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GEORGE WELLS BEADLE October 22, 1903-fune 9, l 989 BY NORMAN H. HOROWITZ GEORGE BEADLE WAS A GIANT in the field of modern genetics. He initiated the great series of advances made between 1941 and 1953 that brought the era of classical ge- netics to a close and launched the molecular age. For this achievement he received many honors, including the Nobel Prize. He was elected to the National Academy of Sciences in 1944 and served on its Council from 1969 to 1972. BeacIle also had a clistinguished career as an academic administrator. When he retired in 1968, he was President of The University of Chicago. Long years in administration, however, dial not dampen his love of experimental genetics, and after his retirement he resumed experimental work on a favorite subjectthe origin of maize. In 1981, he gave up research altogether because of increasing disability from the Alzheimer's disease that eventually ended his life. EDUCATION AND EARLY ElFE Beadle his oldest friends usually called him by his boy- hood nickname, "Beets" was born in Wahoo, Nebraska, to Hattie Albro and Chauncey Elmer Beadle, and he cried in Pomona, California, at age eighty-five. He grew up on his father's forty-acre farm near Wahoo. The farm was a mode] for farms its size and was so designated by the U. S. Depart- 27

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28 BIOGRAPHICAL MEMOIRS ment of Agriculture in 1908. Beets' mother died when he was four years old, and he and his brother ant! sister were raised by a succession of housekeepers. As a boy he worked on the farm, and he retained the skills he learned as a gardener and beekeeper there, and his hand- iness with tools, for the rest of his life. Gardening remained one of his greatest pleasures, and the victory garden he grew around his home at Stanford;during the War produced enough for two families. This garden included beehives, but Beets wouldn't eat the honey, saying he had been stung too many times as a boy. He loved corn, on the other hand, and raised several kinds, including a small Mexican variety that gave his garden the distinction of having the earliest sweet corn at Stanford. After his retirement to Pomona in 1982, he derived much pleasure from growing flowers, a hobby he pursued as long as his health permitted. Beets did well in school and was inspired to go on to col- lege by his high school science teacher, Bess MacDonald (the debt to whom he acknowledged more than once in later years). Despite his father's opinion that a farmer did not need all that education, he entered the University of Nebraska Col- lege of Agriculture in 1922. He graduated in 1926 with a B.S. degree ant! stayed on for another year to work for a master's degree with Franklin D. Keim. His first scientific publication, with Keim, dealt with the ecology of grasses. At some point along the way under Keim's beneficent influence, Beets became interested in fundamen- tal genetics and was persuaded to apply to the graduate school at Cornell University instead of going back to the farm. He entered Cornell in 1927 with a graduate assistant- ship and shortly afterward joined R. A. Emerson's research group on the cytogenetics of maize. Corn genetics was new and exciting for Beets, and Emer- son and his team which inclucled Barbara McClintock and

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GEORGE WELLS BEADLE 29 Marcus Rhoades were inspiring. The result was that in the following five years, Beets published no fewer than fourteen papers dealing with investigations on maize, all begun while he was a graduate student at Cornell. In 1928 he married Marion Hill, a graduate student in botany at Cornell, who assisted him with some of his early corn research. Their son, David, was born in 1931. Beets received his Ph.D. in 1931 and was awarded a Na- tional Research Council Fellowship to do postdoctoral work in T. H. Morgan's Division of Biology at the California Insti- tute of Technology. At Caltech, while finishing the work on maize cytogenetics he had started at Cornell on genes for pollen sterility, sticky chromosomes, failure of cytokinesis, and chromosome behavior in maize-teosinte hybrids (a sub- ject he wouIc! return to in his retirement) Beadle also began doing research on Drosophila. Out of it would come one of the most interesting investigations of his career. DROSOPHILA STUDIES: CROSSING OVER VERMILION AND CINNABAR BeadIe's Drosophila studies at Caltech were concerned with the results of crossing over within various chromosomal rearrangements. The important study of crossing over in attached-X chromosomes he conducted with Sterling Emer- son showed that exchanges occur at random between any two non-sister chromatics. Another, reported jointly with A. H. Sturtevant (in a paper called. "monumental" by E. B. Lewis), was the first systematic investigation of crossing over and dis- junction in chromosomes bearing inversions. In 1934, Boris Ephrussi arrived at Caltech from Paris to study Drosophila genetics with Morgan and Sturtevant. He was just two years oIcler than Beadle and they became close friends. Ephrussi soon communicated to Beadle his own in- terest in the problem of gene action, and the two planned a

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30 BIOGRAPHICAL MEMOIRS collaborative study on Drosophila that would use Ephrussi's skill in the techniques of tissue culture and transplantation. In mid-1935, the two men went to Paris to carry out ex- periments in Ephrussi's laboratory at the Institut de Biologie. Though their attempts to grow imaginal discs in tissue cul- ture failed, they succeeded in devising a method for trans- planting discs from one larva to another that allowed! the discs to continue to develop. Before year's end, they had gone as far as they could with this methodology and had worked out a hypothesis to ac- count for the interaction they observed between the vermi- lion and cinnabar genes in transplanted flies. The results, they showecl, could be explainecl by the following assump- tions: (~) the normal alleles of the two genes control the pro- duction of two specific substances, called the v+- and cn+- substances, both necessary for brown eye-pigment forma- tion; (2) the v+-substance is a precursor of the cn+-substance; and (3) gene mutation blocks formation of the corresponding substance. It was not clear until much later that the two sub- stances are actually precursors of the pigment, and Ephrussi and Beadle frequently referred to them as "hormones." At the time, this small step was a great advance in the science of genetics, for it suggested that development could be broken clown into series of gene-controlled chemical re- actions an idea that cried out for further investigation. It implantec! in Beadle the germ of the one gene-one enzyme idea that he later brought to full flower. But first, the two eye- color substances hac! to be identifiecI, a process that took five years. By that time, Beets was hunting bigger game. Following his return from Paris, Beadle moved to Har- vard University as an assistant professor. There, on a few brief occasions, he met a young woman who wouIc! later be- come my wife and who remembered him fondly afterwards as the only member of the Harvard faculty who spoke to Radcliffe undergraduates at Biology Departmental teas.

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GEORGE WELLS BEADLE BEADLE AND TATUM 31 Beadle left Harvard the following year (1937) for Stan- ford University, where he had accepted an appointment as professor of biology. He was joined by biochemist Edwarc! L. Tatum (1909-1975) as a research associate.' Over the next three years, Tatum contributed his skills to the work of isolating and identifying the two eye-color sub- stances. With others, they established that the two substances were derivatives of tryptophan. By 1940, Tatum had ob- tained a crystalline preparation of the v+-substance, but he and Beadle were beaten to the identification by Butenancit, Weidel, and Becker, who hac! adopted the simple procedure of testing known metabolites of tryptophan for their biolog- ical activity. These researchers found that kynurenine is ac- tive as the v+-substance and that OH-kynurenine is active as the cn+-substance. Much later it was shown that condensation of two molecules of OH-kynurenine forms brown pigment. Despite this setback in the laboratory, the years from 1937 to 1939 were not wasted for Beadle. During this period, he joinec} A. H. Sturtevant in writing a superlative textbook, An Introa~uction to Genetics (1939,5), praised by]. A. Moore as "the complete statement of classical genetics." NEUROSPORA CRASSA AND GENE ACTION As a result of his Drosophila experience it became clear to Beadle that an entirely different method was needed to make headway with the problem of gene action. No other nonautonomous traits were known in Drosophila, and the autonomous onesof which there were many were of such towering complexity from the biochemical standpoint that it was hopeless to attempt to reduce them to their individual chemical steps. Beets enjoyed telling how the solution to this problem came to him while he was listening to Tatum lecture in a ' See p. 356 for Joshua Lederberg's memoir of Tatum.

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32 BIOGRAPHICAL MEMOIRS course on comparative biochemistry. Microbial species, Beets learned, differ in their nutritional requirements even though they share the same basic biochemistry. If these differences were genetic in origin, he thought to himself, it should be possible to induce gene mutations that would produce new nutritional requirements in the test organism. Such an ap- proach, if successful, would allow the researchers to identify genes governing known biochemical compounds immedi- ately, as opposed to the years needed to identify the unknown substances controlled by the usual kinds of genes, including most of those then known. What was needed for such an undertaking was an orga- nism that was genetically workable that could be grown on a chemically defined medium. Beadle knew just the organism. While still a graduate student at Cornell, he had heard about Neurospora crassa, the red bread mold. B. O. Docige had come to the campus from the New York Botanical Garden to give a lecture on Neurospora. Beets remembered clearly that the lecture dealt with the genetics of the organism, including re- sults on first- and second-division segregations of the mating- type and other loci. Even years later Beets was pleased to recall that he and a few other graduate students had been able to explain to the skeptical Dodge that his data could be explained by crossing over or the lack of it between the gene and its centromere. Dodge had played an important role in the history of Neurospora. It was he who discovered that the ascospores could be germinated by heat, thus closing its life cycle and making the organism accessible for genetic study. He also did basic studies on its genetics and was enthusiastic about its possibilities for genetic research. He convinced T. H. Mor- gan, a close friend, to take some cultures with him to Pasa- dena when, in 192S, Morgan went out to found the Division of Biology at Caltech. Dodge, according to Beadle, told

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GEORGE WELLS BEADLE 33 Morgan that Neurospora would be "more important than Drosophila some day," and, in Pasadena, Morgan assigned the cultures to graduate student Car! Linclegren, for his thesis in genetics. Lindegren studier} the relation between first- and second-division segregations anct crossing over. He com- pleted his thesis in 1931, the year Beadle arrived at Caltech. In 1940 the question of the nutritional requirements of Neurospora was still an open one. Previous workers had used nutrient agar as the growth medium, but this would not clo for the experiment Beadle had in mincI. Related fungi, how- ever, were known to have simple requirements, and Tatum soon showed that Neurospora would grow on a synthetic me- dium containing sugar, salts, and a single growth factor- biotinthenceforth referred to as "minimal medium." For- tunately, purified concentrates of biotin tract recently become available, ant] nothing now stood in the way of an experi- mental test of BeacIle's idea. The final step was to clear the Drosophila cultures out of the Stanford lab and convert it into a laboratory for Neuro- spora genetics. The plan was to x-ray one parent of a cross and collect offspring (haploid ascospores isolatecl by hand) onto a medium designed to satisfy the maximum number of possible nutritional requirements (so-called "complete me- lium"~. The resulting cultures wouIc! next be transferred to minimal medium. Growth on complete medium, combined with failure to grow on minimal medium, was to be taken as presumptive evidence of an induced nutritional require- ment. The requirement would be identified, if possible, and the culture would be crossest to wild type to determine its heritability. This scheme, in its time, was breathtakingly claring. Some nongeneticists still suspected that genes governed only trivial biological traits, such as eye color anct bristle pattern, while important characters were determined in the cytoplasm by

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34 BIOGRAPHICAL MEMOIRS an unknown mechanism. Many geneticists believed that gene action was far too complex to be resolved by any simple ex- periment. Indeed, the outcome of Beadle and Tatum's trial run was so uncertain that they agreed at the outset to test 5,000 ascospores before giving up the project and- to avoid early disappointment isolated and stored over a thousand spores before testing any of them. Success came with spore no. 299, which gave rise to a culture that grew on complete but not on minimal medium unless this was supplemented! with pyridoxine. This mutant was followed by others showing requirements for thiamine and p-aminobenzoic acid, respectively. All three require- ments were inherited as single-gene defects in crosses to wild type. These mutants were the subject of the first Neurospora paper by Beadle and Tatum (1941,21. Before long, mutants requiring amino acids, purines, and pyrimidines were also found, and the science of biochemical genetics had been born. Beadle recognized that he and Tatum hac! discovered a new florid of genetics and that more hands would be needed to explore it. Early in the fall of 1941 he came to Caltech to give a seminar on the new discoveries and to recruit a couple of research associates to join the enterprise. Since the first BeadIe-Tatum paper on Neurospora hac! yet to be published, no one in the audience hac! an inkling of what was to come. The seminar was memorable. ~ recorded my recollection of it in an article written in honor of BeadIe's seventieth birthday: "The talk lasted only half an hour, and when it was suddenly over, the room was silent. The silence was a form of tribute. The audience was thinking: Nobody with such a discovery could stop talking about it after just thirty minutes" there must be more. Superimposed on this thought was the realization that something historic had happened. Each one of us, I suspect, was mentally surveying, as best he could, the consequences of the revolution that had just taken place. Finally, when it became clear that

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GEORGE WELLS BEADLE 35 Beadle had actually finished speaking, Prof. Frits Wentwhose father had carried out the first nutritional studies on Neurospora in Java at the turn of the century got to his feet and, with characteristic enthusiasm, ad- dressed the graduate students in the room. This lecture proved, said Went, that biology is not a finished subject- there are still great discoveries to be made!" Neurospora Newsletter 20~1973~:4-6 BEADLE AS LABORATORY HEAD David Bonner and ~ accepted appointments with Beadle and joined his group at Stanford the following year. Later, H. K. Mitchell and Mary Houlahan (Mitchell) came. There were also graduate students (including A. H. Doermann and Adrian Srb) and a steady turnover of visitors in the lab. The next four years were the most exciting of my life, and ~ imagine the same was true for everyone else in the lab. Before the Neurospora revolution, the idea of uniting ge- netics and biochemistry had been only a dream with a few scattered observations. Now, biochemical genetics was a real science, and it was all new. Incredibly, we privileged few had it all to ourselves. Every day brought unexpected new results, new mutants, new phenomena. It was a time when one went to work in the morning wondering what new excitement the clay would bring. Beadle presided over this scientific paradise with the en- thusiasm, intelligence, and good humor that characterized everything he did. He was a popular and much admired boss. He worker! in the lab with everyone else. He especially en- joyed working with his hands, and he had plenty of oppor- tunity to indulge himself in this regard. The laboratories were located in the basement (the "cat- ~ ~ ~ ~ . acombs ) ot Jordan tiall, a location that gave them a certain remoteness from the campus. There were a bench and lathe in the lab, and Beets used these to make small equipment and do minor repairs around the place; he called the campus shops only for major work and did as much as possible him-

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36 BIOGRAPHICAL MEMOIRS self. ~ came to work early one morning and found him paint- ing one of the rooms. All this was in addition to his research and his teaching Buttes as a professor of biology. He always did more than anyone else. ~ recall going to a lab picnic at the beach one summer day, over the coast range of hills from the Stanford campus. We were bicycling to save gas, huffing and wheezing (we had no gears then); Beets differed from the rest of us only in that he was carrying a watermelon on his handlebars. Beets knew his responsibilities and took them seriously. It was wartime, and he concerned himself with all that implied for the pursuit of fundamental research. He had to find fi- nancial support for the program while trying to keep his group together. He succeeded on both scores, obtaining sup- port from both the Rockefeller and Nutrition foundations support that continued throughout the war and even after- wards. The Committee on Medical Research of the Office of Scientific Research and Development classified the Neuro- spora program as essential to the war effort. As ~ recall, no senior researcher or graduate student was drafted, although some of us were called up for physical examinations. Practical applications of Neurospora research were of po- tential utility to the war effort in developing bioassays for vitamins and amino acids in preserved foods, and in search- ing for new vitamins and amino acids. Although the major thrust of the lab remained basic science, we worked on both these applications during the war years. Toward the end of World War Il. Beadle was asked by the War Production Board to devote part of the effort of the lab to seeking mu- tants of Penicillium with increased yields of penicillin. He complied, of course, but we were not successful in this en- deavor. The biochemical and genetic studies carried out between 1941 and 1945 on Neurospora mutants in the Stanford lab- oratory showed that the biosynthesis of any given substance

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GEORGE WELLS BEADLE HONORS AND DISTINCTIONS HONORARY DEGREES Doctor of Science 1947 Yale University 1949 University of Nebraska 1952 Northwestern University 1954 Rutgers University 1955 Kenyon College 1956 Wesleyan University 1959 Birmingham University 1959 Oxford University 1961 Pomona College 1962 Lake Forest College 1963 University of Rochester 1963 University of Illinois lD64 Brown University 1964 Kansas State University 1964 1966 1967 1970 1971 1972 1973 1975 1976 University of Pennsylvania Wabash College Syracuse University Loyola University, Chicago Hanover College Eureka College Butler University Gustavus Adolphus College Indiana State University I-egum Doctor (LL .D.) 1962 University of California, Los Angeles 1963 University of Miami 1963 Brandeis University 1966 Johns Hopkins University 1966 Beloit College 1969 University of Michigan 43

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44 BIOGRAPHICAL MEMOIRS Litterarum Humaniorum Doctor (~.H.D.) 1966 Jewish Theological Seminary of America 1969 DePaul University 1969 University of Chicago 1969 Canisius College 1969 Knox College 1971 Roosevelt University 1971 Carroll College Doctor of Public Service 1970 Ohio Northern University AWARDS 1950 Lasker Award 1951 Dyer Award 1953 Emil Christian Hansen Prize (Denmark) 1958 Albert Einstein Commemorative Award in Science 1958 Nobel Prize in Physiology or Medicine (with E. L. Tatum 1959 1960 1967 1967 and J. Lederberg) National Award, American Cancer Society Kimber Genetics Award Priestley Memorial Award Edison Prize, Best Science Book for Youth (with Muriel Beadle) 1972 Donald Forsha {ones Medal 1984 Thomas Hunt Morgan Medal

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GEORGE WELLS BEADLE PROFESSIONAL AND HONORARY SOCIETIES 45 Genetics Society of America (president, 1945) American Association for the Advancement of Science (president, 1955) National Academy of Sciences (Council, 1969-1972) American Philosophical Society American Academy of Arts and Sciences Royal Society Danish Royal Academy of Sciences Japan Academy Instituto Lombardo di Scienze e Lettre (Milan) Genetical Society of Great Britain Indian Society of Genetics and Plant Breeding Indian Natural Science Academy Chicago Horticultural Society (president, 1968-1971) Phi Beta Kappa Sigma Xi

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46 BIOGRAPHICAL MEMOIRS SELECTED BIBLIOGRAPHY 1927 With F. D. Keim. Relation of time of seeding to root development and winter survival of fall seeded grasses and legumes. Ecology, 8:251-64. 1928 With B. McClintock. A genie disturbance of meiosis in Zea mays. Science, 68:433. 1929 Yellow stripe A factor for chlorophyll deficiency in maize located in the Pr pr chromosome. Am. Nat., 68: 189-192. A gene for supernumerary mitoses during spore development in Zea mays. Science, 70:406-7. 1930 Heritable characters in maize. I. Hered., 21:45-48. Genetical and cytological studies of Mendelian asynapsis in Zea mays. Cornell Univ. Memoir, 129:3-23. A fertile tetraploid hybrid between Euchlaena persons and Zea mays. Am. Nat., 69:190-92. 1931 A gene in Zea mars for failure of cvtokinesis during meiosis. (~lvrol 3: 142-55. A gene in maize for supernumerary cell divisions following meiosis. Cornell Univ. Memoir, 135:3 -12. 1932 A possible influence of the spindle fibre on crossing-over in Dro- sophila. Proc. Natl. Acad. Sci. USA, 18:160-65. A gene in Zea mays for failure of cytokinesis during meiosis. Cytol., 3:142-55. Genes in maize for pollen sterility. Genet., 17:413-31. The relation of crossing over to chromosome association in Zea- Euchlaena hybrids. Genet., 17:481-501. Studies of Euchlaena and its hybrids with Zeal 1. Chromosome be- , c, , ,

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GEORGE WELLS BEADLE 47 havior in Euchlaena mexicana and its hybrids with Zea mays. ZIAVA, 62:291-304. With R. Emerson. Studies of Euchlaena and its hybrids with Zeal II. Crossing over between the chromosomes of Euchlaena and those of Zeal ZIAVA, 62:305 -15. A gene for sticky chromosomes in Zea mays. ZIAVA, 63: 195-217. 1933 Studies of crossing-over in heterozygous translocations in Droso- phila melanogaster. ZIAVA, 65: 111-28. With S. Emerson. Crossing-over near the spindle fiber in attached- X chromosomes of Drosophila melanogaster. ZI AVA, 65: 129-40. Further studies of asynaptic maize. Cytol., 4:269-287. Polymitotic maize and the precocity hypothesis of chromosome conjugation. Cytol., 5: 118-21. 1934 Crossing-over in attached-X triploids of Drosophila melanogaster. ]. Genet., 29:277-309. 1935 Crossing over near the spindle attachment of the X chromosomes in attachecl-X triploids of Drosophila melanogaster. Genet., 20: 179-91. With S. Emerson. Further studies of crossing-over in attached-X chromosomes of Drosophila melanogaster. Genet., 20: 192-206. With R. A. Emerson and A. C. Fraser. A summary of linkage stud- ies in maize. Cornell Univ. Memoir, 180:3-83. With A. H. Sturtevant. X chromosome inversions and meiosis in Drosophila melanogaster. Proc. Natl. Acad. Sci. USA, 21 :384-90. With B. Ephrussi. La transplantation des disques imaginaux chez la Drosophile. C. R. Acad. Sci., 201:98. With B. Ephrussi. Differenciation de la couleur de Ceil cinnabar chez la Drosophile. C. R. Acad. Sci., 201:620. With B. Ephrussi. La transplantation des ovaires chez la Droso- phile. Bull. Biol. Belg., 69:492-502. With B. Ephrussi. Sur les conditions de l'auto-differenciation des caracteres mendeliens. C. R. Acad. Sci., 201:1148. With B. Ephrussi. Transplantation in Drosophila. Proc. Natl. Acad. Sci. USA, 21:642-46.

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~8 BIOGRAPHICAL MEMOIRS 1936 With B. Ephrussi. A technique of transplantation for Drosophila. Am. Nat., 70:218-25. With B. Ephrussi. The differentiation of eye pigments in Droso- phila as studied by transplantation. Genet., 21:225-47. With B. Ephrussi. Development of eye colors in Drosophila: Trans- plantation experiments with suppressor of vermilion. Proc. Natl. Acad. Sci. USA, 22:536-40. With B. Ephrussi and C. W. Clancy. Influence de la lymphe sur la couleur des yeux vermilion chez la Drosophile. C. R. Acad. Sci., 203:545. With A. H. Sturtevant. The relation of inversions in the X chro- mosome of Drosophila melanogaster to crossing-over and disjunc- tion. Genet., 21:554-604. With Th. Dobzhansky. Studies on hybrid sterility IV. Transplanted testes in Drosophila pseudoobscura. Genet., 21:832-40. With B. Ephrussi. Development of eye colors in Drosophila: Stud- ies of the mutant claret. I. Genet., 33:407-10. 1937 With B. Ephrussi. Development of eye colors in Drosophila: Trans- plantation experiments on the interaction of vermilion with other eye colors. Genet., 22:65-75. With B. Ephrussi. Development of eye colors in Drosophila: Dif- fusible substances and their interrelations. Genet., 22:76-85. With B. I:phrussi. Developpement des couleurs des yeux chez la Drosophile: Influence des implants sur la couleur des yeux de l'hote. Bull. Biol. Belg., 71:75-90. With B. Ephrussi. Developpement des couleurs des yeux chez la Drosophile: Revue des experiences de transplantation. Bull. Biol. Belg., 71 :54-74. With C. W. Clancy. Ovary transplants in Drosophila melanogaster: Studies of the characters singed, fused, and female-sterile. B~ol. Bull, 72:47-56. With B. Ephrussi. Development of eye colors in Drosophila: The mutants bright and mahogany. Am. Nat., 71:91-95. The development of eye colors in Drosophila as studied by trans- plantation. Am. Nat., 71:120-26. With K. V. Thimann. Development of eye colors in Drosophila:

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GEORGE WELLS BEADLE 49 Extraction of the diffusible substances concerned. Proc. Natl. Acad. Sci. USA, 23:143-46. Development of eye colors in Drosophila: Fat bodies and Malpigh- ian tubes as sources of diffusible substances. Proc. Natl. Acad. Sci. USA, 23:146-52. With C. W. Clancy and B. Ephrussi. Development of eye colours in Drosophila: Pupal transplants and the influence of body fluid on vermilion. Proc. R. Soc. London, 122:98-105. The inheritance of the color of Malpighian tubes in Drosophila me- lanogaster. Am. Nat., 71:277-79. With B. Ephrussi. Ovary transplants in Drosophila melanogaster. Meiosis and crossing-over in superfemales. Proc. Natl. Acad. Sci. USA, 23:356-60. With B. Ephrussi. Development of eye colors in Drosophila: Pro- duction and release of en+ substance by the eyes of different eye color mutants. Genet., 22:479-83. Chromosome aberration and gene mutation in sticky chromosome plants of Zea ways. Cytol. Fujii Jubilee, pp. 43-56. Development of eye colors in Drosophila: Fat bodies and Malpigh- ian tubes in relation to diffusible substances. Genet., 22:587- 611. 1938 With L. W. Law. Influence on eye color of feeding diffusible sub- stances to Drosophila melanogaster. Proc. Soc. Exp. Biol. Med., 37:621-23. With R. Anderson and J. Maxwell. A comparison of the diffusible substances concerned with eye color development in Drosophila, Ephestia and Habrobracon. Proc. Natl. Acad. Sci. USA, 24:80- 85. With E. L. Tatum. Development of eye colors in Drosophila: Some properties of the hormones concerned. I. Gen. Physiol., 22:239-53. With E. L. Tatum and C. W. Clancy. Food level in relation to rate of development and eye pigmentation in Drosophila melanogas- ter. Biol. Bull., 75:447-62. 1939 Physiological aspects of genetics. Annul Rev. Physiol., 1:41-62. Teosinte and the origin of maize. J. Hered., 30:245-47.

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50 BIOGRAPHICAL MEMOIRS With E. L. Tatum and C. W. Clancy. Development of eye colors in Drosophila: Production of v+ hormone by fat bodies. Biol. Bull., 77:407-14. With E. L. Tatum. Effect of diet on eye-color development in Dro- sophila melanogaster. Biol. Bull., 77:415-22. With A. H. Sturtevant. An Introduction to Genetics. Philadelphia: W. B. Saunders. 1940 With E. L. Tatum. Crystalline Drosophila eye-color hormone. Sci- ence, 91:458. 1941 With E. L. Tatum. Experimental control of development and dif- ferentiation. Am. Nat., 75: 107-16. With E. L. Tatum. Genetic control of biochemical reactions in Neu- rospora. Proc. Natl. Acad. Sci. USA, 27:499-506. 1942 With E. L. Tatum. Genetic control of biochemical reactions in Neu- rospora: An "aminobenzoicless" mutant. Proc. Natl. Acad. Sci. USA, 28:234-43. 1943 With N. H. Horowitz. A microbiological method for the determi- nation of choline by use of a mutant of Neurospora. I. Biol. Chem., 150:325-33. With D. Bonner and E. L. Tatum. The genetic control of biochem- ical reactions in Neurospora: A mutant strain requiring isoleu- cine and valine. Arch. Biochem., 3:71-91. With F. J. Ryan and E. L. Tatum. The tube method of measuring the growth rate of Neurospora. Am. I. Bot., 30:784-99. 1944 With E. L. Tatum and D. Bonner. Anthranilic acid and the biosyn- thesis of indole and tryptophan by Neurospora. Arch. Bio- chem., 3:477-78. With V. L. Coonradt. Heterocaryosis in Neurospora crassa. Genet., 29:291-308.

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GEORGE WELLS BEADLE 51 An ~nositolless mutant strain of Neurospora and its use in bio- assays. I. Biol. Chem., 156:683-89. 1945 With H. K. Mitchell and D. Bonner. Improvements in the cylinder- plate method for penicillin assay. I. Bacterial., 49:101-4. With E. L. Tatum. Biochemical genetics of Neurospora. Ann. Mo. Bot. Gard., 32: 125-29. With N. H. Horowitz, D. Bonner, H. K. Mitchell, and E. L. Tatum. Genic control of biochemical reactions in Neurospora. Am. Nat., 79:304-17. Biochemical genetics. Chem. Rev., 37: 15 -96. Genetics and metabolism in Neurospora. Physiol. Rev. 25:643-63. With E. L. Tatum. Neurospora. II. Methods of producing and de- tecting mutations concerned with nutritional requirements. Am. I. Bot., 32:678-86. The genetic control of biochemical reactions. Harvey Lect., 40: 179-94. Genes and the chemistry of the organism. Am. Sci., 34:31-53, and 76. 1946 With D. Bonner. Mutant strains of Neurospora requiring nicotin- amide or related compounds for growth. Arch. Biochem., 11:319-28. High-frequency radiation and the gene. Science Life in the World, New York: McGraw-Hill, Vol. 2, pp. 163-93. 1959 Genes and chemical reactions in Neurospora. In: Les Prix Nobel, pp. 147-59. Also in: Science, 129: 1715 -19. 1960 Evolution in microorganisms, with special reference to the fungi. ANL, 47:301-19. 1963 Genetics and modern biology. Jayne Lectures for 1962. Am. Philos. Soc., 57:1-73.

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52 BIOGRAPHICAL MEMOIRS 1966 With Muriel B. Beadle. The Language of Life: An Introduction to the Science of Genetics. New York: Doubleday (Doubleday Anchor- book, 1967~. Biochemical genetics: Some recollections. In: Phage and the Origins of Molecular Biology, eds. I. Cairns, G. S. Stent, and ]. D. Watson, Cold Spring Harbor Laboratory of Quantitative Biology, pp. 23-32. 1972 The mystery of maize. Field Mus. Nat. Hist. Bull., 43:2-11. 1973 Thomas Hunt Morgan. In: Dictionary of American Biography, Sup- plement 3 ~ 1941-1945), New York: Chas. Scribners Sons, pp. 538-41. 1974 Recollections. Annul Rev. Biochem. 43:1-13. 1980 The origin of maize. In: Genes, Cells, and Behavior, eds. N. H. Ho- rowitz and E. Hutchings, Jr., San Francisco: W. H. Freeman and Co., pp. 81-87. Ancestry of corn. Sci. Am., 242(1): 112-19. 1981 Origin of corn: Pollen evidence. Science, 213:890-92.

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