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EDWARD ARTHUR STEINHAUS 317 not dwell on conflicts between religion and science but rather that the two philosophies should team up in their approach to a solution of "what goes wrong" in the world. His viewpoint on this subject was in keeping, with his philosophy of life that moti- vated him to accomplish so much through science and education for the welfare of mankind. IN PREPARING this biographical memoir, I am especially indebted to l\labry C. Steinhaus for providing me with reference material not otherwise available. This was largely in the nature of manuscripts of biographies written by others, including those prepared by E. Gorton Linsley and Ray F. Smith of the University of California, Berkeley; fames L. McGaugh, Howard A. Schneiderman, and John E. Smith of the University of California, Irvine; John D. Briggs, Ohio State University, and Mauro Martignoni and Ken Hughes of the U.S. Forestry Sciences Laboratory, Oregon State University, Corvallis. I also drew from biographical information obtained from the National Academy of Sciences. An unpublished docu- ment prepared by E. A. Steinhaus himself contained most valu- able autobiographical information, which was kindly included in the material supplied by Mrs. Steinhaus. I wish also to express my appreciation for the assistance by my daughter, Edwina, in the organization and editing of the biographical memoir.

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318 BIOGRAPHICAL MEMOIRS BIBLIOGRAPHY KEY TO ABBREVIATIONS Bacteriol. Rev. _ Bacteriological Reviews Bull. Entomol. Soc. Am. Bulletin of the Entomological Society of America Calif. Agr. California Agriculture I. Bacteriol. journal of Bacteriology I. Econ. Entomol. _ journal of Economic Entomology I. Insect Pathol. Journal of Insect Pathology I. Invert. Pathol. Journal of Invertebrate Pathology J. Parasitol. Journal of Parasitology Public Health Rept. Public Health Reports 1936 The effect of Escherichia cold on the growth of Bacillus subtilis when grown in mixed cultures. North Dakota State College Thesis (B.S.), Fargo, North Dakota. 31 pp. 1938 With I. M. Birkeland. teriol., 36:216. (A) "Cannibalism" among bacteria. l. Bac- 1939 With I. M. Birkeland. Selective bacteriostatic action of sodium lauryl sulfate and of "Dreft." Proceedings of the Society of Experimental Biology and Medicine, 40:86-88. With I. M. Birkeland. Studies on the life and death of bac- teria. I. The senescent phase in aging cultures and the probable mechanisms involved. I. Bacteriol., 38: 249-61. 1940 Studies on the life and death of bacteria. Abstracts of Doctoral Dis- sertations, No. 31, pp. 325-31. Columbus, Ohio State University Press. A discussion of the microbial flora of insects. 62. (A) J. Bacteriol., 40: 161- . The microbiology of insects with special reference to the biologic relationships between bacteria and insects. Bacteriol. Rev., 4: 17-57. 1941 A study of the bacteria associated with thirty species of insects. l. Bacteriol., 42: 757-89.

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EDWARD ARTHUR STEINHAUS 1942 319 Note on a toxic principle in eggs of the tick, Dermacentor andersoni Stiles. Public Health Rept., 57: 1310-12. Rickettsia-like organism from normal Dermacentor andersoni Stiles. Public Health Rept., 57:1375-77. The microbial flora of the Rocky Mountain wood tick, Dermacentor and ersoni Stiles. I. Bacteriol., 43:91-92. (A) The microbial flora of the Rocky Mountain wood tick, Dermacentor and ersoni Stiles. l. Bacteriol., 44: 397-404. With G. M. Kohls. Isolation of an acid-fast bacillus from a hawk. Journal of the American Veterinary Medical Association, 51:502. Catalogue of Bacteria Associated Extracellularly with Insects and Ticks. Minneapolis, Burgess Publishing Company. 206 pp. 1943 With R. R. Parker and G. M. Kohls. Tularemia in beavers and muskrats and contamination of natural waters and mud by Pasteurella tularensis in the northwestern United States. l. Bacteriol., 45: 56-57. A new bacterium, Corynebacterium lipoptenae, associated with the louse fly, Lipoptena depressa Say. i. Parasitol., 29:80. With R. R. Parker. Rocky Mountain spotted fever: duration of potency of tick-tissue vaccine. Public Health Rept., 58:230-32. With R. R. Parker. Experimental Rocky Mountain spotted fever: results of treatment with certain drugs. Public Health Rept., 58:351-52. With R. R. Parker and G. M. Kohls. Amblyomma americanum a vector of Rocky Mountain spotted fever. Public Health Rept., 58:491. With R. R. Parker. American and Australian Q fevers: persistence of the infectious agents in guinea pig tissues after defervescence. Public Health Rept., 58:523-27. With R. R. Parker and G. M. Kohls. Rocky Mountain spotted fever: spontaneous infection in the tick Amblyomma ameri- canum. Public Health Rept., 58:721-29. With G. M. Kohls. Tularemia: spontaneous occurrence in shrews. Public Health Rept., 58:842. With R. R. Parker. Salmonella enteritid is: experimental trans- mission by the Rocky Mountain wood tick Dermacentor ander- soni Stiles. Public Health Rept., 58: 1010-12.

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320 BIOGRAPHICAL MEMOIRS With H. B. Foote, W. L. Allison, and G. M. Kohls. Effect of chlorination on Pasteurella tularensis in aqueous suspension. Journal of the American Water Works Association, 35:902-10. 1944 With R. R. Parker and M. T. McKee. Cultivation of Pasteurella tularensis in a liquid medium. Public Health Rept., 59:78-79. The isolation of a filter-passing agent from the rabbit tick Haemaphysalis leporis-palustris Packard. Public Health Rept., 59: 1528-29. With T. L. Perrin. Pathologic reaction in guinea pigs to Humphreys' virus strain. Public Health Rept., 59: 1603-9. With R. R. Parker. 1945 Insect pathology and biological control. If. Econ. Entomol., 38:591- 96. Bacterial infections of potato tuber moth larvae in an insectary. i. Econ. Entomol., 38:718-19. 1946 Insect Microbiology. Ithaca, Comstock Pub. Co., Inc. 763 pp. An orientation with respect to members of the genus Bacillus patho- genic for insects. Bacteriol. Rev., 10:51-61. 1947 With L. E. Hughes. Isolation of an unidentified spirochete from hen's eggs after inoculation with liver tissue from hens. Public Health Rept., 62: 309-11. A new disease of the variegated cutworm, Peridroma margaritosa (Haw). Science, 106: 323. Control of insect pests by means of disease agents. Calif. Agr., 1:~. A coccidian parasite of Ephestia kuhniella Zeller and of Plod ia interpunctella (Hbn. ~ (Lepidoptera, Phycitidae>. i. Parasitol., 33:29-32. 1948 Polyhedrosis ("wilt disease") of the alfalfa caterpillar. i. Econ. Entomol., 41:859-65. With Albert Abdel-Malek. Invasion route of Nosema sp. in the potato tuberworm, as determined by iigaturing. J. Parasitol., 34:452-53.

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EDWARD ARTHUR STEINHAUS 1949 321 With K. hi. Hughes. Two newly described species of Microsporidia from the potato tuberworm, Gnorimoschema operculella (Zeller) (Lepidoptera, Gelechiidae). l. Parasitol., 35: 67-75. With K. M. Hughes and H. B. Wasser. Demonstration of the granu- losis virus of the variegated cutworm. l. Bacteriol., 57:219-24. Edith C. G. Thompson. Alfalfa caterpillar tests; biological control by artificial spread of virus disease. Calif. Agr., 3:~-6. With C. G. Thompson. Preliminary field tests using a polyhedrosis virus to control the alfalfa caterpillar. i. Econ. Entomol., 42: 301-~. Insect pathology: the field concerned, training required, and oppor- tunities possible. Canadian Entomologist, 81:53-57. Principles of Insect Pathology. New York, McGraw-Hill Book Co., Inc. 757 pp. M7ith C. G. Thompson. Granulosis disease in the buckeye cater- pillar, f unonia coenia Hubner. Science, 110: 276-78. Nomenclature and classification of insect viruses. Bacteriol. Rev., 13:203-23. 1950 With C. G. Thompson. Further tests using a polyhedrosis virus to control the alfalfa caterpillar. Hilgardia, 19:411-45. \'\lith C. G. Thompson. Alfalfa caterpillar control; treatment of fields by airplane application of spray advances destruction of pest. Calif. Agr., 4:8, 16. Diagnoses of insect diseases; microbial infections in insects diagnosed as part of the research in developing new ways of controlling crop pests. Calif. Agr., 4: 11, 15. 1951 Possible use of Bacillus thuringiensis Berliner as an aid in the bio- logical control of the alfalfa caterpillar. Hilgardia, 20:359-81. Report on diagnoses of diseased insects, 1944-1950. Hilgardia, 20: 629-78. Edith R. R. Parker, G. M. Kohls, and W. L. {ellison. Contamina- tion of natural waters and mud with Pasteurella tularensis and tularemia in beavers and muskrats in the northwestern United States. National Institutes of Health Bulletin, No. 193. Wash- ington, U.S. Govt. Print. Off. 61 pp.

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322 BIOGRAPHICAL MEMOIRS With H. B. Wasser. Isolation of a virus causing granulosis in the red-banded leaf roller. Virginia Journal of Science, 2:91-93. Pest control by bacteria; alfalfa caterpillar in field reduced to subeconomic levels within two days by bacillus applied as spray. Calif. Agr., 5:5. 1952 Microbial infections in European corn borer larvae held in the laboratory. I. Econ. Entomol., 45:48-51. With K. M. Hughes. A granulosis of the western grape leaf skeletonized. l. Econ. Entomol., 45:744-45. The susceptibility of two species of Colias to the same virus. l. Econ. Entomol., 45:897-99. Infectious diseases of insects. In: Insects (Yearbook of Agriculture, 1952), ed. by Alfred Stefferud, pp. 388-94. Washington, U.S. Govt. Print. Off. 1953 Taxonomy of insect viruses. Annals of the New York Academy of Sciences, 56:517-37. With C. Ritchie Bell. The effect of certain microorganisms and antibiotics on stored-grain insects. I. Econ. Entomol., 46:582- 98. Diseases of insects reared in the laboratory or insectary. University of California, College of Agriculture, Leaflet No. 9. 26 pp. lg54 With E. A. Sorrel. Further observations on Bacillus thuringiensis Berliner and other spore-forming bacteria. Hilgardia, 23:1-23. The eRects of disease on insect populations. Hilgardia, 23: 197-261. Insects on stamps. Weekly Philatelic Gossip, 58:172-75. Duration of infectivity of the virus of silkworm jaundice. Science, 120: 186-87. 1955 Observations on the symbioses of certain Coccidae. 185-206. 1956 \Vith M. M. Batey and C. L. Boerke. caeca of certain Heteroptera. Hilgardia, 24:495-518. Hilgardia, 24: Bacterial symbioses from the

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EDWARD ARTHUR STEINHAUS 323 Microbial control the emergence of an idea. 60. Editor, wills R. F. Smith. Annual Review of Entomology, Vol. 1. Stanford, California, Annual Reviews, Inc. 466 pp. (Beginning with Vol. 2, 1957, place of publishers: Palo Alto, California). Vol. 2, 1957, 407 pp.; Vol. 3, 1958, 520 pp.; Vol. 4, 195D, 467 pp.; Vol. 5, 1960, 451 pp.; Vol. 6, 1961, 470 pp.; Vol. 7, 1962, 536 pp. Scientific American, 195:96-104. Living insecticides. Potentialities for microbial control of insects. tural and Food Chemistry, 4:676-80. 1957 Hilgardia, 26: 107- Journal of Agricul- New records of insect-virus diseases. Hilgardia, 26:417-30. With Robert L. Rabb and Frank E. Guthrie. Preliminary tests using Bacillus thuringiensis Berliner against hornworms. l. Econ. Entomol., 50: 259-62. Concerning the harmlessness of insect pathogens and the standard- ization of microbial control products. I. Econ. Entomol., 50: 715-20. Microbial diseases of insects. 165-82. Annual Review of Microbiology, 11: List of insects and their susceptibility to Bacillus thuringiensis Berliner and closely related bacteria. Mimeographed Series, 4: 1-24. Laboratory of Insect Pathology, University of California, Berkeley. With F. I. Brinley. Some relationships between bacteria and certain sewage-inhabiting insects. Mosquito News, 17: 299-302. New horizons in insect pathology. Journal of the New York En- tomological Society, 65: 113-21. 1958 Stress as a factor in insect disease. national Congress on 725-30. Ottawa, The Proceedings of the Xth Inter- Entomology, Montreal, 1956, Vol. 4, pp. Congress, Science Service Building. Crowding as a possible stress factor in insect disease. Ecology, 39: 503-14. 1959 Serratia marcescens Bizio as an insect pathogen. 80. Hilgardia, 28: 351- Granuloses in two Alaskan insects. J. Econ. Entomol., 52:350-52.

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324 BIOGRAPHICAL MEMOIRS On the improbability of Bacillus thuringiensis Berliner mutating to forms pathogenic for vertebrates. J. Econ. Entomol., 52:506- 8. With [. P. Dineen. A cytoplasmic polyhedrosis of the alfalfa cater- pillar. l. Insect Pathol., 1: 1 7 1-83. With I. Lipa. Nosema hippodamiae n. sp., a microsporidian parasite of Hippodamia convergens Guerin (Coleoptera, Coccinel- lidae). [. Insect Pathol., 1:304-8. Possible virus disease in European red mite. 1 :435-37. J. Insect Pathol., Insect pathology and microbial control. Pest Control Review, University of California Agricultural Extension Service, Febru- ary, pp. 1-3. Insect pathology and microbial control. Excerpts from press con- ference. University of California Division of Agricultural Sci- ences. Berkeley, University of California Press. 15 pp. (Special leaflet.) Bacteria as microbial control agents. Transactions of the 1st In- ternational Conference on Insect Pathology and Biological Con- trol, Prague, August 1958, pp. 37-50. 1960 With G. H. Bergold, K. Aizawa, K. M. Smith, and C. Vago. The present status of insect virus nomenclature and classification. International Bulletin of Bacteriological Nomenclature and Taxonomy, 10: 259-62. With i. P. Dineen. Observations on the role of stress in a gran- ulosis of the variegated cutworm. J. Insect Pathol., 2:55-65. With G. A. Marsh. Granulosis of the granulate cutworm. I. Insect Pathol., 2:115-17. The duration of viability and infectivity of certain insect pathogens. J. Insect Pathol., 2:225-29. Notes on polyhedroses in Periodroma, Prodenia, Co lies, Helioth is, and other Lepidoptera. I. Insect Pathol., 2:327-33. With Gertraude Wittig and Joyce P. Dineen. Further studies of the cytoplasmic polyhedrosis virus of the alfalfa caterpillar. J. Insect Pathol., 2: 334~5. Bacterial and viral diseases of insects of medical importance (and other excerpts from Report of Conference on the Biological Con- trol of Insects of Medical Importance, Washington, D.C., Febru-

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EDWARD ARTHUR STEINHAUS ary 1960, pp. 21-27). Biological Sciences. 325 Washington, D.C., American Institute of Insect control, biological. In: McGraw-Hill Encyclopedia of Sci- ence and Technology, Vol. 7, p. 122. New York, McGraw-Hill Book Co., Inc. insect pathology. In: McGraw-Hill Encycloped ia of Science and Technology, Vol. 7, pp. 122a-h. New York, McGraw-Hill Book Co., Inc. Insect pathology: challenge, achievement, and promise. Bull. En- tomol. Soc. Am., 6:9-16. (1959 Entomological Society of America Memorial Lecture.) Tl~e importance of environmental factors in the insect-microbe ecosystem. Bacteriol. Rev., 24:365-73. Some developments in insect pathology and microbial control in the United States. Proceedings of the Society for Study of Plant Protection, 2:151-53. (Translation into Chinese of talk given at National Taiwan University.) 1961 \Vith M. E. Martignoni. Laboratory Exercises in Insect Micro- biology and Insect Pathology. Minneapolis, Burgess Publishing Company. 75 pp. On the correct author of Bacillus sotto. J. Insect Pathol., 3:97-100. 1962 With [. Lipa. Further report on identifications of protozoa patho- genic for insects. Acta Parasitologica Polonica, 10:165-75. Noninfectious disease: an area of neglect in insect pathology. J. Insect Pathol., 4: i-viii. With G. A. Marsh. Report of diagnoses of diseased insects, 1951- 61. Hilgardia, 33: 349-490. 1963 Introduction. Chapter 1 in: Insect Pathology: An Advanced Trea- tise, Vol. I, pp. 1-27. New York, Academic Press, Inc. Background for the diagnosis of insect diseases. Chapter 16 in: Insect Pathology: An Advanced Treatise, Vol. II, pp. 549-89. New York, Academic Press, Inc. Editor. Insect Pathology: An Advanced Treatise. New York, Academic Press, Inc. Vol. I, 661 pp.; Vol. II, 689 pp.

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326 BIOGRAPHICAL MEMOIRS Insect pathology and biomedical research. J. Insect Pathol., 5: i-iv. With Ruth Leutenegger. Icosahedral virus from a scarab (Seri- cesthis). ]. Insect Pathol., 5:266-70. 1964 Microbial diseases of insects. Chapter 18 in: Biological Control of Insect Pests and Weeds, ed. by Paul DeBach and E. I. Schlinger, pp. 515-47. London, Chapman & Hall, Ltd. Pathology, a biological science. J. Insect Pathol., 6:i-v. The day is at hand. Bull. Entomol. Soc. Am., 10:3-7. (Entomo- logical Society of America Presidential Address.) When an insect dies. Bull. Entomol. Soc. Am., 10:183-89. Diagnosis: a central pillar of insect pathology. Colloque interna- tional sur la pathologic des insectes et la lutte microbiologique, Paris, October 1962. Entomophaga Moire No. 2, pp. 7-21. 1965 A new name but the same goals. i. Invert. Pathol., 7:i. External signs of disease and abnormality in the insect egg. J. Invert. Pathol., 7:ii-x. Symposium on microbial insecticides. IV. Diseases of invertebrates other than insects. Bacterial. Rev., 29:388-96. 1966 Greater scholarship in pathology. J. Invert. Pathol., 8:i-ii. With R. D. Zeikus. Observations on a previously undescribed type of cellular degeneration in hydra. I. Invert. Pathol., 8:14-34. Insect control, biological. In: McGraw-Hill Encyclopedia of Sci- ence and Technology, Vol. 7, p. 122. New York, McGraw-Hill Book Co., Inc. Insect pathology. In: McGraw-Hill Encyclopedia of Science and Technology, Vol. 7, pp. 122-122h. New York, McGraw-Hill Book Co., Inc. 1967 Microbial control a comment on its present status in the United States. Bull. Entomol. Soc. Am., 13: 104-8. A guide to the biological sciences. Division of Biological Sciences, University of California, Irvine. 48 pp. (Previous editions: 1965, 1966.) On the importance of invertebrate pathology in comparative pathol-

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EDWARD ARTHUR STEINHAUS and new records. 327 ogy. Revue de Pathologie Comparee, 67:139-42. \\lith G. A. Marsh. Previously unreported accessions for diagnosis I. Invert. Pathol., 9:436-38. With M. E. Martignoni. An abridged glossary of terms used in invertebrate pathology. Pacific Northwest Forest and Range Experiment Station. U.S. Department of Agriculture, Forest Service. 22 pp. 1968 Immunity to infectious diseases in beneficial insects. Abstracts of papers from the XIIIth International Congress of Entomology, Moscow, USSR, p. 258. Hungry people and invertebrate pathology. J. Invert. Pathol., lO:i-iii. Microbial control is not all. Proceedings of the Joint U.S.-lapan Seminar on Microbial Control of Insect Pests, Fukuoka, Kyushu, April 1967, pp. 40-48. With R. D. Zeikus. Teratology of the beetle Tenebrio molitor. I. Gross morphology of certain abnormality types. J. Invert. Pathol., 10:190-210. Centers for pathobiology. l. Invert. Pathol., ll: i-iv. With R. D. Zeikus. Teratology of the beetle Tenebrio molito~r. II. The development and gross description of the pupal-winged adult. l. Invert. Pathol., 11:8-24. With R. D. Zeikus. Teratology of the beetle Tenebrio rnolitor. III. Ultrastructural alterations in the flight musculature of the pupal-winged adult. l. Invert. Pathol., 12:40-52. With R. D. Zeikus. An unusual structural layer in the foregut of the beetle Tenebrio molitor. Submitted for publication in journal of Ultrastructure Research. 1969 With R. D. Zeikus. ~ . Teratology of the beetle Tenebrio molitor. IV. Ultrastructure of the necrotic fat body and foregut associated with the pupal-winged adult. I. Invert. Pathol., 13:337-44. With R. D. Zeikus. Teratology of the beetle Tenebrio molitor. V. Ultrastructural changes and viruslike particles in the foregut epithelium of pupal-winged adults. l. Invert. Pathol., 14:115- 21. Invertebrates as models for the study of diseases of man. Federation Proceedings, 28:1810-14.

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CHESTER HAMLIN WERKMAN 347 with pigeon liver the authors stated that "the dissimilation of pyruvate by pigeon liver occurs with accompanying fixation of carbon dioxide. By use of i3CO2 the fixed carbon has been shown to be exclusively in the carboxyl groups of the 4-carbon dicarboxylic acids (malate, fumarate, and succinate), the carboxyl adjacent to the carbonyl of a-ketoglutarate, and the carboxyl of lactate. Aerobically in the presence of malonate succinate is formed which contains little or no fixed carbon. It is proposed that the 4-carbon dicarboxylic acids are formed by two mechanisms, one reductive through the carbon fixation reaction, the other oxidative by a tentative and modified Krebs cycle which does not involve citric acid. The scheme accounts for the observed positions of the fixed carbon and the aerobic formation in the presence of malonate of succinate not contain- ing fixed carbon." The results provided strong support for Krebs's proposed cycle. H. D. Slade, Wood, Nier, A. Hemingway, and Werkman (J. Biol. Chem., 143: 133-45, 1942) investigated the extent to which other heterotrophic bacteria were capable of carbon dioxide fix- ation. They reported that "fixation of CO2 by C3 and C, addition is apparently a very general reaction," as demonstrated with several genera of heterotrophic bacteria, including A erobacter, Proteus, Staphylococcus, Streptococcus, and Clostridium. In summary, they stated that "the assimilation of CO, is established as a general phenomenon among heterotrophic bacteria. It is shown by the use of heavy carbon, i3C, as a tracer, that the fixed carbon is located in the carboxyl groups of succinic, lactic, and acetic acids. The assimilated CO., is distributed as follows: Aerobacter indologenes, acetate, lactate, and succinate; Proteus vulgaris, Streptococcus paracitrorrorus, and Staphylococcus can- ~ This conclusion was in error and was based on the idea that citrate is a symmetrical molec~le and therefore the ketoglutarate formed from citrate was expected to be labeled in both carboxyl groups. Only when Ogston (Ogston, A. G., Natz~re, 162:4129, 1948) explained that an enzyme can distinguish between the primary carboxyl of citrate did this incorrect conclusion become clarified.

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348 BIOGRAPHICAL MEMOIRS didus, lactate and succinate; Clostridium welchii, acetate and lactate; Clostridium acetobutylicum, lactate." A heat-labile enzyme prepared from ~~icrococcus Iysodeikti- cus was described by L. O. Krampitz and Werkman ( OCR for page 335
CHESTER HAMLIN WERKMAN 349 alcohol fermentation (Wood et al., I. Am. Chem. Soc., 66:1812- 18, 1944) they stated that "for an understanding of the mecha- nism of formation of butyl alcohol, it was necessary to know not only the position of the heavy carbon in the carbon chain but also the concentration of heavy carbon in each position. Such information is essential in deciding whether two molecules of acetic acid unite, for example, as follows: 2CH3-~3COOH ~ CH3-~3CO-CH.,-~3COOH + H2O CH3-~3CO-CH.,-~3COOH + 8H-' CH3-~3CH.,-CHo-~3CH2OH + 2H2O or, whether the acetic acid units with an intermediate compound from the corn starch. In this latter case, probably only one position in the molecule would contain a concentration of heavy carbon in excess of the normal, since that portion of the mole- cule arising from the starch would have a normal concentration of i3C." In order to determine precisely the location of the heavy carbon atoms, it was necessary to develop a method of degrading butyric acid to achieve selective isolation of fragments of the carbon chain. This was accomplished by a modification of the hydrogen peroxide oxidation procedure of R. H. Allen and E. J. Witzemann (I. Am. Chem. Soc., 63: 1922~ 27, 1941~. The oxidation products, which included carbon dioxide, acetic acid, acetone, acetaldehyde, propionaldebyde, and an unidenti- fied non-volatile compound, were determined quantitatively and their ]3C contents were measured. The degradation of butyric acid and the determination of the location of i3C atoms in the oxidation products are excellent examples of the technical pre- cision characteristic of Werkman's laboratory. With the exception of the Booth-Green mill, the work of W. P. Wiggert, Milton Silverman, M. F. Utter, and Werkman (~`Iowa State Coll. I. Sci., 14: 179-86, 1940) was the first to show that extracts could be prepared from bacteria (as had been done with yeast and muscle) which dissimilate carbohydrates. The

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350 BIOGRAPHICAL MEMOIRS availability of these enzyme preparations permitted some of the first studies of glycolytic enzymes in bacteria, especially by Utter and Werkman (J. Bacteriol., 41 :5-, 1940; J. Bacteriol., 42:665-76, 1941; Biochem. J., 36:485-93, 1942~. These studies demonstrated that bacteria have fermentative pathways involving many of the same reactions as yeast and muscle. G. Kalnitsky and Werkman (J. Bacteriol., 44:256-57, 1942, and Arch. Biochem., 2: 113-24,1943) employed a cell-free enzyme preparation obtained by grinding a mass of Escherichia cold cells with powdered glass, with subsequent extraction using phosphate buffer (Wiggert and Werkman's method), in the anaerobic dissimilation of pyruvic acid; the result was the formation of acetic, formic, and succinic acids and carbon di- oxide, and with a trace of lactic acid. The enzyme preparation contained very active formic dehydrogenase and hydro~enase activity. i3CO~ was fixed in formic and succinic acids. The quantity of i3CO-, in formic acid suggested that it was formed from the pyruvic acid. The formation of succinic acid from pyruvate and carbon dioxide, with the ~3C in the carboxyl group, indicated the fixation of CO.' with the formation of a carbon-to-carbon linkage. During his extended career Werkman and his associates investigated a relatively wide range of the biochemical activities of bacteria. It is appropriate to mention briefly some of the other investigations which have not been treated more ex- tensively in this paper. Helen J. Weaver (1927) was perhaps Werkman's first graduate student; she was involved in the study of the bacteriological spoilage of canned vegetables. Shortly thereafter, Gertrude Sunderlin ~ 1928) studied the synthesis of vitamins by microorganisms. Sara Kendall (1931) made a systematic study of the propionic acid bacteria; G. Gillen (1932) studied the production of trimethylene glycol by bacteria; Roger Patrick (1933) studied the xylan-fermenting bacteria; C. A. Johnson and H. D. Coile (1933) devised an electron tube

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CHESTER HAMLIN WERKMAN 351 potentiometer for the determination of oxidation-reduction potentials; and Carl Erb (1936) participated in the design of both a multiple-cup micro- and multiple-cup macro-respirometer, which he employed in studying the aerobic dissimilation of lactic acid by the propionic acid bacteria and which were used in many subsequent investigations by Werkman and his stu- dents. Milton Silverman (1938, 1939) was among the first to demonstrate the role of vitamin Be and cocarboxylase in bac- terial metabolism; A. A. Andersen (1940) studied the growth factor and amino acid requirements of bacteria and described a dextro-lactic acid-forming organism of the genus Bacil lus; Milo N. Mickelson (1940) investigated the mechanism of the dissimilation of glycerol and the formation of trimethylene glycol by organisms related to the coli-aerogenes group of bac- teria; and M. E. Nelson ~ 1940) studied the dissimilation of levulose and other substrates in the lactic acid fermentation. Carl Brewer (1939, 1940) investigated the aerobic and anaerobic dissimilation of citric acid by the cold form bacteria, W. S. Waring (1944) the function of iron in microbial metabolism, David Paretsky (1947, 1950) the mechanism for the conversion of 2,3-butylene glycol to acetylmethylcarbinol in bacterial fer- mentation, and Noel Gross (1947) the isotopic composition of acetylmethylcarbinol produced by yeast juice from ~3C-labelecI acetaldehyde and pyruvate. A. G. C. White (1947) investigated the assimilation of acetate by yeast and the use of fatty acids in fat synthesis; Samuel Ail (1948, 1949) studied the mechanism of carbon dioxide replacement by dicarboxylic acids, which by . . . . . . am~nat~on, transam~nat~on, or similar reactions serve as sub- stitutes for carbon dioxide; and G. E. Wessman (1950) demon- strated the inhibition of carbon dioxide fixation by avidin. During the period of about 1950 until his death in 1962, it is not unlikely that Werkman found it somewhat difficult to maintain the cohesiveness of his research program and the momentum which his laboratory had experienced in earlier

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352 BIOGRAPHICAL MEMOIRS years. His energy and scientific drive were curtailed measurably by a chronic illness which worsened progressively, but never- theless he was active with his students and in the affairs of the Department of Bacteriology almost to the very end. It is evident from the publications coming from his laboratory that he at- tempted to continue work on carbon dioxide fixation as well as to develop some new directions. Examples of the research of his students during this period are as follows: dismutative assimilation of carbon dioxide (Dean Watt, 1950-1954), bac- terial synthesis of purines (W. B. Sutton, 1951-1953), bacterial synthesis of amino acids (Eric Fowler, 1952) bacterial metab- olism of amino acids (Mitchell Korzenovsky, 1953), mecha- nism of aerobic dissimilation of glucose (C. A. Claridge, 1954), formation of adenosine by cell-free bacterial extracts ~ John Ott, 1954), the role of transamination in bacterial metabolism (D. H. Hug, 1958), chemoautotrophic fixation of carbon dioxide by bacteria of the genus Mycobacterium (T. Myoda, 1960>, carbon dioxide fixation by heterotrophic and photosynthetic bacteria (D. S. Bates and C. L. Baugh, 1960), and fatty acid carboxylation by cell-free bacterial extracts (G. W. Claus, 1961- 1962). Seldom does an individual scientist stand alone in his con- tributions to the body of knowledge; more often than not his reputation and recognition result from the force of his leader- ship compounded with the efforts of his graduate students and his younger associates in research, and this was certainly true in regard to C. H. Werkman. In any attempt to review his accomplishments as a distinguished American scientist it would be impossible to dissociate his individual work from that of the numerous younger scientists who were associated with him over a period of several decades. This fact is recognized here, and it should be emphasized, moreover, that in a memoir of this nature it is not possible to record specifically the extensive basic contributions to the reputation of the Werkman labora- tory which were made by his many students. The impressive

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CHESTER HAMLIN WERKMAN 353 bibliography attached to this paper must be viewed as the com- posite contribution of Werkman and his associates, and the research reports cited specifically should serve only to illustrate some of the directions of the research effort stimulated by Werk- man as the leader of his laboratory. Werkman was a member of the faculty of Iowa State Uni- versity in the Department of Bacteriology continuously from 1925 until his death in 1962. He served as Assistant Professor, 1925-1927, and Associate Professor, 1927-1933. He attained the rank of Professor in 1933, became department head in 1945, and continued in this capacity until his death. He served as major professor for more than fifty graduate students of whom . ~ . ~ J ~ thlrty-slx received the Doctor of Philosophy decree; he was author and co-author of at least 275 publications in scientific journals. For various periods he served as an editor of the fol- lowing scientific journals: A rch ices of Biochemistry, A dvances in Enzymology, Proceedings of the Society for Experimental Biology and Medicine. Enz~mologia (assistant editor), Biotek ~_ ~ 7 . ' ru0~catzons Assistant ect~tor', and Iowa State College Journal of Science. In 1944 Werkman received the degree of Doctor of Science honoris cause from Purdue University, and in 1951 he received the Pasteur Award. Dr. Werkman was elected to the National Academy of Sci- ences in 1946; he also held membership in the following or- ganizations: American Society for Microbiology, American Chemical Society, Society of American Biological Chemists, Biochemical Society of Great Britain, American Association for the Advancement of Science (Fellow), Society of Experimental Biology and Medicine, Iowa Academy of Science, Society of the Sigma Xi, Phi Kappa Phi, Phi Lambda Upsilon, and Kappa Delta Pi. In 1958 he was the recipient of the Iowa State Uni- versity Faculty Citation. He served as a member of the Board of Trustees of the Carver Research Foundation of Tuskegee Institute.

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354 KEY TO ABBRE VIA TIONS BIOGRAPHICAL MEMOIRS BIBLIOGRAPHY 9^ntonie van Leeuwenhoek l. Microbiol. Serol. Antonie van Leeuwenhoek journal of Microbiology and Serology Arch. Biochem.Archives of Biochemistry Arch. Biochem. Biophys. _ Archives of Biochemistry and Biophysics Biochem. I.- Biochemical Journal Ind. Eng. Chem. - Industrial and Engineering Chemistry Ind. Eng. Chem., Anal. Ed. Industrial and Engineering Chemistry, An- alytical Edition Iowa Agr. Exp. Sta. Res. Bull. Iowa Agricultural Experiment Station Research Bulletin Iowa State Coll. J. Sci. _ Iowa State College journal of Science I. Agr. Res. journal of Agricultural Research I. Am. Chem. Soc. Journal of the American Chemical Society I. Bacterial. = Journal of Bacteriology i. Biol. Chem. Journal of Biological Chemistry I. Infect. Diseasesjournal of Infectious Diseases Proc. Iowa Acad. Sci. _ Proceedings of the Iowa Academy of Science Proc. Nat. Aced. Sci. Proceedings of the National Academy of Sciences Proc. Soc. Exp. Biol. Med. - Proceedings of the Society for Experimental Biology and Medicine 1921 With William M. Gibbs. Tuberculosis of poultry. tural Experiment Station Bulletin 126. Idaho Agricul- 1922 With W. M. Gibbs. Effect of tree products on bacteriological ac- tivities in soil. I. Ammonification and Vitrification. Soil Sci- ence, 13:303-22. 1923 Immunologic significance of vitamins. I. Influence of the lack of vitamins on the production of specific agglutinins, precipitins, hemolysins and bacteriolysins in the rat, rabbit and pigeon. J. Infect. Diseases, 32:247-54. Immunologic significance of vitamins. . . . ~ II. Influence of lack of vitamins on resistance of rat, rabbit and pigeon to bacterial infection. J. Infect. Diseases, 32:255-62. Immunologic significance of vitamins. III. Influence of the lack of vitamins on the leukocytes and on phagocytosis. I. Infect. Diseases, 32:263-69.

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CHESTER HAMLIN WERKMAN 355 With Max Levine. Bacterium cold and Bacterium aerogenes in swimming pools. Journal of the American Water Works Asso- ciation, 10: 620-22. 1924 Theory of dye utilization in bacterial media. 97. i. Bacteriol., 8:295- With V. E. Nelson and E. I. Fulmer. Immunologic significance of vitamins. IV. Influence of lack of vitamin C on resistance of the guinea-pig to bacterial infection, on production of specific ag- glutinins, and an opsonic activity. J. Infect. Diseases, 34:447-53. With F. M. Baldwin and V. E. Nelson. Immunologic significance of vitamins. V. Resistance of the avitamic albino rat to diphtheria toxin; production of antitoxin and blood pressure effects. I. Infect. Diseases, 35: 549-56. 1925 Continuous reproduction of microorganisms. 1926 Science, 62: 115-16. The role of accessory food factors in the physiology of microorga- nisms. I. Bacteriol., 11:86-87. 1927 Microbiological death rates. Proc. Iowa Acad. Sci., 34:85-87. Vitamin effects in the physiology of microorganisms. J. Bacteriol., 14:335-47. With H. J. Weaver. Studies in the bacteriology of sulphur stinker spoilage of canned sweet corn. Iowa State Coll. i. Sci., 2:57-67. With Helen Weaver. Bacterial blackening of canned vegetables. Proc. Iowa Acad. Sci., 34:92-93. 1928 Factors influencing the death time of microorganisms. Proc. Iowa Acad. Sci., 35:97. With Gertrude Sunderlin. Synthesis of vitamin B by microorga- nisms. J. Bacteriol., 16: 17-33. 1929 Bacteriological studies on sulfid spoilage of canned vegetables. Iowa Agr. Exp. Sta. Res. Bull. 117.

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356 BIOGRAPHICAL MEMOIRS 1930 An improved technic for the Voges-Proskauer test. I. Bacteriol., 20: 121-25. Determination of organic acids in mixtures. I. Determination of fatty acids in mixtures by partition between isopropyl ether and water. Ind. Eng. Chem., 2:302-4. Determination of organic acids. II. Determination of mixtures of two fatty acids by partition between ethyl ether and water. Iowa State Coll. I. Sci., 4:459-64. Determination of organic acids. III. Note on the use of the isoamyl ether-water system in the partition method. Iowa State Coll. l. Sci., 5:1-3. With R. H. Carter. Factors influencing the production of acetic acid from corn stalks by thermophilic bacteria. Proc. Iowa Acad. Sci., 37:51-52. Dimethyl-alpha-naphthylamine for the determination of bacterial reduction of nitrates. Proc. Iowa Acad. Sci., 37:53-55. With E. I. Fulmer. An Index to the Chemical Action of Micro- organisms on the Non-Nitrogenous Organic Compounds. Spring- field, Charles C. Thomas, Publisher. xiii + 198 pp. With O. L. Osburn. Determination of butyl and ethyl alcohols in fermentation mixtures. Proc. Soc. Exp. Biol. Med., 28:241-42. With Roger Patrick. Notes on the bacterial flora of the snake. Proc. Iowa Acad. Sci., 37: 57-58. 1931 Determination of organic acids. IV. A method for the provisional identification and quantitative determination of two fatty acids in a mixture. Iowa State Coll. i. Sci., 5: 121-25. With O. L. Osburn. A method for the determination of ethyl and butyl alcohols in fermentation mixtures. i. Bacteriol., 21: 20-21. With E. I. Fulmer and A. L. Williams. The effect of sterilization of media upon their growth promoting properties toward bac- teria. J. Bacteriol., 21: 299-303. With E. I. Fulmer and A. L. Williams. Production of bacterial growth stimulants by heating the medium under pressure. Proc. Soc. Exp. Biol. Med., 28:462. With Sara Kendall. The propionic acid bacteria. I. Classification and nomenclature. Iowa State Coll. i. Sci., 6:17-32.

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CHESTER HAMLIN WERKMAN 357 Title O. L. Osburn. ~ Determination of organic acids. V. Applica- tion of partition method to quantitative determination of acetic, ~ ~ ~ ~ ~ ~ ~ Ind. Eng. Chem., Anal. propionic and butyric acids in mixture. Ed., 3:264. With O. L. Osburn. Determination of butyl and ethyl alcohols in mixtures. Ind. Eng. Chem., 3:387-89. 1932 With Charles Davis. Apparatus for continuous fractional vacuum distillation. Cl~emist-Analyst, 2 1: 20-2 1. With G. F. Gillen. Bacteria producing trimethylene glycol. T Bacteriol., 23: 167-82. Title O. L. Osburn. Ind. Eng. Chem., 4: 1-6. With Roger Patrick. ~ man. Proc. Iowa Acad. Sci., 39:49-51. Determination of carbon in fermented liquors. A new species of Actinomyces pathogenic in With O. L. Osburn. Comparative dissimilation of xylose and glucose by Escherichia cold and Citrobacter anindolcium. Proc. Iowa Acad. Sci., 39:134-35. NVith O. L. Osburn and H. G. Wood. Determination of formic, acetic and propionic acids in fermenting mixtures. Proc. Soc. Exp. Biol. NIed., 29:294-95. 1933 With M. C. Brockmann. Determination of 2,3-butylene glycol in fermentations. Ind. Eng. Chem., 5: 206-7. With M. C. Brockmann. Oxidation-reduction studies on the 2,3- butylene glycol-acetylmethylcarbinol system in a fermentation. Proc. Soc. Exp. Biol. Med., 30:1146-48. With C. S. McCleskey and Howard Reynolds. Physiology of Shigella paradysenteriae var. Sonnei. Proc. Soc. Exp. Biol. Med., 30: 1050-51. With C. W. Davis and C. A. Tarnutzer. Notes on the trimethylene glycol fermentation. l. Bacteriol., 24:33. \Vith R. W. Brown. The propionic acid l~acteria. II. Classification. J. Bacteriol., 26: 393-417. With Roger Patrick. Bacterial fermenting xylan. Iowa State Coll. J. Sci., 4:407-18. NAlith H. Reynolds and O. L. Osburn. Determination of the fur- fural yielding constituents of plant materials. Iowa State Coll. J. Sci., 4:443-51.