SCIENTIFIC CAREER

J. B. S. Haldane was educated at Eton and Oxford, graduating with distinction in classics in 1915. His further education was interrupted when he joined the Black Watch battalion to fight in the First World War. He fought with bravery and distinction. His life was in constant danger when he was fighting in the trenches in France when almost all the men in his battalion were killed. He was saved mainly because his uncle, Lord Haldane, was Minister for War and managed to transfer his nephew to a safer job in Scotland. After the war in 1919, Haldane received a Fellowship in Physiology at New College, Oxford, where he pursued research in both physiology and genetics. In 1923, he was appointed to the Dunn Readership in Biochemistry under Professor F. G. Hopkins at Cambridge University. In addition to research and teaching in biochemistry, Haldane found time to write most of his important mathematical papers on the “mathematical theory of natural selection,” which helped found population genetics along with the independent contributions of Fisher and Wright. Simultaneously, he conducted outstanding research in physiology and biochemistry, deriving the law of steady-state kinetics in enzyme chemistry. Haldane (1932) summarized his early research in population genetics in his book The Causes of Evolution. He continued to publish brilliant papers subsequently, many of them becoming the foundations for new areas of research: genetic loads and the cost of natural selection (Haldane 1957), infectious disease and selection, estimation of human mutation rates, linkage and human gene mapping, rates of evolution (Haldane 1949), and the biochemistry of gene action, to name a few.

From 1933 to 1937, he was Professor of Genetics at University College, London. In 1937, he was elected to the Weldon Chair of Biometry where he continued to each and direct research in biometry, statistics, human genetics and several other subjects. In 1957, both Haldane and his wife migrated to India to accept an invitation from Prof. P. C. Mahalanobis, Director of the Indian Statistical Institute in Calcutta. Mahalanobis was a close friend and advisor to the then Prime Minister of India, Jawaharlal Nehru.

Haldane's numerous publications include more than 400 papers (for most of which he was the sole author) and 24 books as well as several hundred popular articles. Summaries of his life and work can be found in Dronamraju (1968, 1985, 1987, 1990, 1995, 2009) and Crow (2004).

REASONS FOR MOVING TO INDIA

Complex reasons motivated Haldane to abandon his position at University College London and move to India in 1957. His professional reputation and prestige were at their maximum, yet Haldane chose to turn his back on Europe and other active centers of scientific research in the world. The most important reason was undoubtedly his increasing dissatisfaction with the politics of Great Britain at that time. He was not comfortable living in the east or west block during the cold war years. He was attracted to the neutralist policy of Prime Minister Jawaharlal Nehru of India. Several years earlier, he had abandoned Marxist politics and was not interested in living in any countries of the communist bloc.

Another important reason was his desire to pursue research on Indian plant and animal species and human populations. Ever since his first visit to India in 1917, when he was sent to a British Army Hospital to recuperate from wounds he received in Mesopotamia (Iraq), Haldane frequently mentioned his desire to return to India. Upon his arrival in 1957, he initiated at once several research projects of an ecological and biometrical nature. These are briefly described below. Furthermore, as a classical scholar, he was drawn to India's ancient civilizations, religions, languages, and cultures. He often enjoyed quoting passages and verse in several Indian languages during his scientific lectures, just as he used to quote from Latin and Greek classics in Europe.

The job offers for both Haldane and his wife (his former student and biologist Helen Spurway) came at an opportune time from the Indian Statistical Institute in Calcutta. He was close to retirement from University College London. Furthermore, he was looking forward to enjoying the warm climate and clear sky in India. One of his great passions was observing the stars in the Indian skies.

LIFE AND RESEARCH IN INDIA

Haldane was a Research Professor at the Indian Statistical Institute in Calcutta from 1957 to 1961, where four of us, S. K. Roy, T. A. Davis, S. D. Jayakar and myself, closely worked with him as his research associates. I completed my research for a Ph.D. in genetics, which was awarded later. When political differences with the Director, P. C. Mahalanobis, arose later, Haldane resigned and moved to Bhubaneswar in the State of Orissa, where the Chief Minister (equivalent to a state Governor in the United States) offered to build him an Institute for Genetics and Biometry. However, his remaining life was too short. He died there on December 1, 1964 after cancer surgery. I worked closely with Haldane both at Calcutta (1957–1961) and at Bhubaneswar (1962–1963).

DARWIN AND WALLACE IN INDIA

Haldane's genetic research was primarily influenced by the Darwinian tradition. His major contribution to science was his mathematical theory of evolution (natural selection), which was an interpretation of Darwin's theory of evolution in the context of Mendelian genetics. After his arrival in India, Haldane saw Darwin from a new perspective.

Haldane regarded Darwin's work in botany as his most original and important contribution to science. He wrote: “In my opinion, however, Darwin's most original contributions to biology are not the theory of evolution but the great series of books on experimental botany published in the latter part of his life” (Dronamraju 2009, p. 212). That work includes books on insectivorous plants, climbing plants, sexuality in plants, and inbreeding in plants. Charles Darwin is best known for his work on evolution. However, that subject had also been discussed by many others including his grandfather, long before Darwin. His experiments on plants, although not so well known, are completely original. Haldane encouraged us to follow Darwin's research in botany. But Haldane wondered what Darwin's attitude was that led to his discoveries. Darwin was most open minded in his attitude toward all living beings. Haldane wrote: “Darwin did not draw a sharp line between earthworms and the old gentlemen who had failed to interest him in mathematics at Cambridge” (Dronamraju 2009, p. 213). From the biographical account of his son Francis, we know that Darwin was totally immersed in his experiments and observations. Indeed, he was so dedicated that he used human attributes when describing his experimental animals and plants.

Many of the research projects that were undertaken under Haldane's direction were quantitative biological studies. Haldane had earlier come to the conclusion that the denseness of the tropical flora could result in interdependence involving both competition and cooperation between species. Haldane wrote: “One would expect symbiotic relations between flowering plants to be much commoner in regions with a diverse flora than in those where a natural plant community often consists of few members … if such symbiotic regulations are common, they should be looked for between different genotypes of the same species” (see Dronamraju 1987, p. 221). The experiments of my colleague, S. K. Roy (Roy 1960), were concerned with the effect of symbiosis between different varieties of rice on their grain yields. Two varieties may be planted either fully mixed, in alternate rows, or in separate halves of the same plot surrounded by a dam. In 31 experiments, only 5 showed a gain of >10% over the mean of the two varieties, and all of them showed some gain over the better yielding of the two.

Some research projects followed what Bateson (1894) called “meristic variation,” that is to say, variation in like parts, such as petals, vertebrae, or teeth, in which Haldane (1958) was deeply interested. It remained for Gruneberg (1952), an associate of Haldane's at University College London, to show that it is akin to the variation in size of the third upper molars in some strains of mice, which he termed “quasi-continuous” variation. In India, Haldane's associates studied meristic variation in local plant and animal species. For instance, Roy (1963) studied the variation in petal numbers on plants of Nyctanthes arbor-tristis and Jasminum multiflorum. The plants showed floral teratology toward the end of the flowering season. Double flowers occurred frequently in late season. The variance increased toward the end of the flowering season. One of my studies was concerned with variation in the style lengths of flowers of Bauhinia acuminata. I also noted heterostyly in this species, which involved, unlike the classical heterostyly described by Darwin in Primula, both short- and long-styled flowers occurring on the same plant and the short-styled flowers being female sterile (Dronamraju 1960a). In his Presidential address to the Centenary and Bicentenary Congress at the University of Malaya in Singapore (December 2, 1958), Haldane stated: “It was (Alfred Russell) Wallace, above all, who saw that certain biological problems are forced on one by a study of tropical life, which are by no means so obvious in temperate climate” (Haldane 1960, p. 713).

ANIMALS HAVE RIGHTS AND DUTIES

Haldane stated that Christian theologians drew a sharp distinction between humans and other species. Hence, Darwin's argument that human beings have descended from animals created a furor among Europeans (Dronamraju 2009, p. 211), whereas many eastern religions such as Hinduism, Buddhism, and Jainism have long recognized that animals have rights and duties. Indian art and religion depicted divinity in various animal forms. Kindness toward other species has long been an accepted tradition, at least in theory, if not always in practice, in India. There are no parallels in the traditions of Europe and North America. Furthermore, there is still some opposition to teaching evolution in some Christian communities, especially in the southern United States. This is in contrast to the situation in India where many Hindus see similarities between vertebrate evolution and the successive incarnations of Vishnu.

Not only did Haldane interpret Darwinism from the Indian philosophical and religious point of view, but also he developed several research projects on Indian species of plants and animals that are in the Darwinian tradition. As his students, we were all encouraged to conduct research on different projects in that tradition. There were certain characteristics that are common to all these projects. Quantifying results as much as possible and the utilization of very simple methods were characteristic of Haldane's approach to science: (a) Much of our research involved measuring biological variation, especially meristic variation, that is to say, variation in like parts such as petal numbers in flowers, venation in leaves, number of teeth, or bilateral spotting in caterpillars, etc.; (b) research involved local resources as much as possible; (c) the projects did not require much financial support; (d) the methods employed were simple but the results had scientifically profound consequences; (e) the observations must be quantified wherever possible; (f) the project should be flexible enough to facilitate the incorporation of any new and unexpected results and the resulting change in direction or methods; (g) it is necessary to have several ongoing projects simultaneously to achieve quick results as there will always be some lines of pursuit that may take longer or not bear any fruit at all; and (h) the results should be published immediately.

Some of the projects were an imitation of Darwin's experiments. One such example is a project by Roy (1957) on the estimation of the volume of soil brought to the surface by earthworms per acre per year and its ecological implications. Another project, which I followed, is the investigation of color preferences of lepidoptera among different colored flowers and their impact on speciation (Dronamraju 1960b, 1985). By their preferential visits to different varieties of a plant species differing in flower color, insect pollinators can create reproductive isolation within a species. By means of increasing such isolation (for instance, it could happen if there is a new flower color mutant appearing in the plant population) further divergence can follow, ultimately leading to complete isolation and speciation. Such speciation has been called “sympatric speciation,” as opposed to “allopatric” or “geographic speciation,” where such geographic factors as floods, earthquakes, and simple geographic distance can result in reproductive barriers within a species, as emphasized by Haldane's friend Ernst Mayr (Mayr 1942). Haldane believed that such research projects followed the Darwinian tradition and would have interested Darwin greatly. Another colleague, T. A. Davis (Davis 1962) pursued research on coconut palms, showing that foliar asymmetry in their crowns is not inherited. Surprisingly, Davis (1963) discovered that palms with left-handed coiling in their crowns produce more nuts than those palms with right-handed spiraled crowns.

Among other studies, Helen Spurway (Mrs. Haldane) conducted biometrical investigations of the life cycle of the tussore silk moth, Antheraea mylitta (Spurway and Dronamraju 1962), and the nest-building activity of the solitary wasp, Sceliphron, in collaboration with Jayakar and Dronamraju (Spurway et al. 1964). Anthropologist A. K. Ray conducted population studies of toe anomalies and their possible genetic basis. Haldane commented that such studies are easier to follow in India because many people have bare feet or wear sandals, unlike western countries where shoes and boots are worn by many (Ray and Haldane 1965).

Haldane drew attention to Aristotle's logic and his difficulties in classifying animals, which he based on similarities (Dronamraju 2009, p. 214). Darwin, on the other hand, foreshadowed a logic based on differences. In The Origin of Species, Darwin (1859, p. 268) wrote: “Systematists will have only to decide (not that this will be easy) whether any form be sufficiently constant and distinct from other forms, to be capable of definition, and if definable whether the difference be sufficiently important to deserve a specific name … Hence, without rejecting the consideration of the present existence of intermediate gradations between any two forms, we shall be led to weigh more carefully and to value higher the actual amount of difference between them.” Haldane regarded Darwin's statement as the starting point for a whole new program in statistics (Haldane 1959).

THEORETICAL POPULATION GENETICS

In several papers, some in collaboration with S. D. Jayakar (Haldane and Jayakar 1963), Haldane returned to finding solutions for the unsolved problems of theoretical population genetics that he first initiated in the 1920s. The titles of some of these papers are “The selection of double heterozygotes,” “Natural selection in a population with annual breeding but overlapping generations,” “Polymorphism due to selection of varying direction,” “The elimination of double dominants in large random mating populations,” and “Natural selection in a population with annual breeding but overlapping generations.” At the same time, Haldane enjoyed returning to nature as was evident in his participation, with Helen and Jayakar, in bird watching, observing wasp and fish behavior, and other studies of an ecological nature.

Throughout his life, Haldane enjoyed being the human “calculator” for the data collected by his colleagues. He followed the same practice in India. He enjoyed nothing better than to sit in his arm chair and analyze data collected by others and devise novel statistical tests, often extracting more information than the investigator had hoped for. He was fond of doing complex sums in long hand, refusing to use even the simplest calculators that were available then.

Haldane rarely took time out from his mathematical labors. However, he indulged himself occasionally, for instance, in writing such pieces as “A defense of beanbag genetics” (Haldane 1964a) in response to Mayr's (1959) criticism of the early papers in beanbag genetics (see Dronamraju 2010). Mayr criticized the mathematical theory of natural selection, which was founded by Haldane, Fisher, and Wright, because genes were treated, for the sake of mathematical convenience, as noninteracting independent units. In Mayr's view, this was similar to the work of early Mendelians who kept beans of various colors in different bags while counting Mendelian ratios. With his tongue firmly in cheek, Haldane enjoyed writing a “defense” of his work, arguing that the mathematical theory provided a “scaffolding” within which a reasonably secure theory expressible in words may be built up. It is amusing to read his justification, “Fisher is dead, but when alive preferred attack to defense. Wright is one of the gentlest men I have ever met, and if he defends himself, will not counterattack. This leaves me to hold the fort, and that by writing rather than speech” (Haldane 1964a, p. 344).

HALDANE AT THE END

Haldane's poem, “Cancer is a Funny Thing,” which was written shortly before his death and published in many popular magazines and newspapers, reflected his courage when facing almost certain death. While he was visiting the United States in October 1963 to attend a conference on the origin of life in Florida, Haldane found out that he was suffering from rectal bleeding. On his return journey, stopping in London, Haldane found out that he had rectal cancer and was operated on instantly at University College Hospital. While recuperating in London, he wrote the poem “Cancer is a Funny Thing,” which amused and irritated his readers in equal numbers. I quote a few lines (Haldane 1964b):I wish I had the voice of HomerTo sing of rectal carcinoma,Which kills a lot more chaps, in fact,Than were bumped off when Troy was sacked.Yet, thanks to modern surgeon's skills,It can be killed before it killsUpon a scientific basisIn nineteen out of twenty cases.My final word, before I'm done,Is “Cancer can be rather fun”.Thanks to the nurses and Nye BevanThe NHS² is quite like heavenProvided one confronts the tumourWith 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 bitTo help the surgeons make us fit.

CONCLUDING REMARKS

Haldane's life and activities in India left an indelible impact on Indian science. He was especially inspiring to the younger generation who were happy to see him challenging the authorities and bureaucracy when they were not able to do so themselves. The research projects undertaken by his colleagues provided a new impetus to the development of certain branches of science, such as biometry, population genetics, human genetics, and animal behavior.