Kammerer, Paul

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(b. Vienna, Austria-Hungary, 17 August 1880; d. Puchberg am Schneeberg, Lower Austria, 23 September 1926),

zoology, heredity, evolution.

The main goal of Kammerer’s research was to demonstrate the modifying power of the environment and the heritability of acquired characteristics, using the experimental methodology of twentieth-century biology. Conceptually, he aimed to reconcile this Lamarckian mode of heredity with the new science of genetics and an older interpretation of Darwinism that posited the evolutionary significance of acquired characteristics. Along with many other opponents of neo-Darwinism, Kammerer argued that Darwin had intended natural selection only as a means of eliminating unfavorable characteristics. New and favorable ones had to be acquired in response to environmental changes and to changed habits. Individual improvement and inheritance of acquired skills and behaviors, in addition to physical traits, were also the key to human evolution and cultural progress, according to Kammerer.

Kammerer was a prolific writer and popularizer. His books, articles, and lectures reached beyond Vienna to a broad public, as did his ideas about how to apply his experimental findings to human cultural evolution. Among professional scientists, his work was also well known, and it stimulated heated discussion about the nature of heredity and the causes of variation. But Kammerer’s arguments for the inheritance of acquired characteristics were not widely accepted by scientists at the time, and they are not considered valid today. Moreover, he is suspected of tampering with, or perhaps even fabricating, his experimental evidence.

Family Background and Education . Paul Kammerer grew up in the secure, prosperous, and creative world of upper-middle-class Vienna in the last decades of the Habsburg Empire. His father Karl was a manufacturer whose family came from Siebenbürgen, a German-speaking enclave in what later became Romania. His mother Sophie was from a converted Jewish family on at least one side. His parents had both been married before, and Paul had four much-older half siblings.

Like many others of his generation and social class, Kammerer was intensely devoted to the arts, and even though he chose a scientific career, he always valued creativity and thought of himself as an artist among scientists. Kammerer studied zoology at the Vienna University and music theory at the conservatory, beginning in 1899. He wrote his dissertation on the evolutionary relationships between two species of salamander, under the direction of Berthold Hatschek, a protØgØ of Ernst Haeckel, who taught a comparative approach to morphology and evolution. Kammerer got his introduction to experimental methodology outside the university, under the tutelage of zoologist Hans Przibram, another student of Hatschek’s who was rebelling against the comparative approach.

Kammerer taught biology courses at a variety of places, from the university, to the Volkshochschule (a system of adult-education centers), and a high school for girls. He also traveled to give talks throughout the Austrian Empire and Germany, often to chapters of the Monist League, which was associated with Ernst Haeckel. Kammerer shared the Monists’ goals of popularizing science and especially scientific approaches to social and political problems.

Kammerer was said to be handsome, charismatic, and vain, and stories abound about his attractiveness to the Viennese ladies and his requited and unrequited loves. Kammerer was married in 1906 to Felicitas Maria Theodora (“Dora”) von Wiedersperg, the daughter of Bohemian aristocrats, through whom Kammerer gained entry into Viennese high society. Kammerer left her to marry the painter Anna Walt around 1912 or 1913, but returned to Dora after a short, stormy marriage and a suicide attempt. He and Dora had a daughter named Lacerta, after the genus of mural lizards.

Kammerer also had some success in the arts. He published a book of lieder and had them performed in Vienna. He made the acquaintance of many of the eminent artistic figures of turn-of-the-century Vienna, such as Gustav Mahler, whom he idolized, and Mahler’s wife Alma, whom he courted after the composer’s death and to whom he dedicated a small book on the inheritance of musical talent.

Environmental Effects and Their Inheritance . In 1902, before finishing his dissertation, Kammerer went to work for Przibram at the newly founded Biologische Versuchsanstalt (Institute for Experimental Biology) in Vienna. This laboratory, also known as the Vivarium, after the zoological exhibition hall that previously occupied the same premises, was owned by Przibram and devoted to the new experimentalism, which Przibram wanted to apply to the widest possible range of organisms and scientific problems. As Przibram’s assistant, Kammerer helped acquire the animal stocks, design and build aquaria and terraria, and develop methods of rearing the animals and controlling their environments. He also began doing experiments of his own and added an experimental portion to his dissertation project.

After defending his dissertation in 1904, Kammerer remained at the Vivarium to pursue a multifaceted career, combining experimental work on heredity, evolution, development, regeneration, and symbiosis with forays into evolutionary theory, and sidelines as a composer, teacher, and popular science writer and lecturer.

The themes of his later work were already discernable in Kammerer’s dissertation on the evolutionary relationship between the spotted salamander, Salamandra maculosa and its alpine cousin S. atra. By manipulating their environments in the laboratory, Kammerer induced individuals of each species to take on characteristics of the other: for example, instead of the usual pattern of producing a large brood of eggs that hatched into tadpoles, S. maculosa was induced to give birth ovoviviparously on land to fewer, but fully metamorphosed young. S. atra was induced to make the opposite shift. In a follow-up study, Kammerer bred these salamanders and found that their offspring retained the modified reproductive strategies, even when reared in their original environments. That work earned him the Sömmering Prize from the Senckenberg Society of Naturalists in Frankfurt in 1909 as well as his habilitation, or qualification to teach courses at the university at the rank of privatdozent (lecturer), in 1911.

Kammerer made further demonstrations of the modifying power of the environment, using many of the other species and environments available at the Vivarium. For example, he produced color-variants in the mural lizard Lacerta by varying temperature, and he used an artificial lighting regimen to induce the blind cave salamander Proteus to develop large, functional eyes. More controversial were the experiments in which he also bred his modified individuals and reported inheritance of the new characteristics. In addition to his dissertation project, these breeding experiments included several on the effects of background colors on the patterns of spots on S. maculosa; several on the life cycle and breeding behavior of the midwife toad Alytes obstetricans; the effects of amputation on siphon length in the sea squirt or tunicate Ciona intestinalis; and more besides.

Kammerer argued that in such cases the environmentally induced modification had been communicated to the chromosomes, where incipient genes were forming that would perpetuate the modification, at least partially. Crosses between modified and unmodified stocks sometimes yielded hybrid progeny in Mendelian ratios, apparently supporting the idea that new genes were at least partially formed. Kammerer speculated that hormones might be the medium of communication that allowed the chromosomes to express themselves in the body and carried information about the body back to the chromosomes.

In any case, he argued that his experiments had vindicated the older interpretation of Darwinism that had allowed for inheritance of acquired characteristics, and undercut the neo-Darwinism of August Weismann that ruled it out. In particular, they had falsified Weismann’s conception of an isolated germ plasm or hereditary material that could receive no communication back from the body and was mostly shielded from environmental influences as well. Kammerer felt he had demonstrated that there was no Weismannian barrier to communication with the hereditary material.

Most geneticists, neo-Darwinian evolutionists, and even some proponents of the inheritance of acquired characteristics

Paul Kammerer. © BETTMANN/CORBIS .

received these results with skepticism, if not hostility. They tried to explain away Kammerer’s results as effects of selection in highly variable organisms, atavisms, lucky mutations, poor environmental controls, and other errors. In retrospect, inadvertent selection probably accounts for most of the results. Recording errors and poor environmental controls are also likely.

Opponents also complained that Kammerer’s published documentation was not up to professional standards. His verbal descriptions of modified animals were vague, his drawings sketchy, his photographs fuzzy or retouched, and he had to defend himself repeatedly against accusations of unreliability and insinuations of fraud. Przibram and others who saw Kammerer’s specimens vouched for their authenticity, however.

Human Cultural Evolution . Kammerer’s public lectures and popular writings applied his evolutionary ideas and experimental results to human affairs. Kammerer believed that every individual had the potential to be improved and to contribute to biological or cultural progress. Inherited musical talent, for example, could be improved by practice and passed on in slightly enhanced form to the next generation. Such effects on posterity were what made the individual and his or her upbringing and education most valuable and effective. Education, training, and practice were not Sisyphean chores that had to be repeated entirely from scratch in every generation, as the Weismannian alternative implied.

During World War I Kammerer was called away from the Vivarium to work for the military censor. His experimental work came to a halt and his stocks of modified laboratory animals died out. In his spare time Kammerer wrote antiwar essays, mostly using evolutionary arguments about the benefits of cooperation and the purely eliminative effects of struggle and selection. He also published a textbook of general biology (1915) and a theoretical analysis of coincidences titled Das Gesetz der Serie (The law of series, 1919). The latter book was rather speculative, if not pseudoscientific, and it cost him a promotion at the university to ausserordentlicher Professor (extraordinary professor—just a rank, not a position on the payroll). Kammerer’s rejection for this promotion has also been blamed on anti-Semitism, Kammerer’s socialism and wartime pacifism, and rumors about his unreliability as a scientific observer, but Przibram, who was on the committee, said they were ready to overlook everything else before the book appeared.

After the war, Kammerer did little further experimental work. His writings and lectures shifted their focus to the prospects for getting humanity back on track towards evolutionary progress through the controlled acquisition of desirable characteristics. He now drew on the work of his Vivarium colleague, the endocrinologist Eugen Steinach, on the power of glandular secretions, especially the testicular secretion, to modify both body and behavior. In his lectures and popular writings he touted the “Steinach operation,” a form of vasectomy, as a means of rejuvenating aging men by stimulating testicular secretion. He also favored other kinds of interventions, such as injections with glandular extracts, radiation treatments, and testicle transplants, as means of improving unfit individuals instead of selecting against them. He argued, based on his prewar experimental results, that the hormonal effects would become hereditary and offer a much more effective and humane alternative to negative eugenic measures, such as sterilization or restrictions on marriage.

Scandal and Suicide . Kammerer resigned from the Vivarium in 1921 and lived from lecturing and writing. A grand tour took him to Britain and the United States in 1923 and 1924, where he promoted the Steinach operation and his proposals for an alternative eugenics, and where he also exhibited his one remaining specimen of a midwife toad that had inherited an acquired characteristic: a nuptial pad. Normally absent in midwife toads, which mate on land, these dark and rough patches of skin on the front legs of the male are found in water-breeding frogs as adaptations for clasping the female. Kammerer’s experimental midwife toads had acquired the habit of mating in the water and subsequently developed the pads.

Questions had been raised before about whether the dark patches were nuptial pads, and his tour revived them. The geneticist William Bateson was most aggressive in his questioning, and a testy exchange of letters to Nature ensued after Kammerer’s visit to Cambridge. The British embryologist Ernest William MacBride also joined in on Kammerer’s side.

Kammerer got a much warmer reception in the Soviet Union in May and June 1926. There, Marxist intellectuals at the Communist Academy in Moscow were suspicious of Western genetics and neo-Darwinism, and sympathetic to Kammerer’s alternative. They also regarded him as a fellow socialist, and offered him a job as director of a small institute to be built in Moscow under the academy’s auspices. Kammerer accepted the offer, but never returned to Moscow to take up the position.

Earlier in 1926, the American herpetologist G. Kingsley Noble had come to the Vivarium to examine the midwife toad and found that the dark color of the purported nuptial pad came from an injection of India ink. Separate accounts and interpretations of the finding, by Noble and Przibram, appeared in Nature on 7 August. Kammerer asserted his innocence but committed suicide six weeks later, leaving himself with a posthumous reputation as an ideologically motivated opponent of modern Darwinism and a fraud.

Kammerer’s defenders, among them the novelist Arthur Koestler, claim that he was either framed by an unscrupulous Darwinian, eager to discredit unwelcome evidence for Lamarckism, or that a well-meaning assistant had merely touched up a fading specimen that really did once have a distinct nuptial pad. It is also possible that Kammerer inked it himself in order to reduce the glare when taking a photograph of the wet specimen.



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“Eyes: Newt, Rat, Human.” Time, 18 June 1923, p. 19.

“Famous European Biologist Visits Us: Dr. Paul Kammerer, Associate of Steinach, Driven from Vienna by Poverty.” New York Times, 25 November 1923.

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———.“The Case of Paul Kammerer: Evolution and Experimentation in the Early Twentieth Century.” Journal of the History of Biology 39, no. 3 (2006): 525–563.

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Sander Gliboff