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Reginald Crundall Punnett: First Arthur Balfour Professor of Genetics, Cambridge, 1912

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Reginald Crundall Punnett: First Arthur Balfour Professor of Genetics, Cambridge, 1912 PERSPECTIVES Reginald Crundall Punnett: First Arthur Balfour Professor of Genetics, Cambridge, 1912 A. W. F. Edwards1 Gonville and Caius College, Cambridge CB2 1TA, United Kingdom ABSTRACT R. C. Punnett, the codiscoverer of linkage with W. Bateson in 1904, had the good fortune to be invited to be the first Arthur Balfour Professor of Genetics at Cambridge University, United Kingdom, in 1912 when Bateson, for whom it had been intended, declined to leave his new appointment as first Director of the John Innes Horticultural Institute. We here celebrate the centenary of the first professorship dedicated to genetics, outlining Punnett’s career and his scientific contributions, with special reference to the discovery of “partial coupling” in the sweet pea (later “linkage”) and to the diagram known as Punnett’s square. His seeming reluctance as coauthor with Bateson to promote the reduplication hypothesis to explain the statistical evidence for linkage is stressed, as is his relationship with his successor as Arthur Balfour Professor, R. A. Fisher. The background to the establishment of the Professorship is also described. HE centenary of the foundation of Cambridge Univer- the findings of Gregor Mendel when these were brought to Tsity’s Professorship of Genetics in 1912 provides a timely light in 1900. He lived, therefore, in a period that was filled occasion to recall the contributions of its first holder, with excitement and could rightly feel that he was involved Reginald Crundall Punnett (1875–1967; Figure 1). Over- in a great adventure that would surely lead to a revolution shadowed by his senior colleague William Bateson (1861– in biological thought.” Crew’s memoirs should be consulted 1926), for whom the Professorship had been intended, and for details of Punnett’s life; here I concentrate on his his successor R. A. Fisher (1890–1962), Punnett played an scientific contributions and give only a brief biographical important role in the early days of Mendelian genetics. He summary. wrote the first genetics textbook Mendelism (Punnett 1905), collaborated in the discovery of partial coupling (linkage), asked G. H. Hardy the question that led to the formulation Brief Biography of what became known as Hardy–Weinberg equilibrium, Punnett was born to George and Emily Punnett (née published Mimicry in Butterflies (Punnett 1915) and Hered- Crundall) at Tonbridge, Kent, on June 20, 1875. Both ity in Poultry (Punnett 1923a), and pioneered the use of sex- parents were of Kentish stock. He was educated at Clifton linked markers for sexing poultry chicks. He founded the School, Bristol, and Gonville and Caius College, Cambridge, Journal of Genetics with Bateson in 1911 and edited it alone which he entered as a scholar in 1895. Originally registering after Bateson’s death. He was the first Secretary and was as a medical student, he took the Natural Sciences Tripos, later President of the Genetical Society of Great Britain. His specializing in zoology in his third year and being placed in name is immortalized in “Punnett’s square” (Figure 2). the first class in the Tripos in 1898. He spent the next year at F. A. E. Crew (Crew 1967) wrote Punnett’s biographical the Naples Zoological Station (Naples, Italy) and at Heidel- memoir for the Royal Society, to which Punnett was elected berg University (Heidelberg, Germany) and in September in 1912, and followed this with a shorter account for 1899 accepted the post of Demonstrator in the Natural His- GENETICS (Crew 1968). In the opening paragraph of the tory Department of the University of St. Andrews (St. latter he said that Punnett “had the good fortune to be an Andrews, Fife, Scotland). In October 1901 a Fellowship of active participant in the work that confirmed and extended Gonville and Caius College followed, capped by the Univer- sity post of Demonstrator in Morphology, which he held un- Copyright © 2012 by the Genetics Society of America doi: 10.1534/genetics.112.143552 til 1904, when he became Balfour Student in Zoology. This 1Address for correspondence: [email protected] studentship, in memory of Francis Balfour, Professor of Genetics, Vol. 192, 3–13 September 2012 3 Figure 2 Punnett’s square, from the Second Edition of Mendelism (Punnett 1907). carrying out Mendelian experiments at Merton House, Grantchester, [Punnett] wrote to him suggesting that per- haps his nutritional experiments might be so designed that they would yield information concerning the inheritance of coat colour [in the mouse]” (Crew 1967). When Bateson Figure 1 R. C. Punnett. Courtesy of the Master and Fellows of Gonville fi and Caius College, Cambridge. received an offer of nancial support from his friend Mrs. Christiana Herringham in December 1903, he first thought Animal Morphology, Arthur Balfour’s brother, had been held of Leonard Doncaster as an associate, but Doncaster de- by William Bateson from 1897 to 1900. clined (Cock and Forsdyke 2008, p. 217) and so he wrote Then in 1908 Punnett started a rapid rise up the to Punnett (on Christmas Day), inviting him to come “into academic ladder. Still with his Caius Fellowship (which he partnership in my breeding experiments.”“Mr. Punnett was to retain until his death) he became Demonstrator in joined with enthusiasm, and very generously refused Animal Morphology in the Department of Zoology, Superin- the ... salary” (Bateson 1928, p. 87), “... and so a partnership tendent of the Museum of Zoology in 1909, and, when that was to last six years and that was to make notable and Bateson resigned his Professorship of Biology in 1910 to take enduring contributions to genetics came into being. The two up the Directorship of the John Innes Institute, Punnett men were very different temperamentally, Bateson was succeeded to it. In 1912 the Arthur Balfour Professorship of a forceful personality, combative and stern; Punnett was re- Genetics was founded and, following the failure of the tiring, tolerant and friendly; it was a happy and harmonious University to attract Bateson back from his Directorship, partnership” (Crew 1967). Punnett was appointed. We consider the history of the In 1913 Punnett married Eveline Maude Froude, widow Professorship in a later section. of Sidney Nutcombe-Quicke. They lived in Whittingehame At Naples in 1899 Punnett started to study the morphol- Lodge, Storey’s Way, Cambridge, in the house provided for ogy of nemertine (or “Nemertean”) marine worms, and the Arthur Balfour Professor, until Punnett retired in 1940 at these continued to be his main interest at St. Andrews and the age of 65. He and his wife then moved to Bilbrook, near on his return to Cambridge. In 1903 he embarked on a sta- Minehead, Somerset, where he died on January 3, 1967. tistical study “On nutrition and sex-determination in man,” There were no children. using data for London from the 1901 Census, which Crew’s (1967) biographical memoir contains a list of revealed a modest facility in handling numbers (Punnett Punnett’s publications and summaries of his work beyond 1903). The human sex ratio was my own Ph.D. topic and the topics I discuss in detail here. In the summer of 1909 in 1959 I must have heard about his interest and sent him an Punnett had visited Ceylon to study mimetic butterflies, offprint of one of my articles (Edwards 1958) for I still have where he met his Caius colleague R. H. Lock, then Assistant his letter in reply. Director of the Royal Botanical Gardens at Peradeniya Punnett’s association with Bateson started at the begin- (Sri Lanka). The visit led to a handsomely illustrated book- ning of 1904. Some time earlier, “Knowing that Bateson was Mimicry in Butterflies (Punnett 1915). “... it included a 4 A. W. F. Edwards mutationist’s explanation for the evolution of complex mi- metic resemblances between members of unrelated species” (Bennett 1983, p. 8). R. A. Fisher’s view of their evolution was completely different. He set it out in Fisher (1927) and in Chapter VIII, “Mimicry,” of The Genetical Theory of Natural Selection (Fisher 1930a) with special reference to Punnett’s view in the section “The theory of saltations.” Provine (1971, p. 150) gives an account. On evolution Fisher and Punnett were to cross swords again when Punnett reviewed The Genetical Theory, which we refer to below under Popu- lation Genetics. Punnett’s experience with studying Mendelian characters in poultry led him to invent the method of using sex-linked plumage color factors to sex day-old chicks, thus enabling the unwanted majority of cockerels to be disposed of imme- diately. By 1940 he had published, alone or jointly, 11 “Ge- netic studies in poultry,” with another two to come in retirement in 1948 and 1957. Crew (1968) may be referred to for further details, for unlike the biographical memoir for the Royal Society his memoir in GENETICS contains a sub- Figure 3 Punnett’s square, from Report III (Bateson et al. 1906b). stantial extract by Professor F. B. Hutt “whose Genetics of the Fowl is in the direct line of Punnett’s Heredity in Poultry, so the two developments went hand in hand. Here we 1923a” (Crew 1968). give the salient features of Punnett’ssquare,relyingon Punnett (1928) edited Bateson’s scientific articles for the extended account by Edwards (2012), which is fully Cambridge University Press. T. H. Morgan (1929) reviewed illustrated. the two volumes in Nature, regretting the omission of the The first published diagrams appeared in 1906. On Reports to the Evolution Committee (see below), which February 1 Bateson, in an address to the Neurological were represented only by summaries. After Bateson’s death Society (Bateson 1906), displayed the 9:3:3:1 Mendelian in 1926, Punnett (1926) wrote a memoir of him in the ratio among the F2 for two loci when dominance is complete Edinburgh Review, part of which was reprinted in Notes at both.
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