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J. B. S. Haldane

Geneticist and evolutionary biologist (1892–1964)

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JBS Haldane at UCL in the early 1940s
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Through Physiology, Genetics and Abiogenesis

Haldane's article on abiogenesis in 1929 introduced the "primordial soup theory", which became the foundation for the concept of the chemical origin of life. He established human gene maps for haemophilia and colour blindness on the X chromosome, and codified Haldane's rule on sterility in the heterogametic sex of hybrids in species. He correctly proposed that sickle-cell disease confers 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 and trans-acting regulation, coupling reaction, molecular repulsion, the darwin (as a unit of evolution), and organismal cloning.

In 1957, Haldane articulated Haldane's dilemma, a limit on the speed of beneficial evolution, an idea that is still debated today. He is also remembered for his work in human biology, having coined "clone", "cloning", and "ectogenesis". With his sister, Naomi Mitchison, Haldane was the first to demonstrate genetic linkage in mammals. Subsequent works established a unification of Mendelian genetics and Darwinian evolution by natural selection whilst laying the groundwork for modern synthesis, and helped to create population genetics.

Haldane served in World War I, and obtained the rank of captain. He was a professed socialist, Marxist, atheist, and secular humanist whose political dissent led him to leave England in 1956 and live in India, becoming a naturalised Indian citizen in 1961. Arthur C. Clarke credited him as "perhaps the most brilliant science populariser of his generation". Brazilian-British biologist and Nobel laureate Peter Medawar called Haldane "the cleverest man I ever knew". According to Theodosius Dobzhansky, "Haldane was always recognized as a singular case"; Ernst Mayr described him as a "polymath" (as did others); Michael J. D. White described him as "the most erudite biologist of his generation, and perhaps of the century"; James Watson described him as "England's most clever and eccentric biologist", and Sahotra Sarkar described him as "probably the most prescient biologist of this [20th] century". According to a Cambridge student, "he seemed to be the last man who might know all there was to be known". He willed his body for medical studies, as he wanted to remain useful even in death.

Biography

Early life and education

Haldane was born in Oxford in 1892. His father was the Scottish physiologist, scientist, philosopher, and Liberal, John Scott Haldane, who was the grandson of evangelist James Alexander Haldane. His mother Louisa Kathleen Trotter, was a Conservative of Scottish ancestry. His only sibling, Naomi Mitchison, became a prominent Scottish writer. His uncle was Viscount Haldane and his aunt was the author Elizabeth Haldane. Descended from an aristocratic and secular family of the Clan Haldane, he later claimed that his Y chromosome could be traced back to Robert the Bruce.

Haldane grew up at 11 Crick Road, North Oxford. He learnt to read at the age of three, and at four, after injuring his forehead, he asked the physician treating him about the bleeding, "Is this oxyhaemoglobin or carboxyhaemoglobin?" He was raised as an Anglican Christian. 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 with its own private laboratory. At age 8, in 1901, his father brought him to the Oxford University Junior Scientific Club to listen to a lecture on Mendelian genetics, which had been recently rediscovered. Although he found the lecture given by Arthur Dukinfield Darbishire, Demonstrator of Zoology at Balliol College, Oxford, "interesting but difficult", it influenced him permanently such that genetics became the field in which he made his most important scientific contributions.

His formal education began in 1897 at Oxford Preparatory School (now Dragon School), where he gained a First Scholarship in 1904 to Eton College. In 1905 he joined Eton, where he experienced severe abuse from senior students for allegedly being arrogant, but was befriended by Julian Huxley. The indifference of authority left him with a lasting hatred for the English education system. However, the ordeal did not stop him from becoming captain of the school.

He participated for the first time in scientific research as a volunteer subject for his father in 1906. John was the first to study the effects of decompression (relief from high pressure) in humans. He investigated the physiological condition called "bends", such as when goats lift and bend their legs if discomforted, that also may affect deep-sea divers. In July 1906, on board HMS Spanker off the west coast of Scotland, Rothesay, young Haldane jumped into the Atlantic Ocean with the experimental diving suit. The study was published in a 101-paged article in The Journal of Hygiene in 1908; where Haldane was described as "Jack Haldane (age 13)" for whom it "was the first time [he] had ever dived in a diving dress". The research became a foundation for a scientific theory called Haldane's decompression model.

He studied mathematics and classics at New College, Oxford, and obtained first-class honours in mathematical Moderations in 1912. 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 that same year, as a co-author alongside his father. His mathematical treatment of the study was published in December 1913 in the Proceedings of the Physiological Society.

Haldane did not want his education to be confined to a specific subject; he took up Greats (classics) and graduated with first-class honours in 1914. While he had full intention of studying physiology, his plan was, as he described later, referring to World War I, "somewhat overshadowed by other events". His only formal education in biology was an incomplete course in vertebrate anatomy.

Career

To support the war effort, Haldane volunteered to join the British Army, and was commissioned a temporary second lieutenant in the 3rd Battalion of the Black Watch (Royal Highland Regiment) on 15 August 1914. He was assigned as the trench mortar officer, to lead his team for hand-bombing the enemy trenches, the experience of which he described as "enjoyable". In his article in 1932 he described how "he enjoyed the opportunity of killing people and regarded this as a respectable relic of primitive man". He was promoted to temporary lieutenant on 18 February 1915 and to temporary captain on 18 October. While serving in France, he was wounded by artillery fire and sent back to Scotland, where he served as instructor of grenades for the Black Watch recruits. In 1916, he joined the war in Mesopotamia (Iraq), where an enemy bomb severely wounded him. He was relieved from action and sent to India, where he stayed for the rest of the war. He returned to England in 1919 and relinquished his commission on 1 April 1920, retaining his rank of captain. For his ferocity and aggressiveness in battles, his commander described him as the "bravest and dirtiest officer in my Army". Another senior officer of his regiment called him 'mad' and 'cracked'.

Between 1919 and 1922, he served as Fellow of New College, Oxford, where, despite his lack of formal education in the field, he taught and researched in physiology and genetics. During his first year at Oxford, six of his papers dealing with physiology of respiration and genetics were published. He then moved to the University of Cambridge, where he accepted a newly created readership in biochemistry in 1923 and taught until 1932. During his nine years at Cambridge, he worked on enzymes and genetics, particularly the mathematical side of genetics. While working as a visiting professor at the University of California, Berkeley in 1932, he was elected Fellow of the Royal Society.

Haldane worked part-time at the John Innes Horticultural Institution (later named John Innes Centre) at Merton Park in Surrey from 1927 to 1937. When Alfred Daniel Hall became the director in 1926, one of his earliest tasks was to appoint as assistant director "a man of high quality in the study of genetics" who could become his successor. Upon the recommendation of Julian Huxley, the council appointed Haldane in March 1927, with the terms: "Mr. Haldane to visit the Institution fortnightly for a day and a night during the Cambridge terms, to put in two months also at Easter and long vacations in two continuous blocks and to be free in the Christmas vacation." He was officer in charge of Genetical Investigations. He became the Fullerian Professor of Physiology at the Royal Institution from 1930 to 1932 and in 1933 he became Professor of Genetics at University College London, where he spent most of his academic career. As Hall did not retire until 1939, Haldane did not in fact succeed him, but resigned from the John Innes in 1936 to become the first Weldon Professor of biometry at University College London. Haldane was credited with helping the John Innes become "the liveliest place for research in genetics in Britain". He moved his team to the Rothamsted Experimental Station in Hertfordshire from 1941 to 1944, during World War II, to escape bombings. Reginald Punnett, founder of the Journal of Genetics in 1910 with William Bateson, invited him to become editor in 1933, a post he retained until his death.

In India

In 1956, Haldane left University College London, and joined the Indian Statistical Institute (ISI) in Calcutta where he worked in the biometry unit. Haldane gave many reasons for moving to India. 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. In an article "A passage to India" that he wrote in The Rationalists Annual in 1958, he stated: "For one thing I prefer Indian food to American. Perhaps my main reason for going to India is that I consider that the opportunities for scientific research of the kind in which I am interested are better in India than in Britain, and that my teaching will be at least as useful there as here." The university had sacked his wife Helen for being drunk and disorderly and refusing to pay a fine, triggering Haldane's resignation. He decided that he would no longer wear socks, stating that "Sixty years in socks is enough." Afterwards, he always dressed in Indian attire.

Haldane was keenly interested in inexpensive research. Explaining in "A passage to India", he said, "Of course, if my work required electron microscopes, cyclotrons, and the like, I should not get them in India. But the sort of facilities which Darwin and Bateson used for their researches—such as gardens, gardeners, pigeon lofts, and pigeons—are more easily obtained in India than in England." 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. He took an interest in the study of floral symmetry. In January 1961 he befriended Canadian lepidopterist Gary Botting, the 1960 U.S. Science Fair winner in zoology (who had first visited the Haldanes along with Susan Brown, 1960 U.S. National Science Fair winner in botany), inviting him to share the results of his experiments hybridising Antheraea silk moths. He, his wife Helen Spurway, and student Krishna Dronamraju were present at the Oberoi Grand Hotel in Calcutta when Brown reminded the Haldanes that she and Botting had a previously scheduled event that would prevent them from accepting an invitation to a banquet proposed by the Haldanes in their honour and had regretfully declined the honour. After the two students had left the hotel, Haldane went on his much-publicized hunger strike to protest what he regarded as a "U.S. insult". When the director of the ISI, P. C. Mahalanobis, confronted Haldane about both the hunger strike and the unbudgeted banquet, Haldane resigned from his post (in February 1961), and moved to a newly established biometry unit in Bhubaneswar, the capital of Orissa (Odisha).

Haldane took Indian citizenship; he was 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 responded:

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 found science teaching in India wanting. He said "the stupidity of the mynah shows that in birds, as in men, linguistic and practical abilities are not very highly correlated. A student who can repeat a page of a text book may get first class honours, but may be incapable of doing research."

Personal life

Haldane was married twice, first to Charlotte Franken and then to Helen Spurway. In 1924, Haldane met Charlotte Franken, who was a journalist for the Daily Express and married to Jack Burghes. Following the publication of Haldane's Daedalus, or Science and the Future, she interviewed Haldane and they began a relationship. In order to marry Haldane, Franken filed a divorce suit, which resulted in controversy as Haldane was involved as co-respondent in the legal proceeding. Additionally, as Sahotra Sarkar reported: "For her to secure a divorce, Haldane overtly committed adultery with her". Haldane's conduct was described as "gross immorality", for which he was formally dismissed by Cambridge's Sex Viri (a six-member disciplinary committee) from the university in 1925. Cambridge professors, including G. K. Chesterton, Bertrand Russell, and W. L. George, raised their defence for Haldane insisting that the university should not make such judgements, based solely on a professor's private life. The ouster was revoked in 1926. Haldane and Charlotte Franken were married in 1926. Following their separation in 1942, they divorced in 1945. Later that year he married Helen Spurway, his former PhD student. He also had an affair with Angel Records founder Dorle Soria.

Haldane once boasted about himself, saying, "I can read 11 languages and make public speeches in three; but am unmusical. I am a fairly competent public speaker." He had no children, but he and his father were important influences to his sister Naomi's children, of whom Denis Mitchison, Murdoch Mitchison, and Avrion Mitchison became professors of biology at the University of London, Edinburgh University, and University College London, respectively.

Inspired by his father, Haldane often used self-experimentation and would expose himself to danger in order 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 that resulted in his 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".

Haldane made himself unpopular among his colleagues from the start of his academic career. In Cambridge, he annoyed most of the senior faculty due to his uninhibited behaviour, particularly at dinner. His partisan, Edgar Adrian (a 1932 Nobel laureate), had almost convinced Trinity College to offer him an appointment as a Fellow, but that was ruined by an incident when Haldane arrived at the dining table carrying a gallon jar of urine from his laboratory.

Later life and death

In the autumn of 1963, Haldane visited the US for a series of scientific conferences. At the University of Wisconsin, Sewall Wright introduced him before his speech, noting many of Haldane's achievements, after which Haldane modestly remarked that the introduction would have been more accurate if all the references to "Haldane" were replaced with "Wright". In Florida, he met, for the first and only time, the Russian biochemist Alexander Oparin, who had developed an origin of life theory quite independent of his own in the 1920s. It was while there that he started feeling abdominal pains.

Haldane went to London for a diagnosis. He was found to have colorectal cancer, and had a surgery in February 1964. Around that time Philip Dally was making a BBC documentary about eminent living scientists, which included Sewall Wright and the double Nobel laureate Linus Pauling. Dally's team approached Haldane at the hospital for the documentary profile, but instead of a filmed interview, Haldane gave them a self-obituary, the opening lines of which run:

I am going to begin with a boast. I believe that I am one of the [originally as "I am the most"] most influential people living today, although I haven't got a scrap of power. Let me explain. In 1932 I was the first person to estimate the rate of mutation of a human gene.

He also 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 on 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,
Which kills a lot more chaps, in fact,
Than were bumped off when Troy was sacked ...

The poem ends:

...
Thanks to the nurses and Nye Bevan
The NHS is quite like heaven
Provided one confronts the tumour
With a sufficient sense of humour.
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.

He willed that his body be used for medical research and instruction 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.

His surgery in London was declared successful. But the symptoms reappeared after returning to India in June, and in August, the Indian doctors confirmed that his condition was terminal. Writing to John Maynard Smith on 7 September, he said, "I am not appreciably upset by the prospect of dying fairly soon. But I am very angry [at the English doctor who performed the operation]."

He died on 1 December 1964 in Bhubaneswar. On that day the BBC broadcast his self-obituary as "Professor J.B.S. Haldane, obituary." Following his will, his body was moved to Kakinada where Vissa Ramachandra Rao performed post-mortem and preservation of his body parts. His skeleton and organs are on display to the public in the Haldane Museum, located in the pathology department of Rangaraya Medical College.

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