Hermann Joseph Muller∗ (21 December 1890–05 April 1967)

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ARTICLE-IN-A-BOX Hermann Joseph Muller∗ (21 December 1890–05 April 1967)

Born in 1890, and brought up in the city of New York, Hermann Muller was a third generation American of mixed German and Portuguese/Spanish/English ancestry. His life went into a crisis very early, as his father, a metal-worker, died when the young Muller was just nine and half years old. At this young age, he had to start earning to support his mother and sister. While most would have succumbed to the pressure and given up on dreams of academic excellence, Muller was an exception. He excelled whenever he was subjected to immense pressure – right from his early years, and throughout his life. After finishing his schooling, and having by then become first a pantheist and then an atheist, Muller joined an undergraduate programme (BA) at Columbia University with a scholarship (1907–1910). There, he was introduced to the rapidly growing field of genetics, first through Robert Heath Lock’s book on recent developments in the study of inheritance, and later by a course on genetics by Edmund Beecher Wilson. After finishing his BA, Muller wanted to get to the active centre of genetics research at that time by joining Thomas Hunt Morgan’s Fly lab at Columbia University. However, due to the lack of a suitable scholarship, he had to take up a graduate fellowship at Columbia Medical School, which allowed him to finish his MA (1911), based on a thesis on the physiology of nerve impulses. Muller spent the following year at Cornell Medical School, on a teaching fellowship. By this time, he was working three shifts a day. He had teaching duties at the Medical School during the day, and he was teaching English to foreign students at night. Rest of his time was divided between course work and research in physiological chemistry. Finally, in 1912, Muller found a way to join Morgan’s Fly lab with a laboratory assistantship. This was the beginning of an intellectual journey that helped shape the subsequent progress of the field of genetics. After finishing his doctoral thesis in 1915, Muller was given a position at the Rice Institute in Texas, USA. He quickly invited his childhood friend Edgar Altenburg to join the Institute. This was the beginning of a lasting collaboration, both scientific and personal, that was perhaps the most influential in Muller’s life. Until 1932, Muller stayed at the Rice Institute, and this tenure proved to be one of the most productive phases in his career. While Muller’s story as a scientist (see the General Article in this issue) is exciting, it becomes even more inspiring when put in the context of the social and political backdrop against which his science was done. Muller

∗DOI: https://doi.org/10.1007/s12045-019-0794-4

RESONANCE | April 2019 413 ARTICLE-IN-A-BOX made his exciting discoveries of the 1920s at a time when his personal life was passing through severe emotional ups and downs. He was married to Jessie Marie Jacobs, a mathematician, in 1923, and their son David, was born the following year. During the time of the discovery of induced mutations, Muller was seriously overworked, though neither for the first nor the last time in his life. This was perhaps because Muller had realized that the work on mutagenesis was probably the most important among all that he had done till then, and that he was on the cusp of bringing about a paradigm shift in experimental genetics. There was just too much to be done. As it turns out, this extreme dedication and frenetic work eventually culminated in one of the most exciting discoveries of the time – production of mutations by means of x-ray irradiation. This was the discovery for which Muller was awarded the Nobel Prize (Medicine and Physiology) in 1946. However, these achievements came at a serious cost. Overwork jeopardized his marriage. Caught between the tussle of marriage and work, and saddened by what he perceived as the ethical degradation of society, Muller was driven to a serious mental breakdown during the latter part of the decade. Perhaps to recover from the bruises of his breakdown, he almost instantly immersed himself in work again, upon recovery. By this time, Muller also began to express his political views more openly, something that carried substantial risks in the USA, given Muller’s Marxist sympathies. During his first trip to Europe in the early 1920s, Muller was deeply perturbed by the social inequalities that he experienced on-board the ship. This experience and perhaps his struggle to support his family as a teenager, was enough to turn him into a deeply committed socialist. Towards the end of the 1920s, he became secretly involved in communist activities. Though it was never officially declared as a reason, he had to resign from his position at Rice Institute in 1936 for his active participation in the publication of the communist magazine – The Spark. In 1932, after his relationship with Jessie ended, Muller moved to Europe. He travelled first to Hitler’s Germany and then eventually to the USSR. Deeply troubled by the social and political oppression in Germany, the prospect of settling in a socialist country like Russia attracted Muller. He spent a few nice years working in Leningrad and Moscow with some of the best Russian geneticists of the time. However, the rise of Trofim Lysenko and pseudoscience, fa- mous as ‘Lysenkoism’, proved too much for both Russian genetics and Muller. During those dark days for genetics in Russia, driven by political motivation, followers of Lysenkoism cap- tured, tortured and even executed a large number of geneticists and Darwinists in the USSR. In 1937, Muller had to escape from Russia to save himself and his friends. During the same time, due to his alleged involvement with The Spark, he was forced to resign from the Rice Institute back home. This meant that Muller could not return to his old job in the USA. Therefore, he moved to Scotland and joined the University of Edinburgh. Though he spent almost three

414 RESONANCE | April 2019 ARTICLE-IN-A-BOX fruitful years at Edinburgh, he had to return to the USA when World War II broke out. At Edin- burgh, he met his second wife – Dorothea Johanna Kantorowicz, who brought the much-needed stability to his personal life. The couple later had a daughter, Helen Juliette. An interesting anecdote about Muller in the Indian context is that he met and worked with Sachi Prasad Ray-Chaudhuri at Edinburgh. His work with Ray-Chaudhuri showing the dose dependence of the mutagenic effect of radiation was considered to be very important by Muller himself. Ray-Chaudhuri later came back to India and established genetics research groups, first at the University of Calcutta in 1956, and later at Banaras Hindu University in 1960. This marked the early establishment of genetics and cytogenetics research, particularly using fruit flies, in India, at Mysore [1] and Banaras around the same time. Many contemporary Indian Drosophila geneticists and cytogeneticists, especially in northern India, trace their academic ancestry to Ray-Chaudhuri. After the onset of World War II, Muller had to return to the USA from Edinburgh. To his utter disappointment, after his arrival at New York, he struggled to secure an academic position for a long time. After almost a year’s struggle, he managed to find a temporary position at Amherst College. He continued to work there until 1944 under bad financial conditions. Finally, in 1945, Indiana University at Bloomington recognized the value of bringing Muller to their zoology department and offered him a professorship. He continued to work at Bloomington for the rest of his life, first as a Professor, then as Distinguished Service Professor, and finally as Emeritus Professor. One year after joining Indiana University, he was awarded the Nobel Prize for his work on ionizing radiation induced mutations done at the Rice Institute. He was a civilian advisor for the Manhattan Project, without realizing what the project was all about during 1943–1944, but not in 1945, the year of the bombing of Hiroshima and Nagasaki. He was also involved in assessing whether radar could be mutagenic. After 1945, Muller continued to warn against the use of nuclear weapons through his extensive writings and lectures. He was one of the proactive scientists who compiled the anti-nuclear weapon statement – the Einstein–Russell manifesto – which was eventually signed by thousands of scientists across the world. This manifesto was the first step of a movement that eventually culminated in a worldwide movement that played a crucial role in regulating nuclear proliferation across the world through the United Nations. Through the 1950s and 1960s, Muller continued working on various aspects of mutations, in- cluding some of his long-held ideas. He wrote extensively on the means of improving human qualities by positive eugenics (as opposed to the more commonly held view of negative eugenics) and voluntary germinal selection. He also took up a fundamentally important role to uplift the general standard of biology education in the USA and was involved in the preparation of

RESONANCE | April 2019 415 ARTICLE-IN-A-BOX three biology textbooks. If the modern field of molecular biology had its roots in genetics, Muller can be said to have planted the seed when a young James Watson in 1947 came to In- diana University and learned about genes from Muller. It was in this period that Muller made some prophetic statements about the future of genetics and the chemical nature of genes [2]. Fortunately, many of his predictions were verified experimentally in his lifetime, and he wit- nessed the early development of molecular genetics with great excitement. He passed away on 05 April 1967, of heart disease, leaving a rich legacy of ideas and having strengthened the then young field of genetics as one of the most important disciplines in basic and applied biology research.