PUNNETT, REGINALD CRUNDALL

(b. Ton-bridge, England, 20 June 1875; d. Bilbrook, Somerset, England, 3 January 1967)

morphology, genetics.

Punnett was the elder son of Georage Punnett, the head of a Tonbridge building firm, and Emily Crundal. The famlily was comfortably bourgeois, conservative, and churchgoing. In 1913 he married Evline Maude Froude, widow of Sidney Nut6combe-Quicke and daughter of John Froude, Bellew; they had no children.

Having graduated from a preparatory school in Tonbridge, Punnett, in 1889, went to Clifton, where his interest was the classics. He then obtained a scholarship at Gonville and Caius College, Cambridge, and registered as a medical student. For the first part of the natural science tripos he studied zoology, human anatomy, and physiology, but found the first of these so attractive that he decided to become a zoologist. For the second part of the tripos he offered zoology, with human physiology as his subsidiary subject, and gained a first in the examination and the Walsingham Medal in 1898. With the aid of the Shuttleworth studentship at Caius, he spent six months at the Zoological Station in Neples and a short time at Gegenbaur’s laboratory in Heidelberg. In 1899 he went to St. Andrews University as demonstrator and part-time lecturer in the natural history department. In 1901 he was elected a fellow of Caius and in the following year returned to Cambridge ad demonstrated in morphology in the department of zoology. He was Balfour student from 1904 to 1908, was awrded the Thurston Medal, and in 1909 was appointed super intendent of the Museum of Zoology. In 1910 he succeeded Bateson as professor of bilogy at the University of Cambridge. When this impermanent chair became adequately endowed and was tranformed into the Arthur Balfour chair of genetics (the first such chair in the United Kingdom), Punnett became its first occupant. He held this post until his retirement in 1940. Punnett was elected a fellow of the Royal Society in 1912 and was awarded its Darwin Medal in 1922. He was a founding mamber of the (British) Genetical Society and was one of its secreteries from 1919 to 1930, when he became president. He was also an honoury member of the Genetical Society of Japan and of the Poultry Science Association of America.

While still an undergraduate, Punnett became keenly interested in nemertines; and he pursued this interest at the Nepals Zoological Station, at the Marine Biological Station in Plymouth, at the Marine Labortary at St. Andrwes University, and at Nordgaard’s laboratory in Bergen. Punnett wroter about a dozen papers on the morphology of these worms and his name was given to Cerebratulus punnetti and Punnettia splendida Stasny-Wijinhoff (KEferstein).

In 1901, while convalescing from an appendectomy, Punnett bacame interested in a report that sex ratios can be influenced by diet; and he decided to test this hypothesis experimently by using mice. On his return to Cambridge in 1902, he wrote to Bateson, who was conducting Mendelian experimentation in Granschester, and asked if his proposed nutritional expriment might be tested to yield information concerning the inheritance of coat color. Althoughj he learned that such investigations were already well advanced, this contact with Bateson become the turning in Punnett’s career. In 1903 an anomymous friend offered Bateson £150 a year, for two years, to continue his work. He invited Punnett to join him. Their collaboration was to last six years and produced many notable and enduring contributions to the nascent science of genetics. During the period 1904-1910 they confirmed several basic discoveries of classical Mendelian genetics, including the Mendelian explanation of sex determination, sex linkage, complementary factors and factor interaction, and the first example of autosomal linkage. For these experiments Bateson and Punnett also used the rabbit, and in addition to his hybridization work he was greatly interested in mimicry in the butterfly. In 1911 Bateson and Punnett launched the Journal of Genetics, which they edited jointly until Bateson’s death in 1923. Thereafter Punnett became sole editor until 1946, when he was succeeded by J. B. S. Haldane.

During World War I, Punnett served in the Food Production Department of the Board of Agriculture(he was an authority on the scientific aspects of poultry breeding), and it was here that he conceived using sex-linked plumage-color factors in the fowl to reduce wastage. Feed stuffs were in short supply and the vast majority of male chicks were unwanted. If the sexes could be distinguished among day-old chicks the unwanted males could be destroyed and considerable economy achieved. He suggested various crosses involving such characters as silver and gold, barred and nonbarred; and his method was soon practiced on a huge scale, involving the production of millions of sex-linked chicks every year.

After the war the National Poultry Institute was founded and the Genetical Institute at Cambridge was selected as the center for research in poultry breeding. Michael Pease jointed Punnett as research assistant; and the most dramatic production of this partnership was the Cambar, the first autosexing breed of poultry. This breed was synthesized by transferring the X-borne gene for barring(B)from the Barred Rock to the Golden Campine. Ten years later he produced the Legbar, which resulted from the introduction of the B gene into the genotype of the Brown Leghorn. The production of these autosexing breeds was a fine piece of biological engineering. But in 1930, when the authorities decided that this work should be greatly expanded, Punnett with drew; and a separate center for poultry research was established with Pease as its director. Thereafter Punnett worked alone. He subsequently left Cambridge and went to the village of Bilbrook, where he continued his experimental work with fowl until 1955, when his incubator house was destroyed by fire. Thus, more than half a century of quiet, methodical, and highly intelligent Mendelian experimentation ended.

During the period 1899-1904 Punnett worked alone on various problems. But he was principally concerned with meristic variation in elasmobranchs; the morphology and systematics of nemertines and an expert morphologist. Then, encouraged by Bateson, he abruptly shifted his interest to experimental biology and applied his previous experience to hybridization experiments, which confirmed and extended the findings of Mendel. Punnett defended Mendelism against the criticisms of various biometricians and zoologists, and he was instrumental in introducing genetics to the general public, and especially to fanciers and commercial breeders of livestock. He also made significant contributions to knowledge of the genetics of the fowl, the duck, the rabbit, the sweet pea, and of man; and his first book,Mendelism(1905), did much to make the science known. His Heredity in Poultry(1923)remained the standard work on the subject for nearly thirty years, and our present knowledge of poultry genetics has evolved largely from Punnett’s early investigations. In 1940 he produced a map of the X chromosomes of the fowl showing the tentative arrangement of seven genes. His last paper, which was concerned with recessive, black plumage color, appeared in 1957. Data from nearly thirty years of experimentation with the sweet pea were consolidated in his paper on linkage; and in it he showed that eighteen known recessive mutations fell into seven groups, seven being the haploid number of chromosomes in the sweet pea.

Punnett was a gentle, quiet, cultured man who never strove for prominence or priority. The slow rate of reproduction of the fowl and of the sweet pea suited his temperament. He was a Mendelian throughout his career and remained largely unaffected by the development of the theory of the gene, of cytogenetics, and to biometrical genetics. He belonged to the first group of scientist who carried forward the revolution in biological thought that was inspired by the rediscovery of Mendel’s work.

BIBLIOGRAPHY

A complete bibliography of Punnett’s writings is given in Biographical Memoirs of Fellows of the Royal Society,13 (1967), 323-326. His scientific interests can be clearly discerned in the long series of papers on poultry genetics that appeared from 1911 to 1967 in the Journal of Genetics.

F. A. E. Crew