KALCKAR, HERMAN MORITZ (b.Copenhagen, Denmark, 26 March 1908; d. Cambridge,Massachusetts, 17 May 1991), biochemistry, enzymology,molecular biology.

Kalckar’s biochemical contributions were multifac-eted. His initial work opened an investigative pathway towhat has come to be called oxidative phosphorylation. Asa mature investigator, his utilized his laboratory to play asignificant role in the rapidly evolving field of enzymol-ogy, and Kalckar introduced innovative ways of measuringenzymatic activity. A significant part of his enzymologicalwork focused on galactose metabolism, and he was one ofthe first investigators to clarify the nature of the geneticdisorder galactosemia. Like many twentieth-century bio-chemists, Kalckar did not restrict his work to any singleorganism or tissue, and he contributed equally to theunderstanding of microbial physiology and human diseases.

Early Life and Education. Kalckar was the middle ofthree sons and described his early family life as “a middle-class, Jewish-Danish family—Danish for several genera-tions.” While not financially wealthy, his family life wasintellectually rich and allowed Kalckar’s “interest in thehumanistic disciplines” to develop and thrive. His busi-nessman father, Ludvig, had an avid interest in the theaterand described seeing the 1879 world premiere of HenrikIbsen’s A Doll’s House. His mother, Bertha Rosalie (néeMelchior), was fluent in both German and French andintroduced Kalckar to writers such as Gustave Flaubert,Marcel Proust, Johann Wolfgang von Goethe, HeinrichHeine, and Gotthold Ephraim Lessing. Although the fam-

ily members were “mainly free-thinkers” and his fatherwas primarily secular, his mother was more observant andencouraged some religious education. Thus, Kalckarlearned “a minimum of Hebrew” at the Copenhagen mainsynagogue (Kalckar, 1991, p. 2).

Kalckar attended the Østre Borgerdyd Skole, locateda short distance from home, which he described as “inter-esting and rewarding” and having an “Athenian flavor,” acharacteristic that arose from the headmaster, a world-renowned Greek scholar. His early scientific educationalso benefited from “a formidable and passionatelydevoted teacher in mathematical physics,” who signifi-cantly influenced Kalckar’s decision to pursue a career inresearch and teaching. The Danish Nobel Prize–winning(Physiology or Medicine, 1920) physiologist AugustKrogh shaped Kalckar’s biological interests by somehuman physiology demonstrations. Kalckar commentedthat Krogh was “the only physiologist in the 1920s whotook an active interest in introducing the principles ofhuman physiology to Danish high school boys.” Thedemonstrations were apparently elaborate, as Krogh useda number of research instruments from his own researchprogram (Kalckar, 1991, pp. 2–3).

Kalckar’s younger brother, Fritz, was a physicist and astudent/colleague of Niels Bohr. Herman Kalckar alsoknew Bohr and was close friends with the Bohr family (hisson Niels Kalckar was named after Bohr). In 1937 FritzKalckar died from a status epilepticus attack while in Cal-ifornia. The Bohr family provided personal comfort to theKalckar family, and Niels Bohr gave a eulogy at Fritz’s cre-mation service (Kalckar, 1991, p. 8). The close Bohrfriendship may explain the intense chemical focus thatKalckar brought to biological problems.

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In 1933 Kalckar completed medical studies at theUniversity of Copenhagen and the following year begangraduate studies in the Department of Physiology. Hisresearch mentor was Ejnar Lundsgaard. In the early1930s, Lundsgaard revolutionized the way physiologistsunderstood cellular energetics by demonstrating thatphosphocreatine was involved in muscle contraction. Sev-eral simultaneous events happened when Kalckar joinedLundsgaard’s lab. The biochemist Fritz Lipmann wasforced to leave Germany, and Lundsgaard offered him aposition in Copenhagen. Also, Lundsgaard became physiology department chair, and thus his time was morerestricted. Consequently, Lipmann became Kalckar’s pri-mary mentor and encouraged him to read “the newer lit-erature on carbohydrate and phosphate metabolism inisolated phosphorylation tissue extracts or tissue particlepreparations” (Kalckar, 1991, p. 5).

Kalckar began his research career at an importantperiod in biochemistry’s history. Eugene Kennedyobserved that the central question facing many biologistsat the time was: “How is energy captured by the oxidationof sugars and other foodstuffs linked to the reduction ofmolecular oxygen?” (1996, p. 151). Kalckar very indi-rectly set about to address this question and decided tostudy phosphate and carbohydrate metabolism in kidneycortex extracts.

His use of kidney cortex tissue seems a bit surprisingbecause many workers at the time (including Hans Krebs)were working with minced pigeon breast muscle. Kalckarstated that one reason for his choice of tissue was his men-tor’s interest in carbohydrate transport in the kidney, aprocess thought “to be driven by phosphorylation-dephosphorylation cycle in the membrane.” Thus,Kalckar “set out to scout for a really vigorous phosphory-lation system that could ‘drive’” carbohydrate transport inthe kidney cortex (Kalckar, 1969, p. 171). Later, Kalckarexplained that in order to study pigeon tissue he wouldhave to train himself to decapitate pigeons, however he“greatly preferred to avoid killing animals, probably froma lack of courage.” He was fortunate to obtain fresh catand rabbit kidneys from the weekly physiology depart-ment perfusion experiments (Kalckar, 1991, p. 6).

Using Otto Warburg’s manometric technique andapparatus, Kalckar began to measure relationshipsbetween oxygen utilization and phosphorylation inminced kidney tissue. Few modern investigators canappreciate the multitude of difficulties associated with thisexperimental approach, which required measuringchanges in extremely small volumes of gases. The appara-tus was standard biochemical equipment for much of thetwentieth century; nevertheless, Efraim Racker half jok-ingly referred to the apparatus as “nystagmus inducing”(1965, p. 19). Regardless, Kalckar’s choice of this experi-

mental approach was important, because in the Warburgapparatus the minced tissue was continuously exposed tohigh oxygen levels.

Kalckar noted that his “studies on phosphorylationand respiration in kidney cortex extracts turned out to berewarding” (1991, p. 6). Prior to his work, researcherswere investigating various pathways whereby carbohy-drates are oxidized to products such as lactic acid. Theseoxidations involved a number of phosphorylation reac-tions and produced a phosphorylated compound, ulti-mately identified as adenosine triphosphate (ATP).Referred to as “glycolysis,” these reactions occurred in theabsence of oxygen. While a variety of phosphorylatedcompounds were produced in addition to ATP, the role ofphosphorylation was not understood; most biochemistsbelieved that the phosphate group somehow made thecompound more “fit” for reaction.

In a series of papers published between 1937 and1939, Kalckar established that in kidney cortex tissueslices these phosphorylation reactions were “coupled” withoxygen consumption. In these experiments, Kalckar sus-pended minced kidney cortex in phosphate buffer, whichwas then incubated with various carbohydrates in a War-burg apparatus. At various times O2 consumption wasmanometrically determined and inorganic phosphate (Pi)was chemically measured. In the presence of O2, Pi levelswere significantly reduced, whereas under anaerobic con-ditions no Pi was consumed. Carbohydrate (e.g., glucose)was also required for Pi consumption. The process, whichKalckar referred to as “aerobic phosphorylation,” was thefirst direct demonstration that carbohydrate oxidation wasdirectly “coupled” to carbohydrate phosphorylation(Kalckar, 1937; 1939; 1969, pp. 171–172); the process,later referred to as oxidative phosphorylation, is funda-mental to life, and Kalckar’s work helped establish thebasic phenomenon and opened the way to its systematicexploration (Kennedy, 1996).

Early Career. Kalckar completed work for his PhD in1939 and in January received a Rockefeller research fel-lowship to spend a year at the California Institute of Tech-nology (Caltech). After a brief visit in London, Kalckarspent much of February in St. Louis visiting Carl andGerty Cori’s Washington University laboratory, which wasthen doing some of the most innovative biochemical workin the United States. The Cori lab had, unsuccessfully,tried to reproduce some of Kalckar’s published work.Kalckar noted that in their technique, which involved “theold-fashioned Meyerhof extract technique, using testtubes,” the tissue extracts were static. In Kalckar’s tech-nique, using the Warburg manometer, the tissue extractwas vigorously shaken and thus highly aerated. Kalckarworked with the Coris’ graduate student, Sidney

Kalckar Kalckar

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Colowick, and quickly demonstrated that when the cortexextracts were shaken, oxidative phosphorylation was easilyobserved (Kalckar, 1991, p. 10).

In early March 1939, Kalckar arrived in Pasadena,where he and his wife Vibeke Meyer rapidly becameinvolved in the Caltech intellectual and social community,which included Max Delbrück, Linus Pauling, and twoBonner brothers, James and David. The Delbrück friend-ship, which had initially begun with a brief Copenhagenmeeting, became lifelong. The Bonner brothers intro-duced Kalckar to various members of the Caltech faculty,and helped Herman and Vibeke settle into an Americanlife by arranging housing and the purchase of a car.

Most likely at Delbrück’s encouragement, Kalckarattended Cornelius B. van Niel’s popular microbiologycourse taught at Pacific Grove. Van Niel was responsiblefor introducing numerous American scientists to the com-plexity of the microbial world. As a student of Albert JanKluyver, van Niel spread the gospel of biochemical unity,which Kluyver expressed in the aphorism, “From the ele-phant to the butyric acid bacterium it is all the same!”(Kamp et al., 1959, p. 20). The Pacific Grove experience“may have planted a seed that led later to Kalckar’s inter-est in microbial molecular biology” (Kennedy, 1996, p.153). Throughout his career, Kalckar, like many of hiscontemporaries, frequently turned to bacterial systems toaddress questions unresolvable in more complex biologicalsystems.

The Pauling connection proved especially valuable.Most likely from his familiarity with Bohr’s atomic con-cepts, Kalckar was influenced by the potential impact ofPauling’s chemical ideas on biological problems. At Paul-ing’s encouragement, Kalckar wrote an extensive review inwhich he developed many of our modern concepts ofbioenergetics, including the notion that oxidation reac-tions are “coupled” to phosphorylation reactions via ATP(Kalckar, 1941).

Although not widely appreciated at the turn of thetwenty-first century, Kalckar’s 1941 Chemical Reviewspaper influenced the way many scientists thought aboutthe energetics of life processes. One reason for this lack ofappreciation perhaps lies in the fact that Fritz Lipmannalso published a similar paper, in which he developed the“high energy bond” concept, in 1941. While the Kalckarpaper was influential in shaping scientists’ views aboutbioenergetics, the Lipmann paper was more frequentlycited. Nevertheless, Kalckar viewed his relationship withLipmann as collaborative, stating: “While Lipmann waspreaching the gospel of phosphorylation and the ‘highenergy phosphate bond’ on the eastern seaboard … I wasbeginning my missionary work at the same time in the‘Pacific triangle’—Caltech, Berkeley, and Stanford”(Kalckar, 1992, p. 43).

The outbreak of World War II, and German occupa-tion of Denmark, stranded the Kalckars in the UnitedStates. Because of his earlier visit in the Cori lab, in 1940Kalckar received an appointment as research fellow in thePharmacology Department at Washington University, andhe and Vibeke moved to St. Louis. Kalckar resumed workwith Colowick studying phosphorylation reactions inmuscle tissue. The work was important in terms ofKalckar’s scientific maturation for two reasons: first, hebegan to focus more on individual enzymatic reactions,even partially purifying the responsible protein, than hehad previously; second, he and Colowick discovered theenzyme myokinase (now called adenylate kinase), whichcatalyzed the reversible reaction:

ATP + AMP o 2 ADP (Reaction 1)

Because many biochemical processes led to adenosinemonophosphate (AMP) production, the enzyme wasimportant in helping replenish cellular levels of ATP.Shortly after the collaboration began, Colowick wasdrafted to do war research, and Kalckar finished themyokinase characterization by himself.

In 1943 Oliver Howe Lowry invited Kalckar to joinhis lab at the New York Public Health Institute as aresearch associate. The appointment helped advanceKalckar’s enzymology research in part because, as PaulBerg commented, Kalckar “developed a whole newapproach to being able to use enzymes in a novel way”(2000, p. 26). The lab had a new, and relatively rare,Beckman DU ultraviolet spectrophotometer, and Kalckarrapidly became a virtuoso of the instrument, using it todevelop a number of novel enzyme assay techniques. Thework culminated in a series of three papers on purinemetabolism enzymes (Kalckar, 1947a, b, c). All threepapers were highly cited (3,200 citations in 2006), andthe third paper was recognized as a “Citation Classic” in1984. In a brief commentary on the paper, Kalckar notedthat the paper’s popularity most likely arose from itspotential clinical applications such as diagnosing diseasessuch as arthritic urica or Lesch-Nyhan syndrome, a seriousinborn metabolic error in infants (Kalckar, 1984).

Return to Denmark. When the war ended, Lundsgaardarranged for Kalckar to return to Copenhagen in 1946where he ran the new Cytofysiologisk Institute, funded bythe Rockefeller Foundation, Lederle Laboratories, as wellas the Danish Carlsberg Foundation. His research focusedprimarily on the enzymology of nucleoside and nucleotidemetabolism, and the lab rapidly became a magnet attract-ing numerous bright young biochemists from around theworld, including Americans such as Paul Berg, MorrisFriedkin, Walter McNutt, Günter Stent, and James Watson.

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Shortly after his return to Copenhagen, Kalckar’smarriage to Vibeke dissolved. He married Barbara Wright,a developmental biologist and American postdoctoral fel-low in his lab. Three children, Sonja, Nina, and Niels,were born of this marriage (Kennedy, 1996).

In early 1950, Kalckar suggested that such nucleo-tides as uridine diphosphate (UDP), UDP-glucose, orUDP-galactose were involved in the conversion of glucose-1-P to galactose-1-P, and his laboratory began tofocus on galactose metabolism in microbial and animaltissues, a research program that dominated the rest of his career.

Return to the United States. Bernard Horecker, at theNational Institutes of Health (NIH), invited Kalckar tocome to Bethesda, Maryland, as a visiting scientist in1952; later the appointment was made permanent in theNational Institute of Arthritis and Metabolic Diseases. Atthe NIH, Kalckar expanded his research program on

galactose metabolism and developed an interest in thegenetic disease, galactosemia. The disease, which is char-acterized by liver enlargement, kidney failure, and mentalretardation, occurs because of galactose (formed from themilk component lactose) accumulation. In untreatedinfants, mortality is as high as 75 percent. The prevailingview on galactosemia in 1952 is summarized in Reactions(2) through (4).

Lactose (in milk ) → Glucose + Galactose(Reaction 2)

Galactose + ATP o Galactose-1-P (Gal-1-P) (Reaction 3)

Gal-1-P o Glucose-1-P (Reaction 4)

Reaction 3 is catalyzed by an enzyme called an“epimerase,” and investigators speculated that galac-tosemia arose from a defect in this enzyme (which wouldlead to galactose accumulation via reversal of Reaction 3).

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Kalckar’s “Cytofysiologisk Institut” 1950. Back row, from left: Gunther Stent, Niels Ole Kjeldgaard, Hans Klenow, Jim Watson,and Vincent Price. Front row, from left: Kalckar, Audrey Jarnum, Jytte Heisel, Eugene Goldwasser, Walter McNutt, and E. Hoff-Jorgensen. COURTESY OF STEEN GAMMELTOFT, M.D.

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For a variety of reasons, Kalckar suspected anotherenzyme was defective; he thought the enzyme blocked wasGalactose-1-P uridyl transferase (GALT), which catalyzesReaction 5:

Gal-1-P + UDP-glucose o UDP-galactose + Glucose-1-P (Reaction 5)

If this enzyme was not functioning, Gal-1-P would accu-mulate leading to galactose build up through reversal ofReaction 3. Although galactosemia is relatively rare,Kalckar’s associates were able to identify several afflictedindividuals and obtain blood samples from them. IfKalckar’s hypothesis was correct, the analysis should showtwo features. First, individuals with the disease shouldhave decreased levels of GALT. Second, epimerase levels inafflicted individuals should be normal. Both features wereobserved in blood samples from galactosemia patientswhen compared with normal subjects (Kalckar, et al.,1956a and b).

In addition to a rich scientific environment, the NIHcommunity offered personal advantages. Unlike manyinstitutions at the time, the NIH had no official prohibi-tions regarding employing scientific couples; thusKalckar’s wife, Barbara Wright, also had a staff appoint-ment. The environment was both intellectually andsocially rewarding. Other married couples working at theNIH at the same time included: Thressa and Earl Stadt-man; Marjorie and Evan Horning; and Martha Vaughanand Jack Orloff (Park, 2002).

In 1958 William McElroy offered Kalckar a full pro-fessorship in biology at Johns Hopkins University, wherehis biochemical interests in galactose metabolism contin-ued. Perhaps reflecting the long-term influence of vanNiel’s course, his research focus shifted to bacterial systems.

While at Hopkins, Kalckar suggested (in a Naturepaper) that the effects of isotope fallout from nuclearweapons testing could be measured by analysis of strontium-90 levels in children’s deciduous teeth (Kalckar,1958). The proposal was both important and original andelicited numerous state and local organizations to collectteeth for analysis; the data did indeed show a correlationbetween isotope levels and nuclear testing. Arguably theresulting program helped encourage public sentiment toban atmospheric nuclear weapons testing.

In 1961 Kalckar moved again, to the Harvard Med-ical School where he was Henry S. Wellcome ResearchBiochemist and headed the Massachusetts General Hospi-tal Biochemical Research Laboratory, a position previ-ously held by Fritz Lipmann. His research programcontinued to focus on bacterial galactose utilization, how-ever it tended to become more physiologically oriented(Wellcome Trust, 1963, p. 16). He became interested in

cellular sensory and signaling processes. For example, incollaboration with Winfried Boos, he isolated a galactosebinding protein in Escherichia coli that played an impor-tant role in cellular galactose transport. Later workdemonstrated that the same protein played an importantrole in E. coli ’s chemotactic response to galactose.

The Boston move also had significant personal rami-fications for Kalckar: his marriage to Barbara Wright dis-solved; he renewed a friendship with Agnete Fridericia,whom he had known as a student in Copenhagen. Kalckarcommented that their “cheerful conversations during the1960s, most of it in Danish, changed [his] life”; in 1968,he and Agnete were married (Kalckar, 1991, p. 27).

Kalckar retired as head of the Biochemical ResearchLaboratory in 1974 but remained at Massachusetts Gen-eral as visiting professor. In 1979 he was appointed as adistinguished research professor in the Boston Universitychemistry department, a position that permitted him tocontinue his research work for the rest of his life(Kennedy, 1996, p. 159).

In a scientific career that spanned almost six decadesof the twentieth century, Herman Kalckar witnessedmany fundamental changes in biochemistry. Althoughurease had been crystallized in 1926, in 1934—whenKalckar began his graduate studies—some biochemistswere still debating the nature of enzymes. At his death,after having been a founder of modern bioenergetics andenzymology, he was studying the genes responsible forenzyme action. These achievements received numerousrecognitions: he was elected to the National Academy ofSciences, the Royal Danish Academy, and the AmericanAcademy of Arts and Sciences; he received honorarydegrees from Washington University, the University ofChicago, and the University of Copenhagen.

On a personal level, Kalckar often appears enigmatic.On the one hand, Eugene Kennedy noted “The sweep ofhis intellect was very broad, his spirit was open and gen-erous, and he had a wonderful sense of humor” (Kennedy,1996, p. 160). Former associates speak fondly of excitingscientific discussions, which might be interrupted to talkabout art, music, even Nordic mythology. As his work onstrontium-90 levels in children’s teeth suggests, Kalckarwas passionately concerned about social issues like thespread of nuclear weapons and weapons testing.

However, Kalckar was equally known for his peculiarway of talking, which some hearers found unintelligible.He would, for example, often begin an argument with aconclusion and work backward to the premises. Boos, hisformer associate, referred to this speech mode as a “lan-guage” he called “Kalckarian”: a language that Kalckaroccasionally used to avoid conversations he found uninteresting.

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Boos further observed that Kalckar divided the scien-tific world into two types. The Apollonian was a capabletechnician, who accumulated data to tell clear, logical sto-ries, a perspective Kalckar found boring and lacking anysense of humor. The Dionysian, however, was more inter-ested in the answers, even if they come in a dream, thanin the logic and proof of the answer. For Boos, Kalckar“was the archetype of a Dionysian” (Boos, 1991, p. 8).

The term Dionysian can be used in another sense,related to the Roman incarnation of Bacchus, the God ofwine and other pleasures. In this view, an individual isopen to all that life offers and is not bound in a singleworldview; Kalckar seems to fit this image as well. Sciencewas one of life’s rich rewards; good music, good friends, acornucopia of human experiences were equally valuable.

B I B L I O G R A P H Y

WORKS BY KALCKAR

“Phosphorylation in Kidney Tissue.” Enzymologia 2 (1937):47–53.

“The Nature of Phosphoric Esters Formed in Kidney Extracts.”Biochemical Journal 33, no. 5 (1939): 631–641.

“The Nature of Energetic Coupling in Biological Syntheses.”Chemical Reviews 28 (1941): 71–178.

With Sidney P. Colowick and Carl F. Cori. “GlucosePhosphorylation and Oxidation in Cell-Free Tissue Extracts.”Journal of Biological Chemistry 137 (1941): 343–356.

“The Enzymatic Action of Myokinase.” Journal of BiologicalChemistry 143 (1942): 299–300.

“The Role of Myokinase in Transphosphorylations: II. TheEnzymatic Action of Myokinase on Adenine Nucleotides.”Journal of Biological Chemistry 148 (1943): 127–137.

With Manya Shafran. “Differential Spectrophotometry of PurineCompounds by Means of Specific Enzymes: I. Determinationof Hydroxypurine Compounds.” Journal of BiologicalChemistry 167 (1947a): 429–443.

With Alice Neuman Bessmann. “Differential Spectrophotometryof Purine Compounds by Means of Specific Enzymes: II.Determination of Adenine Compounds.” Journal ofBiological Chemistry 167 (1947b): 445–459.

With Manya Shafran. “Differential Spectrophotometry of PurineCompounds by Means of Specific Enzymes: III. Studies ofthe Enzymes of Purine Metabolism.” Journal of BiologicalChemistry 167 (1947c): 461–475.

With Elizabeth P. Anderson and Kurt J. Isselbacher.“Galactosemia, a Congenital Defect in a NucleotideTransferase.” Biochimica et Biophysica Acta 20 (1956a):262–268.

With Elizabeth P. Anderson, Kurt J. Isselbacher, and KiyoshiKurahashit. “Congenital Galactosemia, a Single EnzymaticBlock in Galactose Metabolism.” Science 123 (1956b):635–636.

“An International Milk Teeth Radiation Census.” Nature 182(1958): 283–284.

Editor. Biological Phosphorylations: Development of Concepts.Englewood Cliffs, NJ: Prentice-Hall, 1969.

“This Week’s Citation Classic.” Current Contents 26 (1984): 148.

“50 Years of Biological Research: From OxidativePhosphorylation to Energy Requiring Transport Regulation.”Annual Review of Biochemistry 60 (1991): 1–37.

“High Energy Phosphate Bonds: Optional or Obligatory?” InPhage and the Origins of Molecular Biology, expanded ed.,edited by John Cairns, Gunther S. Stent, and James D.Watson. Cold Spring Harbor, NY: Cold Spring HarborLaboratory Press, 1992.

OTHER SOURCES

Berg, Paul. “A Stanford Professor’s Career in Biochemistry,Science Politics, and the Biotechnology Industry.” Berkeley:Regional Oral History Office, Bancroft Library, University ofCalifornia, 2000. Available fromhttp://www.calisphere.universityofcalifornia.edu. An oralhistory conducted in 1997 by Sally Smith Hughes.

Boos, Winfried. “Farewell to a Dionysian.” Biological ChemistryHoppe-Seyler 372, no. 11 (1991): 8–9. Original in German;translation kindly provided by Dr. Boos.

Garfield, Eugene. “Highly Cited Articles: 35. BiochemistryPapers Published in the 1940s.” Current Contents 8 (1977):5–11. Reprinted in: Essays of an Information Scientist, Vol. 3.Philadelphia, PA: ISI Press. Available fromhttp://www.garfield.library.upenn.edu/essays.html.

Kamp, A. F., J. W. M. La Rivière, and W. Verhoeven. “Kluyveras Professor: Chronicles of the Laboratory.” In their AlbertJan Kluyver: His Life and Work, pp. 14–48. New York:Interscience Publishers, 1959.

Kennedy, Eugene P. “Herman Moritz Kalckar, March 26,1908–May 17, 1991.” Biographical Memoirs, NationalAcademy of Sciences (U.S.) 69 (1996): 148–165.

Lipmann, F. “Metabolic Generation and Utilization ofPhosphate Bond Energy.” Advances in Enzymology 1 (1941):99–162.

Park, Buhm Soon. “Historian’s Perspective: NIH Offered Havenfrom Antinepotism Rules.” Newsletter of the NIH AlumniAssociation 14, no. 2 (2002): 18–19. Available fromhttp://www.fnih.org/nihaa/NIHAA-Update.pdf.

Racker, Efraim. Mechanisms in Bioenergetics. New York:Academic Press, 1965.

Wellcome Trust. Fourth Report, Covering the Period 1960–1962.London: Author, 1963.

Rivers Singleton Jr.

KAMMERER, PAUL (b. Vienna, Austria-Hun-gary, 17 August 1880; d. Puchberg am Schneeberg, LowerAustria, 23 September 1926), zoology, heredity, evolution.

The main goal of Kammerer’s research was to demon-strate the modifying power of the environment and the heritability of acquired characteristics, using the

Kalckar Kammerer

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