Copyright Ó 2010 by the Society of America DOI: 10.1534/genetics.110.116632

Perspectives

Anecdotal, Historical and Critical Commentaries on Genetics J. B. S. Haldane’s Last Years: His Life and Work in India (1957–1964)

Krishna Dronamraju1 Foundation for Genetic Research, Houston, Texas 77227

ABSTRACT J. B. S. Haldane (1892–1964) was an outstanding scientist and a polymath who contributed significantly to , genetics, biochemistry, statistics, biometry, cosmology, and philosophy, although he himself possessed no formal qualifications in any branch of science. His early science training was provided by his father, University physiologist J. S. Haldane, and the rest was self-taught. The author came to know him well as his student during his last years, which were spent in India (1957–1964). Haldane’s unique intellectual qualities and the research undertaken by his associates in India during those years are described.

OHN Burdon Sanderson ( J. B. S.) Haldane (1892– range evolution in human populations. Human pop- J 1964) was an exceptional scientist whose mental ulation genetics is one of the several fields in which you powers were extraordinary. He possessed both great cannot move without encountering the approaches intelligence and a prodigious memory and was able to and analyses of Prof. Haldane at every step ...Some of divide his attention between two entirely different Prof. Haldane’s ingenious proposals ... have kept us subjects simultaneously. He was able, for instance, to sit busy with discussions for long years ...it is an enormous in a lecture room, writing a mathematical paper on privilege for us to be allowed to carry on this conference evolution while closely paying attention to a lecture in his presence and under his guidance’’ (Goldschmidt on a different topic as he demonstrated during the 1963, p. 5). During the 1930s and 1940s, Haldane was discussion at the end of the lecture. active in finding research positions for Jewish refugees Haldane’s last years in India are of interest because he who were fleeing Nazi . Prominent among them showed how an eminent scientist from the West can was biochemist Boris Chain who was introduced to transplant himself to a less developed country and still Howard Florey by Haldane; Chain and Florey’s sub- lead a productive and creative life during his last years. sequent collaboration led to the Nobel Prize that they He suggested research projects that could be under- shared with Fleming for their work on the discovery and taken by utilizing local resources, requiring no expen- development of penicillin as the first antibiotic (Haldane sive laboratories or equipment, while much of his own 1961). research was of theoretical and mathematical nature J. B. S. Haldane was a great popularizer of science, and hence not dependent on external funding. contributing numerous articles to newspapers and I came to know Haldane intimately during those popular magazines in several countries. A selection of years in India (1957–1964). In Calcutta, I shared a these essays has recently been published (Dronamraju house with the Haldanes in the suburb of Baranagore 2009). Many of these pieces were written in his ‘‘spare near the Indian Statistical Institute. We traveled to- time’’ while traveling on trains and planes. Haldane’s gether all over India and Europe for various scientific ‘‘popular’’ scientific essays are, in fact, much more than meetings. One memorable conference was held in that; they often contained original ideas that remain of on the ‘‘Genetics of Migrant and Isolate Populations’’ interest to scientists. oldschmidt (G 1963). On behalf of the organizing He never received a degree in any branch of science, committee, Elisabeth Goldschmidt invited Haldane to yet he became one of the great and influential scien- preside over the conference, with these words: ‘‘We may tists of the 20th century. Haldane, who was popularly consider ourselves fortunate indeed that Prof. Haldane known as either just ‘‘JBS’’ or ‘‘Prof’’ or simply ‘‘Hal- has accepted the presidency of this conference on short- dane’’ was a true polymath, a genius who possessed intimate knowledge of multiple disciplines. He made 1Address for correspondence: Foundation for Genetic Research, P.O. Box important contributions to several sciences as well as to 27701-0, Houston, TX 77227. E-mail: [email protected] philosophy, religion, logic, popular writing, and ethics.

Genetics 185: 5–10 (May 2010) 6 K. Dronamraju

igure Figure 1.—J. B. S. Haldane. Photograph courtesy of Klaus F 2.—J. B. S. Haldane surrounded by his associates, Patau, University of Wisconsin. on the campus of the Indian Statistical Institute, Calcutta, 1961, the author sitting, third from left. Photograph courtesy of the Indian Statistical Institute. In science, Haldane’s contributions covered physiology, genetics, biochemistry, statistics, biometry, cosmology, close friends, was a precocious child whose brilliance and history and philosophy of science, leaving a signi- was legendary from an early age. His father used to take ficant impact on each of these disciplines. As a popular him down mine shafts where he investigated the writer of science, Haldane covered even more subjects, physiological conditions. At the age of 3 or 4, he was including , physics, chemistry, mathematics, geol- already familiar with the terminology of , ogy, geography, astronomy, meteorology, politics, psychol- which he picked up from his father’s research. Once, ogy, nonviolence, military affairs, theology, and literature. when he fell down and cut his head, a doctor was called In addition to his monumental work in genetics, Haldane to treat the wound, and the young Haldane, looking at made important contributions to enzyme kinetics in the blood, asked whether it was oxyhemoglobin or biochemistry. ! As a young boy, his talent for math- In physiology, in particular, Haldane is best remem- ematics was put to test by his father who, on one occa- bered for his daring experiments in testing the physiolog- sion, forgot his log tables on a field trip and asked his ical effects of poisonous gas mixtures, cold , son to calculate a set. Young Haldane did so promptly. and higher atmospheric in which he employed Haldane’s scientific eminence today is principally himself as his own ‘‘guinea pig.’’ They were often painful based on his extensive series of papers on the mathe- experiments, causing convulsions and injuries. He also matical theory of Darwinian natural selection (evolu- conducted many diving experiments to investigate a tion). The independent contributions of Haldane and submarineaccidentfortheBritishNavy,onceagain R. A. Fisher in and Sewall Wright in the United employing himself as his own guinea pig. He suffered States led to the establishment of modern population permanent injury because of rapid during genetics. the experiments and back pain haunted him the rest of his life (Behnke and Brauer 1968). SCIENTIFIC CAREER Haldane’s distinguished relatives include his father John Scott Haldane, physiologist at New College, J. B. S. Haldane was educated at Eton and Oxford, Oxford University; his uncle Viscount Haldane, scholar graduating with distinction in classics in 1915. His and politician, who was Minister for War and Chancellor further education was interrupted when he joined the of the Exchequer in Prime Minister Asquith’s cabinet; Black Watch battalion to fight in the First World War. his great uncle Burdon Sanderson, first Waynflete He fought with bravery and distinction. His life was in Professor of Physiology at Oxford University; and his constant danger when he was fighting in the trenches in sister (Lady) , prolific writer of numer- France when almost all the men in his battalion were ous books of fiction, nonfiction, travel, history, politics, killed. He was saved mainly because his uncle, Lord and other subjects. (Lady Mitchison’s standing and Haldane, was Minister for War and managed to transfer influence are attested by Jim Watson’s dedicating his his nephew to a safer job in Scotland. After the war in memoir The Double Helix to her.) Haldane’s three 1919, Haldane received a Fellowship in Physiology at nephews are distinguished scientists in the life sciences. New College, Oxford, where he pursued research in Haldane was born in Oxford, England, on November both physiology and genetics. In 1923, he was appointed 8, 1892. Jack, as he was known to family members and to the Dunn Readership in Biochemistry under Pro- Perspectives 7 fessor F. G. Hopkins at Cambridge University. In Upon his arrival in 1957, he initiated at once several addition to research and teaching in biochemistry, research projects of an ecological and biometrical Haldane found time to write most of his important nature. These are briefly described below. Furthermore, mathematical papers on the ‘‘mathematical theory of as a classical scholar, he was drawn to India’s ancient natural selection,’’ which helped found population civilizations, religions, languages, and cultures. He of- genetics along with the independent contributions of ten enjoyed quoting passages and verse in several Indian Fisher and Wright. Simultaneously, he conducted out- languages during his scientific lectures, just as he used standing research in physiology and biochemistry, de- to quote from Latin and Greek classics in Europe. riving the law of steady-state kinetics in enzyme chemistry. The job offers for both Haldane and his wife (his Haldane (1932) summarized his early research in former student and biologist Helen Spurway) came at in his book The Causes of Evolution. an opportune time from the Indian Statistical Institute He continued to publish brilliant papers subsequently, in Calcutta. He was close to retirement from University many of them becoming the foundations for new areas of College . Furthermore, he was looking forward research: genetic loads and the cost of natural selection to enjoying the warm climate and clear sky in India. One (Haldane 1957), infectious disease and selection, esti- of his great passions was observing the stars in the Indian mation of human mutation rates, linkage and human skies. gene mapping, rates of evolution (Haldane 1949), and the biochemistry of gene action, to name a few. LIFE AND RESEARCH IN INDIA From 1933 to 1937, he was Professor of Genetics at University College, London. In 1937, he was elected to Haldane was a Research Professor at the Indian the Weldon Chair of Biometry where he continued to Statistical Institute in Calcutta from 1957 to 1961, where each and direct research in biometry, statistics, human four of us, S. K. Roy, T. A. Davis, S. D. Jayakar and myself, genetics and several other subjects. In 1957, both Haldane closely worked with him as his research associates. I and his wife migrated to India to accept an invitation completed my research for a Ph.D. in genetics, which was from Prof. P. C. Mahalanobis, Director of the Indian awarded later. When political differences with the Di- Statistical Institute in Calcutta. Mahalanobis was a close rector, P. C. Mahalanobis, arose later, Haldane resigned friend and advisor to the then Prime Minister of India, and moved to Bhubaneswar in the State of Orissa, where Jawaharlal Nehru. the Chief Minister (equivalent to a state Governor in the Haldane’s numerous publications include more ) offered to build him an Institute for than 400 papers (for most of which he was the sole Genetics and Biometry. However, his remaining life was author) and 24 books as well as several hundred popular too short. He died there on December 1, 1964 after articles. Summaries of his life and work can be found in cancer surgery. I worked closely with Haldane both at Dronamraju (1968, 1985, 1987, 1990, 1995, 2009) and Calcutta (1957–1961) and at Bhubaneswar (1962–1963). Crow (2004). DARWIN AND WALLACE IN INDIA REASONS FOR MOVING TO INDIA Haldane’s genetic research was primarily influenced Complex reasons motivated Haldane to abandon his by the Darwinian tradition. His major contribution to position at University College London and move to science was his mathematical theory of evolution (natural India in 1957. His professional reputation and prestige selection), which was an interpretation of Darwin’s were at their maximum, yet Haldane chose to turn his theory of evolution in the context of Mendelian genetics. back on Europe and other active centers of scientific After his arrival in India, Haldane saw Darwin from a new research in the world. The most important reason was perspective. undoubtedly his increasing dissatisfaction with the Haldane regarded Darwin’s work in as his most politics of Great Britain at that time. He was not original and important contribution to science. He wrote: comfortable living in the east or west block during the ‘‘In my opinion, however, Darwin’s most original contri- cold war years. He was attracted to the neutralist policy butions to biology are not the theory of evolution but the of Prime Minister Jawaharlal Nehru of India. Several great series of books on experimental botany published years earlier, he had abandoned Marxist politics and was in the latter part of his life’’ (Dronamraju 2009, p. 212). not interested in living in any countries of the commu- That work includes books on insectivorous plants, climb- nist bloc. ing plants, sexuality in plants, and inbreeding in plants. Another important reason was his desire to pursue Charles Darwin is best known for his work on evolution. research on Indian plant and animal species and hu- However, that subject had also been discussed by many man populations. Ever since his first visit to India in 1917, others including his grandfather, long before Darwin. His when he was sent to a British Army Hospital to recuperate experiments on plants, although not so well known, are from wounds he received in Mesopotamia (Iraq), Haldane completely original. Haldane encouraged us to follow frequently mentioned his desire to return to India. Darwin’s research in botany. But Haldane wondered what 8 K. Dronamraju

Darwin’s attitude was that led to his discoveries. Darwin biological problems are forced on one by a study of was most open minded in his attitude toward all living tropical life, which are by no means so obvious in beings. Haldane wrote: ‘‘Darwin did not draw a sharp line temperate climate’’ (Haldane 1960, p. 713). between earthworms and the old gentlemen who had failed to interest him in mathematics at Cambridge’’ ANIMALS HAVE RIGHTS AND DUTIES (Dronamraju 2009, p. 213). From the biographical ac- count of his son Francis, we know that Darwin was totally Haldane stated that Christian theologians drew a immersed in his experiments and observations. Indeed, sharp distinction between humans and other species. he was so dedicated that he used human attributes when Hence, Darwin’s argument that human beings have describing his experimental animals and plants. descended from animals created a furor among Euro- Many of the research projects that were undertaken peans (Dronamraju 2009, p. 211), whereas many eastern under Haldane’s direction were quantitative biological religions such as Hinduism, Buddhism, and Jainism have studies. Haldane had earlier come to the conclusion long recognized that animals have rights and duties. that the denseness of the tropical flora could result Indian art and religion depicted divinity in various animal in interdependence involving both competition and forms. Kindness toward other species has long been an cooperation between species. Haldane wrote: ‘‘One accepted tradition, at least in theory, if not always in would expect symbiotic relations between flowering practice, in India. There are no parallels in the traditions plants to be much commoner in regions with a diverse of Europe and North America. Furthermore, there is still flora than in those where a natural plant community some opposition to teaching evolution in some Christian often consists of few members ... if such symbiotic communities, especially in the southern United States. regulations are common, they should be looked for This is in contrast to the situation in India where many between different genotypes of the same species’’ (see Hindus see similarities between vertebrate evolution and Dronamraju 1987, p. 221). The experiments of my the successive incarnations of Vishnu. colleague, S. K. Roy (Roy 1960), were concerned with Not only did Haldane interpret Darwinism from the the effect of symbiosis between different varieties of rice Indian philosophical and religious point of view, but on their grain yields. Two varieties may be planted either also he developed several research projects on Indian fully mixed, in alternate rows, or in separate halves of species of plants and animals that are in the Darwinian the same plot surrounded by a dam. In 31 experiments, tradition. As his students, we were all encouraged to only 5 showed a gain of .10% over the mean of the two conduct research on different projects in that tradition. varieties, and all of them showed some gain over the There were certain characteristics that are common to better yielding of the two. all these projects. Quantifying results as much as Some research projects followed what Bateson possible and the utilization of very simple methods were (1894) called ‘‘meristic variation,’’that is to say, variation characteristic of Haldane’s approach to science: (a) in like parts, such as petals, vertebrae, or teeth, in which Much of our research involved measuring biological Haldane (1958) was deeply interested. It remained for variation, especially meristic variation, that is to say, Gruneberg (1952), an associate of Haldane’s at Uni- variation in like parts such as petal numbers in flowers, versity College London, to show that it is akin to the venation in leaves, number of teeth, or bilateral spotting variation in size of the third upper molars in some in caterpillars, etc.; (b) research involved local resources strains of mice, which he termed ‘‘quasi-continuous’’ as much as possible; (c) the projects did not require variation. In India, Haldane’s associates studied meristic much financial support; (d) the methods employed variation in local plant and animal species. For instance, were simple but the results had scientifically profound Roy (1963) studied the variation in petal numbers on consequences; (e) the observations must be quantified plants of Nyctanthes arbor-tristis and Jasminum multiflorum. wherever possible; (f) the project should be flexible The plants showed floral teratology toward the end of enough to facilitate the incorporation of any new and the flowering season. Double flowers occurred fre- unexpected results and the resulting change in direction quently in late season. The variance increased toward or methods; (g) it is necessary to have several ongoing the end of the flowering season. One of my studies was projects simultaneously to achieve quick results as there concerned with variation in the style lengths of flowers will always be some lines of pursuit that may take longer of Bauhinia acuminata. I also noted heterostyly in this or not bear any fruit at all; and (h) the results should species, which involved, unlike the classical heterostyly be published immediately. described by Darwin in Primula, both short- and long- Some of the projects were an imitation of Darwin’s styled flowers occurring on the same plant and the experiments. One such example is a project by Roy short-styled flowers being female sterile (Dronamraju (1957) on the estimation of the volume of soil brought 1960a). In his Presidential address to the Centenary and to the surface by earthworms per acre per year and its Bicentenary Congress at the University of Malaya in ecological implications. Another project, which I fol- Singapore (December 2, 1958), Haldane stated: ‘‘It was lowed, is the investigation of color preferences of lep- (Alfred Russell) Wallace, above all, who saw that certain idoptera among different colored flowers and their Perspectives 9 impact on speciation (Dronamraju 1960b, 1985). By in the 1920s. The titles of some of these papers are ‘‘The their preferential visits to different varieties of a plant selection of double heterozygotes,’’ ‘‘Natural selection species differing in flower color, insect pollinators can in a population with annual breeding but overlapping create reproductive isolation within a species. By means generations,’’ ‘‘Polymorphism due to selection of vary- of increasing such isolation (for instance, it could ing direction,’’ ‘‘The elimination of double dominants happen if there is a new flower color mutant appearing in large random mating populations,’’ and ‘‘Natural in the plant population) further divergence can follow, selection in a population with annual breeding but ultimately leading to complete isolation and speciation. overlapping generations.’’ At the same time, Haldane Such speciation has been called ‘‘sympatric speciation,’’ enjoyed returning to nature as was evident in his par- as opposed to ‘‘allopatric’’ or ‘‘geographic speciation,’’ ticipation, with Helen and Jayakar, in bird watching, where such geographic factors as floods, earthquakes, observing wasp and fish behavior, and other studies of and simple geographic distance can result in reproduc- an ecological nature. tive barriers within a species, as emphasized by Haldane’s Throughout his life, Haldane enjoyed being the human friend Ernst Mayr (Mayr 1942). Haldane believed that ‘‘calculator’’ for th e data collected by his colleagues. He such research projects followed the Darwinian tradition followedthesamepracticeinIndia.Heenjoyednothing and would have interested Darwin greatly. Another col- better than to sit in his arm chair and analyze data league, T. A. Davis (Davis 1962) pursued research on collected by others and devise novel statistical tests, often coconut palms, showing that foliar asymmetry in their extracting more information than the investigator had crowns is not inherited. Surprisingly, Davis (1963) dis- hoped for. He was fond of doing complex sums in long covered that palms with left-handed coiling in their hand, refusing to use even the simplest calculators that crowns produce more nuts than those palms with right- were available then. handed spiraled crowns. Haldane rarely took time out from his mathematical Among other studies, Helen Spurway (Mrs. Haldane) labors. However, he indulged himself occasionally, for conducted biometrical investigations of the life cycle of instance, in writing such pieces as ‘‘A defense of the tussore silk moth, Antheraea mylitta (Spurway and beanbag genetics’’ (Haldane 1964a) in response to Dronamraju 1962), and the nest-building activity of the Mayr’s (1959) criticism of the early papers in beanbag solitary wasp, Sceliphron, in collaboration with Jayakar genetics (see Dronamraju 2010). Mayr criticized the and Dronamraju (Spurway et al. 1964). Anthropologist mathematical theory of natural selection, which was A. K. Ray conducted population studies of toe anomalies founded by Haldane, Fisher, and Wright, because genes and their possible genetic basis. Haldane commented were treated, for the sake of mathematical convenience, that such studies are easier to follow in India because as noninteracting independent units. In Mayr’s view, many people have bare feet or wear sandals, unlike this was similar to the work of early Mendelians who western countries where shoes and boots are worn by kept beans of various colors in different bags while many (Ray and Haldane 1965). counting Mendelian ratios. With his tongue firmly in Haldane drew attention to Aristotle’s logic and his cheek, Haldane enjoyed writing a ‘‘defense’’ of his work, difficulties in classifying animals, which he based on arguing that the mathematical theory provided a ‘‘scaf- similarities (Dronamraju 2009, p. 214). Darwin, on the folding’’ within which a reasonably secure theory express- other hand, foreshadowed a logic based on differences. ible in words may be built up. It is amusing to read his In The Origin of Species, Darwin (1859, p. 268) wrote: justification, ‘‘Fisher is dead, but when alive preferred ‘‘Systematists will have only to decide (not that this will attack to defense. Wright is one of the gentlest men I be easy) whether any form be sufficiently constant and have ever met, and if he defends himself, will not distinct from other forms, to be capable of definition, counterattack. This leaves me to hold the fort, and that and if definable whether the difference be sufficiently by writing rather than speech’’ (Haldane 1964a, p. 344). important to deserve a specific name ...Hence, without rejecting the consideration of the present existence of HALDANE AT THE END intermediate gradations between any two forms, we shall be led to weigh more carefully and to value higher the Haldane’s poem, ‘‘Cancer is a Funny Thing,’’ which actual amount of difference between them.’’ Haldane was written shortly before his death and published in regarded Darwin’s statement as the starting point for a many popular magazines and newspapers, reflected whole new program in statistics (Haldane 1959). 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, THEORETICAL POPULATION GENETICS Haldane found out that he was suffering from rectal In several papers, some in collaboration with S. D. bleeding. On his return journey, stopping in London, Jayakar (Haldane and Jayakar 1963), Haldane re- Haldane found out that he had rectal cancer and was turned to finding for the unsolved problems operated on instantly at University College Hospital. of theoretical population genetics that he first initiated While recuperating in London, he wrote the poem 10 K. Dronamraju

‘‘Cancer is a Funny Thing,’’ which amused and irri- Darwin, C., 1859 On the Origin of Species. John Murray, London. tated his readers in equal numbers. I quote a few lines (2009, A facsimile edition of the first edition. Belknap Press of aldane Harvard University Press, Cambridge, MA.) (H 1964b): Davis, T. A., 1962 The non-inheritance of asymmetry in Cocos nu- cifera. J. Genet. 58: 42–50. I wish I had the voice of Homer Davis, T. A., 1963 The dependence of yield on asymmetry in coco- To sing of rectal carcinoma, nut palms. J. Genet. 58: 186–195. Dronamraju, K. R., 1960a Non-genetic polymorphism in Bauhinia Which kills a lot more chaps, in fact, acuminata L. J. Genet. 57: 299–311. ronamraju Than were bumped off when Troy was sacked. D , K. R., 1960b Selective visits of butterflies to flowers, a possible factor in sympatric speciation. Nature 186: 178. Yet, thanks to modern surgeon’s skills, Dronamraju, K. R. (Editor), 1968 Haldane and Modern Biology. Johns Hopkins University Press, Baltimore. It can be killed before it kills Dronamraju, K. R., 1985 Haldane: The Life and Work of J. B. S. Upon a scientific basis HaldaneWithSpecialReferencetoIndia(with a foreword by Naomi Mitchison). Aberdeen University Press, Aberdeen, UK. In nineteen out of twenty cases. Dronamraju, K. R., 1987 On some aspects of the life and work of John Burdon Sanderson Haldane, F.R.S., in India. Notes Rec. R. My final word, before I’m done, Soc. Lond. 41: 211–234. ronamraju Is ‘‘Cancer can be rather fun’’. D , K. R. (Editor), 1990 Selected Genetic Papers of J. B. S. Haldane (with a foreword by James F. Crow). Garland Publish- Thanks to the nurses and Nye Bevan ing, New York/London. ronamraju 2 D , K. R. (Editor), 1995 Haldane’s Daedalus Revisited (with a The NHS is quite like heaven foreword by Joshua Lederberg). Oxford University Press, Oxford. ronamraju Provided one confronts the tumour D , K. R. (Editor), 2009 What I Require from Life: Writings on Science and Life from J. B. S. Haldane (with a foreword by Sir With a sufficient sense of humour. Arthur C. Clarke and preface by James F. Crow). Oxford Univer- sity Press, Oxford. I know that cancer often kills, Dronamraju,K.R.,2010 Haldane, Mayr and Beanbag Genetics. Oxford But so do cars and sleeping pills; University Press, Oxford. Goldschmidt, E. (Editor), 1963 The Genetics of Migrant and Isolate And it can hurt one till one sweats, Populations. Williams & Wilkins, Baltimore. Gruneberg, H., 1952 Genetical studies in the skeleton of the So can bad teeth and unpaid debts. mouse, IV. Quasi-continuous variation. J. Genet. 51: 95–114. A spot of laughter, I am sure, Haldane, J. B. S., 1932 The Causes of Evolution. Longmans, Green, London. Often accelerates one’s cure; Haldane, J. B. S., 1949 Disease and evolution. Ric. Sci. 19(Suppl.): 2–11. aldane So let us patients do our bit H , J. B. S., 1957 The cost of natural selection. J. Genet. 55: 511–524. To help the surgeons make us fit. Haldane, J. B. S., 1958 The theory of evolution, before and after Bateson. J. Genet. 56: 1–17. Haldane, J. B. S., 1959 Natural selection, pp. 101–149 in Darwin’s Biological Work: Some Aspects Reconsidered, edited by P. R. Bell. CONCLUDING REMARKS Cambridge University Press, Cambridge, UK. Haldane, J. B. S., 1960 The theory of natural selection today. Na- Haldane’s life and activities in India left an indelible ture (Lond.) 183: 710–718. impact on Indian science. He was especially inspiring to Haldane, J. B. S., 1961 An autobiography in brief, pp. 1–6 in Illus- the younger generation who were happy to see him trated Weekly of India, Bombay, India. (Reprinted in Dronamraju, challenging the authorities and bureaucracy when they K. R. (Editor) Selected Genetic Papers of J. B. S. Haldane, Foreword by James F. Crow, New York: Garland Publishing, Inc.) were not able to do so themselves. The research projects Haldane, J. B. S., 1964a A defense of beanbag genetics. Perspect. undertaken by his colleagues provided a new impetus to Biol. Med. 7: 343–359. aldane the development of certain branches of science, such as H , J. B. S., 1964b Cancer is a funny thing, p. 298 in The New Statesman, London (21 February 1964). biometry, population genetics, human genetics, and Haldane, J. B. S., and S. D. Jayakar, 1963 Polymorphism due to animal behavior. selection depending on the composition of a population. J. Genet. 58: 318–323. I am most grateful to James F. Crow for his thoughtful comments Mayr, E., 1942 Systematics and the Origin of Species. Columbia Univer- and suggestions. sity Press, New York. Mayr, E., 1959 Where are we? Cold Spring Harbor Symp. Quant. LITERATURE CITED Biol. 24: 1–14. Ray,A.K.,andJ.B.S.Haldane, 1965 The genetics of a common Bateson, W., 1894 Materials for the Study of Variation Treated With Indian digital abnormality. Proc. Natl. Acad. Sci. USA 53: 1050–1053. Especial Regard to Discontinuity in the Origin of Species. Macmillan, Roy, S. K., 1957 Studies on the activity of earthworms. Proc. Zool. London. Soc. Bengal 10: 81–98. Behnke, A. R., and R. W. Brauer, 1968 Physiologic investigations in Roy,S.K,1960 Interactionbetweenricevarieties.J.Genet.57: 137–152. diving and inhalation of gases, pp. 267–276 in Haldane and Mod- Roy, S. K., 1963 The variation of organs of individual plants (with 2 ern Biology, edited by K. R. Dronamraju. Johns Hopkins Univer- plates). J. Genet. 58: 147–176. sity Press, Baltimore. Spurway, H., and K. R. Dronamraju, 1962 The biology of the two Crow, J. F., 2004 Haldane’s ideas in biology with special reference commercial quality cocoons spun by Antheraea mylitta (Drury) to disease and evolution, pp. 11–17 in Infectious Disease and Host- with a note on the cocoons of the related A. assama (Westwood). Pathogen Evolution, edited by K. R. Dronamraju. Cambridge Uni- Genet. Agrar. 45: 175–186. purway ronamraju ayakar versity Press, Cambridge, UK. S , H., K. R. D and S. D. J , 1964 One nest of Sceliphron madraspatanam (Fabr.) (Sphecidae: Hymenoptera). 2Britain’s National Health Service J. Bombay Nat. Hist. Soc. 61: 1–26.