Contents 1 Early life 2 Career 2.1 Discovery of X-ray mutagenesis 2.2 Work in Europe 2.3 Later career 3 Personal life 4 Former graduate students 5 Bibliography 6 See also 7 References 8 External links

Early life[edit] 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.[3] His mother's family was of mixed Jewish (descended from Spanish and Portuguese Jews) and Anglican background, and had come from Britain.[3][4] Among his first cousins are Herbert J. Muller and Alfred Kroeber whose daughter is Ursula LeGuin.[3] As an adolescent, he 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 B.A. degree in 1910.[5] Muller remained at Columbia (the pre-eminent American zoology program at the time, thanks 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).[6] 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.[7]

Career[edit] In 1914, Julian Huxley offered Muller a position at the recently founded William Marsh Rice Institute, now Rice University; he hurried to complete his Ph.D. 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 larmarckiana 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.[8] 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 (who 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.[9] 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,[10] 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 to 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.[11] Discovery of X-ray mutagenesis[edit] 1926 marked the beginning of a series of major breakthroughs. Beginning 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).[12] 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.[13] Work in Europe[edit] 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. But 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 she and their son David E. Muller 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.[14] 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.[15] 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 much 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."[16] Muller—with about 250 strains of Drosophila—moved to Edinburgh in September 1937, after a brief stay 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"[17] 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 there remained little direct physical evidence.[18] Later career[edit] Muller's house in Bloomington, Indiana When Muller returned to the United States in 1940, he took an untenured research position at Amherst College, in the department of Otto C. Glaser. After the U.S. entry into World War II, his position was extended indefinitely and expanded to include teaching. His Drosophila work in this period focused on measuring the rate of spontaneous (as opposed to radiation-induced) mutations. Muller's publication rate decreased greatly in this period, from a combination of lack of lab workers and experimentally challenging projects. However, he also worked as a staff in the Manhattan Project at the University of Chicago in 1942–1943[19] (though he did not know that was what it was), as well as a study of the mutational effects of radar. Muller's appointment was ended after the 1944–1945 academic year, and despite difficulties stemming from his socialist political activities, he found a position as professor of zoology at Indiana University.[20] Here, he lived in a Dutch Colonial Revival house in Bloomington's Vinegar Hill neighborhood.[21] In 1946 Muller was awarded the Nobel Prize in Physiology or Medicine, "for the discovery that mutations can be induced by x-rays". Genetics, and especially the physical and physiological nature of the gene, was becoming a central topic in biology, and X-ray mutagenesis was a key to many recent advances, among them George Beadle and Edward Tatum's work on Neurospora that established in 1941 the one gene-one enzyme hypothesis.[22] In Muller's Nobel Prize lecture he argued that there was no threshold dose of radiation that did not produce mutagenesis, which led to the adoption of the linear no-threshold model of radiation on cancer risks.[23] The Nobel Prize, in the wake of the atomic bombings of Hiroshima and Nagasaki, focused public attention on a subject Muller had been publicizing for two decades: the dangers of radiation. In 1952, nuclear fallout became a public issue; since Operation Crossroads, more and more evidence had been leaking out about radiation sickness and death caused by nuclear testing. Muller—and many other scientists—pursued an array of political activities to defuse the threat of nuclear war. With the Castle Bravo fallout controversy in 1954, the issue became even more urgent.[citation needed] In 1955 Muller was one of eleven prominent intellectuals to sign the Russell-Einstein Manifesto, the upshot of which was the first 1957 Pugwash Conference on Science and World Affairs, which addressed the control of nuclear weapons.[24][25] He was a signatory (with many other scientists) of the 1958 petition to the United Nations, calling for an end to nuclear weapons testing, which was initiated by the Nobel Prize-winning chemist Linus Pauling.[24] It has however been argued that Muller's opinions on the effect of radiation on mutagenesis were not supported by studies on the survivors of the atomic bombings, or in research on mice.[26] Some scientists have criticized the approach of Muller; James F. Crow called Muller's view "alarmist" and wrote that it created in the public "an irrational fear of low-level radiation relative to other risks".[27][28] It is also argued that he was familiar with another study that directly contradicted the linear no-threshold model he supported, leading to a charge in 2011 that Muller misled the public as to the then-current data available and that would have an effect on the formulation of policy that favored his model.[23] Muller was awarded the Linnean Society of London's Darwin-Wallace Medal in 1958 and the Kimber Genetics Award of the U.S. National Academy of Sciences in 1955.[29] He served as president of the American Humanist Association from 1956 to 1958.[30] The American Mathematical Society selected him as its Gibbs Lecturer for 1958.[31] H. J. Muller and science fiction writer Ursula Le Guin were second cousins; his father (Hermann J. Muller Sr.) and her father's mother (Johanna Muller Kroeber) were siblings, the children of Nicholas Müller, who immigrated to the United States in 1848, and at that time dropped the umlaut from his name. Another cousin was Herbert J. Muller, whose grandfather Otto was another son of Nicholas and a sibling of Hermann Sr. and Johanna.[32]

Personal life[edit] Muller is survived by his daughter, Helen J. Muller, now a professor emerita at the University of New Mexico, who has a daughter, Mala Htun. His son, David E. Muller, a professor emeritus of mathematics and computer science at the University of Illinois and at New Mexico State University, died in 2008 in Las Cruces, New Mexico. David's mother was Jessie Jacobs Muller Offermann, Hermann's first wife. Helen's mother was Dorothea Kantorowicz Muller, Hermann's second wife.[3]

Former graduate students[edit] Seymour Abrahamson Raissa L. Berg Elof Axel Carlson Sara Helen Frye H. Bentley Glass C. P. Oliver Irwin I. Oster Abraham P. Schalet Wilson Stone William Edgar Trout III Dale Eugene Wagoner Former post-doctoral fellows George D. Snell Worked in lab as undergraduates Margaret Russell Edmondson Carl Sagan People who worked in his lab in Indiana [1]

Bibliography[edit] Herman Joseph Muller, Modern Concept of Nature (SUNY Press, 1973). ISBN 0-87395-096-8. Herman Joseph Muller, Man's Future Birthright (SUNY Press, 1973). ISBN 0-87395-097-6. H. J. Muller, Out of the Night: A Biologist's View of the Future (Vanguard Press, 1935). H. J. Muller, Studies in Genetics: The Selected Papers of H. J. Muller (Indiana University Press, 1962).

See also[edit] Mutagenesis Bateson–Dobzhansky–Muller model Repository for Germinal Choice Muller's ratchet Muller's morphs History of biology History of genetics History of model organisms

References[edit] ^ Pontecorvo, G. (1968). "Hermann Joseph Muller. 1890-1967". Biographical Memoirs of Fellows of the Royal Society. 14: 348–389. doi:10.1098/rsbm.1968.0015.  ^ Carlson, Elof Axel (1981). Genes, radiation, and society: the life and work of H. J. Muller. Ithaca, N.Y: Cornell University Press. ISBN 0-8014-1304-4.  ^ a b c d Elof Axel Carlson (2009). "Hermann Joseph Muller 1890—1967" (PDF). National Academy of Sciences.  ^ "Hermann J. Muller - Biographical".  ^ Carlson, Genes, Radiation, and Society, pp 17-37 ^ Carlson, Genes, Radiation, and Society, pp 37-69 ^ Carlson, Genes, Radiation, and Society, pp 70-90; for more on the culture and norms of the fly lab, see Kohler, Robert E. (1994). Lords of the fly: Drosophila genetics and the experimental life. Chicago: University of Chicago Press. ISBN 0-226-45063-5. . ^ Carlson, Genes, Radiation, and Society, pp 91-108 ^ Carlson, Genes, Radiation, and Society, pp 109-119 ^ Hamilton, Vivien (2016). "The Secrets of Life: Historian Luis Campos resurrects radium's role in early genetics research". Distillations. 2 (2): 44–45. Retrieved 17 February 2017.  ^ Carlson, Genes, Radiation, and Society, pp 120-140 ^ Carlson, Genes, Radiation, and Society, pp 141-164 ^ Carlson, Genes, Radiation, and Society, pp 165-183 ^ Carlson, Genes, Radiation, and Society, pp 184-203 ^ H. J. Muller, Out of the Night: A Biologist's View of the Future (New York: Vangard, 1935), p. v. ^ Carlson, Genes, Radiation, and Society, pp 204-234; quotation from p 233, correspondence from Muller to Julian Huxley, March 9, 1937 ^ "The 'Geneticists Manifesto'," originally published in Journal of Heredity, 1939, 30:371-73; reprinted in H. J. Muller, Studies in Genetics: The Selected Papers of H. J. Muller (Bloomington: Indiana University Press, 1962), pp. 545-548. ^ Carlson, Genes, Radiation, and Society, pp 235-273 ^ "Our Nobel Laureates". University of Chicago Medical Center. Retrieved 1 December 2017.  ^ Carlson, Genes, Radiation, and Society, pp 274–288 ^ Indiana Historic Sites and Structures Inventory. City of Bloomington Interim Report. Bloomington: City of Bloomington, 2004-04, 90. ^ Carlson, Genes, Radiation, and Society, pp 304–318 ^ a b Calabrese, E. J. (30 June 2011). "Muller's Nobel lecture on dose–response for ionizing radiation:ideology or science?" (PDF). Archives of Toxicology. Springer. 85 (4): 1495–1498. doi:10.1007/s00204-011-0728-8. PMID 21717110. Retrieved 30 December 2011.  ^ a b John Bellamy Foster (2009). The Ecological Revolution: Making Peace with the Planet, Monthly Review Press, New York, pp. 71–72. ^ Carlson, Genes, Radiation, and Society, pp. 336–379. ^ Les leçons inattendues d’Hiroshima par Bertrand Jordan - SPS n° 308, avril 2014 ^ James F. Crow (1987). "Muller, Dobzhansky, and Overdominance". Journal of the History of Biology. 20 (3): 351–380.  ^ "Calabrese says mistake led to adopting the LNT model in toxicology". January 23, 2017.  ^ "Kimber Genetics Award". National Academy of Sciences.  ^ "Past AHA Presidents". American Humanist Association.  ^ Muller, H. J. (1958). "Evolution by mutation". Bull. Amer. Math. Soc. 64 (4): 137–160. doi:10.1090/s0002-9904-1958-10191-3. MR 0095766.  ^ Carlson, Genes, Radiation, and Society, pp 10–11

External links[edit] Nobel Biography Hermann Joseph Muller — Biographical Memoirs of the National Academy of Sciences Hermann Muller and Mutations in Drosophila The Muller manuscripts, 1910–1967 in archives of the Indiana University On the origins of the linear no-threshold (LNT) dogma by means of untruths, artful dodges and blind faith, Edward J. Calabrese, Environmental Research 142 (2015) 432–442. Hermann J. Muller Collection Cold Spring Harbor Laboratory Archives v t e Laureates of the Nobel Prize in Physiology or Medicine 1901–1925 1901 Emil Behring 1902 Ronald Ross 1903 Niels Finsen 1904 Ivan Pavlov 1905 Robert Koch 1906 Camillo Golgi / Santiago Ramón y Cajal 1907 Alphonse Laveran 1908 Élie Metchnikoff / Paul Ehrlich 1909 Emil Kocher 1910 Albrecht Kossel 1911 Allvar Gullstrand 1912 Alexis Carrel 1913 Charles Richet 1914 Róbert Bárány 1915 1916 1917 1918 1919 Jules Bordet 1920 August Krogh 1921 1922 Archibald Hill / Otto Meyerhof 1923 Frederick Banting / John Macleod 1924 Willem Einthoven 1925 1926–1950 1926 Johannes Fibiger 1927 Julius Wagner-Jauregg 1928 Charles Nicolle 1929 Christiaan Eijkman / Frederick Gowland Hopkins 1930 Karl Landsteiner 1931 Otto Warburg 1932 Charles Scott Sherrington / Edgar Adrian 1933 Thomas Morgan 1934 George Whipple / George Minot / William Murphy 1935 Hans Spemann 1936 Henry Dale / Otto Loewi 1937 Albert Szent-Györgyi 1938 Corneille Heymans 1939 Gerhard Domagk 1940 1941 1942 1943 Henrik Dam / Edward Doisy 1944 Joseph Erlanger / Herbert Gasser 1945 Alexander Fleming / Ernst Chain / Howard Florey 1946 Hermann Muller 1947 Carl Cori / Gerty Cori / Bernardo Houssay 1948 Paul Müller 1949 Walter Hess / António Egas Moniz 1950 Edward Kendall / Tadeusz Reichstein / Philip Hench 1951–1975 1951 Max Theiler 1952 Selman Waksman 1953 Hans Krebs / Fritz Lipmann 1954 John Enders / Thomas Weller / Frederick Robbins 1955 Hugo Theorell 1956 André Cournand / Werner Forssmann / Dickinson W. Richards 1957 Daniel Bovet 1958 George Beadle / Edward Tatum / Joshua Lederberg 1959 Severo Ochoa / Arthur Kornberg 1960 Frank Burnet / Peter Medawar 1961 Georg von Békésy 1962 Francis Crick / James Watson / Maurice Wilkins 1963 John Eccles / Alan Hodgkin / Andrew Huxley 1964 Konrad Bloch / Feodor Lynen 1965 François Jacob / André Lwoff / Jacques Monod 1966 Francis Rous / Charles B. Huggins 1967 Ragnar Granit / Haldan Hartline / George Wald 1968 Robert W. Holley / Har Khorana / Marshall Nirenberg 1969 Max Delbrück / Alfred Hershey / Salvador Luria 1970 Bernard Katz / Ulf von Euler / Julius Axelrod 1971 Earl Sutherland Jr. 1972 Gerald Edelman / Rodney Porter 1973 Karl von Frisch / Konrad Lorenz / Nikolaas Tinbergen 1974 Albert Claude / Christian de Duve / George Palade 1975 David Baltimore / Renato Dulbecco / Howard Temin 1976–2000 1976 Baruch Blumberg / Daniel Gajdusek 1977 Roger Guillemin / Andrew Schally / Rosalyn Yalow 1978 Werner Arber / Daniel Nathans / Hamilton O. Smith 1979 Allan Cormack / Godfrey Hounsfield 1980 Baruj Benacerraf / Jean Dausset / George Snell 1981 Roger Sperry / David H. Hubel / Torsten Wiesel 1982 Sune Bergström / Bengt I. Samuelsson / John Vane 1983 Barbara McClintock 1984 Niels Jerne / Georges Köhler / César Milstein 1985 Michael Brown / Joseph L. Goldstein 1986 Stanley Cohen / Rita Levi-Montalcini 1987 Susumu Tonegawa 1988 James W. Black / Gertrude B. Elion / George H. Hitchings 1989 J. Michael Bishop / Harold E. Varmus 1990 Joseph Murray / E. Donnall Thomas 1991 Erwin Neher / Bert Sakmann 1992 Edmond Fischer / Edwin G. Krebs 1993 Richard J. Roberts / Phillip Sharp 1994 Alfred G. Gilman / Martin Rodbell 1995 Edward B. Lewis / Christiane Nüsslein-Volhard / Eric F. Wieschaus 1996 Peter C. Doherty / Rolf M. Zinkernagel 1997 Stanley B. Prusiner 1998 Robert F. Furchgott / Louis Ignarro / Ferid Murad 1999 Günter Blobel 2000 Arvid Carlsson / Paul Greengard / Eric Kandel 2001–present 2001 Leland H. Hartwell / Tim Hunt / Paul Nurse 2002 Sydney Brenner / H. Robert Horvitz / John E. Sulston 2003 Paul Lauterbur / Peter Mansfield 2004 Richard Axel / Linda B. Buck 2005 Barry Marshall / Robin Warren 2006 Andrew Fire / Craig Mello 2007 Mario Capecchi / Martin Evans / Oliver Smithies 2008 Harald zur Hausen / Luc Montagnier / Françoise Barré-Sinoussi 2009 Elizabeth Blackburn / Carol W. Greider / Jack W. Szostak 2010 Robert G. Edwards 2011 Bruce Beutler / Jules A. Hoffmann / Ralph M. Steinman (posthumously) 2012 John B. Gurdon / Shinya Yamanaka 2013 James Rothman / Randy Schekman / Thomas C. Südhof 2014 John O'Keefe / May-Britt Moser / Edvard Moser 2015 William C. Campbell / Satoshi Ōmura / Tu Youyou 2016 Yoshinori Ohsumi 2017 Jeffrey C. Hall, Michael Rosbash, Michael W. 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Hermann Müller (disambiguation)New York City, New YorkIndianapolis, IndianaColumbia UniversityRadiationNewcomb Cleveland PrizeNobel Prize In Physiology Or MedicineLinnean Society Of LondonDarwin–Wallace MedalFellow Of The Royal SocietyAmerican Humanist AssociationGeneticsMolecular BiologyDoctoral AdvisorThomas Hunt MorganH. Bentley GlassJ. T. PattersonGeneticistNobel LaureateRadiationMutagenesisRadioactive FalloutNuclear WarNuclear TestingNew York CityKoblenzHerbert J. MullerAlfred KroeberUrsula LeGuinUnitarianismPantheistAtheistColumbia UniversityMendelian InheritanceChromosomeMutationNatural SelectionEvolutionEugenicsBachelor Of ArtsEdmund Beecher WilsonDrosophilaThomas Hunt MorganAlfred SturtevantCalvin BridgesCornell UniversityDrosophilistJulian HuxleyRice UniversityDoctor Of PhilosophyOenothera LarmarckianaHugo De VriesMutationismChromosomal CrossoverMutation RateWorld War IGenetics (journal)Chromosomal InversionUniversity Of Texas At AustinUniversity Of Texas At AustinJessie Marie Jacobs Muller OffermannRadiumX-raysI.Q.BerlinModel OrganismWaspMaizeGreat DepressionCapitalismUSSRBerlinNikolay Timofeeff-RessovskyNiels BohrMax DelbrückNazi PartyDavid E. MullerLeningradMoscowJoseph StalinLysenkoismTrofim LysenkoLamarckismEdinburghMadridParisRichard GoldschmidtEnlargeBloomington, IndianaAmherst CollegeOtto C. 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