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EDWARD LAWRIE TATUM December 14, 1909-November 7, 1975 BY JOSHUA LEDERB ERG IN THE HISTORY OF BIOLOGY Edward Lawrie Tatum s name is linked with that of George Wells Beadle for their pioneering studies of biochemical mutations in Neurospora.~ First published in 1941, these studies have endures! as the prototype of the investigation of gene action to the present clay. A still more enduring legacy is their development of experimental techniques for the mutation analysis of bio- chemical pathways used claily by modern biologists. Though this sketch is written as a biography of Edward Tatum, these singular scientific accomplishments were in practice and attribution- intimately shared with BeacIle. Ta- tum brought to the work a background in microbiology and a passion for the concept of comparative biochemistry; Beadle, great sophistication in "classical genetics" and the leadership ant! drive to replace the underbrush of vitalistic thinking with a clear-cut, mechanistic view of the gene and the processes of life. Little more than the bare outlines of Edwarct Tatum's per- sonal history can be clocumentecl, because of his own aversion to accumulating paper and the fact that most of his corre- ~ George W. Beadle died on tune 9, 1989, when this essay was in press. His mem- oir, by Norman H. Horowitz, is also included in this volume. 357

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358 BIOGRAPHICAL MEMOIRS sponclence was discarded cluring his various moves. His scientific achievements, however, were largely ant! appro- priately recognized. In 1952 he was elected to the National Academy of Sciences and in 1958, with George Beadle and Joshua Leclerberg, won the Nobel Prize in Physiology or Medicine. Tatum was also known for his commitment to nur- turing younger scientists, with whom he zestfully enjoyed every aspect of laboratory work. A still more enduring legacy of their work has been the everyday use of experimental mu- tation analysis of biochemical pathways in modern biology since then. EDUCATION AND EARLY LIFE Edwarc! Lawrie Tatum was born in BouIcler, Colorado, on December 14, 1909, the first surviving son of Arthur L. (~84-1955) and Mabel Webb Tatum. A twin, Elwood, died shortly after birth. At the time of Ec~ward's birth his father was an instructor in chemistry at the University of Coloraclo at Boulder, where Mabel Webb's father had been Superinten- dent of Schools. Arthur's own father, Lawrie Tatum, a Quaker who hac! settled in the Iowa Territory, hac! been an Indian agent after the Civil War and written a book, Our Red Brothers. In rapic} succession the Tatum family mover! to Madison, Wisconsin; Chicago, Illinois; Philadelphia, Pennsylvania; Vermillion, South Dakota; and, back in 1918 to Chicago. During this period the elcler Tatum helc! a succession of teaching positions while earning a Ph.D. in physiology and pharmacology from The University of Chicago and an M.D. from Rush Meclical College. By 1925 he was settled at the University of Wisconsin at Madison as professor of pharma- cology in a department that was a major center for the train- ing of professors of pharmacology. Among his research ac- complishments were the introduction of picrotoxin as an

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EDWARD LAWRIE TATUM 359 antidote for barbiturate poisoning and the validation of ar- senoxide (mapharsen) for the chemotherapy of syphilis,2 the most effective drug for this purpose until the introduction of penicillin. Edward, having the double advantage of this remarkable family background and the Laboratory School at The Uni- versity of Chicago, continued his education at Wisconsin, earning a bachelor's degree in 1931. At Wisconsin he came upon the tradition of research in agricultural microbiology and chemistry that was then flourishing under the leadership of E. B. Fret! (later president of the University) and W. H. Peterson.3 Tatum's first research was a bachelor's thesis (published 1932) on the effect of associated growth of bacterial species Lactobacillus and Clostrt~ium septicum giving rise to racemic lactic acid. (In 1936 he clemonstratec! that the C. septicum racemized the d-lactic acid! produced by the lactic acid bac- teria.) He continuer! his graduate work at Wisconsin with fi- nancial support from the Wisconsin Alumni Research Foun- dation the beneficiary of royalties from Steenbock's patents on vitamin D milk. His Ph.D. dissertation (1935) concerned the stimulation of C. septicum by a factor isolated from potato, identified as a derivative of aspartic acid and later shown to be asparagine. This was follower! by collaborations with H. G. Wood and Esmond E. Snell in a series of pioneering studies 2.}ohn Patrick Swann, "Arthur Tatum, Parke-Davis, and the Discovery of Ma- pharsen as an Antisyphilitic Agent," Journal of the History of Medicine and Allied Sci- ences, 40(1985):167-87. F. E. Shideman, "A. L. Tatum, Practical Pharmacologist," Science, 123(1956) :449. Anonymous, "Profile of a Research Scientist," Bulletin of Med- ical Research, National Society for Medical Research, 8(1954):7-8. 3 The roots of their work can be traced to Koch, Tollens, and Kossel in Germany. See I. L. Baldwin, "Edwin Broun Fred, March 22, 1887-}anuary 16,1981," Biograph- ical Memoirs of the National Academy of Sciences, Vol. 55, pp. 247-290; and Conrad A. Elvehjem, "Edwin Bret Hart, 1874-1953," Biographical Memoirs, Vol. 28, pp. 117- 161. See also E. H. Beardsley, Harry L. Russell and Agricultural Science in Wisconsin (Madison, Wisconsin: University of Wisconsin Press, 1969).

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360 BIOGRAPHICAL MEMOIRS on the role of vitamins in bacterial nutrition. In 1936 they studier! the growth factor requirements of propionic acic! bacteria, fractionating one factor from an acetone extract of milk powder. Its ohvsical oronerties suggested that the factor might he thiamine, and indeed crystalline thiamine was fully 1 J 1 1- - __,=_. active as an essential growth factor. Vitamins hac! long been recognized to share a role in the nutrition of animals, man, and yeast. Tatum's work with Snell, Peterson, ant! Wood initiates! a genre of studies showing that many bacterial species tract diverse requirements for these identical substances. This was outstanding confirmation of the basic tenet of comparative biochemistry- the evolution- ary conservation of biochemical processes that produced common processes in morphologically diversified species. Tatum's education ant! ctoctoral research coincided with the culmination of understanding that all of the basic building blocks of lifeamino acids, sugars, lipids, growth factors (anc! later nucleic acids) existed in fundamentally similar chemical structures among all forms of life. Hence the most fruitful way to stucly a problem in animal metabolism might be to begin with a microbe, which might well prove more convenient for experimental manipulation and bioassay and as the future would show genetic analysis anc! alter- ation. Tatum then won a General Education Boarcl postdoctoral fellowship that took him, his wife (the former June Alton, a fellow student at Wisconsin), and their infant daughter, Margaret, to Fritz Kog1's laboratory at Utrecht, The Nether- lancis, for a year. Kog] hadjust purified and crystallized biotin as a growth factor for yeast, and this enabled and inspirer! further studies on its nutritional role for other microorga- nisms. (Not until 1940 was the nutritional significance of bio- tin for animals recognizecl.) By Tatum's own account, his brief time at Utrecht, spent in efforts to isolate further growth factors for staphylococci,

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EDWARD LAWRIE TATUM 361 never achieved a sharp research focus. More importantly, he befriended Nils Fries, another research fellow from Uppsala, Sweden, who was using the newly available biotin to define the specific nutritional requirements of an ever wicler range of fungi. Fries anc! Kog! were able to demonstrate striking examples of nutritional symbiosisthe compensation for complementary deficits in mixer! cultures of various fungi. Tatum's report to the General Education Board records his gratification at having been able to meet, as well, A. I. Kluyver at Delft, ant! B. C. I. G. Knight and P. Fildes in En- gland then aIreacly well known as leading investigators of bacterial chemistry and nutrition from a comparative per- spective. ~ I. H. Mueller at Harvarc! anc! A. Ewoff in Paris hac! also stresses} how microbial nutrition reflectec! evolutionary losses of biochemical synthetic competencea concept that can be tracer! to Twort and Ingram in 94though they hac} not as yet acloptec! the language or conceptual frame- work of genetics that would eventually describe such varia- tions as gene mutations affecting biosynthetic enzymes.) THE STANFORD YEARS ~ ~ 937~ 945) That same year, 1937, Beadle was on the point of moving from Harvard to Stanford. His research program In pnys~- ological genetics was to continue the work on the genetics of Drosophila eye pigments that he tract initiated in colIabora- tion with Boris Ephrussi, first at Caltech, then in Paris. The Rockefeller Foundation's support of this enterprise was one of Warren Weaver's most foresighted initiatives in the gesta- tion of molecular biology.5 I.ooking out for a possible position for Tatum, his profes- 4 F. W. Twort and G. L. Y. Ingram, "A Method for Isolating and Cultivating the Mycobacterium enteritidis chronicae pseudotuberculosae fohne," and "Some Experiments on the Preparation of a Diagnostic Vaccine for Pseudo-tuberculous Enteritis of Bo- vines," Proceedings, Royal Society, London, Series B. 84(191 1-12):517-42. 5 See also Mina Rees, "Warren Weaver, July 17, 1894-November 24, 1978," Bio- graphical Memoirs, Vol. 57, pp. 493-530.

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362 BIOGRAPHICAL MEMOIRS sors at Wisconsin forwarcled BeadIe's solicitation for a re- search associate "biochemist to work on hormone-like sub- stances that are concerned with eye pigments in Drosophila." But, practical-minclect, they recommencled that the young man undertake research on the chemical microbiology of butter, writing him that "this field is certainly getting hot." With jobs scarce, economic realities weighed as heavily as intellectual appeal in the choice between insect eyes and dairy microbiology. Arthur Tatum, Edward's father, was much con- cerned that, if his son undertook a hybrid role, he wouIc! Tic! himself an academic orphan, clisowned by each of the disci- plines of biochemistry, microbiology, and genetics. In the event, however, Tatum accepted] BeacIle's offered position, and the multiple challenges of comparative biochemistry that went with it. Though the economic importance of butter re- search was far more obvious at the time, it is certain that Edward! Tatum conic! not have chosen better than Drosophila as a means for contributing to the field of biotechnology. Joining Beadle at Stanford, Tatum was engaged between 1937 and 1941 with the arduous task of extracting pigment- precursors from Drosophila larvae. Ephrussi and BeadIe's earlier transplantation experiments had clemonstratec! that a diffusible substance or hormone produced by witcI-type flies was critically lacking in the mutant strain. Yet Tatum and BeacIle's own experience differed! significantly from the re- port published by Ephrussi and Chevais. According to this report, normal eye color could be restored in cultures sup- plementec! with tryptophane. Tatum, however, could confirm this only with cultures carrying a bacterial contaminant. Far from discarcling such a contaminant as an interfering vari- able, Tatum cultured the organism (a Bacillus species) to prove that it was a source of the elusive hormone. The inter- changeability of growth factors for bacteria and animals and the knowledge that many microbes synthesized vitamins re- quirecl by other species undoubtedly bolstered this theory.

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EDWARD LAWRIE TATUM 363 A. I. Haagen-Smit, whom Beadle hack known at Harvard, was now at the California Institute of Technology, and Tatum visited him to learn microchemical techniques, then set out to isolate the "V + hormone" from the bacterial culture. He succeeded in doing this in 1941, only to be anticipated by Butenancit et at. in the identification of V+ as kynurenine. (Butenanclt, astutely noting from a Japanese publication that kynurenine was a metabolite of tryptophane in clog urine, hac! tested the substance for eye color hormone activ- ity.) The jarring experience of having their painstaking work overtaken in so facile a way impellecl Beadle and Tatum to seek another organism more tractable than Drosophila for biochemical studies of gene action. Neurospora and the One Gene~ne Enzyme Theory In winter quarter 1941, Tatum (although a research associate without teaching responsibilities) volunteered to de- velop and teach a then unprecedented comparative biochem- istry course for both biology and chemistry graduate stu- clents. In the course of his lectures he described the nutrition of yeasts and fungi, some of which exhibited well-defined blocks in vitamin biosynthesis. Attending these lectures, BeacIle recalled B. O. Dodge's elegant work on the segrega- tion of morphological mutant factors in Neurospora that he hac! heart! in a seminar at Cornell in 1932,6 work that was followed up by C. C. Lindegren at Caltech. Neurospora, with its immediate manifestation of segre- gating genes in the string of ascospores, has an ideal life-cycle for genetic analysis. Fries's work suggested that Neurospora might also be cultured reaclily on a well definer! medium. It was soon established that Neurospora required only biotin as 6 See also W. J. Robbins, "Bernard Ogilvie Dodge, April 18, 1872-August 9, 1960," Biographical Memoirs, Vol. 36, pp. 85-124.

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364 BIOGRAPHICAL MEMOIRS a supplement to an inorganic salt-sucrose medium and did indeed prove an ideal organism in which to seek mutations with biochemical effects demonstrated by nutritional require- ments. By February 1941,7 the team was X-raying Neuro- spora and seeking these mutants. ~ . . . . . ~ .arvestlng nutrltlona mutants in microorganisms in those days was painstaking hand labor; it meant examining single-spore cultures isolated from irradiated parents for their nutritional propertiesone by one. No one could have predicted how many thousands of cultures would have to be tested to discover one that would have a biochemical defect marked by a nutritional deficiency. Isolate #299 proved to be the first recognizable mutant, requiring as it did pyridoxine. The trait, furthermore, seg- regated in crosses according to simple Mendelian principles, which foretold that it could in due course be mapped onto a specific chromosome of the fungus. Therewith, Neurospora moved to center stage as an object of genetic experimenta- tion. By May of the same year, Beadle and Tatum were ready to submit their first report of their revolutionary methods to the Proceedings of the National Academy of Sciences. In that report they noted "there must exist orders of di- rectness of gene control ranging from one-to-one relations to relations of great complexity." The characteristics of mu- tations affecting metabolic steps suggested a direct and simple role for genes in the control of enzymes. The authors 7 G. W. Beadle, "Recollections," Annual Revue of Biochemistry, 43 (1974):1-13. In his chapter, "Biochemical Genetics, Some Recollections," in Phage and the Origins of Molecular Biology, eds. J. Cairns, G. S. Stent, and J. D. Watson (Cold Spring Harbor, New York: C. S. H. Biol. Labs, 1966), Beadle confused the 1940-41 meeting of the Society of American Naturalists in Philadelphia, which made no reference to Neu- rospora, with that of the Genetics Society in Dallas in December 1941. The net effect is to date the Neurospora experiments to 1940 rather than to 1941. H. F. Hudson repeated the error in The Eighth Day of Creation (New York: Simon & Schuster, 1979), and it is bound to plague future historians.

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EDWARD LAWRIE TATUM 365 hypothesized, therefore, that enzymes were primary prod- ucts of genes. Indeed, in some cases, genes themselves might be enzymes. This was what came to be labelled the one gene- one enzyme theory, the precursor of today's genetic dogma. We shall return to it later. In that same year Tatum was recruited as an assistant pro- fessor to the regular faculty of Stanford's Biology Depart- ment, where he developed an increasingly independent re- search program exploiting the use of Neurospora mutants for the exploration of biochemical pathways. Despite the ex- igencies of the war effort, an increasing number of talented graduate students and postdoctoral fellows flocked to Stan- ford to learn the new discipline. Their participation rapidly engendered a library of mutants blocked in almost any ana- bolite that could be replaced in the external nutrients. Today, that catalog embraces over 500 distinct genetic loci and well over a thousand publications from laboratories the world over.8 Anticipating the One Gene-One Enzyme Theory Would that contemporaries could anticipate what future historians will ask or what errors they will promulgate! How many simple questions we neglect to ask, or fad! to record the answers, that might have settled continuing controversies. Among these is the place of Archibald E. Garrod's work and thought in anticipation of the one gene-one enzyme hypoth- esis. The following discussion is offered in some detail in order to correct some prevalent misconstructions of that his- tory. In 190S, Garrod published his study of what was then called "inborn errors of metabolism," including alcaptonuria ~ D. D. Perkins, A. Radford, D. Newmeyer, and M. Bjorkman, "Chromosomal loci of Neurospora crassa," Microbiological Reviews, 46 (1982):426-570.

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366 BIOGRAPHICAL MEMOIRS in many This work is sometimes portrayer! as a forgotten precursor of Beadle and Tatum's investigation of gene action. Indeed, many geneticists who specialized in maize or Dro- sophila, including Beadle himself, lamented not knowing of this pioneering work earlierit having received remarkably little comment from geneticists until after Neurospora was launched in ~ 94 ~ . ~ Yet Garrocl's basic findings on alcaptonuria, which parallel the metabolic blocks in Neurospora mutants, were widely quoted in medical texts. I. B. S. Haldane cited them in a well- reac! essay in 1937. Tatum likewise referred to them in his course in comparative biochemistry before beginning his own experiments on Neurospora. BeacIle, in his Nobel Prize lec- ture in INS, was careful to acknowledge these antececlents, though widely quotes! reminiscences have blurred the cletaits of just when Beadle and Tatum became aware of Garrocl's work. Halclane, in his 1937 article, cited the (1ifficulty of exper- imentation on rare human anomalies as an important reason to seek other research paradigms- which Neurospora wouIc! eventually provide. But Garrod himself never quite made 9 "The Croonian Lectures of the Royal College of Physicians," Lancet 2(1908): 1- 7, 73-79, 142-148, 214-220. A H. Harris, ea., Garrod's Inborn Errors of Metabolism (Oxford: Oxford University Press, 1963); and B. Childs and C. R. Scriver, eds., Inborn Factors in Disease by A. E. Garrod (Oxford: Oxford University Press, 1989), include extensive discussion and bibliography on the history of his ideas. On the neglect of Garrod's work, see also R. Olby, The Path to the Double Helix (London, Macmillan Press, 1974). ~ ~ Though G. W. Beadle implies in PATOOMB (Phage and the Origins of Molecular Biology, see footnote 7 above), that he and Tatum were unaware of Garrod until perhaps 1945, they referred to Garrod in a paper on their Drosophila-pigment work delivered January 1, 1941 (see American Naturalist, 75[1941]: 107-16). Garrod's find- ings were also prominent in Tatum's winter 1941 course on comparative biochem- istry at Stanford. I first read about Garrod in Meyer Bodansky's Introduction to Phys- iological Chemistry (New York: Wiley & Sons, 1934), and the late Sewall Wright advised me that he had taught that material in Chicago since 1925. ]2~. B. S. Haldane, "The Biochemistry of the Individual," in Perspectives in Bio- chemistry, J. Needham and D. E. Green, eds. (Cambridge: Cambridge University

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EDWARD LAWRIE TATUM 377 complex multi-chain ensembles ant! can contain nonprotein cofactors requiring the participation of many genes. Uncler- standing the role of RNA as a message intermediary between DNA and protein, the complexities of intervening sequences in RNA, RNA-processing, and post-translational processing came later and required more sophisticated biochemical anal- ysisbut all clerivec! from the concepts and the tools of the Neurospora studies. Beadle and Tatum's contribution, then, comprised the fol- lowing: 1) A methodology for the investigation of gene-enzyme relationships that exploited experimentally-acquired genetic mutations affecting specific biosynthetic steps. 2) A conceptual framework the one gene-one enzyme theory from which to search for and characterize these mutants. This framework was derived from the model that chromosomal genes contain (substantially) all of the blueprints for development and that enzymes (and other proteins) are the mediators of gene action. 3) The dethronement of Drosophila as the prime experimental ma- terial for physiological genetic research in favor of the fungus Neurospora. This further helped open the way to use of bacteria and viruses in genetic research and the culture of tissue cells as if they were microbes. These methods and concepts have been the central paradigm for experimental biology since 1941. Beadle ant! Tatum sharer! many awards in adclition to the 1958 Nobel Prize in recognition of these innovations. In 1952, Tatum was indiviclually honoree! by election to the Na- tional Academy of Sciences. In 1953 he received the Remsen Aware! of the American Chemical Society and was elected to the American Philosophical Society. He was president of the Harvey Society (1964-65) and the recipient of at least seven honorary clegrees. He served on the NAS Carty Fund Committee from 1956 to 1961. For the NRC, he took part in a number of panels

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378 BIOGRAPHICAL MEMOIRS and committees having to do with genetics and biology and was a member of the Advisory Committee on the Biological Effects of Ionizing Radiations from 1970 to 1973. He also dice yeoman service on advisory committees for the National Institutes of Health, American Cancer Society, the National Foundation (March of Dimes), ant! other booties concerned with the award of fellowships and grants. He was chairman of the Scientists' Institute for Public Information and an advisor to the City of Hope Meclical Center, Rutgers University Institute of Microbiology, and SIoan-Kettering In- stitute for Cancer Research, and a consultant in microbiology for Merck and Co. He worked actively on many scientific publications, inclucling Annual Reviews, Science, Biochemica et Biophysica Acta, Genetics, and the Journal of Biological Chemistry. Testifying to a Congressional committee on behalf of the National Science Foundation in 1959, Tatum said: "The general philosophy [of the NSF] is concentration on excellence . . . making it possible for [the scientist] to use his capacities, both for research and for training the next generation . . . whether it is a particular research program in a given area, whether it may or may not be immediately prac- ticable in its application . . . freedom to develop the intellectual curiosity and abilities of the individual...." At this time Beadle and Tatum's legacy is embodied in published work that has influencer! biological research through several scientific generations. The original papers are "classics" ant! taken for granted. Personal recollections of Tatum are facling, and this re- port can hardly clo justice to his humor, his hobbies (includ- ing the French horn), his zest for experiments, his love of microbes, his attachment to students, friends, ant! family the trauma of divorce notwithstanding the tragedy of his final year of bereavement and of an illness that left him gasp- ing for breath. He touched the lives of many young scientists.

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EDWARD LAWRIE TATUM 379 The enduring appreciation of his role in their clevelopment is the memorial he would have cherished most. THE TANTALIZINGLY FEW personal papers of Edward Tatum now extant are on deposit at the Rockefeller University Archive Center. I am particularly indebted to Professor Carlton Schwerdt for hav- ing preserved and made available his lecture notes on Tatum's 1941 course on comparative biochemistry, to June Alton Tatum for making available to me materials regarding Tatum's life before 1946, and to the staff of the Rockefeller University Archive Center. I am also indebted to the following important studies for infor- mation that appears in this account: R. M. Burian, lean Gayon, and Doris Zallen, "The Singular Fate of Genetics in the History of French Biology," Journal of the History of Biology, 21~19881:357-402, on the Beadle-Ephrussi collaboration that led directly to Beadle and Tatum's work on Drosophila eye color "hormones" and dis- cusses the use of that terminology for what would later be termed "precursors." Lily E. Kay, "Selling Pure Science in Wartime: The Biochemical Genetics of G. W. Beadle," journal of the History of Biology, 22~1989~:73-101, reviews the Beadle-Tatum work on pen- icillin improvement during World War II.

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380 BIOGRAPHICAL MEMOIRS SELECTED BIBLIOGRAPHYl6 1932 With W. H. Peterson and E. B. Fred. Effect of associated growth on forms of lactic acid produced by certain bacteria. Biochem. J., 26:846-52. 1934 Studies in the biochemistry of microorganisms. Ph.D. Dissertation, University of Wisconsin, Madison. 1936 With H. G. Wood and W. H. Peterson. Essential growth factors for propionic acid bacteria. II. Nature of the Neuberg precipitate fraction of potato: Replacement by ammonium sulphate or by certain amino acids. I. Bacteriol., 32: 167-74. With H. G. Wood and W. H. Peterson. Growth factors for bacteria. V. Vitamin Be, a growth stimulant for propionic acid bacteria. Biochem. }, 30: 1 898-1 904. 1937 With E. E. Snell and W. H. Peterson. Growth factors for bacteria. III. Some nutritive requirements of Lactobacillus delbruckii. ]. Bacteriol., 33:207-25. With W. H. Peterson and E. B. Fred. Enzymatic racemization of optically active lactic acid. Biochem. l., 30:1892-97. 1938 With G. W. Beadle. Development of eye colors in Drosophila: Some properties of the hormones concerned. l. Gen. Physiol., 22:239-53. 1939 Development of eye colors in Drosophila: Bacterial synthesis of v+ hormone. Proc. Natl. Acad. Sci. USA, 25:486-90. Nutritional requirements of Drosophila melanogaster. Proc. Natl. Acad. Sci. USA, 25:490-97. }6 A complete bibliography can be found in the Archives of the National Academy of Sciences and in the Rockefeller University Archive Center.

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EDWARD LAWRIE TATUM 1940 381 With G. W. Beadle. Crystalline Drosophila eye color hormone. Sci- ence, 91:458. 1941 With G. W. Beadle. Experimental control of development and dif- ferentiation. Am. Nat., 75: 107-16. Vitamin B requirements of Drosophila melanogaster. Proc. Natl. Acad. Sci. USA, 27:193-97. With A. I. Haagen-Smit. Identification of Drosophila v+ hormone of bacterial origin. I Biol. Chem., 140: 575 -80. With G. W. Beadle. Genetic control of biochemical reactions in Neurospora. Proc. Natl. Acad. Sci. USA, 27:499-506. 1942 With G. W. Beadle. Genetic control of biochemical reactions in Neurospora: An "aminobenzoicless" mutant. Proc. Natl. Acad. Sci. USA, 28: 234-43. 1943 With L. Garnjobst and C. V. Taylor. Further studies on the nutri- tional requirements of Colpoda duodenar?a. ]. Cell. Comp. Phys- iol., 21: 199-212. With F. I. Ryan and G. W. Beadle. The tube method of measuring the growth rate of Neurospora. Am. J. Bot., 30:784-99. With D. Bonner and G. W. Beadle. The genetic control of biochem- ical reactions in Neurospora: A mutant strain requiring isoleu- cine and valine. Arch. Biochem., 3:71-91. With D. M. Bonner. Synthesis of tryptophan from indole and serine by Neurospora. I. Biol. Chem., 151 :349. 1944 With D. Bonner. Indole and serine in the biosynthesis and break- down of tryptophan. Proc. Natl. Acad. Sci. USA, 30:30-37. Biochemistry of fungi. Annul Rev. Biochem., 13:667-704. With C. H. Gray. X-ray induced growth factor requirements in bacteria. Proc. Natl. Acad. Sci. USA, 30:404-10.

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382 BIOGRAPHICAL MEMOIRS 1945 With N. H. Horowitz, D. Bonner, H. K. Mitchell, and G. W. Beadle. Genic control of biochemical reactions in Neurospora. Ann. Nat., 79:304-17. With G. W. Beadle. Biochemical genetics of Neurospora. Ann. Mo. Bot. Garden, 32:125-29. X-ray induced mutant strains of E. coli. Proc. Natl. Acad. Sci. USA, 31 :215-19. With G. W. Beadle. Neurospora II. Methods of producing and de- tecting mutations concerned with nutritional requirements. Am. I. Bot., 32:678-86. 1946 With T. T. Bell. Neurospora III. Biosynthesis of thiamin. Am. J. Bot., 33:15-20. With I. Lederberg. Novel genotypes in mixed cultures of biochem- ical mutants of bacteria. Cold Spring Harbor Symp. Quant. Biol., 11:113-14. Induced biochemical mutations in bacteria. Cold Spring Harbor Symp. Quant. Biol., 11:278-84. 1947 Chemically induced mutations and their bearing on carcinogene- sis. Ann. N.Y. Acad. Sci., 49:87-97. With I. Lederberg. Gene recombination in the bacterium Esche- r~chia coli. ]. Bacteriol., 53:673-84. 1950 With R. W. Barratt, N. Fries, and D. Bonner. Biochemical mutant strains of Neurospora produced by physical and chemical treat- ment. Am. I. Bot., 37:38-46. With R. C. Ottke and S. Simmonds. Deuteroacetate in the biosyn- thesis of ergosterol by Neurospora. l. Biol. Chem., 186:581-89. With D. D. Perkins. Genetics of microorganisms. Annul Rev. Mi- crobiol., 4:129-50. With E. A. Adelberg. Characterization of a valine analog accumu- lated by a mutant strain of Neurospora crassa. Arch. Biochem., 29:235-36.

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EDWARD LAWRIE TATUM 383 1951 With E. A. Adelberg and D. M. Bonner. A precursor of isoleucine obtained from a mutant strain of Neurospora crassa. ]. Biol. Chew., 190:837-41. With E. A. Adelberg. Origin of the carbon skeletons of isoleucine and valine. I. Biol. Chem., 190:843-52. 1954 With S. R. Gross, G. Ehrensvard, and L. Garnjobst. Synthesis of aromatic compounds by Neurospora. Proc. Natl. Acad. Sci. USA, 40:271-76. With D. Shemin. Mechanism of tryptophan synthesis in Neuro- spora. I. Biol. Chem., 209:671-675. 1956 With S. R. Gross and R. D. Gaylord. The metabolism of proto- catechuic acid in Neurospora. I. Biol. Chem., 219: 781-96. With S. R. Gross. Physiological aspects of genetics. Ann. Rev. Phys- iology, 18:53-68. With R. A. Eversole. Chemical alteration of crossing-over fre- quency in Chlamydomonas. Proc. Nat. Acad. Sci. USA, 42:68- 73. With L. Garnjobst. A temperature independent riboflavin requir- ing mutant of Neurospora crassa. Am. I. Bot., 43: 149-57. With R. C. Fuller. Inositol-phospholipid in Neurospora and its re- lationship to morphology. Am. J. Bot., 43:361-65. 1958 With R. W. Barratt. Carcinogenic mutagens. Ann. N.Y. Acad. Sci., 71: 1072-84. Molecular basis of the cause and expression of somatic cell varia- tion. J. Cell Comp. Physiol., 52:313-36. 1959 A case history in biological research. Science, 129:1711-15. Also in: Les prix Nobel en 1958, Stockholm, pp. 160-9. With A. J. Shatkin. Electron microscopy of Neurospora crassa my- celia. I. Biophys. Biochem. Cytol., 6:423-26.

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384 BIOGRAPHICAL MEMOIRS 1961 With James F. Wilson and Laura Garnjobst. Heterocaryon incom- patibility in Neurospora crassa Micro-injection studies. Am. i. Bot., 48:299-305. With Noel de Terra. Colonial growth of Neurospora. Science, 134: 1066-68. With E. Reich, R. M. Franklin, and A. I. Shatkin. Effect of actino- mycin D on cellular nucleic acid synthesis and virus production. Science, 134:556-57. 1962 Biochemical genetics and evolution. Comp. Biochem. Physiol., 4:241-48. With A. {. Shatkin, E. Reich, and R. M. Franklin. Effect of mito- mycin C on mammalian cells in culture. Biochem. Biophys. Acta, 55:277-89. With E. Reich, R. M. Franklin, and A. J. Shatkin. Action of acti- nomycin D on animal cells and viruses. Proc. Nat. Acad. Sci. USA, 48:1238-45. 1963 With Noel de Terra. A relationship between cell wall structure and colonial growth in Neurospora crassa. Am. l. Bot., 50:669-77. With B. Mach and E. Reich. Separation of the biosynthesis of the antibiotic polypeptide tyrocidine from protein biosynthesis. Proc. Nat. Acad. Sci. USA, 50:175-81. 1965 Perspectives from physiological genetics. In: The Control of Human Heredity and Evolution, ed. E. Sonneborn, New York: Macmillan, pp. 20-34. With E. G. Diacumakos and L. Garnjobst. A cytoplasmic character in Neurospora crassa. The role of nuclei and mitochondria. I. Cell Biol., 26:427-43. With C. W. Slayman. Potassium transport in Neurospora. III. Iso- lation of a transport mutant. Biochem. Biophys. Acta, 109: 184- 93.

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EDWARD LAWRIE TATUM 1966 385 With Z. K. Borowska. Biosynthesis of edeine by Bacillus brevis Vm4: In viva and in vitro. Biochem. Biophys. Acta, 114:206-9. The possibility of manipulating genetic change. In: Genetics and the Future of Man, First Nobel Conference, Gustavus Adolphus Col- lege.. Ed., J. D. Roslansky, New York: Appleton-Century-Crofts, pp. 51-61. With B. Mach. The biosynthesis of antibiotic polypeptides. In: Ninth International Congress for Microbiology, Moscow, London: Pergamon Press, pp. 57-63. With S. Brody. The primary biochemical effect of a morpholog- ical mutation in Neurospora crassa. Proc. Nat. Acad. Sci. USA, 56: 1290-7. Molecular biology, nucleic acids, and the future of medicine. Per- spec. Biol. Med., 10: 19-32. 1967 With B. Crocken. Sorbose transport in Neurospora crassa. Biochem. Biophys. Acta, 135: 100-5. With E. Pina. Inositol biosynthesis in Neurospora crassa. Biochem. Biophys. Acta, 136:265-71. With S. Brody. Phosphoglucomutase mutants and morphological changes in Neurospora crassa. Proc. Nat. Acad. Sci. USA, 68:923-30. With L. Garnjobst. A survey of new morphological mutants in Neu- rospora crassa. Genet., 57:579-604. With M. P. Morgan and L. Garnjobst. Linkage relations of new morphological mutants in linkage group V of Neurospora crassa. Genet., 57:605-12. With P. R. Mahadevan. Localization of structural polymers in the cell wall of Neurospora crassa. ]. Cell Biol., 35:295-302. 1970 With N. C. Mishra. Phosphoglucomutase mutants of Neurospora sitophila and their relation to morphology. Proc. Nat. Acad. Sci. USA, 66:638-45. With L. Garnjobst. New crisp genes and crisp modifiers in Neuro- spora crassa. Genetics, 66:281-90.

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386 BIOGRAPHICAL MEMOIRS 1971 With W. A. Scott. Purification and partial characterization of glucose-6-phosphate dehydrogenase from Neurospora crassa. ]. Biol. Chem., 246:6347-52. 1972 With E. G. Diacumakos. Fusion of mammalian somatic cells by mi- crosurgery. Proc. Nat. Acad. Sci. USA, 69:2959-62. 1973 With N. C. Mishra. Non-Mendelian inheritance of DNA-induced inositol independence in Neurospora. Proc. Nat. Acad. Sci. USA, 70:3875-79. 1974 With C. R. Wrathall. Hyphal wall peptides and colonial morphol- ogy in Neurospora crassa. Biochem. Genet., 12:59-68.

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