Hermann Joseph Muller (December 21, 1890 – April 5, 1967) was an American geneticist who was awarded the 1946 Nobel Prize in Physiology or Medicine "for the discovery of the production of mutations by means of X-ray irradiation". Muller warned of long-term dangers of radioactive fallout from nuclear war and nuclear testing, which resulted in greater public scrutiny of these practices.

Early life Muller was born in New York City, the son of Frances (Lyons) and Hermann Joseph Muller Sr., an artisan who worked with metals. Muller was a third-generation American whose father's ancestors were originally Catholic and came to the United States from Koblenz.. His mother's family was of mixed Jewish (descended from Spanish and Portuguese Jews) and Anglican background, and had come from Britain. Among his first cousins was Alfred Kroeber (father of Ursula Le Guin) and first cousins once removed was Herbert J. Muller. As an adolescent, Muller attended a Unitarian church and considered himself a pantheist; in high school, he became an atheist. He excelled in the public schools. At 16, he entered Columbia College. From his first semester, he was interested in biology; he became an early convert of the Mendelian-chromosome theory of heredity—and the concept of genetic mutations and natural selection as the basis for evolution. He formed a biology club and also became a proponent of eugenics; the connections between biology and society would be his perennial concern. Muller earned a Bachelor of Arts degree in 1910.

Muller remained at Columbia (the pre-eminent American zoology program at the time, due to E. B. Wilson and his students) for graduate school. He became interested in the Drosophila genetics work of Thomas Hunt Morgan's fly lab after undergraduate bottle washers Alfred Sturtevant and Calvin Bridges joined his biology club. In 1911–1912, he studied metabolism at Cornell University, but remained involved with Columbia. He followed the drosophilists as the first genetic maps emerged from Morgan's experiments, and joined Morgan's group in 1912 (after two years of informal participation).

In the fly group, Muller's contributions were primarily theoretical—explanations for experimental results and ideas and predictions for new experiments. In the emerging collaborative culture of the drosophilists, however, credit was assigned based on results rather than ideas; Muller felt cheated when he was left out of major publications.. For more on the culture and norms of the fly lab, see

Career In 1914, Julian Huxley offered Muller a position at the recently founded William Marsh Rice Institute, now Rice University; he hurried to complete his Doctor of philosophy degree and moved to Houston for the beginning of the 1915–1916 academic year (his degree was issued in 1916). At Rice, Muller taught biology and continued Drosophila lab work. In 1918, he proposed an explanation for the dramatic discontinuous alterations in Oenothera lamarckiana that were the basis of Hugo de Vries's theory of mutationism: "balanced lethals" allowed the accumulation of recessive mutations, and rare crossing over events resulted in the sudden expression of these hidden traits. In other words, de Vries's experiments were explainable by the Mendelian-chromosome theory. Muller's work was increasingly focused on mutation rate and lethal mutations. In 1918, Morgan, short-handed because many of his students and assistants were drafted for the U.S. entry into World War I, convinced Muller to return to Columbia to teach and to expand his experimental program.

At Columbia, Muller and his collaborator and longtime friend Edgar Altenburg continued the investigation of lethal mutations. The primary method for detecting such mutations was to measure the sex ratios of the offspring of female flies. They predicted the ratio would vary from 1:1 due to recessive mutations on the X chromosome, which would be expressed only in males (which lacked the functional allele on a second X chromosome). Muller found a strong temperature dependence in mutation rate, leading him to believe that spontaneous mutation was the dominant mode (and to initially discount the role of external factors such as ionizing radiation or chemical agents). In 1920, Muller and Altenburg coauthored a seminal paper in Genetics on "modifier genes" that determine the size of mutant Drosophila wings. In 1919, Muller made the important discovery of a mutant (later found to be a chromosomal inversion) that appeared to suppress crossing over, which opened up new avenues in mutation-rate studies. However, his appointment at Columbia was not continued; he accepted an offer from the University of Texas and left Columbia after the summer of 1920.

Muller taught at the University of Texas from 1920 until 1932. Soon after returning to Texas, he married mathematics professor Jessie Marie Jacobs, whom he had courted previously. In his early years at Texas, Muller's Drosophila work was slow going; the data from his mutation rate studies were difficult to interpret. In 1923, he began using radium and X-rays, but the relationship between radiation and mutation was difficult to measure because such radiation also sterilized the flies. In this period, he also became involved with eugenics and human genetics. He carried out a study of twins separated at birth that seemed to indicate a strong hereditary component of I.Q. Muller was critical of the new directions of the eugenics movement (such as anti-immigration), but was hopeful about the prospects for positive eugenics. In 1932, at the Third International Eugenics Congress, Muller gave a speech and stated, "eugenics might yet perfect the human race, but only in a society consciously organized for the common good".

Discovery of X-ray mutagenesis In 1926, a series of major breakthroughs began. In November, Muller carried out two experiments with varied doses of X-rays, the second of which used the crossing over suppressor stock ("ClB") he had found in 1919. A clear, quantitative connection between radiation and lethal mutations quickly emerged. Muller's discovery created a media sensation after he delivered a paper entitled "The Problem of Genetic Modification" at the Fifth International Congress of Genetics in Berlin; it would make him one of the better-known public intellectuals of the early 20th century. By 1928, others had replicated his dramatic results, expanding them to other model organisms, such as wasps and maize. In the following years, he began publicizing the likely dangers of radiation exposure in humans (such as physicians who frequently operate X-ray equipment or shoe sellers who radiated their customers' feet).

His lab grew quickly, but it shrank again following the onset of the Great Depression. Especially after the stock market crash, Muller was increasingly pessimistic about the prospects of capitalism. Some of his visiting lab members were from the USSR, and he helped edit and distribute an illegal leftist student newspaper, The Spark. It was a difficult period for Muller both scientifically and personally; his marriage was falling apart, and he was increasingly dissatisfied with his life in Texas. Meanwhile, the waning of the eugenics movement, ironically hastened by his own work pointing to the previously ignored connections between environment and genetics, meant that his ideas on the future of human evolution had reduced impact in the public sphere. Muller's speech before the Third International Eugenics Conference in New York has been credited with marking the end of Galtonism, and perhaps even eugenics itself, as a popular movement in the sciences. H. Bentley Glass, a contemporary observer and Ph.D. student of Muller's, would say Muller's speech "just about finished the activity of the Eugenics Society"., Glass, Bentley. (Discussion) The American Journal of Human Genetics, Volume 6: pp.187-188. (1954). Muller told the assembled that eugenic ideals could no longer be achieved, because the capitalistic system produces the wrong motives of individual action, and he disdained the natures of the dominant class, and the type of society they were creating.Hardin, Garrett. Nature and Man's Fate, pp.228-229, Rinehart & Company, Inc., New York, Toronto

Work in Europe In September 1932, Muller moved to Berlin to work with the Russian expatriate geneticist Nikolay Timofeeff-Ressovsky; a trip intended as a limited sabbatical stretched into an eight-year, five-country journey. In Berlin, he met two physicists who would later be significant to the biology community: Niels Bohr and Max Delbrück. The Nazi movement was precipitating the rapid emigration of scientific talent from Germany, and Muller was particularly opposed to the politics of National Socialism. The FBI was investigating Muller because of his involvement with The Spark, so he chose instead to go to the Soviet Union (an environment better suited to his political beliefs). In 1933, Muller and his wife reconciled, and their son David E. Muller and she moved with Hermann to Leningrad. There, at the Institute of Genetics, he imported the basic equipment for a Drosophila lab—including the flies—and set up shop. The institute was moved to Moscow in 1934, and Muller and his wife were divorced in 1935.

In the USSR, Muller supervised a large and productive lab, and organized work on medical genetics. Most of his work involved further explorations of genetics and radiation. There he completed his eugenics book, Out of the Night, the main ideas of which dated to 1910.H. J. Muller, Out of the Night: A Biologist's View of the Future (New York: Vangard, 1935), p. v. By 1936, however, Joseph Stalin's repressive policies and the rise of Lysenkoism was making the USSR an increasingly problematic place to live and work. Muller and many of the Russian genetics community did what they could to oppose Trofim Lysenko and his Larmarckian evolutionary theory, but Muller was soon forced to leave the Soviet Union after Stalin read a translation of his eugenics book and was "displeased by it, and...ordered an attack prepared against it."; quotation from p 233, correspondence from Muller to Julian Huxley, March 9, 1937 By this time, Muller had already asked for a leave of absence. News of the Lysenko trials had reached the United States, and his son David was being raised there, after his divorce. In the official declaration by the institute, biological determinism was rejected: "The development of society is subject not to biological laws but to higher social laws. Attempts to spread to humanity the laws of the animal kingdom are an attempt to lower the human being to the level of beasts."Hardin, Garrett. Nature and Man's Fate, pp.217, Rinehart & Company, Inc., New York, Toronto

Muller, with about 250 strains of Drosophila, moved to University of Edinburgh in September 1937, after brief stays in Madrid and Paris. In 1938, with war on the horizon, he began looking for a permanent position back in the United States. He also began courting Dorothea "Thea" Kantorowicz, a German refugee; they were married in May 1939. The Seventh International Congress on Genetics was held in Edinburgh later that year; Muller wrote a "Geneticists' Manifesto""The 'Geneticists Manifesto'," originally published in Journal of Heredity, 1939, 30:371–373; reprinted in H. J. Muller, Studies in Genetics: The Selected Papers of H. J. Muller (Bloomington: Indiana University Press, 1962), pp. 545–548. in response to the question: "How could the world's population be improved most effectively genetically?" He also engaged in a debate with the perennial genetics gadfly Richard Goldschmidt over the existence of the gene, for which little direct physical evidence existed at the time.

Later career