Chapter II

A. Introduction

In November 1928, C.B. van Niel and his family departed from the Netherlands by steamship. After six weeks at sea, they arrived at San Francisco, California. They then traveled by train one hundred and twenty miles south through fields and farmlands.⁽¹⁾ Van Niel's destination was Stanford University's marine biological laboratory, located on a small, rocky point of land near the southwestern tip of the Monterey Bay. A new laboratory building bearing the name "Jacques Loeb" overlooked the sea. For the next thirty-five years, van Niel worked in this laboratory. There, he developed and practiced his distinctive conception of microbiology.

Van Niel joined this laboratory in connection with a decisive shift toward "experimental biology" at Stanford in the 1920's and 1930's. No single overriding agenda accounted for this shift. Rather, it occurred because it served a variety of ends. Local institutional goals and a general project of medical reform made Ray Lyman Wilbur, President of Stanford, a champion of experimental biology. In the expectation of vaguely and variously defined "social returns," the Rockefeller Foundation (hereafter "RF") provided crucial financial support for experimental biology. Research scientists became enamored of experimental biology for a variety of reasons. For the director of Stanford's marine station, Walter K. Fisher, the new emphasis on experiment provided the rationale for expanding the size and prestige of his small institution, even at the risk of overshadowing his own interests in invertebrate taxonomy. Protozoologist C.V. Taylor, Wilbur's chief ally at Stanford, considered experimental and expensive research to be identical with modern biology. Taylor promoted the view that modern biological research should adopt the methods and logical rigor of chemistry and physics in order to identify what living things had in common, rather than catalogue their differences. Van Niel shared this view of biology. Only experimental research, he believed, could reveal the universal features of life. Belief in a fundamental "unity of life" motivated van Niel's enthusiasm for experiment. Several different but complementary agendas, then, drove the concerted expansion of experimental biology at Stanford in the 1920's and 1930's. Despite differences in purpose and outlook, administrators, patrons, and researchers found common ground in experimental biology.⁽²⁾

Stanford University's marine station provided an institutional base on which the ambitions for experimental biology could be transformed into practice. In 1928, the new Jacques Loeb Laboratory, paid for by the Rockefeller Foundation, became the most important site for experimental biology at Stanford. This laboratory provided an ideal institutional location for van Niel. Because it was not organized along established disciplinary lines, the laboratory accommodated his innovative research program that would bring together two previously disconnected fields, bacterial physiology and photosynthesis. Because traditional natural historical and modern experimental modes of scientific research co-existed harmoniously at the marine station, van Niel could pursue his broadly biological approach to microbiology.

The circumstances that made the marine station a congenial and productive laboratory lasted through the 1930's. For the most part, the different participants achieved their goals with little conflict.

B. A Seaside Laboratory for Stanford University

"Science" was a preeminent value at the Leland J. Stanford Junior University from its beginnings in the 1890's. Its founders, Leland and Jane Stanford, intended their new institution to be modern, progressive, and all that the word "scientific" conveyed in the late nineteenth century. In 1891, they chose a man of science, David Starr Jordan (1851-1931), to be Stanford's first president.⁽³⁾

The appointment of Jordan, a world-famous ichthyologist, assured a prominent place at Stanford for the life sciences, especially marine biology. A specialist in the taxonomy of fish, Jordan had discovered his vocation in 1873. That year, he attended Louis Agassiz's School in Natural History on Penikese Island, the first seaside summer school held in the United States.⁽⁴⁾ He was later wont to refer to this informal arrangement as the "greatest college in the world."⁽⁵⁾ It is not surprising that Jordan considered a marine laboratory to be essential for the new university, whereas he saw no pressing need for a philosophy department.⁽⁶⁾ Jordan's zoological colleagues, whom he had appointed, shared his enthusiasm. "The necessity for seaside laboratories for advancement in biological science has been thoroughly discussed and practically settled," wrote a Stanford professor in 1893.⁽⁷⁾

By definition located at the seashore, marine stations literally and figuratively exist at the boundary between nature and the laboratory. First established in nineteenth century Europe, they have been important locations for the development of the life sciences for two centuries. Most were founded in connection to fishing and other marine industries. Poised between seashore and city, beach and lab bench, marine stations are distinctive institutions in which the practice of scientific research coexists with the contemplation of nature and summertime recreation.⁽⁸⁾

France, with its long and complicated coastline, vigorous fishing industry, and strong tradition in zoology, was the leader in founding marine laboratories. The first permanent laboratory in that country was founded in 1859 in the village of Concarneau. Others were established at at Arachon (1867), Roscoff (1874), Wimereux (1873), Villefranche (1880), and Banyuls-sur-Mer (1882).⁽⁹⁾ The British and Scandanavians followed suit and founded a fleet of marine stations of various sizes between about 1870 and 1910.⁽¹⁰⁾

In the 1870's, the example of the French marine stations inspired Anton Dohrn, a German zoologist, to organize a major research laboratory. Excited by Darwin's theory of evolution and blessed with an ample personal fortune, Dohrn planned to build a large, well-equipped institution dedicated to research. In 1874, Dohrn's vision became a reality when the Naples Zoological Station opened its doors on the coast of Italy. It soon became the most prestigious marine laboratory in the world. Investigators from around the world, including the United States, arranged to spend time at the Naples station to take advantage of the excellent conditions for work, including the availability of marine life of the Bay of Naples. During the laboratory's first quarter century, researchers investigated problems in phylogeny, morphology, and embryology.⁽¹¹⁾

In the United States, the first permanent marine laboratory evolved from Agassiz's summer school and others like it.⁽¹²⁾ The Marine Biological Laboratory at Woods Hole, Massachusetts (hereafter "MBL"), received its charter in 1888. Modeled in part on the Naples Station, the MBL retained a strong commitment to teaching along with research, as Agassiz had advised. Two years later, the Biological Laboratory was established at Cold Spring Harbor in New York by a triumvirate which included a graduate of Agassiz's school.

For more than a century, the MBL has played a defining role in American biology.⁽¹³⁾ Historian Philip Pauly has argued that "biology" as an intellectual and academic category emerged most forcefully and rapidly in the United States because of the existence of the MBL.⁽¹⁴⁾ Historians Ronald Rainger, Keith R. Benson, and Jane Maienschein have concluded that the core issues of American biology at the turn of the century were defined around the study of marine organisms.⁽¹⁵⁾

Observing the importance of the Naples Station and the MBL, Jordan and his colleagues became intent on having a seaside laboratory for Stanford. In 1892, they appealed to Timothy Hopkins, the adopted son and heir of railroad magnate Mark Hopkins, for financial support. Like his father, Hopkins was a close friend of the Stanfords and a partner in the Pacific Improvement Company, the syndicate that built the Southern Pacific Railroad. During a European sojourn, Hopkins was greatly impressed by a visit to the Naples Zoological Station. In 1892, Hopkins agreed to pay for a small building on the pristine Monterey Bay, about sixty miles south of the main Stanford campus. The Pacific Improvement Company donated a parcel of land on Point Aulon (the present location of the Monterey Bay Aquarium).⁽¹⁶⁾ The wood-framed building was named the Hopkins Marine Station in honor of its patron, despite the risk of confusion with an eminent model (and rival) institution on the East Coast.⁽¹⁷⁾ Providing funds for books, equipment, and a new building in 1893, Hopkins supported the laboratory until his death in 1936.

Like Stanford, the Universities of California and Washington took action to establish marine laboratories on the Pacific Coast. In 1890's, the University of California considered founding a laboratory on the Monterey Bay, but chose a site in southern California instead. That laboratory became the Scripps Institute for Oceanography in 1912. The University of Washington set up its first seaside summer program in 1894, and built a permanent laboratory at Friday Harbor in 1906. Undergraduate teaching remained its principle function for several decades.⁽¹⁸⁾

From its inception, the Hopkins laboratory was committed to two endeavors, teaching and research in biology. The first twenty years emphasized programs for secondary school and college teachers. Over time, its teaching emphasis gradually shifted toward training advanced students for research. Undergraduates were encouraged to spend their summers there to learn about marine organisms from direct experience. Ray Lyman Wilbur, a Stanford undergraduate from 1892 to 1896, spent three summers at the marine laboratory as part of a program in physiology. As a medical student, he returned to the seaside laboratory to assist in physiology courses. In 1898, a dramatic physiological demonstration by the noted physiologist Jacques Loeb made a vivid impression on Wilbur.⁽¹⁹⁾ Loeb came to Pacific Grove in 1898 and 1900 in search of sea urchins spawning in winter. There, he carried out some of his sensational experiments in artificial parthenogenesis. Loeb again took frequent refuge in the Monterey area while professor at the University of California from 1903 to 1910.⁽²⁰⁾ Loeb brought considerable attention to the new experimental biology in Stanford circles, not all of which was appreciated, especially by adherents of more traditional approaches.⁽²¹⁾ Later, as Stanford's third president, Wilbur described his summers at the marine station as "one of the great experiences of my University career."⁽²²⁾

C. Experimental Biology -- "New Data, New Methods, New Objectives"

The expansion of experimental biology at the expense of natural history and field sciences began in the late nineteenth century and has continued to the present. Describing, explaining, and analyzing the shift toward experimentation has become a major historiographical issue for historians of the life sciences. Historian Garland Allen's proposal that a general "revolt against morphology" accounts for the turn toward experimental has been much criticized, but no convincing alternative has yet been proposed.⁽²³⁾ While the broad scale of this change implies that general pressures are at work, the specific timing, course, and facility of this shift varied according to local institutional conditions. More detailed studies of specific cases are needed before a satisfactory explanation for the general trend can be advanced. Historian Jane Maienschein's study of biology between 1880 and 1915 is an important contribution to that end. In her analysis of four prominent biologists of the period, she describes gradual, evolutionary transformation, rather than discontinuous change, in the kinds of questions asked, organisms studied, and methods used. To account for this transformation, she appeals to "a confluence of intellectual, institutional, social, and personal factors."⁽²⁴⁾

A study of the expansion of experimental biology at Stanford provides the opportunity to identify specific factors at work in a specific institution. Medical reform provided the first impetus. In 1911, Wilbur, a strong supporter of medical reform, became the Dean of Stanford's new and controversial medical school. Five years later, he accepted the Stanford presidency, in part to ensure the survival of the medical school. The second president, a geologist, had called it a "menace." In contrast, Wilbur considered a first-rate medical school to be essential for Stanford.⁽²⁵⁾

Wilbur's commitment to the ideal of research exerted a second stimulus. Wilbur's ambitious plan for Stanford was to transform the small, private college into a powerful research university. This required changing the purpose of Stanford from purveyor of moral education to gentlemen to producer of knowledge for society.⁽²⁶⁾ To this end, he cultivated an ethic of research among his faculty. He sought to instill the belief that research was not a mere adornment of the academic career, but a basic obligation. The experimental sciences offered new possibilities for conducting research.

Wilbur gave the reformation of the life sciences special attention. Like other advocates of "scientific medicine," Wilbur believed that the biological sciences provided the essential foundation for medicine. He sought to expand and modernize both medicine and the life sciences at his institution.⁽²⁷⁾ Wilbur had considerable sympathy for the study of nature for its own sake, but his overriding interest was human welfare.

Wilbur's modernization project entailed changes in the basic definition, organization, and practice of the life sciences at Stanford. First, as in all fields, he supported a shift in emphasis toward research. Second, he promoted the development of experimental research over observational and descriptive work. Third, he sought to establish "biology" as both a meaningful intellectual category and legitimate academic unit, in place of the disorderly disciplines and departments inherited from the nineteenth century. Fourth, he actively promoted the introduction of the experimental methods and logical rigor of chemistry and physics into biological research. Like other medical reformers of the era, Wilbur considered biochemistry and biophysics to be the core disciplines for bringing the momentum of medical reform to the life sciences.⁽²⁸⁾

For Wilbur, the development of experimental biology and the medical school were interrelated and mutually advantageous projects. At the same time, Wilbur did not consider biology to be exclusively a resource for the improvement of medical training and practice. He was well aware of the importance of California's fishing industry and the strong tradition in fish biology at Stanford.

Wilbur made the Hopkins Seaside Laboratory into a salient for the advancement of his administrative projects. A personal tie reinforced his interest in the marine station; he and Hopkins were close friends. Under Wilbur's presidency, the laboratory expanded in size and status. With the assistance of Hopkins, Stanford acquired eleven acres of land known as China Point on the Monterey Bay. Legally owned by the Pacific Improvement Company, this land was occupied by a Chinese fishing village until 1903 when it was destroyed by a fire the origin of which was never officially determined. Rumors have persisted to the present that the fire was intentionally set by the landowners to dislodge the Chinese residents.⁽²⁹⁾ In 1917, Stanford built a two-story brick laboratory on this location. Wilbur gave the institution a full-time director, the aptly-named Walter K. Fisher, and a new name with a technical connotation suitable for a research laboratory. In 1917, the "seaside laboratory" became the Hopkins Marine Station (hereafter "HMS").⁽³⁰⁾

Wilbur's main administrative project in the 1920's was to simplify the baroque departmental structure of the campus by establishing a smaller number of "Schools," one each for Humanities, Physical Sciences, Biological Sciences, and Social Sciences. For Wilbur, "Biology" was both a logically defined domain of knowledge and a useful instrument for administrative reform. Under his presidency, it acquired institutional reality at Stanford. Wilbur himself, along with faculty members from Botany, Entomology, Physiology, and Zoology, taught the first course at Stanford called Biology during the academic year 1919 to 1920. Wilbur planned to integrate the diverse life sciences departments into a single administrative unit to be called "Biology."⁽³¹⁾

Wilbur's project was part of a second wave of attempts to establish schools or departments of biology in American universities. The first wave occurred between 1870 and 1900, with Johns Hopkins leading the way. Columbia and the University of Chicago also established biology departments before 1900. As historian Philip Pauly has pointed out, at these institutions schools of medicine were relatively weak, a situation that allowed general biological programs to gain a foothold.⁽³²⁾ After about 1915, however, the medical reform movement created new service roles for teaching basic sciences courses, including biology.⁽³³⁾ In the 1920's and 1930's, Harvard, Caltech, and Stanford established schools or departments of biology.

At Stanford, opposition to the establishment of "biology" as an academic and intellectual category derived from the traditional departments of the life sciences like Botany and Zoology. Establishing Biology as stable administrative unit required undermining the power of these departments, where morphological and taxonomic approaches were still strong. Wilbur emphasized experimental biology to undermine the legitimacy of the traditional departments as logically defined units. In 1922, he made a first, and only partially successful, attempt to consolidate the various life science departments. He created a School of Biology, but uneasy faculty in Botany, Zoology, and Physiology, fearful of losing power and status, managed to keep the School on the same administrative level as the traditional departments.⁽³⁴⁾ At this time, the HMS was also administratively equivalent to a department.

In 1925, Wilbur made a key appointment for his campaign to advance experimental biology. Taking advantage of recent retirements, Wilbur appointed C.V. Taylor (1885-1946) to the Stanford faculty. A former student of the classical protozoologist Charles Kofoid at Berkeley, Taylor had become a strong advocate of experimental biology while carrying out his dissertation research, in which he proved his professor wrong. On morphological grounds, Kofoid had proposed a neuromuscular function for a prominent group of fibrils in the protozoan Euplotes. Taylor became convinced that their function could not be determined by observation alone. Using technically advanced micro-dissection techniques, Taylor showed that the fibrils did not act like contractile muscles, but functioned in the conduction of sensory signals. During the next twenty years, his enthusiasm for experiment only increased.⁽³⁵⁾

Taylor became a strong and effective ally for Wilbur in building experimental biology at Stanford. Like Wilbur, Taylor believed that the aim of modern research should be to reveal the "unity of life" rather than to emphasize diversity. Taylor also shared the view that the traditional departmental structure obstructed progress toward this goal.⁽³⁶⁾ Wilbur positioned Taylor to lead his ongoing but still unsuccessful efforts to consolidate the departments into a single School of Biology.⁽³⁷⁾ In 1926, Wilbur appointed Taylor associate director of the Hopkins Marine Station to build experimental research programs there.

Wilbur and Taylor were firmly convinced that progress in biology and physiology required the use of chemistry and physics, a widely shared view in the 1920's.⁽³⁸⁾ In his presidential report for 1925, Wilbur noted the new direction in biology, presumably with satisfaction: "There is a shift of interest in zoology and botany from systematic and morphological studies to general physiology and particularly to the applications of physics and chemistry."⁽³⁹⁾

An important patron supported this new direction in the 1920's and 1930's: the Rockefeller Foundation. The motives of the RF in supporting experimental biology are open to debate. They were perhaps too diffuse and various (and possibly nefarious) to be easily characterized. However, there is general agreement that the expectation of "social returns," however vaguely defined, motivated the RF's substantial investment in experimental biology.⁽⁴⁰⁾

As a member of the RF board of trustees from 1923 to 1940, Wilbur could be said to have shared the Foundation's goals, including "social control." "It is imperative for us to try to gain control of our complicated emotional setup," he wrote in his memoirs, "Research in the social sciences has become of transcendental importance if we are to know how to guide society into safer ways."⁽⁴¹⁾ He believed that, ultimately, the social sciences should be based on biology: "We can in no way escape the implications of biology. In social science, the reactions of all living organisms must be considered." Wilbur's interest in social problems co-existed with other motivations for modernizing the life sciences at Stanford, the reform of medicine, institutional expansion, administrative efficiency, and the study of nature for its own sake.

In 1925, Edwin Embree became head of the newly established and vaguely defined "Division of Studies" at the RF. To secure his division within the Foundation, Embree sought important projects to fund.⁽⁴²⁾ In the ppring of 1925, he visited the major academic institutions on the West Coast, including Stanford. He was especially interested in meeting with Fisher, the director of the HMS, to discuss "human biology" and "the biological questions which lie next door" to the Foundation's traditional interests in public health and medical education.⁽⁴³⁾

Trained as a field naturalist and a specialist in invertebrate anatomy and taxonomy, Fisher nonetheless welcomed Embree's overtures and the opportunity to expand experimental research at the HMS. Emphasizing the Station's commitment to research in physiology, biochemistry, and biophysics, Fisher described his "most pressing needs" in detail to Embree.⁽⁴⁴⁾ By emphasizing his interest in experimental biology and physiology, Fisher distinguished the Hopkins Marine Station from others on the West Coast. The laboratory operated by the University of Washington, Embree noted, was still engaged in undergraduate teaching, whereas the Scripps Institute in southern California was turning more and more toward oceanography. A Canadian government laboratory on Vancouver Island was most concerned with the fishing industry. Embree concluded that RF should support the Stanford marine station precisely because it was "the only one on the Pacific Coast devoting itself largely to experimental physiology." He recommended that the RF provide funding for a new research laboratory at the HMS.⁽⁴⁵⁾

President Wilbur was delighted by this development.⁽⁴⁶⁾ The financial support of the RF made possible a major advance in his project to build up experimental biology at Stanford. To consummate the partnership between patron and beneficiary, Wilbur suggested a name for the new building: "the Jacques Loeb Laboratory for Marine Physiology." "Do you think there would be any objection to this on the part of the Foundation?" he asked disingenuously. In the same note, he casually suggested that more funding would be needed to complete the new laboratory.⁽⁴⁷⁾ As plans for the new building developed, the costs escalated and Wilbur requested a second grant of $50,000.⁽⁴⁸⁾ In the official proposal for the new grant, Wilbur again invoked the scientific legacy and legitimating power of the name of Jacques Loeb.⁽⁴⁹⁾

In the late 1920's, Loeb's name evoked a variety of meanings in American scientific culture. From 1910 until his death in 1924, Loeb was a leading proponent of experimental physiology at the Rockefeller Institute for Medical Researcher. Trained by the elite of German physiology, Loeb served as a kind of icon for American science. For some practitioners, Loeb and his work represented the importance of "right method" in biology: the use of rigorous experimentation based on chemistry and physics. For others, his name meant a radically reductionist philosophy of biology, a commitment to promote a mechanistic conception to life and to eradicate vitalism. Loeb's name could also invoke the potential power of physiology to create new life forms.⁽⁵⁰⁾

For many scientists of the era, the core of Loeb's significance lay not in the substance of his scientific work, but rather in his cultural meaning. Reincarnated by author Sinclair Lewis as the physiologist Max Gottlieb in the popular novel Arrowsmith (1924), Loeb represented the many dimensions of the ideal of pure science. Lewis won the Pulitzer prize in 1926 for this novel, which suggests that its themes were especially high on the cultural agenda at the time.

Naming the new laboratory for Jacques Loeb associated these various meanings with the marine station. It conveyed the impression that the institution was a temple both to pure science and to experimental physiology of the highest quality. In the spring of 1926, the Foundation agreed to Wilbur's proposed name and to the second grant of $50,000, assuring completion of the building.⁽⁵¹⁾ Bearing the name of Jacques Loeb engraved in stone above the entranceway, the new laboratory would be equipped with the latest physiological equipment and serve eight full-time researchers dedicated to chemo-physical biology.

To lead the development of experimental research at the new laboratory, Wilbur chose L.G.M. Baas Becking (1895-1963), a talented and temperamental polymath from the Netherlands.⁽⁵²⁾ Becking had studied one year at the THD before earning a Ph.D. in physics from the University of Utrecht in 1919. He then moved to California in the hope that the climate would ease his chronic asthma. He continued scientific research and earned a Ph.D. in Botany from Stanford in 1921.⁽⁵³⁾ His training and interests made him an excellent candidate to bring the methods of physics to problems in biology. In 1924, he was appointed professor of Biology at Stanford. He was interested in comprehending the flow of matter and energy from the biophysical level to the earth as a whole.

Becking envisioned a comprehensive research program at the new laboratory that would integrate chemistry, physics, biology, geology, and oceanography. He became intimately involved in designing the new laboratory. He took considerable trouble, for example, to find pipes that would not contaminate seawater pumped into the aquaria. In his reports to the RF, Wilbur made frequent reference to his research star's activities. Becking was one of the first to get new equipment with the Rockefeller money.⁽⁵⁴⁾

When the new Jacques Loeb Laboratory neared completion in 1928, Wilbur suggested that the older building be officially named as well. The scientists at the HMS unanimously rejected Wilbur's first suggestion, the name of a recently retired Stanford professor. Fisher was adamant that the building be named for an internationally known figure in research, ideally in oceanography. The name "Alexander Agassiz" satisfied both requirements and was the unanimous choice of the marine station scientists.⁽⁵⁵⁾ The names of the two buildings, Jacques Loeb and Alexander Agassiz, symbolized the coexistence of two different programs in biology at the HMS, one oriented toward experimental biology based on chemistry and physics, the other toward oceanography and organismal biology. Both programs expressed the increasing importance of research at Stanford, even if they differed as to subject of study and methodologies.

A great success for Director Fisher, the new laboratory for the marine station was merely a small prize for Wilbur. A more ambitious plan for the biological sciences failed when he was unable to raise funds to match a grant of $750,000 from the RF.⁽⁵⁶⁾ Wilbur's partial success, however, had important consequences. A new laboratory was built that provided a location for the expansion of experimental biology at Stanford.

Lasting only one year, Embree's initiative was unsuccessful in comparison to subsequent programs at the RF, especially those established under Warren Weaver, director of the Natural Sciences division in the 1930's. Nonetheless, Embree's support of "human biology" as a subject distinct from medical research per se and his promotion of experimental biology based on physics and chemistry anticipated elements of Weaver's program. From the point of view of the HMS, Embree's small program produced enduring results. It provided for a new place for scientific work, richly endowed with the latest equipment for experimental physiology. It was up to Stanford to find scientists capable of using it.

D. The Jacques Loeb Laboratory

In principle, the two laboratories at the marine station complemented rather than competed with each other. In practice, the potential for conflicts of various kinds was in place. Officially, Fisher was the director of the station as a whole, but Becking wielded considerable influence over the affairs of the Jacques Loeb Laboratory. As the local expert on experimental biology, Becking enjoyed considerable support from Wilbur. In 1927, Becking was appointed to first "Dr. Morris Herzstein Professorship in Biology," made possible by a recent bequest to Stanford.⁽⁵⁷⁾ To Fisher's considerable irritation, Becking was given substantial authority in choosing researchers for the new laboratory.⁽⁵⁸⁾

In the spring of 1928, Becking returned to the Netherlands on sabbatical. While there, he looked for promising candidates for the new laboratory at the marine station and paid several visits to Kluyver in Delft. On one occasion, Becking met Kluyver's assistant, the talented and underemployed van Niel. Becking saw in van Niel a fellow Dutchman who was talented researcher with solid training in chemistry, substantial experience with microorganisms, and a broadly biological conception of microbiology. These qualities made van Niel an ideal prospect for the grand, collaborative program Becking envisioned for the Jacques Loeb Laboratory. Becking even shared van Niel's specific interest in the physiology of the purple bacteria. Later that spring, Becking asked van Niel if he would accept a position at the Jacques Loeb Laboratory.⁽⁵⁹⁾ Becking's proposal was probably the single best opportunity in the world for a scientist with van Niel's interests and disposition. This was not immediately obvious to van Niel, however, and he hesitated to accept Becking's offer. Like many other European intellectuals, he considered the United States to be corrupted by crass commercialism. Recalling his decision later, van Niel wrote, "There were two places in the world we never wanted to go -- the Netherlands East Indies and America. People went there to make money, and I was not interested in making money."⁽⁶⁰⁾ After further consideration, van Niel perceived the advantages of relocating to the HMS. Kluyver assured van Niel that he could return to the laboratory in Delft if he did not like life in California. Van Niel wrote to Becking that he would accept an offer with one important stipulation, that he be completely free to study the purple bacteria.⁽⁶¹⁾

When he returned to Stanford from Europe, Becking praised van Niel's potential and exaggerated his accomplishments to Wilbur.⁽⁶²⁾ An appointment for van Niel was soon arranged. In the fall of 1928, he received official word from Wilbur that he was appointed "Acting Assistant Professor of Microbiology" at Stanford as of January 1, 1929.⁽⁶³⁾ In November of 1928, he and his family departed from Antwerp to make the crossing to a new world.

On December 28, 1928, van Niel woke to see a brilliant sunrise over the Golden Gate, the beautiful point of entry to the San Francisco Bay.⁽⁶⁴⁾ Becking met van Niel and his family as they disembarked. They then drove to Stanford for an introductory audience with Wilbur. They continued by train through the rich farmlands of the Salinas valley to the cluster of small towns and fishing villages along the crescent-shaped Monterey Bay. Becking took the van Niels to the nearby village of Carmel where he had rented a house for them. Van Niel marveled at the mild climate, brilliant sunshine, and profusion of lovely plants in bloom. The exquisite setting of the Hopkins Marine Station, located in the village of Pacific Grove, "enchanted" him.⁽⁶⁵⁾

In 1928, Pacific Grove consisted of clusters of summer cottages, a smaller number of permanent residences, and several large fish canneries. Sardines were the base of the local economy. The Marine Station was situated on a rocky promontory variously known as China, Mussel, or Cabrillo Point, a stone's throw away from Cannery Row.⁽⁶⁶⁾ The Alexander Agassiz Laboratory faced north, overlooking a sheltered landing place and small harbor for boats. Three stories high, it contained large rooms for sorting collections and a few small research rooms. The large general laboratories were mainly used for teaching. The Agassiz laboratory housed "the staff concerned with the hydrobiological survey of the Monterey Bay and with the more traditional lines of research in zoology and botany."⁽⁶⁷⁾

The Jacques Loeb Laboratory (or "JLL") faced toward the west. Every aspect of the building from its name to its plumbing announced that experimental biology had arrived at Stanford. It was two stories high in the middle section with two wings one story in height on each side. The eastern wing contained four laboratories for general use, four rooms for private investigations, two storage rooms, and an office. Specialized laboratories were located in the western wing: a chemistry laboratory, a thermostat room, a balance room, and a special apparatus room. Two rooms could be maintained at constant temperatures. Laboratories were specially set up for spectroscopy, polarimetry, and photometry. A "heliostat room" was illuminated directly through the roof of the building. Two more rooms were set up for micromanipulation. A general photographic darkroom, a small store room and general office completed the west wing. The second story contained a reading room, two private rooms, and two stack rooms. A full machine shop and storeroom were located in the adjacent boiler house.⁽⁶⁸⁾

Most of the laboratories and work rooms were supplied with sea water, hot and cold fresh water, distilled water, gas, alternating and direct current up to 200 volts, compressed air up to 100-pound pressure, and vacuum outlets. The larger laboratories were provided with variable currents and special hoods for chemical work. Sea water was pumped from a natural upwelling in the ocean into a reservoir situated on the top of a rocky outcropping. It then flowed into the laboratories by gravity.

Even after the building was complete, equipping and organizing the laboratories continued to be a major undertaking for the staff. Becking complained to the President of Stanford that the $14,000 provided by the university for equipment and instruments was sadly inadequate. He pointed out that the Scripps Institution in Southern California had five times as much money to support a comparable staff. In contrast, the scientists at the HMS were obliged to construct themselves many of the devices they needed. The resident chemist, for example, made dispersion gratings, a radio micrometer, and a high dispersion densitometer. An apparatus for determining boiling, melting, and freezing points and a compensation calorimeter were also home-made. Perhaps Becking was not exaggerating when he grumbled, "Due to the fact that all of the members of the staff have sacrificed a large amount of valuable time to act as laborers, carpenters and mechanics, the place looks well equipped."⁽⁶⁹⁾

A lavish array of the latest instruments and equipment had, in fact, been purchased outright. Becking admitted that the microscopic instrumentation included "practically every microscopic accessory one might need, such as dark field, and ultra-violet systems, a monochrometer, microspectral equipment, and micro-manipulators." Five micro-photographic cameras, a polarization microscope, and special microscope stages that could be temperature controlled were also available. Instruments for biophysical work included an Abbe refractometer, several colorimeters and potentiometers, and a conductivity apparatus. The bacteriological laboratory was provided with the essentials: an autoclave, ovens, and incubators. Among the lower priced items indispensable for experimental work were grinders, mills, shakers, centrifuges, and balances. A special cabinet with lights inside was purchased for van Niel. Despite Becking's complaints, the JLL was by far the best equipped facility for experimental biology at Stanford.⁽⁷⁰⁾

The laboratory constituted a distinctive place of scientific work. It provided the advantages of an ambitious university building its research programs with the isolation and tranquillity of a small community set apart from the complex distractions of urban life. A second notable feature of the marine station as place for biological research was that it had almost no contact with medicine; its practical ties were to the fishing industry and oceanography. It was an environment where traditional natural historical and modern experimental modes of practice could co-exist. Because it was not organized along traditional disciplinary lines or departmental boundaries, research topics could be more freely chosen.

For Fisher, supporting the expansion of experimental biology brought him a new building that more than doubled the size his institution. At the same time, he could continue to direct programs in oceanography and traditional marine biology. Enjoying a period of institutional expansion, Fisher did not have to sacrifice one for the other.

Although there was no mistaking the power and status of the new experimental biology embodied in the Jacques Loeb Laboratory, Fisher remained committed to a broadly defined program he called "Oceanic Biology." In official bulletins, Fisher described the research of the station in broad terms:

The research in a general way will concern itself with oceanic biology....Special attention will be paid to the relations between the organism and its environment, including a consideration of the composition of sea-water....A systematic survey of the fauna and flora will be undertaken and any promising leads in the natural history of marine organisms will be followed.⁽⁷¹⁾

The program outlined by Fisher included both new experimental projects and more traditional lines of investigation. Despite (or because of ) the new emphasis on experimental biology, Fisher was careful to make clear that descriptive studies, especially taxonomy, were valued aspects of the Station's work. Fisher's field of expertise was "Marine Zoology," by which he meant the anatomy, taxonomy, and natural history of oceanic invertebrates. Representing oceanography, Tage Skogsberg studied "Ecological problems pertaining to open waters, and comparative morphology and taxonomy of planktonic organisms." His principal obligation was to carry out a hydrobiological survey of the Monterey Bay in cooperation with the California State Fish and Game Commission. Harold Heath, professor of Zoology, was an accomplished ornithologist and studied embryology from a morphological viewpoint.⁽⁷²⁾

The new experimentalists were represented by embryologist A.R. Moore; Becking, who traversed botany, biophysics, and bacteriology without much regard for disciplinary boundaries; C.V. Taylor who investigated the physiology of protoplasm; and Harold Mestre, charged with carrying out spectroscopic studies.⁽⁷³⁾

The diversity of research interests at the HMS suited Becking's purposes. He saw the new laboratory as an opportunity to advance his grand and ambitious synthetic program of research. He envisioned that the group of specialists he had helped to assemble would work closely together on common problems that he would choose. His goal was to study the geological, physical-chemical, and biological features of oceans and salt lakes and their inter-relations.⁽⁷⁴⁾ He had high hopes that the Hopkins Marine Station would become "the Naples of the United States," as prestigious and productive as the most famous marine laboratory in the world, the Zoological Station in Italy.⁽⁷⁵⁾

When van Niel arrived in 1929, the station was admirably set up to fulfill this dream. Van Niel joined eight scientists in residence at the HMS and six associated faculty from Berkeley, Stanford, and Harvard. In contrast to the laboratory at Delft, all of the scientists were concerned with biological problems. A wide range of research interests was represented. Van Niel was delighted with his new situation. Writing to Kluyver in Delft, van Niel described the marine station as a "wonderland" where a "holiday atmosphere" prevailed, although everyone worked very hard. He often ate lunch on the rocky shore outside the laboratory, amid gulls and pelicans, seaweeds and shellfish. Underneath the idyllic surface, though, van Niel sensed political undercurrents from which he resolved to keep his distance.⁽⁷⁶⁾

For the first year, the laboratory lived up to its promise. Becking instigated several collaborations and organized a weekly tea where the scientists discussed their research informally. He was said to be much like Jacques Loeb, temperamental but brilliant and stimulating.⁽⁷⁷⁾ Underneath the surface, however, tensions between Becking and Fisher were intensifying. Irreconcilable conflict between them was perhaps inevitable. Each had strong views about the proper direction of research and Becking constantly challenged Fisher's authority. Fisher was adamant that marine biology should be the central focus of research at the Station. Becking's "program" was so broadly defined that almost any work in experimental biology could be undertaken in its name.

Scientifically, van Niel impressed his colleagues soon after his arrival. In June of 1929, he presented a seminar on "The Unity of Biochemistry," in which he outlined Kluyver's general theory of metabolism as comprised of a small set of dehydrogenation reactions.⁽⁷⁸⁾ Afterwards, a colleague described van Niel as "brilliant."⁽⁷⁹⁾ Writing to Wilbur, Fisher called van Niel "a real 'find,'" and "a scientist of the first rank with a keen, original, and critical mind."⁽⁸⁰⁾ Becking introduced van Niel to Hermann Spoehr, an expert in the chemistry of photosynthesis and director of a small laboratory funded by the Carnegie Institutes of Washington.⁽⁸¹⁾

Soon Becking and Fisher were fighting over van Niel. Fisher was furious to learn that van Niel had never been informed by Becking that he should expect to work on oceanological problems.⁽⁸²⁾ Recognizing van Niel's potential, Fisher attempted to direct him toward marine biology. "Dr. van Niel has indicated he will later join the oceanological program," he wrote to Wilbur, "-- an event of importance to the Station for which I have been particularly waiting."⁽⁸³⁾ Van Niel made some attempt to comply with Fisher's wishes and gave two lectures in which he considered bacteria in relation to marine biology: "Bacteria and their role in the household of the ocean," and "The bacteriological side of marine biology."⁽⁸⁴⁾ Before long, however, van Niel pursued his research without regard to whether it connected to marine biology or not.

Friction between Becking and Fisher over control of the laboratory's budget and over the affairs of the HMS in general reached a breaking point.⁽⁸⁵⁾ Despite his instrumental role in its creation, Becking did not remain long at the Jacques Loeb Laboratory. In 1930, he resigned from Stanford and returned to the Netherlands to become Professor of Botany at the University of Leiden.⁽⁸⁶⁾ He became a prominent figure in Dutch science and politics.⁽⁸⁷⁾ Becking's vision of a grand synthetic program departed from the marine station with him and the few collaborations he had instigated soon collapsed into independent projects. He left behind him, however, a new laboratory with an excellent stock of equipment for physiology and several lively research projects underway.⁽⁸⁸⁾ He left microbiology secure in the hands of van Niel, whose very first year of research yielded exceptionally important results.

The departure of Becking, while disappointing in some ways to Wilbur, provided him with an extremely valuable bargaining chip, the Herzstein Professorship. Wilbur was quick to use it to secure the return of Taylor to Stanford. Spending the year 1930 to 1931 at the University of Chicago, Taylor was seriously considering an attractive offer from the University of Michigan.⁽⁸⁹⁾ His year at Chicago had only reinforced his conviction that experimental biology was the key to the future of the life sciences. "A new day has come," Taylor wrote to Swain, the acting president of Stanford, "visible at every turn here at Chicago. New data, new emphases, new methods, and to some extent new objectives are being placed before the students."⁽⁹⁰⁾ The base for experimental biology established at the marine station was attractive to Taylor. "With the splendid opportunities now afforded at the Hopkins Marine Station," he wrote to Swain, "and with the close sympathies and affiliations between the biological and the physical sciences on the Campus, what other university in the land can offer as much? The only remaining need is more funds, but the ready and very effective answer to that is a group of hard-working, thoroughly capable, harmonious souls who see and drive toward a common goal. Funds must come our way."⁽⁹¹⁾ The promise of the chaired professorship and renewed commitments from Wilbur to both experimentation and biology were ultimately successful in convincing Taylor to return to Stanford.⁽⁹²⁾

Van Niel began several projects early in 1929, but soon his attention was monopolized by the purple bacteria.⁽⁹³⁾ Since his student days in Delft, van Niel had been intrigued by the confused and conflicting interpretations of the metabolism of the purple bacteria. Several reports claimed that the purple bacteria were capable of photosynthesis; others heatedly claimed the opposite. The possibility of resolving this confused state of affairs attracted van Niel's interest. Inconveniently, however, the purple bacteria were not marine organisms. They lived in places rich in hydrogen sulfide (H₂S), like sulfurous hot springs or the malodorous canals of Delft.

Nonetheless, van Niel advanced his project by taking full advantage of the local opportunities available to him: his new status as an independent researcher, the excellent conditions for experimental research at the Jacques Loeb Laboratory, and Wilbur's emphasis on biology based on chemistry and physics. These circumstances permitted van Niel to launch a research program in general microbiology in a marine station. He undertook a comprehensive study of the general biology of the purple bacteria and a group of organisms often found cohabiting with them in nature, "the green bacteria." His work led him to conclude that these organisms carry out a special form of photosynthesis. That conclusion led him to a fundamentally important insight into the biochemistry of this process.

E. Photosynthesis

In the eighteenth century, chemists, not botanists, conducted the first investigations into the physiological activities of plants. While botanists were preoccupied with describing, naming, and classifying plants, chemists investigated the problems of plant nutrition.⁽⁹⁴⁾ Their studies led to the remarkable conclusion that plants derived their material substance not from soil, as Aristotelian doctrine maintained, or exclusively from water, as van Helmont thought he had proved, but also from air. In 1771, Joseph Priestley, one of the principals in this drama, reported that green plants had the special capacity to restore air that had been "vitiated" by animal respiration. In his terms, green plants evolved "dephlogisticated air." By the early nineteenth century, this capacity of plants was re-interpreted in terms of the "new chemistry;" in the presence of light, plants converted carbon dioxide and water into sugar and oxygen.⁽⁹⁵⁾ Quantitative analysis led to the formulation of the following chemical equation that was generally accepted as correct by the 1860's:⁽⁹⁶⁾

6CO₂ + 6H₂O C₆H₁₂O₆ + 6O₂

Variously known as "carbon fixation," "carbon assimilation," or "photosynthesis," several mechanisms for this process were proposed by plant physiologists and chemists in the late ninteenth century. These generally assumed that the decomposition of carbon dioxide was the key event, and attempted to identify a specific role for chlorophyll in the chemical reaction. A proposal in the 1860's by the German chemist Baeyer was especially influential. Baeyer's central idea was that carbon dioxide became adsorbed to chlorophyll in the presence of light. It was then supposed to be converted to formaldehyde by addition of water. Further as yet unknown chemical reactions were supposed to transform formaldehyde into sugar. An important implication of this theory was that the oxygen evolved in photosynthesis derived from the decomposition of the carbon dioxide. In various forms, the "formaldehyde theory" dominated thinking about the chemistry of photosynthesis for more than half a century.⁽⁹⁷⁾

In 1913, after years of research, the German chemists Willstätter and Stoll published the chemical structure of chlorophyll. They showed that it consisted of four pyroll groups united into a porphyrin ring binding a central magnesium atom. Five years later, they proposed a mechanism for the assimilation of carbon dioxide. Like Baeyer forty years earlier, they proposed that carbon dioxide became adsorbed to chlorophyll in the presence of light, and was then converted to formaldehyde by addition of water. Enzymes were supposed to breakdown the chorophyll-formaldehyde complex and catalyze condensation reactions that could in theory generate almost any carbohydrate. Despite the paucity of experimental evidence for it, this mechanism was widely accepted. The prestige of German chemistry compensated for the lack of experimental results.⁽⁹⁸⁾

Investigators in Germany continued to dominate the field into the 1920's, by which time "photosynthesis" had become the customary term of art. The biochemist-physiologist Otto Warburg (1883-1970) made influential practical and theoretical contributions to the field in the 1920's. He introduced a new technique, manometric measurements, and a new organism to study, the unicellular green alga Chlorella. The manometric method of measuring gas exchange made it possible to measure the rate of photosynthesis over a time range of minutes rather than hours. Biochemical analysis of the algae was much easier than of whole plants.⁽⁹⁹⁾

Although his initial studies on the problem were biochemical, Warburg soon made a sharp biophysical turn. Warburg formulated a problem that attracted the attention of both biologists and physicists, the determination of the quantum efficiency of photosynthesis. In 1922, he posed the simple question: How many quanta of light were needed to produced one molecule of oxygen in photosynthesis? Warburg gave a simple answer: Four.⁽¹⁰⁰⁾ Many researchers interpreted Warburg's number in terms of Willstätter and Stoll's theory of the chlorophyll-formaldehyde complex. Warburg proposed his own scheme. He proposed that carbon dioxide was taken up chemically and then transferred to a hypothetical acceptor activated by light. According to his scheme, too, the oxygen evolved in photosynthesis derived from the decomposition of carbon dioxide.⁽¹⁰¹⁾ Warburg became deeply attached to this idea and never relinquished it, even after three decades of research disproved it later.⁽¹⁰²⁾

In the 1920's, the chemist and plant physiologist Hermann Spoehr (1885-1954) became one of the first Americans to make photosynthesis a central research topic. The son of a Chicago candy manufacturer, Spoehr became interested in the biochemical formation of sugar in plants. In 1926, he published a major monograph to draw attention to the urgent need for rigorous experimental work on the chemistry of photosynthesis. Desperately worried about a "profligate and squandrous" civilization recklessly wasting coal and oil, or "man's patrimony," Spoehr considered photosynthesis research to be crucial for developing solar power to secure the future for human civilization.⁽¹⁰³⁾ In 1928, he became head of a newly-formed Division of Plant Biology in a laboratory funded by the Carnegie Foundation as part of its Carnegie Institutes of Washington. This laboratory moved to the Stanford campus in 1929.⁽¹⁰⁴⁾

A central objective of the chemical analysis of photosynthesis was to identify all of the chemical reactions by which carbon dioxide and water were converted into sugar and oxygen. Despite the simplicity of the overall equation for photosynthesis, almost nothing about its reaction mechanism had been established definitively by the mid-1920's. As Spoehr pointed out, the chemical aspects of photosynthesis were both intriguing and difficult to study, a situation that invited undisciplined theorizing. In his monograph, he castigated chemists, especially Willstätter and Stoll, for proposing untestable theories of the mechanism of photosynthesis. Railing against speculative theorizing, Spoehr sounded a clarion call for careful experimental work.

Oxygen evolution was the easiest aspect of photosynthesis to measure experimentally. Investigators equated the rate of oxygen evolution with the rate of photosynthesis.⁽¹⁰⁵⁾ As a result, oxygen evolution became defining feature of photosynthesis. An obvious and important question, then, was to determine the source of oxygen evolved in photosynthesis. Did it derive from carbon dioxide or water? The simplest and most obvious hypothesis proposed that the carbon dioxide was split into carbon and oxygen. The carbon would then be bound to the water, forming glucose. The remaining oxygen would be released. As Spoehr recorded, however, convincing experimental evidence for this attractive hypothesis was still lacking in the 1920's. This lack did not prevent wide acceptance of this apparently plausible idea.

Photosynthesis research was carried out with almost no contact with bacterial physiology. It was generally assumed that photosynthesis was the exclusive prerogative of the green plants and their "primitive" relatives, the algae, mosses, and lichens. Although bacteria were generally classified within the plant kingdom, they were considered to be too primitive to carry out photosynthesis. Because the study of bacteria occurred predominantly within the domain of medical bacteriology, it was isolated from the major branches of the life sciences. This isolation was especially stark in the case of photosynthesis. In Spoehr's authoritative monograph, bacteria appear only as diagnostic tools, not as organisms to be investigated. The same is true of the British counterpart, Photosynthesis: The Assimilation of Carbon by Green Plants, by Walter Stiles. The sub-title illustrates the point that photosynthesis was assumed to be a function only of green plants.⁽¹⁰⁶⁾ The separation of photosynthesis research from bacterial physiology was mirrored on the bacteriological side. For example. Marjory Stephenson's Bacterial Metabolism (1930), one of the first authoritative monographs on bacterial physiology, made no mention of photosynthesis at all.

Bacteria entered the domain of photosynthesis research as diagnostic tools. In the 1880's, the German botanist Englemann relied on the motility of bacteria to indicate the occurrence of oxygen in his now famous studies on photosynthesis in green algae. Similarly, Beijerinck used luminous bacteria as indicators of the presence of oxygen. ⁽¹⁰⁷⁾ During his studies, Englemann noticed that purple bacteria migrated toward light and gathered along wavelengths matching their absorption spectra. In 1883, he suggested that these bacteria could carry out the light-dependent assimilation of carbon. Assuming the process would occur as in green plants, Englemann reported in 1888 that purple bacteria produced oxygen in the light. No other researchers could confirm this finding, casting serious doubt on the idea of bacterial photosynthesis.

A second observation implied that the purple bacteria did not really carry out photosynthesis. They required H₂S to live. This fact implied that the organisms were chemosynthetic, not photosynthetic. This conclusion was based on a discovery, quite astonishing at the time, made by the Russian microbiologist Sergei Winogradsky in the 1880's. He found that some bacteria could obtain all of their energy from simple inorganic compounds. They could grow in the dark with CO₂ as their only carbon source, while they obtained energy by oxidizing H₂S, ultimately to sulfuric acid (H₂SO₄). This mode of metabolic life came to be called "chemosynthesis," in contrast to photosynthesis.⁽¹⁰⁸⁾ If the purple bacteria could use H₂S for energy, as seemed to be the case, then on energetic grounds they had no need for photosynthesis.

Nonetheless, the question was not resolved. Scattered reports on the subject occurred at irregular intervals into the 1920's. There was no sustained work on the subject, or real controversy, despite sharp disagreements. The investigations were usually carried out by botanists or plant physiologists, not bacteriologists or microbiologists. Two German investigators reopened the question of the seemingly paradoxical requirements of the purple bacteria.⁽¹⁰⁹⁾ Van Niel carefully studied these works, published in 1919 and 1924, and found a fatal flaw in their hypotheses. These investigators proposed that chemosynthesis and photosynthesis exist side-by-side in the purple bacteria. The idea was that the bacteria produced oxygen through photosynthesis, but the oxygen was usually not detectable because it was immediately taken up to oxidize the H₂S to H₂SO₄. This suggestion seemed to explain all of the observations -- except, van Niel noticed, for one thing. If the purple bacteria really could carry out oxygenic photosynthesis, they should be able to live in the absence of H₂S. This was not the case. Van Niel believed that understanding the metabolism of the purple bacteria required accounting for the fact that they required both light and H₂S to live.⁽¹¹⁰⁾

He was also convinced that understanding this problem would have wider implications than were obvious at first sight. Committed to the ideal of the unity of life, van Niel assumed that a seemingly specialized problem in the metabolism of an obscure group of organisms could yield results of general importance. He had the technical background, the intellectual disposition, the institutional support, and the freedom of action to investigate the problem. Taking full advantage of these circumstances, he made photosynthesis in bacteria the focus of his first research program.

F. Van Niel's General Equation

Van Niel's basic approach to the problem closely resembled his dissertation research. He set out to survey the occurrence in nature of a specific group of bacteria, then undertake systematic morphological and physiological studies, and integrate the new information into a more general framework of biochemistry and microbiology. His training at Delft strongly influenced how he did his research. At the HMS, however, he was unconstrained by direct practical concerns in his choice of problem. He chose the topic because it was intellectually interesting and aesthetically appealing. It was also likely to be tractable with the kind of training and techniques he had available.

Van Niel conceived of the problem in terms of the physiology of bacteria in their natural environments. He wanted to understand what biochemical processes provided matter and energy for their basic metabolic reactions in relation to their lives in nature. For van Niel, the organisms were not (or perhaps, not yet) tools for investigating the behavior of molecules. He undertook biochemical analysis to understand the behavior of the organisms.

At the marine station, it was easy for van Niel to combine the sensibilities of a naturalist and experimentalist. He framed his study in terms of general ecological questions: How are these organisms distributed in nature? Where are they found in abundance, where are they rare, and why? What ecological role do they play? He would then turn to a detailed study of their physiology and morphology. Observation, description, and classification were as important to van Niel's investigation as experiment and analysis.

For van Niel, however, collecting bacteria from their natural environment was not an uninteresting distraction necessary to produce an object to study. In contrast, this process was a crucial part of the investigation itself. It provided valuable information about the distribution of microorganisms in nature and their ecological relationships.

Van Niel took samples of soil and water from gardens, beaches, canals, ditches, lakes, and the salt-flats of the San Francisco Bay. Some of his samples came from Holland, others from various places on the Monterey Peninsula.

Following procedures developed by Beijerinck, van Niel prepared "enrichment cultures" from the samples he collected. Material from a sample was placed in a series of culture flasks, each containing a different chemically defined medium. The flasks were then left for a few days, in the light or dark, and either with or without access to air, to see if anything would grow. Van Niel then examined samples from these cultures to see what kinds of microorganisms had appeared. He noted that organisms appeared in different proportions in different kinds of growth media. He concluded that organisms reproduce at different rates in different media. Like Beijerinck, Van Niel interpreted this fact in evolutionary terms. He considered enrichment cultures to be a direct experimental demonstration of natural selection.

The next step in van Niel's approach was to obtain pure cultures from the enrichment cultures. Since the nineteenth century, the preparation of pure cultures had been a major activity for microbiologists. There were some dissenters from the cult of pure cultures, but most microbiologists and bacteriologists accepted them as necessary for biochemical studies.

Van Niel emphasized that a culture was pure only if it derived from one single cell. Because the organisms were so small, this was more easily advocated than accomplished. He went to great lengths to ensure he obtained a pure culture and was a sharp critic of the dubious procedures of others. The tiny green bacteria were particularly difficult to separate from their larger relatives, the purple bacteria, among the cell walls of which they could easily be hidden. The standard procedure was to begin by determining the density of cells in a liquid culture. This could be achieved by counting the number of bacterial cells in a sample on a special glass slide that held a specific volume and was engraved with a grid pattern, easily visible in the light microscope. The culture would then be diluted to a density of one cell per milliliter. Then, a sample from this liquid culture would be spread out over a plate of agar-agar, and incubated under defined conditions of light, temperature, and oxygen availability until colonies appeared on the surface. The presumption was that each colony derived from a single cell. A single colony was then used to prepare a new liquid culture. The entire process was then repeated. Van Niel believed that repeating this process several times ensured that the resulting culture derived exclusively from one cell. Careful technique throughout was crucial for avoiding contamination by other micro-organisms, or the mixing of more than one cell in the starter colony.

Once van Niel had prepared a pure culture, he began the next phase of investigation: a careful study by light microscopy of the morphology of the bacteria. He made systematic observations of the effects of the age of the culture, the pH of the medium, and the concentration of Na₂S on the size and shape of the bacterial cells. Making simple drawings of his observations, he recorded substantial morphological variation in his cultures.

At this point, van Niel was very concerned with the problem of bacterial variation. One of the key debates among late nineteenth century bacteriologists concerned the stability and variability of bacteria. Pleomorphists argued that bacteria were infinitely variable and did not exist as stable species. Monomorphists claimed that bacteria were, in fact, well-behaved, stable organisms. By 1900, the controversy had been decisively won by the monomorphists.⁽¹¹¹⁾ In the 1920's, however, the pleomorphist position was revived with new claims that bacteria undergo complex life cycles.⁽¹¹²⁾

Van Niel observed a variety of forms in a culture he was certain derived from a single cell. However, he was reluctant to interpret these forms as different stages in a presumed life cycle and was careful to distance himself from the "new pleomorphists." At the same time, he chided the dogmatism of the absolute monomorphists. This view, he said, had led to the ostensible identification of a profusion of new species of bacteria. Some of these alleged new species, van Niel argued, were variant forms of one species. Van Niel suggested that his observations pointed "toward another possibility, namely, the change of form with change in the environmental conditions."⁽¹¹³⁾ This conclusion expressed a theme van Niel returned to often, the flexibility and adaptability of living things, especially microorganisms.

He then turned to the question of their physiology and the baffling facts of their metabolism. A simple technical change accelerated the rate of work. Following a suggestion of Kluyver, van Niel found that the bacteria grew more rapidly if illuminated continuously rather than under a more natural regime of periods of light followed by darkness. This small change meant that enough material for experiments could be available within a few days rather than in weeks.

The purple bacteria were especially intriguing because their metabolism appeared paradoxical. The problem was to explain why these organism required both H₂S and light to live, when either by itself was sufficient for other organisms. He began testing the proposed explanations. His experimental approach was to make quantitative determinations of the organisms' starting materials and products. He showed that the amount of CO₂ produced was directly proportional to the amount of H₂S which disappeared and to the amount of elemental sulfur produced.

In the summer of 1929, van Niel interrupted his laboratory work for a trip to Yellowstone National Park. There, van Niel could observe microbial ecology in the context of the dramatic landscapes of the American West. At the Mammoth Hot Springs, van Niel observed striking displays of color accompanying the flowing and bubbling hot sulfurous water. Based on the colors he saw, he predicted that three groups of microbes lived in the hot springs, including the purple bacteria. When he collected samples the next day, he found no purple bacteria at all; only blue-green algae were present. Mystified, van Niel took samples of the water back to the laboratory for later study. Years later, students of van Niel showed that the lack of fixed nitrogen in the hot springs accounted for the absence of purple bacteria, and the abundance of blue-green algae. The latter are capable of converting atmospheric nitrogen to biologically usable form, while the former are not.⁽¹¹⁴⁾ In December 1929, the Western Society of Naturalists held its winter meeting in Pacific Grove. Organized by the enterprising and indefatigable Taylor, the conference assembled many of the prominent researchers from the California universities. Traditional and modern research approaches were represented. The conference included six papers on oceanography, four on permeability and the ionic relations of cells, three papers on early development, two on marine algae, and three on photosynthesis. One paper, by Berkeley protozoologist Charles Kofoid, concerned the taxonomy and evolution of pelagic ciliates. Becking reported on quantitative biophysical studies on a species of green algae capable of living in highly concentrated salt solutions.⁽¹¹⁵⁾

In his paper at the conference, van Niel proposed an important new interpretation of photosynthesis, based on experimental results he had just recently obtained and some from his years at Delft. Van Niel provided compelling evidence that bacteria could carry out a bona fide photosynthesis, if this process were understood in terms of oxidation-reduction reactions. Van Niel's experimental work showed that the green and purple bacteria carried out light dependent reactions, according to either this equation:

CO₂ + 2H₂S (CH₂O) + 2S

or, this equation:

2CO₂ + H₂S + 4H₂O 2(CH₂O) + H₂SO₄

Van Niel wrote these equations this way to emphasize their resemblance to the standard equation for green plant photosynthesis:

CO₂ + H₂O (CH₂O) + O₂

Recognizing the striking parallels between the bacterial and plant equations, van Niel formulated a new equation for photosynthesis in a more general form:

CO₂ + 2H₂A (CH₂O) + H₂O + 2Awhere "A" represented a general hydrogen acceptor.⁽¹¹⁶⁾

Van Niel's equation had important implications for understanding photosynthesis. First, it showed that oxygen evolution was not the sine qua non of photosynthesis. Viewed in the light (as it were) of van Niel's interpretation, green plant photosynthesis was a special case of a more general process. Second, it demonstrated that the key chemical event in photosynthesis could be understood as an oxidation-reduction reaction. That is, van Niel's equations implied that the basic biochemical event was the transfer of hydrogen atoms from a donor to CO₂ thereby producing glucose. He concluded that H₂0 and H₂S were simply sources of hydrogen atoms in plants and bacteria respectively for the reduction of CO₂ . In bacteria, sulfur or sulfate was left over; in plants, oxygen was the by-product.

Van Niel's formulation entered the long-standing argument over the source of oxygen in photosynthesis. It implied that the oxygen evolved in photosynthesis derived exclusively from the water and not from the carbon dioxide.

In making this formulation, van Niel combined a direct application of Kluyver's unified theory of biochemistry as based on hydrogen transfer reactions, Kluyver's "comparative biochemical" approach, and his own original research results.

Van Niel's conclusion made clear that his purpose in studying bacteria was to arrive at fundamental principle:

And here, especially lies the importance of the study of these "abnormal" photosynthetic processes, because a comparison of the factors and conditions which are required for their accomplishment will enable us to find those characteristics which are common to all. It will then be possible to derive the fundamental laws underlying all photosynthetic processes and to correlate these into a general view.⁽¹¹⁷⁾

The general equation reinforced the view, to which van Niel was already committed, that the broadest and therefore most meaningful generalizations in biology could be ascertained only through the study of both the microscopic and macroscopic worlds. In this case, the underlying unity of photosynthesis was revealed only by studying bacteria.

In 1932, van Niel compiled the results of the research he had carried out the previous three years into a major 112-page article.⁽¹¹⁸⁾ Its title, "On the Morphology and Physiology of the Purple and Green Sulfur Bacteria," makes clear that he conceived this research fundamentally as a contribution to general microbiology, not to the biochemistry of photosynthesis. In other words, van Niel investigated photosynthesis to understand the physiology of bacteria, not the other way around. Van Niel's work made a crucial link between the physiology of bacteria and higher plants and opened a new line of investigation in photosynthesis. It brought considerable attention to the value of working with bacteria for biological studies. It addressed a diverse audience of scientists, including chemists, biophysicists, plant physiologists, and microbiologists.

Van Niel's study of the purple bacteria made bacterial photosynthesis into a visible phenomenon and into a new research topic. British biochemist Stephenson, for example, who had made no mention of photosynthesis in the first edition of Bacterial Metabolism, in 1929 gave it a prominent place in the second edition. She called the discovery of bacterial photosynthesis "an event in bacterial chemistry of first-class biological importance." In her assessment, van Niel's demonstration that bacteria, like plants, could use light to reduce carbon dioxide to sugar amounted to "the revelation of a new mode of life" that "ranks with Winogradsky's discovery of chemosynthetic organisms at the end of the last century."⁽¹¹⁹⁾ Stephenson wrote a whole new chapter on bacterial photosynthesis and extolled van Niel's contribution. "The task of ... disclosing the true character of the photosynthetic bacteria was reserved for van Niel, to whose masterly researches and penetrating insight we owe our present knowledge on this subject."⁽¹²⁰⁾

Others followed his example. In 1931, Kluyver himself took up the subject of bacterial photosynthesis, in part because of van Niel's success.⁽¹²¹⁾ He launched a new collaborative project with Lenard Ornstein and the Physics Laboratory at the University of Utrecht. The project was funded by the RF, which had made the promotion of joint projects between physical and biological scientists an explicit policy in the 1930's. As historian Doris Zallen has shown, the collaboration between Kluyver's biological group at Delft and Ornstein's physics group at Utrecht was cordial and productive for decades.⁽¹²²⁾

Despite the great geographic distance between them, van Niel and Kluyver remained in close contact through frequent correspondence. They discussed the research in their respective laboratories in detail and attempted to arrange for exchanges, though the economic situation in the early 1930's made this difficult. While on a brief American tour in 1932, Kluyver was able to visit van Niel and the Jacques Loeb Laboratory. F.M. Muller, technically a student at the University of Utrecht, came to van Niel's laboratory in the early 1930's to conduct experiments on the metabolism of the purple bacteria in organic media.⁽¹²³⁾ P.A. Roelefson, another student officially enrolled at Utrecht, worked closely with Kluyver on photosynthesis of the Thiorhodaceae. Van Niel was consulted about his work throughout.⁽¹²⁴⁾

By 1936, four more laboratories had begun research on bacterial photosynthesis. In a review of another proposal to research the subject made to the RF, Kluyver vigorously defended van Niel's leadership of the field. "The outstanding characteristic of the [proposed] programme," he wrote to RF officer Tisdale, "is its nearly complete lack of originality." He emphasized that van Niel's innovative research on the topic began during "an intimate cooperation" with Kluyver from 1925 to 1927.⁽¹²⁵⁾

Closer to home, van Niel maintained a good working relationship with Spoehr. After an undistinguished year as director of the Natural Sciences Division of the RF, Spoehr returned to California in 1931 to resume his position as chairman of the Division of Plant Biology of the Carnegie Institute of Washington. With Spoehr's laboratory located adjacent to the Stanford campus, it was relatively easy for Spoehr and van Niel to arrange collaborative research. In the springs of 1933 and 1934, van Niel worked frequently in Spoehr's laboratory.⁽¹²⁶⁾ With J.H.C. Smith, an associate of Spoehr, van Niel worked on the isolation and chemical analysis of the pigments in the puple bacteria. These studies showed that the light absorbing molecules in photosynthetic bacteria were different from those in green plants. These differences did not, however, undermine van Niel's basic claim that the fundamental chemical reaction of photosynthesis in plants and bacteria could be expressed in a single general equation.

G. A School for Biology and Success for van Niel

The promising beginning of the Jacques Loeb Laboratory and van Niel 's position there were endangered almost immediately by the economic crisis of the early 1930's which hit Stanford hard. Expensive programs like experimental research were especially vulnerable to the decrease in available funds. Two researchers who left the marine station because they were not promoted were not immediately replaced.⁽¹²⁷⁾ Fisher was not at all displeased to see Mestre, one of Becking's selections, leave for a teaching position, but the departures nearly depleted the Jacques Loeb Laboratory.⁽¹²⁸⁾ Between 1933 and 1935, van Niel and a postdoctoral fellow were the only full-time researchers there.⁽¹²⁹⁾ Van Niel even began to worry that the entire laboratory might be closed. He confided to Kluyver that he might even be reduced to applying for a position at the once-scorned Gist-en-Spiritus Fabrik in Delft. Van Niel was also forced to discourage a promising student of Kluyver from coming to California because it would be impossible to buy the equipment necessary for the proposed study on the quantum efficiency of photosynthesis in purple bacteria.⁽¹³⁰⁾

One beneficial outcome of the economic crisis from the point of view of the Stanford campus was that Wilbur resumed his presidency on a full-time basis in 1933. Between 1929 and 1933, Wilbur had served as Secretary of the Interior under President Herbert Hoover, a close friend since their undergraduate days at Stanford. Robert E. Swain, Acting President during this period, consulted with Wilbur regularly, but there was considerable criticism of the official president's prolonged absence from the campus. The defeat of Hoover in the presidential election of 1932 brought Wilbur's tenure in Washington to an end.

On his return to Stanford, Wilbur found a dire budget situation. He was forced to make faculty pay cuts, tuition increases, and urgent appeals to private donors.⁽¹³¹⁾ The situation was desperate enough that Wilbur resorted to less conventional tactics. At the end of 1933, he simply took over the balances of the solvent departments, like the marine station, for the general funds of the University. Fisher was incensed by this maneuver, but powerless to stop it.⁽¹³²⁾

Despite the economic problems, Wilbur and Taylor continued to make strong appointments in experimental biology. They took advantage of a large number of retirements in the 1930's and the dire employment situation to obtain top-notch researchers. In 1932, Victor C. Twitty, an experimental embryologist, joined the still extant but weakening Zoology Department.⁽¹³³⁾ Douglas M. Whitaker, a Ph.D. student under Taylor and therefore an avid experimentalist, was appointed to the faculty in 1932. Whitaker replaced Heath, who had studied embryology from the morphological viewpoint. Whitaker continued work on the micromanipulation of protozoa and began research in experimental embryology. His first studies in this area investigated the effects of temperature and oxygen concentration on the development of polarity in the animal egg.⁽¹³⁴⁾ In 1933, Leonard Blinks, a plant physiologist trained at the Rockefeller Institute, became associate professor of botany, replacing outgoing George J. Pierce. Blinks confessed to feeling some dismay at leaving the Rockefeller Institute for "a well-known but financially straitened university ... on a large farm," but considered himself fortunate to have found any job in the depths of the Depression.⁽¹³⁵⁾ Assigned to the Stanford campus, Blinks spent most of his summers at the HMS. The equipment for physiological experiments available there was vastly superior to the facilities in the Botany Department. The laboratory he inherited from Pierce had only one "antique" balance, and a single spectroscope.⁽¹³⁶⁾ When Blinks attempted to take equipment from the marine station to the main campus, he incurred the wrath of Fisher.⁽¹³⁷⁾ Blink's specialty in electrical phenomena and algae suited the direction of research at Stanford. He became a productive and valued member of the team of experimentalists.

Wilbur's return in 1933 meant renewed efforts to establish the incipient School of Biology. With the support of Taylor, Fisher and other faculty sympathetic to the project, Wilbur gradually transferred power over courses, degrees, students, and budgets from the old departments to the School. In 1934, he struck the fatal blow to his adversaries, weakened by steps during the previous decade. After forty-three years on the Stanford campus, the departments of Botany and Zoology ceased to exist, absorbed completely into the new School of Biological Sciences. The status of the former pre-medical departments was less clear. While officially part of the new School, Anatomy, Bacteriology, and Physiology were entitled to one representative each to the executive committee of the School. The HMS continued to exist as an autonomous unit within the School of Biology, as did the Natural History Museum.⁽¹³⁸⁾ Twelve years in the making, the School of Biology with Taylor as its Chairman assumed authority over the life sciences at Stanford. "Congratulations on a chemical miracle," Fisher wrote to Wilbur, "i.e., changing a crystalloid (School of Biology) to a colloidal sol. We shall now get somewhere."⁽¹³⁹⁾

Taylor was quick to take advantage of the new organization and his position of authority. Within the School of Biology, he assembled the most productive experimentalists into a research group to pursue common interests, including funding. While each investigator pursued independent projects, all could be construed as "chemophysical biology," a priority for RF funding in the 1930's. Blinks investigated bio-electric phenomena in plant and algal cells. The focus of Twitty's work in experimental embryology was the expression of larval pigmentation in the salamander. Whitaker studied early development in the egg of the marine alga, Fucus. Taylor investigated the effects of X-rays and other agents on free-swimming and encysted Colpoda. Van Niel 's work was the most biochemical. He continued to investigate the metabolism of the purple and green bacteria, especially their photosynthetic processes. In 1934, the RF awarded Taylor $10,000 to direct research on "the effects of radiation and other chemophysical agents on unicellular organisms." A year later, an RF report called Taylor's team "a brilliant young group of investigators all of whom are working completely within the spirit and range of our program interests." The group was awarded a second grant of $12,500.⁽¹⁴⁰⁾

This success encouraged Taylor to make more requests for funds, especially for increasingly complex equipment. Taylor detailed to Wilbur the items "most needed" by the experimentalists: constant temperature cold rooms for the HMS, a spectrophotometer for Blinks and van Niel, respiration equipment for Twitty, an ultracentrifuge for the group as a whole, and ultraviolet equipment, including a quartz microscope, for Giese and Whitaker. Blinks and van Niel were also hoping for a monochromator and a cathode ray oscilloscope.⁽¹⁴¹⁾ The funding from the RF and the investment in complex equipment gave Taylor and colleagues tangible support for their conviction that experimental biology represented the proper direction of research in the life sciences. Increases in capital meant increasing power and it helped solidify the School of Biology.

While Fisher was an enthusiastic supporter of the School of Biology and expansion of experimentation, he continued to encourage more traditional lines of investigation at the marine station as well. Oceanographic surveys and studies of fish continued to occupy an important place among the marine station's activities. Bolin studied the systematics, morphology, embryology, and ecology of cottoid fishes, while Fisher specialized in hydrocorals and sea stars. Skogsberg continued the hydrobiological survey of the Monterey Bay. Visiting investigators carried out all sorts of projects, some directly related to marine biology, and others distinctly not.⁽¹⁴²⁾ Life at the station was lively, fun, and interesting. In the fall of 1934 Fisher reported to Wilbur, 'The Summer Session is one of the best we have ever had. There is a full house and everyone seems to be happy."⁽¹⁴³⁾

Van Niel distinguished himself among the marine station's researchers. His research on photosynthesis propelled him into the first rank of a field that continued throughout the 1930's to be a dynamic research frontier, squarely within the bounds of "chemophysical biology." His research began to attract the attention of prominent institutions. A invitation from Harvard to give a course in general microbiology alarmed the Stanford administration. When it became clear that Harvard was seriously attempting to recruit van Niel, Stanford immediately began the process of promoting him to full professor.

When Taylor asked for assessments of van Niel, everyone at the marine station regardless of research interests praised the microbiologist in the highest terms. Skogsberg, the resident oceanographer, found van Niel 's outlook stimulating but critical, and considered it a "privilege" to work with him.⁽¹⁴⁴⁾ Whitaker was even more laudatory. "He is a man of very outstanding brilliance," he wrote to Taylor. "I have rarely encountered so logical and penetrating a mind, and one which is at the same time so objectively self critical, honest, and generous." On numerous occasions, Whitaker said, faculty members including himself and graduate students in various fields of biology consulted van Niel. "In all these cases," continued Whitaker, "he has been a veritable warehouse of information and ingenuity, delivered with truly remarkable powers of clear exposition. His teaching is as excellent as his research." Further, Whitaker judged that van Niel 's field of interest was rapidly increasing in importance and that it related in fundamental ways to a wide variety of fields in biological sciences, particularly those on the borderline between biology and the physical sciences. He told Taylor that it was well known that van Niel was held in very high esteem by "our research neighbor in the south, Caltech" as well as by Harvard.⁽¹⁴⁵⁾ Weymouth and Blinks concurred and gave their own highly laudatory reviews of their microbiologist colleague.⁽¹⁴⁶⁾ The result was never in doubt. In March of 1935, van Niel was promoted to full professor.⁽¹⁴⁷⁾ Fisher reported to Wilbur that van Niel was exuberant. There were no further discussions of a move to Harvard.⁽¹⁴⁸⁾

After his first six years at Stanford, van Niel had achieved a considerable scientific reputation, not only in the United States, but internationally. As he informed Wilbur, van Niel was invited to speak at the most important scientific meetings in his area of research. In 1935, he accepted invitations to speak at the symposium on Quantitative Biology at Cold Spring Harbor, the International Botanical Congress, and the Second International Congress for Microbiology.⁽¹⁴⁹⁾

Wilbur, too, had advanced his projects in the early 1930's, despite the economic problems facing the university. By 1935, Wilbur's ambition for a School for Biology had become a reality. At the same time, the departments affiliated with the Medical School continued to resist full integration and frequent squabbles over curriculum and courses broke out.⁽¹⁵⁰⁾ Nonetheless, a small but productive group of experimental biologists became well established within the School. Continued financial support from the RF for their research appeared likely.

For Taylor, too, establishing the School of Biology was a victory. The victor missed a crown, however, and Taylor advised his president that he would "be willing" to be named Dean of the School if that would help ensure its permanence.⁽¹⁵¹⁾ In 1936, he received the word. "You are hereby appointed Dean of the School of Biological Sciences," wrote Wilbur.⁽¹⁵²⁾

While the School of Biology represented success for Wilbur, the HMS did not live up to the potential envisioned for it in 1928. It remained a small and quiet institution, never gaining the stature or size of the MBL in Woods Hole or the Zoological Station in Naples. Still, important research was carried out in the Jacques Loeb Laboratory. For Wilbur and Taylor, the new laboratory provided a highly valuable place in which to develop experimental biology free from the constraints of the established departments on the campus where resistance to the new biology was strong. The HMS advanced Wilbur's goals to reform medicine and to build Stanford into a major research university. As an RF officer, Embree advanced his own interests and the larger goals of the RF by funding the HMS. Fisher more than doubled the size of the institution under his direction by joining in the support of experimental biology. Stanford faculty Blinks, Taylor, and Whitaker launched sustained research programs that relied heavily on biophysical and experimental physiological approaches. In summers, a whole host of investigators and students arrived at the Station and conducted a wide variety of research projects.

The Jacques Loeb Laboratory proved to be an ideal institutional setting for van Niel. There, he could pursue a broadly biological approach to microbiology and conduct his path-breaking work on photosynthesis in bacteria. His highly original work took what appeared to be an obscure problem and transformed it into a new and productive research topic, bacterial photosynthesis. His research brought together subjects that had previously been divided by disciplinary boundaries. By studying what seemed to be an idiosyncratic process, he achieved an important insight into a fundamental life process. His work made possible the formulation of new experimentally tractable problems. In the process, he brought new attention to the utility of bacteria as experimental objects for fundamental research in biology. It allowed van Niel to formulate a general equation that subsumed both plant and bacterial photosynthesis.

The specific characteristics of this small institution, in concert with van Niel's intellectual outlook and the more general purposes of both Stanford University and the RF permitted the process of scientific innovation and expansion to occur. Van Niel's successful research validated many of the principles upon which the heavy investment in experimental biology at Stanford was made. It demonstrated the practical value of physics and chemistry for investigating biological problems. Because his work revealed fundamental similarities between bacteria and plants, it validated the principle of "the unity of life" and the concept of "biology" as a meaningful scientific and institutional category. To explain why the unity of life gained ascendance over the exploration life's diversity for life scientists in the 1920's and 1930's remains an important issue for future study.

The research of van Niel and colleagues, the expansion of experimentation, and the establishment of the School of Biology were mutually supportive endeavors in the 1930's. The Jacques Loeb Laboratory at the HMS provided an institutional basis for the consolidation of biology and the expansion of experimentation at Stanford.

The marine station was an ideal environment for van Niel in another important sense. The specific configuration of institutional location and source of financial support permitted van Niel to adhere with little compromise to the ideal of pure science. Ensconced with a small group of researchers in a beautiful and remote setting, van Niel could conceive of science as a spiritual pursuit to advance the rational comprehension of the natural world. Neither curing disease nor the production of useful chemicals could compare with that magnificent project.

Index | Previous | Next

1. C.B. van Niel, "The Education of a Microbiologist: Some Reflections," Annual Review of Microbiology, 21 (1967), 1-30.

2. My argument contrasts with that made by historian Lily E. Kay in her study of the history of molecular biology. Kay argues that an agenda of "social control" advocated by conservative American elites associated with the RF was either explicitly or tacitly shared by influential scientists. In her view, this alleged shared vision accounts for the dominance of experimental biology based on chemistry and physics in American life sciences in this century. In contrast, the approach I have taken here reveals the diversity of interests that were served by the expansion of experimental biology at one specific institution. Cf. Lily E. Kay, The Molecular Vision of Life: Caltech, The Rockefeller Foundation and the Rise of the New Biology (New York: Oxford University Press, 1993), especially p. 3 and pp. 6-11.

3. Laurence R. Vesey, The Emergence of the American University (Chicago: University of Chicago Press, 1965) pp. 104-08; Percy Baumberger, "A History of Biology at Stanford University," Bios, 25 (1954), pp. 123-26.

4. Baumberger, "Biology at Stanford University," (1954), pp. 123-25; David Magnus, In Defense of Natural History: David Starr Jordan and the Role of Isolation in Evolution (Dissertation: Stanford University, 1993), pp. 48-58; Jane Maienschein, Transforming Traditions in American Biology, 1880-1915 (Baltimore: Johns Hopkins University Press, 1991), especially pp. 47-50.

5. Bailey Mallard, "Jordan of Stanford," L.A. Times Sunday Magazine, January 21, 1934.

6. On Jordan's disinterest in philosophy, see Veysey, Emergence of the American University, (1965), p. 80.

7. O.P. Jenkins, "The Hopkins Seaside Laboratory," Zoe, 4 (1893), 58-63.

8. See Philip J. Pauly, "Summer Resort and Scientific Discipline: Woods Hole and the Structure of American Biology, 1882-1925," in Ron Rainger, Keith R. Benson and Jane Maienschein, eds., The American Development of Biology (Philadelphia: University of Pennsylvania Press, 1988), 121-150 for interesting reflections on this suggestion.

9. Charles Atwood Kofoid, The Biological Stations of Europe (Washington, D.C.: Government Printing Office, 1910), pp. 35-143.

10. Ibid., pp. 144-217, 273-316.

11. Ibid., pp. 7-32; Keith R. Benson, "The Naples Stazione Zoologica and its Impact on the Emergence of American Marine Biology," Journal of the History of Biology, 21 (1988), 331-341.

12. On the marine laboratory directed by W. K. Brooks for Johns Hopkins, see Maienschein, Transforming Traditions, (1991) p. 50.

13. See, e.g., Jane Maienschein, "Agassiz, Hyatt, Whitman, and the Birth of the Marine Biological Laboratory," Biological Bulletin, 168 (1985), 26-54 ; and Maienschein, Transforming Traditions, (1991) for the history of the MBL and its importance in American biology.

14. Pauly, "Summer Resort and Scientific Discipline," (1988), pp. 121-150.

15. Ron Rainger, Keith R. Benson, and Jane Maienschein, "Introduction," in The American Development of Biology (Philadelphia: University of Pennsylvania Press, 1988).

16. W.K. Fisher, "The Hopkins Marine Station at Stanford University," The Collecting Net, 6 (1931), 65-67.

17. On the central importance of the Johns Hopkins University to American biology, see, e.g., Keith R. Benson, "American Morphology in the Late Nineteenth Century: The Biology Department at Johns Hopkins University," Journal of the History of Biology, 18 (1985) 163-205; Pauly, "Appearance of Academic Biology," (1984); and Maienschein, Transforming Traditions (1991), chapter 2.

18. Keith R. Benson, "From Teaching to Research: Marine Biological Laboratories of the Pacific Coast," paper delivered at the annual meeting of the American Association for the Advancement of Science, February 20, 1994, San Francisco, CA.

19. Edgar E. Robinson and Paul C. Edwards, eds., The Memoirs of Ray Lyman Wilbur, 1875-1949 (Stanford: Stanford University Press, 1960), p. 78.

20. Philip J. Pauly, Controlling Life: Jacques Loeb and the Engineering Ideal in Biology (Berkeley: University of California Press, 1987), pp. 99, 107-113, 135.

21. Baumberger, "Biology at Stanford," (1954), p. 128.

22. Robinson and Edwards, eds., Memoirs of Wilbur (1960), p. 66.

23. See Garland E. Allen, Life Sciences in the Twentieth Century (Cambridge: Cambridge University Press, 1975) for the original formulation of this hypothesis. See Jane Maienschein, Ronald Rainger, and Keith R. Benson, "Introduction: Were American Morphologists in Revolt?" Journal of the History of Biology, 14 (1981) 83-87 for an overview of critiques of Allen's thesis.

24. Maienschein, Transforming Traditions (1991), p. 295.

25. Robinson and Edwards, eds., Memoirs of Wilbur (1960), pp. 158-180, especially pp. 178-79.

26. See Roger L. Geiger, To Advance Knowledge: The Growth of American Research Universities, 1900-1940 (New York: Oxford University Press, 1986) on the development of American research universities generally in the period between 1900 and 1940.

27. Robinson and Edwards, eds., Memoirs of Wilbur (1960), p. 291; Robert E. Kohler, Partners in Science: Foundations and Natural Scientists (Chicago: University of Chicago Press, 1991), pp. 213-14.

28. Kohler, Partners in Science (1991), pp. 213-14.

29. Lydon, The Chinese in the Monterey Region (1985), pp. 350-55; Patrick Hathaway, personal communication, July 1996.

30. Robinson and Edwards, eds., Memoirs of Wilbur (1960), p. 45. From 1906 to 1917, the laboratory was called "The Marine Biological Laboratory of Stanford." See W.K. Fisher, "The New Hopkins Marine Station of Stanford University," Science N.S., 47 (1918), 410-13.

31. Robinson and Edwards, eds., Memoirs of Wilbur (1960), p. 290.

32. Pauly, "Appearance of Academic Biology," (1984).

33. See Robert E. Kohler, From Medical Chemistry to Biochemistry: The Making of a Biomedical Discipline (Cambridge: Cambridge University Press, 1982), Chapter 6 for discussion of the movement to reform medical education and training.

34. Wilbur to the Honorable Board of Trustees, February 10, 1922; SUT 15/11. See also Robinson and Edwards, eds., Memoirs of Wilbur (1960), pp. 290-92.

35. C.H. Danforth, "C.V. Taylor, 1885-1946," National Academy of Sciences Biographical Memoirs, 25 (1948), pp. 215-16.

36. Ibid.

37. Wilbur to the Honorable Board of Trustees, February 2, 1934; SUT 20/22.

38. Martin to Swain, March 12, 1931; RLW 75/ Biology, School of.

39. Quoted in Baumberger, "Biology at Stanford" (1954), p. 133. See also van Niel to Stanier, April 25, 1945; RYS 6/21.

40. For contrasting interpretations, see Pnina Abir-Am, "The Discourse of Physical Power and Biological Knowledge in the 1930's: A Reappraisal of the Rockefeller Foundation's 'Policy' in Molecular Biology," Social Studies of Science, 12 (1982) 341-382; Kay, Molecular Vision of Life (1993), and Kohler, Partners in Science, (1991).

41. Robinson and Edwards, eds., Memoirs of Wilbur (1960), p.577.

42. See Kohler, Partners in Science (1991), pp. 53-57 and 125-28 for discussion of Embree and his failed attempt to build a strong division of human biology at the RF.

43. Embree to Fisher, February 26, 1925; RF 1.1/200D 157/1925 RAC.

44. Fisher to Embree, March 26, 1925; RF 1.1/200D 157/1925 RAC.

45. Embree to Wilbur, April 27, 1925; RF 1.1/200D 157/1925 RAC. The initial grant was for $50,000 on a matching basis. The trustees approved the award in May, and Stanford had the funds by July 1925. See "Historical Record, Stanford University-Hopkins Marine Station, 1925-27, 1929-34," pp. 21-22, 28; RF 1.1/200D 157/1925 RAC.

46. Wilbur to O'Connor, January 29, 1926; "Historical Record, Stanford University-Hopkins Marine Station, 1925-27, 1929-34," p. 38; RF 1.1/200D 157/1925 RAC.

47. Wilbur to Embree, March 19, 1926; RF 1.1/200D 157/1925 RAC. Next to Wilbur's suggestion, a marginal note presumably written by an RF official, possibly Embree, reads, "On the contrary."

48. Wilbur to Embree, April 8, 1926; RF 1.1/200D 157/1925 RAC.

49. Wilbur to the RF, April 8, 1926; RF 1.1/200D 157/1925 RAC.

50. See Pauly, Controlling Life: Jacques Loeb (1987), chapter 8, for an interpretation of efforts to "construct" images of Loeb after his death.

51. Embree to Wilbur, May 27, 1926; RF 1.1/200D 157/1925 RAC.

52. Kohler, Partners in Science (1991), pp. 213-14.

53. Lourens G.M. Baas Becking, The Origin of the Vascular Structure of the Genus Botrychium (Dissertation, Stanford University, 1921); Radiation and Vital Phenomena (Dissertation, Rijksuniversiteit Utrecht, 1921).

54. Kohler, Partners in Science (1991), pp. 213-14; Wilbur to O'Connor, January 29, 1926; RF 1.1/200D 157/1925 RAC.

55. Fisher to Wilbur, December 18, 1928; and Becking, Fisher, MacGinitie, Skogsberg, and Taylor to Wilbur, December 21, 1928; RLW 70/HMS.

56. Kohler, Partners in Science (1991), pp. 213-14.

57. Swain to G.E. Jennings, July 25, 1929; RLW 70/Herzstein.

58. Fisher to C.L. Alsberg, August 25, 1930; RLW 76/HMS.

59. Van Niel, "Education," (1967), pp. 13-14.

60. "Van Niel of Pacific Grove," RF Trustees Confidential Report, March 1956; RF 1.2/205D 6/43 RAC.

61. Van Niel, "Education," (1967), p. 14.

62. Becking to Wilbur, October 8, 1928; RLW 70/HMS. Becking wrote that van Niel was "in charge of Beyerinck's and Kluyver's laboratories."

63. Wilbur to van Niel, October 12, 1928; RLW 70/HMS.

64. Van Niel, "Education," (1967), pp. 14-15.

65. Ibid.

66. See John Steinbeck, Cannery Row (New York: The Viking Press, 1945) for a fictional account of life in the Monterey area in the 1930's.

67. W.K. Fisher, "The Hopkins Marine Station of Stanford University," Scientific Monthly, 29 (1929), 298-303 on p. 301.

68. Bulletin of the Hopkins Marine Station (1930). HMS.

69. Becking to Swain, April 5, 1929, and attached "Memo. of Equipment, Jacques Loeb Lab;" RLW 70/HMS.

70. Becking to Swain April 5, 1929, and attached "Memo. of Equipment, Jacques Loeb Lab;" RLW 70/HMS. Lawrence R. Blinks, "A Golden Anniversary: The Hopkins Marine Station and Its Connections with L.R. Blinks (or vice versa) at Stanford Since September 1, 1933," (1983), p. 9; LRB.

71. Bulletin of the Hopkins Marine Station (1930); HMS.

72. W. K. Fisher, "The Hopkins Marine Station," (1929), pp. 298-303. Bulletin of the Hopkins Marine Station (1930), pp. 9-11; HMS.

73. Bulletin of the Hopkins Marine Station (1930), pp. 9-11; HMS.

74. Van Niel, "Education," (1967), p. 16.

75. Becking to Wilbur, February 10, 1929; RLW 70/HMS.

76. Van Niel to Kluyver, February 28, 1929; AJK.

77. Heath to Wilbur, July 10, 1929; RLW 70/HMS.

78. Van Niel, unpublished manuscripts; VNS 7/1.

79. Heath to Wilbur, July 10, 1929; RLW 70/HMS. See also Fisher to Swain, November 12, 1929; RLW 73/HMS.

80. Fisher to Wilbur, August 28, 1930; RLW 76/HMS.

81. Van Niel to Kluyver, February 28, 1929; AJK.

82. Fisher to C.L. Alsberg, August 25, 1930; RLW 76/HMS.

83. Fisher to Wilbur, Aug. 28, 1930; RLW 76/HMS.

84. Van Niel, unpublished manuscripts; VNS 7/1.

85. Becking to Swain, April 8, 1929; Fisher to E.S. Erwin, April 6, 1929; RLW 76/HMS. Blinks, "A Golden Anniversary: the HMS," (1983), p. 6; LRB; Fisher to Wilbur, Aug. 28, 1930; RLW 76/HMS.

86. Swain to Becking, March 20, 1930; Swain to Fisher, March 20, 1930; RLW 73/HMS.

87. R. N. Robertson, "Dr. L.G.M. Baas Becking," Australian Journal of Science, 26 (1963), 15-16. F.W. Went, "Dr. L.G.M. Baas Becking," Nature, 198 (1963), 134.

88. Blinks, "A Golden Anniversary: the HMS," (1983), p. 9; LRB.

89. Taylor to Wilbur, July 8, 1930; RLW 73/Biology-School of.

90. Taylor to Swain, May 13, 1931; RLW 75/Biology-School of.

91. Taylor to Swain March 27, 1931; RLW 75/Biology-School of.

92. Wilbur to Taylor, August 1, 1930; RLW 73/Biology-School of.

93. Van Niel, "Education," (1967), pp. 14-16.

94. See John E. Lesch, "Systematics and the Geometric Spirit," in Tore Frngsmyr, J.L. Heilbron, and Robin Rider, eds., The Quantifying Spirit in the Eighteenth Century (Berkeley: University of California Press, 1990), pp. 73-112 for discussion of the preoccupation of botanists with classification in the eighteenth century.

95. Julius Sachs, Lectures on the Physiology of Plants (Oxford: The Clarendon Press, 1887), pp. 296-97.

96. Walter Stiles, Photosynthesis: The Assimilation of Carbon by Green Plants (London: Longmans, Green, and Co., 1925), p. 4.

97. Herman Augustus Spoehr, Photosynthesis (New York: The American Chemical Catalogue Company, Inc., 1926), pp. 256-278.

98. Ibid., pp. 264-85.

99. Jack Myers, "Conceptual Developments in Photosynthesis, 1924-1974," Plant Physiology, 54 (1974), 420-466; Doris T. Zallen, "Redrawing the Boundaries of Molecular Biology," Journal of the History of Biology, 26 (1993), 65-87; idem., "The 'Light' Organism for the Job: Green Algae and Photosynthesis Research," Journal of the History of Biology, 26 (1993), 269-279.

100. This research is summarized in Myers, "Conceptual Developments in Photosynthesis, 1924-1974," (1974), pp. 420-26.

101. Spoehr, Photosynthesis, (1926), pp. 286-87.

102. Hans Gaffron, "Van Niel's Theory: Thirty Years After," in Howard Gest, Antonio San Pietro, and L. P. Vernon, eds. Bacterial Photosynthesis (Yellow Springs, Ohio: Antioch Press, 1963), 3-14.

103. Spoehr, Photosynthesis (1926), pp. 13-26.

104. J.H.C. Smith and C. Stacy French, "H. Spoehr," Annual Review of Biochemistry, 24 (1955), xi-xvi.

105. Spoehr, Photosynthesis (1926), pp. 84-85.

106. Stiles, Photosynthesis (1925).

107. Spoehr, Photosynthesis (1926), p. 229.

108. See C.B. van Niel, "On the Morphology and Physiology of the Purple and Green Sulphur Bacteria," Archiv fr Mikrobiologie, 3 (1931), 1-112, for discussion of chemosynthesis.

109. Reviewed in C.B. van Niel, "Photosynthesis of Bacteria," in Contributions to Marine Biology (Stanford: Stanford University Press, 1929), 161-169.

110. Ibid., p. 163.

111. See Pauline M. H. Mazumdar, Species and Specificity:An Interpretation of the History of Immunology (Cambridge: Cambridge University Press, 1995) for extensive, insightful analysis of the philosophical and political dimensions of debates about the stability of bacterial species in the context of nineteenth century German biology.

112. Olga Amsterdamska, "Stabilizing Instability: The Controversy over Cyclogenic Theories of Bacterial Variation during the Interwar Period," Journal of the History of Biology, 24 (1991), 191-222.

113. Van Niel, "On the Morphology and Physiology of the Purple and Green Sulfur Bacteria," Archiv fr Mikrobiologie, 3 (1931), p. 60.

114. Van Niel, "Education," (1967), pp. 17-18.

115. See the twenty-three papers collected in Contributions to Marine Biology, (Stanford: Stanford University Press, 1929), including Charles Atwood Kofoid, "Factors in the Evolution of the Pelagic Ciliata, the Tintinnoinea," and L.G.M. Baas Becking, "Observations on Dunaliella viridis, Teodoresco," pp. 102-114.

116. Van Niel, "Photosynthesis of Bacteria," (1929), pp. 161-169.

117. Ibid., p. 168.

118. Van Niel, "On the Morphology and Physiology of the Purple and Green Sulfur Bacteria," Archiv fr Mikrobiologie, 3 (1931), pp. 1-112.

119. Marjory Stephenson, Bacterial Metabolism (London: Longmans, Green, and Co. 1939), pp. vi-vii.

120. Ibid., p. 292.

121. Kluyver to van Niel, May 16, 1935; AJK.

122. This collaboration is analyzed in detail in Doris T. Zallen, "The Rockefeller Foundation and Spectroscopy Research: The Programs at Chicago and Utrecht," Journal of the History of Biology, 25 (1992), 67-89.

123. F.M. Muller, "On the Metabolism of the Purple Sulfur Bacteria in Organic Media," Archiv fr Mikrobiologie, 4 (1933), p. 131.

124. P.A. Roelefson, "On the Metabolism of the Purple Sulfur Bacteria," Proc. Koninkl. Akda. Wetenschap., 37 (1934), 660-69.

125. Kluyver to W.E. Tisdale, February 18, 1936; AJK.

126. Van Niel, "Education," (1967), p. 19.

127. Fisher to Swain, March 19, 1932; RLW 78/HMS; Swain to Mestre, April 18, 1932; RLW 78/HMS.

128. Fisher to Wilbur, May 29, 1933; RLW 81/HMS.

129. Van Niel, "Education," (1967), p. 24; Kluyer to van Niel, January 10, 1933; AJK.

130. Van Niel to Kluyver, February 5, 1933; AJK.

131. Robinson and Edwards, eds., Memoirs of Wilbur (1960), pp. 571-74.

132. Wilbur to Fisher, December 27, 1933; RLW 85/HMS.

133. Twitty studied under Ross G. Harrison at Yale, see his Of Scientists and Salamanders, (San Francisco: W.H. Freeman, 1966) for a retrospective account of his career at Stanford.

134. Baumberger, "Biology at Stanford," (1954), pp.141-42.

135. Blinks, "A Golden Anniversary: the HMS," (1983), p. 8; LRB.

136. Ibid., p. 8.

137. Ibid., p. 9.

138. Wilbur to Honorable Board of Trustees, February 2, 1934; SUT 20/2.

139. Fisher to Wilbur, March 8, 1934; RLW 85/HMS.

140. Natural Sciences Report, April 17, 1935; RF 1.1 205D, 8/5, RAC.

141. Taylor to Wilbur, November 27, 1935; RLW 90/Biology--School of.

142. Fisher to D.L. Fox, August 28, 1934; RLW 85/HMS.

143. Fisher to Wilbur, August 4, 1934; RLW 85/HMS.

144. Skogsberg to Taylor, November 16, 1934; RLW 88/HMS.

145. Whitaker to Taylor, November 16, 1934; RLW 88/HMS.

146. Weymouth to Taylor, November 15, 1934; Blinks to Taylor, November 12, 1934; RLW 88/HMS.

147. Van Niel to Wilbur, March 24, 1935; RLW 88/HMS.

148. Fisher to Wilbur, March 16, 1935; RLW 88/HMS; van Neil suggested that Harvard go after Kluyver and vice versa. Van Neil to Kluyver, February 10, 1935.

149. Van Niel to Wilbur, May 12, 1935; RLW 88/HMS.

150. Van Niel to Stanier, April 25, 1945; RYS 6/21.

151. Taylor to Wilbur, April 8, 1936, Taylor to Wilbur, May 15, 1936; RLW 90/ Biology - School of.

152. Wilbur to Taylor May 29, 1936; RLW 90/ Biology - School of.