The Third Base

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Appendix

The Third Base

Donald Forsdyke

If I thought that by learning more and more I should ever arrive at the knowledge of absolute truth, I would leave off studying. But I believe I am pretty safe. Samuel Butler, Notebooks

Darwin’s mentor, the geologist Charles Lyell, and Darwin himself, both considered the relationship between the evolution of biological species and the evolution of languages [1]. But neither took the subject to the deep informa- tional level of Butler and Hering. In the twentieth century the emergence of a new science – Evolutionary Bioinformatics (EB) – was heralded by two discoveries. First, that DNA – a linear polymer of four base units – was the chromosomal component conveying hereditary information. Second, that much of this information was “a phenomenon of arrangement” – determined by the sequence of the four bases. We conclude with a brief sketch of the new work as it relates to William Bateson’s evolutionary ideas. However, imbued with true Batesonian caution (“I will believe when I must”), it is relegated to an Appendix to indicate its provisional nature. Modern languages have similarities that indicate branching evolution from common ancestral languages [2]. We recognize early variations within a language as dialects or accents. When accents are incompatible, communication is impaired. As accents get more disparate, mutual comprehension de- creases and at some point, when comprehension is largely lost, we declare that there are two languages where there was initially one. The origin of language begins with differences in accent. If we understand how differences in accent arise, then we may come to understand something fundamental about the origin of language (and hence of a text written in that language). It was for this reason that an abstraction was introduced in Chapter 5 – the notion that hereditary information could include an “accent” that might be the key to understanding the origin of species. A distinction was made between a message itself (primary information) and the accent with which it was spoken (secondary information). In other words, multiple forms of information (e.g. message and accent) can share a common text and we can consider them in hierarchical fashion (primary, secondary, etc.). 668 Appendix

There is little new in this. In 1978 in Mind and Nature Gregory Bateson referred to the hierarchical levels of hereditary information postulated by the anti-Darwinian “typologists,” which contrasted with the “synthetic” views of those favoring the “modern synthesis” of Darwinian and Mendelian ideas [3]: It is interesting to note the current controversy between the upholders of ‘synthetic’ theory in evolution (the current Darwinian orthodoxy) and their enemies, the ‘typologists.’ Ernst Mayr, for example, makes fun of the blindness of the typologists: ‘History shows that the typologist does not and cannot have any appreciation of natural selection’ [[4]]. ... Do any of the genetic messages and static signs that determine the phenotype have the sort of syntax ... which would divide the ‘typological’ from ‘synthetic’ thinking? Can we recognize, among the very messages which create and shape the animal form, some messages more typological and some more synthetic? Here Gregory Bateson was thinking of two physical classes of message each conveying a distinct class of information, rather than of one physical class of message capable of conveying two distinct classes of information. He acknowledged possible conflicts between different classes (layers) of information: “Every evolutionary step is an addition of information to an al- ready existing system. Because this is so, the combinations, harmonies, and discords between successive pieces and layers of information will present many problems of survival and determine many directions of change.” One such direction of change would lead to establishment of a new species. Since hereditary information is transmitted in the form of DNA, the task is twofold: First, to determine what is the “accent” of DNA. Second, to determine how changes in that accent can bring about an incompatibility between members of a species (“discord”) such that they do not effectively reproduce with each other. Being reproductively isolated they would be, by definition, independent species even if, in the extreme case, they showed no anatomical or physiological differences. What we see (the conventional phenotype) is largely determined by proteins. These are linear polymers of basic units – the amino acids. The sequence of amino acids distinguishes one protein from another and determines how each folds into a complex unique structure. Thus, its sequence determines its properties, which may be structural (e.g. tendons, muscle) or cata- lytic (e.g. bringing about chemical changes so that muscles contract). What determines the sequence of amino acids, determines the nature of the result- ing protein, and hence determines the organism. We can equate phenotype with protein and genotype with DNA. The linear sequence of the bases in an organism’s DNA codes for the linear sequences of amino acids in its proteins. If we know its DNA sequence we The Third Base 669 know which proteins the organism can make. Hence, if we fully understand the properties of those proteins, we can predict the organism’s anatomical and physiological characteristics. In short, DNA (the genotype) determines protein (the phenotype). The code which relates a particular base sequence to a particular amino acid is a triplet code. Each amino acid corresponds to a three base sequence (“codon”) in DNA. For example, the amino acid glycine is encoded by GGT, and the amino acid threonine is encoded by ACT. So a protein sequence that happened to consist just of these two amino acids – say glycine, alanine, glycine, glycine, alanine, glycine – could be encoded by the sequence: GGTACTGGTGGTACTGGT ………… (A-1) Where is the accent of DNA? It turns out that the code is degenerate so that each amino acid has four possible codons. Glycine can be encoded by GGT, GGC, GGA or GGG. Threonine can be encoded by ACT, ACC, ACA or ACG. As far as an amino acid is concerned, what matters is the first two bases of its codon. The third base indicates the accent of DNA. Thus, two organisms may make the above amino acid sequence, but their corresponding DNA sequences may have different accents – for example, instead of the above sequence, one might have the sequence: GGCACCGGCGGCACCGGC ………… (A-2) This also encodes the sequence glycine, alanine, glycine, glycine, alanine, glycine. In this particular case in every codon the third base is C (under- lined). But any of the four bases in the third codon position will suffice as far as the “primary information” – i.e. for the amino acid sequence – is concerned. So organisms can vary in their accent (“secondary information”) while maintaining the same primary information (and hence the same phenotype). Although somewhat simplified, the above account is uncontentious and widely agreed upon. More contentious is whether, in the general case, it is this difference in accent that initiates reproductive isolation between two groups living in the same territory. Furthermore, even if this accent difference did bring about reproductive isolation, what would be the mechanism? William Bateson, under some prompting from Crowther (Chapter 17), agreed that the most fundamental form of reproductive isolation, manifest at an early stage as the hybrid sterility seen when recently diverged (allied) species are crossed, was due to an incompatibility that could be characterized cytologi- cally as an incompatibility between paternal and maternal chromosomes when attempting to pair during meiosis (see Fig. 9-5). Thus, if we can understand what makes chromosomes incompatible, we can understand hybrid sterility. And if we understand hybrid sterility, we can understand an origin of species. 670 Appendix

We seek to understand how chromosomes that are homologous (i.e. are alike) can pair with each other. Do they pair by virtue of this likeness (like- with-like), or by virtue of some key-in-lock (sword-in-scabbard) complementarity, which implies that they are not really alike? One must be the sword, the other the scabbard. This paradox was resolved when it was appre- ciated that chromosomal DNA is a duplex consisting of two complementary strands – a “Watson” strand and a “Crick” strand – that pair with each other on the basis of their complementarity. So, in Crowther’s terminology, potentially the sword of one strand can pair with the scabbard of the other (and vice versa). For this swords have to be unsheathed from their own scab- bards and then each inserted into the scabbard of the other. Thus, the Watson strand of one chromosome would pair with the Crick strand of its homologous chromosome (and vice versa). This would require that the Watson strand be displaced from pairing with the Crick strand of its own chromosome. Likewise, the Crick strand of the homologous chromosome would be displaced from pairing with the Watson stand of its own chromosome. Then the cross-pairing could occur. It was Francis Crick, a codiscoverer of the structure of DNA, who worked it out. Most pictures of DNA molecules show them as double-helices with two strands of DNA containing inward-looking bases, which pair with each other, thus joining the two strands to form a duplex. Crick proposed that under certain circumstances parts of the strands would separate (“unpair”) from each other and become outward-looking. In this circumstance, the outward-looking bases in the DNA of a maternal chromosome could pair with complementary outward-looking bases of a paternal chromosome [5]. What about the accent of DNA? It has been suggested that for chromosomes to pair (as when a sword “pairs” with its scabbard) their DNA’s must have similar accents [6–8]. When accents differ (a potentially complement- tary scabbard has been misshapen by many small blows), pairing is dis- rupted. How does accent affect pairing? It has been shown that the degree of unpairing from the parental duplex would be extremely sensitive to the accent of the corresponding DNA. Specifically, the strands in a segment of DNA that is rich in the bases G and C will unpair less readily than those in a segment of DNA that is rich in the bases A and T. If (through multiple small blows, i.e. mutations) maternal and paternal DNAs have begun to differ very slightly in accent (i.e. in their degree of richness in G and C), this will disturb the synchrony of their unpairing from the conventional duplex configuration. Hence they will not display their outward-looking bases in a manner (time and configuration) permitting cross-pairing between maternal and paternal chromosomes.

The Third Base 671

Fig. A-1. Stem-loop extrusion from duplex DNA is exquisitely sensitive to difference in base composition (e.g. X% compared with (X + 1)%). This prevents pairing between otherwise nearly identical DNA sequences. At the left, paternal (P) and maternal (M) duplexes have the same value (X). During meiosis the strands of each duplex syn- chronously open and equivalent stem-loop structures are extruded. Interactions between the loops can then progress to pairing. At the right, the duplexes differ slightly in their base compositions. The maternal duplex, of slightly higher value (X + 1), opens less readily and stem-loop structures are not equivalent. Thus, it is not the lack of identity itself, but the fact that this lack of identity is accompanied by minute differences in base composition, that leads to mispairing. In other words, differences in base composition provide a sensitive index of underlying homology differences 672 Appendix

The parents are reproductively isolated from each other since their hybrid offspring are unable to continue the line by virtue of their sterility (whether or not they flourish due to natural selection). This is shown in Figure A-1, where the “accent” is quantified as the proportion of two bases (G and C) expressed as the G + C percentage among the four bases (A, C, G and T). Darwin’s error was in regarding sterile hybrid offspring in the context of natural selection (acting on their conventional phenotypes), rather than as manifestations of interacting parental genome phenotypes. Galton in Natural Inheritance suggested that an organism has a certain stability, so that internal changes must build up over several generations (i.e. remain latent) until there is a sudden change to a new position of stability. Likewise, Romanes (Chap- ter 5) pointed to a “collective variation” building up in “a whole race or strain.” This can now be seen as a cumulation of differences in percentage G + C building up to a threshold beyond which reproductive isolation becomes increasingly apparent. Future changes in phenotype (that might be favored by natural selection) are then conserved since effective crossing with the main type (that might result in blending of characters in fertile offspring) is impaired. It is of interest that William Bateson, having at last found the evidence convincing that genes (primary information) were located on chromosomes (Chapter 17), looked there in vain for the location of his postulated “residue,” and instead pondered a cytoplasmic location (Chapter 13). He sometimes referred to the “residue” as a “base” – but, of course, this was a different usage from that employed here. We now see that Bateson’s “residue,” Johannsen’s “Grundstock” (a “great central something”) and Guyer’s “general substratum” are likely to be actually within DNA itself – the third base of each codon (in protein-encoding regions). In this context we can see the exon of a gene not as a sequence of consecutive bases, but as a sequence of sets of two adjacent bases, each set being separated from the next by a single “residue base” that is essentially non-genic (despite its location in a gene). Why has this solution been so elusive? Apart from the political games described in this book, the answer is that definitive evidence has been hard to obtain. A simple metaphor may help. Your newly acquired dog must be kept from running off. A barrier is needed. At first you tether it in your garden by a leash. The leash is the first barrier. Then you build a fence. So the leash is no longer necessary. If the first barrier (the leash) is damaged or lost, it may not be noticed. The second barrier (fence) should suffice. On the other hand, the leash then becomes available for other purposes. Thus, following establishment of a second barrier (between species), a first barrier (between species) may degenerate or change in a random way, or may find another role. If a Sherlock Holmes then tried to discern whether there had been an earlier barrier, and what form it had taken, there might be a problem.

Publications of William Bateson

Bateson’s papers are in Punnett’s Scientific Papers of William Bateson (1928), or in Beatrice Bateson’s Memoir (1928). Some are available in Forsdyke’s web-pages: http://post.queensu.ca/~forsdyke/bateson1.htm https://qspace.library.queensu.ca/handle/1974/421 Another resource is the collected papers of Charles Hurst (Experiments in Genetics, Cambridge University Press, 1925).

Bateson, W. (1884) The early stages in the development of Balanoglossus. Quarterly Journal of Microscopical Science 24, 208–236. Bateson, W. (1884) On the development of Balanoglossus. Annals and Magazine of Natural History 13, 65. Bateson, W. (1885) The later stages in the development of Balanoglossus kowalevskii, with a suggestion as to the affinities of the Enteropneusta. Quarterly Journal of Microscopical Science 25(Supplement), 81–122. Bateson, W. (1886) Continued account of the later stages in the development of Balanoglossus kowalevskii, and of the morphology of the Enteropneusta. Quarterly Journal of Microscopical Science 26, 511–533. Bateson, W. (1886) The ancestry of the chordata. Quarterly Journal of Microscopical Science 26, 535–571. Bateson, W. (1888) Suggestion that certain fossils known as Bilobites may be regarded as casts of Balanoglossus. Proceedings of the Cambridge Philosophical Society 6, 298. Bateson, W. (1889) On some variations of Cardium edule, apparently correlated to the conditions of life. Philosophical Transactions of the Royal Society, B. 180, 297–330. Bateson, W. (1889) On some variations of Cardium edule, apparently correlated to the conditions of life. Proceedings of the Royal Society, B. 46, 204–211. Bateson, W. (1889) Notes and memoranda: notes on the senses and habits of some Crustacea. Journal of the Marine Biological Association 1, 211–217. Bateson, W. (1890) The sense organs and perceptions of fishes: with remarks on the supply of bait. Journal of the Marine Biological Association 1, 225–256. Bateson, W. (1890) On some cases of abnormal repetition of parts in animals. Proceed- ings of the Zoological Society, p. 579. Bateson, W. (1890) On some skulls of Egyptian mummified cats. Proceedings of the Cambridge Philosophical Society 7, 68. [Abstract] Bateson, W. (1890) On the nature of supernumary appendages in insects. Proceedings of the Cambridge Philosophical Society 7, 159. [Abstract] 674 Publications of William Bateson

Bateson, W. (1890) Application for Deputy Linacre Professorship, Oxford. [See C. B. Bateson (1928).] Bateson, W. (1890) For Greek. Cambridge University Press, Cambridge. [Fly-sheet] Bateson, W. & Bateson, A. (1891) On variations in floral symmetry of certain plants having irregular corollas. Journal of the Linnean Society, Botany 28, 386–421. Bateson, W. (1892) On variation in the colour of cocoons of Eriogaster lanestris and Saturnia carpini. Transactions of the Entomological Society, London, pp. 45–52. Bateson, W. (1892) On numerical variation in teeth, with a discussion of the conception of homology. Proceedings of the Zoological Society, pp. 102–115. Bateson, W. (1892) On variation in the colour of cocoons, pupae and larvae. Further experiments. Transactions of the Entomological Society, London, pp. 205–214. Bateson, W. (1892) The alleged “aggressive mimicry” of Volucellae. Nature 46, 585. Bateson, W. (1892) The alleged “aggressive mimicry” of Volucellae. Nature 47, 77. Bateson, W. (1892) Exhibition of, and remarks upon, some crab’s limbs bearing supernumary claws. Proceedings of the Zoological Society, p. 76 [A five line note.] Bateson, W. & Brindley, H. H. (1892) On some cases of variation in secondary sexual characters statistically examined. Proceedings of the Zoological Society, pp. 585–594. Bateson, W. (1893) Exhibition of and remarks upon an abnormal foot of a calf. Pro- ceedings of the Zoological Society, pp. 530–531. Bateson, W. (1894) Materials for the Study of Variation, Treated with Especial Regard to the Discontinuity in the Origin of Species. Macmillan, London. Bateson, W. (1894) Exhibition of specimens of the common pilchard (Clupea pilchardus) showing variation in the number and size of the scales. Proceedings of the Zoological Society, p. 164. Bateson, W. (1894) Exhibition of specimens and drawings of a phytophagus beetle, in illustration of discontinuous variation in colour. Proceedings of the Zoological Society, p. 391. Bateson, W. (1894) On two cases of colour-variation in flat-fishes, illustrating principles of symmetry. Proceedings of the Zoological Society, pp. 246–249. Bateson, W. (1895) Note in correction of a paper on colour-variation in flat-fishes. Proceedings of the Zoological Society, pp. 890–891. Bateson, W. (1895) The origin of the cultivated Cineraria. Nature 51, 605–607. Bateson, W. (1895) The origin of the cultivated Cineraria. Nature 52, 29. Bateson, W. (1895) The origin of the cultivated Cineraria. Nature 52, 103. Bateson, W. (1895) Notes on hybrid Cinerarias produced by Mr. Lynch and Miss Pertz. Proceedings of the Cambridge Philosophical Society 10, 308. Bateson, W. (1895) On the colour variations of a beetle of the family Chrysomelidae statistically examined. Proceedings of the Zoological Society, pp. 850–860. Bateson, W. (1896) Exhibition of, and remarks upon, three pigeons showing webbing between the toes. Proceedings of the Zoological Society, pp. 989–990. Bateson, W. (1897) Habits of Zygaena exulens. Entomological Record 9, 328. [Short note on date of emergence and relation to snow.] Bateson, W. (1897) On progress in the study of variation. Science Progress 6, 554–568. Bateson, W. (1898) On progress in the study of variation. Science Progress 7, 53–68. Bateson, W. (1898) Experiments in the crossing of local races of Lepidoptera. Entomo- logical Record 10, 241. [Report of an exhibition of napi, egeria, etc., at the Publications of William Bateson 675

Fourth International Zoological Congress, Cambridge. This, and a passage in Problems in Genetics, form the only published account of this work.] Bateson, W. (1898) Protective colouration of Lepidopterous pupae. Entomological Record 10, 285. Bateson, W. (1900) Hybidization and cross-breeding as a method of scientific investigation. Journal of the Royal Horticultural Society 24, 59–66. [From the First International Conference on Plant Breeding and Hybridization, London, 1899.] Bateson, W. (1900) On a case of homoeosis in a crustacean of the genus Asellus. Antennule replaced by a mandible. Proceedings of the Zoological Society, pp. 268–271. Bateson, W. (1900) British lepidoptera. Entomological Record 12, 231. [Review of volume 2 of Tutt’s British Lepidoptera.] Bateson, W. (1900) Collective inquiry as to progressive melanism in moths. Entomo- logical Record 12, 140. [Memorandum from the Evolution Committee of the Royal Society, London; also in Entomology Monthly Magazine, p. 139.] Bateson, W. & Pertz, D. (1900) Notes on the inheritance of variation in the corolla of Veronica buxhaumii. Proceedings of the Cambridge Philosophical Society 10, 78–92. Bateson, W. (1901) Problems of heredity as a subject for horticultural investigation. Journal of the Royal Horticultural Society 25, 54–61. [Read May 8, 1900] Bateson, W. (1901) Heredity, differentiation, and other conceptions of biology: a consideration of Professor Karl Pearson’s paper ‘On the principle of Homoty- posis.’ Proceedings of the Royal Society 69, 193–205. Bateson, W. (1902) Introductory note to the translation of ‘Experiments in plant hybridization’ by Gregor Mendel. Journal of the Royal Horticultural Society 26, 1–3. Bateson, W. (1902) Mendel’s Principles of Heredity. A Defence. Cambridge University Press, Cambridge. [Reprinted (1903) as: The Problems of Heredity and Their Solution. Smithsonian Museum, Washington.] Bateson, W. (1902) Note on the resolution of compound characters by cross-breeding. Proceedings of the Cambridge Philosophical Society 12, 50–54. Bateson, W. (1902) British lepidoptera. Entomological Record 14, 320. [Review of Vol. 3 of Tutt’s British Lepidoptera.] Bateson, W. & Saunders, E. R. (1902) Experimental studies in the physiology of heredity. Reports to the Evolution Committee of the Royal Society I, pp. 1–160. Bateson, W. (1903) Variation and Differentiation in Parts and Brethren. Cambridge University Press, Cambridge. Bateson, W. (1903) On Mendelian heredity of three characters allelomorphic to each other. Proceedings of the Cambridge Philosophical Society 12, 153–154. Bateson, W. (1903) The present state of knowledge of colour-heredity in mice and rats. Proceedings of the Zoological Society, London 2, 71–99. Bateson, W. (1903) Mendel’s principles of heredity in mice. Nature 67, 462, 585; 68, 33. Bateson, W. (1904) Practical aspects of the new discoveries in heredity. Memoirs of the Horticultural Society of New York 1, 1–9. [Opening paper at the Second Interna- tional Conference on Plant Breeding and Hybridization, New York, September, 1902.] 676 Publications of William Bateson

Bateson, W. (1904) Presidential address to Section D (Zoology) of the British Asso- ciation, Cambridge. [Reprinted in part as: Heredity and evolution. Popular Science Monthly. New York, pp. 522–531.] Bateson, W. (1904) Albinism in Sicily. A correction. Biometrika 3, 471–472. [Letter correcting Weldon.] Bateson, W. (1904) A natural history of British lepidoptera. Entomological Record 16, 234. [Review of volume 4 of Tutt’s British Lepidoptera.] Bateson, W. (1904) Exhibition of a series of Primula sinensis. Linnean Society. Report of General Meeting. Feb. 18, pp. 2–3. Bateson, W. (1905) Experimental Studies in the Physiology of Heredity. Report to Committee. British Association 1904 pp. 346–348. Bateson, W. (1905) Evolution for amateurs. The Speaker, June 24. [A review of Weismann’s The Evolution Theory (1904). Edward Arnold, London.] Bateson, W. (1905) Compulsory Greek at Cambridge. Nature 71, 390. Bateson, W. (1905) Heredity in the physiology of nations. The Speaker, Oct. 14. [A review of Archdall Reid’s The Principles of Heredity.] Bateson, W. (1905) Practical aspects of the new discoveries in heredity. Journal of the Royal Horticultural Society 29, 417–419. [Abstract of paper from Second Conference on Plant-Breeding and Hybridization, New York, 1902.] Bateson, W. (1905) The exhibition of, and remarks upon specimens of fowls, illustrating peculiarities in the heredity of white plumage. Proceedings of the Zoological Society, London 2, 3. Bateson, W. (1905) Albinism in Sicily. A further correction. Biometrika 4, 231. Bateson, W. (1905) Experimental Studies in the Physiology of Heredity. Report to Committee. British Association 10. Bateson, W. & Punnett, R. C. (1905) A suggestion as to the nature of the ‘walnut’ comb in fowls. Proceedings of the Cambridge Philosophical Society 13, 165–168. Gregory, R. P. & Bateson, W. (1905) On the inheritance of heterostylism in Primula. Proceedings of the Royal Society, B. 76, 581–586. Bateson, W., Saunders, E. R., & Punnett, R. C. (1905) Experimental studies in the physiology of heredity. Reports to the Evolution Committee of the Royal Society II, pp. 1–131. [In his listing in Scientific Papers, Punnett adds H. Killby to the author list, and omits C. C. Hurst whose “Experiments with Poultry,” although separate, formed part of the report (pp. 131–154).] Bateson, W. (1906) Science of sorts. The Speaker, Apr. 14. [Review of Burke’s The Origin of Life.] Bateson, W. (1906) The progress of genetic research. Gardeners’ Chronicle. Aug. 4. [An inaugural address to the Third Conference on Hybridism and Plant Breeding. Also published (1907) as the Report of the Third International Conference 1906 on Genetics. Edited by W. Wilks. Royal Horticultural Society, London. pp. 90–97.] Bateson, W. (1906) Mendelian heredity and its application to man. Brain, Part 114, pp. 1–23. [An address to the Neurological Society, London.] Bateson, W. (1906) The progress of genetics since the rediscovery of Mendel’s papers. Progressus Rei Botanicae 1, 368–418. [Jena] Bateson, W. (1906) A text-book of genetics. Nature 74, 146. [Review of Lotsy’s Vorlesungen über Descendenz-theorien. I.] Publications of William Bateson 677

Bateson, W. (1906) Coloured tendrils of sweet-peas. Gardeners’ Chronicle (Series 3) 39, 333. Bateson, W., Saunders, E. R., & Punnett, R. C. (1906) Further experiments on inheritance in sweet peas and stocks: preliminary account. Proceedings of the Royal Society, B. 77, 236–238. Bateson, W., Saunders, E. R., & Punnett, R. C. (1906) Experimental studies in the physiology of heredity. Reports to the Evolution Committee of the Royal Society III, pp. 1–53. Bateson, W. (1907) Facts limiting the theory of heredity. Science 26, 649–660. [Also (1912) Proceedings of the International Congress of Zoology (Boston, 1907) 7, 306–389.] Bateson, W. (1907) Trotting and pacing: dominant and recessive. Science 26, 908. Bateson, W. (1908) The Methods and Scope of Genetics. Cambridge University Press, Cambridge. [Inaugural lecture, Oct. 23.] Bateson, W. (1908) Lectures on evolution. Nature 78, 386. [Review of Lotsy’s Vor- lesungen über Descendenz-theorien, II.] Bateson, W. (1908) British Association discussion on sex-determination. Correction. Nature 78, 665. Bateson, W. & Punnett, R. C. (1908) The heredity of sex. Science 27, 785–787. Bateson, W., Saunders, E. R., & Punnett, R. C. (1908) Experimental studies in the physiology of heredity. Reports to the Evolution Committee of the Royal Society IV, pp. 1–59. [H. B. Killby is a coauthor in Saunder’s section. F. M. Durham has sections on coat color in mice (pp. 41–53), and sex in canaries (pp. 57–59). L. Doncaster has a section on sex-inheritance in moths (pp. 53–57)]. Bateson, W. (1909) Summary of “Mendel’s Principles.” Harmsworth’s World’s Great Books. Bateson, W. (1909) Heredity and variation in modern lights. In: Darwin and Modern Science. Edited by A. C. Seward. Cambridge University Press, Cambridge, pp. 85–101. Bateson, W. (1909) Mendel’s Principles of Heredity. Cambridge University Press, Cambridge. Conklin, E. G. et al. (1909) William Keith Brooks. A sketch of his life by some of his former pupils and associates. Journal of Experimental Zoology 9, 1–51. Wheldale, M., Marryat, D. C. E., & Sollas, I. B. J. (1909) Experimental studies in the physiology of heredity. Reports to the Evolution Committee of the Royal Society V, pp. 1–78. [Bateson will have read and approved this report.] Bateson, W. (1910) Boyle lecture. Delivered at Oxford 1909. [Manuscript not found.] Bateson, W. (1911) An appreciation of J. W. Tutt. Entomological Record 23, 123–124. Bateson, W. (1911) The John Innes Horticultural Institute Report for 1910. Gardeners’ Chronicle (Series 3) 49, 179. [Subsequent annual reports were privately printed for the Council of the Institution.] Bateson, W. (1911) Recent advances in the genetics of plants. Nature 88, 36–37. [Review of Baur’s Einführung in die experimentelle Vererbungslehre.] Bateson, W. (1911) Presidential address to the agricultural subsection, British Associa- tion, Portsmouth. British Association Report, pp. 587–596. Bateson, W. & de Vilmorin, P. L. (1911) A case of gametic coupling in Pisum. Pro- ceedings of the Royal Society, B. 84, 9–11. 678 Publications of William Bateson

Bateson, W. & Punnett, R. C. (1911) On the interrelations of genetic factors. Pro- ceedings of the Royal Society, B. 84, 3–8. Bateson, W. & Punnett, R. C. (1911) Reduplication of terms in series of gametes. 4th Conference Internationale de Génétique, Paris. [Conference Proceedings published in 1913, pp. 99–100.] Bateson, W. & Punnett, R. C. (1911) The inheritance of the peculiar pigmentation of the silky fowl. Journal of Genetics 1, 185–203. Bateson, W. & Punnett, R. C. (1911) On gametic series involving reduplication of certain terms. Journal of Genetics 1, 293–302. [Also in Verh. Naturforsch. Verein. Brünn, 49, 324–334.] Bateson, W. (1912) Biological Fact and the Structure of Society. Clarendon Press, Oxford. [Herbert Spencer lecture, delivered February 28.] Bateson, W. (1912) Lectures to Royal Institution (Fullerian Professorship). Gardeners’ Chronicle 51, pp. 57, 74, 89, 104, 120, 123, 139. Bateson, W. (1913) Problems of the cotton plant. Nature 90, 667–668. [Review of Balls’ The Cotton Plant in Egypt.] Bateson, W. (1913) Problems in Genetics. Yale University Press. [Silliman Memorial lectures.] Bateson, W. (1913) Heredity. British Medical Journal 2, 359–362; Lancet 2, 451–454. [Address to Seventeenth International Congress of Medicine.] Bateson, W. (1913) Oenothera crosses. Gardeners’ Chronicle, Series 3. 54, 406. Bateson, W. (1913) Discussion of Lotsy’s theory of evolution by hybridization. Proceedings of the Linnean Society, p. 89. [A paragraph.] Bateson, W. (1914) Presidential addresses, British Association Meetings at Melbourne and Sydney. Nature 93, 635–642, 674–681. [Also (1916) Smithsonian Report, pp. 359–394.] Bateson, W. (1914) Royal Institution and Fullerian lectures. Gardeners’ Chronicle 55, pp. 74, 92, 112, 131, 149, 171, 332–33. Bateson, W. (1915) Address to Salt Schools. [In C. B. Bateson, 1928.] Bateson, W. & Pellew, C. (1915) On the genetics of “rogues” among culinary peas, Pisum sativum. Journal of Genetics 5, 13–36. Bateson, W. & Pellew, C. (1915) Note on an orderly dissimilarity in inheritance from different parts of a plant. Proceedings of the Royal Society, B. 89, 174–175. Bateson, W. (1916) Review of The Mechanism of Mendelian Heredity by Morgan et al. Science 44, 536–543. Bateson, W. (1916) Notes on experiments with flax at the John Innes Horticultural Institution. Journal of Genetics 5, 199–201. Bateson, W. (1916) Root-cuttings, chimaeras and “sports.” Journal of Genetics 6, 75–80. Bateson, W. (1917) Gamete and zygote. A lay discourse. [Henry Sidgwick lecture, Newnham College. In C. B. Bateson (1928).] Bateson, W. (1917) Philippe Leveque de Vilmorin. Proceedings of the Linnean Society of London, Session 130, pp. 44–46. Bateson, W. (1917) The place of science in education. Cambridge Essays on Education. Cambridge University Press, Cambridge. Bateson, W. (1917) The ear of Dionysius. The Times Literary Supplement, May 3 and May 17. Publications of William Bateson 679

Bateson, W. (1917) Is variation a reality? Nature 99, 43. [Review of Lotsy’s Evolution by Means of Hybridization.] Bateson, W. & Thomas, R. H. (1917) Note on a pheasant showing abnormal sex- characters. Journal of Genetics 6, 163–164. Bateson, W. (1918) Gamete and zygote. Proceedings of the Royal Institute of Great Britain. Feb. 15, pp. 1–4. [Abstract. Also in C. B. Bateson, 1928.] Bateson, W. (1919) Science and nationality. Edinburgh Review 229, 123–138. [Inaugural address to the Yorkshire Natural Science Association.] Bateson, W. (1919) Progress in Mendelism. Nature 104, 214. Bateson, W. (1919) Linkage in the silk worm. A correction. Nature 104, 315. Bateson, W. (1919) Dr. Kammerer’s testimony to the inheritance of acquired characters. Nature 103, 344. Bateson, W. (1919) Studies in variegation. I. Journal of Genetics 8, 93–99. Bateson, W. & Sutton, I. (1919) Double flowers and sex-linkage in Begonia. Journal of Genetics 8, 199–207. Bateson, W. (1920) Classical education and science men. Précis of evidence offered to the Prime Minister’s Committee on Classics, June 1920. [In C. B. Bateson (1928).] Bateson, W. (1920) Genetic segregation. Proceedings of the Royal Society, B. 91. 358–368. [Croonian lecture. Also in (1920) Nature 105, 531 [abstract] with a correction (1921) Nature 107, 233; and in (1921) American Naturalist 55, 1–19.] Bateson, W. (1920) Organization of scientific work. Nature 105, 6. [Letter concerning state organization of science in India.] Bateson, W. (1920) Prof. L. Doncaster, FRS. Nature 105, 461–462. Bateson, W. & Pellew, C. (1920) The genetics of “rogues” among culinary peas, Pisum sativum. Proceedings of the Royal Society, B. 91, 186–195. Bateson, W. (1921) Common sense in racial problems. Eugenics Review13, 325–338 [Galton lecture.] Bateson, W. (1921) Leonard Doncaster, 1877–1920. Proceedings of the Royal Society, B. 92, 41–46. Bateson, W. (1921) Root cuttings and chimaeras. II. Journal of Genetics 11, 91–97. Bateson, W. (1921) The determination of sex. Nature 106, 719. [Review of Gold- schmidt’s Mechanismus und Physiologie der Geschlechtsbestimmung.] Bateson, W. (1921) Variegation in a fern. Nature 107, 233. [Correction to Croonian lecture.] Bateson, W. (1921) Classical and modern education. Nature 108, 308. [Review of Classics in Education, the report of the Prime Minister’s Committee to inquire into the position of classics in the educational system of the United Kingdom.] Bateson, W. (1922) Articles on “Genetics,” “Mendelism,” and “Sex.” In the twelfth edition of the Encyclopedia Britannica. Bateson, W. & Gairdner, A. E. (1921) Male sterility in flax, subject to two types of segregation. Journal of Genetics 11, 269–275. Bateson, W. (1922) Evolution and education. In: Ideals, Aims and Methods in Edu- cation. The New Educator’s Library. Isaac Pitman Ltd., London. [Written 1915.] Bateson, W. (1922) Evolutionary faith and modern doubts. Science 55, 55–61. [Address to the American Association for the Advancement of Science, Toronto. Also in Nature 109, 553–556.] 680 Publications of William Bateson

Bateson, W. (1922) Genetical analysis and the theory of natural selection. Science 55, 373. [A correction of misquotation from Historia Plantarum, and a reply to Osborn.] Bateson, W. (1922) Interspecific sterility. Nature 110, 76. [Controversy following the Toronto address.] Bateson, W. (1922) Darwin and evolution, limits and variation. The Times, Apr. 13. [Letter to the Editor concerning the Toronto address.] Bateson, W. (1923) Area of distribution as a measure of evolutionary age. Nature 111, 39. [Review of Willis’s Age and Area. Cambridge University Press, 1922.] Bateson, W. (1923) Somatic segregation in plants. In: Report of International Horti- cultural Congress, Amsterdam, pp. 155–156. Bateson, W. (1923) The revolt against the teaching of evolution in the United States. Nature 112, 313–314. Bateson, W. (1923) Note on the nature of plant chimaeras. Studia Mendeliana, Brünn, pp. 9–12. Bateson, W. (1923) Dr. Kammerer’s Alytes. Nature 111, 738, 878. Gregory, R. P., de Winton, D., & Bateson, W. (1923) Genetics of Primula sinensis. Journal of Genetics 13, 219–253. Bateson, W. (1924) Progress in biology. Nature 113, 644–646, 681–682. [Address given (Mar. 12) at the Centenary of Birkbeck College.] Bateson, W. (1925) Huxley and evolution. Nature 115, 715–717. Bateson, W. (1925) Mendeliana. Nature 115, 827–830. [Review of Corren’s collected papers and Iltis’s Life of Mendel.] Bateson, W. (1925) Evolution and intellectual freedom. Nature 116, 78. [Supplement] Bateson, W. (1925) Science in Russia. Nature 116, 681. [Account of visit.] Bateson, W. & Bateson, G. (1925) On certain aberrations of the red-legged partridges Alectoris rufa and saxatilis. Journal of Genetics 16, 101–123. Bateson, W. (1926) Segregation: being the Joseph Leidy Memorial Lecture of the University of Pennsylvania, 1922. Journal of Genetics 16, 201–235. Bateson, C. B. (1928) Letters from The Steppe Written in the Years 1886–1887 by William Bateson. Cambridge University Press. Bateson, C. B. (1928) William Bateson, F. R. S. Naturalist. His Essays and Addresses Together with a Short Account of his Life. Cambridge University Press. [Includes Beatrice’s Memoir.] Hall, A. D. (1928) Bateson’s experiments on bolting in sugar beet and mangolds. Journal of Genetics 20, 219–232. Punnett, R. C. (1928) Scientific Papers of William Bateson, Vols. 1 and 2. Cambridge University Press, Cambridge. References and Notes

In works of this nature there is usually a detailed list of references. But much of the early literature is now available on line, in word- or phrase-searchable form, through such operations as Robert Robbin’s Electronic Scholarly Publishing (ESP) project (history of genetics), the JSTOR scholarly journals archive, the Project Gutenberg, and Google Scholar. Furthermore, primary sources such as the Darwin Correspon- dence Online Database, and archives, such as that at the John Innes Centre, can be (or will soon be) accessible for searching. In this circumstance other considerations, such as the need for conciseness, come into play. Accordingly, the minimal information for retrieving most references is given in the text (usually the date of a letter, or of one of Bateson’s publications). Other pertinent references are listed below on a chapter by chapter basis.

Prologue 1. Darwin, C. (1959) The Origin of Species by Natural Selection, or The Pre- servation of Favoured Races in the Struggle for Life. John Murray, London. 2. Mendel, G. (1866) Experiments in plant-hybridization. Verhandlungen des naturforschenden Vereines in Brunn 4, 3–47. 3. Forsdyke, D. R. (2001) The Origin of Species, Revisited. McGill-Queen’s University Press, Montreal, Quebec, Canada. 4. Forsdyke, D. R. (2006) Evolutionary Bioinformatics. Springer, New York. 5. Gould, S. J. (2002) The Structure of Evolutionary Theory. Harvard Univer- sity Press, Cambridge, MA. 6. Lesch, J. E. (1975) The role of isolation in evolution: George J. Romanes and John. T. Gulick. Isis 66, 483–503. 7. Provine, W. B. (1986) Sewell Wright and Evolutionary Biology. University of Chicago Press, Chicago, IL. pp. 207. 8. Bateson, C. B. (1928) William Bateson, F. R. S. Naturalist. His Essays and Addresses Together with a Short Account of his Life. Cambridge University Press, Cambridge. [Beatrice’s memoir constitutes the first 160 pages.] 9. Bateson, C. B. (1928) Letters from the Steppe Written in the Years 1886- 1887 by William Bateson. Cambridge University Press, Cambridge. 10. Coleman, W. (1968) The Bateson papers. The Mendel Newsletter 2, 1–3. 11. Provine, W. (1971) The Origins of Theoretical Population Genetics. University of Chicago Press, Chicago, IL. 12. Coleman, W. (1970) Bateson and chromosomes: Conservative thought in science, Centaurus 15, 228–314. 13. Lipset, D. (1980) Gregory Bateson. The Legacy of a Scientist. Prentice-Hall, Englewood Cliffs, NJ. 14. Cock, A. G. (1973) William Bateson, Mendelism and biometry. Journal of the History of Biology 6, 1–36. 682 References and Notes

15. Cock, A. G. (1977) The William Bateson papers. Mendel Newsletter 14, 1–4. 16. Pais, A. (1982) The Science and Life of Albert Einstein. Oxford University Press, New York. 17. Forsdyke, D. R. (2000) Tomorrow’s Cures Today? Harwood Academic, Newark, NJ. 18. Harrison, E. (1987) Whigs, prigs and historians of science. Nature 329, 213–214. 19. Bateson, C. B. (1927) Letter to C. C. Hurst, June 13. In: Hurst, R. (1974) The Evolution of Genetics. Unpublished manuscript in Cambridge University Library. 20. Wyhe, J. van (2007) Mind the gap: Did Darwin avoid publishing his theory for many years? Notes and Records of the Royal Society 61, 177–205.

1 A Cambridge Childhood (1861–1882) 1. Jones, H. F. (1919) Samuel Butler, Author of Erewhon (1835-1902): A Memoir, Vols. 1 and 2. Macmillan, London. 2. Weismann, A (1909) Charles Darwin. Contemporary Review 96, 1–22. 3. Huxley, J. (1959) The evolutionary vision. In: Evolution After Darwin. Edited by S. Tax (1960). University of Chicago Press, Chicago, IL. 4. Smocovitis, V. B. (1999) The 1959 Darwin Centennial celebration in America. Osiris 14, 274–323. 5. Bateson, M (1895) Professional Women and their Professions. Horace Cox, London. 6. Heitland, W. E. (1926) After Many Years. Cambridge University Press, Cambridge. 7. Forsdyke, D. R. (2001) The Origin of Species, Revisited. McGill-Queen’s University Press, Montreal, Quebec, Canada. 8. Geison, G. L. (1978) Michael Foster and the Cambridge School of Physiology. Princeton University Press, Princeton, NJ. 9. Scharpey-Schafer, E. (1927) History of the Physiological Society during its First Fifty Years. The Physiological Society, London. 10. Harris, H. (1999) The Birth of the Cell. Yale University Press, New Haven, CT. 11. Haeckel, E. (1909) Charles Darwin as an anthropologist. In: Darwin and Modern Science. Essays in Commemoration of the Centenary of the Birth of Charles Darwin and of the Fiftieth Anniversary of the Publication of the Origin of Species. Edited by A. C. Seward. Cambridge University Press, Cambridge. 12. Lankester, E. R. (1890) The history and scope of zoology. In: The Advance- ment of Science. Macmillan, London. [Reproduced from the ninth edition of the Encyclopaedia Britannica.] 13. Raverat, G. (1952) Period Piece: A Cambridge Childhood. Faber & Faber, London. 14. Heitland, W. E. (1926) William Bateson 1861-1926. The Eagle 44, 227–230. [St. John’s College magazine] References and Notes 683

15. Garnett, R. (1991) Constance Garnett: A Heroic Life. Sinclair-Stevenson, London.

2 From Virginia to the Aral Sea (1883–1889) 1. Brooks, W. K. (1883) The Law of Heredity. A Study of the Cause of Variation and the Origin of Living Organisms. Murphy, Baltimore, MD. 2. Hall, B. K. (2005) Betrayed by Balanoglossus: William Bateson’s rejection of evolutionary embryology as the basis for understanding evolution. Jour- nal of Experimental Zoology 304B, 1–17. 3. Punnett, R. C. (1926) William Bateson. Edinburgh Review 244, 71–86. 4. Conklin, E. G. et al. (1910) William Keith Brookes: A sketch of his life by some of his former pupils and associates. Journal of Experimental Zoology 9, 1–51. 5. Lankester, E. R. (1890) The history and scope of zoology. In: The Advance- ment of Science. Macmillan, London. [Reproduced from the ninth edition of the Encyclopaedia Britannica.] 6. Garnett, D. (1954) The Golden Echo, Vol. 1. Intimations of Mortality. Chatto & Windus, London. 7. Fowler, H. (1996) Cambridge Women: Twelve Portraits. Edited by E. Shils & C. Blacker, Cambridge University Press, Cambridge. 8. Lankester, E. R. (1890) The scientific results of the international fisheries expedition. In: The Advancement of Science. Macmillan, London.

3 Galton 1. Wallace, A. R. (1889) Darwinism: An Exposition of the Theory of Natural Selection with Some of its Applications. Macmillan, London. 2. Galton, F. (1889) Natural Inheritance. John Murray, London. 3. Weismann, A. (1889) Essays upon Heredity and Kindred Biological Prob- lems. Translated by E. B. Poulton, S. Schönland, & A. E. Shipley. Clarendon, Oxford. [Bateson’s copy is inscribed “From A. E. Shipley, June 4, 1889.”] 4. Perhaps Bateson is here referring to the fact that in ancient Greek the “n” of pan changes to “m” before a word beginning with “m”, so it should not be “panmixia,” but “pammixia.” There is no Greek noun corresponding to this. There is a Greek adjective from the same roots – pammiges – meaning “mixed of all sorts” as in an army made up of men from different social statuses and ethnic origins. 5. Galton, F. (1872) On blood-relationship. Proceedings of the Royal Society 20, 394–402. 6. Galton, F. (1865) Hereditary talent and character. Macmillan’s Magazine 12, 157–166, 318–327. 7. Galton, F. (1875) A theory of heredity. Contemporary Review 27, 80–95. 8. Galton, F. (1875) A theory of heredity. Journal of the Anthropological Institute 5, 329–348. 684 References and Notes

9. Romanes, G. J. (1893) An Examination of Weismannism. Open Court Pub- lishing, Chicago, IL. 10. Wilson, E. B. (1896) The Cell in Development and Inheritance. Macmillan, New York. 11. Darwin, C. (1868) The Variation of Animals and Plants under Domestication. John Murray, London. 12. Sanders, A. (1870) On Mr. Darwin’s hypothesis of pangenesis as applied to the faculty of memory. Journal of Anthropology 1, 144–149. 13. Galton, F. (1871) Experiments in pangenesis, by breeding from rabbits of a pure variety, into whose circulation blood taken from other varieties had previously been largely transfused. Proceedings of the Royal Society 19, 393–410. 14. Darwin, C. (1871) Pangenesis. Nature 3, 502–503. 15. Romanes, E. (1896) The Life and Letters of George John Romanes. Long- mans, Green & Co., London. 16. Betteridge, K. (2003). A history of farm animal embryo transfer and some associated techniques. Animal Reproduction Science 79, 203–244. 17. Galton, F. (1897) The average contribution of each of several ancestors to the total heritage of offspring. Proceedings of the Royal Society 61, 401–413. 18. Jenkin, F. (1867) The origin of species. The North British Review 46, 277–318. 19. Galton, F. (1892) Finger Prints. Macmillan, London. 20. Yule, G. U. (1902) Mendel’s laws and their probable relations to intra-racial heredity. The New Phytologist 1, 193–207, 222–238. 21. De Vries, H. (1889) Intracellulare Pangenesis. Gustav Fischer, Jena. Translated by C. S. Gager (1910). Open Court Publishing, Chicago. 22. Roll-Hansen, N. (1978) The genotype theory of Wilhelm Johannsen and its relation to plant breeding and the study of evolution. Centaurus 22, 201–235. 23. Johansson, W. (1903) Ueber Erblichkeit in Populationen und in reinen Linien. Gustav Fischer, Jena. 24. Weldon, W. F. R. & Pearson, K. (1903) Inheritance in Phaseolus vulgaris. Biometrika 2, 499–503. 25. Yule, G. U. (1904) Professor Johannsen on heredity. Nature 69, 223–224. 26. Shull, G. H. (1905) Galtonian regression in the ‘pure line’. Torreya 5, 21–25.

4 Variation (1890–1894) 1. Barnes, E. (1998) The early career of George John Romanes 1867–1878. Unpublished undergraduate thesis, Cambridge University Press, Cambridge. 2. Romanes, E. (1896) The Life and Letters of George John Romanes. Longmans, Green & Co., London. 3. Romanes, G. J. (1873) Permanent variation of colour in fish. Nature 8, 101. 4. Romanes, G. J. (1896) A Selection from the Poems of George John Romanes. Edited by T. H. Warren. Longmans, Green & Co., London. 5. Bateson, A. & Darwin, F. (1888) The effect of stimulation on turgescent vegetable tissues. Linnean Journal (Botany) 24, 1–27. [Read at the Linnean Society on January 20th 1887; Proceedings of the Linnean Society 99, 8.] References and Notes 685

6. Bateson, A. & Darwin, F. (1888) On a method of studying geotropism. Annals of Botany 2, 65–68. 7. Bateson, A. (1888) The effect of cross-fertilization of inconspicuous flowers. Annals of Botany 1, 255–261. 8. Bateson, A. (1889) On the change of shape exhibited by turgescent pith in water. Annals of Botany 4, 117–125. 9. Bateson, W. & Bateson, A. (1891) On the variation in the floral symmetry of certain plants having irregular corollas. Linnean Society Journal 28, 386–421. 10. Raverat, G. (1952) Period Piece: A Cambridge Childhood. Faber & Faber, London. 11. Romano, T. M. (2002) Making Medicine Scientific. John Burdon Sanderson and the Culture of Victorian Science. Johns Hopkins University Press, Baltimore, MD. 12. Lankester, E. R. (1890) The history and scope of zoology. In: The Advancement of Science. Macmillan, London. [Reproduced from the ninth edition of the Encyclopaedia Britannica.] 13. Huxley, T. H. (1859) Letter to Lyell, June 25. In: Life and Letters of Thomas Henry Huxley. Appleton (1901), New York. 14. Darwin, C. (1875) The Variation of Animals and Plants under Domestica- tion. John Murray, London. [Here Darwin cites Owen’s contrast between “metamorphosis” and “metagenesis.” In the latter case “the new parts are not moulded upon the inner surfaces of the old ones. The plastic force has changed its mode of operation. The outer case, and all that gave form and character to the precedent individual, perish, and are cast off; they are not changed into the corresponding parts of the new individual. These are due to a new and distinct developmental process.”] 15. Goldschmidt, R. (1940) The Material Basis of Evolution. Yale University Press, New Haven, CT. [Silliman Lectures] 16. MacAlister, A. (1894) Materials for the study of variation. The Expositor 53, 375–380. 17. Galton, F. (1894) Discontinuity in evolution. Mind (new series) 3, 362–372. 18. Pearson, K. (1906) Walter Frank Raphael Weldon 1860-1906. Biometrika 5, 1–52. 19. Jones, H. F. (1919) Samuel Butler, Author of Erewhon (1835-1902): A Memoir, Vols. 1 and 2. Macmillan, London. 20. Weldon, W. F. R. (1890) The variations occurring in certain decapod crustacean: 1. Crangon vulgaris. Proceedings of the Royal Society 47, 445–453. 21. Weldon, W. F. R. (1892) On certain correlated variations in Crangon vulgaris. Proceedings of the Royal Society 51, 2–21. 22. Delboeuf, J. (1877) A law of mathematics applicable to the theory of transformation. Kosmos 2, 105–127. [An English translation of this paper appears in Forsdyke’s web-pages.] 23. Provine, W. B. (1986) Sewall Wright and Evolutionary Biology. University of Chicago Press, Chicago, IL. 24. Weldon, W. F. R. (1894) The study of animal variation. Nature 50, 25–26.

686 References and Notes

5 Romanes 1. Forsdyke, D. R. (2001) The Origin of Species, Revisited. McGill-Queen’s University Press, Montreal, Quebec, Canada. 2. Romanes, G. J. (1886) Physiological selection: An additional suggestion on the origin of species. Nature 34, 314–316, 336–340, 362–365. 3. Romanes, G. J. (1897) Darwin, and After Darwin. 3. Isolation and Physio- logical Selection. Longmans, Green & Co., London. 4. Romanes, G. J. (1874) Rudimentary organs. Nature 9, 441–442; Disuse as a reducing cause in species. Nature 10, 164. 5. Wells, H. G. (1934) Experiment in Autobiography. Macmillan, New York. 6. Catchpool, E. (1884) An unnoticed factor in evolution. Nature 31, 4. 7. Romanes, E. (1896). The Life and Letters of George John Romanes. Long- mans, Green & Co., London. 8. Crichton, M. (1990) Jurassic Park. Knopf, New York. 9. Dyer, W. T. (1888) Opening Address to the British Association, Section D. Nature 38, 473–480. 10. Darwin, F. (1886) Physiological selection and the origin of species. Nature 34, 407. 11. Belt, T. (1874) A Naturalist in Nicaragua. Dent, London. 12. Milner, R. (1999) Huxley’s bulldog: The battles of E. Ray Lankester (1846- 1929). The Anatomical Record 257, 90–95. 13. Meadows, A. J. (1972) Science and Controversy. A Biography of Sir Norman Lockyer. MIT Press, Cambridge, MA. [The Lankester letter (Sept. 25) was probably written in 1888.] 14. Lankester, E. R. (1889) Darwinism. Nature 40, 566–570. 15. Lankester, E. R. (1905) Nature and Man. Nature 72, 184. [Abstract of his Romanes Lecture.] 16. Huxley, T. H. (1888) Obituary Notices of Fellows Deceased. Proceedings of the Royal Society, 44. [Obituary of Charles Darwin; also in Darwiniana, Collected Essays. Macmillan, London, 1893.] 17. Huxley, T. H. (1894) Letter to Bateson, (Feb. 20). [Nephelococcygia de- rives from The Birds, a play by Aristophanes (414 B. C.), where birds imag- ine shapes in clouds.] 18. Ralston, H. J. (1944) Neuromuscular physiology. G. J. Romanes on the Excitability of Muscle. Science 100, 123–124. 19. Eimer, G. H. T. (1890) Organic Evolution as the Result of the Inheritance of Acquired Characters According to the Laws of Organic Growth. Translated by J. T. Cunningham. Macmillan, London. 20. Cunningham, J. T. (1895) The origin of species among flat-fishes. Natural Science 6, 169–177, 234–239. 21. Lankester, E. R. (1896) The utility of specific characters. Nature 54, 365–366. 22. Buch, C. L. von (1825) Physikalische Beschreibung der Canarische Inseln Druckerei der Koniglichen Akademie der Wissenschaften, Berlin, Germany. 23. Gulick, J. T. (1905) Evolution, Racial and Habitudinal. Carnegie Institution of Washington. References and Notes 687

24. Romanes, G. J. (1895) Darwin, and After Darwin. 2. Post-Darwinian Ques- tions. Heredity and Utility. Longmans, Green & Co., London.

6 Reorientation and Controversy (1895–1899)

1. Poulton, E. B. (1890) The Colours of Animals: Their Meaning and Use, Especially Considered in the Case of Insects. Kegan Paul, Trench, Trubner & Co., London. 2. Pasteur, G. (1982) A classificatory review of mimicry systems. Annual Re- views of Ecology and Systematics 13, 169–199. 3. Weldon, W. F. R. (1895) Variation in animals and plants. Nature 51, 449–450. 4. Dyer, W. T. (1895) Variation and specific stability. Nature 51, 459–461. 5. Darwin, C. (1876) The Effects of Cross and Self Fertilization in the Vegetable Kingdom. John Murray, London. [Footnote: pp. 335] 6. Weldon, W. F. R. (1895) Attempt to measure the death rate due to the selec- tive destruction of Carcinus moenas with respect to a particular dimension. Proceedings of the Royal Society 57, 360–379. 7. Pearson, K. (1906) Walter Frank Raphael Weldon 1860-1906. Biometrika 5, 1–52. 8. Pearson, K. (1930) The Life, Letters and Labours of Francis Galton. Cam- bridge University Press, Cambridge. 9. Edelston, R. S. (1864) Amphidasis betularia. Entymologist 2, 150. 10. Tutt, J. W. (1896) British Moths. Routledge, London. 11. Doncaster, L. (1906) Collective inquiry as to progressive melanism in lepidoptera. The Etomologist’s Record 18, 1–16. 12. Majerus, M. E. N. (1998) Melanism: Evolution in Action. Oxford University Press, New York. 13. Pearson, K. (1901) On the principle of homotyposis and its relation to heredity, to the variability of the individual, and to that of the race. Part 1. Homotyposis in the vegetable kingdom. Philosophical Transactions of the Royal Society, A. 197, 285–379. 14. Davenport, C. B. (1899) Statistical Methods with Special Reference to Bio- logical Variation. Wiley, New York. 15. Pearson, K. (1902) On the fundamental conceptions in biology. Biometrika 1, 320–344.

7 What Life May Be 1. Lago, M. (1996) Christiana Herringham and the Edwardian Art Scene. Lund Humprhies, London. 2. Rothenstein, W. (1931, 1939). Men and Memories. Faber, London. 3. Blackman, F. F. (1926) The Eagle 44, 225–233. [St. John’s College magazine.] 4. Romanes, G. J. (1887) Mental differences between men and women. Nine- teenth Century 21, 654–672. 5. Richmond, M. L. (1997) A lab of one’s own. The Balfour biological laboratory for women at Cambridge University, 1884-1914. Isis 88, 422–455. 688 References and Notes

6. Durham, C. B. (1895) At a conversazione. The English Illustrated Maga- zine. September, pp. 547–552. 7. Bedford, H. A. R. & Pickering, P. S. U. (1919) Science and Fruit Growing. Macmillan, London. 8. Darby, R. (2005) The Surgical Temptation. The Demonization of the Fore- skin and the Rise of Circumcision in Britain. University of Chicago Press, Chicago, IL. 9. Cock, A. G. (1977) Bernard’s symposium. The species concept in 1900. Biological Journal of the Linnean Society 9, 1–30.

8 Rediscovery (1900–1901)

1. Olby, R. C. (1985) Origins of Mendelism. Second edition. University of Chicago Press, Chicago, IL. 2. Olby, R C. (1987) William Bateson’s introduction of Mendelism to England: A reassessment. British Journal of the History of Science 20, 399–420. 3. Olby, R. C. (2000) Horticulture: The font for the baptism of genetics. Nature Reviews Genetics 1, 65–70. 4. Vorzimmer, P. J. (1968) Darwin and Mendel: The historical connection. Isis 59, 77–82. 5. Naudin, C. (1856) Observations constatant le retour simultané de la descen- dance d’une plante hybride aux types paternels et maternels. Compte Rendu Academie Sciences, Paris 42, 628–636. 6. Romanes, G. J. (1881) Hybridism. Encyclopaedia Britannica. Ninth edition. 7. Romanes, E. (1896) The Life and Letters of George John Romanes. Longmans, Green & Co., London. 8. Romanes, G. J. (1894) Letter to Schafer, May 18. Welcome Museum of the History of Medicine, London. 9. Cock, A. G. (1973) William Bateson, Mendelism and biometry. Journal of the History of Biology 6, 1–36. 10. Weldon, W. F. R. (1902) Mendel’s laws of alternative inheritance in peas. Biometrika 1, 228–254. 11. Guyer, M. F. (1900) Spermatogenesis in hybrid pigeons. Science 21, 248–249, 312. 12. Hurst, R. (1974) The Evolution of Genetics. Unpublished manuscript in Cambridge University Library. 13. Lipset, D. (1980) Gregory Bateson. The Legacy of a Scientist. Prentice- Hall, Englewood Cliffs, NJ. 14. Keynes, G. (1981) The Gates of Memory. Clarendon, Oxford. 15. Wilmott, A. J. (1926) Obituary. William Bateson. The Journal of Botany 64, 78–80. 16. Lago, M. (1996) Christiana Herringham and the Edwardian Art Scene. Lund Humprhies, London. 17. Punnett, R. C. (1950) Early days of genetics. Heredity 4, 1–10.

References and Notes 689

9 Mendel’s Bulldog (1902–1906) 1. Mitchison, N. (1968) Beginnings. In: Haldane and Modern Biology. Edited by K. R. Dronamraju. Johns Hopkins University Press, Baltimore, MD. 2. Yule, G. U. (1906) On the theory of inheritance of quantitative compound characters on the basis of Mendel’s laws – a preliminary note. Report on the Third International Conference 1906 on Genetics. Edited by W. Wilks. Royal Horticultural Society, London, pp. 140–142. 3. Fisher, R. A. (1918) The correlation between relatives on the supposition of Mendelian inheritance. Transactions of the Royal Society of Edinburgh 52, 399–433. 4. Castle, W. E. (1903) The laws of heredity of Galton and Mendel, and some laws governing race improvement by selection. Proceedings of the American Academy of Arts and Sciences 39, 223–242. 5. Weldon, W. F. R. (1902) On the ambiguity of Mendel’s categories. Biometrika 2, 44–55. 6. Yule, G. U. (1902) Mendel’s laws and their probable relations to intra-racial heredity. The New Phytologist 1, 193–207, 222–238. 7. Ankeny, R. A. (2000) Marvelling at the marvel: The supposed conversion of A. D. Darbishire to Mendelism. Journal of the History of Biology 33, 315–347. 8. Weldon, W. F. R. (1903) Mr. Bateson’s revisions of Mendel’s theory of heredity. Biometrika 2, 286–298. 9. Garrod, A. E. (1902) Incidence of alkaptonuria: A study in chemical individuality. Lancet 2, 1616. 10. Bearn, A. G. (1994) Archibald Edward Garrod, the reluctant geneticist. Genetics 137, 1–4. 11. Gregory, R. P. (1903) The seed characters of Pisum sativum. The New Phytologist 2, 226–228. 12. Linnean Society (1904) Report of meeting, (Feb. 18). 13. Punnett, R. C. (1950) Early days of genetics. Heredity 4, 1–10. 14. Pearson, K. (1906) Walter Frank Raphael Weldon 1860-1906. Biometrika 5, 1–52. 15. Rushton, A. R. (2000) Nettleship, Pearson and Bateson: The biometric- Mendelian debate in a medical context. Journal of the History of Medicine and Allied Sciences 55, 134–157. 16. Lankester, E. R. (1906) Inaugural address by Professor E. Ray Lankester, President of the Association. Nature 74, 321–335. [Address to the BA at York.] 17. Schuster, E. H. J. (1906) Results of crossing grey (house) mice with albinos. Biometrika 4, 1–12. 18. Darbishire, A. D. (1905) The mutation theory of the origin of species. Nature 72, 314–316. [Review of Species and Varieties by de Vries (1905).] 19. Darbishire, A. D. (1906) On the difference between physiological and statistical laws of heredity. Memoirs and Proceedings of the Manchester Literary and Philosophical Society 50 (11), 1–44. [July 6]. 20. Schuster, E. H. J. (1913) Eugenics. Collins, London. 690 References and Notes

21. Lister, J. J. (1906) The life history of the Foraminifera. Nature 74, 400–404. [Address to the BA at York.] 22. Pearson, K. (1906) The latest critic of biometry. Nature 74, 465–466. 23. Lister, J. J. (1906) Biometry and biology: A reply to Professor Pearson. Nature 74, 584–585. 24. Pearl, R. (1911) Inheritance of fecundity in the domestic fowl. American Naturalist 45, 321–345. 25. Galton, F. (1901) On the probability that the son of a very highly gifted father will be no less gifted. Nature 65, 79. 26. Weldon, W. F. R. (1902) Professor de Vries on the origin of species. Biometrika 1, 365–374. 27. Mayr, E. (1982) The Growth of Biological Thought. Harvard University Press, Cambridge, MA. 28. Dawkins, R. (1986) The Blind Watchmaker. Longmans Scientific, Harlow, Essex, England. 29. Spillman, W. J. (1909) Application of some of the principles of heredity to plant breeding. Bulletin no. 165, Bureau of Plant Industry, US Department of Agriculture. 30. Anonymous (1906) Review by “N. R. C” of J. B. Burke’s The Origin of Life, (May 10). The Cambridge Review, Cambridge, pp. 375–376.

10 Bateson’s Bulldog 1. Haldane, J. B. S. (1926) The Nation and the Atheneum. Feb. 20, p. 713. 2. Spillman, W. J. (1901) Bulletin no. 115. Office of Experimental Stations, Washington, DC, pp. 88–98. 3. Hurst, C. C. (1903) Mendel’s principles applied to wheat hybrids. Journal of the Royal Horticultural Society 27, 876–893. 4. Davenport, C. B. (1901) Mendel’s law of dichotomy in hybrids. Biological Bulletin 2, 307–310. 5. Spillman, W. J. (1902) Exceptions to Mendel’s law. Science 16, 794–796. 6. Carlson, L. (2005) William J. Spillman: Scientific Agriculture in the Pro- gressive Era. University of Missouri Press, Columbia, MO. 7. Hurst, C. C. (1897) Curious crosses. The Orchid Review 5, 179–180. [This and other “curiousities of orchid breeding,” are in Hurst’s collected papers Experiments in Genetics. Cambridge University Press (1925), Cambridge, pp. 1–53.] 8. Hurst, C. C. (1900) Notes on some experiments in hybridization and cross- breeding. Journal of the Royal Horticultural Society 24, 90–126. 9. Hurst, C. C. (1904) Notes on Mendel’s methods of cross-breeding. Memoirs of the Horticultural Society of New York 1, 11–15. [Read at the Second International Conference on Plant Breeding and Hybridization, New York, Sept. 1902.] 10. Roberts, H. F. (1929) Plant Hybridization before Mendel. Princeton University Press, Princeton, NJ. References and Notes 691

11. Hurst, C. C. (1903) Mendel’s principles applied to orchid hybrids. Journal of the Royal Horticultural Society 27, 614–624. 12. Hurst, C. C. (1904) Mendel’s discoveries in heredity. Transactions of the Leicester Literary and Philosophical Society 8, 121–134. 13. Dawkins, R. (1976) The Selfish Gene. Oxford University Press, Oxford. 14. Castle, W. E. 1903. The heredity of Angora coat in mammals. Science 18, 760–761. 15. Hurst, C. C. (1905) Experimental studies on heredity in rabbits. Journal of the Linnean Society, Zoology 29, 283–324. 16. Hurst, C. C. (1905) Experimental studies on heredity in rabbits. Transac- tions of the Leicester Literary and Philosophical Society 9, 110–117. 17. Pearson, K. & Lee, A. (1900) Mathematical contributions to the theory of evolution, VIII. On the inheritance of characters not capable of exact quantitative measurement. Part 1. Introductory. Part 2. On the inheritance of coat colour in horses. Part 3. On the inheritance of eye colour in man. Philoso- phical Transactions of the Royal Society, A 195, 79–150. 18. Pearson, K., Yule, G. U., Blanchard, N., & Lee, A. (1903) Law of ancestral heredity. Biometrika 2, 214–215. 19. Hurst, R. (1974) The Evolution of Genetics. Unpublished manuscript in Cambridge University Library. [See also: Hurst, R. (1975) The Hurst collection of genetic letters now in the Cambridge University Library. Heredity 34, 279–280; (1977) Further note on the Hurst collection of genetical material in the Cambridge University Library. Heredity 38, 259–160.] 20. Hurst, C. C. (1906) On the inheritance of coat colour in horses. Proceedings of the Royal Society, B. 77, 388–394. 21. Punnett, R. C. (1950) Early days of genetics. Heredity 4, 1–10. 22. Weldon, W. F. R. (1906) Note on the offspring of thoroughbred chestnut mares. Proceedings of the Royal Society, B 77, 394–398. 23. Hurst, C. C. (1907) Mendelian characters in plants and animals. Report of the Third International Conference 1906 on Genetics. Edited by W. Wilks. Royal Horticultural Society, London, pp. 114–128. 24. Shull, G. H. (1909) The ‘presence and absence’ hypothesis. American Naturalist 43, 410–419. 25. Hurst, C. C. (1910) Mendel’s law of heredity and its application to horticulture. Journal of the Royal Horticultural Society 36, 22–52. 26. Shull, G. H. (1910) Germinal analysis through hybridization. Proceedings of the American Philosophical Society 49, 281–290. 27. Shull, G. H. (1915) Genetic definitions in the new standard dictionary. American Naturalist 49, 52–59. 28. Shull, G. H. (1911) Reversible sex mutants in Lychnis dioica. Botanical Gazette 52, 329–368. 29. Johannsen, W. (1911) The genotype conception of heredity. American Naturalist 45, 129–159. 30. Shull, G. H. (1912) ‘Genotypes,’ ‘Biotypes,’ ‘Pure Lines,’ and ‘Clones.’ Science 35, 27–29. 692 References and Notes

31. Wilson, E. B. (1914) Croonian Lecture: The bearing of cytological research on heredity. Proceedings of the Royal Society of London, B 88, 333–352. 32. Punnett, R. C. (1905) Mendelism. Macmillan & Bowes, Cambridge. 33. Darbishire, A. D. (1907) Nature 76, 73–74. [Anonymous review of second edition of reference 32]. 34. Hardy, G. H. (1908) Mendelian proportions in a mixed population. Science 28, 49–50. 35. Weinberg, W. (1908) Ueber den Nachweis der Vererbung beim Menchen. Jahreshefte des Vereins für Vaterländische Naturkunde in Württemberg 64, 368–382. 36. Lock, R. D. (1906) Recent Progress in the Study of Variation, Heredity and Evolution. John Murray, London. 37. Carlson, E. A. (1966) The Gene: A Critical History. W. B. Saunders, Phila- delphia, PA.

11 On Course (1907–1908) 1. Weinstein, A. (1980) A note on W. L. Tower’s Leptinotarsa work. In: The Evolutionary Synthesis. Edited by E. Mayr & W. Provine. Harvard Univer- sity Press, Cambridge, MA. 2. Hurst, C. C. (1908) On the inheritance of eye-colour in man. Proceedings of the Royal Society, B 80, 85–96. 3. Davenport, C. B. (1907) Heredity of eye colour in man. Science 26, 589–592. 4. Report concerning a Chair in Genetics. (1908) Cambridge University Reporter, March 3, p. 632. 5. Olby, R. (1989) Scientists and bureaucrats in the establishment of the John Innes Horticultural Institution under William Bateson. Annals of Science 46, 497–510. 6. Pearson, K. (1930) The Life, Labour, and Letters of Francis Galton, Vol. 3. Cambridge University Press, Cambridge, pp. 340. 7. Wallace, A. R. (1908) The present position of Darwinism. Contemporary Review 94, 129–141. 8. Poulton, E. B. (1908) Essays on Evolution, 1889-1907. Clarendon, Oxford. 9. Punnett, R. C. (1908) Old bottles. The New Quarterly, October. 10. Rothenstein, W. (1939) Men and Memories, Vol. 2. Faber, London. 11. Morgan, T. H. (1909) What are “factors” in Mendelian explanations? American Breeders Association Reports 5, 365–368. 12. (1908) The influence of heredity on disease, with special reference to tuber- culosis, cancer and diseases of the nervous system. Royal Society of Medi- cine General Report, pp. 9–142.

12 Darwin Centenary (1909) 1. Haldane, J. B. S. (1957) The theory of evolution, before and after Bateson. Journal of Genetics 56, 11–27. References and Notes 693

2. Nogler, G. A. (2006) The lesser known Mendel: His experiments with Hieracium. Genetics 172, 1–6. 3. Bateson. W. (1909) Boyle lecture. [There is no indication that this was formally published. The manuscript has not been found.] 4. Richmond, M. (2006) The 1909 Darwin celebration. Reexamining evolution in the light of Mendel, mutation and meiosis. Isis 97, 447–484. 5. Rothenstein, W. (1931) Men and Memories, Vol. 1. Faber, London. 6. Meldola, R. (1909) Evolution: Old and new. Nature 80, 481–485. 7. Gulick, J. T. (1872) On the diversity of evolution under one set of external conditions. Journal of the Linnean Society, Zoology 11, 496–505. 8. Gould, S. J. & Lewontin, R. C. (1979) The spandrels of San Marco and the Panglossian paradigm: A critique of the adaptionist program. Proceedings of the Royal Society of London. B. 205, 581–598. 9. Forsdyke, D. R. (2001) The Origin of Species, Revisited. McGill-Queen’s University Press, Montreal, Quebec, Canada. 10. Orr, H. A. (1996) Dobzhansky, Bateson and the genetics of speciation. Genetics 144, 1331–1335. 11. Shipley, A. E. (1910) Report of the British Association for the Advancement of Science 1909. Murray, London, pp. 484–502. 12. Haldane, J. B. S., Sprunt. A. D., & Haldane. N. M. (1915) Reduplication in mice. Preliminary Communication. Journal of Genetics 5, 133–135. 13. Haldane, J. B. S. (1926) The Nation and the Atheneum. Feb. 20, p. 713. [Obituary of William Bateson.]

13 Chromosomes

1. Boveri, T. H. (1888) Zellen-Studien II. Die Befruchtung und Teilung des Eies von Ascaris megalocephala. Jenaischen Zeitschrift für Medicin und Naturwissenschaft 22, 685–882. 2. Boveri, T. H. (1889) Translated by T. H. Morgan (1893) An organism produced sexually without characteristics of the mother. American Naturalist 27, 222–232. 3. Roux, W. (1883) Uber die Bedeutung der Kerntheilungsfiguren: Eine hypothetische Erörterung. Wilhelm Engelmann, Leipzig. 4. Stern, C. (1950) Boveri and the early days of genetics. Nature 166, 446. 5. Cremer, T. & Cremer, C. (1988) Centennial of Wilhelm Waldeyer’s introduction of the term “chromosome” in 1888. Cytogenetics and Cell Genetics 48, 65–67. 6. Farmer, J. B. & Moore, J. E. S. (1905) On the meiotic phase (reduction division) in animals and plants. Quarterly Journal of Microscopical Science 48, 489–557. 7. Weismann, A. (1887) On the significance of the polar globules. Nature 36, 607–609. 8. Mitchell, P. C. (1892) The new volume of Weismann. Nature 46, 558–559. 9. Romanes, G. J. (1893) An Examination of Weismannism. Open Court Pub- lishing, Chicago, IL. 694 References and Notes

10. Wilson, E. B. (1900) The Cell in Development and Inheritance. Second edition. Macmillan, New York. 11. Cannon, W. A. (1904) Some cytological aspects of hybrids. Memoires of the Horticultural Society of New York 1, 89–92. [Read at the International Conference on Plant Breeding and Hybridization, New York, September, 1902.] 12. Sutton, W. S. (1900) The spermatogonial divisions in Brachystola magna. Kansas University Quarterly 9, 135–160. 13. Sutton, W. S. (1902) On the morphology of the chromosome group in Brachystola magna. Biological Bulletin 4, 24–39. 14. Cannon, W. A. (1902) A cytological basis for Mendelian Laws. Bulletin of the Torrey Botanical Club 29, 657–661. 15. Sutton, W. S. (1903) The chromosomes in heredity. Biological Bulletin 4, 231–251. 16. McCusick, V. A. (1960) Walter Sutton and the physical basis of Mendelism. Bulletin of the History of Medicine 34, 487–497. 17. Vries, H. de (1910) Fertilization and hybridization. In the English translation of Intracellular Pangenesis (1910) Open Court Publishing, Chicago, IL, pp. 217–263. 18. Boveri, T. H. (1904) Ergebnisse über die Konstitution der chromatischen Substanz des Zellkerns. Gustav Fischer, Jena. 19. Montgomery, T. H. (1901) A study of the germ cells of the metazoa. Trans- actions of the American Philosophical Society 20, 154–236. 20. Paulmier, F. C. (1899) The Spermatogenesis of Anasa tristis. Ginn & Co., Boston, MA. 21. McClung, C. E. (1899) A peculiar nuclear element in the male reproductive cells of insects. Zoological Bulletin 2, 187. 22. Stevens, N. M. (1905) Studies in spermatogenesis with especial reference to the “accessory chromosome.” Carnegie Institution of Washington, 36, 1–32. 23. Boveri, T. H. (1901) Das Problem der Befruchtung. Gustav Fischer, Jena. 24. Boveri, T. H. (1902) Über mehrpolige Mitosen als Mittel zur Analyse des Zellkerns. Verhandlungen der Physicalisch-Medizinischen Gesselschaft zu Würzburg. Neu Folge 35, 67–90. 25. Herringham, W. P. (1888) Muscular atrophy of the peroneal type affecting many members of a family. Brain 11, 230–236. 26. Doncaster, L. & Raynor, G. H. (1906) Breeding experiments with lepidoptera. Proceedings of the Zoological Society London 1, 125–133. 27. McKusick, V. A. (1962) On the X chromosome of man. Quarterly Review of Biology 37, 69–175. 28. Wilson, E. B. (1909) Recent researches on the determination and heredity of sex. Science 29, 53–70. 29. Smith, G. (1906) Rhizocephala. Fauna und Flora des Golfes Von Neapel 29, 89. [Published by R. Friedlander & Sohn, Berlin.] 30. Hurst, C. C. (1909) Mendelism and sex. In: Experiments in Genetics. Cambridge University Press (1925), Cambridge, pp. 331–350. 31. Gregory, R. P. (1904) The reduction division in ferns. Proceedings of the Royal Society 73, 86–92. References and Notes 695

32. Guyer, M. F. (1900) Spermatogenesis in hybrid pigeons. Science 11, 248–249, 312. 33. Guyer, M. F. (1899) Ovarian structure in an abnormal pigeon. Zoological Bulletin 2, 211–224. 34. Guyer, M. F. (1902) Hybridism and the germ cell. Series 2, Vol. 2. Bulletin no. 21 of the University of Cincinnati, OH [Based on Guyer’s thesis (1900) Spermatogenesis in Normal and Hybrid pigeons. University of Chicago, Chicago, IL.] 35. Bungener, P. & Buscaglia, M. (2003) Early connection between cytology and Mendelism: Michael F. Guyer’s contribution. History and Philosophy in the Life Sciences 25, 27–50. 36. Guyer, M. F. (1903) The germ cell and the results of Mendel. The Cincin- nati Lancet Clinic 53, 490–491. 37. Guyer, M. F. (1907) Do offspring inherit equally from each parent? Science 25, 1006–1010. 38. Guyer, M. F. (1911) Nucleus and cytoplasm in heredity. American Naturalist 45, 284–305. 39. Guyer, M. F. (1909) Deficiencies of the chromosome theory of heredity. Series 2, Vol. 5, no. 3. University Press, Burnet Woods, Cincinnati, OH, pp. 1–19. [Report of paper presented in Boston in 1907.] 40. Godlewski, E. (1906) Die hybridisation der Echiniden und Crinoiden familie. Archiv für EntwicklungsMechanik der Organismen 20, 579. 41. Spillman, W. J. (1910) Heredity. American Naturalist 44, 504–512. 42. Whitman, C. O. (1909) cited in Riddle, O. (1919) Orthogenic Evolution in Pigeons. Posthumous Works of C. O. Whitman. Carnegie Institution of Washington. 43. Hagemann, R. (2000) Erwin Baur or Carl Correns: Who really created the theory of plastid inheritance? Journal of Heredity 91, 435–440. 44. Fangerau, H. & Müller, I. (2007) Scientific exchange: Jaques Loeb and Emil Godlewski as representatives of a transatlantic developmental biology. Studies in the History and Philosophy of Biology and Biomedical Sciences 38, 608–617. 45. Johannsen, W. (1923) Some remarks on units in heredity. Hereditas 4, 133–141. 46. Forsdyke, D. R. (2006) Evolutionary Bioinformatics. Springer, New York. 47. Shull, G. H. (1909) The ‘presence and absence’ hypothesis. American Natu- ralist 43, 410–419. 48. Moore, A. R. (1910) A biochemical conception of dominance. University of California Publications in Physiology 4, 9–15. 49. Forsdyke, D. R. (1994) The heat-shock response and the molecular basis of genetic dominance. Journal of Theoretical Biology 167, 1–6. 50. Galton, F. (1897) The average contribution of several ancestors to the total heritage of the offspring. Proceedings of the Royal Society 61, 401–413. 51. Pearson, K. (1906) Walter Frank Raphael Weldon 1860-1906. Biometrika 5, 1–52. 696 References and Notes

52. Castle, W. (1906) Effects of inbreeding, cross-breeding and selection on the fertility and variability of Drosophila. Proceedings of the American Acad- emy of Arts and Sciences 41, 731. 53. Davenport, C. B. (1941) The early history of research with Drosophila. Science 93, 305–306. 54. Morgan, T. H. (1910) Chromosomes and heredity. American Naturalist 44, 449–496. 55. Morgan, T. H. (1909) For Darwin. Popular Science Monthly 70, 367–380. 56. Morgan, T. H. (1911) Random segregation versus coupling in Mendelian inheritance. Science 34, 384. 57. Haldane, J. B. S. (1919) The combination of linkage values, and the calculation of distance between linked factors. Journal of Genetics 8, 299–309. 58. Bridges, C. B. (1914) The chromosome hypothesis of linkage applied to cases of sweet peas and primula. American Naturalist 48, 524–534. 59. Sturtevant, A. H. (1914) The reduplication hypothesis as applied to Droso- phila. American Naturalist 48, 535–549. 60. Muller, H. J. (1914) A gene for the fourth chromosome of Drosophila. Journal of Experimental Zoology 17, 325–336. 61. Sturtevant, A. H. (1915) The behaviour of the chromosomes as studied through linkage. Zeitschrift für induktiv Abstammungs- und Vererbungslehre 13, 234–287. 62. Punnett R. C. (1923) Linkage in the sweet pea. Journal of Genetics 13: 101–123. 14 Passages (1910–1914) 1. Butler, S. (1912) The Notebooks of Samuel Butler. Edited by H. F. Jones. Fifield, London. 2. Olby, R. (1989) Scientists and bureaucrats in the establishment of the John Innes Horticultural Institution under William Bateson. Annals of Science 46, 497–510. 3. Rothenstein, W. (1939) Men and Memories, Vol. 2. Faber, London. 4. Lago, M. (1996) Christiana Herringham and the Edwardian Art Scene. Lund Humprhies, London. 5. Anonymous (1910) Our Portrait. The Gownsman, Cambridge, May 5. 6. Crew, F. A. E. (1967) Reginald Punnett. Biographical Memoirs of Fellows of the Royal Society 13, 309–326. 7. Lipset, D. (1980) Gregory Bateson. The Legacy of a Scientist. Prentice-Hall, Englewood Cliffs, NJ, p. 55. 8. Balls, W. L. (1912) The Cotton Plant in Egypt. Macmillan, London. 9. Olby, R. (1991) Social imperialism and state support for agricultural research in Edwardian Britain. Annals of Science, 48, 509–526. 10. Hurst, C. C. (1911) The government grant for horse-breeding: The question of eugenics. The Times, March 4. 11. Hurst, C. C. (1911) The application of genetics to horse-breeding. In: Experiments in Genetics. Cambridge University Press (1925), Cambridge. References and Notes 697

12. Hurst, C. C. (1912) Mendelian experiments with thoroughbred horses. Bloodstock Breeders’ Review 1, 86–90. 13. Wilson, J. (1914) Unsound Mendelism developments, especially as regards the presence and absence theory. Scientific Proceedings of the Royal Dublin Society 8, 399–421. 14. Cunningham, J. T. (1914) Biological Agnosticism. [This is a newspaper cutting the source of which has not been identified.] 15. Harwood, J. (1993) Styles of Scientific Thought. The German Genetics Community 1900-1933. University of Chicago Press, Chicago, IL. 16. Gates, R. R. (1915) The Mutation Factor in Evolution. Macmillan, London. 17. Goldschmidt, R. (1955) Theoretical Genetics. University of California Press, p. 478. 18. Winge, Ö. (1917) The chromosomes. Their numbers and general importance. Comptes Rendus des Travaux du Laboratoire Carlsberg 13, 131–275. 19. Smolin, L. (2007) The other Einstein. The New York Review of Books 54 (10), 76–83.

15 Eugenics 1. Galton, F. (1865) Hereditary talent and character. Macmillan’s Magazine 12, 157–166, 318–327. 2. Galton, F. (1873) On the causes which operate to create scientific men. Fortnightly Review 13, 345–351. 3. Fancher, J. (1979) A note on the origin of the term “nature and nurture” Journal of the History of Behavioural Sciences 15, 321–322. 4. Pearson, K. (1903) Inheritance of psychical and physical characters of man. Nature 68, 607–608. [Abstract of his Huxley lecture (Oct. 16).] 5. Gilham, N. W. (2001) A Life of Sir Francis Galton: From African Explora- tion to the Birth of Eugenics. Oxford University Press, Oxford. 6. Harper, P. S. (2005) Julia Bell and the treasury of human inheritance. Human Genetics 116, 422–432. 7. Lipset, D. (1980) Gregory Bateson. The Legacy of a Scientist. Prentice-Hall, Englewood Cliffs, NJ. 8. Reid, G. A. (1907) The interpretation of Mendelian phenomena. Nature 76, 566. 9. Lawrence, W. (1823) Lectures on Physiology, Zoology and the Natural History of Man. Third edition, London. 10. Hurst, C. C. (1908) Mendel’s law of heredity and its application to man. Transactions of the Leicester Literary and Philosophical Society 12, 35–48. 11. Hurst, C. C. (1912) Mendelian heredity in man. The Eugenics Review 4, 1–25. 12. Fisher, R. A. (1911) Address on “Heredity” recorded in the minute book of the Cambridge Eugenics Society meeting held at Trinity College on 10th November 1911. In: Norton, B. & Pearson, E. S. (1976) Notes and Records of the Royal Society of London 31, 151–162. [A note on the background to, and refereeing of, R. A. Fisher’s 1918 paper ‘On the correlation between relatives on the supposition of Mendelian inheritance’.] 698 References and Notes

13. Darwin, L. (1912) The first international eugenics congress. Nature 89, 558–561. 14. Punnett, R. C. (1917) Eliminating feeblemindedness. Journal of Heredity 8, 464–465. 15. Pilkington, D. (1996) Follow the Rabbit-Proof Fence. University of Queen- sland Press, Saint Lucia, QLD, Australia. 16. Baur, E., Fischer, E. & Lenz, F. (1921) Grundriss der menschlichen Erblichkeitslehre und Rassenhygiene. J. F. Lehmann, Munich.

16 War (1915–1919) 1. Butler, S. (1912) The Notebooks of Samuel Butler. Edited by H. F. Jones. Fifield, London. 2. Cock, A. G. (1983) Chauvinism and internationalism in science: The international research council, 1919-1926. Notes and Records of the Royal Soci- ety of London 37, 249–288. 3. Haldane, J. B. S. (1925) Callinicus: A Defence of Chemical Warfare. Kegan Paul, London. 4. Smith, N. (1916) The place of natural science in secondary education. The Times Educational Supplement. (May 6). 5. Morgan, T. H., Sturtevant, A. H., Muller, H. J., & Bridges, C. B. (1915) The Mechanism of Mendelian Heredity. Henry Holt & Co., New York. 6. Bridges, C. B. (1916) Non-disjunction as proof of the chromosome theory of heredity. Genetics 1, 1–52, 107–163. 7. Lotsy, J. P. (1916) Evolution by Means of Hybridisation. Nijhoff, The Hague. 8. Brink, R. A. (1958) Paramutation at the R locus in maize. Cold Spring Harbor Symposium on Quantitative Biology 23, 379–391. 9. Stam, M. & Mittelsten Scheid, O. (2005) Paramutation: An encounter leaving a lasting impression. Trends in Plant Science 10, 283–290. 10. Gregory, R. P. (1914) On the genetics of tetraploid plants in Primula sinensis. Proceedings of the Royal Society, B. 87, 484–492. 11. Wilson, E. B. (1914) The bearing of cytological research on heredity. Proceedings of the Royal Society, B. 88, 333–352. 12. Winge, Ö. (1917) The chromosomes. Their numbers and general importance. Comptes Rendus des Travaux du Laboratoire Carlsberg 13, 131–275. 13. Rosenberg, O. (1907) Cytological investigations of plant hybrids. Report of the Third International Conference 1906 on Genetics. Edited by W. Wilks. Royal Horticultural Society, London. pp. 289–291. 14. Herringham, W. P. (1919) A Physician in France. Edward Arnold, London.

17 My Respectful Homage (1920–1922) 1. Haldane, J. B. S. (1919) The combination of linkage values and the calculation of distances between the loci of linked factors. Journal of Genetics 8, 299–309. References and Notes 699

2. Forsdyke, D. R. (2000) Haldane’s rule: Hybrid sterility affects the heterogametic sex first because sexual differentiation is on the path to species differentiation. Journal of Theoretical Biology 204, 443–452. 3. Provine, W. (1971) The Origins of Theoretical Population Genetics. The University of Chicago Press, Chicago, IL. 4. Haldane, J. B. S. (1924) A mathematical theory of natural and artificial selection. Transactions of the Cambridge Philosophical Society 23, 19–41. 5. Wilson, E. B. & Morgan, T. H. (1920) Chiasmatype and crossing over. American Naturalist 54, 193–219. 6. Johannsen, W. (1923) Some remarks about units of heredity. Hereditas 4, 133–141. 7. Muller, H. J. (1934) Lenin’s doctrines in relation to genetics. In: Graham, L. R. (1972) Science and Philosophy in the Soviet Union. A. Knopf, New York, pp. 451–469. 8. Darlington, C. D. (1979) Morgan’s crisis. Nature 278, 786–787. 9. Provine, W. B. (1986) Sewall Wright and Evolutionary Biology. The University of Chicago Press, Chicago, IL, p. 104. 10. Cock, A. G. (1989) Bateson’s two Toronto addresses, 1921. 1. Chromosomal skepticism. Journal of Heredity 80, 91–95. 11. Cock, A.G. (1989) Bateson’s two Toronto addresses. 1921, 2. Evolutionary faith. Journal of Heredity 80, 96–99. 12. Osborn, H. F. (1922) William Bateson on Darwinism. Science 55, 194–197. 13. Mitchell, P. C. (1922) Unfashionable darwinism: No alternative solution. The Times, April 11. 14. Mitchell, P. C. (1922) Darwin and evolution. The Times. April 15. 15. Cunningham, J. T. (1922) Species and adaptations. Nature 109, 775–777. 16. Crowther, C. R. (1922) Evolutionary faith and modern doubts. Nature 109, 777. 17. Cock, A. G. (1983) William Bateson’s rejection and eventual acceptance of chromosome theory. Annals of Science 40, 19–59. 18. Harman, O. S. (2004) The Man Who Invented the Chromosome. A Life of Cyril Darlington. Harvard University Press, Cambridge, MA. 19. Forsdyke, D. R. (2001) The Origin of Species, Revisited. McGill-Queen’s University Press, Montreal, Quebec, Canada. 20. Dobell, C. (1925) The chromosome cycle of the sporozoa considered in relation to the chromosome theory of heredity. La Cellule 25, 167–192. 21. Harwood, J. (1993) Styles of Scientific Thought. The German Genetics Community 1900-1933. University of Chicago Press, Chicago, IL. 22. Johannsen, W. (1922) Hundert Jahre Vererbungsforschung. Verhandlungen der Gesellschaft deutscher Naturforscher und Arzte 87, 70–104. 23. Russell, E. S. (1930) The Interpretation of Development and Heredity. Clarendon, Oxford. 24. Punnett, R. C. (1922) Nature 110, 85–86. [Obituary of Onslow.]

700 References and Notes

18 Limits Undetermined (1923–1926) 1. Haldane, J. B. S. (1926) The Nation and Atheneum. Feb. 20, p. 713. 2. Hurst C. C. (1921) The genetics of egg-production in white leghorns and white wyandottes. The National Poultry Journal. (Sept. 2 and Dec. 9). 3. Hurst C. C. (1923) The genetics of fecundity in the domestic hen. In: Eugenics, Genetics and the Family. Williams & Wilkins, Baltimore, MD, pp. 212–217. 4. Hurst, R. (1974) The Evolution of Genetics. Unpublished manuscript in Cambridge University Library. 5. Hurst, C. C. (1925) Chromosomes and characters in Rosa and their significance in the origin of species. In: Experiments in Genetics, Cambridge University Press, Cambridge, pp. 534–550. 6. Williams, G. C. (1966) Adaptation and Natural Selection. Princeton University Press, Princeton, NJ. 7. Forsdyke, D. R. (2006) Evolutionary Bioinformatics. Springer, New York. 8. Roll-Hansen, N. (2005) The Lysenko effect: Undermining the autonomy of science. Endeavour 29, 144–147.

19 Butler 1. Butler S. (1887) Luck or Cunning as the Main Means of Organic Modifica- tion? Trübner, London. 2. Forsdyke, D. R. (2006) Heredity as transmission of information: Butlerian “intelligent design.” Centaurus 48, 133–148. [see also: Forsdyke, D. R. (2008) Samuel Butler and long-term memory. Is the cupboard bare? Centaurus (submitted for publication).] 3. Forsdyke, D. R. (2006) Evolutionary Bioinformatics. Springer, New York. 4. Darwin C. (1867) Letter to Canon Farrar, March 5. In: More Letters of Charles Darwin, Vol. 2. Edited by F. Darwin (1903). Appleton, New York, pp. 441–442. [Comments on a lecture the Canon had given at the Royal Institution “On Some Defects in Public School Education.”] 5. Butler, S. (1896) The Life and Letters of Dr. Samuel Butler, Vol. 1. John Murray, London, p. 353, pp. 376–391. 6. Butler, S. (1903) The Way of All Flesh. Grant Richards, London. 7. Heitland, W. E. (1926) After Many Years. Cambridge University Press, Cambridge. 8. Jones, H. F. (1919) Samuel Butler, Author of Erewhon (1835-1902): A Memoir, Vols. 1 and 2. Macmillan, London. 9. Bowlby, J. (1990) Charles Darwin. A New Life. Norton, New York. 10. Iltis, H. (1932) Life of Mendel. Allen & Unwin, London. 11. Orel, V. (1984) Mendel. Translated by S. Finn. Oxford University Press, Oxford. 12. Butler, S. (1914) A First Year in Canterbury Settlement with Other Earlier Essays. Edited by R. A. Streatfeild. Fifield, London, pp. 149–185. 13. Butler, S. (1872) Erewhon or Over the Range. Trübner, London. 14. Butler, S. (1880) Unconscious Memory. David Bogue, London. 15. Romanes, G. J. (1881) Unconscious memory. Nature 23, 285–287. References and Notes 701

16. Butler, S. (1914) The Humour of Homer and Other Essays. Kennerley, New York. 17. Forsdyke, D. R. (2001) The Origin of Species, Revisited. McGill-Queen’s University Press. 18. Butler, S. (1878) Life and Habit. Trübner, London. [“We are such stuff as dreams are made on,” are the words of Prospero in Shakespeare’s The Tempest.] 19. Hering E. (1870) Über das Gedachtniss als eine allgemeine Function der organisirten Materie. Karl Gerold’s Sohn, Vienna. [“On memory as a universal function of organized matter.” A lecture delivered at the anniversary meeting of the Imperial Academy of Sciences at Vienna. Butler learned German in order to understand Hering, and there is a translation in Butler’s Unconscious Memory.] 20. Butler, S. (1912) The Notebooks of Samuel Butler. Edited by H. F. Jones. Fifield, London. 21. Mivart, St. G. J. (1871) On the Genesis of Species. Macmillan, London. 22. Galton, F. (1875) A theory of heredity. Contemporary Review 27, 80–95. 23. Forsdyke, D. R. & Ford, P. M. (1983) Segregation into separate rouleaux of erythrocytes from different species: Evidence against the agglomerin hypothesis of rouleaux formation. Biochemical Journal 214, 257–260. 24. Haeckel, E. (1876) Die Perigenesis der Plastidule oder die Wellenzeugung der Lebenstheilchen. Reimer, Berlin, Germany. 25. Darwin, C. (1876) Letter to Romanes, May 29. In: More Letters of Charles Darwin, Vol. 1. Edited by F. Darwin (1903). Appleton, New York, pp. 363–365. 26. Lankester, E. R. (1876) Perigenesis versus pangenesis – Haeckel’s new theory of heredity. Nature 14, 235–238. 27. Clodd, E. (1916) Memories. Chapman & Hall, London. 28. Brücke, E. (1861) Die Elementarorganismen. Sitzungsberichte der Akademie der Wissenschaften Wein, Mathematische-wissenschaftliche Classe, 44, 381–406. 29. Butler, S. (1879) Evolution, Old and New. Hardwicke & Bogue, London. 30. Lamarck, J. B. (1809) Philosophie Zoologique. Translated as Zoological Philosophy. University of Chicago Press, Chicago, IL, 1984. 31. Butler, S. (1884) Selections from Previous Works, with Remarks on Mr. G. J. Romanes’ “Mental Evolution in Animals,” and a Psalm of Montreal. Trübner, London. 32. Bernstein, C. & Bernstein, H. (1991) Aging, Sex and DNA Repair. Academic, San Diego, CA. 33. Romanes, E. (1896) The Life and Letters of George John Romanes. Longmans, Green & Co., London, p. 132, 153. 34. Haeckel, E. (1909) Charles Darwin as an anthropologist. Darwin and Mod- ern Science. Cambridge University Press, Cambridge. 35. Waterhouse, P. M., Wang, M.-B. & Lough, T. (2001) Gene silencing as an adaptive defence against viruses. Nature 411, 834–841. 36. Wallace, A. R. (1879) Organization and intelligence. Nature 19, 477–480. 37. Dawkins, R. (1976) The Selfish Gene. Oxford University Press, Oxford. 702 References and Notes

38. Anonymous (1879) Review of Evolution, Old and New. The Atheneum, London. July 26, pp. 115–117. 39. Romanes G J. (1884) Mental Evolution in Animals, with a Posthumous Essay on Instinct by Charles Darwin. Appleton, New York. 40. Anonymous (1884) Review of Mental Evolution in Animals. The Atheneum, London. Mar. 1, pp. 282–283. 41. Anonymous (1884) Review of Selections from Previous Works. The Atheneum, London. Mar. 23, pp. 378–379. 42. Salisbury, L. (1994) Inaugural address of the Most Hon. the Marquis of Salisbury, K. G., D. C. L., F. R. S., Chancellor of the University of Oxford, President. Nature 50, 339–343. [Salisbury (Robert Cecil) ceded his Prime Ministership to his nephew, Arthur Balfour, in 1902; in other words, “Bob’s your uncle.”] 43. Darwin, C. (1958) The Autobiography of Charles Darwin 1809-1882. The First Complete Version. Edited by N. Barlow. Collins, London. 44. Keynes, G. & Hill, B. (1935) Letters between Samuel Butler and Miss E. M. A. Savage. Jonathan Cape, London. [These editors also facilitated the production of Bartholomew’s Further Extracts from the Note-Books of Samuel Butler (1934).] 45. Darwin, F. (1878) The analogies of plant and animal life. Nature 17, 388–391, 411–414. 46. Darwin, F. & Pertz, D. F. M. (1892) On the artificial production of rhythm in plants. Annals of Botany 6, 245–264. 47. Hartog, M. (1892) Problems of reproduction: Conjugation, fertilization and rejuvenescence. Contemporary Review 62, 92–104. 48. Hartog, M. (1897) Fundamental principles of heredity. Natural Science 11, 233–239, 305–316. 49. Hartog, M. (1914) Samuel Butler and recent mnemic biological theories. Scientia 15, 38–52. [An attack on Richard Semon who committed suicide in 1918.] 50. Russell, E. S. (1916) Form and Function. John Murray, London, pp. 335– 344. 51. Dawkins, R. (2003) The Ancestor’s Tale. Weidenfeld & Nicolson, London. [Dawkins was unaware of Semon’s word “mneme” when he suggested the word “meme” as a self-replicating element of culture, passed on by imitation.] 52. Jones, H. F. (1911) Charles Darwin and Samuel Butler. A Step Towards Reconciliation. Fifield, London. 53. Galton, F. (1910) Kantsaywhere. Unpublished novel. Galton Archives, University College, London. 54. Bateson, M. C. (1984) With a Daughter’s Eye. A Memoir of Margaret Mead and Gregory Bateson. William Morrow, New York. 55. Bateson, G. (1962) The role of somatic change in evolution. Evolution 17, 529–539. 56. Bateson, G. (1971) A re-examination of Bateson’s rule. Journal of Genetics 60, 230–240. 57. Bateson, G. (1979) Mind and Nature. A Necessary Unity. Dutton, New York. References and Notes 703

58. Bateson, G. & Bateson, M. C. (1987) Angels Fear. Towards an Epistemol- ogy of the Sacred. Macmillan, New York. 59. Cock, A. C. (1975) Taped interview with Gregory Bateson. Queen’s University Archives, Kingston. 60. Bateson, G. (1970) The Alfred Korzybski memorial lecture 1970. General Sematics Bulletin 37. [A lecture given at the Harvard Club of New York, 9th January.]

20 Pilgrimages 1. Cock, A. G. (1980) William Bateson’s pilgrimages to Brno. Folia Mende- liana 15, 243–250. [Volume 65 in the series Acta Musei Moraviae.] 2. Cock, A. G. (1982) Bateson’s impression at the unveiling of the Mendel monument at Brno in 1910. Folia Mendeliana 17, 217–223. [Volume 67 in the series Acta Musei Moraviae.] 3. Hurst, C. C. (1911) Mendelian characters in plants, animals and man. In: “Festschrift zum Andenken an Gregor Mendel,” Verhandlungen des Natur- forshenden Vereines in Brünn 49, 192–213. 4. (1925) Memorial Volume in Honor of the 100th Birthday of J. G. Mendel. Fr. Borovy, Prague. 5. Correns, C. (1924) Carl Correns. Julius Springer, Berlin, Germany. [Collected papers.] 6. Iltis, H. (1924) Gregor Johann Mendel, Leben, Werk und Wirkung. Julius Springer, Berlin, Germany. 7. Nivet, C. (2004) Une maladie énigmatique dans la vie de Gregor Mendel. Médecine Sciences 20, 1050–1053; (2006) 1848: Gregor Mendel, le moine qui voulait être citoyen. Médecine Sciences 22, 430–433. 8. Orel, V. (1965) Editorial. Folia Mendeliana, Vol. 1.

21 The Kammerer Affair 1. Lamarck, J. B. (1809) Philosophie Zoologique. Translated as Zoological Philosophy. University of Chicago Press, Chicago, IL, 1984. 2. Noble, G. K. (1926) Kammerer’s Alytes. Nature 118, 209–210. 3. Koestler, A. (1971) The Case of the Midwife Toad. Hutchinson, London. 4. Gould, S. J. (1972) Zealous advocates. Science 176, 623–625. 5. Bateman, K. G. (1959) The genetic assimilation of four venation pheno- copies. Journal of Genetics 56, 443–474. 6. Cannon, H. G. (1959) Lamarck and Modern Genetics. Charles C Thomas, Springfield, IL. 7. Aronson, L. R. (1975) The case of The Case of the Midwife Toad. Behavior Genetics 5, 115–125. 8. Gliboff, S. (2005) ‘Protoplasm … is soft wax in our hands’: Paul Kammerer and the art of biological transformation. Endeavour 29, 162–167.

704 References and Notes

22 Science and Chauvinism 1. Nuttall, G. H. F. (1901) On the question of priority with regard to certain discoveries upon the aetiology of malarial diseases. Quarterly Journal of Microscopical Science 44, 429–441. 2. Sleigh, M. A. & Sutcliff, J. F. (1966) The Origins and History of the Society for Experimental Biology. [A 32 page booklet printed for the Society for Experimental Biology. Tollfree Ltd., London.] 3. Cock, A. G. (1979) The Genetical Society in 1924 – a near demise. Heredity 42, 113–117. 4. Cock, A. G. (1983) Chauvinism and Internationalism in science. Notes and Records of the Royal Society of London 37, 249–288. 5. Larmor, J. (1917) Papers in Royal Society Collection. Letter from H. E. Armstrong to Larmor. (July 8). 6. Hampson, G. (1918) A classification of the Pyralidae. Proceedings of the Zoological Society, pp. 55–131. 7. Jones, H. S. (1960) The early history of the ICSU. ICSU Review 2, 169–187.

23 Degrees for Women 1. Punnett, R. C. (1926) William Bateson. Edinburgh Review 244, 71–86. 2. Sidgwick, E. (1897) University Education for Women. Macmillan & Bowes, Cambridge. 3. Gardner, A. (1921) A Short History of Newnham College. Bowes & Bowes, Cambridge. [See also Newnham College Register.] 4. Cock, A.G. (1979) Anna Bateson of Bashley – Britain’s first professional woman gardener. Hampshire 19, 59–62. 5. Bateson, W. (1897) Cambridge University Reporter. March 26, pp. 797–799. 6. This chapter was one of the first completed by Alan Cock on Bateson since “it was relatively easy to write up in isolation.” In a letter to Darlington (Mar. 8, 1977) Alan said that it had been sent, as a sample chapter, to a potential publisher (Cambridge University Press). Having seen an early draft, Gregory Bateson commented (Sept. 3, 1974): “You seem unaware that WB almost totally reversed his position on femi- nism at the end of his life. He became aware that women’s colleges were a mix of matrimonial market with blue-stocking, and he did not approve of either as components in a university. For better or worse, he enjoyed the flavor of male monasticism in St. Johns. He knew very well that the flavor of female monasticism would be very different. He was after all surrounded by female celibacy – his sisters, Beatrice’s sisters, Saunders, Pellew, etc., etc. His own marriage had been prevented for some years by the crazy “Dick” (Edith Durham) – and so on. I recall (but cannot date) his giving me a little lecture on women in the university – that of course they could learn whatever they might want to – but not to be a part of the institution which might decide the values and style of the institution. If and when the women’s colleges install cellars of good wine like other colleges, then they might be accepted as policy makers in the References and Notes 705

university. ‘Why should the men invite into their society people who will certainly change the character of that society’.”

24 Bashing 1. Morgan, T. H. (1926) William Bateson. Proceedings of the Linnean Society, Session 138, pp. 66–74; Science 63, 531–535. 2. Coleman, W. (1970) Bateson and chromosomes: Conservative thought in science. Centaurus 15, 228–314. 3. Morgan, T. H. (1932) The Scientific Basis of Evolution. Faber & Faber, London. 4. Forsdyke, D. R. (2001) The Origin of Species, Revisited. McGill-Queen’s University Press, Montreal, Quebec, Canada. 5. Farmer, J. B. (1927) Obituary notice of fellows deceased. 1927. William Bateson, 1861-1926. Proceedings of the Royal Society, B. 101, i-v. 6. Haldane, J. B. S. (1926) The Nation and the Atheneum. Feb. 20, p. 713. 7. Haldane, J. B. S. (1957) The theory of evolution before and after Bateson. Journal of Genetics 56, 11–27. 8. Punnett, R. C. (1950) Early days of genetics. Heredity 4, 1–10. 9. Fisher, R. L. (1930) The Genetical Theory of Natural Selection. Oxford University Press, Oxford. 10. Darwin, L. (1929) Letter to Fisher, Jan. 19. In: Natural Selection, Heredity and Eugenics. Edited by J. H. Bennett (1983). Oxford University Press, Oxford, pp. 95–96. 11. Fisher, R. A. (1936) Has Mendel’s work been rediscovered? Annals of Science 2, 225–137. 12. Cock, A. G. (1985) Mendel’s honesty. The Biologist 35, 5. 13. Cock, A. G. (1973) William Bateson, Mendelism and biometry. Journal of the History of Biology 6, 1–36. 14. Stern, C. (1974) The domain of genetics. Presidential address. Genetics 78, 21–33 15. Qin, J., Calabrese, P., Tiemann-Boege, I., Shinde, D. N., Yoon, S.-R., Gelfand, D., Bauer, K., & Arnheim, N. (2007) The molecular anatomy of spontaneous germline mutations in human testis. PLoS Biology 5, e224 16. Haldane, J. B. S. (1956) The theory of selection for melanism in Lepidoptera. Proceedings of the Royal Society, B. 144, 217–220. 17. Majerus, M. E. N. (1998) Melanism. Evolution in Action. Oxford University Press, Oxford. 18. Hooper, J. (2002) Of Moths and Men. An Evolutionary Tale. Norton, New York. 19. Muller, H. J. (1934) Lenin’s doctrines in relation to genetics. In: Graham, L. R. (1972) Science and Philosophy in the Soviet Union. A. Knopf, New York, pp. 451–469. 20. Huxley, J. (1942) Evolution the Modern Synthesis. Allen & Unwin, London. 21. Huxley, J. (1970) Memories. Allen & Unwin, London. 22. Darlington, C. D. (1964) Genetics and Man. Allen & Unwin, London. 706 References and Notes

23. Weiss, K. (2002) Good vibrations: The silent symphony of life. Evolutionary Anthropology 11, 176–182. 24. Mallet, J. (2004) Species problem solved 100 years ago. Nature 430, 503. 25. Mayr. E. (1973) The recent historiography of genetics. Journal of the History of Biology 6, 125–154. 26. Dawkins, R. (1986) The Blind Watchmaker. Longmans, Harlow, Essex, England. 27. Dawkins, R. (1983) Universal darwinism. In: Evolution from Molecules to Man. Edited by D. S. Bendell. Cambridge University Press, Cambridge. 28. Dawkins, R. (1982) The Extended Phenotype. W. H. Freeman, San Francisco, CA. 29. Wilson, D. S. (2001) Evolutionary biology. Struggling to escape exclusively individual selection. Quarterly Review of Biology 76, 199–205. 30. Dawkins, R. (1998) Unweaving the Rainbow. Houghton Mifflin, New York. 31. Lamphere, L. (2001) Letter from the President of the American Anthropo- logical Association. New York Times. (Aug. 12). 32. Dawkins, R. (1998) Postmodernism disrobed. A review of Intellectual Impostures. Nature 394, 141–143. 33. Gould, S. J. (1972) Zealous advocates. Science 176, 623–625. 34. Gould, S. J. (1980) Is a new and general theory of evolution emerging? Paleobiology 6, 119–130. 35. Orr, H. A. (1996) Dobzhansky, Bateson and the genetics of speciation. Genetics 144, 1331–1335. 36. Coyne, J. A. & Orr, H. A. (1998) The evolutionary genetics of speciation. Philosophical Transactions of the Royal Society 353, 287–305. 37. Lynch, M. & Force, A. G. (2000) The origin of interspecies genomic incompatibility via gene duplication. American Naturalist 156, 590–605. 38. Wilkins, A. (2001) The Evolution of Developmental Pathways. Sinauer, Sunderland, MA. 39. Gavrilets, S. (2004) Fitness Landscapes and the Origin of Species. Princeton University Press, Princeton, NJ. 40. Coyne, J. A. & Orr, H. A. (2004) Speciation. Sinauer, Sunderland, MA. 41. Johnson, N. A. (2002) Sixty years after “Isolating mechanisms, evolution and temperature”: Muller’s legacy. Genetics 161, 939–944. 42. Navarro, A. & Barton, N. (2003) Chromosomal speciation and molecular divergence—accelerated evolution in rearranged chromosomes. Science 300, 321–324. 43. Crow, J. F. (2005) Herman Joseph Muller, evolutionist. Nature Genetics 6, 941–945. 44. Scannell, D. R., Byrne, K. P., Gordon, J. L., Wong, S., & Wolfe, K. H. (2006) Multiple rounds of speciation associated with reciprocal gene loss in polyploid yeasts. Nature 440, 341–344. 45. Mackenzie, D. (1978) Statistical theory and social interests. Social Studies in Science 8, 35–83. 46. Allen, G. E. (1978) Opposition to Mendelian-chromosome theory: The physiological and developmental genetics of Richard Goldschmidt. Journal of the History of Biology 7, 49–92. References and Notes 707

47. Cock, A. G. (1983) William Bateson’s rejection and eventual acceptance of chromosome theory. Annals of Science 40, 19–59. 48. Smolin, L. (2007) The other Einstein. The New York Review of Books 54 (10), 76–83. 49. Bowler, P. (1983) The Eclipse of Darwinism. Anti-Darwinian Evolution Theories in the Decades around 1900. John Hopkins University Press, Baltimore, MD. 50. Adams, M. B. (1990) La génétique des populations était-elle une génétique évolutive? In: Histoire de la Génétique, pp. 153–171. Edited by J. -L. Fischer & W. H. Schneider, ARPEM, Paris. [There is also: “Little Evolu- tion, Big Evolution. Rethinking the History of Population Genetics,” a 34 page unpublished manuscript kindly sent to DRF by MBA in 2003.] 51. Wells, H. G., Huxley, J., & Wells, G. P. (1931) The Science of Life. Cassell, London. 52. Henig, R. M. (2000) The Monk in the Garden. The Lost and Found Genius of Gregor Mendel, the Father of Genetics. Houghton Mifflin, New York. 53. Richmond, M. (2001) Women in the early history of genetics. William Bateson and the Newnham College Mendelians, 1900-1910. Isis 92, 55–90. 54. Richmond, M. (2006) The “domestication of heredity”: the family organization of geneticists at Cambridge University, 1895-1910. Journal of the His- tory of Biology 39, 565–605. 55. Bateson, P. (2002) William Bateson. A biologist ahead of his time. Journal of Genetics 81, 49–58.

25 Epilogue 1. Keilin, D. (1966) The History of Cell Respiration and Cytochrome. Cambridge University Press, Cambridge. 2. Mathew, P. (1831) Naval Timber and Arboriculture. Longmans & Co., London. 3. Lock, R. D. (1906) Recent Progress in the Study of Variation, Heredity and Evolution. John Murray, London. 4. Darwin, C. (1874) The Descent of Man. John Murray, London. 5. Forsdyke, D. R. (2001) The Origin of Species, Revisited. McGill-Queen’s University Press, Montreal, Quebec, Canada. 6. Forsdyke, D. R. (2006) Heredity as transmission of information: Butlerian “intelligent design.” Centaurus 48, 133–148. 7. Olby, R. (1971) The influence of physiology on heredity theories in the nineteenth century. Folia Mendeliana 6, 99–103. 8. Forsdyke, D. R. (2000) Tomorrow’s Cures Today? Harwood Academic, Amsterdam, The Netherlands. 9. Punnett, R. C. (1950) Early days of genetics. Heredity 4, 1–10. 10. Forsdyke, D. R. (2004) Grant Allen, George Romanes, Stephen Jay Gould and the evolution establishments of their times. Historical Kingston 52, 95–103. 708 References and Notes

11. Morton, P. (2005) The Busiest Man in England: Grant Allen and the Writing Trade, 1875-1900. Palgrave-Macmillan, New York. 12. Provine, W. (1971) The Origins of Theoretical Population Genetics. The University of Chicago Press, Chicago, IL. 13. Crowther, J. G. (1952) British Scientists of the Twentieth Century. Routledge & Kegan Paul, London. 14. Forsdyke, D. R. (2006) Evolutionary Bioinformatics. Springer, New York. 15. Gulick, A. (1932) John Thomas Gulick. Evolutionist and Missionary. University of Chicago Press, Chicago, IL. 16. Gulick, J. T. (1891) Remarks on Mr. Moulton’s reasonings and calculations. Unpublished paper (142/2B). Galton Archives, University College, London. 17. Dobzhansky, T. (1962–1963) Oral memoir quoted in Provine, W. B. (1994) The origin of Dobzhansky’s Genetics and the Origin of Species. In: The Evolution of Theodosius Dobzhansky. Edited by M. B. Adams, Princeton University Press, Princeton, NJ. 18. Provine, W. B. (1986) Sewall Wright and Evolutionary Biology. The University of Chicago Press, Chicago, IL. 19. Cock, A. G. (1980) Faking and the intent to deceive. British Medical Journal 281, 1214–1215. 20. Meijer, O. G. (1982) The essence of Mendel’s discovery. In: Gregor Mendel and the Foundation of Genetics, the Past, and Present of Genetics. Edited by V. Orel. The Mendelianum of the Moravian Museum, Brno, Czech Republic, pp. 173–200. 21. Cock, A. G. (1988) Mendel’s honesty. The Biologist 35 (1). 22. Baumgartner, A. & Ettinghausen, A. von (1842) Die Naturlehre nach ihrem gegenwärtigen Zustand mit Rucksicht auf mathematische Begrundung. Seventh edition. Carl Gerold, Vienna, Austria. 23. Turing, A. M. (1952) The chemical basis of morphogenesis. Philosophical Transactions of the Royal Society of London. B. 237, 37–72. 24. Hurst, C. C. (1932) A genetical formula for the inheritance of intelligence in man. Proceedings of the Royal Society, B. 112, 80–97. 25. Hurst, R. (1951) What’s All This About Genetics? Watts & Co, London. 26. Crew, F. A. E. (1967) Reginald Crundall Punnett. Biographical Memoirs of Fellows of the Royal Society 13, 309–326. 27. Muggeridge, M. (1936) The Earnest Athiest. Eyre & Spottiswoode, London. 28. Bateson, M. C. (2006) Personal communication to DRF. 29. Heitland, W. E. (1908) A Letter to a Lady, or a Word with the Female Anti-Suffragists. E. Johnson, Trinity Street, Cambridge, MA. 30. Jones H. F. (1919) Samuel Butler, Author of Erewhon (1835-1902): A Memoir, Vols. 1 and 2. Macmillan, London. 31. Butler, S. (1914) The Humour of Homer and Other Essays. Kennerley, New York. 32. Voltaire (1770) Letter to F. L. H. Leriche. In: The Complete Works of Voltaire, Vol. 120. The Voltaire Foundation, Banbury, Oxfordshire, (1975), p. 18. 33. Coleman, W. (1970) Bateson and chromosomes: Conservative thought in science. Centaurus 15, 228–314. References and Notes 709

34. Fowler, H. (1996) In: Cambridge Women: Twelve Portraits. Edited by E. Shils & C. Blacker. Cambridge University Press, Cambridge. 35. Cock, A. G. (1980) Obituaries of Rona Hurst and Gregory Bateson. Mendel Newsletter 19, 5–7.

Appendix 1. Lyell, C. (1863) The Geological Evidences of the Antiquity of Man with Remarks on Theories of the Origin of Species by Variation. G. W. Childs, Philadelphia, PA. 2. Körner, K. (1983) Linguistics and Evolution Theory (Three Essays by August Schleicher, Ernst Haeckel and Wilhelm Bleek). J. Benjamin, Amsterdam, The Netherlands. 3. Bateson, G. (1979) Mind and Nature. Dutton, New York. 4. Mayr, E. (1970) Populations, Species and Evolution. Harvard University Press, Cambridge, MA. 5. Crick, F. (1971) General model for chromosomes of higher organisms. Nature 234, 25–27. 6. Forsdyke, D. R. (2006) Evolutionary Bioinformatics. Springer, New York. 7. Forsdyke, D. R. (2007) Calculation of folding energies of single-stranded nucleic acids: Conceptual issues. Journal of Theoretical Biology 248, 745–753. 8. Forsdyke, D. R. (2007) Molecular sex. The importance of base composition rather than homology when nucleic acids hybridize. Journal of Theoretical Biology 249, 325–330.

Acknowledgements

Donald Forsdyke

The “repatriation” of the Bateson papers was approved by Gregory Bateson and by Mary Catherine Bateson, who is now Bateson’s closest surviving relative. With Stephen Jay Gould and David Lipset, she assisted Alan Cock in the collecting and sorting for shipment to the UK. The paper’s first home was the University of Southampton where cataloguing, supported by the Wellcome Trust, was carried out by Alan with secretarial help from Carol Newhouse and Patricia Hughes. Sources of other Bateson-relevant materials included the American Philosophical Society, Baltimore (the William Cole- man microfilm), St. John’s College, Cambridge, the Galton Collection at University College, London, the University of Cambridge Library, and the Cambridge Department of Genetics (the Bateson-Punnett notebooks). Ar- rangements were made for the transfer of materials to the John Innes Centre (Norwich), where photocopying and further cataloguing and indexing were carried out in collaboration with Brian Harrison, Peter Young and Rosemary Harvey. Alan was granted interviews by Nora Barlow, Patrick Bateson and his father F. W. Bateson (a nephew of William Henry Bateson), Cyril Darlington, Edmund B. Ford, Geoffrey Keynes, Lily Newton (wife of W. C. Frank Newton) and Helen Pease. His studies of the Pearson-Weldon-Galton correspondence in the Watson Library at University College were facilitated by Maxine Merrington. Alan’s wife, Marta Cock, helped me trace him in 2004 to his address in London, where he and his four children – Sybil, Hannah, Christina and Oliver – approved the transfer of his unfinished manuscript and his copy of the Bateson papers to Canada. Sybil did the packing and arranged the shipment. Christina authorized documentation relating to Alan’s estate. In 2005 the Librarian at the John Innes Centre, Kenneth Dick, facilitated my examination of the Bateson Library, which the Trustees had purchased from relatives in the USA. In the course of this flying visit I came across an unpublished five volume treatise – William Bateson and the Emergence of Genetics (2000) – composed by Rosemary Harvey (retired). The index indicated a different approach to the present work. My plan to photocopy the Harvey treatise did not come about, and I reluctantly decided that this would be one promising stone I would leave unturned. Among those who helped me with the relevant literature, particular thanks are due to Mark Adams and Marsha Richmond for 712 Acknowledgements making available advanced copies of various texts, Christiane Nivet for comments on Chapter 20, and Patrick Bateson for advanced copies of papers. Joan Horton kindly searched Bateson family UK census records. My getting up-to-speed in the history of science was assisted by Gerard Wyatt’s dona- tion of books from his collection. Permission to quote from Rona Hurst’s unpublished text was given by Patrick Zutshi on behalf of the Syndics of Cambridge University Library. The figures were either drawn especially for this volume, or were taken from my previous works with the approval of the publishers (McGill- Queen’s University Press, Montreal, and Springer, New York). The various photographs were either taken from sources where copyright had expired, or were part of Alan Cock’s set of the Bateson papers’ which I have now depos- ited in the Archives of Queen’s University at Kingston, with the kind assistance of Paul Banfield. The photograph of Samuel Butler is reproduced with the permission of the Master and Fellows of St. John’s College. Queen’s University provided invaluable library resources and host’s my web-pages where further information on Bateson and his fellow “Mendels” may be found. Special thanks are due to Emily Blackie and others at the Bracken Library for assistance in hunting down evasive references. Andrea Macaluso, and Saladi Gunabala and her team at SPi Technolo- gies in Pondicherry, India, and Melanie Wilichinsky of Springer (New York), Jeffrey Ciprioni, James Russo smoothed the passage from typescript to final copy. Charlotte Forsdyke assisted me with the copy-editing. Polly Forsdyke advised on French and German usages. Sara Forsdyke and Caroline Falkner advised on Latin and Greek usages. Ruth Forsdyke provided helpful texts that I would not have otherwise found. My wife Patricia was a constant source of inspiration and advice. Alan was close to his family. Had he written an acknowledgement, I am sure it would have revealed an indebtedness to them that is no less than I to my own.

Index

A population may contain multiple, 475, A Naturalist in Nicaragua (Belt), 126 507 A Passage to India (Forster), 9, 385 shortened to “alleles”, 208 A Period Piece (Raverat), 13, 422 their linkage to others, 372 academic career Allen, G, 6, 275, 530, 648 influence of degree on, 76 Alytes obstetricans. See midwife toad is like musical chairs, 296 American Museum of Natural History, 263, negative findings do not help, 38 633 academic examinations amoeba, 12, 531 as genetic calibration, 553 amphibia, 446, 569, 582 conduct of examiners in, 522 An Examination of Weismannism high mathematical content of, 611 (Romanes), 59, 68 Littlego. See Littlego examination An Introduction to Biology (Darbishire), 247 poor performance of Mendel in, 565 analogy poor performances of the eminent, 421 between biological species and a pressure on Anna before, 77 discipline, 653 Previous. See Previous Examination between hereditary and mental accent information, 90, 544, 551 as base composition of DNA, 123, 668, between hybrid sterility and not 670 tuningin mentally, 544 as methylation of DNA, 453 between Mendel and Morgan, 560 differences in make species of key and lock, 325, 487, 529 incompatible, 668 of light and sound transmission, 528 reproductive, 123 of playing cards, 248, 343, 344, 532 acorn worm, xviii, 17, 18, 20, 35 of shooting at a mark, 103 Ad Eundem Club, 226 of spandrels, 330 Adams, M. B, 638 of sword and scabbard, 325, 487, 648, 670 Agar, W, 214 ancestor age contribution of, 66, 223, 227 correlation with size, 156 reversion to, 137, 201, 249, 530, 533 deferred, 31, See rejuvenation ancestral series of reproduction, 532 successive terms of, 92, 108, 198, 518 of the earth, 495, 542 Ancestrians. See Biometricians Age and Area (Willis), 503 Animal Colouration (Beddard), 147 aggregation Animal Life and Intelligence of like with like, 529 (C. L. Morgan), 148 Agricultural Chemistry (Liebig), 524 anthropology, xiv, 140, 556, 635 Aiken, Anna. See Bateson, Anna (Annie) anti-Darwinian Aiken, J, 6, 386, 603 advocates, 5, 39, 135, 372, 514, 530 Alice in Wonderland (Carroll), 553 typologists, 668 Allard, Mr, 383, 390, 461 Aral Sea, xxii, 20 allelomorphs Aristotle, 28, 633 are alternative forms of an element, 475 armed forces are on homologous chromosomes, 282 Bateson’s lectures to, 460, 657 come in pairs, 297, 324 conscription to, 57, 422 introduction of the term, 209 health of, 419 714 Index

horses for, 396 B Military Service Tribunal for, 659 Bach, J. S, 545 weapons for, 440 Backhouse, W, 383, 390, 391, 462 Armstrong, H, 440, 444, 592 Baer, K. E. von, 12 Arnold, M, 612 Bailey, A, 462 art Bailey, P. C, 385, 390 as one of two cultures, 407, 444, 556, Balanoglossus. See acorn worm 616, 617 Balfour Biological Laboratory for Women, Buddhist wall paintings, 384 179 Cennini’s painting in tempera, 175 Balfour Chair in Genetics, 386, 425 chaos acclaimed as, 436 Balfour Fund, 232, 348 contempt for, 435, 616 Balfour Studentship, 33, 36, 102, 219, 244 copying in Florence, 181 Balfour, A, 6, 12 emancipation in, 432 as anonymous donor, 306, 382 enjoyment of, 14, 392 as Prime Minister, 419 Hermitage, St. Petersberg, 511 at the Atheneum, 387 Hornsey School of, 384 nephew of Lord Salisbury, 702 Madonna di San Siste, Dresden, 14, 562 on eugenics, 429 male domination of, 216 President at BA, 233 medicine as an, 84 President of Genetical Society, 469, 498 National Arts Collections Fund, 216 Balfour, Eleanor. See Sidgwick, Eleanor post-impressionism, 216, 392 Balfour, Evelyn, 12 practice of, 388 Balfour, F, 11, 17, 33, 35, 96, 101 Raphael, 14, 465 Balfour, G, 12, 457 Rembrandt, van R, 465 Balls, W, 214, 391 sales reported in the Morning Post, 233 Balzac, H. de, 187, 212 Slade Professorship at Cambridge, 527 Bařina, F. S, 562 Society of Painters in Tempera, 216 Baring, E, 31 trade in, 186, 203, 295, 326, 388, 407 Barker, B. T, 382 Van Gogh, 652 Barlow, A, 461, 624 Wallace Gallery, 326 Barlow, N. See Darwin, N with science is extra-national, 463 barrier, 134, 672 Artemia salina. See brine shrimp as natural property of organisms, 234 artificial selection, 115, 423, 542. See developmental, 119, 121 breeding geographical, 117 asexual reproduction, 138, 165, 261, 287, 324, 339, 456 linguistic, 123, 667 Asiatic Mediterranean Sea, 22, 29 of Weismann, 650 Asquith, H, 443, 615 originating, 211 atheism, xx, 11, 42, 188, 554, 624, 633, 662 sterility, 119, 121, 211, 403 Atheneum, 387, 649 temporal, 117, 122 Austin, J, 386 transmission, 118, 121 Australia, 409, 432, 440 Bartholemew, A. T, 543, 702 Authoress of the Odyssey (Butler), 548 Bashley, 83, 279, 659 authority in science, 392 Bateman, K. G, 573 Butler’s view on, 535, 538 Bates, H, 148 publication obstructed by, 274, 375, 399, Bateson, Anna, 8, 131, 659 638, 647, 655 academic pressure on, 77 review of research by, 207, 623 as business woman, 160 role of mentor, 201, 247, 565, 647 encouraged by Bateson, 617 the dynasty of selection, 234, 502, 647, 664 publications of, 80 the role of bluff, 152 second class degree of, 78 Weldon’s misuse of, 157, 222 studies of cross-fertilization by, 81 Index 715

Bateson, Anna (Annie), 7, 24, 310, 398, Bateson, R, 6 461, 602 Bateson, W Bateson, C. Beatrice, 384, 409, 471, 624, agreement with Wallace, 309 640 allies of, 75, 144, 222, 265, 269, 283 as author, xx ancestors of, 419 as hostess, 499 anticipations of Mendel by, 197 collection of letters by, xiv, xxi, 638, 657 apparent disregard of Romanes by, 133, kept from William, xx, 180, 704 135, 406 malchaperoning of, 40 appointed College Fellow, 20 marriage of, 180 appointed College Steward, xxii, 664 role in research, 219, 656, 657 appointed Newton’s Deputy, 186 role in writing, 387 appointed Reader, 303 Bateson, Edith, 9, 184, 389 appointed to Chair in Biology, 305 Bateson, Edward, 9, 25, 80, 461, 624, 640, 660 appointed to John Innes Institute, 329, Bateson, F. W, 711 336 Bateson, G, xiv, xv, 624, 641, 711 as a “Mendel”, 644 as Cambridge spy, 503 attacked by Coleman, 638 at St. John’s College, 501, 556 attacked by Darlington, 632 birth of, 188, 232, 420 attacked by Dawkins, 635 mimicry by, 421 attacked by Fisher, 629 on his father, 44, 181, 219, 556, 649, attacked by J. Huxley, 632 662, 665, 704 attacked by L. Darwin, 629 on information, 369, 668 attacked by Mayr, 633 on the departure from Grantchester, 389 attacked by Muller, 631 role in Kammerer visit, 577 Centenary essay of, 329, 356, 365, 483, transient engagement of, 556 547, 637, 665 visits Galapagos Islands, 510 chromosome doubts of, 268, 297, 339, Bateson, H, 6, 95, 245 347, 353, 365, 447, 451, 459, Bateson, J, 554 476, 489, 508, 517, 628, 638 birth of, 183 chromosome role affirmed by, 342, 478, death of, 461, 554 489 enlistment of, 439 collaboration with Saunders, 160, 219, military cross awarded, 460 249, 640 Bateson, M. C, xiv, xv, 556, 658, 711 disparagement of Butler, 552, 555 Bateson, Margaret, 39 disparagement of Galton, 140, 202, 323, as prompter of female initiative, 83, 601 434 disapproval of marriage by, 42 disparagement of Romanes, 142 on privacy of letters, xxi disparagement of Semon, 551 work for Queen magazine, 8, 660 disparagement of Weismann, 142 Bateson, Martin, 461, 554 dispute with de Vries, 70, 260, 329, 365, at St. John’s College, 555 403, 503, 510 birth of, 188 dispute with Morgan, 376 enters RADA, 473, 555 dispute with Pearson, 164, 222 his deep reading of Butler, 555 dispute with Poulton, 147, 309 misidentified as Mr. Martin, 576 dispute with Weldon, xxiii, 36, 102, 152, Royal Air Force training of, 439, 554 157, 222, 232, 323, 664 suicide of, 492, 555 dominance explained by, 369 Bateson, Mary elected FRS, 96 book on medieval England by, 242 elected to Russian Academy, 511 death of, 262 error corrected by Nettleship, 351 friend of C. J. Herringham, 8, 173 eugenic caution of. See eugenics speech to the Prime Minister by, 601 eureka moment of, 305, 354 success in examinations by, 80 first class degree of, 77 Bateson, P, 641, 660, 711 Garrod contacted by, 230 716 Index

health of, 299, 314, 408, 481, 508, 624, Bateson, W. H, 43, 601, 711 664 clash with Dr. S. Butler, 522 his debt to Weldon, 152, 245, 246 death of, 12 his feelings of alienation, 35, 554 elected Master of St. John’s, 5, 523 his heroes, 38 enters St. John’s College, 3, 523 his ignorance of Butler-Darwin dispute, on conduct of examinations, 522 545, 546 on women’s education, 602 his patriotism, 585 Bathurst Studentship, 79, 179, 295 his role in SEB, 589 Baur, E, 321, 374, 387, 397, 432, 474, 561 knighthood declined by, xviii, 492, 636 critic of Kammerer, 575 lecture notes of, 135, 545 first genetics journal begun by, 337 lectures on physiological selection, 140 first meeting with, 329 love letter from, 40 his narrowness of interests, 407, 561 mistaken optimism of, 20, 86, 215, 226 post-war activities of, 592, 593 on democracy and socialism, 393, 423, Bayly, Mr, 46 426, 436, 464, 500 Beddard, F. E, 147, 159 on F. Darwin, 544 Bedford College, 181 on Haldane’s rule, 471 Beethoven, L. van, 432, 465 on Johannsen’s terminology, 287, 504 Bell, J, 421 on laboratory studies, 80, 98, 441, 449, Belt, T, 126 477, 610, 653 Ben Battle, 278, 315 on liberty, 501, 513 Berlin, 320, 329, 439 on M. Foster, 215 Agricultural College, 407 on materialism. See heredity, Fifth Genetics Congress at, 594 transmitting agency in Institut für Biologie, 394, 413 on medical ethics, 431 storm trooper raid in, 631 on one sex as heterogametic, 348 Bernard, B, 190, 336 on pacifism, 439 bible, xx, 3, 441, 554 on pangenesis, 504 bible classes, 13, 213 on prohibition, 435, 478 Biffen, R, 214, 231, 244, 274, 381, 444, on religion, 11, 42, 311, 428, 431, 436, 492, 654 480, 508, 554, 556, 565, 624 Binyon, L, 384, 392, 478, 552 on scientific instruments, 508, 651, 653 biochemistry on species classification, 192 20th century dominance foreseen, 402, on study of zoology, 86, 508 509, 515, 638, 649, 665 on W. H. Bateson, 43 genetic engineering derived from, 506 open-mindedness of, 376, 382, 415, 472, of inherited disease. See Garrod 475, 515, 628, 662 of plant pigments, 231, 335 philosophy of, 135, 141, 311, 427, 442, struggle for recognition of, 230 464, 466, 555, 662 biology praise of Butler by, 547 as part of “natural knowledge”, 442 praise of Morgan by, 445, 478, 517 Chair of at Cambridge, xii, 305, 546 praise of Morgan School by, 482, 509 consequences of unapplied knowledge praise of physiological selection by, 131, of, 436 333, 645 five branches of, 85 praise of Weismann by, 329 fundamental goal of, 233 praised by Stern, 630 future of, 165, 509 regrets of, 623 origin of the word, 508 research agenda of, 85, 136, 159, 160, 189 revolution in, xix, 30, 233, 419, 508, 590 retirement of, 623 society for. See Society for Experimental short-hand of, xv, 204, 237, 243, 270, Biology 272, 275, 353 steam introduced into, 20 writing style of, 387, 576, 640 wholesome teaching of, 435

Index 717

Biometricians brine shrimp, 23, 39 contribution to statistics by, 221, 281, 653 Bristol, 161, 391 dispute with Mendelians, xxiii, 102, 232, British Association 240, 273, 276, 280, 315, 323 1887 (Manchester), 49, 341 new, 247, 632, 637 1888 (Bath), 123 Biometrika, 205, 222, 241, 245 1894 (Oxford), 144, 542 editorial board of, 247, 421 1898 (Bristol), 161 establishment of, 169 1903 (Southport), 226, 229 biotype 1904 (Cambridge), 142, 233, 241, 273, definition of, 287 647, 657 term introduced, 286 1906 (York), 244, 247, 444 Birkbeck College, 508, 553, 588, 654 1907 (Leicester), 302 Black, C. See Garnett, C 1908 (Dublin), 310, 544 Blackman, F. F, 176, 296 1909 (Winnipeg), 334 Blake, W, xxiv, 13, 183, 213 1910 (Sheffield), 389 blending inheritance, 62, 64, 70, 116, 189, 1911 (Portsmouth), 393, 397 198, 269, 533, 672 1912 (Dundee), 398 Bloodstock Breeders’ Review, 397 1913 (Birmingham), 408 Bloomsbury, 8, 30, 40 1914 (Australia), xxiv, 357, 407, 430 Board of Agriculture, 385, 394, 396, 439 1921 (Edinburgh), 495 Boer War, 188, 396, 419 1922 (Hull), 502 Bonnet, C, 415 1923 (Liverpool), 496, 502 Botanic Gardens funding by, 244, 273 Cambridge, 22, 79, 160, 182, 188, 244, British Museum, xxiv, 102, 190, 384, 492, 385, 496, 618 511, 525, 526 Copenhagen, 508 Brno. See Brünn London. See Kew Gardens Brooklyn Institute of Arts and Sciences, New York, 142 225, 296, 301 Paris, 200 Brooks, W, 6, 17, 18, 39, 225, 663 Peradeniya, 291, 382 Brown, F. A, 13, 14 Boulenger, G, 575, 593 Brown, R. Staples, 226 Boveri, T, 234, 235, 324, 339, 341, 347, Brücke, E, 530 348, 359, 394 Brunftschwielen. See nuptial pads Bowler, P, 638 Brünn, xi, 242, 273, 389, 559, 560, 562 Brainerd, E, 299 Brussels, 279, 594 breeding. See artificial selection Bryan, W. Jennings, 485, 500 as a promising start, 19, 87, 100, 312 Buch, L. von, 144 Bateson’s questions on, 137 Buffon, G. L. L. de, 19, 521, 535, 547 cross-bred, definition of, 248, 313 bulldog does not give a “physiological species”, Bateson’s, 265, 654 509 Darwin’s, 4 must begin with pure lines, 229, 271 Mendel’s, xxiii, 197 of horses, 396 Pearson’s, 247 of orchids, 267, 272 Bunyard, E, 469 pure-bred, definition of, 248, 313, 412 Burbage, 212, 265, 278, 302, 395, 408, 425, restricted by contraception, 389, 435 439 studies neglected post-1859, 204, 233, Burke, J. B, 261 321, 329, 372, 649 Burkill, J. H, 154 the dihybrid case, 250, 251 Burney Prize Essay, 76 the monohybrid case, 250, 251, 258, 362 Butler, Dr. S, 521 true, 70, 209, 256, 258, 261, 269 appointed Bishop, 523 Breeding and the Mendelian Discovery headmaster of Shrewbury, 3 (Darbishire), 246 Butler, S, 57, 293, 367, 443, 508, 557, 639 Bridges, C, 4, 375, 417, 447, 451, 478 anticipation of Semon by, 145, 549 718 Index

anticipation of Weismann by, 547 emigration to, 513 as a “Mendel”, 644 Toronto Conference in, 481 at Weldon’s lecture, 101 Canary Islands, 144, 409 attacked by Fisher, 629 Candolle, A. de, 419 Bateson’s reading of, 320, 321, 329, Cannon, H. G, 578 546, 646 Cannon, W, 271, 342, 344 birth of, 3, 523 Cardium edule, 87, See cockles Darbishire as disciple of, 247 Carlyle, T, 612 death of, 534 Carnegie Institution, 224, 664 enters St. John’s College, 4, 524 Carnegie Station for Experimental friendship with R. Garnett, 39 Evolution, 232, 283, 307 his ghost, xxiv Castle, W. E, 226, 275, 349, 355, 371, 481 his land of Erewhon, 311, 426, 546 Catchpool, E, 115, 123, 648, 664 his reading of Darwin, 524 Cayley, D, 390, 439, 461 his reading of history, 643 Cecil, Robert. See Lord Salisbury lecture at Working Men’s College by, Cennini, 175 101, 527 Ceylon, 35, 291, 313, 385 materialist viewpoint of, 531, 661 Chalmers-Mitchell, P, 444 Newth as disciple of, 39 critic of Materials, 98 on origin of variation, 90, 537 critic of the IRC, 597 on true greatness, 559, 661 critic of Toronto Address, 485 refuses to be ordained, 524 Secretary of Zoological Society, 225, 327 sheep farmer in New Zealand, 524 character Butler, S (1612-80), 508, 546, 654 acquired through use/disuse, 113, 516, Butler, T, 3, 523 534, 541, 550, 562 ancestral, 50, 55, 65, 210 C as a unit, 204, 207, 235, 271, 285, 322, Calandruccio, S, 586 366 Caldwell, W. H, 33, 35, 128, 467 blending. See blending inheritance Callinicus: a Defence of Chemical Warfare blurred distinction from “allelomorph”, (Haldane), 440 243 Cambridge Essays on Education, 444 blurred distinction from “determinant”, Cambridge Instrument Company, 35 272 Cambridge University blurred distinction from “element”, 204, Department of Genetics, xv, 219, 386, 657 237, 248, 314, 372 Eugenic Society, 425 compound, 232, 235, 272, 281, 298, 324 expansion of curriculum at, 611 coupled. See character, linkage Farm, 385 dependent on “ultimate factors”, 322 Financial Board, 618 determinants of, 267, 272, 366, 474 Genetical Research Institute, 495 dominant. See dominance influence of agricultural depression on, 613 gametic, embryonic or adult, 50, 273 Natural History Society, 501 generic (of a genus), 269, 354 Philosophical Society, 319 incidental or decorative, 330, 516, 625 Plant Breeding Institute, 394 independent assortment of, 344 Press, xii, 263, 319, 391 inheritance of indirect, 341, 528 Press Syndicate, 47, 72, 319 inutility of, 97, 406 Rede Lecture, 444, 554 latent, 50 School of Agriculture, 394 latent in F1 generation, 205 Senate, 179, 305, 603, 605, 609 linkage, 209, 234, 253, 343, 345, 358, Sidgwick Memorial Lecture, 448 360, 363, 372, 430, 445, 448, Canada 517 BA meeting in, 334 loss per generation, 51, 269, 341 emigration from, 648 mean value of, 151

Index 719

mid-parent value of, 62, 66 as individuals, 341, 343, 347, 349, 367 non-blending, 108, 179, 198, 200 as unique entities, 354, 415 patent, 50 conjugate in meiosis, 341, 346, 361, 368, qualitative, 67, 199 487 quantitative, 199, 258, 475 constancy within a species, 267, 341 repelled, 372, See character, linkage contain blocks representing characters, represented by a chromosome particle, 342, 482, 517 342, 449, See particulate contain nucleic acid and protein, 414 theory of heredity correlation with “chromatin”, 343, 362 segregation of. See segregation crossing-over between. See sex-linked. See sex-linkage recombination skipping generations, 50 differentiation of, 349, 351, 353, 359 solely zygotic, not gametic, 250 direct inheritance of, 341, 367, 416 specific (of a species), 269, 353, 354, disjunction of in meiosis, 340 364, 406, 412 if complementable can error-correct, transferable in Mendelian analysis, 489, 369, 455 490, 509 linear arrangement of genes on, 235, transitional forms of. See intermediate 339, 346, 374, 445, 447, 517 forms loose pairing in autopolyploids, 414, 457 transmission of, 223, 267, 298, 346 mapping gene location on, 447, 517 two classes of, 198 mechanism of pairing of, 487, 670 utility of, 516 no tissue-specific differentiation of, 372, varietal (of a variety), 269, 354, 364, 373, 491 412, 497 non-disjunction of in meiosis, 376, 447 Chargaff, E, 649 number equals linkage group number, Charles Darwin and Samuel Butler (Jones), 375, 417, 446, 517 550 number halved in gametes, 268, 269, Charles, E, 587 341, 356 Chicago pairing fails in species hybrids, 361, 626, Anniversary Meeting at, 5 669 painful visit to, 300 pairing of complement with chimaeras, xxiv, 381, 453 complement, 487, 670 China, 437, 439 pairing of like with like, 256, 334, 343, Chinese primrose 344, 346, 361, 414, 454, 670 as tetraploid, 454, 467 pairing usually succeeds in race-hybrids, polymorphism of, 516 361 Christ Church College, Oxford, 84 random orientation in meiosis, 344, 363 Christ’s College, Cambridge recombination between. See Charles Darwin as student at, 3, 522 recombination Shipley as Master of, 444 reduction division of, 60, 341, 356 chromatin that determine sex. See sex as contrasted with cytoplasm, 365 chromosomes bears “heredity qualities”, 366 church bears pangens, 510 as a place for marriage, 42, 181 homogenous appearance of, 448 domination by, 436 maintains its individuality, 342 where people stare dully, 393 Chromosome Atlas of Flowering Plants Churchill, W, 429 (Darlington and La Cour), 154 Cineraria, 83, 151 chromosomes Ciona intestinalis. See sea-squirt as determining reproductive isolation, circumcision, 183, 188 497, 669 classification as determining varietal differences, 354, as a genealogical tree, 35, 85 486 discord with hybrid sterility of, 269, 333, as fibrous gels, 477 406, 509, 516 720 Index

of criminals, 434 crime of modes of variation, 87 of pulmonary consumption, 426 of species, 190, 269, 364, 404 sterilization of those committing, 427, 434 of types of sterility, 405, 414 white collar, 420, 434 Clausen, R, 508 Crofts, E, 15, 82, 664 Clodd, E, 530, 661 Croonian Lecture, 289, 474 clone,definition of, 287 Crowther, C. R, 487, 648, 664, 669 Cock, A. G, xiv, xviii, xxi, 181, 437, 484, Crudeli, C. T, 587 490, 556, 630, 634, 636, 638, crustacea, 23, 28, 30, 46, 87, 102 649, 650, 655, 659, 704, 711 Cuénot, L, 229, 232, 251, 273, 275, 279, 561 cockles, 31, 38, 87 Cunningham, J, 44 Coleman, W, xiv, 638, 663, 711 critic of Bateson’s residue (base), 486 Colenso, J. W, 7 critic of Weldon, 152 color-blindness, 234, 349, 357, 429, 446 member of Mendel Society, 336 Columbia University, 4, 292, 342, 371, 445, notes Bateson ignores Romanes, 133, 560 406 competition on hormones, 470 among Galtonian elements, 53, 59, 97 physiological selection praised by, 132 among genes, 52 tirade at Zoological Society by, 327 among ideas, xii, 146 Curie, M, 511 among individual humans, xii, 428 Cuvier, G, 85 between Genetical Society and SEB, 588 cytoplasm, 366, 387, 509 between newspapers, 172 as location of “substratum”, 364 for means of living, 130 as location of Johannsen’s for students with Cambridge, 389 “Grundstock”, 492, 505 of women with men, 178 Czechoslovakia, 564 winners of are often distinct, 436 Comte, A, 545 D conflict. See competition Dallas, W. S, 539, 547 between elements, 210 Darbishire, A. D, 228, 241, 242, 246, 289, between evolutionary forces, 88 336 between types of information, 668 Darlington, C. D, xxi, 711 Conn, H. W, 98 appointed to John Innes, 490 conservative thought, xiii, xiv, 221, 625, attack on Bateson by, 632 638, 655 attack on Hurst by, 655 Coombe-Tennant, W, 457 misrepresentation of Bateson by, 632 Cope, E, 128, 308 Darwin and Modern Science (Seward), xii, copying. See mimicry 4, 319, 328 as an error-prone process, 90 Darwin Centenary, xxiii, 306, 327, 532 by the masses, 464 Darwin, and After Darwin (Romanes), 72, Cornell University, 300, 479 111, 132, 145, 293 Correns, C, xi, 81, 202 Darwin, B, 82 collected papers of, 564 Darwin, C, xi, 612 discovery of Mendel’s paper by, 197 birth of, 3 on heterogamety, 355 clash with Dr. S. Butler, 521 on linkage, 253 clash with S. Butler, 539, 543, 553, 660 counter-selection, 112 false book of, 28 Coutagne, G, 275 greatest scientific work of, 510 Coyne, J, 637 his delay in publication, xxv crab, 151, 155 his Lamarckian later works, 113, 544, 553 Cramer, F, 100 his views not superceded, 244, 308, 328, Crangon vulgaris. See shrimp 467, 625 Crew, F, 588 inability to understand Galton, 49, 645 Crick, F, 670 on continuity of variation, 448 Index 721

research associates of, xxii, 11, 121 doubts on chromosomes of, 365 shadow of, xii, xxv, 84, 204, 319, 553 scale of operations of, 296 supported by Meldola, 328 visits Brünn, 564 Darwin, Dr. E, 521, 535, 539 visits Cambridge, 321 Darwin, F, 22, 91, 404, 421 Dawkins, R, xvi, 60, 145, 272, 535, 702 advice on dealing with Butler, 540 Dean, F. E, 500 Butler’s ideas recognized by, 544 decoys critic of physiological selection, 126 cytoplasmic inheritance as, 365 death of, 511 Darwinism as, 665 dispute with Butler, 542 descriptive studies as, 233 Dublin BA address by, 310, 544 eugenics as, 419 first class degree of, 75 genic paradigm as, 665 friendship with Butler, 530, 534 hybridizations as, 253 funding by, 244 Mendelism as, 665 his father’s research associate, xxii, 11 non-allelic interactions as, 324 moderating influence of, 154, 304 works of elders as, 72 offer to Anna, 78 degenerate organs, 112, 143 praise of Semon by, 544, 549, 551 Delboeuf, J, 106 rivalry with Romanes, 76 Department of Agriculture, Washington DC, role in Centenary, 319 225, 267 support for women’s education by, 607 depression support of Foster by, 215 A. G. Cock, xvi wives of, 13, 15, 82 Bateson, 263, 383, 492, 664 Darwin, Frances, 82, 664 Beatrice, 183 Darwin, G, 13, 189, 421 C. J. Herringham, 384 Darwin, Gwen. See Raverat, G E. Baur, 593 Darwin, H, 13, 35 Ellen Crofts, 82, 664 Darwin, Henrietta, 543 F. Darwin, 664 Darwin, Major L, 13, 425, 429, 649 Frances Darwin, 664 Darwin, N, 461, 548, 624, 711 H. Muller, 631 C. Darwin’s autobiography revised by, Martin Bateson, 555 543 descent recollection of lectures by 13, 213 doctrine of, 92, 94, 108, 199, 485 work at John Innes of, xxi, 390 evidence for, 86 Darwin, W, 13 from monkeys, 460 Darwinism genetics as physiology of, 329 alleged union with Mendelism, 5, 565, development 632, 650 catapillar as arrested, 205 eclipse of, 632 Galton’s usage of the term, 53 neo-, 5 mechanism of, 215, 373, 402, 685 paradigm of, 23 mutations during, 348, 627, 645 reinterpretation needed, 639 needs cooperation of parental triad becomes a tetrad, 120 chromosomes, 487 ultra-. See ultra-Darwinians of an embryo (ontogeny), 53, 489, 525 universal, 635 seed as arrested, 205 Darwinism (Wallace), 49, 128 Development Commission, 394, 460, 495 Darwinism and the Problems of Life Dew-Smith, A, 82 (Guenther), 261 Dew-Smith, A. G, 15, 35 Darwin-Wallace Celebration (1908), 307 dice Davenport, C, 267, 295, 329, 362, 371, 425, balanced, 88 493, 664 without play, 532 becomes a Mendelian, 232, 273 Die Mneme (Semon), 544, 549 coeditor of Biometrika, 169, 421 Die Mutationstheorie (de Vries), 69, 70, confirms Hurst, 303, 315 246, 260, 261, 291, 364 722 Index

Die Perigenesis der Plastidule (Haeckel), as a dosage effect, 369 529 ignored by Biometricians, 323 Die Pflanzenmischlinge (Focke), 201 in horse coloration, 277 differentiation in mouse coat color, 232 as asymmetric cell division, 297, 401 in sex determination, 355 as differential expression of normal in the presence and absence hypothesis, genes, 69, 239, 491 283, 297, 314 homo- and heterotypic, 167 incomplete, 209, 210, 269, 370, 515 dimorphism, 62, 95, 351, 355, 550 refers to allelic characters, 297 diploidy, 52, 340 unpredictability of, 250, 345 discontinuity Doncaster, L, 214, 461, 654 as chance event, 106 collaboration with Raynor, 371 definition of, 92 coopted to Evolution Committee, 162 in dice scores, 106 failed recruitment of, 217 newspaper metaphor for, 172 move back to Cambridge by, 383 of environment, 99, 109 on sex chromosomes, 335, 351, 354, of species, 93, 94 358, 413, 448 of variation, 89, 92, 94, See variation promise from Przibram to, 574 road as metaphor for, 92 Doyle, A. C, 311, 672 disease Dresden, xx, 14, 40, 562 eugenics as a remedy for, 424 Drosophila melanogaster. See fruit fly inherited. See genetic disease Druery, C, 70, 203 mental, 422, 430, 434, See depression Drummond, H, 275 pulmonary consumption, 426, 432 Duke of Disraeli, B, 386 Argyll, 128, 242 distribution Bedford, 186, 664 bimodal, 62 Devonshire, 613 of a trait about the mean, 111 Duncan, C, 40 of accents in a population, 124, 667 Durham, A, 44 of heights in a population, 62 Durham, C. Beatrice. See Bateson, DNA C. Beatrice accent of, 668 Durham, E, 44, 704 as a duplex, 670 Durham, F. H, 181, 398 as an arrangement of bases, 325, 411, Durham, F. M, 181, 213, 228, 231, 251, 476, 667 278, 335, 355, 390, 398, 439, as genotype, 668 470, 472 as polymer of four bases, 288, 667 Durham, H, 608 base differences between alleles, 297 Durham, M. E, 44, 181 damaged by radiation, 532 like a written text, 647 E mutation in bases of, 415, 507 economics primary information in, 669 and biology, 588 replication of, 529 of growth, 112 secondary information in, 669 of plant breeding, 508 separation of strands of, 326, 670 education sequencing of, 117, 653 “natural knowledge” as a basis for, 441, stem-loop extrusion from, 671 463 third codon base as “residue”, 672 as nurture, 433, 443 triplet codons in, 669 better in Europe, 441 Dobell, C, 39, 385, 459, 490 classical, 442, 473, 610, 612, 616 Dobzhansky, T, 5, 633, 637, 651 goals of, 442, 616 Dogiel, V. M, 512 limits of, 423 dominance, 137, 189, 203 patriotism as indoctrination, 440, 441, absence of, 267 442, 463 Index 723

science neglected in, 442, 444 as “national physiology”, 422 to sway public opinion. See public as applied genetics, 425 opinion Bateson’s caution, 423, 424 women’s. See women capital as an institution of, 433 Egypt, 31 Galton’s hopes for, 420 Eimer, T, 31, 132, 133, 147 International Congresses in, 429, 437, Einstein, A, xvi, 402, 417, 555, 598, 638, 495 640 Eugenics Education Society, 425, 429, 433 elements Eugenics Record Office, 225, 421 as “genes”, 445 evening primrose, 256, 403, 413, 453, 467, as “unit factors”, 281, 330, 410 489, 503, 646 as determining characters, 204, 206, 270, Evidences of Christianity (Paley), 611, 644 450, 474, 632 evolution as representing characters, 53, 58, 210 “synthetic” messages in, 668 competition between, 52 “typological” messages in, 668 different from “characters”, 237 as descent from lower forms, 485, 516 formative, 69, 204, 528 as gain or loss of factors, 323 of Galton, 51, 210 branching, 115, 119 perhaps non-segregating, 364, 474 convergent, 23 embryo, 12 cross between allied forms in, 200, 405 develops itself, 59, 528, 531 discontinuous, 660 morphogenic gradients in, 633 from simple to complex, 412 recalls parental information, 528 Huxley’s remaining problems in, 130 embryology, 85, 560 linear, 117, 119, 126 failure of, xxii, 75, 86, 376 my brain boils with, 32 lectures on by Balfour, 11 of culture, 145 lectures on by Foster, 11 of language, 667 of tissue formation, 449 of mental functions, 58, 159, 542 Russian doll model of, 415 opposition to teaching of, 485, 495, 500 Emerson, A, 5, 480 orthogenetic, 414, 480, 568 Emma Darwin (Darwin), 543 steps in, 64, 86, 199, 405, 482 Encyclopaedia Britannica Wallace’s remaining problems in, 158 Bateson’s article in, 514 Evolution and Adaptation (Morgan), 372 Huxley’s article in, 535 Evolution and Religion in Education Lankester’s article in, 13, 85 (Osborn), 627 Romanes article in, 201 Evolution Committee of the RS, 157 Endless Forms. Species and Speciation, 637 enquiry on melanism by, 163, 631 Entomological Society, 309 establishment of, xxiii environment reports to, 162, 207, 249, 274, 335, 348, adaptation to is only approximate, 91 351, 358, 371 continuous, 139 research agenda of, 158 discontinuities in, 99, 109 resignations from, 161 man’s control over, 464 Evolution Racial and Habitudinal (Gulick), psychological, 146 404 salinity of, 22, 136 Evolution, Old and New (Butler), 329, 535, simulated alpine, 569 539, 546, 643 enzymes, 230, 299, 311, 325, 335, 662 evolutionary bioinformatics (EB), xiii, 521, Erasmus Darwin (Krause), 539 665, 667 Erewhon (Butler), 426, 525, 545, 553 Evolutionary Bioinformatics (Forsdyke), Erewhon dinners, 549 xiii, 526, 553 Essays on Evolution (Poulton), 309 evolutionary biology, xiii, 504, 632, 644 Essays upon Heredity (Weismann), 49, 147 factions in, xvi, 553 eugenics, 246, 293 revolution in, 3 application to “half-castes”, 432 Ewart, J. Cossar, 224, 397, 444 724 Index

Bateson’s disparagement of, 84 Serajevo murder, 409 on Evolution Committee, 159 The Central Powers, 594 exceptions USA joins, 443 study of, xxiv, 644 fish to be treasured, xi, 311, 361, 381, 451, 665 bait for, 46 experimental station proposal coloration of, 76 at Down House, 157, 189 irradiation of, 132 at Wisley, 190, 214 Fisher, R. L, 224, 247, 470 at Woburn Abbey, 186 alliance with L. Darwin, 629 Bateson not supportive of, 161, 395 appointed to Chair in genetics, 657 Davenport’s, 225, 664 his role in SEB, 589 Hurst’s, 395, 396 on dominance, 370 eye color in humans, 276, 289, 302, 412, 425 on eugenics, 425 on Mendel’s data, 652 F Flahault, C, 595 Fabian Society, 39 Floral Morphology (Saunders), 654 factor fluctuations ambiguous use of term, 249, 288, 311 are not inherited, 70 as a “gene”, 445 distinction from mutations, 334 as a gene product, 249 Focke, W, 201 as an element, 250, 281, 330, 410 Follow the Rabbit-Proof Fence (Pilkington), as unit transmitted by gamete, 249, 272, 432 273, 297, 322 Ford, E. B, 5, 630, 649, 711 correspondence with a character, 297 Forsdyke, D. R Haldane’s interpretation of, 322, 333 web-pages of, xvi, 673, 686, 712 latency of, 326 Forster, E. M, 9, 385 may be positive or negative, 515 Foster, M, 11, 17, 22, 34, 162 Shull’s definition of, 289 advice on candidacy from, 84 with its complement inhibits meiosis, recruitment of Hopkins by, 230 331, 405 role in Quick Bequest decision, 215 Fanny’s First Play (Shaw), 548 support for women’s education by, Farabee, W, 243, 299 607 Farmer, J, 39, 234, 341, 381, 387, 413, 627 support of Bateson’s travel by, 22 ferment. See enzymes fowl, 275 fertility an egg’s way of reproducing itself, 535 impairment of. See hybrid sterility Andalusian, 250, 251, 256, 270 of different social groups, 426, 435 inheritance of comb types, 250, 324 fertilization sexing of newborn of, 495 as union of haploid chromosome sets, 268 Fowler, C. Herbert, 245 chromosomes remain separate during, 368 France, 177, 200, 229, 295 specificity of, 529 in World War I, 338, 393, 407, 460 two steps in, 324 fraud in science, 300, 490, 567, 574, 578, Fertilization of Orchids (Darwin), 554 582, 586, 630 Field, Reverend T, 40, 181 Freeman, D, 635 finger-prints, 96 Froude, E, 386 First World War, 268 fruit fly, 141, 445 declared, 413 four chromosomes of, 446, 473 German scientists’ manifesto, 592 four linkage groups of, 446 mobilization for, 247, 439 genetic assimilation in, 573 pacifists, 440, 592 gradations in eye color, 448, 515 preparations for, 388, 396 technology of their breeding, 371 roles of contractors and publishers in, white eye mutant of, 357, 371 434, 440, 465 Fry, R, 216, 392 Index 725

G correspondence between gene and Gadow, H, 577, 604 enzyme proposed by, 230 Galton Laboratory, 440, 633 friendship with Hopkins, 230 Galton Lecture Gärtner, C. F. von, 201, 204, 234 by Bateson, 433, 437 Gates, R. R Galton, F, xii, 49, 280, 307, 319, 367 dispute with Bateson, 413, 470, 515 advocate of biometrics, xxii, 140 role in SEB, 588 as a “Mendel”, 644 Geddes, P, 128 as consulting editor, 169, 222 gemmules, 56, 69 chairman of RS Committee, 156, 646 as “fragments” or “particles”, 343, 528 curve of error, 82, 103, 104, 223 as pangens, 510 death of, 421 capable of “self-division”, 528 dedication of paper to, 71 combinations of different types of, 210 derivation of Mendal’s ratios by, 210 combinations of same type of, 201 his opposition to Bohemian elements, 434 each run by a memory, 528, 538 Huxley Lecture of, 419 in brain, 58 insect studies of, 370 named by Darwin, 58, 527 latent and patent characters of, 50 rabbit transplant experiments on, 58 on “organic units”, 529 gene review of Materials by, 96 allelic and non-allelic, 250 rolling of rough stone, 95, 636, 653 as “ultimate factor”, 325 Savile Club meeting, 102 as a phenomenon of arrangement, 287, support of women’s education by, 604 288, 325, 410 supported by Meldola, 328 as a segregable unit, 322 tribute to Butler by, 553 as an unknown physiological unit, 207 unreliable data of, 302, 316 as superficial, not fundamental, 285, gamete 339, 364, 365, 504 aberrant, 209, 211 as ultimate indivisible fragment, 506 constant size of, 51 correspondence with enzyme, 230 differentiation of, 168 dosage of, 369 equal numbers for each allele, 237, 298, homeotic, 94, 663 359 homology needed for recombination, 346 purity of transmission of, 203, 235, 342, Johannsen’s definition of, 286, 504 360, 362, 363 linkage on chromosome, 236, 375 that fails, 172, 626 mapping of position, 447 type, 209 meristic, 94 gametogenesis, 235 name suggested, 4, 286, 510 impaired by non-homology, 346 pairing with its allele, 343 impaired when hybrids are sterile, 211 questions concerning, xii, 237, 285 loss of chromosomes during, 269 rearrangement due to recombination, 237 rejuvenates cell structures, 506 regulatory, 299 symmetry of division in, 237, 341, 359 selfish, 60, 97, 130, 272, 466, 506, 534, treats “power to produce” as a unit, 325 See individuals as transitory treats characters as units, 237 William’s definition of, 506 Gardiner, J. S, 180, 273, 305, 307, 498, General Stud Book of Race Horses 577 (Weatherby), 277 Gardiner, W, 37 generation Garnett, C, 39, 390, 657, 662 F1, 208, 230 Garnett, D, 39, 526, 657 F1 does not breed true, 256 Garnett, E, 39, 657 F1 of uniform type, 239 Garnett, R, 39, 526, 657 F1 sometimes breeds true, 260 Garrod, A F2, 208 chemical sports of, 402 F2 of non-uniform type, 250, 251

726 Index

Generelle Morphologie (Haeckel), 85 militarism, 393, 407, 434, 460 genetic scientific journals, 440 assimilation, 573 scientists repatriated, 413 disease, xiii, 230, 349, 424, 427, 429, translations, 408, 476, 539, 544, 577 630 treatment of Czechs, 562 fitness, 532 Germany, 263, 320, 443 inequality of individuals, 465 analine dye industry of, 309, 585 information, 89, 667 exclusion from IUBS of, 595 linkage, xviii, 209, See character linkage Giekie, A, 274 relations, 86 Gilman, D. C, 17, 224, 225, 664 Genetical Society, 143, 469, 496, 498 Glasgow, 316 its near demise, 588 Godlewski, E, 365 role of Punnett in, 657 Godman, F. D, 159 genetics Godron, D. A, 509 as including development, 215, 589 Goethe, J. W. von, 85, 407 as name of new science, xviii, 248 Goldschmidt, R, 94, 413, 415, 476, 507, 511 as the physiology of descent, 248 as Bateson’s intellectual heir, xiii, 649 as the physiology of reproduction, 143, praised by Gould, 636 590 visits England, 470 classical, 212, 657 gonad developmental, 470 as seat of the germ-line, 56 does not define a nation, 465 as site of meiosis, 60 does not sanction eugenics, 424, 427, entry of gemmules into, 58 430 finite storage capacity of, 55 of populations, 470, 506, 650 maldevelopment of, 211 struggle for recognition of, 230, 304 with hybridized gemmules, 201 Genetics and Man (Darlington), 632 Gordon, L. W, 6 genic Gould, S. J, xvi, 638, 711 constituency, 53 critic of Bateson, 568, 636 juggernaut, xiii, 509 critic of Koestler, 568 paradigm, xii spandrel metaphor of, 330 genome, 286, 515, 635 supporter of Goldschmidt, xiii as a chromosome set, 256 Government Grant Committee, 22, 161 phenotype, 672 grandparent sequencing projects, xiii reversion to, 201, 363, 533 genotype, 50, 285, 326 Grassi, G. B, 586 as parental information, 171, 528 Gregory Bateson. The Legacy of a Scientist as total information in DNA, 286 (Lipset), xiv differences in impede chromosome Gregory, R. P, 214, 231, 375, 382, 390, 413, pairing, 454 454, 457, 461, 467 fundamental part does not segregate, 504 Guaita, G. von, 229 Johannsen’s definition of, 286, 491, 515 Gulick, J, 133, 293, 651 meiosis as disrupter of, 506 ally of Romanes, 129, 144 of beans, 71 attack on natural selection by, 111 only superficial part of segregates, 505 founder postulate of, 144 Shull’s articulation of, 71, 287, 288 on random drift, 106, 330, 404 term introduced, 286 on reproductive isolation, 637 George, D. Lloyd, 394, 443 Guyer, M, 296, 456, 472, 473 germ cell. See gamete chromosome doubts of, 365 soma communicates to, 544 chromosome studies of, 212 German early citation by Bateson, 339, 360 education of children, 441 Mendel’s laws derived by, 362 governess, 242, 295 substratum postulate of, 364, 672 liberal party, xi Gwynne-Vaughan, G, 489 Index 727

H theory of. See particulate theory of Haeckel, E, 12, 19, 85, 190, 307, 529, 532, heredity 535 transmitting agency in, 203, 237, 238, Hagedoorn, A, 561 287, 325, 353, 362, 412, 449, Haldane, J. B. S, 247 451, 455, 475, 477, 528, 538, Bateson Lecture of, 322 632, 667 editor of Journal of Genetics, 338 Hereditary Genius (Galton), 545 his role in SEB, 588 Heredity in Poultry (Punnett), 492, 657 linkage experiments of, 337, 375 Heredity in the Light of Recent Researches obituary of Bateson by, 628 (Doncaster), 383 on dominance, 370 Heribert-Nilsson, N, 462, 503 on industrial melanism, 630 Hering, E, 19, 57, 443, 521, 528, 544, 644 rule of, 268, 469, 471 hermaphrodism, 205 Haldane, N. See Mitchison, N Herringham, C. J, xxiii, 9, 215, 278, 296, 307 Hall, D, 394, 460, 461, 507, 655 her role as intermediary, 180 Hamilton, W, 60 research support from, 215 Handbook of Invertebrate Zoology schizophrenia of, 384 (Brooks), 18 strange letter from, 383 haploidy, 341 travels with Bateson, 173 Happisburgh, 244, 296, 386, 388 Herringham, W. P, 173, 222, 349, 384, 460 Hardy, G. H, 289, 504, 592, 597, 615, 652 Hertwig, O, 327 Hardy, W. B, 190, 492 Hertwig, R, 367, 476 Hardy-Weinberg equilibrium, 289 heterogametic sex Harmer, S, 12, 29, 36, 263, 304, 586, 607 as “heterozygous sex”, 354 Harriman, M, 225 distinct from homogametic sex, 348, 354 Harrison, A. E, 6 heterozygote Harrison, J, 39, 82, 527, 664 as allelomorphically mixed, 208, 285 Hartog, M, 549, 646 forms two dissimilar products, 475 Harvard University, xv, 300, 371, 633 no distinction from homozygote when Harvey Road, 13, 24, 26, 30 dominance complete, 369 Harvey, W, 633 heterozygous sex, 335, 471 Hawaii. See Sandwich Islands Hickson, S, 241, 301 Heape, W, 12, 37, 59, 91, 159, 222 hierarchical height agencies in evolution, xvi as continuous trait, 223 levels of information, 667 as discontinuous trait, 223, 255, 258, 314 Hilaire, Geoffrey St, 627 distribution in a population, 63, 223 Hill House Hotel, 244, 296, 311, 327, 388 regression analysis of, 66 Hill, A. V, 14 Heitland, W, 8, 25, 523, 660 Hill, B, 543 hemophilia, 234, 349 historians Henig, R, 639 biases of, xxv Henking, H, 347 difficulties of, xiii, 586, 634, 638, 643 Herbert, D, 137, 201 silence of, 553 heredity history actuarial side of, 420 and popular media, 639 as case of homotyposis, 168 and scientific research, xi, xii, xvii, 535, as continuity of form, 239 560, 623, 634, 639, 643, 646 as memory, 528, 536, 544, 551, 663 depends on conservation, 272 as transfer of information, 3, 521, 526, 532 its role in education, 442 cytoplasmic. See cytoplasm retrospective vindication by, 281, 631 knowledge of as a social force, 423, 424, Whig, xviii 426 Hitler, A, 631 legal origin of the term, 223 Hogben, L, 143, 231, 498, 588

728 Index

Holmes, E, 552 misrepresented by Darwin, 660 Homer, 441, 527 proposal for discontinuous variation, 89 homotyposis, 66, 164, 373 saw sterility as the weak spot, 115, 130, homozygote 329, 330, 483, 509 as allelomorphically pure, 209, 241, 284 support of Bateson by, 22 forms two similar products, 475 support of the MBA by, 44 Hooker, J, 89, 307, 327, 612 support of women’s education by, 604 Hope Professorship, 149, 471 hybrid inviability, 119, 121, 331, 348, 456 Hopkins, F. G, 230, 391 hybrid sterility, 119, 140, 405, 467, 532 Horace, 308 as a non-Mendelian phenomenon, 212, Horticultural Society of New York 358, 483 Conference on Plant Breeding (1902), as facilitator of divergence, 115, 125, 224, 270, 341, 362 211, 330, 483, 509 Hudibras (Butler), 546, 654 as weak spot in Darwin’s theory, 509 Hurst, C. C, 229, 241, 251, 255, 461, 561, cured by polyploidization, 256, 454 563, 624 defines a “physiological species”, 115, as Johah, 302 269, 510 enters Cambridge Ph.D. program, 496 discord with classification, 269, 516 first appearance at RS, 277 due to non-homology, 346 fowl, genetic studies of, 495 due to summation of variations, 487 his precision of terms, 249 Haldane’s rule for, 470 letters from, xxi, 212, 265 indicates “remoteness of kinship”, 359 lost manuscript of, 299 inversely related to conjugative power, on chromosomes, 357 456, 626 on eugenics, 425 is Mendelian when sporadic, 358 on eye colors, 301 linked to failed meiosis, 141, 212, 331, on sex determination, 356 342, 626 persecution of, 654 hybrid vigor, 81, 201, 371, 456 rabbit experiments of, 273

rabbit skins of, 243, 274 renaming of Burbage Nursery by, 395 I role in Mendel Society, 336, 356 Ibsen, H, 546 rose cytogenetic studies of, 496 Iltis, H, 561 shielding role of, 265, 423 Imperial College of Science, 4, 25, 307, war-time duties of, 439 507, 588 Hurst, R, 212, 266, 471, 691 Inborn Errors of Metabolism (Garrod), 230 marriage of, 496 inbreeding meeting with Saunders, 498 defective offspring results from, 371, 456 Hutchinson, G. E, 577 India, 338, 384, 413, 611 Huxley, J, 5, 438, 449, 469 individuals appointed to Zoology Society, 632 as members of a population, 52 attack on Bateson by, 632 as transitory constellations of genes, collaboration with Haldane, 651 466, 506 criticized by Bateson, 472 correlation between, 66 his “eclipse of Darwinism”, 632 correlations within, 67 his “modern synthesis”, 632, 638, 650, latent and patent parts of, 50 668 information his role in SEB, 588 analog or digital, 450 misrepresentation of Bateson by, 632 conflict between forms of, 668 Huxley, T, 4, 75, 241, 327, 612 cultural transmission of, 145 advice on dealing with Butler, 540, 550 for a character, 53 advice on pangenesis from, 510 hereditary, 90, 366 advice on public speaking from, 485 hierarchical levels of, xvi, 667 attack on Romanes by, 129 manipulation of for propaganda, 464 Index 729

mental, 90 Issayev, W, 513 preservation of, 90, 369 Italy primary, as content, 126, 667 Butler’s annual visit to, 525 secondary, as accent, 126, 667 delight of, 174 template for, 90 Ithaca two levels of, 60 and “the corn men”, 480 when stored is “memory”, 506 International Congress of Genetics at, infusoria. See organisms, unicellular 631, 655 inheritance alternative (non-blending), 198, 203, 534 J as continuity of gene not organism, 506 Jackson, H, 611 biparental, 365 Jackson, L, 542, 624 blending. See blending inheritance Janssens, F, 446 epigenetic transgenerational, 453 Jenkin, F, 60, 533, 535 integral, 200, 206, 533 Jenkinson, F, 604 legal origin of the term, 223 Jennings, H. S, 287, 448 non-Mendelian, 365, 412 Jodrell Chair in Zoology, 11, 102 of acquired characters. See Lamarckism Johannsen, W, 329, 397, 475, 491, 508, 511 instinct attacked by Muller, 632 added to character list, 321 chromosome doubts of, 491, 505 due to “hereditary memory”, 536 his non-genic “Grundstock”, 367, 504, due to “ready-formed information”, 536 672 due to “unconscious memory”, 526 introduces new terminology, 4, 286 examples of, 536, 552 on Bateson’s terminology, 507 simplification of, 428 pure lines of, 70 transfer of by gemmules, 58 visit to England by, 493 interaction John Innes Institute, xiv, 40, 293, 460 epistatic, 241, 249, 298, 326 Bateson’s appointment to, xxiv, 381 hypostatic, 249, 298 Bateson’s successor, 655 involving distinct allele pairs, 251, 297 local discontent, 392 involving one allele pair, 251 meeting of Genetical Society at, 470, laws for, 322 498 of gene products, 249, 251 microscopist appointed, 489 intermediate forms Norwich, xv, 19, 547, 643, 711 as a series in time, 92 studentships at, 383 as a series of combinations, 411, 518 Johns Hopkins University, xiv, 327 as evolutionary intermediates, 65, 89, Jones, H. F, 101, 539, 543, 549 92, 95, 138, 139, 411, 448 Jones, Lewis, 470 characters masked in, 50 Jordan, K, 470 in linear evolution, 117 Journal of Genetics International Congress of Medicine, 429 editorship of, 657 International Congresses of Genetics, 200, establishment of, 337 249, 397, 407, 437, 591, 593, 631 Jurassic Park (Crichton), 117 Intracellular Pangenesis (de Vries), 68,

235, 322, 341, 346, 491, 510, 646 Introduction to the Study of Cytology K (Doncaster), 448 Kammerer, P, 301, 404, 477, 561 isolation address at Cambridge by, 501, 577 due to a barrier, 134, 403 cave newt studies of, 572 geographical, 64, 98, 106, 117, 121, 127 first communication from Bateson, 573 of fit from fit, 122 first meeting with Bateson, 562 reproductive. See reproductive isolation green fingers of, 569 temporal, 122 midwife toad studies of, 571 without adaptation, 133, 333 military censor in World War I, 462 730 Index

offered post in Russia, 514 language rapid experimental success of, 573 accent in, 123, 667 salamander studies of, 569 alternatives to the classical, 612 sea-squirt studies of, 572 change in meaning of, xxv suicide of, 514, 567 English, 180, 286 visit to USA by, 579 evolution of, 667 Kantsaywhere (Galton), 553 first, grammar second, 616 Karkaralinsk, 28 French, 242, 290 Kassarjian, M. C. See Bateson, M. C German, 68, 242, 291, 295, 329, 592, Kazakhstan, 20 646, 701 Kazalinsk, 24, 34, 80, 131 Greek, 179, 442, 444, 610 Keilin, D, 643 its power to conceal, 286, 663 Kennedy, B, 183, 523 Kirghiz, 26 Kenyon, F. G, 444 Latin, 442, 444, 610 Kettlewell, B, 5, 630 of medieval abuse, 170, 222 Kew Gardens, 96, 192, 269, 381, 382, 390, origin of, 667 473, 623 Russian, 26, 511 Keynes, G, 13, 14, 213, 392, 543, 711 Lankester, E. R, 11, 307, 320, 327, 509 Keynes, J. M, 14, 392, 511, 588 Bernard Symposium, 191 Keynes, J. N, 13 critic of A. Schuster, 592 Keynes, M, 13, 14, 543 critic of Bateson, 244, 308 Killby, H, 213, 335 critic of de Vries, 308 King’s College, Cambridge, 9 critic of Haeckel’s perigenesis, 530 King’s College, London, 101, 588 critic of Mendelism, 467 Kingsley, C, 4, 536 critic of Romanes, 128, 308, 467 Klebs, E, 587 critic of Weldon, 155 Koestler, A, 567, 623, 649 Marine Biological Association, 45 Kölreuter, J. G, 201, 204, 234 on education in science, 444 Krause, E, 539, 543, 547, 660 role in Quick Bequest decision, 215 Kruschev, N, 566 signed memorial to, 84, 88 Kusano, T, 392 supported by Meldola, 328 supporter of Caldwell, 35 L supporter of Poulton, 128, 308 Lake Balkhash, 22, 27 supporter of Wallace, 128 Lakin, Miss F, 9, 242 Larcher, D, 384 Lamarck, J. B, 3, 85, 521, 535, 567 Larmor, J, 226, 604 Lamarckism, 86, 113, 308, 371, 531 Law of as epigenetic inheritance, 453 Ancestral Heredity, 67, 221, 228, 269, colonial insects contradict, 534 323, 370, 420 eliminated from pangenesis, 510 Error, 81 giraffe example of, 537 inheritance, 190, 203 mate location facilitated by, 533, 543 interaction, 322 opposed by Bateson, xxiv, 321, 329, probability, 362 443, 575 recapitulation, 85 revived by Butler, 521, 530, 550 Regression, 66 revived by Darwin, 58 von Baer, 86 revived by Haeckel, 530 Law of Heredity (Brooks), 663 revived by Kammerer, 567 Lawrence, W, 424, 508 revived by Koestler, 650 Leicester, 302 revived by Lysenko, 650 Literary and Philosophical Society, 272 revived by Semon, 404 Leidy Lecture, 90, 475, 489 revived by Spencer, 429 Leighton, G, 265 Lane, B, 659 Lenin, V, 512, 631 Lang, A, 397 Lesch, J, xiii Index 731

Letters between Samuel Butler and Miss E. meeting (1884), 541 M. A. Savage (Keynes), 543 meeting (1886), 26, 121, 131, 645 Letters from the Steppe (Bateson), xiv, 24 meeting (1904), 231, 273 Lewes, G, 536, 544 meeting (1923), 577, 578 Liberal Party Lipset, D, xiv, xxi, xxiv, 553, 629, 711 Bateson a supporter of, 262, 433 Lister, J. J, 44, 91, 188, 247, 303, 393 Cambridge elections, 14 Littlego examination, 10, 176 Marylebone Women’s Association, 30 Liverpool, 6, 295 vote for Lloyd George, 443 School of Tropical Medicine, 586 women’s section of, 7 Lloyd, A. See Dew-Smith, A library Lloyd, R. E, 662 Bateson’s, 19, 547, 554, 643 Lock, R. H, 214, 232, 242, 382, 423 metaphor for collection of genes, 399 comment on Erewhon by, 313 of British Museum, 39, 525, 526 early writers dismissed by, 643 Liebig, J. von, 524 obituary of, 459 life praise of de Vries by, 290 a begger’s, 30, 648 Locke, J, 611, 632, 633 as memory, 538 Lockyer, N, 128, 526 as preserving “selfish genes”, 466 Loeb, J, 296, 367 conferred by macromolecules, 527 Lord cycles of plants versus animals, 205, 367 Alverstone, 429 occult views of, 431 Avebury, 9, 214, 308, 327 study of English, 545 Cavendish, 394 when married, 180 Esher, 386 Life and Habit (Butler), 528, 530, 535, 539, Kelvin, 542 544, 549 Lister, 44 Life and Letters of Charles Darwin, 542 Peckover, 307 Life and Word (Lloyd), 662 Raleigh, 276, 450 Life of Samuel Butler (Jones), 550 Rothschild, 470 light response of Ullswater, 610 cave newts to, 572 Walsingham, 592 crustacea to, 46 Lotsy, J, 250, 260, 397, 411, 413, 451, 474, fish to, 132 561, 596 flagellates to, 459 Lubbock, J. See Lord Avebury plants to, 79 Luck or Cunning (Butler), 539, 542, 629 salamanders to, 570 Lutz, A. M, 256 limits Lyell, C, 85, 667 of a species, xxiv, 60 Lynch, R. E, 22, 154, 387 of education. See education Lysenko, T, 566 of fruit fly experiments, 467 ascendence of, 650 of Mendelian analysis, 285, 322, 416, interrogation of Vavilov by, 514 495, 514 of natural selection, 234 of recombination, 326, 412, 515 M of segregation, 474, 515 MacBride, E, 39, 509, 575, 577, 608 of statistics, 171 MacDougal, D, 142 of the genome, 515 MacDougal, W. T, 404, 477 of variation, xxii, 139 Madame Bovary (Flaubert), 460 on human population size, 437 Magnus, Sir P, 459 undetermined, 495, 517 Maitland, F, 8, 13, 263, 611 Linacre Professorship, 84, 102, 245 Major Barbara (Shaw), 545 Linnaeus, C, 85, 191, 639 malaria Linnean Society attributed to “Bacillus malariae”, 587 celebration, 307 Leveran’s discovery of agent of, 587

732 Index

life cycle of the agent of, 586 Maupas, E, 367, 549 link with mosquito, 586 Mayer, A, 225, 296 Malthus, T, 426 Mayo, J, 606 Man and Superman (Shaw), 423 Mayr, E, 644 Manchester, 49, 137, 163, 228, 386 attack on Bateson by, 5, 633, 641 Literary and Philosophical Society, 246 disdain for “typological” thinking by, Marine Biological Association, 44 668 marine biological laboratory misrepresentation of Bateson by, 633 Chesapeake Bay, 17 McClung, C, 235, 347, 359, 369, 481 Naples, 17, 101 Mead, M, xiv, 556, 635 of Romanes, 59 Mechanisus und Physiologie der Plymouth, 44, 102 Geschlechtsbestimmung Woods Hole, 296 (Goldschmidt), 477 marketing medal of genius, 545, 662 Darwin, 142, 174, 190, 242, 492, 586, of scientific ideas, xii, 565, 632, 644 657 Marquis of Salisbury, 12, 111, 144, 542 from RHS, 249, 397 marriage nomination process for, 492, 586, 644 and property, 175 Royal, 469 arranged in Westminster, 427, 430 meiosis, 60 as a mixer of nationalities, 442 disjunction in, 340, 376, 447 as happily enduring, xx, 180, 393 equational division in, 340 choice of partner in, 420 homo- and heterotypic divisions in, 341 college fellows prohibited from, 4, 523, named, 341 603 postulate of two kinds, 298 of Beatrice and William, 180 reduction division in, 235, 344, 349, 359 of Edward Bateson and M. Corbett, 393 melanism in moths, 112, 163, 472, 630 of G. Bateson and Margaret Mead, 556 Meldola, R of Hurst, 281, 496 Centenary book reviewed by, 328 of Lister and Marryat, 188 member of RS Committee, 157 of M. Darwin and G. Keynes, 13 on the aniline dye industry, 309, 585 of Punnett and Froude, 386 resignation from Evolution Committee, of Wheldale and Onslow, 459 161 of William Henry and Anna, 5 Savile Club meeting, 102 Marryat, D, 188, 214, 335, 390, 472 meme concept, 702 Marshall, A, 177, 426 BDM, 637, 644 Martin, H, 6, 17 Romanes’ anticipation of, 145 Marx, E, 39 memory Masters, M. T, 154, 159, 161, 187 as a deep mystery, 551 Materials for the Study of Variation as a mental rhythm, 544 (Bateson), xxii, 92, 170, 545 as stored information, 90, 526 mathematics. See statistics between generations, 525 alliances with biology of, 629, 651 its location in a cell, 505 applied to geometry of life, 402 modes of, 538 as queen of the sciences, 232 runs its gemmule, 528, 538 coaching in, 10 unconscious, 443, 526, 530, 551 failure to see context in, 109, 168, 479 Men and Memories (Rothenstein), 549 modelling in, 24, 66, 504 Men and Women’s Club, 39 of patterns in biology, 652 Mendel Journal, 336 of probability theory, 108 Mendel Memorial Symposium, 566 senior wrangler in, 178 Mendel Society, 336, 508 Mathew, P, 643 Mendel, G Matsui, K, 392 as a liberal, 559, 563 Matthews, L. H, 577 celebrations of, 562, 564, 576 Index 733

character of, 565 kaleidoscope as, 255, 256 death of, xi military, 56 delay in recognition of, 204, 281, 559 of bottles in medicine chest, 313 genius of, 394 of descent of property, 223 his data questioned, 203, 206, 630, 652 of dog leash, 672 his disagreement with Darwin, 559 of genes as books, 399 his reading of Darwin, 524, 559 of motion as evolution, 24 laws of, xviii, 204, 228, 231, 271, 342, of organism as bookcase, 399 344, 362, 445, 534 of ships at sea, 455 nephews of, 559 political, 52 rediscovery of work of, xi, 197, 646 Metchnikoff, E, 327 shadow of, xii, xxv, 649 Metz, C. W, 446 thread where he dropped it, 204, 665 mice translation of, 203 amputation of tails of, 568, 572 Mendel’s Principles of Heredity (Bateson), coat color of, 232 xii, 244, 321 Japanese waltzing, 228, 244 Mendel’s Principles of Heredity. A Defence Michurin, I, 514 (Bateson), 207, 222, 270 microscopy Mendelians difficulties of, 496 as transmission geneticists, 470 histology, 234 attacked by Poulton, 309, 648 limitations of, 297, 366, 527 attacked by Wallace, 308, 648 of chromosomes, 353, 478 Bateson school of, xxiii, 212, 265 of germ cells, 268 dispute with Biometricians, 102, 221, reveals cell as fundamental unit, 508 232, 240, 276, 314 microtome, 17 in France, 229, 232 automatic, 35, 653 in Germany, 329, 407 development of, 467 in USA, 226, 232, 267 sections of nuptail pads, 576 their alleged mathematical naivete, 226, midwife toad, 501, 567, 569, 570, 575 241, 629 Miers, H, 511 Mendelism mimicry. See copying a red herring, 309, 649 aggressive, 147 alleged union with Darwinism, 632 at school, 422 applicable to quantitative traits, 223, disguising mediocrity, 433, 444 258, 475 driven by fashion in humans, 464 as theory of heredity, not origin, 416, in butterflies, 385 482, 628 protective, 148 as transmission genetics, 502 Mimicry in Butterflies (Punnett), 149, 386 concerns superficial varietal characters, Mind and Nature (G. Bateson), 668 211, 504 Minot, C. S, 241, 273 first mentioned in lectures, 140 Mitchison, N. M, 221, 337 fundamental questions in, 141, 339 mitosis, 68, 339 opposed by Biometricians. See Mivart, St. G. J, 19, 529, 663 Biometricians model opposed by Lankester, 467 BDM, 637 opposed by Reid, 423 mathematical, 66 opposed by Rolfe, 336 Russian doll, 415 Mendelism (Punnett), 289, 312 wax, 90, 366, 528, 685 Mental Evolution in Animals (Romanes), Montgomery, T, 343, 347, 367, 449, 455 536 Moore, A. R, 370 Merton House, Grantchester, xxiii, 187, Moore, J. E, 341 328, 386, 389 Moore, Mr, 153 metaphor More Letters of Charles Darwin, 81 Galton’s use of, 50, 93, 636 Morgan, C. L, 111, 132, 148, 509 734 Index

Morgan, J. P, 216 located at point on chromosome, 414, Morgan, T, 4, 17, 190, 373, 408, 421, 443, 507, 626 451, 475, 511 of organic compounds, 89 at Genetical Society, 498 reversion as back, 415 begins breeding fruit fly, 357, 371 sporadic. See sporadic chromosome doubts of, 372 synthetic lethal, 209, 232 distrusted by Dobell, 490 mutationists, 5, 260, 308, 635 eureka moment of, 374, 560 obituary of Bateson by, 625 N on eugenics, 437 Nachtsheim, H, 593 on Kammerer, 580 Nägali, C. von, 201, 565, 647 school of, xxiv, 445 National Poultry Institute, 495 view on factor terminology, 312 National Union of Scientific Workers, 598 visit to, 478 Natural History Museum, 184, 461 morphology, xxii, 11, 18, 37, 372, 507, 509, natural hybrids, 268 560, 590 Natural Inheritance (Galton), 49, 59, 62, 65, Morrell, Lady O, 40 68, 95, 223, 370, 545, 672 mosaicism natural selection genetically transmissible, 450 and bell curves, 64 germ-line derived, 210 as “classical Darwinism”, 23, 541 somatically-derived, 210, 239, 275, 345, as an isolating agent, 121 377, 475 as major agency, xvi, 130, 234, 244, Moscow, 22, 511 328, 625 Moseley, H, 84, 128 assisted by intelligence, 541 Moulton, F, 651 attack on, xxii, 64, 81, 91, 96, 111, 133, Mousliketeff, M, 23 199, 329, 400, 473, 516 Mudge, G. P, 314, 315, 336, 390, 425 cessation of, 111, 143 Müller, F, 12, 85 correlation with target of, 155, 406 Muller, H, 4, 293, 448, 481 Darwin’s caveat on, 4 emigration to Russia, 513, 631 Darwin’s delayed publication on, xxv finds genes on chromosome 4, 375 defeated by philanthopy, 423 role in BDM, 637 does not give a physiological species, Museum of Zoology, 184, 217, 305, 321, 510 383, 574 many steps in, xxii, 411, 481 museum specimens, 103, 109, 148 melanism as example of, 5, 163, 630 Musset, A. de, 546 not understood by “typologists”, 668 mutation of fit over less fit, 119, 121 as a discontinuous origin, 258, 416, 515 pre-Darwinian proposers of, 535, 643 as a disintegration, 411 statistical analysis of, 151 as a germinal disturbance, 414 Wiesmann’s acceptance of, 111, 542 as a pathological accident, 239 young are more susceptible to, 532 as addition or subtraction, 330 Nature as another word for “variation”, 209, anti-Mendelian posture of, 289, 648 445 delay in reporting by, 241 as independent of crossing, 413 Huxley celebration of, 509 as reversible modification, not jubilee issue of, 467 innovation, 285 legally threatened by Butler, 526 as segregation of new gamete type, 239 nature and nurture, 61, 70, 311, 411, 419, as stereochemical rearrangement in 433, 545, 647 nucleic acid, 414 Naudin, C, 19, 141, 200, 201, 204, 533 by radiation, 225, 404, 414, 532 Naval Timber and Arboriculture (Mathew), distinct from fluctuation, 334 643 in bacteria, 414 Nettleship, E, 243, 349, 391, 395 J. Watt as example of, 426 Neurological Society, 243, 349 Index 735

New York, 270, 296, 478, 500, 580 as a physiological phenomenon, 122, New Zealand, 4, 524, 557 509 Newnham College as an avoided problem, 482, 487, 517, Council of, 7, 601 638, 649 fellowship at, 321, 383 as group selection, 635 Mary Bateson Fellowship at, 9, 263 as mystery of mysteries, 308, 376, 635 Newton, A, 12, 33, 197, 296, 608 by polyploid hybridization, 253, 256, Newton, I, 539, 555 467, 496, 503, 646 Newton, L, 711 defects in evidence for, 516 Newton, W. C. F, 471, 489, 497, 588 explained by natural selection, 130, 328, Nietzsche, F, 546 399, 625 Nilsson-Ehle, H, 397, 508, 561, 593 follows changes in chromosomes, 417 Noble, G. K, 567, 581 no Mendelian explanation for, 309, 339, Norwich House, Cambridge, 182, 185, 186 364, 509 nucleus Orr, H. Allen, 637 as pangen reservoir, 69 orthodoxy contains chromatin threads, 510 Darwinian, 28, 668 primary role in heredity of, 267, 339 religious, 4, 432, 532, 560 reproductive isolation within, 497 Osborn, H. F, 263, 299, 327, 480, 484, 492, nuptial pads, 571, 574, 581 509, 510 Nuttall, G, 587 critic of Materials, 100 saw Bateson as “Great Commoner”, 627 O supported by Meldola, 328 Oenothera Lamarckiana. See evening Osler, W, 231, 444 primrose Ostenfeld, C, 440, 454, 459, 461, 593, 597 Olby, R, 646 Outline of Human Genetics and Racial Omega Workshops, 40 Hygeine (Baur), 432 Omsk, 30, 35 ovary Onslow, H, 358, 492 as seat of the germ-line, 55 Onslow, M. See M. Wheldale tranplantation of, 58 ontogeny, 12 ovum Ontology and Phylogeny (Gould), 636 as conserving element in evolution, 18, Orenburg, 77 365 organic stability, 401 as the primordial cell, 547 Galtonian doctrine of, 65, 98, 672 cytoplasm of, 339, 365 oscillations around position of, 103 finite storage capacity of, 55 related to discontinuous variation, 93, 138 neither male nor female, 348 organism structureless, 51, 170 “nature of” as a cop-out, 139 Owen, R, 90, 685 as a set of characters, 69 Oxford University as an aggregate of genes, 517 admission of women by, 609 as product of genes, 204 Boyle Lecture at, 327 inseparable from its characters, 211, 486 H. Spencer Lecture at, 397, 426, 429 intrinsic nature of, 94, 234, 330, 504, See resistance to science of, 84 organic stability Rolleston Prize of, 37 pathogenic, 402, 586 Romanes Lecture at, 129 polarity of, 377 unicellular, 367, 449, 459, 530 P origami, 94 Paine, T, 430 origin of species pangenesis, 56, 58, 291 as a fundamental problem, 91, 139, 199, and selfish genes, 131 211 criticized by Bateson, 504 as a genic problem, 637, 649 criticized by Haeckel, 529 as a non-genic problem, 211, 364, 403 proposed by Darwin, 527 736 Index

research on, 58, 321 Peck, J, 635 updated by de Vries, 68, 646 Pellew, C, 390, 393, 398, 408, 443 pangens beneficiary of will, 624 as “fragments” or “particles”, 343 discovery of rogues, 451 as determiners of characters, 510 role in Genetical Society, 469 differential tissue expression of, 69 Perkins, J. B, 495 no differential elimination of, 70 Pertz, D, 83, 154, 548 strict intracellularity of, 68 phenotype, 50, 171, 200 Pangloss, Dr, 5, 142, 192, 330, 428, 585, blurred distinction from genotype, 204, 636 208, 314 Pankhurst, E, 398 Johannsen’s definition of, 286, 515 panmixia of beans, 71 as unfettered reproduction, 65, 112 produced by macromolecules, 527, 668 coinage by Weismann, 49, 76, 683 sterility-of-offspring, 332 stopped by natural selection, 121 term introduced, 286 parentage Philiptschenko, J, 512 need to ascertain, 103 Philosophie Zoologique (Lamarck), 3, 568 of eminent offspring, 419 phylogeny, 12, 88, 136 reemergence in F2 generation, 323 physiological reemergence in gametes, 360 complements, 123 parthenogenesis, 261, 324, 339 units, 527, 529 particulate theory of heredity, 59, 97, 289, physiological selection, 400 342, 366, 410, 446, 449, 451 attacked by Dyer, 123 Pasteur, L, 215, 226, 371, 394, 459 attacked by Francis Darwin, 126 Paulmier, F, 347 attacked by Huxley, 129 pea attacked by Lankester, 128 crosses of Mendel, xi, 202, 205, 342 attacked by Wallace, 123 Gradus case, 72, 627 in Bateson’s lectures, 140 height as a discontinuous character in, praised by Bateson, 131, 645 223, 255, 258, 283 praised by Cunningham, 132 round and wrinkled, 206, 231, 266 proposed by Romanes, 111, 121 yellow and green, 205, 227, 283, 374 physiology Pearl, R, 247, 387, 421 genetical, 517 Pearson, K, 39, 275, 307, 327, 444 laboratory of Foster, 11, 35, 75 appointed to Biometric Laboratory, 421 national, 422 appointed to Galton Chair, 421 school at Cambridge, 10 collaboration with Weldon, xxiii, 102, pilgrimage 629 1904, 560 dealings with Hurst, 276, 277, 316 1910, 561 development of Galton’s ideas by, 65 planet earth disparagement of brain-power by, 399 age of, 495 dispute with Bateson, 164, 178, 222, as an island, 443 241, 319 plants dispute with Johannsen, 71 chloroplasts of, 509 dispute with The Lancet, 316 distributed potential for gametogenesis, distrusted by J. M. Keynes, 392 450 funding of, 216 early flowering, 122 his error in correlation tables, 280 have memory, 544 Huxley Lecture by, 420 immobility of, 205 on RS Committee, 157 location of germinal tissue in, 374 supported by Meldola, 328 pigments of, 231, 335 Pease, H, 440, 711 self-fertilization of, 205 Pease, M, xiv, 413, 437, 461, 495, 564, suitability for study, 205 576 variegation of, xxiv, 381 Index 737

Plato, 28 Problems of Life and Mind (Lewes), 536 Plymouth, 31, 44, 648 process polyallelism. See polymorphism as either chemical or mechanical, 325, 401 polymorphism, 52, 401, 426, 475, 507, error-prone, 90 516 evolutionary and mental are similar, 528 polyploidy of division, 401 as intracellular twinning, 456 of mutation, 94 F1 may breed true, 256, 646 of scientific discovery, xvii in roses, 496 related to gap between forms, 93, 412, 627 large cells in, 256, 453 Professional Women and their Professions transient tetraploidy in mitosis, 340 (Bateson), 8, 83, 602 post-war proteins boycott of foreign scientists, 592 as “physiological units”, 527 isolation of the Central Powers, 594 as amino acid sequences, 668 poverty, 579 as polymeric macromolecules, 527 restoration of communication, 586, 591, limited versatility of, 20, 527 595 protozoology, 39, 214, 490 post-war reform Provine, W, xiii, xiv, 629, 638, 650 Bateson as IRC delegate, 594 Przibram, H, 561, 562, 569, 580 extra-nationalism, 463, 595 Psychological Club, 59 League of Nations, 463, 599 public opinion role of the media, 464 Huxley’s direction of, 509 the CBSS, 596 marketing of ideals to sway, 463 the IRC, 591, 594 Punnett, R, xiv, xix, 214, 241, 277, 289, the IUBS, 595 307, 309, 355, 376, 399, 615, 654 Treaty of Versailles, 564, 594 appointed to Chair in Biology, 385, 429 World State, 464 appointed to Chair in Genetics, 386 Poulton, E. B, 59, 444, 471, 509 as right hand man, 655 critic of Bateson, 309 awarded Darwin medal, 492 on mimicry, 147 dispute with Poulton, 385 on RS Committee, 157 editor of Journal of. Genetics, 337 praised by Lankester, 128, 308 his data reinterpreted, 375 praised by Wallace, 309 his doubts on SEB, 589 praiser of Weldon, 148 his square tabulation of characters, 289, visit to Ceylon of, 385 421 visit to USA of, 327 on sex of newborn fowl, 495 Poultry Genetics Unit, Cambridge, xiv recruiting of, 216 Poultry Research Centre, Edinburgh, xv role in Genetical Society, 469 Powell, T, 175 sweet pea studies of, 358, 375 power to produce, 19, 299, 312, 325, 335, views on Bateson, 18, 601, 628 405, 406 views on Hurst, 654 Practical Plant Biochemistry (Wheldale), pure lines, 70, 247, 287 477 Pygmalion (Shaw), 423 Prain, D, 381, 391, 508, 594, 596, 598, pygmies, 63 623 as a pure line, 70 prepotency. See dominance presence and absence hypothesis, 237, 250, Q 282, 283, 285, 293, 297, 298, Quakers, 44, 218 398, 415, 475, 506, 517, 633 Quastel, J. H, 577 Previous Examination, 180, 611, 612 Queen Victoria, 512 Principles of Biology (Spencer), 527 Queen’s University Principles of Psychology (Spencer), 536 Archives, xvii, 703, 712 Problems of Genetics (Bateson), xxiv, 399, for women, proposal of, 606 575 Quick Professorship, 214, 248 738 Index

R due to geographical barrier, 117, 121 random drift, 106, 144 due to hybrid sterility, 342 ratio F2 due to intrinsic causes, 122, 211, 333 depends on degree of dominance, 251 due to temporal barrier, 122, 404 in the dihybrid case, 251 first gives a “physiological species”, 509 in the monohybrid case, 205, 209, 210, precedes natural selection, 115, 126 250 proposed by Gulick, 145, 637 irregular in evening primrose, 260 rapid in polyploids, 255, 260 Raverat, G, 13, 422, 461, 473 separates the fit from the fit, 120 Raynor, Reverend G, 335, 371 term used by Dyer, 123 Reading reproductive selection, 62, 64, See breeding recruits from, 390 research Sutton’s Nurseries, 270, 454 accolades for fact or for theory, 654 recapitulation applied threatens pure, 214, 434 as an early Victorian notion, 487 as a way of life, 556 Lankester’s caveats on, 13, 85 assessment of, 623, 644, 664 of phylogeny, 12 avoidance of dogmatism in, 534 recombination, 235 centralization of, 395 as a rearrangement, 204, 239, 261, 411 credit to junior colleagues in, 588 as sole source of variation, 411 delay in correcting errors in, 586, 643, between homologous chromosomes, 644 343, 347, 360, 446 funding of, 47, 159, 166, 198, 216, 225, is absent in male fruit fly, 446 273, 306, 385, 394, 396, 460, mapping by frequency of, 447 664 of whole chromosomes, 344 politically correct path of, 204, 233, 394, product is stably transmitted, 237 436, 565, 623, 644, 648 suspected when linkage is incomplete, practical results of, 45, 394 446 priority of discovery in, 586, 643 within a gene, 235 reductionist approach to, 509 reduplication hypothesis, 357, 373, 377, support for megaprojects, 656 430, 452, 469, 474, 560, 630 switch to new approach in, xix, 20, 36, regression 171, 534 to the mean, 61, 108 the role of genius in, 132, 394, 545 Reid, G. Archdall, 336, 420, 422 residue rejuvenation, 665 as a “base” or “basis”, 211, 326, 474, as basis of sex, 368, 549 484, 486, 517 during gametogenesis, 367, 506 Bateson’s use of the term, 141, 210, 285, involves exchange (recombination), 368 313, 326, 339, 364, 488, 495, relationship 515, 650 circuitous between parent and child, 56, Galton’s use of the term, 51, 210 59, 367 location of, 339, 672 path of is indeterminate, 67 may escape segregation, 474 Remak, R, 12 related to Guyer’s substratum, 364 Renner, O, 503 related to hybrid sterility, 211, 333, repetition 484 as symmetry or likeness, 297 related to Johannsen’s “Grundstock”, of parts, 94, 136, 164, 402 504 reproductive isolation Romanes’ use of the term, 57 as conserver of adaptations, 126 Shull’s use of the term, 285 as part of a tetrad, 120 reviews due to accent differences, 668 obstructive, 274 due to chromosomes themselves, 497 of Materials, 96 due to extrinsic causes, 122 press clippings of, 326

Index 739

Richmond, M, 640 International Congress (1906), 248, 249, rogue 281 as a “paramutation”, 453 Scientific Committee of, 161, 202, 269 as a recessive, 209 Royal Institution peas, xxiv, 451 Fullerian Lectures, 397 that always breeds true, 451 lecture by Bateson, 386, 450, 473 that will not breed true, 270 lecture by F. Darwin, 544 Rolfe, R. A, 154, 187, 269, 336 lecture by Weldon, 101, 104 Romanes, G. J, xxii, 6, 10, 293, 319, 554 Royal Society anticipation of Bateson by, xiii, xxiv Calandruccio Enquiry Committee of, anticipation of Mendel by, 200 586 as a “Mendel”, 644 Committee for Statistical Enquiries, 102, books ignored by Bateson, 72, 142, 157 406 Council of, 225, 586, 592 collaboration with C. Darwin, 76 Evolution Committee. See Evolution criticism of Butler by, 526, 535, 538, Committee of the RS 660 financial support from, xxiii, 22 death of, 647 medal awards of, 492 doubts on female intelligence, 178 Royal Society of Medicine friendship with Galton, 59, 645 debate on heredity, 314 interpretation of Weismann by, 57 Rugby school, xxii, 9, 554 investigation of pangenesis by, 58, 145, Rushdie, S, 439 322, 532, 646 Russell, B, 592 Linnean Society address, 26, 122, 645 Russell, E, 393, 394 move to Oxford by, 84 Russell, J, 469 nicknamed “The Philosopher”, 122, 131, Russia 142, 145 Academy of Sciences Bicentenary, 511 physiological selection theory of, 467 ascendance of Lysenko, 650 second class degree of, 75 literature of, 39 Roosevelt, President T, 299 Muller emigrates to, 631 rose reason to visit, 38 cytogenetics of, 496 shabbiness of clothing, 511 development from seed, 525 Siarus document, 513 Rosenberg, O, 456 Rutherford, E, 327, 496 Ross, Major R, 586, 593, 644 Rothamsted Experimental Station, 382, 394, S 469 Salaman, R, 337, 395, 398 Rothenstein, W Salt Schools, 9, 440, 610, 665 at Centenary celebrations, 327 Salvin, O, 157, 159 friendship with C. J. Herringham, 175 Samuel Butler’s Notebooks (Keynes), 543 friendship with F. Darwin, 549 Sand, G, 546 friendship with H. F. Jones, 549 Sanders, A, 58 his passage to India, 384 Sanderson, J. B, 11 supported by Bateson, 549 Sandwich Islands, 111, 144 Roux, W, 341 Saunders, E. R, 186, 207, 242, 277, 335, Royal Anthropological Society, 419 390, 409, 654 Royal Army Medical Corps, 439, 460 collaboration with Bateson, xxiii, 219 Royal College of Science, 39, 509 first class honors degree OF, 179 Royal Commission inebriation of, 183 on education system, 7 leave fellowship of, 295 on Universities, 609 linkage in stocks found by, 209, 241, Royal Horticultural Society 344, 565 International Congress (1899), 186, 199, misrepresented by Henig, 640 253, 255, 268 RHS medal awarded to, 249 740 Index

role in Genetical Society, 469, 590, 654 sex chromosomes visits John Innes, 393, 499 discovery of, 235, 347 Savage, Eliza M. A, 526, 543, 545, 657, 662 X chromosome, 349, 353, 429, 447 Savile Club, 32, 102, 387, 555 Y chromosome, 447 Schiller, J. C. von, 407, 563 sex determination Schindler, F, 559 a Mendelian phenomenon, 234, 348, Schmankewitsch, W. J, 23, 27, 30, 39, 136 351, 353, 355, 356 Schübeler, F. C, 551 analogous to species determination, 484 Schuster, A, 592 sex-linkage, 446, 448 Schuster, E, 229, 246, 421 as “sex-limited”, 349, 445 Scientific Papers of William Bateson of disease, 234, 349, 429 (Punnett), 657 of eye color, 371 Sclater, P. J, 185 of feather color, 349, 628 Scopes, J, 500 sexual characters, 477 Seaman, O, 398 secondary, 55, 112, 470 sea-squirt sexual reproduction amputations of, 572 destroys the individual, 506 regenerations of, 572 origin of, 367, 532, 549 Sedgwick, A, 17, 30, 34, 35, 37, 38, 39, 319, Shakespeare, 24, 83, 320, 415, 420, 425, 604 432, 465, 466, 529, 554, 701 appointed to Chair at Imperial College, Sharp, D, 91, 184, 193, 319, 321, 493, 586 307 Sharpey, W, 11 appointed to Chair in Zoology, 303 Sharpley, E, 15 as supporter of Weldon, 101 Shaw, G. B, 420 on Anna’s degree, 79 friendship with Garnetts, 39 on John Innes Advisory Council, 381 influence of his plays, 433 on Quick bequest, 214 on eugenics, 423 political skills of, 304, 320 praise of Butler by, 545, 549 review of Materials, 96 Shipley, A, 12, 34, 263, 319 Seebohm, H, 542 BA Address of, 334 seed collaboration with Poulton, 149 as a new individual, 205 Master of Christ’s College, 444 as an arrested developmental form, 205 women’s education resisted by, 608 coat maternally-derived, 205, 223 Shrewsbury School, 3, 521 cotyledon colors, 205, 223 shrimp, 82, 102 germination of, 525 Shull, G, 283, 310 segregation, 474, 489 dominance explained by, 369 Bateson’s definition of, 287 editor of Genetics, 376 defines ultimate units, 322 supporter of Johannsen, 71 location of, 239, 345, 373, 430, 452 Sidgwick, Eleanor, 12, 44, 179 mechanism of, 313, 325, 363, 449 Silliman Lectures, xii, 263, 295, 300, 389, of characters, 212, 365, 449 399 of Galtonian elements, 52 Smith, G. C. Moore, 393, 548 time of, 235, 349, 356, 373, 451, 452, Smith, Geoffrey, 337, 356, 457 467 Snow, C. P, 444 Selections from Previous Works (Butler), social Darwinism, 428, 433 538 Society for Experimental Biology, 588, 591 self-fertilization, 205, See asexual Society for Psychical Research, 44 reproduction, See Sociological Society, 232, 420 parthenogenesis Socrates, 9, 506 Semon, R, 145, 404, 544, 549, 551, 702 Sollas, I, 214, 335 Semper, K, 136, 147 soma Sense and Sensibility (Austin), 386 communication with germ cells, 544 Seward, A, 244, 304, 307, 319, 321, 496 disposable, 56 Index 741

formula different from gametic, 274 portrait of Butler, 549 in colonial insects, 534 stewardship of, 75, 176, 185, 306, 601 separate from germ cells, 55, 140, 143, St. Petersburg, 22, 26, 39, 77, 511 568 Stalin, J, 514, 566 Spallanzani, L, 633 Standfuss, M, 471 species statistics, 151, 189, 200, 201, 221, 281 as contained within a sphere, 60 Stebbing, Reverend R. T, 241 definitions of, 119, 191, 211, 510 Stebbins, L, 5 differences between are inherent, 234, Stephen, L, 550 498, 638 Steppes, 20, 29, 38, 77, 131 differences in chromosome number of, Stern, C, 630 353 Stevens, N, 354, 446 differences seldom quantative, 199 stirp theory, 49, 645 discontinuity of, xxii Stopes, M, 389, 413 distinction from varieties, 117, 119, 193, Strange, M, 389 330, 399, 411, 462, 482, 625, Strasburger, E, 234, 307, 324 639 Streatfeild, R. A, 548 induced transformation between, 24 strike of railway workers, 393 sibling, 117, 140, 191 struggle varieties are not steps between, 405 between elements, 97 Species and Varieties, their Origin by between ideas, 146 Mutation (de Vries), 364 for existence, 28, 65, 68, 130, 428 Spencer, H, 128, 366, 410 for recognition of biochemistry, 230 as a Lamarckist, 225 for recognition of genetics, 197, 230, as a philosopher, 85 304 physiological units of, 19, 527 to be understood, xxiv, 565, 623, 645, views on religion, 426 646 spermatozoon Stuart, J, 614 as progressive element in evolution, 18, Studies on the Theory of Descent 365 (Weismann), 309 structureless, 51, 170 Sturtevant, A, 4, 375, 478, 498 Spillman, W. J, 261, 266, 270, 271, 281, suffragette movement, 7, 216, 262, 310, 345 spiritualism, 128, 432, 457 398, 601, 659 sporadic Suharov, C, 513 mutations are statistically irregular, 239, survival of 271 the fittest, 428, 537 sterility as a Mendelian phenomenon, the survivors, 404 358, 405 Sutton, A, 469 sterility versus consistent sterility, 100, Sutton, L, 162 333, 358, 405 Sutton, W, 343, 345, 359, 363 variation is of low frequency, 205, 209 Swingle, W. F, 397 sports symmetry as discontinuous variations, 99 control by, 136 as extreme variants, 64, 97 subverted, 88 bud, 345, 453, 508 types of, 164 often non-viable, 99 Sprunt, A. D, 338 T Sri Lanka. See Ceylon Tebb, F. See Weldon, F St. John’s College technology engagement party at, xx, 44 of DNA sequencing, xvi fellowship of, 20, 75, 228, 386 of insect breeding, 370 memorial service at, 624 of mouse breeding, 228 pillar box removal, 177 of plant breeding, 223 742 Index

of rapid section cutting, 35 The Scientific Basis of Evolution (Morgan), statistics as, 221 626 teleology, 85, 89, 330, 532, 540 The Structure of Evolutionary Theory Tennessee trial of Scopes, 500 (Gould), xiii, 636 testis The Times, 123, 245, 396 as seat of the germ-line, 55, 343 publishes “Oxford Letter”, 593, 597 The Ancestor’s Tale (Dawkins), 636 spawns Daily Mail, 171, 411 The Anthocyanin Pigments of Plants The Variation of Animals and Plants (Wheldale), 391 (Darwin), 91, 201, 213, 510, 554 The Art of Travel (Galton), 21 The Way of All Flesh (Butler), 523, 526, The Burlington Magazine, 216 533, 545, 554, 555, 559, 585, 661 The Case of the Midwife Toad (Koestler), The Wizard of Oz (Baum), 10 567 Theophrastus, 507 The Cell in Development and Inheritance Theories of Development and Heredity (Wilson), 57, 348 (Russell), 492 The Colours of Animals (Poulton), 147 theory of heredity. See particulate theory of The Cotton Plant in Egypt (Balls), 391 heredity The Eclipse of Darwinism (Bowler), 638 Third International Conference on Genetics. The Effects of Cross- and Self-Fertilization See RHS International Congress (Darwin), 201 (1906) The English Illustrated Magazine, xx, 180 Thiselton-Dyer, W, 96, 231, 358, 509, The Englishwoman, 216 542 The Evolution of Genetics (R. Hurst), 496, definition of species by, 191 655 on Evolution Committee, 159 The Evolution Theory (Weismann), 142 praised by Wallace, 309 The Extended Phenotype (Dawkins), 635 praiser of Weldon, 151 The Genesis of Species (Mivart), 530 Thomas, R. Haig, 336, 337 The Genetical Theory of Natural Selection Thompson, D’Arcy, 511 (Fisher), 629 Thomson, J. A, 142, 309, 444, 648 The Grammar of Science (Pearson), 66 Thomson, J. J, 47, 441 The Hour Glass, 8, 26 Thorpe, W. H, 577 The Law of Heredity (Brooks), 18 Threlfall, R., 35 The Mechanism of Creative Evolution Thucydides and Dionysius, 457 (Hurst), 655 Tischler, G, 457 The Midwife Toad (Koestler), 623, 649, Tjebbes, K, 576 650 Toronto, 479, 489 The Monk in the Garden (Henig), 639 Tower, W, 283, 300, 309, 404, 477, 644 The Mutation Factor in Evolution (Gates), trade 413 in art. See art The Orchid Stud Book (Rolfe and Hurst), relapse into, 83 336 scorn for, 10, 44, 433, 436 The Origin of Life (Burke), 261 transmission. See particulate theory of The Origin of Species (Darwin), 4, 10, 126, heredity 129, 213, 233, 406, 524 from parent to child, 50, 66, 203, 223, The Origin of Species Revisited (Forsdyke), 231, 346, 372, 410, 419, 516, xiii, xvi 528 The Origins of Theoretical Population genetics, 502 Genetics (Provine), xiv, 638 of “power to produce”, 325 The Power of Movement in Plants (Darwin), Treasury of Human Inheritance, 421 81 Treviranus, G, 508 The Present Evolution of Man (Reid), 422 Trinity College, Cambridge, 11, 425, 496, The Principles of Heredity (Reid), 422 592 The Problems of Genetics (Bateson), 263 Tschermak, E, xi, 202, 397, 561 The Queen, 8, 39, 83, 602 discovery of Mendel’s work, 197 Index 743

Turgenev, I, 39 dice-throwing model of, 103 Turing, A, 653 discontinuity of, xxii, 82, 98, 139, 199, Turkestan, 22, 26 239, 482 Tutt, J. W, 163 frequent type is due to rearrangement, Tylor, A, 542 209, 239, 402, 410 Tyndall, J, 535, 612 homeotic, 92, 108 type in elements of reproduction, 127 as typical form in a species, 200, 234 in generative elements, 116, 121 in printer’s block, 90 in kind and in degree, 82, 405, 639 in the germinal tissues, 412 U in the reproductive system, 122, 134 ultra-Darwinians, xiii, 111, 144, 626 latent, 116, 137, 200 Unconscious Memory (Butler), 526, 536, meristic, 92, 401 540, 544, 548, 549 of repeated parts, 87 University College, London, 11, 17, 37, 65, originating, 209, 411, 451 101, 141, 247, 391, 421 originating a species, 90, 139, 192, 330, University Correspondence College, 114 483 University Grants Committee, 462, 492 primary and secondary parts of, 32 University of qualitative difference of inter-specific, California, Santa Cruz, xv 650 Chicago, 169, 300, 361 rare type due to nascent mutation, 209, Cincinnati, 363 239 Copenhagen, 70 reversibility of, 27, 38 Edinburgh, xv, 514, 522, 588 scope of, 89 Southampton, xv, 711 selection preceded by, 113 Toronto, 481 spontaneous, 89, 121 Vienna, 569 sporadic. See sporadic Unweaving the Rainbow (Dawkins), 635 substantive, 92, 401 swamping of, 198 V term replaced by “mutation”, 90, 261, Vanderbilt, Mrs, 271 445 variation, 88 that produces hybrid sterility, 483, 486 as a non-inherited fluctuation, 330, 334 transilient of Galton, 97 as a novel cell division, 69, 239, 297, two classes of Bateson, 401 412 two classes of Darwin, 99 as a rearrangment of the preexisting, 451 two factors of de Vries, 69 as an addition or elimination, 137, 330, two types of Bateson, 93 402, 410 utility of, 88 as an error in cell division, 518 when information is not preserved, 90 as an unpacking of the previously within species gives varieties (races), existing, 402, 412, 415, 515 405, 483 as evolution, 75 Variation and Differentiation (Bateson), 169 as fundamental property, 89, 133, 330 Variation, Heredity and Evolution (Lock), at somatic or chromosome levels, 487 xii, 4, 292, 309 bias in, 88 Vavilov, N. I, 392, 511, 552 causes of, 139, 240, 249, 344, 402, 412, Vavilov, S. I, 513 537, 626 vernalization of wheat, 552 collective, 134, 672 Verrall, A, 457 congenital, 98, 145, 645 Verrall, G. H, 279 continuity of, xxii, 92, 448 vestigial. See degenerate organs correlation with other variations, 32, vibratory theory of 113, 155 Bateson, 83, 387, 450, 477, 480, 529, cumulative, 116, 483, 672 633

744 Index

F. Darwin, 544 his “ids” as particles, 97, 343 Hering, 19, 528 immortal germ-line postulate of, 55, 547, Mivart, 19, 529, 663 568 Vienna, 242, 467, 501, 561, 564, 574 Welby, V, 420 Vilmorin, L. de, 70, 200 Weldon, F, 101, 245, 247 Vilmorin, P. de, 187, 397, 548 Weldon, W. F. R, xxii, 12, 36, 420, 604, 664 Virchow, R, 12, 239 analysis of Mendel’s work by, 204 vivisection, 11, 111, 382 as popular lecturer, 46, 242 Voltaire, 5, 142, 386, 661 collaboration with Pearson, xxiii Vries, E. de, 393 critic of Johannsen, 71 Vries, H. de, xi, 6, 10, 68, 225, 256, 366 critic of Materials, 101, 102, 150 as a “Mendel”, 644 critic of Mendelism, 206, 207, 227, 241 attacked by Dawkins, 635 death of, 245, 280, 664 attacked by Mayr, 634 definition of species by, 191 criticized by Bateson, 503 insect breeding by, 371 decoyed by polyploid hybridization, 256 simulation of variation with dice, 104 discovery of Mendel’s work by, 197, supporter of Dyer, 152 646 supporter of Lankester, 84 disenchantment with Mendelism, 260 view of Romanes, 132, 143 dominance explained by, 369 Weldon, Walter, 101 mutation theory of, 141, 403, 415, 467 Wells, H. G, 114, 420, 445, 478, 588, 639 praised by Bateson, 322 Wells, W. C, 643 praised by Lock, 292 Wheeler, Mrs (Emerson), 481 social standing of, 186 Wheldale, M, 213, 295, 305, 335, 502 view on recombination, 346 account of Kammerer visit by, 577 visits Cambridge, 186, 200, 262, 321, appointed to John Innes, 383 329 correspondence with Baur, 321 Vries, M. de, 187, 305, 329 independence of, 391 marriage to Onslow, 459 W pigment studies of, 231 Wagner, R, 14, 95, 408 training with Nierenstein, 389 Wallace, A. R, 49, 307 Whitehead, A. N, 44, 180, 242 critic of Materials, 98 Whitman, C, 361, 366, 472 critic of Mendalism, 648 Wilcox, Miss, 481 definition of species by, 193 Wilks, W, 214, 248, 266, 270 his term “physiological complements”, will of 123 Bateson, 624 key evolutionary questions of, 158 F. Galton, 421 praised Butler, 535 J. Innes, 381 supported by Meldola, 328 T. Powell, 175 Ward, H. Marshall, 618 Willett, Mrs. See Coombe-Tennant, W Ward, J, 184, 310, 552, 662 Willey, A, 385 Washington State Agricultural College, 266 William Bateson, F. R. S. Naturalist Watson, J. B, 580 (Bateson), xiv Watson, W, 386 Williams, Carrington B, 390, 462 Watt, J, 426 Williams, G. C, 60, 506 Weismann, A, 5, 307, 429 Willis, J. C, 261, 291, 502 acceptance of natural selection by, 111, Willmott, A. J, 213, 390 542 Willmott, Miss, 296, 308 address to BA, 49, 341 Wilson, D. Sloan, 635 anticipated by Galton, 59 Wilson, E. B, 4, 17, 57, 234, 289, 296, 321, barrier of, 651 343, 344, 351, 355, 454, 475 compared with Hering, 531 Wilson, H. V, 449

Index 745

Wilson, J, 398, 411, 415 Z Winge, Ö, 416, 454, 508 Zacharias, Frau, 326 women. See Men and Women’s Club Zacharias, G, 393 low-paying studentships for, 383 Zacharias, O, 295 proposed Queen’s University for, 606 Zeitschrift für induktive Abstammungs und right to university education of, 7, 177, Vererbungslehre, 337 601, 704 Zionism, 459 spirit of initiative of, 83, 602 Zoological Congress Wood, T, 214 Boston (1907), 296, 353 words Cambridge (1898), 185 consistency of usage, 272 Zoological Society hormone jargon, 502 move to Regent’s Park, 327 legal origin of, 223 secretaryship of, 184, 225, 632 new coinage of, xviii, 50, 92, 208, 298, support by, 198 419, 507, 508, 510, 663 zygote, 51 Wright, S, 5, 247, 469, 479 Batesonian usage of the term, 56, 68, alliance with Dobzhansky, 629, 651 272 concept of random drift, 106 characters of, 240 concept of shifting balance, 652 developmental arrest of, 205 dispute with Fisher, 370 Wyhe, J. van, xxv

Y Yale University, xii, 263, 314, 389, 481 Yanks, 14, 42 Yule, G. U, 66, 228, 503, 652