J. B. S. Haldane

5 Nov 1892
1 Dec 1964
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J. B. S. Haldane, born John Burdon Sanderson Haldane, known as Jack, FRS (/ˈhɔːldeɪn/; 5 November 1892 – 1 December 1964) (but who used ‘J. B. S.’ in his printed works), was a British-Indian scientist, well known for his works in physiology, genetics and evolutionary biology, mathematics where he made innovative contributions to statistics and biostatistics. He was son of the equally famous John Scott Haldane. He was a socialist, Marxist, atheist, and humanist whose political dissent led him to leave England in 1956 and live in India, and became a naturalised Indian citizen in 1961.

His first paper in 1915 demonstrated genetic linkage in mammals while subsequent works helped to create population genetics, establishing a unification of Mendelian genetics and Darwinian evolution by natural selection and laying the groundwork for modern evolutionary synthesis. His article on Abiogenesis in 1929 introduced the “Primordial Soup Theory”, and is the foundation of modern understanding of the chemical origin of life. Haldane established human gene maps for haemophilia and colour blindness on the X chromosome, and formulated Haldane’s rule on sterility in the heterogametic sex of hybrids in species. He correctly proposed that sickle-cell disease gives some immunity to malaria. He was the first to suggest the central idea of in vitro fertilisation, as well as concepts such as hydrogen economy, cis trans, coupling, repulsion, darwin (as a unit of evolution) and cloning. In 1957 he articulated Haldane’s dilemma, a limit on the speed of beneficial evolution which subsequently proved incorrect. He willed his body for medical studies, as he wanted to remain useful even in death.

Arthur C. Clarke credited him as “perhaps the most brilliant science populariser of his generation”. Nobel laureate Peter Medawar called Haldane “the cleverest man I ever knew”

With his sister Naomi Mitchison he started investigating Mendelian genetics in 1908, used guinea pigs and mice, publishing Reduplication in mice in 1915, demonstrating genetic linkage in mammals,as well as in chickens.

Following his father’s footsteps, Haldane’s first publication was on the mechanism of gaseous exchange by haemoglobin. and subsequently worked on the properties of blood.He investigated several aspects of kidney functions and mechanism of excretion.

In 1925, with G. E. Briggs, Haldane derived a new interpretation of the enzyme kinetics law described by Victor Henri in 1903, different from the 1913 Michaelis–Menten equation. Leonor Michaelis and Maud Menten assumed that enzyme (catalyst) and substrate (reactant) are in fast equilibrium with their complex, which then dissociates to yield product and free enzyme. The Briggs–Haldane equation was of the same algebraic form, but their derivation is based on the quasi steady state approximation, that is the concentration of intermediate complex (or complexes) does not change. As a result, the microscopic meaning of the “Michaelis Constant” (Km) is different. Although commonly referring it as Michaelis–Menten kinetics, most of the current models actually use the Briggs–Haldane derivation

He was one of the three major figures to develop the mathematical theory of population genetics along with Ronald Fisher and Sewall Wright, and thus playing an important role in the modern evolutionary synthesis, the concept more popularly known as “neo-Darwinism” (popularised by Richard Dawkins’ 1976 work titled The Selfish Gene), which re-established natural selection as the premier mechanism of evolution by explaining it in terms of the mathematical consequences of Mendelian inheritance.He wrote a series of ten papers called A Mathematical Theory of Natural and Artificial Selection, on the mathematical theory behind natural selection. It showed the direction and rates of changes of gene frequencies and pioneered the interaction of natural selection with mutation and animal migration. Haldane’s book, The Causes of Evolution (1932), summarised these results, especially in its extensive appendix.

His contributions to statistical human genetics included the first methods using maximum likelihood for estimation of human linkage maps, and pioneering methods for estimating human mutation rates. He was the first to estimate rate of mutation in humans, at 2 × 10−5 mutations per gene per generation for the X-linked haemophilia gene, and to introduce the idea of a “cost of natural selection”.At the John Innes Horticultural Institution he developed the complicated linkage theory for polyploids,and extended the idea of gene/enzyme relationships with the biochemical and genetic study of plant pigments. In his essay On Being the Right Size he outlines Haldane’s principle, which states that the size very often defines what bodily equipment an animal must have: “Insects, being so small, do not have oxygen-carrying bloodstreams. What little oxygen their cells require can be absorbed by simple diffusion of air through their bodies. But being larger means an animal must have complicated oxygen pumping and distributing systems to reach all the cells.” The conceptual metaphor to animal body complexity has been of use in energy economics and secession ideas.

Haldane introduced the modern concept of abiogenesis in an eight-page article titled The origin of life (1929), describing the primitive ocean as a “vast chemical laboratory” containing a mixture of inorganic compounds – like a “hot dilute soup” in which organic compounds could have formed. Under the solar energy the oxygenless atmosphere containing carbon dioxide, ammonia and water vapour gave rise to a variety of organic compounds, “living or half-living things”. The first molecules reacted with one another to produce more complex compounds, and ultimately the cellular components. At some point a kind of “oily film” was produced that enclosed self-reproducing molecules, thereby becoming the first cell. J.D. Bernal named the hypothesis biopoiesis or biopoesis, the process of living matter evolving from self-replicating but nonliving molecules. The idea was generally dismissed as “wild speculation”. In 1924 Alexander Oparin had suggested similar idea in Russian, and in 1936 he introduced it to the English-speaking people. The experimental basis of the theory began in 1953 with the classic Miller–Urey experiment. Since then the primordial soup theory became a dominant theory in the chemistry of life, and often is attributed as Oparin-Haldane hypothesis.

He proposed in 1949 that genetic disorders in humans living in malaria-endemic regions provide beneficial effects in making people immune to malaria itself, noting that red blood cell mutations such as sickle-cell anemia and various thalassemias were prevalent only in tropical regions where malaria was endemic and were favourable traits for natural selection that prevented individuals from receiving malarial infection. This concept of resistance to malaria was eventually confirmed by A.C. Allison in 1954

Haldane was born in Oxford to John Scott Haldane, a physiologist, scientist, a philosopher and a Liberal, and Louisa Kathleen Haldane (née Trotter), a Conservative. His younger sister, Naomi Mitchison, became a writer while his uncle was Richard Haldane, 1st Viscount Haldane and his aunt the author Elizabeth Haldane. Descended from an aristocratic and secular Haldane family of the Clan Haldane, he would later claim that his Y chromosome could be traced back to Robert the Bruce.

He grew up at 11 Crick Road, North Oxford. He learnt to read age three and age four, after he injured himself on the forehead he asked the doctor, “Is this oxyhemoglobin or carboxyhemoglobin?”. From age eight he worked with his father in their home laboratory where he experienced his first self-experimentation, the method he would later be famous for. He and his father became their own “human guinea pigs”, such as in their investigation on the effects of poison gases. In 1899 his family moved to “Cherwell”, a late Victorian house at the outskirts of Oxford having its private laboratory.

His formal education began in 1897 at Oxford Preparatory School (now Dragon School), where he gained a First Scholarship in 1904 to Eton. In 1905 he joined Eton, where he experienced severe abuse from senior students on his part as being arrogant. The indifference of the authority left him with a lasting hatred for English education system. However, the ordeal did not stop him from becoming Captain of the school. He studied mathematics and classics at New College at the University of Oxford and obtained first-class honours in mathematical moderations in 1912 and a first-class honours in 1914. With his father he published his first scientific paper, age 20. He became engrossed in genetics and presented a paper on gene linkage in vertebrates in the summer of 1912. His first technical paper, a 30-page long article on haemoglobin function, was published, as a co-author alongside his father.

His education was interrupted by the First World War during which he fought in the British Army, being commissioned a temporary second lieutenant in the 3rd Battalion of the Black Watch (Royal Highland Regiment) on 15 August 1914.He was promoted to temporary lieutenant on 18 February 1915 and to temporary captain on 18 October.He served in France and Iraq, where he was wounded. He relinquished his commission on 1 April 1920, retaining his rank of captain. Between 1919 and 1922 he was a Fellow of New College, Oxford, where he researched physiology and genetics. He then moved to the Cambridge University, where he accepted a readership in Biochemistry at Trinity College in the University of Cambridge and taught until 1932.From 1927 until 1937 he was also Head of Genetical Research at the John Innes Horticultural Institution. During his nine years at Cambridge, Haldane worked on enzymes and genetics, particularly the mathematical side of genetics.He was the Fullerian Professor of Physiology at the Royal Institution from 1930 to 1932 and in 1933 he became full Professor of Genetics at University College London, where he spent most of his academic career. Four years later he became the first Weldon Professor of Biometry at University College London.

In 1924, Haldane met Charlotte Haldane. So that they could marry, Charlotte divorced her husband, Jack Burghes, causing some controversy. Haldane was almost dismissed from Cambridge for the way he handled his meeting with her, which led to the divorce. They married in 1926. Following their separation in 1942, the Haldanes divorced in 1945. He later married Helen Spurway.

Haldane, inspired by his father, would expose himself to danger to obtain data. To test the effects of acidification of the blood he drank dilute hydrochloric acid, enclosed himself in an airtight room containing 7% carbon dioxide, and found that it ‘gives one a rather violent headache’. One experiment to study elevated levels of oxygen saturation triggered a fit which resulted in him suffering crushed vertebrae. In his decompression chamber experiments, he and his volunteers suffered perforated eardrums, but, as Haldane stated in What is Life, “the drum generally heals up; and if a hole remains in it, although one is somewhat deaf, one can blow tobacco smoke out of the ear in question, which is a social accomplishment.

In 1956, Haldane left University College London, and joined the Indian Statistical Institute (ISI) in Kolkata, India,where he headed the biometry unit. Officially he stated that he left the UK because of the Suez Crisis, writing: “Finally, I am going to India because I consider that recent acts of the British Government have been violations of international law.” He believed that the warm climate would do him good, and that India shared his socialist dreams.The university had sacked his wife Helen for excessive drinking and refusing to pay a fine, triggering Haldane’s resignation. He declared he would no longer wear socks, “Sixty years in socks is enough.” and always dressed in Indian attire.

He was keenly interested in inexpensive research. He wrote to Julian Huxley about his observations on Vanellus malabaricus, the Yellow-wattled Lapwing. He advocated the use of Vigna sinensis (cowpea) as a model for studying plant genetics. He took an interest in the pollination of Lantana camara. He lamented that Indian universities forced those who took up biology to drop mathematics.Haldane took an interest in the study of floral symmetry. In January 1961 he befriended Gary Botting, visiting ISI to share his experiments hybridising Antheraea silk moths. The United States cultural attache prevented Botting’s attendance at a banquet to which the Haldanes had invited him to meet biologists from all over India and Haldane protested by going on a hunger strike.When the director of the ISI, P. C. Mahalanobis, confronted Haldane about both the hunger strike and the unbudgeted banquet, Haldane resigned his post, in February 1961, and moved to a newly established biometry unit in Odisha.

Haldane became an Indian citizen.He was also interested in Hinduism and became a vegetarian. In 1961, Haldane described India as “the closest approximation to the Free World.” Jerzy Neyman objected that “India has its fair share of scoundrels and a tremendous amount of poor unthinking and disgustingly subservient individuals who are not attractive.”Haldane retorted:

“ Perhaps one is freer to be a scoundrel in India than elsewhere. So one was in the U.S.A in the days of people like Jay Gould, when (in my opinion) there was more internal freedom in the U.S.A than there is today. The “disgusting subservience” of the others has its limits. The people of Calcutta riot, upset trams, and refuse to obey police regulations, in a manner which would have delighted Jefferson. I don’t think their activities are very efficient, but that is not the question at issue. ”
When on 25 June 1962 he was described in print as a “Citizen of the World” by Groff Conklin, Haldane’s response was as follows:

“ No doubt I am in some sense a citizen of the world. But I believe with Thomas Jefferson that one of the chief duties of a citizen is to be a nuisance to the government of his state. As there is no world state, I cannot do this. On the other hand, I can be, and am, a nuisance to the government of India, which has the merit of permitting a good deal of criticism, though it reacts to it rather slowly. I also happen to be proud of being a citizen of India, which is a lot more diverse than Europe, let alone the U.S.A, the U.S.S.R or China, and thus a better model for a possible world organisation. It may of course break up, but it is a wonderful experiment. So, I want to be labeled as a citizen of India. ”

Haldane became a socialist during the First World War, supported the Second Spanish Republic during the Spanish Civil War, and then became an open supporter of the Communist Party in 1937, being a pragmatic, dialectical materialist Marxist, and writing many articles for the Daily Worker. In On Being the Right Size, Haldane wrote: “while nationalization of certain industries is an obvious possibility in the largest of states, I find it no easier to picture a completely socialized British Empire or United States than an elephant turning somersaults or a hippopotamus jumping a hedge.”

In 1938, he proclaimed enthusiastically that “I think that Marxism is true.” He joined the Communist Party of Great Britain in 1942. The first edition of his children’s book My Friend Mr. Leakey contained an avowal of his party membership which was removed from later editions. Pressed to speak out about the rise of Lysenkoism and the persecution of geneticists in the Soviet Union as anti-Darwinist and the denouncement of genetics as incompatible with dialectical materialism, Haldane shifted the focus to the United Kingdom and a criticism of the dependence of scientific research on financial patronage. In 1941 Haldane wrote about the Soviet trial of his friend and fellow geneticist Nikolai Vavilov:

“ The controversy among Soviet geneticists has been largely one between the academic scientist, represented by Vavilov and interested primarily in the collection of facts, and the man who wants results, represented by Lysenko. It has been conducted not with venom, but in a friendly spirit. Lysenko said (in the October discussions of 1939): ‘The important thing is not to dispute; let us work in a friendly manner on a plan elaborated scientifically. Let us take up definite problems, receive assignments from the People’s Commissariat of Agriculture of the USSR and fulfil them scientifically. Soviet genetics, as a whole, is a successful attempt at synthesis of these two contrasted points of view.’ ”
By the end of the Second World War Haldane had become an explicit critic of the regime. He left the party in 1950, shortly after considering standing for Parliament as a Communist Party candidate. He continued to admire Joseph Stalin, describing him in 1962 as “a very great man who did a very good job”.

Haldane was parodied as “the biologist too absorbed in his experiments to notice his friends bedding his wife” by his friend Aldous Huxley in the novel Antic Hay (1923). His essay Daedalus; or, Science and the Future (1924), about ectogenesis and in vitro fertilisation was an influence on Huxley’s Brave New World (1932). Haldane’s work was also admired by Gerald Heard.Various essays on science were collected and published in a volume titled Possible Worlds in 1927. His book, A.R.P. (Air Raid Precautions) (1938) combined his physiological research into the effects of stress upon the human body with his experience of air raids during the Spanish Civil War to provide a scientific explanation of the air raids that Britain was to endure during the Second World War.

Along with Olaf Stapledon, Charles Kay Ogden, I. A. Richards, and H. G. Wells, Haldane was accused by C. S. Lewis of scientism. Haldane criticised the Ransom Trilogy and Lewis for the “complete mischaracterisation of science, and his disparagement of the human race”. He wrote a book for children titled My Friend Mr Leakey (1937), containing the stories “A Meal With a Magician”, “A Day in the Life of a Magician”, “Mr Leakey’s Party”, “Rats”, “The Snake with the Golden Teeth”, and “My Magic Collar Stud” with illustrations by Quentin Blake. Haldane edited Gary Botting’s manuscript on the genetics of giant silk moths with margin notes.

In 1923, in a talk given in Cambridge titled “Science and the Future”, Haldane, foreseeing the exhaustion of coal for power generation in Britain, proposed a network of hydrogen-generating windmills. This is the first proposal of the hydrogen-based renewable energy economy.

In his An Autobiography in Brief, written shortly before his death in India, Haldane named four close associates as those showing promise to become illustrious scientists: T. A. Davis, K. R. Dronamraju, S. D. Jayakar and S. K. Roy.

Haldane was the first to have thought the genetic basis of creating cloning of humans, and eventually super-talented individuals. For this he coined the term “clone”,though the synonym “clon” (from the Greek word klon meaning “twig”) was used in horticulture since the 19th century. He introduced the term in his speech on “Biological Possibilities for the Human Species of the Next Ten Thousand Years” at the Ciba Foundation Symposium on Man and his Future in 1963.

Shortly before his death from cancer, Haldane wrote a comic poem while in the hospital, mocking his own incurable disease; it was read by his friends, who appreciated the consistent irreverence with which Haldane had lived his life. The poem first appeared in print in the 21 February 1964 issue of the New Statesman, and runs:

“Cancer’s a Funny Thing:
I wish I had the voice of Homer
To sing of rectal carcinoma,
This kills a lot more chaps, in fact,
Than were bumped off when Troy was sacked…”

The poem ends:

“… I know that cancer often kills,
But so do cars and sleeping pills;
And it can hurt one till one sweats,
So can bad teeth and unpaid debts.
A spot of laughter, I am sure,
Often accelerates one’s cure;
So let us patients do our bit
To help the surgeons make us fit.”

Haldane died on 1 December 1964. He willed that his body be used for study at the Rangaraya Medical College, Kakinada.

“My body has been used for both purposes during my lifetime and after my death, whether I continue to exist or not, I shall have no further use for it, and desire that it shall be used by others. Its refrigeration, if this is possible, should be a first charge on my estate.

Haldane was elected a Fellow of the Royal Society in 1932.The French Government conferred him its National Order of the Legion of Honour in 1937. In 1952, he received the Darwin Medal from the Royal Society. In 1956, he was awarded the Huxley Memorial Medal of the Anthropological Institute of Great Britain. He received the Feltrinelli Prize from Accademia Nazionale dei Lincei in 1961. He also received an Honorary Doctorate of Science, an Honorary Fellowship at New College, and the Kimber Award of the US National Academy of Sciences. He was awarded the Linnean Society of London’s prestigious Darwin–Wallace Medal in 1958.

He is famous for the (possibly apocryphal) response that he gave when some theologians asked him what could be inferred about the mind of the Creator from the works of His Creation: “An inordinate fondness for beetles.” This is in reference to there being over 400,000 known species of beetles in the world, and that this represents 40% of all known insect species (at the time of the statement, it was over half of all known insect species).
Often quoted for saying, “My own suspicion is that the universe is not only queerer than we suppose, but queerer than we can suppose.”
“It seems to me immensely unlikely that mind is a mere by-product of matter. For if my mental processes are determined wholly by the motions of atoms in my brain I have no reason to suppose that my beliefs are true. They may be sound chemically, but that does not make them sound logically. And hence I have no reason for supposing my brain to be composed of atoms.”
“Teleology is like a mistress to a biologist: he cannot live without her but he’s unwilling to be seen with her in public.”
“I had gastritis for about fifteen years until I read Lenin and other writers, who showed me what was wrong with our society and how to cure it. Since then I have needed no magnesia.”
“I suppose the process of acceptance will pass through the usual four stages: i) This is worthless nonsense, ii) This is an interesting, but perverse, point of view, iii) This is true, but quite unimportant, iv) I always said so.”
“Three hundred and ten species in all of India, representing two hundred and thirty-eight genera, sixty-two families, nineteen different orders. All of them on the Ark. And this is only India, and only the birds.”
When asked whether he would lay down his life for his brother, Haldane, preempting Hamilton’s rule, supposedly replied “two brothers or eight cousins”.

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