Thirty-five years ago, a renowned British biologist branded England "a criminal state" and migrated to India. Today, in the year of his birth centenary, J B S Haldane is remembered as a scientist …

-- JOHN BURDON Sanderson Haldane, who was born November 5, 1892, is remembered today mainly for developing a quantitative theory of evolution. He was also a superb populariser of science, a strong believer in rationality and a Marxist. Exuberantly boyish in many ways, he was also kind and gentle.

When Haldane was born, his father, John Scott Haldane, was well on his way to being recognised as an authority on physiology. The young Haldane's first steps in science were taken as an infant "guinea pig" in his father's experiments. Haldane was a precocious child who could read by age three and write by five. Precocity was combined with a remarkable memory (even in later years, it is said, Haldane had no need to look up a scientific reference when writing his papers), intellectual brilliance and a strongly pugnacious nature. A fight was never far off when Haldane was around -- a quality he retained all his life.

After preparatory school, where he earned a reputation of teaching mathematics to his teacher, Haldane went on to Eton, where he was extremely unhappy. This may be a reason for his strong anti-establishment attitude in later life. Oxford followed and he enjoyed his sojourn there, studying mathematics, European classics and philosophy.
Love of Arts His love of what we would call the Arts was enduring and he enbellished his writings with quotations from Greek and Latin, Christian theology and even the Hindu scriptures. A string of academic appointments in some of Britain's most prestigious educational institutions followed, including a fellowship at Oxford (1919-1922) and the Weldon Professorship of Biometry at University College, London (1937-1957).

Haldane married twice. His first wife, Charlotte, was a novelist and journalist; his second wife, Helen Spurway, was a geneticist and student of animal behaviour. Haldane and Helen Spurway migrated to India in 1957 and stayed there for seven years until his death in December 1964. Helen Spurway carried on with her research in India and survived her husband by 12 years. The Haldanes did not have any children.

A remarkable piece of research carried out by the young Haldane derived from his recognition, when just 16, that some of the data presented in a paper by the mouse geneticist A D Darbishire did not fall into the Mendelian expectation of independent assortment (the chance of any gene being passed down from parent to offspring is independent of the chance of any other gene being passed down). Haldane had discovered the phenomenon of linkage, one of the first hints that genes, the particles of heredity, were not free to "move" independently of one another but were instead physically associated in the manner of beads on a string. He proceeded to verify this hypothesis by means of breeding experiments carried out at home with the assistance of his sister Naomi and a friend. Unfortunately, his paper on this was delayed by the First World War and by then, evidence for linkage had been derived independently by T H Morgan in New York.
Evolutionary theory Charles Darwin's main claim to greatness is that he put forward an overwhelmingly plausible theory of evolution. Since then, two distinct lines of research have fuelled what has come to be called "post-Darwinian synthesis". On the one hand, there was field work aimed at strengthening the base that Darwin had built; on the other, there was the need to prove beyond all doubt that the principles of genetics enunciated by a Bohemian monk, Gregor Mendel, provided the logical underpinning for natural selection (as Darwin's theory is called). This task, known as the quantitative theory of evolution, or population genetics, was accomplished by the early 1930s by four men: S S Chetverikov in Russia, Sewall Wright in USA and R A Fisher and Haldane in England.

Starting in 1924, Haldane began to develop in a series of papers a comprehensive theory of population genetics. These papers were later compiled and published as a book in 1932 entitled The Causes of Evolution. Even though he continued to work in the field for the rest of his life, he does not seem to have attempted again a comparably wide-ranging look at evolution.

The aim of his early papers was set out clearly. "In order to establish the view that natural selection is capable of accounting for the known facts of evolution, we must show not only that it can cause a species to change, but that it can cause it to change at a rate which will account for present and past transmutation," he wrote.

In his evolutionary studies, Haldane often had in mind a practical problem to which he wanted a specific, preferably numerical, answer. For example, how many generations would it need for a beneficial trait with a selective advantage of one-tenths of 1 per cent to lead to a gene frequency change from .001 to .01? Or, what were the rates of evolution in the lineage leading to the modern horse?

Though such questions might lead one to think that Haldane's evolutionary studies had a narrow focus, such thinking would be unfair. But because the problems tackled by Haldane were highly diverse, he has left a legacy that is both rich and disjointed. Population genetics has not solved all the problems of evolutionary biology. What it has done -- and this is a tremendous achievement -- was to show that a certain set of approaches led to possible solutions to the problems of evolution.

"Mapping" genes
Haldane was also the first to derive a formula for "mapping" genes along the chromosome. This is a process comparable approximately to drawing a geographical map on the basis of distances measured locally (the formula popularly used today is based on a modification evolved by D D Kosambi). This, too, was with a potential application in mind as, for example, if a gene with a benign but visible effect (for instance, colour blindness) is closely associated with another gene that can have severe deleterious effects but whose presence is masked (such as haemophilia), the presence of the first gene can alert a medical practitioner to look out for the second, thereby enabling early therapy or genetic counselling.

Haldane was a pioneer in the application of genetics to biochemistry. His early appreciation of the importance of A E Garrod's studies on inborn metabolic deficiencies and, much later, his association with the work on the colours of flowers at the John Innes Horticultural Institution, led him to consider the "one gene, one enzyme" hypothesis well before the individuals usually credited with its enunciation, US geneticists G W Beadle and E Tatum. In fact, Haldane raised the possibility as early as 1920 that a gene makes a particular chemical species of enzymes, conceding, nevertheless that he got the idea from Cuenot.

In biochemistry, Haldane (together with G E Briggs) correctly interpreted the kinetics of the simplest enzyme reaction and wrote an influential textbook on enzymes. Among non-evolutionary biologists, Haldane's name is most widely associated with the theory he developed -- independently of the Soviet biologist A I Oparin -- for the origin of life. Haldane's basic idea was that in the primeval oxygen-free (and so reducing) atmosphere, the action of ultraviolet light on a mixture of carbon dioxide, water and ammonia could lead to the formation of more complex molecules, which, given a sufficient concentration with "the consistency of hot dilute soup", might lead to the formation of proteins. Typically for Haldane, the hypothesis is tossed off as an aside in two paragraphs of an article on Louis Pasteur, fermentation and other matters, written for a general readership in The Rationalist Annual.

Had Haldane not taken to biology, he might have become a professional mathematician for his deep interest in mathematics was an outgrowth of problems in genetics or evolution. To a large extent, his interest involved devising improved statistical methods for analysing data and estimating "inverse" probabilities. Both Haldane and his wife Helen were interested in animal behaviour and in a marvellous joint paper, they analysed the dance language of the honey bee, from the perspective of information theory.

A born communicator
Haldane was a peerless communicator of scientific ideas. Besides the money it brought him, he wrote extensively because he had a passionate belief in the value of knowledge and the duty of sharing it with others. His topics covered a wide range from the uses of philosophy to how bees communicate, because he was convinced that there were no bounds to science. Haldane had the ability to convey meaning in simple but vivid language as in this example: ".... I find it no easier to picture a completely socialised British Empire or United States than an elephant turning somersaults or a hippopotamus jumping a hedge."

Haldane could tackle almost any subject and could also weave a number of themes together within the same piece. In a perceptive essay on animal communication and the possible origin of the human language, he offered a stage direction from Shakespeare as support for his suggestion that muscular activity is not only a reflection of a mental state but also serves to reinforce that state.

Haldane was physically a large man, and he described himself as "a man of violence by temperament and training". He could be both friendly and insulting, but he contended that he insulted everyone impartially. He applied a strict test of rationality not only to his own words and actions but also to the actions of others, which often led to problems. And, though he would pay from his own pocket the airfare for a student wanting to attend a conference, he could fumble successfully when paying for tea in the canteen. As he was never shy of expressing his opinion in public on any topic, he was perennially quotable.

Extremely stubborn and argumentative in scientific matters, he was unselfish and generous in giving credit to colleagues and students. There are many instances of papers by others, whose implications were appreciated and referred to first by Haldane.

Like many intellectuals of his time, Haldane was attracted by the egalitarian philosophy of socialism. Always materialistic and uncompromisingly rational, he was once asked if he believed in God. Prompted by the knowledge that insects form an overwhelming majority of animal species, his reply was, "I do not know if God exists, but if He does, He must be inordinately fond of beetles." When John Maynard Smith (Haldane's most distinguished student) was in India many years ago, he related a story illustrating Haldane's ability to respond rapidly to a demanding situation. Prof (as Haldane was known) and his students had been out drinking in a pub and on the way back, a carelessly thrown match set the car's upholstery alight. Without wasting any time, Prof stopped the car, ordered the students to jump out and look the other way, and then proceeded to douse the fire by a judicious spraying of the contents of his bladder.

The Lysenko affair
Though Haldane remained a Marxist all his life, he maintained formal allegiance to the Communist Party of Great Britain for just eight years, giving it up in 1950. Events in the then USSR forced a difficult choice on Haldane: sympathy with the Soviet people versus increasing evidence of the serious damage caused to Soviet biology by the activities of Trofim Lysenko, a favourite of Joseph Stalin.

Lysenko gained notoriety with a sensational claim that he never justified that by inducing certain seeds to germinate earlier than usual, their descendants would be better able to survive the early frosts in the extreme northerly latitudes. Not content with striking out in this manner at the heart of conventional Darwinian thinking, Lysenko also suggested that by performing grafts he could cause different species of plants to hybridise. This added another blow at genetic orthodoxy. Not only were these theories of Lysenko disastrous for Soviet agriculture, but in the attempt to justify his beliefs, genetical thinking was ridiculed, the Soviet Union lost its place in contemporary biology and many leading Soviet biologists were deprived of work, security and, on occasion, their lives.

In this situation, Haldane's attitude was at best equivocal. Though he never abandoned his scientific views, he bent over backwards to suggest that Lysenko might be on to something after all. Eventually, he left the Communist Party because functioning within the rigid norms of official Marxist dogma and as part of a highly organised structure was much too un-Haldanian. Even so, he never dismissed Lysenko's claims in their entirety and, interestingly, he may even have been technically correct in holding to this attitude.

JBS in India
The Anglo-French invasion of Suez provided a convenient and dramatic stimulus for the Haldanes to leave England, branding it "a criminal state." Also, Haldane had developed a genuine interest in India and the prospects for doing useful scientific work here. His readings in Hindu, Buddhist, Jain and Islamic philosophy bolstered this interest. His friendship with P C Mahalonobis led him to join the Indian Statistical Institute (ISI) in Calcutta and he went on to become an Indian citizen.

In India, Haldane collected a small band of enthusiastic young students and put them on to interesting problems. Though he continued to travel and lecture, he also maintained the high standards of the Journal of Genetics, which he had brought with him to India. However, problems were not long in coming, caused in part by a bureaucratic structure that was also quite vague, resulting in the nature of his relationship with the ISI remaining hazy throughout his time there.

As was often the case with Haldane, when the break came, it was over an episode that need not have been more than an irritation by itself. It involved Mahalonobis countermanding, without any warning and at the last moment, Haldane's arrangements for displaying the work of his group during a visit by Alexei Kosygin who was soon to become prime minister of the USSR. This was provocation enough for Haldane to resign.

What followed was even worse than his ISI experiences. At the suggestion of then vice president S Radhakrishnan, the head of the Council for Scientific and Industrial Research (CSIR), M S Thacker, persuaded Haldane to start a genetics and biometry unit in Calcutta. He was assured that buildings, laboratory and other facilities would follow in due course, but nothing was done and haldane began referring to CSIR as the Council for the Suppression of Independent Research. Within a year, Haldane left the CSIR stating in his letter of resignation something that remains depressingly true even today, "It is the intolerable conditions imposed by bureaucrats and not the low salaries or the lack of equipment which cause so many Indians to take up posts abroad."

Haldane's next posting was in Bhubaneswar, thanks to the enterprise and initiative of chief minister Biju Patnaik. Haldane was appointed head of an entirely independent research establishment, where he worked with his young colleagues from Calcutta, including S D Jayakar, with whom he published papers on population genetics that are as enduring as the best of Haldane's early work.

During his stay at Bhubaneswar, Haldane, who had always stressed the suitability of ethology, or the study of animal behaviour, as a subject of biological research for countries lacking funds for elaborate technical equipment, and his wife, Helen, who was a behavioural biologist herself, worked with their students on such problems as nest-building of wasps and the consequences for evolution of selective visits by butterflies to flowers.

On December 1, 1964, Haldane died of rectal carcinoma. Out of misplaced kindness, the doctors who operated on him in Britain had left him with the impression he was out of immediate danger. When the truth about his condition became known, he complained he would have planned his last days quite differently had he been told the truth. In keeping with his express instructions that he wanted his body to be put to use, it was sent to a medical college for dissection.

Haldane's life was one of child-like wonder, described aptly in his own words: "Our only hope of understanding the universe is to look at it from as many different points of view as possible. This is one of the reasons why the data of mystical consciousness can usefully supplement those of the mind in its normal state. Now, my own suspicion is that the universe is not only queerer than we suppose, but queerer than we can suppose.

"I suspect there are more things in heaven and earth than are dreamed of, or can be dreamed of, in any philosophy. That is the reason why I have no philosophy myself, and must be excused for dreaming."

---Vidyanand Nanjundiah is with the Centre for Theoretical Studies and Developmental Biology and Genetics Laboratory at the Indian Institute of Science, Bangalore. This article is based on a recent lecture that he had delivered on J B S Haldane at the Indian Institute of World Culture, Bangalore.

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