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KALCKAR, HERMAN MORITZ (b. ily members were “mainly free-thinkers” and his father Copenhagen, , 26 March 1908; d. Cambridge, was primarily secular, his mother was more observant and Massachusetts, 17 May 1991), , enzymology, encouraged some religious education. Thus, Kalckar molecular biology. learned “a minimum of Hebrew” at the Copenhagen main synagogue (Kalckar, 1991, p. 2). Kalckar’s biochemical contributions were multifac- eted. His initial work opened an investigative pathway to Kalckar attended the Østre Borgerdyd Skole, located what has come to be called oxidative phosphorylation. As a short distance from home, which he described as “inter- a mature investigator, his utilized his laboratory to play a esting and rewarding” and having an “Athenian flavor,” a characteristic that arose from the headmaster, a world- significant role in the rapidly evolving field of enzymol- renowned Greek scholar. His early scientific education ogy, and Kalckar introduced innovative ways of measuring also benefited from “a formidable and passionately enzymatic activity. A significant part of his enzymological devoted teacher in mathematical physics,” who signifi- work focused on galactose metabolism, and he was one of cantly influenced Kalckar’s decision to pursue a career in the first investigators to clarify the nature of the genetic research and teaching. The Danish Nobel Prize–winning disorder galactosemia. Like many twentieth-century bio- (Physiology or Medicine, 1920) physiologist August chemists, Kalckar did not restrict his work to any single Krogh shaped Kalckar’s biological interests by some organism or tissue, and he contributed equally to the human physiology demonstrations. Kalckar commented understanding of microbial physiology and human that Krogh was “the only physiologist in the 1920s who diseases. took an active interest in introducing the principles of human physiology to Danish high school boys.” The Early Life and Education. Kalckar was the middle of demonstrations were apparently elaborate, as Krogh used three sons and described his early family life as “a middle- a number of research instruments from his own research class, Jewish-Danish family—Danish for several genera- program (Kalckar, 1991, pp. 2–3). tions.” While not financially wealthy, his family life was Kalckar’s younger brother, Fritz, was a physicist and a intellectually rich and allowed Kalckar’s “interest in the student/colleague of Niels Bohr. Herman Kalckar also humanistic disciplines” to develop and thrive. His busi- knew Bohr and was close friends with the Bohr family (his nessman father, Ludvig, had an avid interest in the theater son Niels Kalckar was named after Bohr). In 1937 Fritz and described seeing the 1879 world premiere of Henrik Kalckar died from a status epilepticus attack while in Cal- Ibsen’s A Doll’s House. His mother, Bertha Rosalie (née ifornia. The Bohr family provided personal comfort to the Melchior), was fluent in both German and French and Kalckar family, and Niels Bohr gave a eulogy at Fritz’s cre- introduced Kalckar to writers such as Gustave Flaubert, mation service (Kalckar, 1991, p. 8). The close Bohr Marcel Proust, Johann Wolfgang von Goethe, Heinrich friendship may explain the intense chemical focus that Heine, and Gotthold Ephraim Lessing. Although the fam- Kalckar brought to biological problems.

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In 1933 Kalckar completed medical studies at the mental approach was important, because in the Warburg and the following year began apparatus the minced tissue was continuously exposed to graduate studies in the Department of Physiology. His high oxygen levels. research mentor was Ejnar Lundsgaard. In the early Kalckar noted that his “studies on phosphorylation 1930s, Lundsgaard revolutionized the way physiologists and respiration in kidney cortex extracts turned out to be understood cellular energetics by demonstrating that rewarding” (1991, p. 6). Prior to his work, researchers phosphocreatine was involved in muscle contraction. Sev- were investigating various pathways whereby carbohy- eral simultaneous events happened when Kalckar joined drates are oxidized to products such as lactic acid. These Lundsgaard’s lab. The biochemist Fritz Lipmann was oxidations involved a number of phosphorylation reac- forced to leave Germany, and Lundsgaard offered him a tions and produced a phosphorylated compound, ulti- position in Copenhagen. Also, Lundsgaard became mately identified as (ATP). physiology department chair, and thus his time was more Referred to as “,” these reactions occurred in the restricted. Consequently, Lipmann became Kalckar’s pri- absence of oxygen. While a variety of phosphorylated mary mentor and encouraged him to read “the newer lit- compounds were produced in addition to ATP, the role of erature on carbohydrate and phosphate metabolism in phosphorylation was not understood; most biochemists isolated phosphorylation tissue extracts or tissue particle believed that the phosphate group somehow made the preparations” (Kalckar, 1991, p. 5). compound more “fit” for reaction. Kalckar began his research career at an important In a series of papers published between 1937 and period in biochemistry’s history. Eugene Kennedy 1939, Kalckar established that in kidney cortex tissue observed that the central question facing many biologists slices these phosphorylation reactions were “coupled” with at the time was: “How is energy captured by the oxidation oxygen consumption. In these experiments, Kalckar sus- of sugars and other foodstuffs linked to the reduction of pended minced kidney cortex in phosphate buffer, which molecular oxygen?” (1996, p. 151). Kalckar very indi- was then incubated with various carbohydrates in a War- rectly set about to address this question and decided to burg apparatus. At various times O2 consumption was study phosphate and carbohydrate metabolism in kidney manometrically determined and inorganic phosphate (Pi) cortex extracts. was chemically measured. In the presence of O2, Pi levels His use of kidney cortex tissue seems a bit surprising were significantly reduced, whereas under anaerobic con-

because many workers at the time (including Hans Krebs) ditions no Pi was consumed. Carbohydrate (e.g., glucose) were working with minced pigeon breast muscle. Kalckar was also required for Pi consumption. The process, which stated that one reason for his choice of tissue was his men- Kalckar referred to as “aerobic phosphorylation,” was the tor’s interest in carbohydrate transport in the kidney, a first direct demonstration that carbohydrate oxidation was process thought “to be driven by phosphorylation- directly “coupled” to carbohydrate phosphorylation dephosphorylation cycle in the membrane.” Thus, (Kalckar, 1937; 1939; 1969, pp. 171–172); the process, Kalckar “set out to scout for a really vigorous phosphory- later referred to as oxidative phosphorylation, is funda- lation system that could ‘drive’” carbohydrate transport in mental to life, and Kalckar’s work helped establish the the kidney cortex (Kalckar, 1969, p. 171). Later, Kalckar basic phenomenon and opened the way to its systematic explained that in order to study pigeon tissue he would exploration (Kennedy, 1996). have to train himself to decapitate pigeons, however he “greatly preferred to avoid killing animals, probably from Early Career. Kalckar completed work for his PhD in a lack of courage.” He was fortunate to obtain fresh cat 1939 and in January received a Rockefeller research fel- and rabbit kidneys from the weekly physiology depart- lowship to spend a year at the California Institute of Tech- ment perfusion experiments (Kalckar, 1991, p. 6). nology (Caltech). After a brief visit in London, Kalckar Using Otto Warburg’s manometric technique and spent much of February in St. Louis visiting Carl and apparatus, Kalckar began to measure relationships Gerty Cori’s Washington University laboratory, which was between oxygen utilization and phosphorylation in then doing some of the most innovative biochemical work minced kidney tissue. Few modern investigators can in the United States. The Cori lab had, unsuccessfully, appreciate the multitude of difficulties associated with this tried to reproduce some of Kalckar’s published work. experimental approach, which required measuring Kalckar noted that in their technique, which involved “the changes in extremely small volumes of gases. The appara- old-fashioned Meyerhof extract technique, using test tus was standard biochemical equipment for much of the tubes,” the tissue extracts were static. In Kalckar’s tech- twentieth century; nevertheless, Efraim Racker half jok- nique, using the Warburg manometer, the tissue extract ingly referred to the apparatus as “nystagmus inducing” was vigorously shaken and thus highly aerated. Kalckar (1965, p. 19). Regardless, Kalckar’s choice of this experi- worked with the Coris’ graduate student, Sidney

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Colowick, and quickly demonstrated that when the cortex The outbreak of World War II, and German occupa- extracts were shaken, oxidative phosphorylation was easily tion of Denmark, stranded the Kalckars in the United observed (Kalckar, 1991, p. 10). States. Because of his earlier visit in the Cori lab, in 1940 In early March 1939, Kalckar arrived in Pasadena, Kalckar received an appointment as research fellow in the where he and his wife Vibeke Meyer rapidly became Pharmacology Department at Washington University, and involved in the Caltech intellectual and social community, he and Vibeke moved to St. Louis. Kalckar resumed work which included Max Delbrück, , and two with Colowick studying phosphorylation reactions in Bonner brothers, James and David. The Delbrück friend- muscle tissue. The work was important in terms of ship, which had initially begun with a brief Copenhagen Kalckar’s scientific maturation for two reasons: first, he meeting, became lifelong. The Bonner brothers intro- began to focus more on individual enzymatic reactions, duced Kalckar to various members of the Caltech faculty, even partially purifying the responsible protein, than he and helped Herman and Vibeke settle into an American had previously; second, he and Colowick discovered the life by arranging housing and the purchase of a car. enzyme myokinase (now called adenylate kinase), which Most likely at Delbrück’s encouragement, Kalckar catalyzed the reversible reaction: attended Cornelius B. van Niel’s popular microbiology ATP + AMP o 2 ADP (Reaction 1) course taught at Pacific Grove. Van Niel was responsible for introducing numerous American scientists to the com- Because many biochemical processes led to adenosine plexity of the microbial world. As a student of Albert Jan monophosphate (AMP) production, the enzyme was Kluyver, van Niel spread the gospel of biochemical unity, important in helping replenish cellular levels of ATP. which Kluyver expressed in the aphorism, “From the ele- Shortly after the collaboration began, Colowick was phant to the butyric acid bacterium it is all the same!” drafted to do war research, and Kalckar finished the (Kamp et al., 1959, p. 20). The Pacific Grove experience myokinase characterization by himself. “may have planted a seed that led later to Kalckar’s inter- In 1943 Oliver Howe Lowry invited Kalckar to join est in microbial molecular biology” (Kennedy, 1996, p. his lab at the New York Public Health Institute as a 153). Throughout his career, Kalckar, like many of his research associate. The appointment helped advance contemporaries, frequently turned to bacterial systems to Kalckar’s enzymology research in part because, as Paul address questions unresolvable in more complex biological Berg commented, Kalckar “developed a whole new systems. approach to being able to use enzymes in a novel way” The Pauling connection proved especially valuable. (2000, p. 26). The lab had a new, and relatively rare, Most likely from his familiarity with Bohr’s atomic con- Beckman DU ultraviolet spectrophotometer, and Kalckar cepts, Kalckar was influenced by the potential impact of rapidly became a virtuoso of the instrument, using it to Pauling’s chemical ideas on biological problems. At Paul- develop a number of novel enzyme assay techniques. The ing’s encouragement, Kalckar wrote an extensive review in work culminated in a series of three papers on purine which he developed many of our modern concepts of metabolism enzymes (Kalckar, 1947a, b, c). All three bioenergetics, including the notion that oxidation reac- papers were highly cited (3,200 citations in 2006), and tions are “coupled” to phosphorylation reactions via ATP the third paper was recognized as a “Citation Classic” in (Kalckar, 1941). 1984. In a brief commentary on the paper, Kalckar noted Although not widely appreciated at the turn of the that the paper’s popularity most likely arose from its twenty-first century, Kalckar’s 1941 Chemical Reviews potential clinical applications such as diagnosing diseases paper influenced the way many scientists thought about such as arthritic urica or Lesch-Nyhan syndrome, a serious the energetics of life processes. One reason for this lack of inborn metabolic error in infants (Kalckar, 1984). appreciation perhaps lies in the fact that Fritz Lipmann also published a similar paper, in which he developed the Return to Denmark. When the war ended, Lundsgaard “high energy bond” concept, in 1941. While the Kalckar arranged for Kalckar to return to Copenhagen in 1946 paper was influential in shaping scientists’ views about where he ran the new Cytofysiologisk Institute, funded by bioenergetics, the Lipmann paper was more frequently the Rockefeller Foundation, Lederle Laboratories, as well cited. Nevertheless, Kalckar viewed his relationship with as the Danish Carlsberg Foundation. His research focused Lipmann as collaborative, stating: “While Lipmann was primarily on the enzymology of nucleoside and preaching the gospel of phosphorylation and the ‘high metabolism, and the lab rapidly became a magnet attract- energy phosphate bond’ on the eastern seaboard … I was ing numerous bright young biochemists from around the beginning my missionary work at the same time in the world, including Americans such as , Morris ‘Pacific triangle’—Caltech, Berkeley, and Stanford” Friedkin, Walter McNutt, Günter Stent, and James (Kalckar, 1992, p. 43). Watson.

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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.

Shortly after his return to Copenhagen, Kalckar’s galactose metabolism and developed an interest in the marriage to Vibeke dissolved. He married Barbara Wright, genetic disease, galactosemia. The disease, which is char- a developmental biologist and American postdoctoral fel- acterized by liver enlargement, kidney failure, and mental low in his lab. Three children, Sonja, Nina, and Niels, retardation, occurs because of galactose (formed from the were born of this marriage (Kennedy, 1996). milk component lactose) accumulation. In untreated In early 1950, Kalckar suggested that such nucleo- infants, mortality is as high as 75 percent. The prevailing tides as uridine diphosphate (UDP), UDP-glucose, or view on galactosemia in 1952 is summarized in Reactions UDP-galactose were involved in the conversion of (2) through (4). glucose-1-P to galactose-1-P, and his laboratory began to focus on galactose metabolism in microbial and animal Lactose (in milk ) → Glucose + Galactose tissues, a research program that dominated the rest of (Reaction 2) his career. Galactose + ATP o Galactose-1-P (Gal- 1-P) (Reaction 3) Return to the United States. Bernard Horecker, at the Gal-1-P o Glucose-1-P (Reaction 4) National Institutes of Health (NIH), invited Kalckar to come to Bethesda, Maryland, as a visiting scientist in Reaction 3 is catalyzed by an enzyme called an 1952; later the appointment was made permanent in the “epimerase,” and investigators speculated that galac- National Institute of Arthritis and Metabolic Diseases. At tosemia arose from a defect in this enzyme (which would the NIH, Kalckar expanded his research program on lead to galactose accumulation via reversal of Reaction 3).

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For a variety of reasons, Kalckar suspected another cellular sensory and signaling processes. For example, in enzyme was defective; he thought the enzyme blocked was collaboration with Winfried Boos, he isolated a galactose Galactose-1-P uridyl transferase (GALT), which catalyzes binding protein in Escherichia coli that played an impor- Reaction 5: tant role in cellular galactose transport. Later work demonstrated that the same protein played an important Gal-1-P + UDP-glucose o UDP- role in E. coli ’s chemotactic response to galactose. galactose + Glucose-1-P (Reaction 5) The Boston move also had significant personal rami- If this enzyme was not functioning, Gal-1-P would accu- fications for Kalckar: his marriage to Barbara Wright dis- mulate leading to galactose build up through reversal of solved; he renewed a friendship with Agnete Fridericia, Reaction 3. Although galactosemia is relatively rare, whom he had known as a student in Copenhagen. Kalckar Kalckar’s associates were able to identify several afflicted commented that their “cheerful conversations during the individuals and obtain blood samples from them. If 1960s, most of it in Danish, changed [his] life”; in 1968, Kalckar’s hypothesis was correct, the analysis should show he and Agnete were married (Kalckar, 1991, p. 27). two features. First, individuals with the disease should Kalckar retired as head of the Biochemical Research have decreased levels of GALT. Second, epimerase levels in Laboratory in 1974 but remained at Massachusetts Gen- afflicted individuals should be normal. Both features were eral as visiting professor. In 1979 he was appointed as a observed in blood samples from galactosemia patients distinguished research professor in the Boston University when compared with normal subjects (Kalckar, et al., chemistry department, a position that permitted him to 1956a and b). continue his research work for the rest of his life In addition to a rich scientific environment, the NIH (Kennedy, 1996, p. 159). community offered personal advantages. Unlike many In a scientific career that spanned almost six decades institutions at the time, the NIH had no official prohibi- of the twentieth century, Herman Kalckar witnessed tions regarding employing scientific couples; thus many fundamental changes in biochemistry. Although Kalckar’s wife, Barbara Wright, also had a staff appoint- urease had been crystallized in 1926, in 1934—when ment. The environment was both intellectually and Kalckar began his graduate studies—some biochemists socially rewarding. Other married couples working at the were still debating the nature of enzymes. At his death, NIH at the same time included: Thressa and Earl Stadt- man; Marjorie and Evan Horning; and Martha Vaughan after having been a founder of modern bioenergetics and and Jack Orloff (Park, 2002). enzymology, he was studying the genes responsible for enzyme action. These achievements received numerous In 1958 William McElroy offered Kalckar a full pro- recognitions: he was elected to the National Academy of fessorship in biology at , where Sciences, the Royal Danish Academy, and the American his biochemical interests in galactose metabolism contin- Academy of Arts and Sciences; he received honorary ued. Perhaps reflecting the long-term influence of van degrees from Washington University, the University of Niel’s course, his research focus shifted to bacterial systems. Chicago, and the University of Copenhagen. While at Hopkins, Kalckar suggested (in a Nature On a personal level, Kalckar often appears enigmatic. paper) that the effects of isotope fallout from nuclear On the one hand, Eugene Kennedy noted “The sweep of weapons testing could be measured by analysis of his intellect was very broad, his spirit was open and gen- strontium-90 levels in children’s deciduous teeth (Kalckar, erous, and he had a wonderful sense of humor” (Kennedy, 1958). The proposal was both important and original and 1996, p. 160). Former associates speak fondly of exciting elicited numerous state and local organizations to collect scientific discussions, which might be interrupted to talk teeth for analysis; the data did indeed show a correlation about art, music, even Nordic mythology. As his work on between isotope levels and nuclear testing. Arguably the strontium-90 levels in children’s teeth suggests, Kalckar resulting program helped encourage public sentiment to was passionately concerned about social issues like the ban atmospheric nuclear weapons testing. spread of nuclear weapons and weapons testing. In 1961 Kalckar moved again, to the Harvard Med- However, Kalckar was equally known for his peculiar ical School where he was Henry S. Wellcome Research way of talking, which some hearers found unintelligible. Biochemist and headed the Massachusetts General Hospi- He would, for example, often begin an argument with a tal Biochemical Research Laboratory, a position previ- conclusion and work backward to the premises. Boos, his ously held by Fritz Lipmann. His research program former associate, referred to this speech mode as a “lan- continued to focus on bacterial galactose utilization, how- guage” he called “Kalckarian”: a language that Kalckar ever it tended to become more physiologically oriented occasionally used to avoid conversations he found (Wellcome Trust, 1963, p. 16). He became interested in uninteresting.

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Boos further observed that Kalckar divided the scien- Editor. Biological Phosphorylations: Development of Concepts. tific world into two types. The Apollonian was a capable Englewood Cliffs, NJ: Prentice-Hall, 1969. technician, who accumulated data to tell clear, logical sto- “This Week’s Citation Classic.” Current Contents 26 (1984): 148. ries, a perspective Kalckar found boring and lacking any “50 Years of Biological Research: From Oxidative sense of humor. The Dionysian, however, was more inter- Phosphorylation to Energy Requiring Transport Regulation.” ested in the answers, even if they come in a dream, than Annual Review of Biochemistry 60 (1991): 1–37. in the logic and proof of the answer. For Boos, Kalckar “High Energy Phosphate Bonds: Optional or Obligatory?” In Phage and the Origins of Molecular Biology, expanded ed., “was the archetype of a Dionysian” (Boos, 1991, p. 8). edited by John Cairns, Gunther S. Stent, and James D. The term Dionysian can be used in another sense, Watson. Cold Spring Harbor, NY: Cold Spring Harbor related to the Roman incarnation of Bacchus, the God of Laboratory Press, 1992. wine and other pleasures. In this view, an individual is open to all that life offers and is not bound in a single OTHER SOURCES worldview; Kalckar seems to fit this image as well. Science Berg, Paul. “A Stanford Professor’s Career in Biochemistry, was one of life’s rich rewards; good music, good friends, a Science Politics, and the Biotechnology Industry.” Berkeley: Regional Oral History Office, Bancroft Library, University of cornucopia of human experiences were equally valuable. California, 2000. Available from http://www.calisphere.universityofcalifornia.edu. An oral BIBLIOGRAPHY history conducted in 1997 by Sally Smith Hughes. Boos, Winfried. “Farewell to a Dionysian.” Biological Chemistry WORKS BY KALCKAR Hoppe-Seyler 372, no. 11 (1991): 8–9. Original in German; “Phosphorylation in Kidney Tissue.” Enzymologia 2 (1937): translation kindly provided by Dr. Boos. 47–53. Garfield, Eugene. “Highly Cited Articles: 35. Biochemistry “The Nature of Phosphoric Esters Formed in Kidney Extracts.” Papers Published in the 1940s.” Current Contents 8 (1977): Biochemical Journal 33, no. 5 (1939): 631–641. 5–11. Reprinted in: Essays of an Information Scientist, Vol. 3. “The Nature of Energetic Coupling in Biological Syntheses.” Philadelphia, PA: ISI Press. Available from Chemical Reviews 28 (1941): 71–178. http://www.garfield.library.upenn.edu/essays.html. With Sidney P. Colowick and Carl F. Cori. “Glucose Kamp, A. F., J. W. M. La Rivière, and W. Verhoeven. “Kluyver Phosphorylation and Oxidation in Cell-Free Tissue Extracts.” as Professor: Chronicles of the Laboratory.” In their Albert Journal of Biological Chemistry 137 (1941): 343–356. Jan Kluyver: His Life and Work, pp. 14–48. New York: Interscience Publishers, 1959. “The Enzymatic Action of Myokinase.” Journal of Biological Chemistry 143 (1942): 299–300. Kennedy, Eugene P. “Herman Moritz Kalckar, March 26, 1908–May 17, 1991.” Biographical Memoirs, National “The Role of Myokinase in Transphosphorylations: II. The Academy of Sciences (U.S.) 69 (1996): 148–165. Enzymatic Action of Myokinase on Adenine .” Journal of Biological Chemistry 148 (1943): 127–137. Lipmann, F. “Metabolic Generation and Utilization of Phosphate Bond Energy.” Advances in Enzymology 1 (1941): With Manya Shafran. “Differential Spectrophotometry of Purine 99–162. Compounds by Means of Specific Enzymes: I. Determination of Hydroxypurine Compounds.” Journal of Biological Park, Buhm Soon. “Historian’s Perspective: NIH Offered Haven Chemistry 167 (1947a): 429–443. from Antinepotism Rules.” Newsletter of the NIH Alumni Association 14, no. 2 (2002): 18–19. Available from With Alice Neuman Bessmann. “Differential Spectrophotometry http://www.fnih.org/nihaa/NIHAA-Update.pdf. of Purine Compounds by Means of Specific Enzymes: II. Racker, Efraim. Mechanisms in Bioenergetics. New York: Determination of Adenine Compounds.” Journal of Academic Press, 1965. Biological Chemistry 167 (1947b): 445–459. Wellcome Trust. Fourth Report, Covering the Period 1960–1962. With Manya Shafran. “Differential Spectrophotometry of Purine London: Author, 1963. Compounds by Means of Specific Enzymes: III. Studies of the Enzymes of Purine Metabolism.” Journal of Biological Chemistry 167 (1947c): 461–475. Rivers Singleton Jr. With Elizabeth P. Anderson and Kurt J. Isselbacher. “Galactosemia, a Congenital Defect in a Nucleotide Transferase.” Biochimica et Biophysica Acta 20 (1956a): 262–268. KAMMERER, PAUL (b. , -Hun- With Elizabeth P. Anderson, Kurt J. Isselbacher, and Kiyoshi Kurahashit. “Congenital Galactosemia, a Single Enzymatic gary, 17 August 1880; d. Puchberg am Schneeberg, Lower Block in Galactose Metabolism.” Science 123 (1956b): Austria, 23 September 1926), zoology, heredity, evolution. 635–636. The main goal of Kammerer’s research was to demon- “An International Milk Teeth Radiation Census.” Nature 182 strate the modifying power of the environment and (1958): 283–284. the heritability of acquired characteristics, using the

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