John Burdon Sanderson Haldane

John Burdon Sanderson Haldane was the son of the Oxford physiologist John Scott Haldane (1860-1936), 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 self-confidence that enabled him to tackle the problems of science in the belief that to a Haldane nothing was impossible.

Haldane’s first scientific training was provided by his father, whom he assisted from the age of eight in the latter’s private laboratory and whom he accompanied on work as government investigator of mining accidents and as physiologist for the admiralty. His first formal training was in the Dragon School in North Oxford, before he went to Eton.

In 1901 Haldane’s 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 Arthur Dukinfield Darbishire (1879-1915), 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 went to war in World War I, as an officer in The Black Watch (Royal Highland Regiment) on the Western front and in Mesopotamia, and was wounded in both campaigns. For a short while he worked with his father and Claude Gordon Douglas (1882-1963), 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 afterwards 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 blood-stream 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 at Cambridge under Frederick Gowland Hopkins (1861-1947), the 1929 recipient of the Nobel Prize for Physiology or Medicine, jointly with Christiaan Eijkman (1858-1930) for the discovery of vitamin.

At Cambridge, he concentrated on the study of enzymes. Using some elegant mathematics, he calculated the rates at which enzyme reactions take place (1931); with George Edward Briggs (1883-1985) 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.

Meanwhile Haldane had been continuing his investigations 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 Aylmer Fischer (1890-1962) and Sewall Green Wright (1889-1988) were working along similar lines. They 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 examples to which he applied his theory, the replacement of the light-coloured 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 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 colour 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 (1879-1979, of Martin-Bell syndrome), began an extensive investigation which showed the genetic linkage between haemophilia and colour 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 part-time 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 (1861-1926).

Shortly before the outbreak of World War II, Haldane was retained by the Amalgamated Engineering Union to represent the interest of their members at the public inquiry into the loss of ninety-nine 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, and much other underwater work. For considerable periods Haldane and a band of personally recruited colleagues, including his future second wife, Helen Spurway (1917-1928), risked their lives regularly. Many of their results were described by Haldane and E. M. Case in 1941.

This work, and later statistical investigations for the government, was carried out despite the fact that Haldane had announced himself a Marxist in the 1930s and for several years was editor of the communist Daily Worker (London). 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 government which before the outbreak of World War II demanded better air raid precautions in Britain.

Haldane stressed the social responsibilities of science and contributed more than 300 articles on popular science to the Daily Worker. 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 Thomas Henry Huxley (1825-1895).

He later became disillusioned with the official communist party line and with the rise of the Soviet biologist Trofim Denisovich Lysenko (1898-1976).

A passage to India

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 cross-fertilization 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 of 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.

Haldane was widely decorated and honoured in academic circles. He possessed an unusual combination of analytic powers, literary abilities, a wide range of knowledge, and a force of personality that produced numerous discoveries in several scientific fields and proved stimulating to an entire generation of research workers.

Haldane was married twice: first to Charlotte Franken, a journalist; and then to Helen Spurway, a fellow biologist, who survived him.

«Science is vastly more stimulating to the imagination than are the classics. »
Daedalus

«I am quite sure that our views on evolution would be very different had biologists studied genetics and natural selection before and not after most of them were convinced that evolution had occurred. »

«Illiteracy in England is mainly determined by congenital weak-mindedness, in India by parental poverty. » Possible Worlds. «Eugenics and Social Reform.»

I have no doubt that in reality the future will be vastly more surprising than anything I can imagine. Now, my own suspicion is that the universe is not only queerer than we suppose, but queerer than we can suppose. Possible Worlds, 1927.

«Early diagnosis of disease is the business of the general public even more than of the medical profession. Possible Worlds, «The Time Factor in Medicine»

«I have never yet met a healthy person who worried very much about his health, or a really good person who worried much about his own soul.»
The inequality of man

«There are a few honest antivivisectionists . . . . I have not met any of them, but I am quite prepared to believe that they exist»
Possible Worlds, «Some Enemies of Science»

“All that biology tells us about the nature of God is that he has "An inordinate fondness for beetles".
In George Evelyn Hutchison (1903-1991): Homage to Santa Rosalia or why are there so many kinds of animals?, The American Naturalist, Chicago, 1959, 93:145-159.

• Reduplication in Mice. With A. D. Sprunt and N. M. Haldane.
  Journal of Genetics, 1915, 5: 133-135.
• The Combination of Linkage Values, and the Calculation of Distances Between the Loci of Linked Factors. Journal of Genetics, Cambridge, 1919, 8: 299–309.
• Sex Ratio and Unisexual Sterility in Hybrid Animals.
  Journal of Genetics, Cambridge, 1922, 12: 101-109.
• A Mathematical Theory of Natural and Artificial Selection.
  Part 1 in: Transactions of the Cambridge Philosophical Society, 1924, 23: 19-41.
  Part 2: The influence of partial self-fertilisation, inbreeding, assortative mating and selective fertilisation on the composition of Mendelian populations and on natural selection.
  In: Proceedings of the Cambridge Philosophical Society, 1924, 1: 158-163.
  Part 3: Proceedings of the Cambridge Philosophical Society, 1926, 23.363-372.
  Part 4: Proceedings of the Cambridge Philosophical Society, 1927, 23: 607-615.
  Part 5: Selection and mutation.
  Proceedings of the Cambridge Philosophical Society, 1927, 23: 838-844.
  Part 6: Isolation. Proceedings of the Cambridge Philosophical Society, 1930, 26: 220-230.
  Part 7: Selection intensity as a function of mortality rate.
  Proceedings of the Cambridge Philosophical Society, 1931, 27: 131-136.
  Part 8: Metastable populations.
  Proceedings of the Cambridge Philosophical Society, 1932, 27: 137-142.
  Part 9: Rapid selection.
  Proceedings of the Cambridge Philosophical Society, 1932, 28: 244-248.
  Part 10: Some theorems on artificial selection. Genetics, 1934, 19: 412-429.
• The effect of reaction changes on human inorganic metabolism.
  J. B. S. Haldane, V. B. Wigglesworth and C. E. Woodrow:
  Proceedings of the Royal Society of London. Series B, London, 1924, 96: 1-14.
• The effect of reaction changes human carbohydrate and oxygen metabolism.
  J. B. S. Haldane, V. B. Wigglesworth and C. E. Woodrow:
  Proceedings of the Royal Society of London. Series B, London, 1924, 96: 15-28.
• Callinicus – a Defence of Chemical Warfare. London, 1925.
• Change of Linkage Poultry With Age.
  With F. A. E. Crew. Nature, 1925; 115: 641.
• A Note on the Kinetics of Enzyme Reaction.
  With G. E. Briggs. Biochemical Journal, 1925, 29: 338-339.
• Possible Worlds, and Other Essays. London, 1927.
• Animal Biology. With Julian Henry Huxley. 1927.
• Science and Ethics. 1928.
• The Origin of Life. Rationalist Annual, 1929: 3-10.
• Enzymes. London, 1930.
• Linkage in the Tetraploid Primula sinensis.
  Written with D. De Winton. Journal of Genetics, 1931, 24: 121-144.
• The Molecular Statistics of an Enzyme Action.
  Proceedings of the Royal Society of London, 1931, 108B: 559–567.
• Linkage in the Tetraploid Primula sinensis.
  With D. de Winton. Journal of Genetics, 1931, 24: 121-144.
• The Causes of Evolution. London, 1932.
• The principles of plant breeding, illustrated by Chinese Primrose.
  Proceedings of the Royal Institution of Great Britain, London, 1931, 26: 199-208.
• Prehistory in the light of genetics. London 1931
  Proceedings of the Royal Institution of Great Britain, London, 1931, 26: 355-370.
• A note on inverse probability.
  Proceedings of the Cambridge Philosophical Society, 1932, 28: 55-61.
• The Inequality of Man. 1932.
• On the nonlinear difference equation Dch=kf(ch). Cambridge 1932
  Proceedings of the Cambridge Philosophical Society, 1932, 28: 234-243
• The Causes of Evolution. London, 1933.
• The Genetics of Primula sinensis: Segregation and Inter-action of Factors in the Diploid. Written with D. De Winton. Journal of Genetics, 1933, 27: 1-44.
• A Provisional Map of a Human Chromosome. Nature, London, 1935; 137: 397.
• Some problems of mathematical biology.
  Journal of Mathematics and Physics, Cambridghe, Massachusetts, 1935, 14: 125-136.
• Natural Selection. Nature, London, 1936; 138: 1053.
• My Friend Mr. Leakey. London, 1937.
• The Effect of Variation on Fitness. American Naturalist, 1937; 71: 337-349.
• The Linkage Between the Genes for Colour-Blindness and Haemophilia in Man.
  With Julia Bell. Proceedings of the Royal Society of London. Series B, 1937, 123: 119-150.
• Heredity and Politics. 1938.
• The Marxist Philosophy and the Sciences. 1939.
• New Paths in Genetics. 1941.
• Human Physiology Under High Pressure. 1. Effects of Nitrogen, Carbon Dioxide, and Cold. Written with E. M. Case. Journal of Hygiene, 1941, 41: 225-249.
• A labour-saving method of sampling. Nature, London, 1945, 155: 49-50.
• Science Advances. 1947.
• The formal genetics of man. Croonian Lecture.. London 1947-1948
  Proceedings of the Royal Society of London. Series B, 1947-1948, 135: 147-170.
• In Defense of Genetics. Modern Quarterly, 1949; 4: 194.
• The Biochemistry of Genetics. 1954.
• Une Discussion sur l'origine de la vie. Translated from English.
  J. D. Bernal, J. B. S. Haldane. Paris, 1955. Publications de l'Union rationaliste.
• Genetical effects of radiation from products of nuclear explosions.
  Nature, London, 1955, 176: 115.
• Origin of man. Nature, London, 1955, 176: 169-170.
• Educational problems of the Colonial Territories.
  Nature, London, 1955, 176: 750
• The maximisation of national income. Sankhya, Calcutta 1955-1956, 16: 1-2.
• Biometry. Sankhya, Calcutta 1955-1956, 16: 207-214.
• On the biochemistry of heterosis, and the stabilization of polymorphism.
  Proceedings of the Royal Society of London. Series B, 1955-1956,, 144: 217-220.
• The Biochemistry of Genetics. 2. imprint, London, etc. 1956: 144.
• A Provisional Map of a Human Chromosome. Nature, 1935, 137: 397.
• Natural Selection. Nature, 1936, 138: 1053.
• My Friend Mr. Leakey. London, 1937.
• The Effect of Variation on Fitness. American Naturalist, 1937, 71: 337-349.
• The Linkage Between the Genes for Colour-Blindness and Haemophilia in Man.
  With Julia Bell. Proceedings of the Royal Society, 1937, 123B: 119-150.
• Human Physiology Under High Pressure. 1 . Effects of Nitrogen, Carbon Dioxide, and Cold. With E. M. Case. Journal of Hygiene, 1941, 41: 225-249.
• In Defence of Genetics. Modern Quarterly, n.s. 1i939, 4: 194.
• Imprinting and the evolution of instincts. With Helen Spurway. Nature, London, 1956, 178: 85-86.
• The theory of selection for melanism in Lepidoptera.
  Proceedings of the Royal Society of London. Series B, 1956, 145: 303-306.
• The relation between density regulation and natural selection.
  Proceedings of the Royal Society of London. Series B, 1956, 145: 306-308.
• Almost unbiassed estimates of functions of frequencies.
  Sankhya, Calcutta, 1956-1957, 17: 201-208.
• Graphical methods in enzyme chemistry. Nature, London, 1957, 179: 832.
• The Syadvada system of predication. Sankhya, Calcutta, 1957, 18: 195-200,
• The quantitative study of animal behaviour in an approximately steady state.
  With Helen Spurway. V pamet na Metodij Popov: Sbornik of naucni trudove po biologija i medicina, Sofija, 1958: 89-93.
• The Scope of biological statistics. Sankhya, Calcutta, 1958, 20: 195-206.
• The theory of natural selection to-day. Nature, London, 1959, 183: 710-713.
  
  
• Parthogenese. Naturwissenschaftliche Rundschau, Stuttgart, 1959, 12: 453-458.
  
  
• The analysis of heterogenity. I. Sankhya, Calcutta, 1959, 21: 209-216.
• Pasteur and cosmic asymmetry. Nature, London, 1960, 185: 87.
• The scientific work of J. S. Haldane (1860-1936).
  Nature, London, 1960, 187: 102-105.
• 'Dex' or 'Order of magnitude'? Nature, London, 1960, 187: 879.
• The addition of random vectors. Sankhya, Calcutta, 1960, 22: 213-220.
• Simple approximations to the probability integral and P ( c2,1) when both are small.
  Sankhya, Calcutta, 1961, 23: 9-10.
• The proper social application of the knowledge of human genetics.
  In: The Science of science. Edited by Maurice Goldsmith and Alan Mackay, London 1964.
  
  
• The genetics of a Common Indian digital abnormality.
  With Ajit Kisor Ray. Proceedings of the National Academy of Sciences of the United States of America, Washington, 1965, 53: 1050-1053.
• Science and human life.
  Freeport, New York, 1971. 287 pages. (Essay Index Reprint Series)

• K. R. Dronamraju, editor:
  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, 1966, 12: 219–249; and Science Reporter (Delhi), 2 1965, a special Haldane number containing articles on his life and work.
• Ronald William Clark, 1916-1987:
  J. B. S.: The life and work of J. B. S. Haldane.
  London : Hodder & Stoughton Ltd, 1968. 286 pages.
• Ronald William Clark:
  Haldane, John Burdon Sanderson. In: Charles Coulston Gillispie, editor in chief: Dictionary of Scientific Biography. Charles Scribner’s Sons. New York, 1970.
• Encyclopædica Britannica. 1986.
• Maurice B. Strauss, editor:
  Familiar Medical Quotations.
  Little, Brown and Company (Inc). Boston, 1968.
• Wikipedia.