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Haldane, JBS

John Burdon Sanderson Haldane FRS (/ˈhɔːldeɪn/; 5 November 1892 – 1 December 1964[1][2]), nicknamed "Jack" or "JBS",[3] was a British-Indian scientist who worked in physiology, genetics, evolutionary biology, and mathematics. With innovative use of statistics in biology, he was one of the founders of neo-Darwinism. He served in the Great War, and obtained the rank of captain.[4] Despite his lack of an academic degree in the field,[1] he taught biology at the University of Cambridge, the Royal Institution, and University College London.[5] Renouncing his British citizenship, he became an Indian citizen in 1961 and worked at the Indian Statistical Institute for the rest of his life.

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.[6][7] 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.[8] He willed his body for medical studies, as he wanted to remain useful even in death. 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 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".[9][10] Brazilian-British biologist and Nobel laureate Peter Medawar called Haldane "the cleverest man I ever knew".[11] According to Theodosius Dobzhansky, "Haldane was always recognized as a singular case"; Ernst Mayr described him as a "polymath" (as did others);[12]

Michael J. D. White described him as "the most erudite biologist of his generation, and perhaps of the century";[13] James Watson described him as "England's most clever and eccentric biologist",[14] and Sahotra Sarkar described him as "probably the most prescient biologist of this [20th] century".[15] According to a Cambridge student, "he seemed to be the last man who might know all there was to be known".[12]

Biography

Early life and education

Haldane was born in Oxford in 1892. His father was John Scott Haldane, a physiologist, scientist, a philosopher, and a Liberal who was the grandson of evangelist James Alexander Haldane.[16] His mother Louisa Kathleen Trotter, was a Conservative, and descended from Scottish ancestry. His only sibling, Naomi, became a writer and married Dick Mitchison, Baron Mitchison (thereby becoming Naomi Mary Margaret Mitchison, Baroness Mitchison), who was his best friend at Eton College.[17] His uncle was Viscount Haldane and his aunt was the author Elizabeth Haldane. Descended from an aristocratic and secular family of the Clan Haldane,[18] he would later claim that his Y chromosome could be traced back to Robert the Bruce.[19]

Haldane grew up at 11 Crick Road, North Oxford.[20] 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?" As a youth he was raised as an Anglican.[21] 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.[22]

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.[23] Although he found the lecture given by Arthur Dukinfield Darbishire, Demonstrator of Zoology at Balliol College, Oxford, "interesting but difficult",[10] it influenced him permanently such that genetics became the field in which he made his most important scientific contributions.[13]

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 for allegedly being arrogant. 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.[24]

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.[25] He investigated the physiological condition called "bends", such as when goats lift and bend their legs if discomforted, that is also experienced by deep-sea divers.[26] 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".[26]: 436  The research became a foundation for a scientific theory called Haldane's decompression model.[27]

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.[28] He presented the mathematical treatment of the study on 19 October in the Proceedings of the Physiological Society and was published in December 1913.[29]

Haldane did not want his education to be confined to a specific subject. He took up Greats and graduated with first-class honours in 1914. While he had full intention of studying physiology, his plan was, as he described later, "somewhat overshadowed by other events" (referring to World War I).[24] His only formal education in biology was an incomplete course in vertebrate anatomy.[1]

Career

To support the war effort, Haldane volunteered for and joined 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.[30] He was assigned as the trench mortar officer, to lead his team for hand-bombing the enemy trenches, the experience of which he remarked "enjoyable".[24] 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".[1] He was promoted to temporary lieutenant on 18 February 1915 and to temporary captain on 18 October.[31][32] While serving in France, he was wounded by an artillery fire for which he was sent back to Scotland.

There 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 war fronts and was sent to India and stayed there for the rest of the war.[24] He returned to England in 1919 and relinquished his commission on 1 April 1920, retaining his rank of captain.[4] For his ferocity and aggressiveness in battles, his commander described him as the "bravest and dirtiest officer in my Army".[33]

Between 1919 and 1922, he served as Fellow of New College, Oxford,[34] where, despite his lack of formal education in the field, he taught and researched in physiology and genetics. During his first year at Oxford, he published six papers dealing with physiology of respiration and genetics.[1] He then moved to the University of Cambridge, where he accepted a newly created readership in Biochemistry in 1923 and taught until 1932.[18] During his nine years at Cambridge, he worked on enzymes and genetics, particularly the mathematical side of genetics.[18] While working as a visiting professor at the University of California in 1932, he was elected Fellow of the Royal Society.[35]

Haldane worked part-time at the John Innes Horticultural Institution (later named John Innes Centre) at Merton Park in Surrey from 1927 to 1937.[36] When Alfred Daniel Hall became the director in 1926,[37] 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. Recommended by 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."[38] He was officer in charge of Genetical Investigations.[1]

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.[39] As Hall did not retire so early as expected – retiring in 1939,[37] Haldane had to resign from the John Innes in 1936 to become the first Weldon Professor of Biometry at University College London.[18] Haldane's service was recorded to have helped the John Innes as "the liveliest place for research in genetics in Britain".[38] At the height of World War II, he moved his team to the Rothamsted Experimental Station in Hertfordshire during 1941 to 1944 to escape bombings.[1] 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.[2]

In India

In 1956, Haldane left University College London, and joined the Indian Statistical Institute (ISI) in Calcutta (later renamed Kolkata), India, where he worked in the biometry unit.[1] 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.[40] 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."[41] The university had sacked his wife Helen for being drunk and disorderly and refusing to pay a fine, triggering Haldane's resignation. He declared he would no longer wear socks, "Sixty years in socks is enough."[42] and he always dressed in Indian attire.[10]

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."[41] 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.

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Personal life

Haldane was married twice, first to Charlotte Franken and then to Helen Spurway.[49] 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.[24] 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.[1] Additionally, as Sahotra Sarkar reported: "For her to secure a divorce, Haldane overtly committed adultery with her".[15]

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.[35] 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.[50]

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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.[51] In his decompression chamber experiments, he and his volunteers suffered perforated eardrums. But, as Haldane stated in What is Life,[52] "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".[53]

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Malaria and sickle-cell anemia

Haldane was the first to realise the evolutionary link between genetic disorder and infection in humans. While estimating the rates of human mutation in different situations and diseases, he noted that mutations expressed in red blood cells, such as thalassemias, were prevalent only in tropical regions where deadly infection such as malaria has been endemic. He further observed that these were favourable traits (heterozygous inheritance of sickle cell trait) for natural selection that protected individuals from receiving malarial infection.[87] He introduced his hypothesis at the Eighth International Congress of Genetics held in 1948 at Stockholm on a topic "The Rate of Mutation of Human Genes".[88] He proposed that genetic disorders in humans living in malaria-endemic regions provided a condition (phenotype) that makes them relatively immune to malarial infections. He formalised the concept in a technical paper published in 1949 in which he made a prophetic statement: "The corpuscles of the anaemic heterozygotes are smaller than normal, and more resistant to hypotonic solutions. It is at least conceivable that they are also more resistant to attacks by the sporozoa which cause malaria."[89] This became known as "Haldane's malaria hypothesis", or concisely, the "malaria hypothesis".[90] This hypothesis was eventually confirmed by Anthony C. Allison in 1954 in the case of sickle-cell anemia.[91][92]

Population genetics

Further information: Modern synthesis (20th century)

Haldane was one of the three major figures to develop the mathematical theory of population genetics, along with Ronald Fisher and Sewall Wright. He thus played an important role in the modern evolutionary synthesis of the early 20th century. He re-established natural selection as the central mechanism of evolution by explaining it as a mathematical consequence of Mendelian inheritance.[93][94] He wrote a series of ten papers, A Mathematical Theory of Natural and Artificial Selection, deriving expressions for the direction and rate of change of gene frequencies, and also analyzing the interaction of natural selection with mutation and migration. The series consists of ten papers published between 1924 and 1934 in journals such as Biological Reviews (part II), Mathematical Proceedings of the Cambridge Philosophical Society (parts I and from III to IX), and Genetics (part X).[95][96][97] He gave a set of lectures based on this series at the University of Wales in 1931, and they were summarised in his book, The Causes of Evolution in 1932.[23]

His first paper on the series in 1924 specifically deals with the rate of natural selection in peppered moth evolution. He predicted that environmental conditions can favour the increase or decline of either the dominant (in this case the black or melanic forms) or the recessive (the grey or wild type) moths. For a sooty environment such as Manchester, where the phenomenon was discovered in 1848, he predicted that the "fertility of the dominants must be 50% greater than that of the recessives".[46] According to his estimate, assuming 1% dominant form in 1848 and about 99% in 1898, "48 generations are needed for the change [for the dominant to appear]... After only 13 generations the dominants would be in a majority."[95] Such mathematical prediction was considered improbable for natural selection in nature,[15] but it was subsequently proven by an elaborate experiment (named Kettlewell's experiment) that was performed by an Oxford zoologist Bernard Kettlewell between 1953 and 1958.[98][99][100] Haldane's prediction was proven further by a Cambridge geneticist Michael Majerus in his experiments conducted between 2001 and 2007.[101]

His contributions to statistical human genetics included: the first methods using maximum likelihood for the estimation of human linkage maps; pioneering methods for estimating human mutation rates; the first estimates of mutation rate in humans (2 × 10−5 mutations per gene per generation for the X-linked haemophilia gene); and the first notion that there is a "cost of natural selection".[102] He was the first to estimate the rate of human mutation in his 1932 book entitled The Causes of Evolution.[103] At the John Innes Horticultural Institution, he developed the complicated linkage theory for polyploids;[36][104] and extended the idea of gene-enzyme relationships with the biochemical and genetic study of plant pigments.[105][13]

Political views

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 of Great Britain in 1937. A pragmatic dialectical-materialist Marxist, he wrote many articles for the Daily Worker. In On Being the Right Size, he wrote that "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".

)A David Low cartoon featuring Haldane – "Prophesies for 1949"

In 1938, Haldane proclaimed enthusiastically: "I think that Marxism is true." He joined the Communist Party in 1942. He was pressed to speak out about the rise of Lysenkoism and the persecution of geneticists in the Soviet Union as anti-Darwinist and the political suppression of genetics as incompatible with dialectical materialism. He shifted his polemic focus to the United Kingdom, criticizing the dependence of scientific research on financial patronage. In 1941, he 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 Soviet 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".[33] Haldane has been accused by authors including Peter Wright and Chapman Pincher of having been a Soviet GRU spy codenamed Intelligentsia.[106][107]

Social and scientific views

Human cloning

Haldane was the first to have thought of the genetic basis for human cloning, and the eventual artificial breeding of superior individuals. For this he introduced the terms "clone" and "cloning",[108] modifying the earlier "clon" that had been used in agriculture since the early 20th century (from Greek klōn, twig). 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. He said:[109]

It is extremely hopeful that some human cell lines can be grown on a medium of precisely known chemical composition. Perhaps the first step will be the production of a clone from a single fertilized egg, as in Brave New World...

On the general principle that men will make all possible mistakes before choosing the right path, we shall no doubt clone the wrong people [such as Hitler]...

Assuming that cloning is possible, I expect that most clones would be made from people aged at least fifty, except for athletes and dancers, who would be cloned younger. They would be made from people who were held to have excelled in a socially acceptable accomplishment.

Ectogenesis and in vitro fertilisation

His essay Daedalus; or, Science and the Future (1924) posited the concept of in vitro fertilisation, which he called ectogenesis. He envisioned ectogenesis as a tool for creating better individuals (eugenics).[110] Haldane's work was an influence on Huxley's Brave New World (1932) and was also admired by Gerald Heard.[111] Various essays on science were collected and published in a volume entitled 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 account of the likely effects of the air raids that Britain was to endure during the Second World War.

Criticism of C. S. Lewis

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 Lewis and his Ransom Trilogy for the "complete mischaracterisation of science, and his disparagement of the human race".[112] Haldane wrote a book for children entitled 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". Later editions featured illustrations by Quentin Blake. Haldane also wrote an essay criticising Lewis's arguments for the existence of God, entitled "More Anti-Lewisite", a reference to the poison gas and its antidote.[113]

Hydrogen-generating windmills

In 1923, in a talk given in Cambridge entitled "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.[114][115][116]

Scientists

In his An Autobiography in Brief, published shortly before his death in India, Haldane named four close associates as showing promise to become illustrious scientists: T. A. Davis, Dronamraju Krishna Rao, Suresh Jayakar, and S. K. Roy.[117]

Awards and honours

Haldane was elected a Fellow of the Royal Society in 1932.[39] 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, Oxford, and the Kimber Award of the National Academy of Sciences in the United States. He was awarded the Linnean Society of London's prestigious Darwin–Wallace Medal in 1958.[58]

Legacy

The Haldane Lecture at the John Innes Centre,[118] where Haldane worked from 1927 to 1937 is named in his honour.[38] The JBS Haldane Lecture[119] of The Genetics Society is named in his honour as well.

In the novel Antic Hay (1923) Haldane was parodied by his friend Aldous Huxley as an obsessive self-experimenter described as "the biologist too absorbed in his experiments to notice his friends bedding his wife".[120]

Quotations

Oxford University Museum of Natural History display dedicated to Haldane and his reply when asked to comment on the mind of the Creator

Publications...

https://en.wikipedia.org/wiki/J._B._S._Haldane

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