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Journal of the History of Biology33: 141–180, 2000.© 2000Kluwer Academic Publishers. Printed in the Netherlands.

141

From Bacteriology to Biochemistry: Albert Jan Kluyver andChester Werkman at Iowa State

RIVERS SINGLETON, JR.Departments of Biology and EnglishUniversity of DelawareNewark, DE 19716-2590, U.S.A.E-Mail: [email protected]

Abstract. This essay explores connections between bacteriology and the disciplinary evolu-tion of biochemistry in this country during the 1930s. Many features of intermediarymetabolism, a central component of biochemistry, originated as attempts to answer funda-mental bacteriological questions. Thus, many bacteriologists altered their research programsto answer these questions. In so doing they changed their disciplinary focus from bacteriologyto biochemistry. Chester Hamlin Werkman’s (1893–1962) Iowa State career illustrates theresearch perspective that many bacteriologists adopted. As a junior faculty member in theBacteriology Department in the late 1920s, Werkman faced a powerful professional dilemma:establishing a research identity that distinguished him from his colleagues with flourishingnational and international reputations. His solution was to radically alter his research programfrom traditional bacteriology to a biochemistry program, which reflected the influence of theDutch microbiologist/biochemist, Albert Jan Kluyver (1888–1956). Werkman was extremelysuccessful in this career change. His laboratory made significant contributions to biochem-istry, and Werkman achieved a notable degree of personal success. His career began in theshadow of his departmentalbacteriologicalcolleagues; within a decade he became the depart-ment’s dominant research figure, as abiochemist. Werkman’s personal success, however, hadprofound consequences for the disciplinary future of bacteriology at Iowa State.

Keywords: bacteriology, biochemistry, Robert Earle Buchanan (1883–1973), Delft, disci-plinary evolution, Iowa State College/University, Albert Jan Kluyver (1888–1956), scientificcareers, scientific progress, Chester Hamlin Werkman (1893–1962)

Even the simplest story in the history of science has as many characters as aRussian novel.

David Hull, Science as a Process1

Science is not done by logically omniscient lone knowers but by biologicalsystems with certain kinds of capacities and limitations.

Philip Kitcher,The Advancement of Science2

1 Hull, 1988, p. 35.2 Kitcher, 1993, p. 59.

142 RIVERS SINGLETON, JR.

Introduction

How do scientific disciplines form? For example, today we recognize acollected body of knowledge, and associated experimental practice, thatwe call “biochemistry.” Furthermore, we also distinguish “biochemistry” asunique from related disciplines, such as “organic chemistry” or “physiology.”But, how did the disciplinary uniqueness of “biochemistry” come to be? Whatwere the forces that led to the unique “body of knowledge and experimentalpractice” so clearly identifiable from “physiology” or “organic chemistry”?What were the consequences of these forces on both practitioner and theirinstitutions?

In this essay, I seek to explore these questions regarding scientificdisciplinary evolution. More specifically I intend to focus on the way thatbiochemistry, especially notions of intermediary metabolism, evolved frombacteriology during the 1930s. I have several objectives in this study. First, anobvious goal is to understand ways that one discipline can effect the develop-ment of another. For example, Neil Morgan noted that many features of inter-mediary metabolism originated as attempts to answer fundamental questionsabout bacteria.3 To answer these questions many bacteriologists substan-tially changed their research programs, i.e. they moved from practicingbacteriology to practicing biochemistry.

A second essay objective is more sociological. Disciplinary changes donot occur in vacuo; the are brought about by human actors and can haveprofound effects reaching far beyond disciplinary evolution. When indi-vidual scientists significantly change their research practice, the changes cansuccessfully alter their careers and the broader disciplines within which theyoperate. Furthermore, achieving scientific success may allow an individual togain institutional power that they may wield either skillfully or poorly, whichcan profoundly affect their own institutions.

To explore these themes, I will focus on the career of Chester HamlinWerkman (1893–1962) and his interactions with the Dutch microbiologistand biochemist, Albert Jan Kluyver (1888–1956) in the bacteriology depart-ment at Iowa State; Werkman shifted his research program away from thetraditional bacteriology practice of his colleagues to the rapidly evolvingpractice of biochemistry. As a case study, Werkman’s career, illustratesboth the complexity of decisions about research programs and the personal,disciplinary, and institutional consequences of those decisions.

To place these objectives in context, however clarification of the terms“discipline” and “biochemistry” is vital. Both terms are complex, thus simpli-fication is essential. To address the first issue, i.e. the nature of discipline, I use

3 Morgan, 1990, p. 496.

FROM BACTERIOLOGY TO BIOCHEMISTRY 143

Philip Kitcher’s idea that disciplines can be defined in terms of the consensuspractices that community members see at its core.

Biochemistry: Discipline and Origin

Disciplinary Definition

Kitcher’s notion of consensus practice describes general principles that unitescientific communities around such shared attributes as language, notionsof significant problems, standards for justification and methodology, andacceptable explanatory schemes.4 Importantly, like Kohler’s notion of disci-pline,5 consensus practice is not fixed in time but rather undergoes successivechange. As disciplines evolve and mature over time, their consensus practicesalso change. Sub-disciplines may develop unique consensus practices thatdistinguish them from other disciplines. The notion of consensus practice isheuristically useful because it infers what practitioners within a disciplineactually do thus is helpful to achieve disciplinary clarity.

At a minimum, four such practices, i.e. enzyme theory, non-linear (cyclic)processes, metabolic pathways, and biological unity of processes, seeminglyprovide the core of biochemistry’s attempt to deal with questions about “themultitude of chemical reactions that occur within the living cells.”6

i) Enzyme theory: This notion postulates that all biological processes arecatalyzed by unique protein molecules. The postulate implicitly assumesthat understanding the nature of protein molecules would illuminate theunderlying biological process.7

ii) Integrated cycles: Many biochemical processes are not linear but ratherare integrated cycles, in which various central intermediates – indispens-able for cellular structure and function, energy transfer, or oxidationand reduction balances – must be constantly regenerated in order formetabolism to continue.8 The term cycle can have different mean-ings in biochemical parlance. For example, some processes occur asmajor metabolic cycles, such as the citric acid cycle discovered byKrebs and elegantly described by Holmes.9 However, all metabolicprocesses are ultimately driven by two critical cyclic processes. First,metabolic compounds are activated by ATP phosphorylation, which must

4 Kitcher, 1993, p. 87.5 Kohler, 1982, p. 1.6 Allen, 1978, p. 147.7 Holmes, 1991, p. 17; Kohler, 1973.8 Bechtel, 1986a; Bechtel, 1986b; Bechtel, 1988.9 Holmes, 1991; Holmes, 1993


be continuously regenerated. Second, low molecular weight compounds(cofactors such as NAD and NADH), which serve as cellular elec-tron donors and acceptors have limited concentrations; thus they mustbe continuously oxidized and reduced in a cyclic fashion in order forbioenergetic and biosynthetic reactions to proceed effectively.

iii) Metabolic pathway: This concept is implied by the previous two notions.Although the view that metabolism occurred via pathways of discretechemical steps involving transitory intermediates was conceptually estab-lished by 1900,10 the actual role of enzymes and reaction sequencesslowly evolved through the first half of this century. This work led to theview thatall metabolic processes occur in discrete chemical reactions,each catalyzed by a unique protein or enzyme molecule. Each step in theprocess is integrated and linked to another; the product of one reactionbecomes the substrate for the next. Metabolic pathways are essential forenergy conservation, accumulation of cellular products, or degradation ofcell materials.

iv) Biochemical unity: Biochemical research programs are shaped by theview that all living organisms share common chemical attributes, i.e.chemical mechanisms underlying biological activities are common fromone organism to another. Thus, biochemists could reasonably assumethat most organisms oxidizing glucose to carbon dioxide most likely didso via a metabolic pathway such as the Krebs cycle. Although manybiochemists shared this view during the early 1900s, A. J. Kluyver helpedform the concept and was one of its most articulate proponents.11

Modern biochemistry is structured around these four consensus practices,however they were not universally accepted and integrated into biochem-istry’s disciplinary structure until the middle decades of this century. Forexample, as late as 1938 Haldane noted that biochemistry was an emergingdiscipline in which metabolism, especially “intermediary” metabolism, hada central focus. He observed: “The ultimate aim of biochemistry may bestated as acomplete account of intermediary metabolism, that is to say, of thetransformations undergone by matter in passing through organisms.”12 Formuch of the century, that “complete account of metabolism” was achieved byindividual scientists seeking to understand the nature of enzymes and theirrole in the cyclic metabolic processes of a variety of organisms; they referredto their practice asbiochemistry.

Particular practices guided research in many laboratories, however theyoften had not achieved consensus status. The enzyme theory, for example,

10 Holmes, 1986; Holmes, 1992.11 See, for example, his classical paper: Kluyver and Donker, 1926.12 Haldane, 1938, p. 1 (emphasis added).


guided many biochemical research programs, nevertheless a solid consensusaround the theory did not develop until mid century. In 1933 Willstätter couldclaim that enzymes were low molecular weight compounds,13 even thoughprotein molecules were crystallized and demonstrated to possess uniquecatalytic activities in the mid to late 1920s.14 A decade later, Harrow’sText-book of Biochemistrystated tentatively: “Some of the enzymes, at least, areprobably proteins. . . . ”15 Ultimately the enzyme theory attained consensusstatus, because numerous biochemical research programs during this centuryisolated enzyme molecules in highly purified form and described the unique-ness, structure, and function of those proteins.16 In a similar fashion, indi-viduals who professionally identified themselves asbiochemistsbegan anexpansion of these central concepts by applying them to new organismsand different metabolic processes. Chester Werkman’s research program andthose of the students he trained, for example, increasingly reflected and weredriven by all four of the consensus practices summarized above.

Biochemistry’s Origin

Like its disciplinary structure, biochemistry’s origin is similarly complex.Traditionally, biochemistry’s disciplinary development emphasizes an exper-imentally driven evolution from physiological chemistry.17 As nineteenth-and early twentieth-century physiologists began to ask more complex andsophisticated questions about living processes, they increasingly turnedto the methods and rigor of chemistry for answers.18 The pervasivenessof this research approach was reflected even in popular culture, such asSinclair Lewis’ novel,Arrowsmith.19 When the student, Martin Arrowsmith,approached research professor, Max Gottlieb, to do research in microbiology,he was dismissed and told to come back to Gottlieb’s laboratory when he had“learned physical chemistry.”20

Organic chemical methods, especially analytical methods, providednumerous ways to isolate and characterize various compounds involvedin living processes.21 When Buchner demonstrated that a yeast cell-freeextract could convert glucose to ethanol and carbon dioxide, he simultan-

13 Laszlo, 1986, p. 445.14 Fruton, 1992.15 Harrow, 1943, p. 2 (emphasis added).16 Kornberg, 1989.17 Teich, 1970.18 Kay, 1989, p. 4.19 Rosenberg, 1976.20 Lewis, 1980, p. 15.21 Florkin, 1975; Fruton, 1990; Fruton, 1992; Teich, 1970.

Rivers Singleton

Highlight

Rivers Singleton

Sticky Note

see: Willsätter, Richard. 1933. Problems of Modern Enzyme Chemistry. Chemical Reviews 13: 501 - 512. (p. 508-509)


eously resolved the contentious Liebig/Pasteur debate over the nature offerments and, according to Teich, helped distinguish biochemistry as a uniquediscipline.22

Kohler has argued that biochemistry’s origin was more institutionallydriven. While he concurs that Buchner’s demonstration ofin vitro fermen-tation played an important role in the discipline’s evolution, Kohler contendsthat biochemistry resulted from “a reorganization of professional boundaries,whereby specialists in a wide variety of fields found common ground.”23 Partof the driving force behind this change was the reorganization and profession-alization of American medicine. As medical schools revised their curricula toincrease offerings in basic science during the early decades of this century,there was a rapid displacement of “medical chemists” (either physicianswho developed chemical skills or chemists who applied their skills to medi-cine) by “biological” or “physiological chemists.” The latter were no longerconcerned with analytical questions of medicine but rather addressed basicquestions about physiology. The major rationale for this shift in focus wasthat “physiological chemistry” potentially provided more insight into patho-logical processes than could be achieved by “medical chemistry,” which wasan applied science focused primarily on clinical, forensic, and toxicologicalanalysis.

In many respects, the traditional view, as explored by writers suchas Teich, and Kohler’s more institutionally driven view are both correct;however neither adequately explains the diversity of research inquiriespursued by individuals who called themselves “biochemists” during thiscentury. While both views provide insight, both ignore the important roleof bacteriology (especially agricultural bacteriology) and its associated ques-tions that biochemistry helped answer.24 Both perspectives fail to recognizethe important role that microbial metabolism played in biochemical under-standing.25 As physiologists changed their study approach, microbiology alsobegan to flourish. By the early part of this century, isolating and efficientlygrowing pure cultures of microorganisms in the laboratory was relativelyeasy. It soon became apparent that, because of their relatively homoge-neous character and ease of growth under precisely defined conditions,microorganisms were ideal subjects to understand the chemical mechan-isms of living processes.26 For example, the noted biochemist Fritz Lipmann

22 Teich, 1970.23 Kohler, 1973, p. 183.24 Kohler, 1985; Morgan, 1990.25 Morgan, 1985; Morgan, 1990.26 Werkman, 1939; Werkman and Wood, 1942; Kluyver and van Niel, 1956; Kohler, 1985;

Morgan, 1985; Morgan, 1990.


once remarked: “One of the lessons I had learned in Meyerhof’s labora-tory was that if mammalian systems show difficulties, one should turn tomicroorganisms.”27

A central premise of this essay is that bacteriologists, shifting theirresearch inquiry from the organism to the chemical processes involving theorganism,28 fostered biochemistry’s disciplinary evolution during the middledecades of this century. This shift may have been – in part – encouraged bythe increased service role that bacteriologists were called to serve. AlthoughPatricia Gossel’s work concerns an earlier period, she has demonstratedthat during the late nineteenth- and early twentieth-century bacteriologistsdeveloped numerous standardized techniques and practices to solve a varietyof public health issues. As this professionalization occurred, bacteriolo-gists increasingly assumed a utilitarian role in diverse areas ranging fromsanitation and public health to agriculture and food protection.29 While theinference is admittedly speculative, many of these individuals may have beenintellectually dissatisfied with this service role and sought more challenging“basic” research problems.30 This seems to have been the case with Werkmanwho saw biochemistry as a “more basic science.” When he was made Head ofthe Bacteriology Department (see below), a major rationale for the appoint-ment was that his research was “more modern” than that of his departmentalcolleagues.

Biochemistry: Kluyver and Werkman

Like Morgan, I argue here that biochemistry was significantly influenced byan attempt to understand fundamental microbial processes. I will examineconnections between A. J. Kluyver and changes in Chester Werkman’sresearch program to illustrate an evolutionary process whereby scientificdisciplines grow and mature. During the 1920s and 1930s Kluyver was avocal advocate for studying microbial metabolism in order to understandfundamental biochemistry. For example, in a 1932 lecture at the University ofPennsylvania Medical School entitled “Microbial metabolism and its bearingon the cancer problem” (later published inScience),31 Kluyver argued that

27 Quoted in Gunsalus, 1976, p. 127.28 Morgan, 1985; Morgan, 1990. While some bacteriologists shifted their major research

focus to biochemistry, other researchers, such as Marjory Stephenson, attempted to forge anew, middle ground discipline. “Bacterial physiology” was shaped by the “idea that bacteriawere organisms with a distinctive physiology worth investigating for its own sake” (Kohler,1985, p. 163).

29 Gossel, 1992.30 For a popular rendering of these potential intellectual conflicts, see Martin Arrowsmith’s

attempts to work as a public health bacteriologist (Lewis, 1980, pp. 186–257).31 Kluyver, 1932a.


Figure 1. A. J. Kluyver in 1926.

while many advances in biomedical sciences were made by studying thebiochemistry of – what we now call – eukaryotic organisms, bacteria alsoserved a major role in helping answer significant biochemical questions.A central premise in my argument is that Kluyver’s views, both indirectly(through his published work) and directly (through a visit to the Amescampus), were an important evolutionary influence on Werkman’s researchprogram.

A legitimate question is: “Why study Werkman?” He was not well knownoutside of a narrow scientific community and was one of many scientistsmaking similar transitions from bacteriology to biochemistry in the 1930s.History, however is often the sum story of many “not well known” indi-viduals, but whose unique stories provide valuable insights into the pressuresthat shaped their careers, their lives, the disciplines they practiced, and theinstitutions within which they worked. Holmes succinctly stated the point:“The context [of any historical narrative] is abackgroundwhich situates aforeground. Without a clear image of the foreground, sophisticated analysesof the background will remain barren.” Furthermore, thatforegroundmustbe sharply focused on “the immediate activities of creative scientists ‘doing


Figure 2. C. H. Werkman ca. early 1930s.

science’.”32 Undeniably, there were many “Chester Werkmans” scatteredacross American campuses during this century’s middle decades. But, ifwe are to fully understand the nature of the science they created, we mustunderstand their choices about research paths, as well as the personal andinstitutional consequences of those choices.

Werkman’s research program at Iowa State during the decade of 1930–1940 provides an excellent case to explore both the impact of personalchoices about research focus and disciplinary connections between bacteri-ology and biochemistry. Werkman’s laboratory produced several preeminentbiochemists, three of whom (Lester Krampitz, Merton Utter, and HarlandWood), as well as their mentor, Chester Werkman, were elected to theNational Academy of Sciences. Krampitz, Utter, Werkman, and Wood allmade significant contributions to our understanding of the chemistry of lifeprocesses, especially intermediary metabolism. Thus, a reasonable inferenceis that Werkman’s research program played an important role in shaping thedisciplinary evolution of biochemistry.

The focus of Werkman’s research program is seemingly associated withKluyver’s tenure as a visiting professor at Iowa State during the Spring andSummer terms of 1932, when he presented a series of twenty-four lectures on

32 Holmes, 1981, p. 61.


the “Physiology and Biochemistry of Microorganisms.” The lectures werelisted as graduate courses in both the bacteriology and chemistry depart-ments for both terms and were taken by Werkman’s students. Kluyver alsoorganized and led seminars on “The Physiology of Microorganisms,” whichmet twice weekly in the evenings, and were also listed as graduate courses.33

Bacteriology department faculty, in addition to graduate students, were activeparticipants in the seminar series.

Other questions arise from this brief summary of biochemistry’s struc-ture and origins: i) how do individuals develop a disciplinary identity?; ii)how and why do individuals make shifts in their research inquiry?; and iii)how does an individual’s disciplinary identity influence broader disciplinaryconsensus issues? These are questions about the ways consensus practicesdevelop and shape broader disciplinary evolution; the research program thatevolved in Werkman’s laboratory helps to illuminate them. Werkman changedhis research focus from the conventional bacteriology practice of his depart-mental colleagues to a program focused on problems of bacterial metabolism,a research agenda reflecting the ideas of Albert Jan Kluyver.

Albert Jan Kluyver and the “Delft Tradition” of Bacteriology

In popular opinion, Delft has been identified with bacteriology sinceLeeuwenhoek’s early descriptions of bacteria. However, for most modernmicrobiologists the city, and its Technical School, assumed prominence withthe work of Martinus Beijerinck (1851–1931). Beijerinck’s major contri-bution was in the area of microbial physiology,34 and as Bert Theunissenobserved, his “research canwith hindsight be characterized as chemicalmicrobiology.”35 Beijerinck’s disciplinary legacy was to demonstrate thechemical diversity of microbial life, but as Theunissen further commented,that legacy did not emerge from a “coherent physiological or chemicalprogram of bacteriological research.”

The Delft “coherent program” was established by Beijerinck’s successor,A. J. Kluyver. However, when Kluyver was appointed to the chair of generaland applied microbiology vacated by Beijerinck’s retirement he lacked thecredentials (in terms of formal training) to lead a department of “generaland applied microbiology,” certainly not one of the stature developed byBeijerinck. Although skilled as a chemist, Kluyver was “by no means an

33 Iowa State College, 1932, p. 130.34 Bennett and Phaff, 1993; Bos and Theunissen, 1995.35 Theunissen, 1996, p. 199 (emphasis added).


accomplished or broadly experienced microbiologist.”36 As an undergraduateand graduate student with G. van Iterson, Jr. (1878–1972) he mastered suffi-cient microbiology for instructional purposes. His doctoral thesis utilizeda variety of yeast strains, which also presumably introduced Kluyver tosome fundamental bacteriological practices. The major focus of his thesis,however was a correlation of CO2 production with sugar concentration, andit developed an analytical method for quantitative analysis of individualsugars in complex mixtures.37 Thus, while Kluyver’s graduate study utilizedmicroorganisms and made some contributions to yeast physiology, it wasfundamentally a chemical study.

Despite these disciplinary inadequacies, however, within a decade Kluyverdeveloped a unique research program at Delft that complemented andrivaled, in its imaginative sweep, his predecessor’s. Beijerinck contributedto our understanding about the diversity of microbial life. That knowledgeemerged from research concerned with “the heredity and variability of micro-organisms” and careful use of the enrichment culture technique,38 whichdemonstrated microbial activities in a wide spectrum of previously unex-pected ecological niches.39 Kluyver, however, built on Beijerinck’s vision ofmicrobial diversity, and developed a chemically based microbial physiologyresearch program that recognized the underlying chemical unity ofall livingorganisms. He often expressed this notion of biochemical unity in theaphorism: “From elephant to butyric acid bacterium – it is all the same!”40

Kluyver organized a vigorous research group committed to understandingthe chemical unity of microorganisms, and by the late 1920s and early 1930shis ideas were beginning to achieve international recognition. In 1930 he wasinvited to deliver a series of lectures at the University of London entitled“The chemical activities of micro-organisms.” The “Preface” to these lectures

36 van Niel, 1959, p. 70. See also: Amsterdamska, 1995, p. 196. In this regard, Kluyverwas not unlike Beijerinck, who, according to Theunissen, required several years “to famili-arize himself with bacteriological research, in which he had no experience at the time of hisappointment” (Theunissen, 1996, p. 208).

37 van Niel, 1959, p. 71.38 Theunissen, 1996, p. 199.39 Bennett and Phaff, 1993.40 Kamp, La Riviere and Verhoeven, 1959, p. 20. van Niel (1959, pp. 68–69) observed that

by mid-century many of Kluyver’s ideas were such established dogma that he was no longercited as their source. For example, Kluyver’s aphorism gained sufficient cachet that by theearly ’60s, Jacob and Monod paraphrased it, without attribution as “thatold axiom‘what istrue for bacteria is also true for elephants’ ” to justify the universality of the genetic code.[Monod and Jacob, 1961, p. 393 (emphasis added)]. Lily Kay (personal communication tothe author) very reasonably suggested that Jacob and Monod probably picked up Kluyver’saphorism from their close association with individuals who trained in van Niel’s microbialphysiology program at the Hopkin’s Marine Station.


clearly stated Kluyver’s perspective on the connection between biochemistryand microbiology:

The aim of the lectures was to show that a judicious selection of the exper-imental material available concerning microbial metabolism allowed oflogical deductions as to the fundamentals of biochemistry.A study of thechemical activities of micro-organisms reveals all the advantages whichmay be derived from “comparative biochemistry.”Although this line ofstudy has not as yet been much developed, it may in future win the samesignificance for biochemistry as “comparative anatomy” has already longago attained for anatomy.41

Thus, Kluyver, with his broad chemical vision of micro-organisms andnotions about the “unity of biochemistry” rapidly became a leading advocatefor the important role of micro-organisms as a means to understand funda-mental life processes on the molecular level. He brought this perspective toChester Werkman’s research programs and to his Iowa State students andcolleagues. To contextualize the changes in Werkman’s research agenda,an understanding of how bacteriology at Iowa State initially developed isimportant.

Early Bacteriology at Iowa State

Robert Earle Buchanan (1883–1973)

The Bacteriology Department at Iowa State College (ISC) in Ames, Iowaoriginated as a “spin-off” from the Botany Department, headed by LouisHermann Pammel (1862–1931). In 1888, Pammel introduced a bacteri-ology course, which was required for students in agriculture and veterinarymedicine.42 According to R. E. Buchanan, who wrote a brief departmentalhistory,43 Pammel’s introduction of this and other bacteriology courses intothe Iowa State curriculum made the college one of the first American agricul-tural institutions to offer formal course work in bacteriology. In 1910 Pammelrecommended that Bacteriology should become a unique department and thatBuchanan, his former student, should be its Head.

As an Iowa State undergraduate, Buchanan competed several coursesin bacteriology and assisted in bacteriology laboratories. He completed aB. S. degree in botany in 1904, and upon graduation Pammel asked that he

41 Kluyver, 1931, p. 5 (emphasis added).42 Packer, 1958; Packer, 1983.43 Buchanan, 1959.


remain as an instructor to teach bacteriology; during this time he completeda M.S. in Botany (in 1906). Over the next two years he completed a Ph.D.in soil bacteriology at Chicago, returned to Ames in 1908 as an AssociateProfessor of Bacteriology, and soon found himself “professor and head ofthe newly separated Department of Bacteriology.”44 Although he served avariety of other administrative roles, including Dean of the Graduate Schooland Director of the Agricultural Experimental Station, he retained the title ofDepartmental Head until 1945.

Pammel’s confidence in Buchanan’s abilities was not misguided, forBuchanan rapidly became one of this century’s foremost American micro-biologists.45 Widely recognized for his significant work in bacterialtaxonomy, Buchanan was also the first person to define bacterial growthprocesses in precise mathematical language. In 1918, he published a seminalpaper describing the various stages of bacterial growth.46 The paper’s centralfigure, and the accompanying mathematical equations, are virtually identicalto those found in modern microbiology textbooks. Further evidence ofBuchanan’s success was his election as President of the Society of AmericanBacteriologists [(SAB), now American Society for Microbiology (ASM)] in1918 at age 35,47 one of the youngest persons elected to that office.

Max Levine (1899–1967)

As Head of the new department, Buchanan believed it needed to developthe field of sanitary bacteriology, “[S]o we took up the matter with Dr.Sedgewick of the Massachusetts Institute of technology who recommendedhisstar pupil, Max Levine, who came to us to teach and to carry on research inthis field.”48 The choice was fortunate, for Levine rapidly became one of the

44 Buchanan, 1959.45 Singleton, 2000.46 Buchanan, 1918.47 American Society for Microbiology, 1992, pp. viii–xiii.48 Buchanan, 1959, emphasis added. While Levine may have been one of Sedgewick’sstar

pupils in terms of technical abilities, it is not clear that Sedgewick held him in high regard inother areas. In a July 25, 1913 letter to Levine apparently regarding Levine’s failure to obtainanother faculty position, Sedgewick noted:

I dare say that . . . you were the best qualified of the Tech candidates for Dr. Frost’s posi-tions so far as mere attainments go. In all such things, however, personality plays a greatpart, and I have no doubt that the fact that you are a Jew counted against you. . . .

It is, of course, unfortunate on some accounts that people have to pay attention topersonality, race, social and marital conditions, etc., etc., but I am finding that we havegot to take this old world as it is and not as it ought to be, and you must expect, I think,to run counter to a good deal of race prejudice.. . . it is a fact that most of our educationalinstitutions are run on what I might call the American plan, under which a young man


country’s leading water quality/sanitary microbiologists. Levine completedhis undergraduate training in sanitary bacteriology at MIT with WilliamThompson Sedgwick (1855–1921) in 1912 and briefly held a faculty positionin the Department of Biology and Public Health at MIT. In 1913 he moved toIowa State first as an Instructor and then as Assistant Professor. He completeda Ph.D. at the University of Iowa in 1922 and was promoted to full professorat Iowa State that year.

Levine’s career reflects many aspects of the professionalization of bacteri-ology discussed by Patricia Gossel (see above).49 His research activitycombined basic research on micro-organisms in parallel with applied bacteri-ology; indeed, it is often difficult to draw distinctions between the basic andapplied elements in his research program. He developed new methods toisolate and grow bacteria and turned those methods to utilitarian benefit. Forexample, Levine’s eosine methylene-blue medium allows one to isolate andpartially characterize coliform bacteria as indicators of the fecal contamina-tion of water;50 although developed in 1918, the medium is still frequentlyused to culture coliform bacteria in water samples. The pragmatic aspects ofhis work is reflected in a project to develop more efficient ways to sterilizere-usable glass bottles, work which was partially funded by commercialorganizations such as the American Bottlers of Carbonated Beverages.51

takes a position and works along at small pay, unmarried, waiting until he has got a goodsalary before he changes his social status. Jews, like yourself, on the contrary, seem to feelat liberty to marry, even before they graduate, and this I think, – and I am not sure butrightly, – produces real prejudice against them in ordinary American minds.

In other words, we cannot play two games in this world, the Jewish game and theAmerican game, but have got to be chiefly either one thing or the other; and when the twoclash, as they sometimes do, one must take the consequence of being either a Yankee or aJew as the case may be.

I fear, therefore, that you will find some difficulty in securing any educational posi-tion, and accordingly had better turn your attention to technical positions, such as that ofSanitary Bacteriology, in which I hope you may soon secure something.

Sedgewick closed the letter on a somewhat more friendly note but nevertheless felt compelledto reinforce his philosophical perspective: “Please give my kindest regards to Mrs. Levine andpardon me if I have spoken too frankly in this matter. Personally, I think that it is generally bestto face the music and to look facts, especially when they are disagreeable, squarely in the face,so that we may regulate our lives according to natural conditions” (Sedgwick, 1913). DespiteSedgewick’s anti-Semitic sentiments and desire to live lives regulated by “natural conditions,”he apparently did not express these feelings in his correspondence with Buchanan regardingthe faculty position at Iowa State.

49 Gossel, 1992.50 Levine, 1918a, 1918b.51 For example, see the “acknowledgment” in publications such as: Levine, Toulouse, and

Buchanan, 1928 or Levine and Buchanan, 1928.


Levine also played a central administrative role in Iowa State’s bacteri-ology department. In the early 1930s, as Buchanan began to assume thevarious higher level administrative functions discussed previously, Levinewas designated “Professor in Charge of Department.” The historical recordis vague regarding the precise duties associated with this ambiguous title butsuggests that Levine was the de facto department head. The annual budgetfor many years distinguished Levine’s appointment as “Professor in Charge”separately from other salary lines,52 and he signed graduate theses in thespace normally designated for the department head. Levine recognized hissomewhat unique role and in biographical publications, such asAmericanMen of ScienceandWho’s Who in America, designated himself as “Professorin Charge of Department.” He retained the title until 1945 when Buchananwas forced to give up the formal title of Department Head and was replacedby Chester Werkman.53

Chester Werkman

In his historical commentary Buchanan does not mention why, in the mid-1920s, he hired Chester Werkman – his former graduate student – as a facultymember in this developing bacteriology department. Werkman completed aB.S. in chemistry at Purdue in 1919. After a brief appointment as a researchassociate at the University of Idaho, he began graduate studies at Iowa Stateand completed a Ph.D. with Buchanan in 1923. Werkman’s thesis researchwas in immunology and vitamins and led to three publications dealing withthe effects of vitamin deficiency on the immune response.54 Werkman helda brief faculty appointment at the University of Massachusetts, howeverBuchanan may have brought him back to Ames, as an assistant professorin 1925, for this immunological expertise.

Upon returning to Iowa State, Werkman’s research interests began toundergo a slow evolution. Initially, he continued to publish papers on bothimmunology and vitamins. Possibly because of support by the agriculturalexperimental station, which Buchanan directed, he developed an interest infood microbiology and the role of vitamins as growth factors for bacteria.

52 Budget: Iowa State College.53 Because of the way he was later treated, this relationship may have caused discomfort

with the Iowa State administration. Levine’s administrative role at the institution has neverbeen officially recognized. The Index to the Official Archives at Iowa State University providesa brief history of all academic units and names of their administrative leaders, including indi-viduals who served as “acting” or “professor in charge.” Levine’s name is notably absent fromthis list for the Microbiology Department, which evolved from the Bacteriology Department(Archival Index).

54 For a complete list of Werkman’s publications, see: Brown, 1974.


During the early 1930s the Iowa State agricultural experimental station wasincreasingly interested in utilizing bacterial fermentation to dispose of farmwaste (see below). Werkman was involved in this effort, and he began topublish, with O. L. Osburn (one of his most productive graduate students),55

an extensive series of papers describing organic chemical techniques toisolate and quantify products of various fermentation processes.

The Evolving ISC Bacteriology Department

Although there were other bacteriology programs at Iowa State (therewere strong programs in dairy and veterinary bacteriology, for example),Buchanan, Levine, and Werkman constituted thecore ISC BacteriologyDepartment.56 The department was responsible for most of the generalbacteriology curriculum as well as for training graduate students. The latterrole was no small task, for as Rossiter has noted, Iowa State had “the largestgraduate enrollment of any of the ‘separate’ land grant colleges” in the late1920s.57

The role of chemistry in such a program cannot be undervalued. Likemany biologists in the early 1900s, Buchanan, Levine, and Werkmanallused chemical approaches to understand living organisms. At Iowa State thatchemical perspective was supported by powerful and productive chemistssuch as Henry Gilman.58 Buchanan relied on chemistry as a means to identifyand systematize bacteria to foster his interest in bacterial taxonomy. ForLevine, chemistry served dual purposes. Like Buchanan, Levine dependedupon chemical methods to identify unique organisms by their physiologicalproperties and to isolate those organisms using chemically defined selectivemedia. However, for Levine – as a sanitary bacteriologist – chemistry wasequally important as a powerful way to control bacteria in food and watersupplies. For Werkman, chemistry moved from serving as an analytical toolto the grounding for his disciplinary perspective.

By the mid-1920s both Buchanan and Levine were establishing nationaland international reputations on the leading edges of, what was then,modern bacteriology. In this rapidly evolving departmental structure ChesterWerkman presumably faced a powerful professional dilemma: establishingresearch independenceand avoiding duplication of his colleagues’ work.Possibly as a reaction to this pressure, by the 1930s his research emphasisbecame increasingly more chemical, and he used organic chemical techniquesto analyze bacterial fermentation mixtures. Kluyver may have provided

55 Singleton, 1997a.56 Packer, 1958, 1983.57 Rossiter, 1986, p. 49.58 Eaborn, 1990.


Figure 3. Iowa State Bacteriology Faculty. From right to left: Buchanan, Werkman, Levine,and graduate student, Harland Wood. Original undated but probably taken around 1935.

a potent solution to any professional quandary by opening the notion ofstudying bacterial metabolism simply in order to understand the chemicalprocess itself. Kluyver’s vision validated chemistry as a means to understandfundamental life processes in bacteria that transcended utilitarian purposes ofsystematics or sanitation; his perspective may have provided Werkman witha route to a professional identity unique from his departmental colleagues.

A. J. Kluyver at Iowa State

By the early years of his fourth decade, Kluyver had rapidly developed aninternational reputation in the biochemical community; he was, for example,a member of the editorial board of the influential journalBiochemische Zeits-chrift from the early 1920s until just before the start of the Second WorldWar. His influence reached even to the corn fields of America; Buchanan andFulmer opened the third volume of their comprehensive study of bacterialphysiology and biochemistry59 with an epigram from Kluyver and Donker’sclassical paper, “Die Einheit in der Biochemie,” and they cite many ofKluyver’s papers throughout the book.

59 Buchanan and Fulmer, 1930b.


Nevertheless, despite Kluyver’s growing biochemical influence, reason-able questions arise: i) why would a bacteriology department in a relativelysmall agricultural college in the American mid-West invite him for anextended visit?; ii) what were the personal driving forces behind the invita-tion?; and iii) what was Kluyver’s impact on such an institution? Thesequestions seem especially compelling, since the invitation was extendedin the midst of the American Depression, which imposed serious financialexigencies on all institutions. Although faculty salaries (at least for Buchanan,Levine, and Werkman) at Iowa State compared favorably with national aver-ages, they were static, and occasionally decreased, during the depressionyears. For example, Werkman’s salary annually varied by several hundreddollars from 1929–1940,60 and during one two-year period decreased almost$600.

Why Kluyver?

The institutional rationale to invite Kluyver seems clear. As Graduate CollegeDean, Buchanan wrote a series of “Annual Reports to the Graduate Faculty”in which he summarized various graduate program activities. In his 1931Report, Buchanan stated that the College hoped to have Kluyver as a“Visiting and Exchange Professor” during the Spring and Summer termsof that academic year, as part of a general agricultural research initiative.He discussed Kluyver’s pending visit in the following context: “As many ofyou are aware the Iowa State College is centering a considerable part of itsresearch around the utilization of farm waste. Among the possibilities of suchutilization are those in which fermentations are used.” Buchanan further notedthat Kluyver’s laboratory had intensely specialized in fermentation studies,and he made the curious observation – perhaps to justify the chemistry depart-ment co-sponsorship – that while Kluyver was “primarily a bacteriologist hehas an excellent grasp of the chemical phases as well.”61

Personal Motives Driving the Invitation

While the institutional rationale for inviting Kluyver seems clear, the personalmotives behind the invitation are complex. Buchanan said that the invitationwas extended by the Research Council, which he chaired, in the names ofthe Departments of Bacteriology (which he also led) and Chemistry. But thereport does not identify the person in either department responsible for initi-

60 Salary data from Budget: Iowa State College. Comparative data from: Stigler, 1956,p. 129.

61 Buchanan, 1931, p. 8.


ating the invitation. Arguably, both Buchanan and Werkman would benefitfrom the Kluyver visit.

Buchanan was deeply influenced by Kluyver’s work and may have beenthe driving force behind the invitation. As I noted previously, the third volumeof Buchanan and Fulmer’s bacterial physiology text opened with an aphorismfrom Kluyver and Donker’s “Die Einheit in der Biochemie.” Furthermore,Buchanan was both academic administrator and research scientist; Kluyver’svisit may have represented an opportunity to “show off” his Midwestern, agri-cultural college to an international scientist of increasing stature. Buchananenthusiastically expressed this perspective in his “Report to the GraduateFaculty” in 1932 (see below).62

Alternatively, evidence supports arguments that Werkman may have beeninvolved in inviting Kluyver. A compelling inferential argument is based onan undated typescript in Werkman’s papers.63 Although not labeled as such,the text appears to be a manuscript for comments Werkman made duringone of Kluyver’s biweekly “Microbial Physiology” seminars. Both contentand references date the text to the early 1930s. In the opening paragraphs,Werkman grandiosely commented that “before the evening is closed, thewhole subject of microbiology may be redefined and all the trouble of theworld except the depression may be eradicated.” The typescript’s connec-tion to Kluyver’s seminar series is evident, for in a subsequent paragraph,Werkman stated that “Prof. Kluyver suggested this topic for a seminar” and“I wish to provide Prof. Kluyver with the opportunity of placing before uscertain of his own reflections. . . . Wehope to find Prof. Kluyver in a discursiveand confidential mood tonight.”64 The typescript is important because itprovides insight into Werkman’s personality, his research thinking, as wellas possible motives for the Kluyver invitation.

As a reflection of Werkman’s personality, his comments give an appear-ance of ingratiation to Kluyver, and he seems to accept many of Kluyver’sideas in an uncritical fashion. Indications of editing suggest that attemptswere made not to offend Kluyver, as Werkman deleted portions of thewritten text that might have been perceived as critical when stated verbally.Werkman’s stance seems a bit unusual, especially when compared withBuchanan’s comments, because the discussion (which involved presentationsby Kluyver, Werkman, and Buchanan) was intended as a debate on “TheSignificance to be Attributed to the term ‘Fermentation’.”

While Werkman’s comments to Kluyver seem almost unctuous, Buchananforcefully challenged many of Kluyver’s views. For example, Kluyver

62 Buchanan, 1932, p. 7.63 Werkman, “Critique.”64 Werkman, “Critique,” p. 1.


developed and defended a traditional Pasteurian definition of fermentationin his presentation. He noted that much confusion had developed about theterm, but that, if it was to retain any semblance of meaning, there must besome return to Pasteur’s original concept of “all energy yielding metabolicprocesses proceeding without the cooperation of free oxygen.”65

Buchanan’s response was deferential to Kluyver’s position both as adistinguished scientist and as a guest. A hand written notation on his typedtext reads: “Respectfully submitted to Dr. A. J. Kluyver, with the hope thatit be suitable expurgated, corrected and emended, and in the end lead to aclarified vision on the part of the present writer.”66 Nevertheless, Buchananthen critically analyzed and dissected Kluyver’s entire argument. Kluyverquoted sentences from Pasteur in the original French; Buchanan quoted entireparagraphs. Buchanan did not appear to want to demonstrate the inadequacyof Kluyver’s discussion but rather to illustrate numerous alternative under-standings of the term “fermentation.” After a devastating critique of “thepresent status of French bacteriology” and the Pasteur Institute, which areconcerned with “too much retrospection, too much hero worship,” Buchanansuggested “several possible definitions of fermentation” based on “the ‘Amer-ican’ concept.” He then outlined a series of possible understandings ofthe term “fermentation,” many of which begin to verge on our modernunderstanding.67

As a representation of Werkman’s thinking about microbial metabolism,his notes reflect a sophisticated understanding, for an agricultural bacteri-ologist, of the then significant biochemical problems. The major focus is oncontemporaneous metabolic concepts in the late 1920s, and the most currentreference cited is 1932. Werkman critically analyzed the confusion thenrampant over the metabolic role of the Harden-Young ester, often referred toas “co-enzyme.” He noted that the compound called “co-enzyme” had beeninvoked to serve at least six different functions, so that the term was mean-ingless.68 Werkman also critically analyzed the Warburg-Weiland controversyover biological oxidation processes and elaborated on Kluyver’s role in tryingto resolve the controversy.

Despite the seemingly ingratiatory aspect of his comments, which maysimply reflect the “junior status” he felt in this “debate,” the typescriptindicates that Werkman was seriously thinking about significant biochem-

65 Kluyver, “Significance,” p. 4.66 Buchanan, “Significance,” p. 1.67 Buchanan, “Significance,” p. 3. Despite Buchanan’s criticism of his argument, the text

sufficiently impressed Kluyver that he retained the copy Buchanan gave him, which was filedwith Kluyver’s own text in Delft.

68 Werkman, “Critique,” p. 2.


ical problems, especially metabolic problems, prior to Kluyver’s visit. LikeKluyver, he was also thinking about ways that bacteria might be used as toolsto address those problems. If Werkman was seeking to identify himself witha clear research program separate from his already distinguished colleagues,support from a world-renowned figure like Kluyver would lend powerfulcredence to that goal. Thus, Werkman’s seminar comments perhaps reflecthis growing need for a research program unique from his colleagues’, and theKluyver invitation helped foster this budding research agenda.

As further support for Werkman’s growing biochemical interest, he appar-ently had begun to establish ties with the biochemical community priorto Kluyver’s visit. For example, Werkman apparently wrote to the highlyrespected German biochemist, Carl Neuberg, to discuss Kluyver’s criticismof fermentation theory. Werkman’s letter is not available, but in August 1932Neuberg replied: “I regard . . . Kluyver as an excellent researcher and anextremely pleasant man; I don’t know what he said in his critique of fermen-tation theory, but I start from the conviction that theories are unimportant andonly experiments have value.”69 Although Werkman’s motives for writing arenot readily apparent, the tone of Neuberg’s letter implies that Werkman wascultivating contacts in the biochemistry community by 1932.

The previous discussion suggests that personal motives driving theKluyver invitation were complex. Perhaps Buchanan invited him, as anadministrative maneuver, to both enhance the status of the Ames campusas well as to provide intellectual enrichment for the college community. Ifso, it seems a somewhat extravagant expenditure of funds during a time offinancial exigency. Kluyver received $500 in remuneration and an additional$75.43 was spent for clerical and other expenses connected with his Amestenure. To place Kluyver’s stipend in perspective, it constitutes between 12%to 15% of a median annual salary for an Associate or Full professor in a landgrant institution in 1929. However, Kluyver was not well compensated, ona monthly basis, in comparison with Werkman and Levine; his $500 stipendwas between 60–70% of their monthly compensation.70 Nevertheless, sinceBuchanan, Levine, and Werkman were all experiencing static or decliningsalaries, the $500 expenditure for Kluyver seems a bit of a luxury.

Werkman, however, by the late 1920s may have become increasinglyaware of his research situation in the bacteriology department and realized

69 Neuberg, 1932.70 Kluyver expenses from: Buchanan, 1932. Comparative data from: Stigler, 1956 and

Budget: Iowa State College. Kluyver also very likely received compensation from othersources during this time. For example, he gave a lecture on “The Chemistry of Microbes”to the Milwaukee Section of the American Chemical Society (undated type script, Kluyverpapers), and a hand-written rail itinerary outlines visits to the Universities of Chicago, Illinois,and Wisconsin (Kluyver papers).


that he needed to stake out new intellectual turf to distinguish himself fromBuchanan and Levine. His commentary in the Kluyver seminar and corre-spondence with Neuberg both suggest that Werkman was well-read in theevolving biochemical literature. A reasonable inference is that Werkmanhelped advance the Kluyver invitation in order to create an environmentfavorable for his evolving personal research interests.

Obviously, these motives are not mutually exclusive. Most likely, the ulti-mate answer is that both Buchanan and Werkman collaborated on Kluyver’svisit as a way to foster their mutual, albeit disparate, interests.

Kluyver’s Impact at Iowa State

Regardless of the rationale for his visit, Kluyver’s tenure on the Ames campuswas important for three major reasons. First, as a distinguished internationalscholar, Kluyver brought a broad cosmopolitan world view to both the collegecampus and to the broader social community. His visit did not go unnoticed.The Ames daily newspaper ran an article announcing his first lecture, “Unityand Diversity of Microbial Metabolism.”71 He was interviewed by a reporterfrom the campus daily newspaper, which ran a front page story under theheadline “Kluyver Here From Holland for Lectures” the day after his lecture.The article noted that Kluyver was “a world famous expert in the fieldsof fermentations and physiology of bacteria.”72 Iowa State awarded him anhonorary D.Sc. at its Spring commencement, the first of many internationalawards Kluyver received during his life, a further reflection of Kluyver’s wideimpact on the college community.

Second, through his lectures and seminars Kluyver introduced adifferent biochemical perspective of bacteriology to the Iowa State scientificcommunity. To place Kluyver’s perspective into proper context, readersshould remember that bacteriology at Iowa State already contained a strongchemical dimension prior to his visit. Kluyver was very much aware ofthis fact, and in his opening lecture, he commented: “bringing microbi-ological knowledge . . . to Ames is very much the same as the traditionalbringing of coals to Newcastle. . . . there will be but a few students of generalmicrobiology in Europe who will not often consult the standard treatiseon the physiology and biochemistry of bacteria by Dean Buchanan andProf. Fulmer.”73 Buchanan, Levine, and Werkman all utilized chemical tech-niques in their bacteriological research programs. Furthermore, students wereinstructed using the three volume treatise,Physiology and Biochemistry of

71 Anonymous, 1932a, p. 1.72 Anonymous, 1932b, p. 1.73 Kluyver, “Microbial Metabolism,” p. 1.


Figure 4. Werkman and Kluyver at Iowa State commencement.

Bacteriaby Buchanan and Fulmer,74 which constituted “a combination andrevision of the lectures on physiology of bacteria. . . given at the Iowa StateCollege to students of bacteriology.” The texts resulted from “a belief on thepart of the authors that there exists a need for a compilation and system-atization of material relating to the physiology of microörganisms.”75 TheBuchanan and Fulmer work, which is heavily grounded in physical andorganic chemistry, is very modern in its perspective and was a departure frommany contemporary bacteriology texts. Thus, Kluyver’s audience in Ameswas not unfamiliar with the basic content of his presentations.

However, a vital difference in perspective emerges from Kluyver’s lecturetopics (summarized in Table 1) and notes76 and the metabolic view reflectedin the Buchanan and Fulmer texts. For bacteriologists reading Buchananand Fulmer, metabolism consisted of observations that different organismsconsumed specific substances (e.g. glucose) and excreted specific products

74 Buchanan and Fulmer, 1928, 1930a, 1930b.75 Buchanan and Fulmer, 1928, p. xi.76 Kluyver, “Lectures Ames”; Kluyver, “Microbial Metabolism”.


Table 1. Kluyver lecture schedule at ISC1

Spring term lectures Summer term lectures

Lect. Date Topic Lect. Date Topic

no. no.

May 9 Introductory lecture:

Unity and Diversity

1 May 11 Introduction and alcoholic 11 June 15Summaryfermentation

fermentation I Harden byB. aerogenes

and Young 13 June 20 Propionic acid fermination

2 May 13 Alcoholic fermentation: 14 June 22 Cellulose fermentation

Alcoholic II First Phases 15 June 24 Methane fermentation

3 May 16 Alcoholic III Final Phases 16 June 27 Denitrification

4 May 18 Alcoholic IV Kv. and Struyk 17 June 29 Desulfuration

5 May 20 Alcoholic V Recent advances 18 July 1 Respiration: acetic acid bacteria

6 May 23 Alcoholic VI Co-enzyme 19 July 6 Respiration: yeast; respiration

7 May 27 Fermentations closely and fermentation

related to alcoholic. 20 July 8 Respiration: other aerobic

Lactic acid fermentation microbes; nitrogen fixation

8 June 1 Butyric acid and butanol 21 July 11 Assimilation I

fermentations I 22 July 13 Assimilation II Purple bacteria,

9 June 3 Butyric acid and butanol carbon dioxide fixation

fermentations II 23 July 15 Specificity in biocatalysis

10 June 6 Fermentation byB. coli 24 July 18 Redox potential and metabolism

1 Based on his notes at Delft; all notes are in AJK’s hand writing.

(e.g. ethanol) during growth. A central problem for bacteriologists was anability to distinguish between two types of micro-organism; differences inmetabolic capabilities served as a potent tool for this problem. Thus, themetabolic focus in Buchanan and Fulmer’s treatise provided vital chem-ical insights to aid bacteriologists’ need to differentiate micro-organisms butprovided little insight into the metabolic process itself.

Kluyver’s biochemical perspective, however, reflected the evolvingconsensus practice about metabolic processes, which had to be understood asintegrated sequences of reactions. For most bacteriologists, the observationthat an organism produced ethanol when growing on glucose was sufficientfor identification purposes. For Kluyver, however, such an observation wasinadequate as long as a series of reactions connecting growth substrate andproduct remained absent, and he devoted six lectures to the topic of “alcoholicfermentation.”

In these lectures he speculated on various aspects of the rapidly evolvingarea of intermediary metabolism, such as the metabolic role of phosphate.


The metabolic importance of phosphorylated esters77 had been recognizedfor decades. Most biochemists, however, had little idea of their chemical role,and the generally accepted view was that phosphate some how “enhanced themetabolic reactivity” of various intermediates.78 Kluyver, however, continu-ously sought to demonstrate a more direct chemical function for phosphate.79

Many of these ideas were wrong, but their hypothetical nature challengedothers to consider alternative hypotheses. Thus, by his strong emphasis on thecurrent biochemical questions about micro-organisms, Kluyver introducedhis Ames audience, in a first hand and immediate fashion, to a new anddifferent rationale for studying bacteria.

Third, Kluyver’s tenure in Ames coincided with a radical alteration ofChester Werkman’s research program. In the post-Kluyver years, his researchfocus underwent a dramatic disciplinary shift away from his colleagues’traditional bacteriology to a research inquiry structured by the then evolvingbiochemical consensus practices. We cannot answer issues of whether: i)Kluyver’s visit motivated this shift in Werkman’s research program; or ii)Werkman maneuvered the Kluyver invitation to foster a campus atmosphereconducive to this new area of research inquiry. We can demonstrate, however,that a profound disciplinary shift occurred, that this shift was coincidentalwith Kluyver’s visit, and then examine some of the consequences of thatchange.

Evolution of Bacteriology to Biochemistry at Iowa State

Kluyver and Iowa State Bacteriology

Kluyver’s influence on Iowa State’s bacteriology program was complex andinvolved social concerns in addition to research activities. Assessing hisinfluence is also contingent on the perspective from which questions areraised.

From Buchanan’s administrative and consequently social perspective,Kluyver’s visit was both a political and academic success. In addition toforcefully advocating his biochemical perspective of microbial life, Kluyver’svisit served two important social purposes: he helped place Iowa State intothe international arena, and he brought an international scientific perspectiveto Buchanan’s Midwestern, agricultural college. With apparent pleasure,Buchanan commented on the value of Kluyver’s visit in his 1932 report to the

77 Organic compounds in which a hydroxy lmoiety is phosphorylated, e.g.glucose-6-phosphate.

78 Korman, 1974.79 van Niel, 1959, pp. 91–93 and cited references.


graduate faculty. He noted that Kluyver: “proved himself to be a real asset,and the inspiration of his contact will be long with us. It means, further, thatwe have in Holland now a man quite conversant with the Iowa State College,who can interpret to his European colleagues the work we are doing. It hasmade possible and available to our graduate students an important Europeanscholar, and made real to them the existence and importance of Europeanscholarship.”80

Kluyver was apparently also pleased about the Iowa – Delft connection.After finishing the Ames lectures, the Kluyvers traveled across the country,ultimately visiting Kluyver’s former student, C. B. van Niel, Director of theHopkins Marine Station in California. In an informal and chatty letter to theWerkmans, written in Chicago on the return trip back east, Kluyver notedthat: “Obviously Van Niel has been a very efficient propagandist for ourDelft ideals, and I trust that in the future there will be a very good mutualunderstanding between the bacteriological departments of Ames and of theHopkins Marine Station.” On a more personal note, Kluyver concluded: “Iwill always keep unforgettable souvenirs of my Ames stay, and wheneverthese memories will be before me, I will see you and your friendship in thosedays in the center of my picture. But before all, I hope that you, too, willconsider my visit just an episode in our friendly relations.”81

From a research activity perspective, Kluyver probably had little influenceon either Levine or Buchanan. Both men had well established, nationallyrecognized research programs prior to Kluyver’s visit. Furthermore, whileboth Levine and Buchanan used chemical approaches in their researchinquiry, their research questions involved the entire micro-organism and itsenvironment. Levine had published almost forty papers and two books priorto Kluyver’s visit,82 but most of this work focused on characterizing bacteriawith infectious or public health implications. Buchanan was principally inter-ested in developing a coherent and unified taxonomic scheme for bacteria.Kluyver’s highly reductionist biochemical perspective on micro-organismsserved little to advance the research programs of either Levine or Buchanan.

Ultimately Kluyver’s visit to the Iowa State Bacteriology Departmentappears as a watershed event in the department’s history and coincideswith profound shifts in the way the department viewed itself. This alteredview came about primarily through the change in Werkman’s disciplinaryfocus; a shift that led to a successful scientific career for Werkman but alsoprecipitated profoundly negative institutional consequences.

80 Buchanan, 1932, p. 7, emphasis added.81 Kluyver, 1932b, pp. 2 and 4.82 Fleming, 1956.


Kluyver’s Influence on Werkman

Whether his research interests began to shift prior to Kluyver’s visit orwhether Kluyver created a new research vision for him to follow, Werkman’sresearch focus radically changed to a more biochemical perspective in thepost-Kluyver years. Despite the absence of Kluyver citations in Werkman’spublications,83 topical analysis of his publication record illustrates both theprofound changes that occurred in his research work, as well as the closecoincidence of those changes with Kluyver’s visit.

From 1924, when he was initially appointed as a faculty member atIowa State, until 1931, Werkman published twenty-seven papers, some ofwhich extended his doctoral studies. Other papers were loosely connectedto various projects of the Agricultural Experimental Station, which providedWerkman with both salary and research support.84 Although these papersdealt with bacteriological topics, they were poorly focused and do not reflecta unified research program. Werkman’s biographer, R. W. Brown, noted:“Publications bearing Werkman’s name during the period 1920–1930 exhibita variety of unrelated research efforts and suggest that he was searchingfor an area that would become his primary focus.”85 In 1930 and 1931,Werkman began to publish a series of papers that developed methods toisolate and quantify organic compounds from fermentation mixtures, perhapsreflecting Iowa State’s “centering a considerable part of its research aroundthe utilization of farm waste.”86 After Kluyver’s visit, however, Werkman’slaboratory increasingly concentrated on the metabolic processes responsiblefor the fermentations rather than simply analyzing fermentation products. Hisresearch assumed a more biochemical rather than bacteriological or chemicalfocus, and this change is clearly reflected in his career publication record.

The journals in which scientists publish can help identify their disci-plinary perspective and identification, and the journal venue of Werkman’spublications illustrates his disciplinary research focus. During his career,Werkman’s laboratory published 230 papers or monographs in twenty-ninedifferent journals or book chapters.87 Based on journal title, the disciplinarybasis of these publications varied greatly, however almost 40% of Werkman’s

83 Because scientists generally acknowledge the influence of each other’s ideas in theirpublications (Hull, 1988), one might expect to see an increased reference to Kluyver’s work inWerkman’s publications. Yet, except in rare instances, neither Werkman nor his students citedKluyver in their publications. (I thank Ms. Jennifer Smith who completed this analysis as partof an undergraduate study project.)

84 Budget: Iowa State College.85 Brown, 1974, p. 337.86 Buchanan, 1931, p. 8.87 Brown, 1974.


Table 2. Chester Werkman’s publications organized by topic discipline

Discipline topic No. papers %

Biochemistry 136 59.1

Bacteriology 67 29.1

Chemistry 22 9.6

Immunology 5 2.2

research reports were in distinctly biochemical journals.88 While this obser-vation suggests a biochemical disciplinary identity in Werkman’s researchprogram, it is insufficient evidence to fully clarify either his disciplinaryidentity or changes that may have occurred in it. Ambiguity arises because formuch of this century both biochemistry and bacteriology have been markedlyinterdisciplinary. Journals in each discipline frequently publish papers thatdeal with the other discipline. These difficulties are further complicated bythe fact that almost a quarter of Werkman’s publications were in a variety of“general science” journals, which by definition can have various disciplinaryfoci. Consequently, clarification of Werkman’s disciplinary identity requiresa more subtle analysis of his publication record.

Based on content, Werkman’s publications can be organized into fourtopical categories: bacteriology, biochemistry, chemistry, and immunology.Data in Table 2 summarizes the distribution of papers in each disciplinarycategory and clearly illustrate the overall biochemical nature of Werkman’sresearch inquiry.89 The data, however, do not support the hypothesis that hisresearch focus changed over time, nor does this observation support or refuteKluyver’s connection with that investigation. The change in Werkman’sresearch program, and Kluyver’s connection to that change, is reflected by theway these research topics changed over time, as shown by data in Figure 5.90

88 This analysis was obtained as follows. Citation data, including individual co-author, date,journal, journal discipline, and paper topic for each of Werkman’s publications (Brown, 1974,pp. 328–370) were entered into a spreadsheet (Quarto QPro©) database. This database wassorted by journal discipline, and the number of papers in each category counted using thespreadsheet “@COUNT.”

89 Data in Table 2 were obtained as described previously, except that the database was sortedby paper topic, which was assessed based on title, abstract, or if necessary, critical reading ofthe full paper.

90 Data in Figure 5 were obtained from the database described previously. Publications wereinitially sorted by publication year and secondarily by paper topic. The number of papers ineach topical category, for each year, and the total annual publications were calculated usingthe spreadsheet “@COUNT” function.


Figure 5. Chester Werkman’s publication record: chronological and topical perspectives.

Werkman’s immunology research was relatively minor, resulting primarilyfrom his thesis research, and appeared over a very short time. During thelate 1920s, he began to develop a series of chemical analytical methods touse in fermentation analysis. This interest is seen in the increased number of“chemistry papers” beginning around 1930 and continuing for a major part ofthe decade. The interest was relatively short lived, and after 1940 Werkman’slaboratory published no papers with a uniquely chemical focus.

Werkman published work with a specific bacteriological focus from hisearliest career. During the 1930s, however, these “bacteriological” papersbegan to decline, replaced in part by papers with a more “chemical” focus.Around the same time papers identifiable as “biochemical” began to strik-ingly increase from none in the years prior to 1933 to a maximum of fifteenpapers in 1940. By 1936 the major portion of Werkman’s research outputconsistend of biochemical papers, which often constituted 100% of his annualpublications, as data in Figure 5 clearly demonstrate. Figure 5 also illus-


trates that this shift in Werkman’s research interests closely coincided withKluyver’s visit to Iowa State.

The content of Werkman’s biochemical papers reflect the variousconsensus practices shaping biochemistry’s evolving disciplinary structure;the research program was shaped by concepts such as “enzyme theory,”“metabolic pathways,” and “biochemical unity.” For example, as his thesisproblem, Harland Goff Wood (1907–1991) began to study glucose utilizationby the propionic acid bacteria (a problem begun by C. B. van Niel, Kluyver’sgraduate student). His thesis work led to the discovery of heterotrophicCO2-fixation (the concept thatall organisms, not just plants or specializedbacteria can utilize CO2), known as “the Wood-Werkman reaction.” Overthe next decade, Werkman’s laboratory developed a major research effort tostudy the chemical mechanism of this process.91 Wood’s thesis also proposeda scheme of reactions to explain the fermentation of glucose to propionic acid.Over the rest of his career Wood elucidated the detailed pathway of glucoseutilization and isolated and characterized the enzymes involved.92 AnotherWerkman student developed a cell-free bacterial extract thereby providing ameans to purify bacterial enzymes. Other students isolated metabolic inter-mediates from a variety of bacteria thereby confirming the role of metabolicpathways (e.g. the Kreb’s cycle) in these organisms.

While Kluyver’s visit seems clearly associated with Werkman’s shift indisciplinary focus, this change was also fostered by a succession of ener-getic and dedicated graduate students and postdoctoral associates, who wereselected for their strengths in chemistry.93 Harland Wood, for example, joinedthe laboratory in September 1931. During the following twelve years, asgraduate student and post-doctoral research associate, Wood helped developWerkman’s laboratory into one of the foremost facilities in the world forbiochemical research, especially in the developing field of using isotopictracers to understand metabolic processes.94

The combination of refocused research perspective and gifted studentsmade Werkman’s laboratory extremely productive after Kluyver’s visit. Priorto 1931, Werkman published an average of two to three papers per year; from

91 Singleton, 1997a.92 Singleton, 1997b.93 Werkman apparently had little reservations about “raiding” talented students from other

departments. Merton Utter initially began graduate studies in Henry Gilman’s organic chem-istry laboratory. Werkman offered Utter a fellowship that paid more than Gilman’s stipend(it was sufficient to allow the Utters to marry), and Utter began graduate studies in theBacteriology Department the following term. (Mrs. Marjory Utter, personal communication)Werkman’s ability to raid students from Gilman’s laboratory was most likely enhanced byGilman’s miserly treatment of his students (Eaborn, 1990).

94 Singleton, 1997a, 1997b.


1931 to 1943 his research group annually published between six to seventeenpapers. Furthermore, for much of this period the “3-year running average”publication record exceeded ten papers per year.95 As I noted above, thesepublications reflected the evolving biochemical consensus practices, albeitbiochemistry focused on microbial activities. Nevertheless, in many of thesepapers the role of the microbe was a “bag full of enzymes,” a perspectivethat Kluyver alluded to in his first Ames lecture, when he referred to, “. . . theconception that a living cell is to be considered an arsenal filled with theseagents – or enzymes.”96

As his laboratory’s reputation increased, Werkman’s prestige in thescientific community, especially the biochemical community, also increased;he was appointed to the Chemical Section of the War Production Board andto editorial boards of several biochemical publications. He was a foundingeditor of Archives of Biochemistry, where he reviewed “articles dealingparticularly with enzymes and cellular metabolism,”97 as well as an Editorof the annual monograph on enzymes,Advances in Enzymology. Werkmanpersonally began to identify himself with the rapidly developing biochemicalcommunity, rather than as a bacteriologist. In 1946, he was elected to theNational Academy of Sciences, and in his letter accepting Academy member-ship indicated that he wished to be included in the Academy’s “biochem-istry and physiology” section, rather than the “pathology and bacteriology”section.98

Conclusion: Institutional Consequences and Disciplinary Evolution

Despite the powerful but understated role of contingency in this story, twomessages seem clear. If, in 1930, Chester Werkman was a scientist seeking toestablish a unique professional identity in a department dominated by well-established careers, he was very successful; by 1940hewas the preeminentdepartmental research force. As this case study illustrates, he achieved thissuccess by identifying himself with the external biochemical communityrather than in the bacteriology community that was his institutional home.As his significant number of publications suggest, Werkman’s laboratory wasinfluential in shaping modern biochemistry.

The story also illustrates that Werkman’s shift in professional identity wasassociated with Kluyver’s visit. Whether Werkman maneuvered Kluyver’sinvitation to Ames to facilitate his already evolving independent research

95 Singleton, 1997a, pp. 111–112.96 Kluyver, “Microbial Metabolism,” p. 16.97 Anonymous, 1943, p. 4.98 Werkman, 1946.


identity or whether he was swept up by Kluyver’s biochemical perspectiveduring the visit is not clear. Werkman’s shift in disciplinary focus, howeverfrom traditional bacteriology to biochemistry, clearly was connected with theDutch biochemist’s stay on the Ames campus.

While this story about Werkman’s personal success is apparently clear,what does it tell us more broadly about scientists’ careers and scientificprogress? Furthermore, can we learn anything about the impact of thesechanges on the institutions in which they occur? Like a Tolstoy novel, answershere are interconnected, subtle, and convoluted. They connect around thenotion of scientific progress, which requires brief clarification.

Philip Kitcher noted that scientific progress occurs “continuously, weekby week, day by day, because of numerous small incidents and decisionsinvolving small groups of people.”99 In Kitcher’s view, which shares commonfeatures with David Hull’s perspective of scientific progress, individualscientists join together in communities around shared notions of rigor,methodology, and explanatory principles. For both Kitcher and Hull, thesecommunities are dynamic; they change and evolve as new ideas and prin-ciples develop and become established. Despite this communal aspect,individual scientists are central to both perceptions of scientific progress.Kitcher’s notion of consensus practice “changes in response to modifica-tions of individual practices.”100 For Hull, success or failure in science ispartially contingent on the ability of individual scientists to evaluate andutilize the ideas of others. Hull states, “In the ongoing process of sciencethe inherent worth of ideas is far from irrelevant, but it is also far fromsufficient. . . . Being‘right’ is not enough. Scientists must convert their fellowscientists as well.”101

Changes in Werkman’s research program support many of these notionsabout scientific progress. Werkman’s biochemical success illustrates twoimportant points: i) the complex social interactions that occur as individualscientists seek to foster their personal careers; and ii) their potential tofurther progress in disciplines external to their institutional “home.” Werkmanbecame an eminently successful scientist by his disciplinary shift away fromthe traditional bacteriology practice of his departmental colleagues. Of equalimportance, his career shift advanced the disciplinary progress of biochem-istry. Both Werkman’s laboratory and many of his students’ laboratories wereinfluenced by the evolving biochemical consensus practices; moreover, theysignificantly advanced the progress of those practices, especially in the areaof intermediary metabolism. Thus, if scientific success reflects an ability to

99 Kitcher, 1993, p. 58.100 Kitcher, 1993, p. 60.101 Hull, 1988, p. 114.


“convert” one’s “fellow scientists,” Chester Werkman developed a successfulscientific career.

Individual success in advancing scientific progress or disciplinary expan-sion, however, tells us little about the broader institutional impact of thatpersonal achievement. Werkman’s personal success provoked serious andextensive institutional repercussions. These repercussions resulted from adeeply-rooted discontinuity between personal progressiveness in advancingscience and the institutions within which scientists function. Laboratorysuccess does not guarantee administrative success. To understand the insti-tutional impact of Werkman’s scientific success we need to once again turnto the historical narrative.

Dénouement

Ultimately, Werkman’s successful shift in research focus profoundly andlamentably shaped the future of bacteriology as an academic disciplineat Iowa State. As I noted earlier, Buchanan retained the title of Bacteri-ology Department “Head” from its inception in 1910, while he concurrentlyserved various other administrative roles. In 1941, Max Levine, who effec-tively ran the department for many years as “Professor in Charge,” was inmilitary service; in Levine’s absence, Werkman was appointed “Professorin Charge,” effective in the Fall of 1942. As Werkman’s national and inter-national prestige flourished, maneuvering began to make this appointmentpermanent. Correspondence between Charles Friley, then president of IowaState, and Buchanan implies that Werkman’s research was perceived as “moremodern,” and thus Werkman was in a better position to lead the department inthe post-war years.102 Buchanan was forced into retirement, and in July 1945Werkman was formally appointed Head of the Bacteriology Department.Details on how this change came about are sketchy, but the appointment,from Buchanan’s perspective, was contentious.

Buchanan apparently did not object to retirement; however, he vigorouslyobjected to Werkman’s selection as department Head for two reasons. First,he believed that the position rightfully belonged to Levine and objected tothe administration’s precipitous action during Levine’s military absence.103 Instrongly worded letters to President Friley, Buchanan stated that if the insti-

102 Friley, 1945.103 The change was so precipitous that Levine apparently had little warning. The story ofWerkman’s appointment was announced in the Ames daily newspaper (Anonymous, 1945,p. 1). According to his daughter, Levine’s wife knew that he received the paper and wouldbe certain to see the story. So, that evening she called him at Brooke Army Hospital in SanAntonio with the news. In a letter, which enclosed a copy of the news story, Levine’s daughtercommented: “I don’t know if Dad was ever informed officially about this turn of events, but I


tution did not recognize amoral obligation to Levine, it at least had a legalobligation not to abolish Levine’s position while he was in the military.104

Second, Buchanan lacked confidence both in Werkman’s leadership abilityand the direction in which he believed Werkman would lead the department.He stated his position to President Friley: “in my opinion he [Werkman]had not shown the qualities of administrative ability and teaching leader-ship that would justify appointment as head of as important a teachingand research department as Bacteriology.”105 Buchanan’s opinion was evenshared by former Werkman students. For example, Ariel Andersen, at theUSDA Laboratory in Albany, California, wrote to Levine that Werkmanlacked “the confidence and respect of his students.”106 Andersen also depreci-ated Werkman’s teaching abilities, claiming that students “made fun” of himand considered “Werkman one of the poorest teachers on the campus.”

Buchanan’s insight was prescient, for the department did not flourishunder Werkman’s leadership. Many factors, such as a difficult and uncooper-ative administrative system, contributed to this decline, and a full discussionof the decline is beyond the scope of this essay. However, Robert Sinsheimer,who was a faculty member in the Physics Department, has commentedextensively on the many administrative difficulties he encountered at IowaState during this time.107

Perhaps the most significant cause of the department’s eclipse was adrastic difference in Werkman’s leadership style in comparison with hispredecessor. Robert Sinsheimer, for example, held a faculty appointmentin physics where he conducted biochemical research using isotopes. Giventhe fact that Werkman’s laboratory was an early leader in this arena, aready speculation is that Sinsheimer might have established a collaborativeeffort with Werkman. Yet, Sinsheimer commented: “I was not close to him[Werkman]. He was a difficult person and I think it is correct to say heran the department into the ground. He could not cope with other talent orindependence in the department and effectively forced such individuals toleave. . . . I think [that Werkman] believed in the Herr Professor model of adepartment.”108

Sinsheimer’s reflection is shared by former department members whocommented that Werkman was petty and autocratic as department Head, andthat these traits created divisiveness and led to a loss of spirit in the depart-

do know that this was where he first heard about it – that is, he read it after my mother phonedhim” (Worthen, 1995).104 Buchanan, 1945a.105 Buchanan, 1945a, 1945b.106 Andersen, 1945.107 Sinsheimer, 1994, pp. 72–73.108 Sinsheimer, 1998.


ment.109 Several of the younger faculty members left for positions elsewhere.Many highly productive members of Werkman’s own laboratory left for otherpositions, and Werkman could no longer sustain the successful laboratoryoperation of the previous decades.110 As a consequence, the biochemicalstrength that Werkman brought to the Bacteriology Department was dissi-pated. Biochemical focus initially shifted to the Chemistry Department andthen to an independent Biochemistry Department.

Perhaps the ultimate lessons of this complex and convoluted story aresimply these. Scientists can dramatically develop successful careers byfollowing disciplinary avenues outside of their primary disciplinary home. Inpursuing such routes they can both enhance their own career and make signifi-cant contributions to disciplinary progress; certainly this was the case forChester Werkman and biochemistry. Many factors, some of which are totallycontextual and contingent, influence the degree of success that individualsattain in pursuing such an approach. Nevertheless, despite the potentialfor personal success, disciplinary expansion, and scientific progress, suchsuccess has serious potential for institutional damage; certainly this was thecase for Iowa State and bacteriology.

Acknowledgments

This essay was written, in part, during a sabbatical in the Departmentof Biochemistry and the Center for Biomedical Ethics at Case WesternReserve University (CWRU); I thank Richard Hanson, Tom Murray, and theircolleagues for their generosity and support. Few historical stories are possiblewithout access to archival resources, and I thank Betty Erickson, BeckyJordan, Jill Osweiler, and Tyler Walters (Iowa State), Jeff Karr (AmericanSociety for Microbiology), Dan Barberio (National Academy of Sciences),and Geis Kuennen and Lesley Robertson (Technical University, Delft) forassistance with archived material in their institutions. The late Paul Hartman,former Head of the Bacteriology Department at Iowa State, was a valu-able resource of institutional information, as well as a friendly critic ofthis work. My Delaware colleagues, Heyward Brock and David Smith, AlanRocke at CWRU, and three anonymous reviewers atJHBmade many helpfulcomments and criticisms as the work evolved. Finally, I thank the Universityof Delaware – College of Arts and Sciences and General University ResearchFund – and the National Science Foundation (Grant Number SBR 9602023)for financial support.

109 Hartman, 1995.110 Singleton, 1994a, 1994b.


Photograph sources: Figure 1, courtesy of Kluyver Archives, TechnicalUniversity of Delft. Figure 2, courtesy of Iowa State University Library/University Archives. Figures 3 and 4, Courtesy of Mr. Robert Werkman.

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