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CORNELIS BERNARDUS VAN NIEL November 4, I 897March l O. 1 985 BY H. A. BARKER AND ROBERT E. HUNGATE CORNELIS BERNARDUS VAN NIEL Kees to his friends and studentsis best known for his discovery of mul- tiple types of bacterial photosynthesis, his decluction that all types of photosynthesis involve the same photochemical mechanism, and his extraordinary ability to transmit his en- thusiasm for the study of microorganisms to his students. His interest in purple ant! green bacteria developed in his first year as a graduate student. After thoughtful analysis of the confusing literature clearing with these bacteria, he carried out a few simple experiments on their growth requirements. Interpreting the results in accordance with the theories of his professor, A. J. Kluyver, on the role of hydrogen transfer in metabolism, he developer! a revolutionary concept of the chemistry of photosynthesis that was to influence research on the topic for many years. As a teacher he was unsurpassed. Although he taught in a small, somewhat remote institution with modest facilities, the force of his personality, his eloquence and scholarship made the Hopkins Marine Station a mecca for students of general microbiology throughout the western world. EDUCATION AND EARLY LIFE Van Nie} was born in Haarlem, The Netherlands, into a family steeped in a highly conservative Calvinist tradition. 389

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390 BIOGRAPHICAL MEMOIRS His father and several uncles were businessmen and did not have a professional education. His father sties! when he was seven years old, ant! thereafter his mother largely depended on his uncles for acivice in educating her young son. Since family tradition decreer! that a son shouIc! succeed to his fa- ther's business, Kees was sent to a secondary school with a curriculum clesignecl to prepare students for a commercial career. At the end of his third year in high school when he was fifteen years old, an event occurred that changed the course of his education. The family was spending their summer va- cation as guests of a friend on a large estate in northern Holland clevotec! to various agricultural activities. A part was set aside for testing the effectiveness of various soil treat- ments on crop production, and van Nie] has described how his host introclucec! him to the methods of agricultural re- search and how impressed he was to learn that "one could raise a question and obtain a more or less definitive answer to it as a result of an experiment . . . particularly because ~ hac! grown up in a milieu where any kind of question was invariably answered by the stereotyped reply: 'Because some- bocly (usually a member of the family) said so"' (1967, I, p. 2~. Van Niel's interest ant! enthusiasm for these activities lect his family to reevaluate his education, and he was finally al- lowed to transfer to a college preparatory high school. Uncler the influence of one of his teachers in the new school he developed a strong interest in chemistry. He liked analytical chemistry so much that he set up a small laboratory at home and analyzed samples of fertilizer in his spare time. His aca- clemic record in high school was sufficient to obtain acimis- sion to the Chemistry Division of the Technical University in Delft on graduation without taking the usual entrance ex- amlnatlon. He entered the University in autumn 1916 but, after only

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CORNELIS BERNARDUS VAN NIEL 391 three months, was incluctect into the Dutch army, in which he server! until the end of December 1918. Life in the army was both a traumatic and a highly educational experience. Removed from the protective environment of his family for the first time, he was exposed to the rough and impersonal life of military training. He later wrote that up to this time he tract been "utterly unaware of the many problems to which man is exposed and with which he must learn to cope." For- tunately, he receiver! practical and intellectual support from a former high school classmate inclucted at the same time, Jacques de Kacit. After a few clays in a primitive military camp on the out- skirts of Amersfoort, Jacques proposer! that they rent a room in the city where they couIct spend their free time in greater comfort. They were soon joined by a friend of Jacques, ant! the three comrades spent their leisure hours discussing many subjects. Jacques was an intellectual with a cosmopolitan background. He introduced van Nie} to new worIcis of liter- ature, art, ant! philosophy. Under his influence, van Nie! read many of the works of Zola, Anatole France, Ibsen, Strinct- berg, Shaw, and Nietzsche. Their ideas frequently conflicted with van Niel's Calvinist background and lee! to what he later described as the rebellious phase of his life. On returning to the University after army service, van Nie! was undecided! whether he should continue the stucly of chemistry or take up the study of literature. But, discussing the alternatives with an aunt whose judgment he trusted "at least in part because of her unconventional attitudes and be- havior," he was finally persuaded to continue on in chemistry. Still, his mental turmoil was such that he could not immecti- ately switch back into the normal academic routine. He spent the first six months reacting French, English, Scandinavian, and Russian 19th century literature ant! was not prepared to take the first year chemistry examination in June 1919.

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392 BIOGRAPHICAL MEMOIRS In the autumn, however, he finally settled down to serious study and by intensive effort was able in June 1920 to pass both the first- ant! second-year chemistry examinations. Dur- ing the following year, van Nie! took several courses in biol- ogy in addition to the prescribed chemistry program, includ- ing G. van Iterson's courses in genetics anct plant anatomy ant! chemistry ant} M. W. Beijerinck's courses in general ant! applied microbiology. By November 1921, van Nie! had completed all the re- quirements for the chemical engineering degree except a year of work in a specialized area of his own choosing. Al- ready strongly attracted to microbiology from his exposure to Beijerinck's courses, he decided to specialize in it after hearing the inaugural lecture of A. I. Kluyver, who succeeded Beijerinck that year. Kluyver suggested that van Nie] investigate the longevity of yeast in a medium containing sugar but little or no nitro- gen. This problem provided some experience with microbi- ological and analytical methods and met the requirements for the degree, though the results were unimpressive. As a sidle project, van Nie! checked a published report that a nonmotile Sarcina conic! develop flagella and motility by repeated transfer in a special medium. His first publication (van Nie] 1923) showed that the previous author had con- fused Brownian movement with true motility and that his so- callecl flagella were artifacts of the staining method. DELFT: WORKING WITH KLUYVER After receiving his Chem. E. degree van Niel accepted a position as assistant to Kluyver. His duties consisted! of caring for a large, pure culture collection of bacteria, yeasts and fungi; assisting undergraduates; and preparing demonstra- tions for Kluyver's two lecture courses. One of the courses dealt with the microbiology of water ant! sewage in which

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CORNELIS BERNARDUS VAN NIEL 393 iron ant! sulfur bacteria play a role. Since Kluyver was un- familiar with these organisms, he assigned van Nie! the task of learning to culture them so that he could provide material for class demonstrations. To fulfill this assignment, van Nie! read the publications of Winogradsky, Engelmann, Molisch, and Bavenclamm on the colorless and purple sulfur bacteria and concluded that fundamental disagreements concerning the metabolism of these organisms neecled clarification. Finding the purple bacteria "aesthetically pleasing," he con- tinuec! studying them after the lecture demonstrations were completed. During the next two years, while continuing as Kluyver's assistant, and later as conservator of the Institute, van Nie! clemonstrated that purple sulfur bacteria could grow in glass- stopperec! bottles completely filled with a mineral medium containing sulfide and bicarbonate that were exposed to day- light. (No growth occurred in the dark.) He also isolated pure cultures of a Chromatium species and Thiosarcina rosea ant! shower! that the yield of cells was proportional to the amount of sulfide provided and much greater than that of colorless aerobic sulfur bacteria in a similar medium. These observations and the earlier demonstration that O2 is not proclucect by purple bacteria were interpreted (in ac- corciance with Kluyver's theory that most metabolic reactions are transfers of hydrogen between donor and acceptor mol- ecules) to mean that purple sulfur bacteria carry out a novel type of photosynthesis in which carbon dioxide is reclucec! by hydrogen derived from hydrogen sulfide with the air! of en- ergy from light. Mentioner! briefly in Kluyver and Donker's treatise, "The Unity in Biochemistry,"i without supporting evidences this interpretation was probably based on van Niel's ' A. J. Kluyver and H. J. L Donker, "Die Einheit in der Biochemie," Chemie der Zelle und Gewebe, 13(1926): 134-90.

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394 BIOGRAPHICAL MEMOIRS work. Kluyver was not a coauthor of any of van Niel's early papers on photosynthetic bacteria. During this period Kluyver and van Nie! published two papers: one clearing with a new type of yeast, Sporobolomyces (thought on the basis of its mode of spore formation to be a primitive basicliomycete), and another providing an expla- nation for the unusual morphology of a spore-forming bac- terium that grew in liquid! media as a tightly twisted, multi- strancled rope. While van Nie! expected to continue his study of purple bacteria for his Ph.D. dissertation, he also developed, as a side project, an effective methoc! for isolating propionic acid bacteria from Swiss cheese. When Kluyver pointed out that a study of this group would provide a faster path to the ctoc- torate than a completion of his investigations of the sIow- growing purple bacteria, van Nie! reluctantly agreed. He spent the next two years, therefore, studying the biochemis- try ant! taxonomy of the propionic acid bacteria. These bio- chemical studies were the first to provide a quantitative pic- ture of the products derived from the fermentations of lactate, glycerol, glucose, and starch. His taxonomic studies provided a sound basis for recognition of the species of Propionibacteraum. Van Niel's dissertation, written in English, was publisher! in 1928. An unexpected byproduct of the study of the propionic acid bacteria was the identification of ctiacety! as the com- pounc] responsible for the characteristic aroma of high qual- ity butter. Van Nie! noticed that cultures of one of his pro- pionic acid bacteria grown on a glucose medium smeller! like butter, then correlates} this oclor with the distinctive ability of the organism to produce acety~methy~carbinol, an odorless compound that is reaclily oxidizer! to diacety1, the actual source of the aroma. Van Nie} spent almost seven years in the Delft laboratory,

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CORNELIS BERNARDUS VAN NIEL 395 a stimulating period cluring which Kluyver was developing his ideas about the importance of hydrogen-transfer reac- tions in metabolism and the similarity of basic biochemical reactions in clifferent organisms (the "unity in biochemistry" theory). Van Nie] consiclerec! these ideas to be among the most fundamental ant} fruitful of that era. Revering Kluyver (whom he always referrer! to as "the Master"), as one of the great scientists of the age, he was yet able at a later time to point out some of Kluyver's errors in the analysis of specific phenomena and his occasional excessive reliance on gener- alizations lacking adequate experimental support (1959,1~. PACIFIC GROVE: HOPKINS MARINE STATION In late 1927, L. G. M. Baas-Becking of Stanford Univer- sity came to Delft looking for a microbiologist to fill a position at the new Jacques Loeb Laboratory at the Hopkins Marine Station on the Monterey Peninsula. Greatly impressed by van Niel's research accomplishments and his capacity for lucid communication, he offered him an appointment as associate professor. Put off by the reputed materialism of American society, van Nie! was yet attracted by Becking's enthusiasm for the new laboratory and encouraged by Kluyver de- ciclec! to strike out on his own. He arriver! in California at the ens] of December ~ 928 and was immediately impressed by the charm of Carmel, the beautiful site of the Jacques Loeb Laboratory, and the free- dom from outside pressures that the Marine Station pro- vided. In later years he could never be persua(lec! to leave even to succeed Kluyver at the Delft laboratory. PHOTOSYNTHESIS STUDIES At the Hopkins Marine Station van Nie] continued his studies of purple and green bacteria with emphasis on the quantitative relations among substrates consumed and procl-

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396 BIOGRAPHICAL MEMOIRS ucts formed. Progress was accelerates! by the finding that the bacteria grew more rapidly under continuous artificial illu- mination. He clemonstratec! that the green bacteria oxidized hydrogen sulfide only as far as sulfur, whereas the purple sulfur bacteria further oxiclized the sulfur to sulfate. Both coupled these oxidations with an essentially stoichiometric conversion of carbon dioxide to cellular materials in light- clependent reactions. The nonsulfur bacteria (Athiorhoda- ceace, which Molisch hac! grown aerobically on various or- ganic compounds) were shown to develop anaerobically, but only in the presence of carbon dioxicle and light. These ant! other observations led van Nie! to conclude that photosyn- thesis is essentially a light-dependent reaction in which hy- drogen from a suitable oxidizable compound! reduces carbon dioxide to cellular materials having the approximate com- position of carbohydrate. This was expressed by the gener- alized equation: THEA + CO2 g > 2A + (CH2O) + H2O. According to this formulation, H2O is the hydrogen donor in green plant photosynthesis and is oxi(lizecl to O2, whereas H2S or another oxidizable sulfur compound is the hydrogen donor for purple and green sulfur bacteria, and the oxida- tion product is sulfur or sulfate, depencling on the organism. The nonsulfur purple bacteria that require suitable organic compounds in addition to carbon clioxide for anaerobic growth in light were presumed to use these compounds as hydrogen donors ant! to oxidize them either partially or completely. Later, the purple sulfur bacteria were also shown to use some organic compounds in place of H2S in their pho- tometabolism. These observations ant! interpretations, the results of some six years of investigation, were first presented at a small meeting of the Western Society of Naturalists in Pacific Grove

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CORNELIS BERNARDUS VAN NIEL 397 at the end of 1929. Two years later van Nie! published a de- tailecl account of the culture, morphology and physiology of purple and green suIphur bacteria (193l,1), bringing his in- terpretation of their metabolism and its implications for green-plant photosynthesis to the attention of a wider aucli- ence. All of the purple sulfur bacteria he isolated were relatively small organisms, belonging to what he called Chromatium, Thiocystis, and Pseudomonas types. In material collected in nature (and in some enrichment cultures) he observed a number of larger forms but, despite numerous attempts, was unsuccessful in isolating them. The cultivation of these or- ganisms was not accomplished until many years later, when N. Pfennig and H. G. SchIegel, both onetime associates of van Niel, discovered that nutritional and environmental re- quirements are more complex than had been previously rec- ognizecl.2 Van Nie} published a large monograph covering many years of work on the culture, general physiology, morphology and classification of the nonsulfur purple and brown bacteria in 1944 (1944,21. He classified over 150 strains isolated from natural sources into six species in two genera Rhodospeudo- monas and RhodospiraZlum. He clescribec! the morphology of the organisms, their pigments, nutritional requirements, and metabolism in the presence and absence of light. As in all his publications, van Nie] also reviewed the historical back- grounc! and current literature of the subject critically and thoroughly. Following the recognition of several types of photosyn- thesis using different hydrogen donors, van Nie! began to 2 H. G. Schlegel and N. Pfennig, "Die Anreicherungskultur einiger Schwefelpur- purbakterien." Archiv fur Mibrobiologie, 38(1961): 1-39, and N. Pfennig and K. D. Lippert, "Uber das Vitamin Be Bedurfnisphototropher Schwefelbakterien." Archiv fur Mibrobiologie, 55(1966):245-56.

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398 BIOGRAPHICAL MEMOIRS consider how radiant energy participates in these reactions. There appeared! to be two possibilities, both consiclered by earlier investigators: radiant energy could be used to activate either carbon dioxicle or the hydrogen donor. Initially, van Nie} and Muller (1931,2) were incliner! to believe that light is used primarily to activate carbon dioxide, a relatively stable compound and the common reactant in all photosynthetic systems. But they die! not exclude the seconc! possibility, that light also activated the hydrogen donor. In this connection they notes! a correlation between the pres- ence of nonchIorophyI] yellow ant! red pigments ant! the na- ture of the hydrogen (loner used by different organisms. These pigments, lacking in the green sulfur bacteria that uti- lize the easily oxiclizable hydrogen sulfide, occur exclusively . . . - ~n organisms up Zing water or sulfur, then thought to re- quire a greater activation. This led van Nie! to undertake a series of studies of the pigments of the purple and green bacteria. Van Nie} and Arnold (193S,1) developed a convenient spectrophotometric method for determining the amount of bacteriochIorophyI] in photosynthetic purple and brown bac- teria uncler conditions avoiding interference by the rec! car- otinoi(1 pigments. They also reported that van Nie! and E. Wieclemann, working in A. Stoll's laboratory, had examined the green pigments of six clifferent strains of purple and brown bacteria ant! concluded that they were identical with the chlorophyll of the purple sulfur bacterium, Thiocystis, previously stucliect by H. Fischer. Van Nie] and Smith (1935,2) began a study of the chem- istry of the major red pigment of the nonsulfur purple bac- terium, Rhodospirillum rubrum. By solvent extraction and re- peated crystallization, they isolated about 100 milligrams of an apparently homogeneous carotinoid they called "spirillo- xanthin." Its empirical composition was found to be

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CORNELIS BERNARDUS VAN NIEL 413 During the early years, only a few students attended, but as van Niel's reputation as a teacher spread, the class hack to be limited, initially to eight, and later to fourteen students- the number that conic! be accommociated in the small Marine Station laboratory. The students were initially undergraduate or graduate students from Stanford, but later a large pro- portion came from other institutions. In 1950, for example, only one of the thirteen students was from Stanford. The others were from Washington University, Wisconsin, Michi- gan, Missouri, California Institute of Technology, Connecti- cut, Illinois, Cambridge, and the University of California at Los Angeles. In addition there were eleven auditors of the discussions and lectures who clid not do the experiments- mostly postdoctoral fellows or established scientists who wished to extend their background in general microbiology. The lists of students and auditors who attended van Niel's course between 1938 and 1962 reads like a Who's Who of bio- logical scientists in the United States, with several, as well, from other countries. Both clirectly, and indirectly through his students, van Nie! exerted a powerful influence on teach- ing and research in general microbiology for a generation. Although his own research was concerned mainly with photosynthetic bacteria, van Nie! was interested in the biol- ogy and metabolism of many other groups of microorga- nisms. He clicl not believe in directing the research of his younger associates but rather encouraged them to follow their own interests, some of which hac! been stimulated by his lectures and personal discussions. As a consequence, the range of phenomena investigated in his laboratory was ex- ceedingly wide and included the culture and physiology of blue-green algae and diatoms, nutritional and taxonomic studies of plant-pathogenic bacteria, biological methane for- mation, pteridine and carbohydrate metabolism of protozoa, germination of mold spores, biology of caulobacteria, culti- vation of free-living spirochetes, induction of fruiting bodies

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414 BIOGRAPHICAL MEMOIRS in myxobacteria, decomposition of cellulose, the role of mi- croorganisms in the foot! cycle of aquatic environments, ad- aptation of bacteria to high salt concentrations, cultivation of spirilIa anct colorless sulfur bacteria, bacterial fermentations, thermophylic bacteria, denitrification, pyrimidine metabo- lism, and the thermodynamics of living systems. To all stu- dents van Nie} gave freely of his time, advice and enthusiasm, drawing on his own extraordinary knowledge of the litera- ture. RETIREMENT Following his retirement from the Marine Station in 1962, van Nie! held a visiting professorship at the University of California at Santa Cruz from 1964 to 1968, teaching part of a freshman-level biology course in collaboration with K. V. Thimann ant! L. Blinks. After 1972, van Nie! gave up teaching and research en- tirely and clisposec! of his scientific library ant! large collection of reprints. Thereafter he lived quietly with his wife, Mimi, in Carme! ant! spent his leisure reading classical and modern literature and listening to classical music, which he greatly enjoyed. He was often visited by former students who con- tinued to be impressed by the warm hospitality of his home, the charm of his personality, the breadth of his understancl- ing, ant! the comprehensiveness of his memory.

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CORNELIS BERNARDUS VAN NIEL 415 HONORS AND DISTINCTIONS DEGREES AND HONORARY DEGREES 1923 Chemical Engineering, Technical University, Delft 1928 D.Sci., Technical University, Delft 1946 D.Sci. (Honorary), Princeton University 1954 D.Sci. (Honorary), Rutgers University 1968 LL.D., University of California, Davis FELLOWSHIPS AND PROFESSIONAL APPOINTMENTS 1925-1928 Conservator, Laboratorium voor Microbiologie, Delft 1928-1935 Associate Professor of Microbiology, Stanford Uni- versity, Hopkins Marine Station 1935 -1936 Rockefeller Foundation Fellow 1935-1946 Professor of Microbiology, Stanford University 1945 John Simon Guggenheim Fellow 1946-1963 Herstein Professor of Biology, Stanford University 1955-1956 John Simon Guggenheim Fellow 1963-1985 Herstein Professor, Emeritus, Stanford University 1964-1968 Visiting Professor, University of California, Santa Cruz AWARDS AND HONORS 1942 Stephen Hales Prize, American Society of Plant Physiology 1964 Emil Christian Hansen Medalist, Carlsberg Foundation of Copenhagen 1964 National Medal of Science 1966 Charles F. Kettering Award, American Society of Plant Physiology 1967 Rumford Medal, American Society of Arts and Sciences 1967 Honorary Volume, Archiv fur Mikrobiologie 1970 Antonie van Leeuwenhoek Medal, Royal Netherlands Academy of Sciences LEARNED SOCIETIES 1945 National Academy of Sciences 1948 American Philosophical Society 1950 American Academy of Arts and Sciences 1952 Charles Reid Barnes Life Membership, American Society of Plant Physiology

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416 BIOGRAPHICAL MEMOIRS 1954 President, American Society for Microbiology 1954 Corresponding Member, Academy of Sciences, Gottingen, Germany 1958 American Academy of Microbiology 1963 Honorary Member, Societe Franchise de Microbiologie 1967 Honorary Member, Society of General Microbiology 1968 Honorary Member, Royal Danish Academy of Sciences and Letters

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CORNELIS BERNARDUS VAN NIEL 417 SELECTED BIBLIOGRAPHY 1923 Uber die Beweglichkeit und das Vorkommen von Geisseln bei einigen Sarcina Arten. Zentralbl. Bakteriol. Parasietenkd. In- fektionskr. Hyg., Abt. II., 60:289-98. 1924 With A. J. Kluyver. Uber Spiegelbilder erzeugende Hefearten und die neue Hefegattung Sporobolomyces. Zentralbl. Bakteriol. Par- asitenkd. Infektionskr. Hyg., Abt. II., 63:1-20. 1925 With F. Visser't Hooft. Die fehlerhafte Anwendung biologischer Agenzien in der organischen Chemie. Eine Warnung. Ber. Dtsch. Chem. Ges., 58: 1606-10. 1926 With A. }. Kluyver. Uber Bacillus funicularis n.sp. nebst einigen Be- merkungen uber Gallionella ferrug~nea Ehrenberg. Planta, 2:507-26. 1927 With A. ]. Kluyver. Sporoboloymces ein Basidiomyzet? Ann. Mycol. Notitiam Sci. Mycol. Univ., 25:389-94. Notiz uber die quantitativ Bestimmung von Diacetyl und Acetyl- methylcarbinol. Biochem. Z., 187:472-78. 1928 The Propionic Acid Bacterza. (Doctoral Dissertation.) Haarlem, The Netherlands: Uitgeverszaak]. W. Boissevain & Co. 1929 With A. J. Kluyver and H. G. Derx. De bacterien der roomverzur- ing en het boteraroma. Verslag gewone Vergader. Afd. Na- turrkd. Nederl. Akad. Wetensch., 38:61-2. With A. J. Kluyver and H. G. Derx. Uber das Butteraroma. Bio- chem. Z., 210:234-51.

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418 BIOGRAPHICAL MEMOIRS 1930 Photosynthesis of bacteria. In: Contributions to Marine Biology, Stan- ford: Stanford University Press, pp. 161-69. 1931 On the morphology and physiology of the purple and green sulfur bacteria. Arch. Mikrobiol., 3:1-112. With F. M. Muller. On the purple bacteria and their significance , ~ for the study of photosynthesis. Rec. Trav. Bot. Neer., 28:245- 74. 1935 Photosynthesis of bacteria. Cold Spring Harbor Symp. Quant. Biol., 3: 138-50. With J. A. C. Smith. Studies on the pigments of the purple bacteria. I. On spirilloxanthin, a component of the pigment complex of Spirillum rubrum. Arch. Mikrobiol., 6:219-29. A note on the apparent absence of Azotobacter in soils. Arch. Mi- krobiol., 6:215 -18. 1936 On the metabolism of the Thiorhodaceae. Arch. Mikrobiol., 7:323-58. With D. Spence. Bacterial decomposition of the rubber in Hevea latex. Ind. Eng. Chem., 28:847-50. Les photosyntheses bacteriennes. Bull. Assoc. Diplomes Microbiol. Fac. Pharm. Nancy, 13:3-18. With A. J. Kluyver. Prospects for a natural system of classification of bacteria. Zentralbl. Bakteriol. Parasitenkd. Infektionskr. Hyg. Abt. II, 94:369-403. 1937 The biochemistry of bacteria. Ann. Rev. Biochem., 6:595-615. 1938 With W. Arnold. The quantitative estimation of bacteriochloro- phyll. Enzymologia, 5:244-50.

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CORNELIS BERNARDUS VAN NIEL 419 1939 A. l. Kluyver. Als mikrobioloog en als biochemikus. Chem. Weekbl., 36: 1-109. 1940 The biochemistry of microorganisms: An approach to general and comparative biochemistry. Am. Assoc. Adv. Sci. Publ., 14:106- 19. 1941 With E. H. Anderson. On the occurrence of fermentative assimi- lation. I. Cell. Comp. Physiol., 17:49-56. The bacterial photosyntheses and their importance for the general problem of photosynthesis. Adv. Enzymol., 1 :263-328. With R. Y. Stanier. The main outlines of bacterial classification. I. Bacteriol., 42:437-66. 1942 With A. L. Cohen. On the metabolism of Candida albicans. ]. Cell. Comp. Physiol., 20:95-112. With S. Ruben, S. F. Carson, M. D. Kamen, and I. W. Foster. Ra- dioactive carbon as an indicator of carbon dioxide utilization. VIII. The role of carbon dioxide in cellular metabolism. Proc. Natl. Acad. Sci. USA, 28:8-15. With I. O. Thomas, S. Rubin, and M. D. Kamen. Radioactive car- bon as an indicator of carbon dioxide utilization. IX. The assim- ilation of carbon dioxide by protozoa. Proc. Natl. Acad. Sci. USA,28:157-61. 1943 Biochemistry of microorganisms. Ann. Rev. Biochem., 12:551-86. Biochemical problems of the chemo-autotrophic bacteria. Physiol. Rev., 23:338-54. 1944 With A. Polgar and L. Zechmeister. Studies on the pigments of the purple bacteria. II. A spectroscopic and stereochemical inves- tigation of Spirilloxanthin. Arch. Biochem., 5:243-64. The culture, general physiology, morphology, and classification of

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420 BIOGRAPHICAL MEMOIRS the nonsulfur purple and brown bacteria. Bacteriol. Rev., 8:1- 118. Recent advances in our knowledge of the physiology of microor- ganisms. Bacteriol. Rev., 8:225-34. 1946 The classification and natural relationships of bacteria. Cold Spring Harbor Symp. Quant. Biol., 11:285-301. 1947 Studies on the pigments of the purple bacteria. III. The yellow and red pigments of Rhodopseudomonas spheroides. Antonie van Leeu- wenhoek T. Microbiol., 12: 156 - 66. 1948 Propionibacterium, pp. 372-79; Rhodobacterineae, pp. 838-74; Beggiatoaceae, pp.988-96; Achromatiaceae, pp.997-1001. In: Bergey's Manual of Determinative Bacteriology, 6th ea., eds. R. S. Breed, E. G. D. Murray, and A. P. Hitchens, Baltimore: Williams and Wilkins Co. 1949 The kinetics of growth of microorganisms. In: The Chemistry and Physiology of Growth, ed. A. K. Parpart, Princeton: Princeton University Press, pp. 91-105. The comparative biochemistry of photosynthesis. In: Photosynthesis in Plants, eds. I. Franck and W. E. Loomis, Ames: Iowa State College Press, pp. 437-95. Comparative biochemistry of photosynthesis. Am. Sci., 37:371-83. The "Delft school" and the rise of general microbiology. Bacteriol. Rev., 13:161-74. 1952 With M. B. Allen. Experiments on bacterial denitrification. I. Bac- teriol., 64:397-412. Bacterial photosynthesis. In: The Enzymes, vol. 2, part 2, eds. I B. Sumner and K. Myrback, New York: Academic Press, pp. 1074-88. With H. Larsen and C. S. Yocum. On the energetics of the photo- syntheses in green sulfur bacteria. J. Gen. Physiol., 36:161-71.

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CORNELIS BERNARDUS VAN NIEL 421 With M. B. Allen. A note on Pseudomonas stutzeri. 64:413-22. 1953 I Bacterial., With M. B. Allen and B. E. Wright. On the photochemical reduc- tion of nitrate by algae. Biochim. Biophys. Acta, 12:67-74. Introductory remarks on the comparative biochemistry of micro- organisms. J. Cell. Comp. Physiol., 41(Suppl. 1~: 1 1-38. 1954 The chemoautotrophic and photosynthetic bacteria. Annul Rev. Microbiol., 8:105-32. 1955 Classification and taxonomy of the bacteria and bluegreen algae. In: A Century of Progress in the Natural Sciences 1853-1953, ed. E. L. Kessel, San Francisco: California Academy of Sciences, pp. 89-1 14. On radicalism and conservatism in science. Bacterial. Rev., 19: 1-5. < Natural selection in the microbial world. l. Gen. Microbiol.. 13:201-17. The microbe as a whole. In: Perspectives and Horizons in Microbiology, ed. S. A. Waksman, New Brunswick, N.~.: Rutgers University Press, pp. 3-12. 1956 With T. W. Goodwin and M. E. Sissins. Studies in carotenogenesis. 21. The nature of the changes in carotinoid synthesis in Rho- dospirillum rubrum during growth. Biochem. I., 63:408-12. Phototrophic bacteria: Key to the understanding of green plant photosynthesis, pp. 73-92; Trial and error in living organisms: Microbial mutations, pp. 130-54; Evolution as viewed by the microbiologist, pp. 155-76. In: The Microbe's Contribution to Biol- ogy. A. J. Kluyver and C. B. van Niel. Cambridge: Harvard Uni- versity Press. With G. Milhaud and l. P. Aubert. Etudes de la glycolyse de Zymo- sarcina ventriculi, Ann. Inst. Pasteur, 91:363-68. In memoriam: Professor Dr. Ir. A. I. Kluyver. Antonie van Leeu- wenhoek J. Microbiol. Serol., 22:209-17.

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