van Niel’s Course in General Microbiology His course offered an early survey of the microbial world as well as a coherent approach for integrating biology Susan Spath t’s the greatest course in the Technical University in Delft, the Netherlands. world,” said Germain (CohenThere, van Niel adopted the broadly ecological apBazire) Stanier. She was referring proach to microbiology developed by M. W. Beijerto C. B. van Niel’s course in geninck, the first professor of microbiology at Delft. In eral microbiology, given most the 1920s, A. J. Kluyver, the successor to Beijerinck, summers from 1930 –1962 at the Hopkins Maundertook a broad, comparative survey of microbial rine Station (HMS), Stanford University’s mametabolism from which he derived, with H. J. L. rine laboratory on Monterey Bay in California. Donker, the principle of the unity of biochemistry. For some students, taking van Niel’s course was This principle greatly impressed van Niel, and a life-changing experience. For most it was unthroughout his life he sought to cultivate an apforgettable. From the 1930s to the proach to microbiology now known 1950s, van Niel’s course played a as the Delft School, which integrated major role in developing general both Kluyver’s and Beijerinck’s perFrom the 1930s microbiology into a dynamic and pectives. to the 1950s, respected discipline. Graduates of In the 1920s, general microbiolvan Niel’s van Niel’s course include Nobel ogy was a small and fragmented course played a Prize winners Arthur Kornberg, enterprise, especially in the United Konrad Bloch, and Paul Berg as well States. Most scientists who were major role in as leading scientists working in nustudying microorganisms were endeveloping merous fields and on central probgaged in medical, industrial, and general lems in biology, including microbial agricultural branches of bacteriolmicrobiology ecology, photosynthesis research, ogy. Only a few laboratories had into a dynamic plant physiology, bacterial physiolanything like sustained programs and respected ogy, and molecular biology. in general microbiology. In other words, bacteriology was almost discipline van Niel Saw Microbiology as completely isolated from the main a Means for Unifying, branches of the life sciences. At Systematizing Biology HMS, van Niel was ideally situated to pursue microbiology as a general science of life. His When van Niel arrived at HMS in 1928, he was vision of microbiology embraced biochemistry, 31 years old, Monterey was a quiet fishing vilphysiology, morphology, ecology, and evolulage, and Stanford was just beginning to build a tion. Further, van Niel proposed that a unified, research program in experimental biology. A coherent science of general microbiology could new laboratory, named for Jacques Loeb, had provide a foundation for unifying the life scijust been built at HMS, and van Niel joined a ences. They were badly fragmented, he believed, team of researchers who were working on a either along organismal lines or because investibroad range of biological problems. gators followed disparate methodological apvan Niel brought to Stanford the approach to proaches. van Niel held that integrating micromicrobiology he developed during his training at the “I Susan Spath is an independent scholar in Berkeley, Calif. Volume 70, Number 8, 2004 / ASM News Y 359 obtainable only by including bacterial photosynthesis, implied that the biochemical aspects of photosynthesis could be understood in terms of oxidation reduction reactions, an important new insight at the time. van Niel’s Course in General Microbiology Encouraged Broad Thinking View of the Hopkins Marine Station on Monterey Bay in California. biology with other branches of biology would necessarily create a more comprehensive and more coherent science of life. van Niel’s belief in microbiology as a unifying force also rested on his conviction that the simpler forms of life present many of the fundamental features of life most clearly. Moreover, the diversity of the microbial world, he argued, meant that investigators could almost always find ideal experimental material for studying fundamental biochemical processes. Today, some of these ideas may seem selfevident, but in 1929 they were not. Because so many of these ideas are now well accepted, it may be difficult to appreciate how unusual they were at the beginning of van Niel’s career. In the late 1920s, the nature of bacteria was very much debated. Organismal biologists, especially, tended to view bacteria as genetically and cytologically bizarre, rather than as valid living things truly representative of more complex organisms. Moreover, it was not generally accepted that research on bacteria could yield insights of general biological importance. Within his first year at HMS, however, van Niel completed a major study on bacterial photosynthesis that illustrated beautifully how research on bacteria could yield unifying concepts with implications for plant physiology, evolution, and ecology. On the basis of that research, he formulated a general equation that applied equally to plant and bacterial photosynthetic processes. He showed that bacteria, like plants, use light energy to reduce carbon dioxide to sugars. His equation, which was 360 Y ASM News / Volume 70, Number 8, 2004 van Niel’s course was probably the first in the United States to offer a comprehensive survey of the microbial world. From the beginning, it was designed to convince students that microbiology offers marvelous opportunities for research in biology. Fundamentally, its purpose was to give students tools they needed to carry out their own research by teaching them methods and concepts, rather than a fixed body of knowledge. On one level, students learned many useful methods for handling microorganisms. On another level, van Niel endeavored to teach students how to develop their powers of scientific reasoning, emphasizing history and thereby familiarizing students with key observations on which the current body of knowledge rested, rather than asking them simply to accept such knowledge. As Robert McElroy, class of 1941, wrote, “No participant would ever forget this course because in addition to scientific knowledge, it imparted a philosophy and a method of thinking about the approach to biological research.” Overall, van Niel presented research in microbiology as a part of an even greater intellectual project to obtain a unified picture of nature. Students in the course met formally three days a week for 10 weeks. Lectures and laboratory work were artfully integrated. The first lecture began at 8 A.M. and could last as long as four hours. After lunch, van Niel often gave another lecture, again sometimes lasting four or five hours. However, some afternoons were instead occupied with laboratory exercises that could extend until midnight. By all accounts, van Niel brought a spellbinding intensity and passion to his teaching, and his former students remember that his lectures could be mesmerizing, even when they lasted for several hours. A master of the Socratic method, van Niel would relentlessly pose questions to and draw answers from his students. Typically, van Niel began with a discussion of van Niel’s Finishing School The heyday of van Niel’s course in the 1950s and 1960s coincided with the development of molecular biology. Because this was a microbial science, it seemed like a good idea to many— especially those entering from other fields such as physics—to learn something about microbes. A paradox ensued: most of the work in the fledgling science of molecular biology was being done with Escherichia coli and its phages, but these were topics that van Niel barely mentioned. So, why was his course so popular that eminent molecular biologists as well as their apprentices flocked to Pacific Grove as inexorably as the swallows coming (far down the coast) to Capistrano? Mind you, the course was meant principally for Stanford undergraduates, with the heavyweights being allowed as auditors only. Two answers suggest themselves. One is that the master was not just a teacher but also a magician. All who took the course reported that they had come under his spell. To wit, who else could keep an audience at the edge of their seats and with pencils poised for 8 hours or more of lecture in one day? Who else, on other days, would lead the class in a discussion that, hours later, ended with the conclusion that a certain experiment would solve the problem, only to find that the equipment for just THAT experiment was ready at the back of the room? All this was being the nature of science and scientific reasoning. For example, on the first day, van Niel would ask the class an apparently simple question, “What is microbiology?” He would use the answers to lead into a discussion of abstract reasoning and the importance and limitations of definitions. van Niel emphasized that the goal of scientific research was not only to discover facts, but to integrate them into general principles. He also insisted that any scientific conclusions should always be considered as tentative because new facts might require investigators to formulate whole new concepts. The scientific picture thus is not static but always open to change. When teaching the substance of microbiology, van Niel systematically interrelated practical methods and theoretical principles. For instance, he connected use of the microscope to discussions of discovering microorganisms and the debate over spontaneous generation. When he taught the methods for preparing pure cultures, he intro- witnessed by a wise old sea anemone that had been living in a tank in the laboratory for over 20 years. I can attest to the fact that the van Niel spell can last a lifetime. The second reason derives from the meaning of the Delft School of microbiology to which van Niel had made stellar contributions. The early days of microbiology included not only its better-known medical discoveries, but also exceptional work on the role of microbes on the cycles of matter in nature by Beijerinck, Kluyver, and their students. They came to the realization that, in Kluyver’s words, “Civilization owes much to the microbe.” He could have added: “and so does all of life.” Although the Delft school followed a different development than molecular microbiology, it was held in high esteem. It carried so much authority that taking a course taught by one of its eminent members was considered a sure way to learn about microbes in general. There may well have been some unintended consequences. I wonder how many people who came to Pacific Grove to take a course that might give them a license to work with E. coli came away with the big picture of the rest of the microbial world instead. Not a bad bargain. Moselio Schaechter San Diego State University San Diego, Calif. duced discussions of Koch’s postulates and the meaning of causality in science. He would also arrange for the students to use enrichment cultures to demonstrate how environmental conditions contribute to natural selection. The concluding first section of the course then surveyed the classical topics of microbiology, such as morphology, classification schemes, and the ecology of yeasts and several major groups of bacteria. During the second half of the course, van Niel took up the subject of microbial biochemistry. He lectured in detail on the fermentations of various microorganisms such as yeasts, lactic acid- and propionic acid-producing bacteria, and anaerobic sporeformers. Typically, he saved his favorite subjects such as photosynthesis for last. By the end of the course, students had gained considerable experience in handling a wide range of microorganisms, a great deal of substantive content, and a set of concepts with which they could approach a wide variety of Volume 70, Number 8, 2004 / ASM News Y 361 FIGURE 1 Kluyver and van Niel. biological problems. van Niel established the basic structure of the course in the 1930s and modified it very little. Of course, as the years passed, van Niel incorporated new findings into this structure. Between 1947 and 1962, he wrote out new sets of lectures every year. His notes for 1956, for example, contain 186 meticulously detailed, handwritten pages. Students Continue To Praise van Niel’s Course In the spring of 1930, when van Niel first taught his general microbiology course, only one student enrolled, Robert Hungate. At Stanford, Hungate had studied a mixture of general biology, ecology, and botany. He decided to take van Niel’s course on the recommendation of a friend. “I was enthralled,” Hungate wrote years later about the experience. van Niel’s “lucid presentation of ideas, his keen memory and insight, coupled with an intense interest in microbiology and in the student, made an indelible impression.” Whereas Hungate had no prior inclination toward the subject, he decided to investigate the protozoa and bacteria in the termite gut and their role in the digestion of wood. The first of many won over to microbiology by van Niel’s teaching, Hungate went on to a distinguished career in microbiology. 362 Y ASM News / Volume 70, Number 8, 2004 Later, during the summer, Horace A. Barker took the course and he, too, found that van Niel “had a very impressive way of speaking. . .” and that “his hypnotic intensity. . .made a deep impression.” Especially impressed by van Niel’s discussions of the biochemistry of bacterial and yeast fermentations, Barker decided to pursue a Ph.D. in chemistry to prepare for future research on these subjects. He spent 1935 as a postdoctoral fellow with Kluyver in Delft, where he began his important research on methane-producing bacteria. In 1936, after listening to van Niel’s lectures for a second time, Barker joined the faculty of the University of California, Berkeley, where he brought the Delft School approach to a new audience. Throughout the 1930s, a succession of students fell under van Niel’s spell before embarking on careers in microbiology, including Lewis Thayer, Michael Doudoroff, Howard Bliss, Roger Stanier, Steven Carson, E. H. Anderson, and J. O. Thomas. “My fate was decided in a few days,” Stanier wrote about his experience in van Niel’s classroom. “It took less than a week to conclude that van Niel was the ideal master and teacher; general microbiology was to be my domain...” In 1947, Stanier joined Barker and Doudoroff on the faculty of Berkeley. Stanier became a leading proponent of general microbiology in his own right. The widely used textbook, The Microbial World, was orginally coauthored by Stanier, Doudoroff, and Edward Adelberg, and was intended to capture as much as possible van Niel’s approach to microbiology. As the high quality of the course became known, researchers from a range of backgrounds traveled to HMS to take it. Photosynthesis researchers Robert Emerson, C. S. French, and William Arnold all took the course in the 1930s, as did biochemist Hermann Kalckar. Physicist Max Delbrück was a student in 1940 and later became a proponent of carrying out biological research on bacteriophage. Other graduates of the era include Robert McElroy, Garrett Hardin, and Dixie Lee Ray. McElroy, who first learned about bacterial luminescence in van Niel’s course, became one of the world’s experts on the topic. Hardin took van Niel’s teaching in a different direction and pioneered the field of human ecology. Ray eventually became head of the Atomic Energy Commission and then governor of the state of Washington. The 1940s and 1950s witnessed an explosion of interest in research on microorganisms, especially in genetics, making van Niel’s course an ever-morehighly valued resource. van Niel received hundreds of applicants for each session, from which he was limited to choosing only a dozen students along with a dozen auditors. In the first decade after World War II, Wolf Vishniac, Barbara Wright, Helge Larsen, Barbara Bachmann, and F. G. Lara received doctoral degrees under van Niel. All five became distinguished microbiologists who continued research in van Niel’s mode. The course played a significant role in training physical scientists who moved into biology after the war. For instance, Leo Szilard, Aaron Novick, Roderick Clayton, and Seymour Benzer all were participants. While Delbrück’s phage course at Cold Spring Harbor has been justly celebrated for training physicists to study biology, it is important to note that many of its participants, including Delbrück, also took van Niel’s course. Researchers with medical backgrounds also found special value in van Niel’s comprehensive presentation of the microbial world, including Bernard Davis, Harry Eagle, Ole Maaløe, Arthur Kornberg, Paul Berg, and Seymour Lederberg. Other graduates of the course who became leading molecular biologists include Dale Kaiser, Bruce Ames, William Sistrom, and Elie Wollmann. Even experienced microbiologists came to the course. For instance, Ralph Wolfe, who is professor emeritus in the department of microbiology, University of Illinois, Urbana-Champaign, audited the course in 1954. “The class was a fantastic experience that opened my eyes to a microbial world of unfamiliar organisms,” he wrote later in recalling his experiences. “I returned to Illinois with many ideas from van Niel that, together with some from Gunny, Luria, Sherman, and myself, became an organisms course that would be taught for nearly three decades.” van Niel’s Lasting Legacy Following the post-World War II expansion of microbiology, van Niel’s research, teaching, and vision continued to be major influential factors. On one level, van Niel’s course was the direct means by which several hundred students learned the basic methods and substantive content of general microbiology. His students and colleagues, their students and their colleagues, and so on, became an integral part of the worldwide community of microbiologists. On another level, van Niel’s course became a model for many other microbiology courses that adopted its message and methods, in whole or in part. Thus, van Niel and his followers greatly stimulated thousands of others to continue exploring the microbial world. In his teaching, van Niel advocated change, flexibility, and open-mindedness in scientific research. He warned his students against accepting uncritically the findings of other microbiologists, even the most famous. Surely, current innovative approaches in microbial ecology and evolution would delight van Niel if he were alive to learn about them. In some sense, the present era represents a fulfillment of much that he wished for. Recent successful efforts to integrate the microbial world into general biological understanding are now bringing forth some of the profound consequences that van Niel envisioned as early as the 1930s. The success of van Niel’s philosophical agenda is less certain. To van Niel, the deepest importance of science was its capacity to cultivate tolerance, objectivity, and rationality in the individual. He hoped that teaching students how to think, how to evaluate evidence, and how to distinguish between observations and inferences would lead to a better society overall. His philosophy of science and his research were consistent with his philosophy of life. To some extent, that coherence was made possible by the particular circumstances of van Niel’s scientific life. Not everyone, however, takes advantage of such opportunities as fully as van Niel did. For microbiologists who now operate in a world becoming morally, politically, and socially more complex, perhaps van Niel may still serve as a model to emulate. SUGGESTED READING Kluyver A. J., and C. B. van Niel. 1957. The microbe’s contribution to biology. Harvard University Press, Cambridge, Mass. Pfennig, N. 1987. van Niel remembered. ASM News 53:75–77. Bennet, J., and H. Phaff. 1993. Early biotechnology: the Delft connection. ASM News 59:401– 404. Spath, S. B. 1999. “C. B. van Niel and the Culture of Microbiology, 1920–1965.” Dissertation, University of California, Berkeley. Volume 70, Number 8, 2004 / ASM News Y 363