Haldane, John Burdon Sanderson

(b. Oxford, England, 5 November 1892; d. Bhubaneswar, Orissa, India, 1 December 1964)

physiology, biochemistry, genetics.

Haldane was the son of the Oxford physiologist John Scott Haldane, member of a Scottish family that traces its ancestry to the mid-thirteenth century. His mother was Louisa Kathleen Trotter, also a Scottish patrician. From both parents he inherited a selfconfidence that enabled him to tackle the problems of science in the belief that to a Haldane nothing was impossible. An Eton education set him against established authority, and service in World War I confirmed an early tendency toward atheism. At Oxford he turned from mathematics and biology to “greats” and thus left the university without scientific qualification. Haldane was married twice: first to a journalist, Charlotte Franken; and then to Helen Spurway, a fellow biologist, who survived him.

A physiologist in the immediate postwar years, Haldane switched first to biochemistry under Frederick Hopkins at Cambridge and then to genetics at University College, London, where, in the precomputer age, his mathematical talénts were fully employed.

Throughout a varied working life he stressed the social responsibilities of science. He belonged naturally to the radical left and was for some years a member of the Communist party, to whose Daily Worker he contributed more than 300 articles on popular science. Despite his varied work in many fields Haldane is likely to be remembered mainly as a geneticist and as a popular expositor of the unity of science. His extensive writings, which continued until the year of his death, ranged from ten famous papers in which he made mathematical contributions to the theory of natural selection to many volumes of essays explaining science to the layman, an art of which he was one of the greatest practitioners since T. H. Huxley.

Haldane’s first scientific training was provided by his father, whom he assisted from childhood in the latter’s private laboratory and whom he accompanied on work as government investigator of mining accidents and as physiologist for the Admiralty. In 1901 his interest in genetics was aroused by a lecture on the recently rediscovered work of Gregor Mendel; it was increased in 1910, when he began to study the laws of inheritance as revealed by his sister’s 300 guinea pigs. Reading an early paper by A. D. Darbishire, Haldane noted what appeared to be the first example of gene linkage in vertebrates; he later read an undergraduate paper on the subject but delayed publication until he had obtained his own data (1915).

Haldane saw service in World War I on the Western Front and in Mesopotamia, and wounded in both campaigns. For a short while he worked with his father and C. G. Douglas, both hurriedly brought to France from England, on the improvisation of gas masks following the first German gas attacks. This gave him an interest in the physiological problems of respiration, an interest he retained for the rest of his life, and the material for a controversial book on gas warfare (1925).

On demobilization in 1919 Haldane took up a fellowship at New College, Oxford, and shortly afterward began teaching physiology. Respiration was the only part of the subject in which he was well versed, but a crash course provided by his father gave him, as he later wrote, “about six weeks’ start on my future pupils.” With Peter Davies, a young worker in the Oxford physiological laboratory, Haldane began to investigate how carbon dioxide in the human bloodstream enables the muscles to regulate breathing under different conditions. During the experiments both men consumed quantities of bicarbonate of soda and “smuggled” hydrochloric acid into their blood by drinking solutions of ammonium chloride. In further experiments, popularly described in “On Being One’s Own Rabbit” (1927), he measured the changes in the sugar and phosphate content of his blood and urine which could be induced by various means.

In 1921 Haldane accepted a readership in biochemistry under Hopkins at Cambridge, where he concentrated on the study of enzymes. Using some elegant mathematics, he calculated the rates at which enzyme reactions take place (1931); with G. E. Briggs he produced the first proof that enzyme reactions obey the laws of thermodynamics (1925); and in Enzymes (1930), produced largely from his Cambridge lectures, he provided an overall picture of how enzymes work. In this, as in much other comparable work, his knowledge of physiology plus his mathematical expertise enabled him to bring a feeling of practical reality to what had previously been largely biochemical assumptions.

Meanwhile Haldane had been continuing his investigation of linkage, and as early as 1919 he had given a formula relating the extent of linkage to the interval on the chromosome. He investigated the variation of linkage with age (1925) and formulated Haldane’s law (1922), covering the crossing of animal species to produce an offspring of which one sex is absent or sterile. Meanwhile Haldane was producing the first of his ten major papers on the mathematics of natural selection, later reprinted as an appendix to his classic The Causes of Evolution (1932). Both Ronald Fisher and Sewall Wright were working along similar lines; but both introduced novel ideas into their papers while Haldane tended, instead, to reinforce the conservative Darwinian theory that natural selection, rather than mutation, was the driving force behind evolution. The most famous example to which he applied his theory, the replacement of the lightcolored moth Biston betularia by a dark mutant form (1924), was strikingly verified by field studies thirty years later.

In 1933 Haldane left Cambridge for University College, London, where he occupied first the chair of genetics and then that of biometry. Here he gave increasing time to human genetics, preparing in 1935 a provisional map of the X chromosome which showed the positions on it of the genes causing color blindness, severe light sensitivity of the skin, night blindness, a particular skin disease, and two varieties of eye peculiarity. In 1936 he and a colleague, Julia Bell, began an extensive investigation which showed the genetic linkage between hemophilia and color blindness. The same year he gave the first estimate of the mutation rate in man, and in 1937 he described the effect on a population of recurrent harmful mutations.

From 1927 until 1936 Haldane also held a parttime appointment at the John Innes Horticultural Institution, then at Merton, outside London, where he carried on the genetic research of the former director, William Bateson. Here he began the joint work by Hopkins’ Cambridge laboratory and the Institute on variation in flower color; and with D. de Winton he contributed to linkage theory by developing the theory for polyploids to tetraploid Primula sinensis (1931, 1933). But his lack of botanical experience and of experimental dexterity combined with his personal aggressiveness to bring the appointment to an end.

Shortly before the outbreak of World War II, Haldane was retained by the Amalgamated Engineering Union to represent the interests of their members at the public inquiry into the loss of ninetynine lives when the submarine Thetis sank while on trials. The physiological work, during which Haldane and four members of the International Brigade were sealed into a chamber in which conditions in the stricken submarine were simulated, led directly to Haldane’s doing much wartime work for the Admiralty. This involved investigation of the physiological problems concerned in escape from submarines, the operations of midget submarines, and much other underwater work. For considerable periods Haldane and a band of personally recruited colleagues, including his future second wife, Helen Spurway, risked their lives regularly. Many of their results were described by Haldane and E. M. Case (1941).

This work, and later statistical investigations for the government, was carried on despite Haldane’s chairmanship of the editorial board of the Daily Worker. He had joined the Communist party soon after the outbreak of the Spanish Civil War, during which he advised the Republican government on gas precautions; and his scientific experience was conscripted by the predominantly left-wing movement which before the outbreak of World War II demanded better air raid precautions in Britain. He was an early supporter of Trofim Lysenko, fighting a rearguard action in his defense—often, it is clear, against his better scientific judgement—until 1949, when his article “In Defence of Genetics” revealed far less than the unquestioning support that Communist orthodoxy demanded.

In 1957 Haldane emigrated to India, ostensibly in protest against the Anglo-French invasion of Suez but largely, in fact, because he was attracted by the country’s facilities for research in genetics and biometry. He worked for the Indian Statistical Office in Calcutta under Prasanta Mahanalobis and, after a short and unsatisfactory spell with the Council for Scientific and Industrial Research, moved to Orissa, setting up a genetics and biometry laboratory in the state capital of Bhubaneswar, where he died of cancer in December 1964. It was typical that while apparently recovering from a cancer operation in London, Haldane should write for the New Statesman a short poem which he hoped would encourage people not to take the disease too seriously. Entitled “Cancer’s a Funny Thing,” and starting. “I wish I had the voice of Homer/To sing of rectal carcinoma,” it was in many ways the apotheosis of Haldane. It brought him a large postbag of letters which, in almost equal numbers, complained of his lack of feeling and praised him for his courage.

The uniqueness of Haldane’s contribution to science was that for much of his life he was able to bring to fresh fields the equipment and concepts he had acquired in other disciplines; for him “the crossfertilisation of ideas” really worked. This was also true of his papers and books, which range from the highly technical to the popular and include one classic book on science for children (1937). It was typical that he should describe in “The Origin of Life” (1929) a mechanism for the synthesis of organic matter which Darwin had merely assumed, a speculation on the origin of life very comparable with Alexander Oparin’s in Russia. It is no matter for surprise that his bibliography should occupy a dozen closely printed pages in Biographical Memoirs of Fellows of the Royal Society.

BIBLIOGRAPHY

I. Original Works. A scientific bibliography as well as a list of Haldane’s books of essays and a selection of his more important popular articles is in Pirie’s biographical memoir (see below) and is reprinted in Clark’s biography (see below).

Among his writings are “Reduplication in Mice,” in Journal of Genetics,5 (1915), 133–135, written with A. D. Sprunt and N. M. Haldane: “The Combination of Linkage Values, and the Calculation of Distances Between the Loci of Linked Factors,” ibid.,8 (1919), 299–309; “Sex Ratio and Unisexual Sterility in Hybrid Animals,”ibid.,12 (1922), 101–109; the 10-part “A Mathematical Theory of Natural and Artificial Selection": pt. 1. in Transactions of the Cambridge Philosophical Society, 23 (1924), 19–41; pts.2–9 in Proceedings of the Cambridge Philosophical Society, 1 , et seq. (1924–1932); pt. in Genetics 19 (1934), 412–429; Callinicus—A Defence of Chemical Warfare (London, 1925): “Change of Linkage in Poultry With Age,” in Nature, 115 (1925), 641, written with F. A. E. Crew; “A Note on the Kinetics of Enzyme Reaction,” in Biochemical Journal,29 (1925), 338–339, written with G. E. Briggs; Possible Worlds, and Other Essays (London, 1927); “The Origin of Life,” in Rationalist Annual (1929), pp. 3–10; Enzymes (London, 1930); “The Molecular Statistics of an Enzyme Action,” in Proceedings of the Royal Society,108B (1931), 559–567; “Linkage in the Tetraploid Primula sinensis:,” in Journal of Genetics, 24 (1931), 121–144, written with D. de Winton; The Causes of Evolution (London, 1932); “The Genetics of Primula sinensis: Segregation and Inter-action of Factors in the Diploid,” in Journal of Genetics,27 (1933), 1–44, written with D. de Winton; “A Provisional Map of a Human Chromosome,” in Nature, 137 (1935), 397; “Natural Selection,” ibid.,138 (1936), 1053; My Friend Mr. Leakey (London, 1937); “The Effect of Variation on Fitness,” in American Naturalist,71 (1937), 337–349; “The Linkage Between the Genes for Colour-Blindness and Haemophilia in Man,”, in. Proceedings of the Royal Society,123B (1937), 119–150, written with J. Bell; “Human Physiology Under High Pressure. 1 . Effects of Nitrogen, Carbon Dioxide, and Cold,” in Journal of Hygiene,41 (1941), 225–249, written with E. M. Case; and “In Defence of Genetics,” in Modern Quarterly, n.s. 4 (1949), 194.

Haldane’s papers were with him in Calcutta and Bhubaneswar and, following his death, are believed to have been taken to Hyderabad by his widow.

II. Secondary Literature. See Ronald W. Clark, J. B. S.: The Life and Work of J. B. S. Haldane (New York, 1969); K. R. Dronamraju, ed., Haldane and Modern Biology (Baltimore, 1968), edited by one of Haldane’s former students; N. W. Pirie, in Biographical Memoirs of Fellows of the Royal Society, 12 (1966), 219–249; and Science Reporter (Delhi), 2 (1965), a special Haldane number containing articles on his life and work. The reminiscences of Haldane’s mother are in L. K. Haldane, Friends and Kindred (London, 1961).

Ronald W. Clark