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PAUL RUFUS BURKHOLDER February 1, 1903-August 11, 1972 BY JAMES G. HORSFALL DISTINGUISHED SCIENTISTS don t just happen. Paul Rufus Burkholder wasn't just accidentally elected (1949) to the National Academy of Sciences. He didn't have his greatness thrust upon him; he earned it. In him was a concatenation of factors that characterize many members of the Academy: (a) He was born into and raised in an intellectual family. His father had a "library" that young Burkholder devoured. (b) He was a bright boy. (c) He was a prodigious worker as witness the extensive bibliography at- tached. (d) He was an introvert at heart, a loner if you please. This allowed him the time to work. (e) He was stimulated by his early teachers to pursue science. (f) He had a devoted wife who shared his hobbies, his life, and his lab. She worked as hard in biology as he did and raised three sons besides. I knew Burkholder in the graduate school at Cornell. I knew him again when I was an adjunct professor in his depart- ment at Yale. I always admired "Burkie," even though he was difficult to "know" well. When he was in graduate school, he compensated for some of his introvert tendencies by joining Gamma Alpha, a gradu- ate student fraternity. There he found another "Burkie" (no relative), whom I also knew. This may not have helped Paul too much, for the second Burkie was a very tall man and Paul 3

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4 BIOGRAPHICAL MEMOIRS was a little short. Promptly the two were distinguished as "big Burkie" and "little Burkie." It must be discouraging to be called a "little Burkie." On the other hand, it drives the little Burkies to excell the big Burkies. At any rate, Paul fought the little-man complex so hard that he was able to put on giant shoes and take giant strides . in science. HIS INTELLECTUAL BASE The Burkholder forebears, religious refugees from the Ger- man section of Switzerland, came to Pennsylvania during its early colonial period. William Rankin Burkholder, Paul's father, was born in 1857, just before the Civil War, on a farm in central Pennsylvania. As a young man, Paul's father and an older brother operated a successful general store in Middle- burg. William was small, quick, and very strong, taking delight in wrestling, hunting, and driving fast horses. To play cards and drink hard liquor was not to be scorned either. Money was plentiful and life was full, strenuous, and satisfying. In some way, William's older brother became convinced that he should forsake this way of life and become a minister in the church. He brought influence to bear on William to do likewise. This was a difficult problem. Could he give up all those things that he enjoyed? The diary he kept at the time bears witness to his struggle. His brotherpositive, eloquent, and persuasive showed him his duty. Once the decision was made, he went off to college,~as very few did a century ago. In time he was ordained as a minister in the United Brethren Church by Bishop Wright, the father of the famous Wright brothers. As his son was to do later, he pursued a solitary, dedicated life of service to the small communities in south- central Pennsylvania. There, when he was thirty-nine years old, he met and married Mary Ellen Schubert, a young girl of , . . .

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PAUL RUFUS BURKHOLDER 5 twenty-one. Their son, Paul, an only child, was born seven years later, on February 1, 1903, at Orrstown, Pennsylvania. Mary's father, George Schubert, came from Germany as a young boy. Because the children made fun of his accent, he left school to become a farmer, married a girl of French descent, and raised a family of seven children. Mary, the oldest, like her son later, had an alert mind and a desire to become a teacher. Her mother pressured her to stay safely at home, however, to help with the family and the aged grandparents. In spite of her bitter disappointment, her sense of responsi- bility won out and she did not rebel. Thus, her formal school- ing came to an end and she turned to church work for an outlet. Here, William found her. Typical of ministers' families, they had very little money, but a house was always provided, along with space for a garden and a farm to keep a cow and a few chickens. Here a small boy could grow in an atmosphere of freedom and with no real sense of hardship. The environ- ment, if somewhat spartan, was intellectual and enriched by visitors to the home and church. Paul read and reread his father's library, consisting mostly of books on religious subjects. Sundays he spent in church, where three times a day he listened to his father's sermons or found the time well spent in medi- tation. He understood what his father was trying to do and admired him for his tireless, if often frustrating, efforts to help the people in their individual, everyday problems and to keep the churches growing and active. Paul knew that the basic princi- ples were right, but there must be more than this. Questions were left unanswered, and Paul went out to find the answers. School and learning were important. Rightly or wrongly, he felt that he was not so bright and must work extra hard to compensate. He took full advantage of the knowledge exhib- ited by the schoolmasters of the little village schools. Summers,

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6 BIOGRAPHICAL MEMOIRS while still very young, he started working for farmers, weeding and picking vegetables and fruit and helping to harvest hay and grain. He came to be known as swift and dependable, so there was no problem in procuring work. Always he saved the money he earned. His spare time was spent roaming in the mountains with his gun, shooting small game for food, and . enjoying nature. Ready for high school, he walked three miles to a train that took him to Chambersburg and three miles home every day. This schedule kept him busy from early until late, but he enjoyed the work and did well in his classes. COLLEGE DAYS In 1920 he enrolled in Dickinson College in Carlisle, Penn- sylvania. His classes ranged over a great variety of subjects as available in a small liberal arts college. 1 ~ ~ ~ T He worked in the Bleary tor expense money. He appreciated this easy access to the books and he used them to good advantage. Like many boys, he worked summers as a laborer. During his four years at Dickinson, he decided to become a botanist. This decision was influenced no doubt by his early and continuing interest in plants. In 1949 Dickinson College recognized his work with an honorary D.Sc. GRADUATE SCHOOL At Cornell, he came under the enthusiastic tutelage of O. F. Curtis and Louis Knudson and almost automatically became a plant physiologist. Of his contemporaries who took plant physiology with him at Cornell, at least four, including himself, were later elected to the National Academy of Sciences. Cornell was surely the happy hunting ground for biologists. There was the science in the laboratory, but the campus and surrounding hills were laced with gorges through which streams

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PAUL RUFUS BURKHOLDER cascaded down to the lake from "far above Cayuga's waters" to quote the old Cornell song. What vegetation to study! It was different on the sunny north side from the shady south side of the gorge. Paul's Gamma Alpha house was on the south edge of Cascadilla gorge. He could collect specimens for his classes almost at his doorstep, or he could go across campus to the Fall Creek gorge to collect plants on the north edge or swim in Beebe Lake and be eyed by the Tri Delta girls from their house on the rim above. And besides he helped the brothers make wine during prohibition days. He was surely a chip off his father's block. Like all graduate students in the twenties, before National Science Foundation student fellowships, he needed money. A job offer introduced him to the excitement of aquatic biology. The New York State Conservation Department offered him a summer job in the limnological survey of the state water- sheds, beginning with the Cayuga Lake basin, with which he was already familiar. He did the phytoplankton studies. He and his colleagues worked hard, had fun, earned money, and pub- lished their papers. This work was to influence him throughout the rest of his life. THE DEPRESSION He received his Ph.D. in 1929, the year of Black Friday and the collapse of the stock market, but it did not discourage him. He was able to continue with the limnological survey. It was extended to Lake Erie in cooperation with the Buffalo Museum of Natural Sciences, where he became a curator of biology and continued plankton studies on the lake. The government boat, Sheerwater, manned by a discrete crew and carefree young scientists, came to be known to the "rum runners" on Lake Erie as a friend in time of need. There were adventures aplenty on the waters and opportunities for firsthand observa-

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8 BIOGRAPHICAL MEMOIRS tion of an epic in our history. Buffalo society added to the fun and frolic. While at the Museum, he planned and installed exhibits and participated in the teaching program. Continuing his plank- ton studies, he spent one summer working in northern New York waters and another working in some lakes of the upper Hudson watershed. The last summer he did a study of the phytoplankton of Frenchman's and Penobscot bays, in Maine, near Bar Harbor. In 1930 he married Lillian Miller, a lady who shared his biological enthusiasms for the rest of his life and who survives him. With the acquisition of a wife and a little later a son, Franz, life took on a more serious aspect. He wished to return to a more academic environment. In the meantime the depres- sion had deepened and no jobs were available. A National Re- search Council Fellowship saved him, however, and he spent two rewarding years, one at Harvard, where a second son, Peter, was born in 1935, and one at Columbia, where he came under the influence of E. W. Sinnott, a member of the Academy. While at Columbia, he took a course in bacteriology. This became another influence on the rest of his career. TO CONNECTICUT COLLEGE Even after two fellowship years, jobs were still scarce, but one appeared in the nick of time. A young Ph.D., George Avery, had recently begun teaching at the Connecticut College for Women, at New London. (This college has now been "liberated" to Connecticut College and men go there.) Being alone among all those women, Avery shouted for help to Sinnott at Columbia. Sinnott sent him Burkholder. Burkholder promptly made his late-blooming course in bac- teriology pay off. He taught the girls about germs, but now his interest in plant physiology returned. Avery started his work on plant hormones at Connecticut College and since it was a

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PAUL RUFUS BURKHOLDER .9 timely subject, they collaborated on work in this field and produced a book on it. The next three years yielded an im- pressive return in papers published. BACK TO AQUATIC BIOLOGY While at New London, Paul purchased a small farm near Falmouth, Massachusetts, on Cape Cod. Falmouth harbor was near, and so Burkholder bought a sail boat and returned to the water. Wood's Hole was near, too, and here was the oppor- tunity to return to the study of the creatures that live in the water. This lasted only three years, however. In 1938 Dr. Rickett left the post of plant physiologist at the University of Missouri and the boy from the hills of Pennsylvania went to the hills of Missouri. This seemed a logical move. A large well-staffed agricultural school provided many interested students for the botany courses. Despite much teaching to be done, he managed to carry on research work, mostly in plant nutrition. Here the third son, Karl, was born in 1940. When he left, Dr. Tucker, chairman of botany, jokingly said, "I will have to hire two men to take your place, a day man and a night man." A YALE MAN In 1940 Sinnott entered his life again. Dr. Sinnott went to Yale as chairman of the botany department and asked Burk- holder to go with him as plant physiologist. Botany at Yale was at an all-time low ebb, partly because of the depression and partly because of a failure to bring in new faculty members, but mostly because the university had no agriculture program to send the students of botany on to jobs. Zoology was strong at Yale because there was a medical school to beckon the students from that department. The next few years were spent in building up the faculty of botany. Norman Giles, a new Ph.D. in plant genetics, came

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10 BIOGRAPHICAL MEMOIRS from Harvard; Reader came as a taxonomist in charge of the herbarium. Ed Tatum brought his Neurospora work and Dave Bonner followed. Burkholder taught a new course in micro- biology that became very popular. Galston came to teach plant physiology. Paul Sears set up a program in conservation. The department was on the way. Graduate students ap- peared. Joshua Lederberg, who discovered sex in bacteria; Ed Adelberg, now head of microbiology at Yale; Guillard at Wood's Hole; Sy Pomper; Lou Nickell; Ina May Martin from Jamaica; Joyce and Ralph Lewin; and many others received degrees from the department. The years were good at Yale. Burkholder was made Eaton professor of botany. His research moved apace, especially in microbiology, production of vitamins by yeasts, and production of antibiotics by lower fungi. CHLORAMPHENICOL DISCOVERED Burkholder soon made a dramatic discovery. Fleming's penicillin had arrived during the war. Penicillin came from a fungus. What other antibiotics from microorganisms could be found? Waksman had discovered streptomycin and received the Nobel Prize. Burkholder soon discovered chloramphenicol in an actinomycete, which he isolated from a Venezuelan soil. The Parke-Davis Company developed, purified, and synthe- . . sized it. Chloramphenicol was discovered just in time to save the lives of soldiers in the Korean War from scrub typhus, a classical killer of soldiers in wartime. Chloramphenicol is a dramatic case in biology for another reason. Chemists like to chlorinate and nitrate compounds. In 1947, when chloramphenicol was discovered, dogma said that no organism could do these chemical tricks. They couldn't attach a chlorine or a NOR group to carbon as chemists could. Chlorine and nitro were the substituents chemists used to make

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PAUI, RUFUS BURKHOLDER 11 killers for organismsnote dinitrophenol wood preservative or chlorinated hydrocarbon insecticides. Burkholder's actinomycete proved the dogma wrong. Not only could it chlorinate a compound, it could nitrate it, too; and besides it could put both substituents in the same molecule. This only proves that chemists were a million years late in learning these tricks. A PALACE REVOLUTION In the meantime, Yale botany was not all serene. Burk- holder became the unfortunate victim of a palace revolution. Sinnott had held a brilliant group of thinkers together, but then he moved up to be dean of the Sheffield Scientific School. Having always supported Burkholder, Sinnott made him chair- man of the botany department, but Burkholder had no liking for scientific politics and had not developed administrative skills. Severe unhappiness showed in the microbiological con- tingent. And Burkholder didn't really hold a union card in microbiology. He came into it late as a plant physiologist. This problem was tied to the national picture as well. The bac- teriologists of the country had arrogated to themselves the term microbiology. People working with other microorganisms like protozoa, fungi, and one-celled algae were low in the pecking order; and Burkholder's specialty was fungi. Bacteria are mostly dealt with in two university disciplines medicine and agriculture. Yale had no agriculture. Medicine, therefore, became a magnet for the new microbiology created by Burkholder at Yale in the botany department. And so it leas natural for friction to develop within classical botany. Whereupon, botany lost another child that it had begotten. The explosion was severe. Tatum left, Galston left, Naylor left, Burkholder left, and Bonner deserted the botany depart- ment for the medical school and a new department of micro-

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12 BIOGRAPHICAL MEMOIRS biology. It seems ironical that Bonner was later elected to the National Academy of Sciences through the botany section, not through microbiology. GEORGIA BECKONS Georgia beckoned to Burkholder and Burkholder listened. He labored strenuously there to build up science in general and microbiology in particular, but it went too slowly for him. The Sloan-Kettering Cancer Institute in New York thought so, too. Cornelius Rhoades, its director, suggested to George Avery, by then director of the Brooklyn Botanic Garden, that Burkholder could be moved. He could be and was. The next five years at the Garden, with his old friend George Avery and with Sloan-Kettering cooperation, saw an intensive program of screening soil organisms for antibiotics that might have the potential for chemotherapy of cancer. There is a fascinating sequel to the Sloan-Ketter~ng phase. They kept Burkholder's cultures "on deposit" when he left. Rhoades chanced to mention this to Jasper Kane, then a re- search executive with the Pfizer Company. Kane procured the cultures for futher screening. In one of these was the fungus that produces Terramycin for Pfizer. BACK TO COLUMBIA Since the cancer idea did not pay off, Burkholder returned in 1961 to Columbia and to his old love for the sea and marine biology. Perhaps the inhabitants of the sea could provide the hoped-for antibiotics that fungi from the worked-over soil could no longer supply. He affiliated with the new biology laboratory in the Lamont Geological Observatory at Columbia, situated up the Hudson River from New York. He was no longer land- locked. He had come full circle back to the organisms in the water. He spent a season in the Antarctic working with Sieburth

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PAUL RUFUS BURKHOLDER BIBLIOGRAPHY KEY TO ABBREVIATIONS 15 Am. I. Bot. = American Journal of Botany Ann. N.Y. Acad. Sci. -_ Annals of the New York Academy of Sciences Appl. Microbiol. = Applied Microbiology Arch. Biochem.Archives of Biochemistry Bot. Gaz. _ Botanical Gazette Bot. Mar. = Botanica Marina Bull. Buffalo Soc. Nat. Sci. _ Bulletin of the Buffalo Society of Natural Sciences Bull. Mar. Sci. Gulf Caribb. _ Bulletin of Marine Science of the Gulf and Caribbean Bull. Torrey Bot. Club _ Bulletin of the Torrey Botanical Club I. Antibiot. = {ournal of Antibiotics i. Bacterial. = journal of Bacteriology .T Org. Chem. = Journal of Organic Chemistry Limnol. Oceanogr. _ Limnology and Oceanography Mar. Biol. = Marine Biology Plant Physiol. = Plant Physiology Proc. Natl. Acad. Sci. = Proceedings of the National Academy of Sciences Spec. Sci. Rep. Fish. = Special Scientific Report, Fisheries 1930 Microplankton studies of Lake Erie. Bull. Buffalo Soc. Nat. Sci., 14:73-93. The heredity and environment exhibit. Hobbies. The Magazine of the Buffalo Museum of Science. 1931 Studies in the phytoplankton of the Cayuga Lake basin, New York. Bull. Buffalo Soc. Nat. Sci., 15: 181. 1933 A study of the phytoplankton of Frenchman's Bay and Penobscot Bay, Maine. Internationale Revue gesamten Hydrobiologie und Hydrographie, 28:262-84. Movement in cyanophyceae. The effect of pH upon movement in Oscillatoria. Journal of General Physiology, 16:875-81. 1934 Movement in the cyanophyceae. The Quarterly Review of Biology, 9:438-59.

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16 BIOGRAPHICAL MEMOIRS 1936 With R. Pratt. Leaf-movement of Mimosa pud ica in relation to light. Am. I. Bot., 23 :46-52. The role of light in the life of plants. The Botanical Review, 2:1-52, 97-168. With G. S. Avery. Polarized growth and cell studies on the Arena coleoptile, phyto-hormone test object. Bull. Torrey Bot. Club, 63: 1-15. With Boysen-lensen and G. S. Avery. Growth Hormones in Plants. New York: McGraw-Hill Book Co. 268 pp. With G. S. Avery and H. B. Creighton. Plant hormones and mineral nutrition. Proc. Natl. Acad. Sci., 22:673-78. 1937 With E. S. Johnston. Inactivation of plant growth substance by light. Smithsonian Miscellaneous Collections, 95~20~: 1-14. Production and distribution of growth hormone in shoots of Aescu- lus and Malus and its probable role in stimulating cambial activity. Am. l. Bot., 24:51-58. \ With G. S. Avery and H. B. Creighton. Avena coleoptile curvature in relation to different concentrations of certain synthetic substances. Am. l. Bot., 24:226-32. With G. S. Avery and H. B. Creighton. Nutrient deficiencies and growth hormone concentration in Helianthus and Nicotiana. Am. J. Bot., 24: 553-57. With G. S. Avery and H. B. Creighton. Growth hormone in termi- nal shoots of Nicotiana in relation to light. Am. J. Bot., 24:666-73. With G. S. Avery and H. B. Creighton. Polarized growth and cell studies in the first internode and coleoptile of Avena in relation to light and darkness. Bot. Gaz., 99: 125-43. 1938 With G. S. Avery, H. B. Creighton, and B. A. Scheer. Darwin and early discoveries in connection with plant hormones. Science, 87:56.

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PAUL RUFUS BURKHOLDER 1939 17 Production of growth substance by bacteria in media containing specific organic and inorganic nitrogenous compounds. Am. l. Bot., 26:422-28. 1940 With I. McVeigh. Growth and differentiation of maize in relation to nitrogen supply. Am. J. Bot., 27:414-24. With I. McVeigh. Growth of Phycomyces Blakesleeanus in relation to varied environmental conditions. Am. [. Bot., 27:634~0. With I. McVeigh. Studies on thiamine in green plants with the Phycomyces assay method. Am. J. Bot., 27:853-61. 1941 With I. McVeigh. Multinucleate plant cells. Bull. Torrey Bot. Club, 68: 395-96. With C. M. Tucker. Calcium deficiency as a factor of abnormal rooting of Philodendron cuttings. Phytopathology, 31:844~8. Some experiments with growth curvatures and growth substances. Am. l. Bot., 28:911-21. 1942 Pyridoxine as a growth factor for Graphium. Science, 95:127-28. With A. G. Snow, Jr. Thiamine in some common American trees. Bull. Torrey Bot. Club, 69:421-28. Synthesis of vitamins by intestinal bacteria. Proc. Natl. Acad. Sci.. 28:285-89. With I. McVeigh. The increase of B vitamins in germinating seeds. Proc. Natl. Acad. Sci., 28:440-46. 1943 Vitamin deficiencies in yeasts. Am. I. Bot., 30:206-11. Vitamins in dehydrated seeds and sprouts. Science, 97:562-64. Synthesis of riboflavin in a yeast. Proc. Natl. Acad. Sci., 29:166-72. With D. Moyer. Vitamin deficiencies of fifty yeasts and molds. Bull. Torrey Bot. Club, 70:372-77. Vitamins in edible soybeans. Science, 98: 188-90. Influence of some environmental factors upon the production of riboflavin by a yeast. Arch. Biochem., 3: 121-29.

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18 BIOGRAPHICAL MEMOIRS With l. Collier and D. Moyer. Synthesis of vitamins by micro- organisms in relation to vitamin content of fancy cheeses. Food Research, 8:314-22. 1944 With I. McVeigh and D. Moyer. Niacin in maize. Yale [ournal of Biology and Medicine, 16~6~: 659-63. With A. W. Evans, I. McVeigh, and H. K. Thornton. Antibiotic activity of lichens. Proc. Natl. Acad. Sci., 30:250-55. Some growth patterns of bacteria in cylinder plate tests for pro- moting and inhibiting substances. Am. I. Bot., 31:555-58. With I. McVeigh and D. Moyer. Studies on some growth factors of yeasts. I. Bacteriol-, 48: 385-91. 1945 With A. W. Evans. Further studies on the antibiotic activity of lichens. Bull. Torry Bot. Club, 72: 157-64. With E. W. Sinnott. Morphogenesis of fungus colonies. in sub- merged shaken cultures. Am. I. Bot., 32:424-31. With I. McVeigh and K. Wilson. Studies on vitamin "B" produced by microorganisms. Arch. Biochem., 7:287-303. With I. McVeigh. The B vitamin content of buds and shoots of some common trees. Plant Physiol., 20:276-82. With I. McVeigh. Vitamin content of some mature and germinated legume seeds. Plant Physiol., 20:301-6. Studies on the antibiotic activity of actinomycetes. l. Bacteriol., 52: 503~. 1947 With Norman H. Giles, in Induced biochemical mutation in Bacillus sub tills. Am. J. Bot., 34: 345~8. With L. G. Nickell. Inhibition of Azotobacter by soil actinomy- cetes. Journal of the American Society of Agronomy, 39:771-79. Chloromycetin, a new antibiotic from a soil actinomycete. Science, 106:2757. 1948 With [. Ehrlich, D. Gottlieb, L. E. Anderson, and T. G. Pridham. Streptomyces Venezuelae, n. sp., the source of chloromycetin. I. Bacteriol., 56:467-77.

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PAUL RUFUS BURKHOLDER 19 With L. G. Nickell. Growth responses of Rumex virus wound tumor in tissue culture. Am. l. Bot., 35:811. With Mary E. Sanders. Influence of amino acids on growth of Datura embryos in culture. Proc. Natl. Acad. Sci., 34:516-26. 1949 With L. G. Nickell. Atypical growth of plants. I. Cultivation of virus tumors of Rumex on nutrient agar. Bot. Gaz., 110:426-37. With Seymour Pomper. Studies on the biochemical genetics of yeast. Proc. Natl. Acad. Sci., 35:456-64. With R. G. H. Sin, R. T. Darby, and E. S. Barghoorn. Specificity of microbiological attack on cellulose derivatives. Textile Re- search, 29~8~:484-88. 1950 With L. G. Nickell. Ayptical growth of plants. II. Growth in vitro of virus tumors of Rumex in relation to temperature, pH and various sources of nitrogen, carbon, and sulfur. Am. l. Bot., 37:538~7. With L. G. Nickell and P. Greenfield. Atypical growth of plants. III. Growth responses of virus tumors of Rumex to certain nucleic acid components and related compounds. Bot. Gaz., 112:42-52. 1951 Determination of vitamin B-12 with a mutant strain of E. coli. Science, 114:459-60. 1952 Microbial studies on materials which potentiate oral vitamin B-12 therapy in Addisonian anemia. Archives of Biochemistry and Biophysics, 39:322-32. Cooperation and conflict among primitive organisms. (Sigma Xi lecture) American Scientist, 40:601-31. 1954 Speciation and variation in asexual fungi. Ann. N.Y. Acad. Sci., 60: 1-82.

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20 BIOGRAPHICAL MEMOIRS With Sung Huang Sun, l. Ehrlich, and L. E. Anderson. Criteria of speciation in the genus Streptomyces. Ann. N.Y. Acad. Sci., 60:102-23. The spirit of science. Georgia Review, 8~4~:373-82. 1955 With Sung Huang Sun, L. E. Anderson, and I. Ehrlich. The iden- tity of viomycin-producing cultures of Streptomyces. Bull. Torrey Bot. Club, 82:108-17. 1956 With L. E. Anderson, l. Ehrlich, and Sung Huang Sun. Strains of Streptomyces, the sources of azaserine, elaiomycin, griseoviridin, and viridogrisein. Antibiotics and Chemotherapy, 6~2~: 100-115. With L. M. Burkholder. Microbiological assay of B-12 in marine solids. Science, 123: 1071-73. With L. M. Burkholder. Vitamin B-12 in suspended solids and marsh muds collected along the coast of Georgia. Limnol. Oceanogr., 1:202-8. Studies on the nutritive value of spartina grass growing in the marsh areas of coastal Georgia. Bull. Torrey Bot. Club, 83: 327-34. 1957 With G. H. Bornside. Decomposition of marsh grass by aerobic marine bacteria. Bull. Torrey Bot. Club, 84:366-83. 1958 With L. M. Burkholder. Antimicrobial activity of horny corals. Science, 127:1174-75. Studies on B vitamins in relation to productivity of the Bahia Fosforescente, Puerto Rico. Bull. Mar. Sci. Gulf Caribb., 8:201-23. 1959 Some microbiological aspects of marine productivity in shallow waters. In: Proceedings of the Salt Marsh Conference, ed. by R. A. Ragotzkie, pp. 70-72. Sapelo Island, Georgia, Marine Institute, 1958. University of Georgia: Marine Institute.

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PAUL RUFUS BURKHOLDER ~1 With L. M. Burkholder and l. A. Rivero. Some chemical constitu- ents of turtle grass, Thalassia tested inum. Bull. Torrey Bot. Club, 86:88-93. With L. M. Burkholder and I. A. Rivero. Chlorophyll A in some corals and marine plants. Nature, 183: 1338-39. Organic nutrition of some mosses mowing, in nilrP rillt~l'- gist, 62:6-15. Antibiotics -the exploitation of microbial antagonisms is having a challenging impact on medicine and challenging society. Science, 129:1457-65. With H. T. Odum and I. Rivero. Measurements of productivity of turtle grass flats, reefs, and the Bahia Fosforescente of south- ern Puerto Rico. Publications of the Institute of Marine Science, University of Texas, Port Aransas, 6: 159-70. - ~-~ ---a -- ret ace. Bryolo- 1960 Distribution of some chemical values in Lake Erie. In: Limnologi- cal Survey of Eastern and Central Lake E,^ie, 1928-29, pp. 71- 110. Spec. Sci. Rep. Fish. no. 334, June 1960. A survey of the microplankton of Lake Erie. In: Limnological Survey ' of Eastern and Central Lake Erie, 1928-29, pp. 123-44. Spec. Sci. Rep. Fish. no. 334, tune 1960. With L. M. Burkholder and L. A. Almodovar. Antibiotic activity of some marine algae of Puerto Rico. Bot. Mar., 2:149-56. General microbiology of the Antarctic. In: Science in Antarctica. I. The Life Sciences in A ntarctica, pp. 129-37. Washington, D.C.: National Academy of Sciences-National Research Council Publication 839. With M. D. Tendler. Studies on the thermophilic actinomycetes. Appl. Microbiol., 9:394-99. With L. M. Burkholder. Photosynthesis in some alcyonacean corals. Am. [. Bot., 47: 866-72. 1961 With l. M. Siel~urth. Phytoplankton and chlorophyll in the Ger- lache and Bransfield straits of Antarctica. Limnol. Oceanogr., 6~1~:45-52.

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22 BIOGRAPHICAL MEMOIRS 1963 Some nutritional relationships among microbes of sea sediments and waters. In: Symposium on Marine Microbiology, pp. 133-50. Springfield, Ill.: Charles C. Thomas. Radioactivity in some aquatic plants. Nature, 198:601-3. Drugs from the sea. Armed Forces Chemical Journal, 17:6-16. 1965 With C. W. Dodge and L. M. Burkholder. Estudio de los liquenes de Tierra del Fuego con especial consideration de suactividad antibiotica. Centro de Investigacion de Biologia Marina, Buenos Aires, Argentina. Contribuciones Cientificas no. 21, 1-24. With R. M. Pfister. Numerical taxonomy of some bacteria isolated from Antarctica and tropical seawaters. i. Bacteriol., 90: 863-72. With E. F. Mandelli. Carbon assimilation of marine phytoplankton in Antarctica. Proc. Natl. Acad. Sci., 54:437-44. With E. F. Mandelli. Productivity of microalgae in Antarctic sea ice. Science, 149: 872-74. With A. Repak and l. Siebert. Studies on some Long Island Sound littoral communities of microorganisms and their primary pro- ductivity. Bull. Torrey Bot. Club, 92:378~02. 1966 With R. M. Pfister and F. H. Leitz. Production of a pyrole anti- biotic by a marine bacterium. Appl. Microbial., 14:649-53. With L. M. Burkholder and P. Genteno. Nutritive values of shrimp flour. Nature, 211: 860-61. With E. F. Mandelli. Primary productivity in the Gerlache and Bransfield straits of Antarctica. Journal of Marine Research, 24:15-27. 1967 With L. M. Burkholder and L. R. Almodovar. Carbon assimilation of marine flagellate blooms in neritic waters of southern Puerto Rico. Bull. Mar. Sci. Gulf Caribb., 17: 1-15. With E. F. Mandelli and P. Centeno. Some chemical properties of Munida gregaria and Euphausia superba. journal of Agricul- tural and Food Chemistry, 1 ~5 :718-20.

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PAUL RUFUS BURKHOLDER ~3 With G. M. Sharma. Studies on antimicrobial substances of sponges. I. Isolation, purification, and properties of a new bromine-con- taining antibacterial substance. l. Antibiot. (Japan), 20:200-203. With G. M. Sharma. Studies on antimicrobial substances of sponges. II. Structure and synthesis of a bromine-containing antibacterial compound from a marine sponge. Tetrahedron Letters, 42: 4147-50. With L. M. Burkholder. Primary productivity in surface waters of the South Pacific ocean. Limnol. Oceanogr., 12:606-17. 1968 With L. M. Burkholder, A. Chu, N. Kostyk, and O. A. Roels. Fish fermentation. Food Technology, 22: 1278-84. Patterns of B vitamin requirements of neritic Canadian journal of Microbiology, 14:537-43. marine bacteria. With H. G. Udell, I. Zarudsky, and T. E. Doheny. Productivity and nutrient values of plants growing in the salt marshes of the town of Hempstead, Long Island. Bull. Torrey Bot. Club, 96:42-51. With T. E. Doheny. Biology of eelgrass. Contribution no. 3. Hempstead, L.I.: Department of Conservation and Waterways, Marine Laboratory. Antimicrobial substances from the sea. In: Drugs from the Sea, ed. by Hugo D. Freudenthal, pp. 87-112. Transactions of a symposium held at the University of Rhode Island, 1967. Washington, D.C.: Marine Technology Society. With G. M. Sharma and B. Vig. Studies on antimicrobial sub- stances of sponges. III. Chemical properties of some antibacterial compounds from marine sponges. In: Drugs from the Sea, ed. by Hugo D. Freudenthal, pp. 119-26. Transactions of a sym- posium held at the University of Rhode Island, 1967. Wash- ington, D.C.: Marine Technology Society. With G. M. Sharma and L. Michaels. Coniodomin, a new antibiotic from a dinoflagellate. l. Antibiot. Japan), 21:659-64. With O. A. Roels. Biological oceanology at Lamont Geological Observatory, pp. 72-85. New York State Science and Tech- nology Foundation.

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24 BIOGRAPHICAL MEMOIRS 1969 With K. Ruetzler. Antimicrobial activity of some marine sponges. Nature, 222:983-84. (L) With G. M. Sharma. Antimicrobial agents from the sea. Lloydia, 32:466-88. 1970 With G. M. Sharma and B. Vig. Studies on antimicrobial substances of sponges. IV. Structure of a bromine-containing compound from a marine sponge. J. Org. Chem., 35:2823-26. With P. Hargraves and R. Brody. Phytoplankton of the lesser An- tilles region. Bull. Mar. Sci. Gulf Caribb., 20:331-49. Some biomedical aspects of marine microbiology. In: Food-Drugs from the Sea Proceedings 1969, ed. by H. W. Youngken, Jr., pp. 255-79. Held at the University of Rhode Island, 1969. Washington, D.C.: Marine Technology Society. With G. M. Sharma and B. Vig. Antimicrobial substances of marine sponges. IV. In: Food-Drugs from the Sea Proceed ings 1969, ed. by H. W. Youngken, Jr., pp. 307-10. Held at the University of Rhode Island, 1969. Washington, D.C.: Marine Technology Society. With G. Cintron and W. S. Maddux. Consequences of brine pollu- tion in the Bahia Fosforescente, Puerto Rico. Limnol. Oceanogr., 15(2):246~9. With E. F. Mandelli, T. Doheny, and R. Brady. Primary produc- tivity in coastal waters of southern Long Island, New York. Mar. Biol., 7~2~: 153-60. 1971 With L. Almodovar. Species composition and productivity of man- grove algae communities in Puerto Rico. Florida Scientist, 36:66-74. With G. M. Sharma. Structure of dibromophakellin, a new bro- mine-containing alkaloid from the marine sponge Phakellia flabellata. Chemical Communications (London), pp. 151-52. With L. Almodovar and L. M. Burkholder. Nutritive constituents of some Caribbean marine algae. Bot. Mar., 14~2~:132-35.

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PAUL RUFUS BURKHOLDER 25 With T. Doheny. Proximate analysis and amino acid composition of some marine phytoplankton and bacteria. Contribution no. 8. Hempstead, L.I.: Department of Conservation and Waterways, Marine Laboratory. The ecology of marine antibiotics. In: Coral Reefs of The World, ed. by R. Endean and 0. A. tones, pp. 117-82. New York: Academic Press, Inc. 1972 With T. E. Doheny. Use of selected enrichment experiments in predicting the eutrophication of an estuary. Contribution no. 7. Hempstead, L.I.: Department of Conservation and Waterways, Marine Laboratory. With R. Brody and A. E. Dammann. Some phytoplankton blooms in the Virgin Islands. Caribbean Journal of Science, 12:23-28. \