BATESON, WILLIAM

(b. Whitby, England, 8 August 1861; d. Merton, London, 8 February 1926)

embryology, genetics. For the original article on Bateson see DSB, vol. 1.

Since the mid-1970s the increasing attention given to the history of genetics has brought fresh studies on Bateson. A growing interest in the history of scientific institutions and the funding of research has also led to further studies of Bateson’s unusual career. A detailed account is now available of his efforts to gain financial and institutional support in Cambridge and to harness the willing hands of the young women of Newnham College, and of his role as the first director of the John Innes Horticultural Institute at Merton. The discovery in 1983 of homeobox genes that control body plans, and the abrupt alterations to those plans caused by mutations among these genes, is a reminder of the examples of just such mutants described by Bateson in 1894. The manner in which genetics and embryology come together in the homeobox helps us to appreciate the undercurrent of embryological thinking in Bateson’s conception of variability and genetics.

Variations. The first phase of Bateson’s research—using embryology to reveal phylogeny—formed a part of the research program pursued in Cambridge University’s zoology department. His decision to terminate such work can be viewed within the broad context of what Peter Bowler has called the period of “The Eclipse of Darwinism.” Skepticism concerning the role of natural selection had been opening the way to suggestions of the internal source of adaptive variation, a scenario in which natural selection seemed unlikely to have played a creative role. Bateson next turned to variation, calling it “the essential phenomenon of evolution” (Materials, 1894, p. 6). To this new agenda he brought what embryology had taught him: the importance of symmetry in the organization of living things, and the repetition of parts, or “meristic” variation, as he called it, and the substitution of parts, or “homeotic” variation. Then there were variations in “the substance of the parts themselves”; he called these “substantive” variations. The first class of variations invited analysis in terms of patterns of cell division leading to radial and bilateral symmetries. The second suggested the appeal to chemistry—ferments producing pigments that color eyes, flowers, and skin.

Bateson’s enthusiastic study of these variations led to the 600-page Materials for the Study of Variation (1894). The subtitle Treated with Especial Regard to Discontinuity in the Origin of Species announced the message of the book. It was diametrically opposed to the continuity of variation assumed by the Darwinians. But his travels to Russia and Egypt in search of discontinuities in the environment that match discontinuities between species ;proved fruitless. Henceforth he decided that the distinctiveness of species—their discontinuity—was not to be attributed to natural selection having eliminated intermediate forms from a continuously varying population, but to the discontinuity of the variations themselves. Therefore he set out to establish the extent of discontinuity in variation, compiling a long catalog of examples that fill the twenty-one chapters of Materials, from extra antennae, legs, and palpi in insects to supernumerary digits, hands, feet, teeth, and teats in humans. He even reported one case of an insect with an extra antenna issuing from one of its eyes (Materials, fig. 19, p. 151). The discontinuity of such variations, he reasoned, should be manifest in their non-blending character when crossbred. Accordingly, he concluded the book with an appeal for “the organization of systematic experiments in breeding, a class of research that, he judged, calls for more patience and more resources than any other form of biological inquiry” (p. 574).

In 1895 he began such a program, and was soon able to draw on assistance from the women of Newnham College. Here Marsha Richmond’s account of their assistance to Bateson has filled a gap in accounts of Bateson’s informal network of Cambridge researchers. Bateson suggested or assigned breeding experiments: Veronica to Dora Pertz; Mendel’s peas to Hilda Blanch Kilby; Lathyrus to Mary Hart-Davis; Antirrhinum to Muriel Wheldale; mice to his sister-in-law, Florence Durham; and canaries to her and Dorothea Charlotte Edith Marryat. Two events that encouraged this support were the failure of the campaign for degrees for women in 1897 and the ensuing exclusion of women from the Morphological Laboratory and from beginning courses in biology (Richmond, 2001, p. 64). Richmond has also described the important role played by Edith Rebecca Saunders as Bateson’s first and longest-serving collaborator from 1895 until 1910. Only when Bateson began to deputize for Professor Alfred Newton in 1899 did he gain followers among the male students, starting with Reginald Crundell Punnett in 1904, followed by Leonard Doncaster, Robert Lock, and Reginald Philip Gregory.

Promotion of the Mendelian Program. Bateson’s experience in crossbreeding prepared him to take the lead in 1900 in promoting in the English-speaking world the work of Gregor Mendel and his rediscoverers. Bateson first read about what became known as Mendelian segregation in the offspring of hybrids (F₂) when Hugo de Vries sent him his first paper on the subject, published in the Comptes rendus de l’Académie des Science. Here de Vries made no mention of Mendel. Evidence has been presented to claim that this was the paper that Bateson read while traveling to London to give his lecture “Problems of Heredity.” The report of this lecture written by the

respected Dr. Maxwell Masters and published in the Gardeners’ Chronicle four days later also makes no mention of Mendel. Indeed, it appears that Bateson presented Hugo de Vries’s results as modifications of Galton’s law because he was incorporating De Vries’s results into the Galtonian framework of the talk he had prepared (Olby, 1987). This is hardly surprising, for as early as 1897 Galton had suggested how his ancestral law could be applied to non-blending characters, using data from the pedigree of basset hounds. Instead of considering ancestral contributions to offspring to be blended in each individual, he pictured them as being distributed in pure form to members of the offspring, in proportions conforming to the ancestral law, not unlike the effects of Mendelian segregation (Galton, 1897; see also Pearson, 1900). Only on his return to Cambridge did Bateson see De Vries’s second, more detailed paper in which Mendel’s work is acknowledged. Only then did he possess the information leading him to the 1865 volume of the Verhandlungen der Naturforschenden Vereines in Brünn in the Cambridge University Library.

Bateson had turned to crossbreeding experiments in order to establish that new variations, providing they were of a discontinuous character, would not be swamped by crossing with the normal form. Mendel’s paper then provided him with an explanation of the underlying stochastic process at the level of the germ cells. For Bateson, Mendel’s essential discovery was “the purity of the gametes.” Bateson’s insistence on the significance for evolution of discontinuous variations thus became attached to the Mendelian theory, a situation that guaranteed opposition to Mendelism from Darwinians. The infamous controversy between Mendelians and Darwinian biometricians that followed has been the subject of much interest, especially because Donald Mackenzie and S. Barry Barnes attempted a sociological analysis of the controversy. Bateson’s success in taking control of the Royal Society Evolution Committee in 1897—and hence gaining access to Royal Society grants for his research— undoubtedly embittered the biometricians, but it gave Bateson the resources to continue and expand his breeding studies.

Meanwhile, during the fourteen years in which Bateson carried out breeding experiments in Cambridge, apart from the award of the Balfour studentship to his collaborator, Reginald Crundall Punnett, he received no research funds from the university or from any of its associated funding bodies. In 1903, however, Bateson’s friend, Christina J. Herringham, the well-known copyist of Italian masters and wife of the Wimpole Street consultant Sir Wilmot Parker Herringham, offered to furnish the funds needed so that Bateson would be able to pay a salary for an assistant. Two years later Francis Darwin and his brothers supplied funds, enabling Bateson to have a greenhouse

constructed for the hybridization of tender plants. Any other financial support came from outside Cambridge.

In 1907 Bateson spent three months in the United States and gave the Silliman Lectures at Yale University that October. The warm reception that greeted him there raised his self-esteem, but he was not prepared when, on his return to Cambridge, the University offered him not a chair but a readership, and in zoology, not genetics. Although he was persuaded to accept, shortly thereafter a member of the University offered the funds to establish and support for five years a chair of biology that was clearly intended for Bateson. In June 1908 he was elected. The duty of the professor would be to “teach and make researches in that branch of biology now entitled Genetics (Heredity and Variation)” (B. Bateson, p. 112). Bateson now had the status and salary worthy of his achievements and international reputation, but the chair carried no funds for research, and its future after the five years was not assured. A year later, when Bateson learned of the plans for a Horticultural Institution funded by the late John Innes and of the appeal for applications for the post of director, he did not apply. On the twelve-member advisory Council for the Trust, however, were five of Bateson’s friends. Passing over two of Bateson’s excellent former coworkers—Gregory and Lock—they opened negotiations with him. Responding positively, he substantially modified the Council’s plan for the Institute, and on these terms he accepted the position. In 1910, after forty-nine years in Cambridge, he left to begin a new life.

Move to Merton. During the early years at Merton, Bateson was occupied with establishing his staff and instituting his research program. Across the ocean, Thomas Hunt Morgan, Arthur H. Sturtevant, Calvin B. Bridges, and Herman J. Muller were breeding fruit flies (Drosophila) and exploring the relation between heredity and the chromosomes. Their book, The Mechanism of Mendelian Heredity(1915), established the chromosomes as the bearers of the Mendelian factors or genes. As William Coleman has already explained, Bateson vigorously opposed it. For him the fundamental process underlying heredity and development was cell division, not chromosomes. The association of particular Mendelian factors he attributed to differential rates of “reduplication” of the gametes carrying those factors, rather than to the position of the factors close together on the same chromosome. Mechanistic theory he did support, but not the conjectured structural complexities of cell organelles, which at the time were only clearly distinguished with the magic of staining techniques. Nor could he ignore the problems posed by the theory. How can the chromosomes control differentiation if each cell has the same complement? Should not the chromosomes of cells in different tissues differ? And why did the number of chromosomes in different species not vary in accordance with the degree of complexity of the species?

Seven years after the publication of The Mechanism of Mendelian Heredity, Bateson visited Morgan’s laboratory at Columbia University and withdrew his opposition to their theory. Addressing the American Association for the Advancement of Science in Toronto, Bateson declared: “I come at this Christmas season to lay my respectful homage before the stars that have arisen in the west … The arguments of Morgan and his colleagues, and especially the demonstrations of Bridges, must allay all skepticism as to the direct association of particular chromosomes with particular features of the zygote” (B. Bateson, p. 392).

When Bateson retired from the John Innes Horticultural Institute (JIHI) in 1926 he could take special credit for the new knowledge gained concerning compatibility and incompatibility between varieties of apples, pears, plums, and cherries; his work, however, could easily give the impression of a piecemeal collection of studies lacking a clear focus. Actually, Bateson’s several projects—on double flowers, rogue peas, and plant chimeras—were aimed at demonstrating segregation without meiosis. He was following his own advice to “treasure your exceptions.” Such a strategy during his early years of working on Mendel’s laws had revealed one exception after another. Pursuing them had led to his discovery of gene interaction: epistasis and hypostasis, “coupling” (i.e., linkage), differing expressions of dominance, as in sex determination, and much else. Many of these research topics were natural choices for a horticultural institute and arose out of Bateson’s close connections with the Royal Horticultural Society and with prominent members of the trade.

Bateson had succeeded in transferring his researches from Cambridge to Merton, but unlike Morgan, he had neither built up a team of researchers at Merton, nor had he attracted colleagues of the caliber of Bridges and Muller. In altering the Council’s plan, he had abolished the four well-paid positions recommended by Council, choosing instead to offer minor grants to students, some at £50 per year, intended chiefly for “women students who wish to devote themselves to research in various fields but can scarcely afford to earn nothing at all” (Olby, 1989,

p. 506). Cyril Dean Darlington, a future director of the JIHI, began working there without pay, as did the mycologist Dorothy Cayley. On his retirement, Bateson expressed his regret that he had not achieved more at the JIHI. “When the centre of chief interest in genetics shifted away from work of my type to that of the American group,” he wrote, “I was already too old and too much fixed in my ideas to become master of so very new and intricate a development” (p. 507).

Shortly after he became established at the JIHI, Bateson was surprised to be invited back to Cambridge. This invitation resulted from the concern of former prime minister Arthur Balfour, who wished to see genetics properly established at Cambridge and thus took the lead in seeking funds for a chair of genetics there. Aided by Lord Esher and encouraged by Prime Minister Herbert Henry Asquith, a benefactor was found: William George Watson, of the Maypole Dairy company (Opitz, 2005). In 1912, at Balfour’s instigation, the chair was offered to Bateson; by this time, however, he felt well settled at Merton and declined. The call from Cambridge, supported by two prime ministers, speaks to the reputation Bateson had acquired since the early years when the Cambridge Darwinians viewed him as a dissident member of their community who sniped at their work, often in a scornful tone.

Honors. Bateson’s work was recognized by the Royal Society with the award of the Darwin Medal in 1904, the Royal Medal in 1923, and he was the Society’s Croonian lecturer in 1920. The Royal Horticultural Society awarded him the Victoria Medal in 1911 and the Royal Institution elected him Fullerian Professor (1912-1915). At a national level he served in 1912 as chairman of Lloyd George’s important Development Commission and ten years later he was offered a Knighthood which he declined. In 1910 he received an honorary degree from Sheffield University and became an Honorary Fellow of his Cambridge College, St. John's.

SUPPLEMENTARY BIBLIOGRAPHY

Originals and copies of Bateson’s correspondence and papers (1883-1929) are preserved and cataloged at the John Innes Archive, The John Innes Center, Norwich, United Kingdom (Originally the John Innes Horticultural Institution). The originals of part of this large collection are held in the archives of the American Philosophical Society, Philadelphia, and at Cambridge University Library. A small collection of Bateson Family Papers 1829–1921 is held at the American Philosophical Society.

WORKS BY BATESON

Materials for the Study of Variation, Treated with Especial Regard for the Study of Variation. London and New York: Macmillan, 1894. Cited as Materials.

Problems of Genetics. New Haven, CT: Yale University Press, 1913. Reprinted in the Silliman Milestones in Science series, New Haven, CT: Yale University Press, 1979.

OTHER SOURCES

Bateson, Beatrice. William Bateson, F.R.S.: His Essays and Addresses together with a Short Account of His Life. Cambridge: Cambridge University Press, 1928. Cited as “B. Bateson.”

Bowler, Peter. The Eclipse of Darwinism: AntiDarwinian Evolution Theories in the Decades around 1900. Baltimore, MD: Johns Hopkins University Press, 1983.

Cock, Alan. “William Bateson’s Rejection and Eventual Acceptance of Chromosome Theory.” Annals of Science40 (1983): 19–59.

Galton, Francis. “The Average Contribution of Each Several Ancestor to the Total Heritage of the Offspring.” Proceedings of the Royal Society 61 (1897): 403.

Henig, Robin M. The Monk in the Garden: The Lost and Found Genius of Gregor Mendel, the Father of Genetics. Boston and New York: Houghton Mifflin, 2000.

Lipset, David. Gregory Bateson: The Legacy of a Scientist. Englewood Cliffs, NJ: Prentice Hall, 1980.

Mackenzie, Donald, and S. Barry Barnes. “Biometriker versus Mendelianer. Eine Kontroverse und ihre Erklarung.” Kölner Zeitschrift für Soziologie und Sozialpsychologie(1975)

Sonderheft 13, section iv, pp. 165–196, and an abbreviated account: “Scientific Judgement: The Biometry-Mendelism Controversy,” in Natural Order: Historical Studies of Scientific Culture, edited by S. B. Barnes and Steven Shapin. Beverly Hills and London: Sage Publications, 1979.

Morgan, Thomas Hunt; A. H. Sturtevant; H. J. Muller; and C. B. Bridges. The Mechanism of Mendelian Heredity. New York: Henry Holt, 1915.

Olby, Robert. “William Bateson’s Introduction of Mendelism to England: A Reassessment.” British Journal for the History of Science 20 (1987): 399–420.

;——. “Scientists and Bureaucrats in the Establishment of the John Innes Horticultural Institution under William Bateson.” Annals of Science 46 (1989): 497–510.

——. “The Dimensions of Scientific Controversy: The Biometric-Mendelian Debate.” British Journal for the History of Science22 (1988): 299–320. Critiques Mackenzie and Barnes.

——. “Horticulture: The Font for the Baptism of Genetics.” Nature Reviews Genetics1 (2000): 65–70.

——. “The Monk in the Garden” by Robert Henig. Perspectives in Biology and Medicine45 (2002): 142–145. Book review.

Opitz, Donald L. “‘No Doubtful or Uncertain Enterprise’ Balfour, Bateson, and Britain’s First Chair of Genetics at Cambridge, 1894–1914.” History of Science Society Meeting, Minneapolis, MN, 3–6 November 2005.

Pearson, Karl. “Mathematical Contributions to the Theory of Evolution—on the Law of Reversion.” Proceedings of the Royal Society66 (1900): 141.

Richmond, Marsha L. “Women in the Early History of Genetics: William Bateson and the Newnham College Mendelians, 1900–1910.” Isis92 (2001): 55–90.

;——. “The ‘Domestication’ of Heredity: The Familial Organization of Geneticists at Cambridge, 1895–1910.” Journal of the History of Biology 37 (2004): 1–41.

Robert C. Olby