Copyright Ó 2010 by the Genetics 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 physiology, 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, Oxford 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 Germany. 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 Israel 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 zoology, physics, chemistry, mathematics, geol- already familiar with the terminology of hemoglobins, 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. carboxyhemoglobin! 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 temperatures, son to calculate a set. Young Haldane did so promptly. and higher atmospheric pressures 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 England 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 decompression 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) Naomi Mitchison, 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