California Institute of Technology
What was the name of the employer Thomas Hunt Morgan work for from 1928 to 1929?
Thomas Hunt Morgan Thomas Hunt Morgan ( September 25 , 1866 – December 4 , 1945 ) was an American evolutionary biologist , geneticist , embryologist , and science author who won the Nobel Prize in Physiology or Medicine in 1933 for discoveries elucidating the role that the chromosome plays in heredity . Morgan received his Ph.D . from Johns Hopkins University in zoology in 1890 and researched embryology during his tenure at Bryn Mawr . Following the rediscovery of Mendelian inheritance in 1900 , Morgan began to study the genetic characteristics of the fruit fly Drosophila melanogaster . In his famous Fly Room at Columbia University , Morgan demonstrated that genes are carried on chromosomes and are the mechanical basis of heredity . These discoveries formed the basis of the modern science of genetics . During his distinguished career , Morgan wrote 22 books and 370 scientific papers . As a result of his work , Drosophila became a major model organism in contemporary genetics . The Division of Biology which he established at the California Institute of Technology has produced seven Nobel Prize winners . Early life and education . Morgan was born in Lexington , Kentucky , to Charlton Hunt Morgan and Ellen Key Howard Morgan . Part of a line of Southern plantation and slave owners on his fathers side , Morgan was a nephew of Confederate General John Hunt Morgan ; his great-grandfather John Wesley Hunt had been the first millionaire west of the Allegheny Mountains . Through his mother , he was the great-grandson of Francis Scott Key , the author of the Star Spangled Banner , and John Eager Howard , governor and senator from Maryland . Following the Civil War , the family fell on hard times with the temporary loss of civil and some property rights for those who aided the Confederacy . His father had difficulty finding work in politics and spent much of his time coordinating veterans reunions . Beginning at age 16 in the Preparatory Department , Morgan attended the State College of Kentucky ( now the University of Kentucky ) . He focused on science ; he particularly enjoyed natural history , and worked with the U.S . Geological Survey in his summers . He graduated as valedictorian in 1886 with a Bachelor of Science degree . Following a summer at the Marine Biology School in Annisquam , Massachusetts , Morgan began graduate studies in zoology at the recently founded Johns Hopkins University . After two years of experimental work with morphologist William Keith Brooks and writing several publications , Morgan was eligible to receive a master of science from the State College of Kentucky in 1888 . The college required two years of study at another institution and an examination by the college faculty . The college offered Morgan a full professorship ; however , he chose to stay at Johns Hopkins and was awarded a relatively large fellowship to help him fund his studies . Under Brooks , Morgan completed his thesis work on the embryology of sea spiders—collected during the summers of 1889 and 1890 at the Marine Biological Laboratory in Woods Hole , Massachusetts—to determine their phylogenetic relationship with other arthropods . He concluded that with respect to embryology , they were more closely related to spiders than crustaceans . Based on the publication of this work , Morgan was awarded his Ph.D . from Johns Hopkins in 1890 , and was also awarded the Bruce Fellowship in Research . He used the fellowship to travel to Jamaica , the Bahamas and to Europe to conduct further research . Nearly every summer from 1890 to 1942 , Morgan returned to the Marine Biological Laboratory to conduct research . He became very involved in governance of the institution , including serving as an MBL trustee from 1897 to 1945 . Career and research . Morgans career and research can be broken into several phases : Bryn Mawr . In 1890 , Morgan was appointed associate professor ( and head of the biology department ) at Johns Hopkins sister school Bryn Mawr College , replacing his colleague Edmund Beecher Wilson . Morgan taught all morphology-related courses , while the other member of the department , Jacques Loeb , taught the physiological courses . Although Loeb stayed for only one year , it was the beginning of their lifelong friendship . Morgan lectured in biology five days a week , giving two lectures a day . He frequently included his own recent research in his lectures . Although an enthusiastic teacher , he was most interested in research in the laboratory . During the first few years at Bryn Mawr , he produced descriptive studies of sea acorns , ascidian worms and frogs . In 1894 Morgan was granted a years absence to conduct research in the laboratories of Stazione Zoologica in Naples , where Wilson had worked two years earlier . There he worked with German biologist Hans Driesch , whose research in the experimental study of development piqued Morgans interest . Among other projects that year , Morgan completed an experimental study of ctenophore embryology . In Naples and through Loeb , he became familiar with the Entwicklungsmechanik ( roughly , developmental mechanics ) school of experimental biology . It was a reaction to the vitalistic Naturphilosophie , which was extremely influential in 19th-century morphology . Morgan changed his work from traditional , largely descriptive morphology to an experimental embryology that sought physical and chemical explanations for organismal development . At the time , there was considerable scientific debate over the question of how an embryo developed . Following Wilhelm Rouxs mosaic theory of development , some believed that hereditary material was divided among embryonic cells , which were predestined to form particular parts of a mature organism . Driesch and others thought that development was due to epigenetic factors , where interactions between the protoplasm and the nucleus of the egg and the environment could affect development . Morgan was in the latter camp ; his work with Driesch demonstrated that blastomeres isolated from sea urchin and ctenophore eggs could develop into complete larvae , contrary to the predictions ( and experimental evidence ) of Rouxs supporters . A related debate involved the role of epigenetic and environmental factors in development ; on this front Morgan showed that sea urchin eggs could be induced to divide without fertilization by adding magnesium chloride . Loeb continued this work and became well known for creating fatherless frogs using the method . When Morgan returned to Bryn Mawr in 1895 , he was promoted to full professor . Morgans main lines of experimental work involved regeneration and larval development ; in each case , his goal was to distinguish internal and external causes to shed light on the Roux-Driesch debate . He wrote his first book , The Development of the Frogs Egg ( 1897 ) . He began a series of studies on different organisms ability to regenerate . He looked at grafting and regeneration in tadpoles , fish and earthworms ; in 1901 he published his research as Regeneration . Beginning in 1900 , Morgan started working on the problem of sex determination , which he had previously dismissed when Nettie Stevens discovered the impact of the Y chromosome on sex . He also continued to study the evolutionary problems that had been the focus of his earliest work . Columbia University . Later in 1904 , E . B . Wilson—still blazing the path for his younger friend—invited Morgan to join him at Columbia University . This move freed him to focus fully on experimental work . When Morgan took the professorship in experimental zoology , he became increasingly focused on the mechanisms of heredity and evolution . He had published Evolution and Adaptation ( 1903 ) ; like many biologists at the time , he saw evidence for biological evolution ( as in the common descent of similar species ) but rejected Darwins proposed mechanism of natural selection acting on small , constantly produced variations . Extensive work in biometry seemed to indicate that continuous natural variation had distinct limits and did not represent heritable changes . Embryological development posed an additional problem in Morgans view , as selection could not act on the early , incomplete stages of highly complex organs such as the eye . The common solution of the Lamarckian mechanism of inheritance of acquired characters , which featured prominently in Darwins theory , was increasingly rejected by biologists . According to Morgans biographer Garland Allen , he was also hindered by his views on taxonomy : he thought that species were entirely artificial creations that distorted the continuously variable range of real forms , while he held a typological view of larger taxa and could see no way that one such group could transform into another . But while Morgan was skeptical of natural selection for many years , his theories of heredity and variation were radically transformed through his conversion to Mendelism . In 1900 three scientists , Carl Correns , Erich von Tschermak and Hugo De Vries , had rediscovered the work of Gregor Mendel , and with it the foundation of genetics . De Vries proposed that new species were created by mutation , bypassing the need for either Lamarckism or Darwinism . As Morgan had dismissed both evolutionary theories , he was seeking to prove De Vries mutation theory with his experimental heredity work . He was initially skeptical of Mendels laws of heredity ( as well as the related chromosomal theory of sex determination ) , which were being considered as a possible basis for natural selection . Following C . W . Woodworth and William E . Castle , around 1908 Morgan started working on the fruit fly Drosophila melanogaster , and encouraging students to do so as well . With Fernandus Payne , he mutated Drosophila through physical , chemical , and radiational means . He began cross-breeding experiments to find heritable mutations , but they had no significant success for two years . Castle had also had difficulty identifying mutations in Drosophila , which were tiny . Finally in 1909 , a series of heritable mutants appeared , some of which displayed Mendelian inheritance patterns ; in 1910 Morgan noticed a white-eyed mutant male among the red-eyed wild types . When white-eyed flies were bred with a red-eyed female , their progeny were all red-eyed . A second generation cross produced white-eyed males—a sex-linked recessive trait , the gene for which Morgan named white . Morgan also discovered a pink-eyed mutant that showed a different pattern of inheritance . In a paper published in Science in 1911 , he concluded that ( 1 ) some traits were sex-linked , ( 2 ) the trait was probably carried on one of the sex chromosomes , and ( 3 ) other genes were probably carried on specific chromosomes as well . Morgan and his students became more successful at finding mutant flies ; they counted the mutant characteristics of thousands of fruit flies and studied their inheritance . As they accumulated multiple mutants , they combined them to study more complex inheritance patterns . The observation of a miniature-wing mutant , which was also on the sex chromosome but sometimes sorted independently to the white-eye mutation , led Morgan to the idea of genetic linkage and to hypothesize the phenomenon of crossing over . He relied on the discovery of Frans Alfons Janssens , a Belgian professor at the University of Leuven , who described the phenomenon in 1909 and had called it chiasmatypie . Morgan proposed that the amount of crossing over between linked genes differs and that crossover frequency might indicate the distance separating genes on the chromosome . The later English geneticist J . B . S . Haldane suggested that the unit of measurement for linkage be called the morgan . Morgans student Alfred Sturtevant developed the first genetic map in 1913 . In 1915 Morgan , Sturtevant , Calvin Bridges and H . J . Muller wrote the seminal book The Mechanism of Mendelian Heredity . Geneticist Curt Stern called the book the fundamental textbook of the new genetics and C . H . Waddington noted that Morgans theory of the chromosome represents a great leap of imagination comparable with Galileo or Newton . In the following years , most biologists came to accept the Mendelian-chromosome theory , which was independently proposed by Walter Sutton and Theodor Boveri in 1902/1903 , and elaborated and expanded by Morgan and his students . Garland Allen characterized the post-1915 period as one of normal science , in which The activities of geneticists were aimed at further elucidation of the details and implications of the Mendelian-chromosome theory developed between 1910 and 1915 . But , the details of the increasingly complex theory , as well as the concept of the gene and its physical nature , were still controversial . Critics such as W . E . Castle pointed to contrary results in other organisms , suggesting that genes interact with each other , while Richard Goldschmidt and others thought there was no compelling reason to view genes as discrete units residing on chromosomes . Because of Morgans dramatic success with Drosophila , many other labs throughout the world took up fruit fly genetics . Columbia became the center of an informal exchange network , through which promising mutant Drosophila strains were transferred from lab to lab ; Drosophila became one of the first , and for some time the most widely used , model organisms . Morgans group remained highly productive , but Morgan largely withdrew from doing fly work and gave his lab members considerable freedom in designing and carrying out their own experiments . He returned to embryology and worked to encourage the spread of genetics research to other organisms and the spread of the mechanistic experimental approach ( Enwicklungsmechanik ) to all biological fields . After 1915 , he also became a strong critic of the growing eugenics movement , which adopted genetic approaches in support of racist views of improving humanity . Morgans fly-room at Columbia became world-famous , and he found it easy to attract funding and visiting academics . In 1927 after 25 years at Columbia , and nearing the age of retirement , he received an offer from George Ellery Hale to establish a school of biology in California . Caltech . Morgan moved to California to head the Division of Biology at the California Institute of Technology in 1928 . In establishing the biology division , Morgan wanted to distinguish his program from those offered by Johns Hopkins and Columbia , with research focused on genetics and evolution ; experimental embryology ; physiology ; biophysics and biochemistry . He was also instrumental in the establishment of the Marine Laboratory at Corona del Mar . He wanted to attract the best people to the Division at Caltech , so he took Bridges , Sturtevant , Jack Shultz and Albert Tyler from Columbia and took on Theodosius Dobzhansky as an international research fellow . More scientists came to work in the Division including George Beadle , Boris Ephrussi , Edward L . Tatum , Linus Pauling , Frits Went , and Sidney W . Fox . In accordance with his reputation , Morgan held numerous prestigious positions in American science organizations . From 1927 to 1931 Morgan served as the President of the National Academy of Sciences ; in 1930 he was the President of the American Association for the Advancement of Science ; and in 1932 he chaired the Sixth International Congress of Genetics in Ithaca , New York . In 1933 Morgan was awarded the Nobel Prize in Physiology or Medicine ; he had been nominated in 1919 and 1930 for the same work . As an acknowledgement of the group nature of his discovery he gave his prize money to Bridges , Sturtevants and his own children . Morgan declined to attend the awards ceremony in 1933 , instead attending in 1934 . The 1933 rediscovery of the giant polytene chromosomes in the salivary gland of Drosophila may have influenced his choice . Until that point , the labs results had been inferred from phenotypic results , the visible polytene chromosome enabled them to confirm their results on a physical basis . Morgans Nobel acceptance speech entitled The Contribution of Genetics to Physiology and Medicine downplayed the contribution genetics could make to medicine beyond genetic counselling . In 1939 he was awarded the Copley Medal by the Royal Society . He received two extensions of his contract at Caltech , but eventually retired in 1942 , becoming professor and chairman emeritus . George Beadle returned to Caltech to replace Morgan as chairman of the department in 1946 . Although he had retired , Morgan kept offices across the road from the Division and continued laboratory work . In his retirement , he returned to the questions of sexual differentiation , regeneration , and embryology . Morgan had throughout his life suffered with a chronic duodenal ulcer . In 1945 , at age 79 , he experienced a severe heart attack and died from a ruptured artery . Morgan and evolution . Morgan was interested in evolution throughout his life . He wrote his thesis on the phylogeny of sea spiders ( pycnogonids ) and wrote four books about evolution . In Evolution and Adaptation ( 1903 ) , he argued the anti-Darwinist position that selection could never produce wholly new species by acting on slight individual differences . He rejected Darwins theory of sexual selection and the Neo-Lamarckian theory of the inheritance of acquired characters . Morgan was not the only scientist attacking natural selection . The period 1875–1925 has been called The eclipse of Darwinism . After discovering many small stable heritable mutations in Drosophila , Morgan gradually changed his mind . The relevance of mutations for evolution is that only characters that are inherited can have an effect in evolution . Since Morgan ( 1915 ) solved the problem of heredity , he was in a unique position to examine critically Darwins theory of natural selection . In A Critique of the Theory of Evolution ( 1916 ) , Morgan discussed questions such as : Does selection play any role in evolution ? How can selection produce anything new ? Is selection no more than the elimination of the unfit ? Is selection a creative force ? After eliminating some misunderstandings and explaining in detail the new science of Mendelian heredity and its chromosomal basis , Morgan concludes , the evidence shows clearly that the characters of wild animals and plants , as well as those of domesticated races , are inherited both in the wild and in domesticated forms according to the Mendels Law . Evolution has taken place by the incorporation into the race of those mutations that are beneficial to the life and reproduction of the organism . Injurious mutations have practically no chance of becoming established . Far from rejecting evolution , as the title of his 1916 book may suggest , Morgan laid the foundation of the science of genetics . He also laid the theoretical foundation for the mechanism of evolution : natural selection . Heredity was a central plank of Darwins theory of natural selection , but Darwin could not provide a working theory of heredity . Darwinism could not progress without a correct theory of genetics . By creating that foundation , Morgan contributed to the neo-Darwinian synthesis , despite his criticism of Darwin at the beginning of his career . Much work on the Evolutionary Synthesis remained to be done . Awards and honors . Morgan left an important legacy in genetics . Some of Morgans students from Columbia and Caltech went on to win their own Nobel Prizes , including George Wells Beadle and Hermann Joseph Muller . Nobel prize winner Eric Kandel has written of Morgan , Much as Darwins insights into the evolution of animal species first gave coherence to nineteenth-century biology as a descriptive science , Morgans findings about genes and their location on chromosomes helped transform biology into an experimental science . - Johns Hopkins awarded Morgan an honorary LL.D . and the University of Kentucky awarded him an honorary Ph.D . - He was elected Member of the National Academy of Sciences in 1909 . - He was elected a Foreign Member of the Royal Society ( ForMemRS ) in 1919 - In 1924 Morgan received the Darwin Medal . - The Thomas Hunt Morgan School of Biological Sciences at the University of Kentucky is named for him . - The Genetics Society of America annually awards the Thomas Hunt Morgan Medal , named in his honor , to one of its members who has made a significant contribution to the science of genetics . - Thomas Hunt Morgans discovery was illustrated on a 1989 stamp issued in Sweden , showing the discoveries of eight Nobel Prize-winning geneticists . - A junior high school in Shoreline , Washington was named in Morgans honor for the latter half of the 20th century . Personal life . On June 4 , 1904 , Morgan married Lillian Vaughan Sampson ( 1870–1952 ) , who had entered graduate school in biology at Bryn Mawr the same year Morgan joined the faculty ; she put aside her scientific work for 16 years of their marriage , when they had four children . Later she contributed significantly to Morgans Drosophila work . One of their four children ( one boy and three girls ) was Isabel Morgan ( 1911–1996 ) ( marr . Mountain ) , who became a virologist at Johns Hopkins , specializing in polio research . Morgan was an atheist .