Basin and Adjoining Areas, respectively), and to Professor T. N. George and Messrs C. A. Watts and Company Limited for fig. 4 in the same article; to the late Dr W. J. Arkell, F.R.S., and the Council of the Palaeontographical Society for PI. I; to Dr H. B. D. Kettlewell and Mr Michael Lyster, Department of Zoology, University Museum, Oxford, for PI. II, and to Dr H. B. D. Kettlewell and Mr John Haywood for PI. III, both plates illustrating the work of Dr Kettlewell discussed by J. B. S. Haldane; to Dr J. W. S. Pringle, F.R.S., for PI. IV, taken from his article in the Journal ofexperimental Biology; and to Mrs M. P. Whitehouse, A.R.P.S., for PI. V. The Editor would personally like to thank Mr John G. Murray for allowing him to see unpublished letters of Charles Darwin, the Cambridge University Press for the care they have devoted to this volume, and in particular Elizabeth Bell for undertaking much of the preparation of the manuscript and other invaluable assistance. xiii
NOTE Superior figures in the text refer to the Notes and References at the end ofthe book (p. 311).