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The Case of Paul Kammerer: Evolution and Experimentation in the Early 20th Century
Author(s): Sander Gliboff
Source: Journal of the History of Biology, Vol. 39, No. 3 (Autumn, 2006), pp. 525-563
Published by: Springer
Stable URL: http://www.jstor.org/stable/4332032
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Journal of the History of Biology (2006) 39:525-563
DOI 10.1007/s 10739-005-3051-5
? Springer 2006
The Case of Paul Kammerer:Evolution and Experimentation
in the Early 20th Century
SANDER GLIBOFF
Department of History and Philosophy of Science
Intliana University
130 Goodhody'Hall
Bloomington, IN 47405
USA
E-mail: sglihoffta'indiana.edu
Abstract. To some, a misguided Lamarckian and a fraud, to others a martyr in the
fight against Darwinism, the Viennese zoologist Paul Kammerer (1880-1926) remains
one of the most controversial scientists of the early 20th century. Here his work is
reconsidered in light of turn-of-the-century problems in evolutionary theory and
experimental methodology, as seen from Kammerer's perspective in Vienna. Kammerer emerges not as an opponent of Darwinism, but as one would-be modernizer of
the 19th-century theory, which had included a role for the inheritance of acquired
characteristics. Kammerer attempted a synthesis of Darwinism with genetics and the
chromosome theory, while retaining the modifying effects of the environment as the
main source of favorable variation, and he developed his program of experimentation
to support it. Kammerer never had a regular university position, but worked at a
private experimental laboratory, with sidelines as a teacher and a popular writer and
lecturer. On the lecture circuit he held forth on the significance of his science for
understanding and furthering cultural evolution and he satisfied his passion for the arts
and performance. In his dual career as researcher and popularizer, he did not always
follow academic convention. In the contentious and rapidly changing fields of heredity
and evolution, some of his stances and practices, as well as his outsider status and partJewish background, aroused suspicion and set the stage for the scandal that ended his
career and prompted his suicide.
Keywords: Darwinism, evolution, experiment, Hans Przibram, Hugo Iltis, laboratory,
Lamarckism, Mendelism, Paul Kammerer, Vienna, Vivarium, William Bateson
Introduction
From 1904 until the First World War, the Viennese zoologist Paul
Kammerer flooded the technical and popular presses with accounts of
dramatic modifications of animal forms and behaviors, all induced by
environmental influences, and most, apparently, heritable. Spotted
salamanders became striped and striped ones became spotted, land
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526
SANDER GLIBOFF
salamanders produced aquatic larvae and water-breeders skipped their
larval stages, blind cave olms developed functional eyes, and lizards, sea
squirts, frogs, and midwife toads changed their colors, shapes, or lifecycles. The best-known change was in midwife toads, which normally
mate on land and brood their eggs attached to the hind legs of the male.
Kammerer coaxed them to mate in the water instead, where the eggs
floated away. From the few surviving eggs he raised a line of habitual
water-breeders, in which "nuptial pads" even appeared, as known from
frogs: dark, rough patches on the male forelimbs with which to clasp the
slippery female.
Kammerer argued that his experimental environments caused the
animals to acquire the new characteristics and that such effects, along
with the effects of use and disuse of organs, were the principal cause of
organic change, adaptation, and progress. They constituted the creative
side of evolution, in contrast to the eliminative work of natural selection. Further, as Kammerer stressed in popular writings and lectures,
the same creative processes were at work in human affairs, where they
ensured progress in culture and the arts. Education, practice, and
artistic achievement resulted in cumulative hereditary improvements in
human creativity and expression.
Kammerer's popularizing efforts let him combine his love of animals
with his passion for the arts and straddle the worlds of experimental
science and Viennese culture. A trained musician and a charismatic
performer on the lecture circuit, he approached his science with an
artist's flair, shaping animals in the laboratory, showing off his creations
and interpreting them for the public. He also delivered a political
message about the value of the individual for the progress of the species
or nation. Environment and education shaped people, societies, and
culture. After the war, these themes dominated Kammerer's lectures,
which presented a vision for healing and renewed progress by means of
a new eugenics that would improve the unfit rather than select the fit.
Kammerer's experiments were a signature achievement of the
Biologische Versuchsanstalt[Institute for Experimental Biology], where
he did them all. Known informally as the "Vivarium," it was founded
and directed by Hans Przibram. It supported the experimental trend in
organismal biology by giving it a home outside the university and
providing research and employment opportunities for experimentalists.
Przibram shared many of Kammerer's interests in the arts, was active in
adult education, and he encouraged Kammerer's popularizing work.
Kammerer's high public profile helped to advertise the Vivarium's
technical capabilities, its accomplishments in heredity and evolution,
and the power of its experimental methodology.
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THE CASE OF PAUL KAMMERER: EVOLUTION AND EXPERIMENTATION
527
However, skepticism about Kammerer's work and credibility was
strong, for various reasons. Some were theoretical. August Weismann
had been campaigning against the inheritance of acquired characteristics since the 1880s, with his model of a hereditary germplasm, largely
isolated from environmental influences. After 1900 Mendelian genetics
left even less room for "Lamarckian" heredity. Other problems were
personal. Kammerer's popularizing, his showy lectures, and public
overstatement of his experimental accomplishments, all detracted from
his reputation among academic scientists. His technical publications
failed to help matters, because the descriptions of modified organisms
were often vague, and the photographs of poor quality. In some cases,
anti-Semitism came into play, for Kammerer was at least partly of
Jewish descent, as were most of his Vivarium colleagues. Rumors circulating about Kammerer's "unreliability" as a scientist found ready
believers, even before Kammerer became embroiled in the scandal for
which he is best remembered.
Kammerer never preserved his modified animals systematically, and
his live stocks died off during the war. Thereafter Kammerer was
reduced to displaying just a few preserved specimens, among them one
midwife toad with one intact nuptial pad, on lecture tours and at the
Vivarium. The scandal broke in 1926, when that last toad was shown to
have been injected with India ink where the nuptial pad was supposed to
be. Kammerer denied having done it, but within weeks he committed
suicide. The act was widely interpreted as an admission of guilt, not only
of tampering with the specimen, but of fabricating all of his results, and
it cast doubt on all claims about Lamarckian heredity.'
The charges are serious, and Kammerer's guilt continues to be
asserted,2 but the recklessness of the fraud - if fraud it was - makes it
difficult to comprehend. Kammerer's friend Hugo Iltis, the plant
geneticist, Mendel biographer, and early promoter of Mendelism, could
not imagine him being so foolish. Iltis recalls discussing the impending
scandal with him, and says Kammerer convinced him of his innocence
with a rhetorical question: "'Do you think I'm a Dummkopf or an idiot?
Because that's what I would have to be if I left a forgery with ink
standing around openly in the laboratory, where so many of my enemies
have entry, or if I even sent it to a scientific congress.' I could not escape
the logic of this argument."3
1
E.g., Jennings, 1930, p. 345.
E.g., Whittaker, 1985; Aronson, 1975.
3 Iltis, 1951, "Paul Kammerer," Paul Kammerer Papers, American Philosophical
Society Library, Philadelphia, B/K128 [hereinafter, "Kammerer Papers"].
2
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528
SANDER GLIBOFF
I agree with Iltis that the pad was not fabricated. Kammerer worked
with highly plastic organisms that probably did exhibit the traits he
reported, and there was more evidence for the existence of the nuptial
pad than just the doctored specimen. However, I suspect Kammerer of
injecting the specimen anyway, only to enhance, not create, its
appearance. But why would Kammerer have done even that? And why,
to paraphrase his question to Iltis, did he show that specimen so willingly, for so many years, to increasingly skeptical audiences?
One of my purposes is to answer these questions, but there are also
larger historical issues to address, which have been obscured by disproportionate attention to the charge of fraud. What was Kammerer's
role in the development of evolutionary theory and experimentalism
before the scandal, especially before 1914? What exact mechanism of
variation, heredity and evolution was he defending, what made it
plausible, and what alternatives were available in the early days of
genetics? Who were his allies, who his opponents, and what did they see
in Kammerer's work? Further: what is the significance of the local
context? How did Viennese culture and the arts, the lecture circuit, and
the internal culture of the Vivarium shape Kammerer's approach?
There have been several attempts to explain the scandal and put
Kammerer's work into perspective. In 1926, Przibram vouched for
Kammerer's results generally, but not for the inked specimen, and not
exactly for his conclusions, either. It was a limited effort at rehabilitating Kammerer, for Przibram feared that a more vigorous approach
would be too provocative, given the hostile atmosphere. As late as 1930
he advised Iltis to hold back, too: "As long as no clarification of the
blackening of the preparations is available, I consider a Kammerer
biography or any other steps toward honoring him to be out of the
question. His friends in England have the same impression."4
Kammerer did not find a bold champion until 1971, when Arthur
Koestler published The Case of the Midwife Toad,s a pro-Lamarckian
counterattack. Koestler accuses "orthodox Darwinians," led by the
British Mendelian William Bateson, of trying to suppress Kammerer's
results, undermine his credibility, and prevent his advancement at the
University. He suggests that a Darwinian enemy inked the toad and
framed him, or else that a well-meaning laboratory worker touched up
the deteriorating specimen.
Several authors have criticized Koestler for grinding his own antiDarwinian ax, not Kammerer's, as well as for oversimplifying the
4 Przibram to Iltis, 9 February 1930, Kammerer Papers.
S
Koestler, 1971.
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THE CASE OF PAUL KAMMERER: EVOLUTION AND EXPERIMENTATION
529
scientific issues, taking them out of context, and making Bateson the
orthodox Darwinian. Bateson's saltational view of evolution and
rejection of the chromosome theory would seem to disqualify him for
the role. Koestler also exaggerated the threat that Kammerer's results
posed to the theory of natural selection, for most of them can be
explained in selectionist terms.6
There are also serious omissions in Koestler's story, such as Kammerer's Jewish background and the possible role of anti-Semitism in the
story. Also missing are two disputes with scientists over purported
inaccuracies in his articles and retouching in his photos, which cast
suspicion on Kammerer and foreshadowed the midwife-toad scandal.
Kammerer's penchant for hyperbole and self-promotion is downplayed
by Koestler, and his campaign for an alternative eugenics is not mentioned.
Albrecht Hirschmuillerfills out the story with archival research and
dispels much of Koestler's mythology. He shows, for example, that
university committees considering Kammerer for promotions were not
as ill-disposed towards him or his theories as Koestler assumes. They
included Przibram and others sympathetic to his evolutionary views and
willing to look past his controversial public persona.7 Still,
Hirschmuiller,like Koestler and Koestler's critics, labels Kammerer a
"Lamarckian" and an anti-Darwinian without capturing the nuances of
his position.
As I shall argue, Kammerer did not oppose Darwinism, but sought
to retain the inheritance of acquired characteristics as a mechanism of
change within a Darwinian framework. Darwin himself had accepted
environmental effects and use-inheritance as causes of variation, and
Ernst Haeckel had made them central in his influential interpretation of
1866.8 Kammerer was trying to update the case for them, by taking
Mendelism, genes, and chromosomes into account, providing experimental demonstrations, and developing a Darwinian world-view as
grand as Haeckel's.
A Biologist in Vienna
Paul Kammerer was born on 17 August 1880, in Vienna. His mother
was from a converted Jewish family on at least one side, a point that is
'
7
8
[Waddington], 1971; Gould, 1972; Bowler, 1983.
Hirschmiiller, 1991.
Haeckel, 1866.
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530
SANDER GLIBOFF
important because his difficulties at the University have been blamed on
anti-Semitism or a special bias against Jewish socialists.9 These claims
are not well supported by Przibram's recollections or the minutes of
promotion committees, as cited by Hirschmiiller. However, a better case
can be made for anti-Semitism having colored perceptions, by some
German biologists, of Kammerer's results.
The son of a manufacturer, young Kammerer cultivated an intense
interest in the arts, like so many members of his generation and class in
Vienna.'0 When he entered the University in 1899, Kammerer studied
Music in addition to Zoology." He composed Lieder and tried to be
one of the Viennese cultural elite, mingling with the Mahlers and other
artistic figures, and joining the eminent crowd courting the widowed
Alma after Gustav Mahler's death in 1911. Throughout his career, he
tried to bring his science to the art world, through his popular writings,
teaching at various levels (high school, university, and adult education),
and especially his "big show-lectures" [gro,ie Schauvortrdge].12
Kammerer began his dissertation research under Berthold Hatschek,
a comparative morphologist and a protege of Haeckel's, who set him to
work on phylogenetic questions. Kammerer found it boring and
unsatisfying: it "did not fully suit me in my individuality," he said.'3 He
complained about academic convention and the emphasis on dead
forms from the evolutionary past,'4 and wished for a chance to do
experimental work on the causes of form, using live animals. Kammerer
ended up riding the wave of experimentalism that Garland Allen has
characterized as a "revolt from morphology." Allen's conception has
been challenged on various grounds, but mainly because of difficulty in
drawing the line between experimentalists and traditional morphologists, or between turn-of-the-century experimentalism and older experimental approaches.'5 However, the revolt metaphor is apt for
Kammerer, and for Przibram as well. A spirit of rebelliousness pervades
their writings, and Przibram explicitly states his wish to depart from
9 Iltis, 1951, "Paul Kammerer," Kammerer Papers; Hirschmuller, 1991, p. 31. Also
"Dr. Kammerer und die Wiener Universitat," Neue Freie Presse, 1924; Bernfeld, Anne,
"Ein Wiener Gelehrter," 1926; both newspaper clippings in the Kammerer Papers.
10
Zweig, 1944, pp. 16-86; Schorske, 1981.
1 Kammerer, [ca. 1910], "Curriculum Vitae," Kammerer Papers.
12 Kammerer to Iltis, 6 July 1910, Kammerer Papers.
3 Kammerer, [ca. 1910], "Curriculum Vitae," Kammerer Papers.
14 Kammerer, 1914b.
15 Allen, 1978. For the critique and further discussion: Maienschein, 1981; Maienschein, Rainger and Benson, 1981; Allen, 1981. For an analysis in terms of gradual generational, institutional, and disciplinary change: Nyhart, 1995, Chs. 9 & 10.
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THE CASE OF PAUL KAMMERER: EVOLUTION AND EXPERIMENTATION
531
academic morphology and promote experimental approaches and
institutions.
The Vivarium Laboratory
Experimentalism was already making strides in other countries, where
calls could be heard for new laboratories devoted to experimentation in
whole-organism research.'6 In Germany, Wilhelm Roux, a student of
Haeckel's, was one of the leading promoters of experimentalism, and
Przibram enlisted his moral and practical support for a laboratory in
Vienna. Przibram's prospects for a university career were bleak, in part
because he was Jewish, but also because of a tight academic job market,
and in 1902, on Hatschek's suggestion, he used some of his family
fortune to buy the "Vivarium," a defunct exotic-animal exhibit in the
Prater amusement park. He enlisted two other underemployed Jewish
experimentalists as co-directors, hired Kammerer as an assistant, and
converted the building into a laboratory. Kammerer then moved his
dissertation work to the Vivarium and added an experimental component to his project.'7
Roux arranged to publish all the Vivarium's research reports in his
journal, the Archiv fur Entwicklungsmechanik [Archive for developmental mechanics, hereinafter, "Roux's Archiv"], which sometimes
devoted entire numbers just to Vivarium papers. The best-known
Vivarium authors during Kammerer's time there were Przibram, whose
wide-ranging work covered regeneration, the effects of environmental
changes on development and heredity, and crystal formation as an
analog of organic growth; and Eugen Steinach, who worked on glandular secretions, sex determination and aging, and who gained notoriety
for his "rejuvenation" surgery. Wolfgang Pauli, father of the famous
physicist, ran a small physico-chemical department and worked on
colloid chemistry; and Przibram's co-founders, Wilhelm Figdor and
Leopold von Portheim headed two botanical departments. Other staff
members and visitors worked on regeneration, artificial
parthenogenesis, symbiosis, the role of gravity growth and development,
1999.
Przibram, 1926b, 1906, 1959; Hofer, 2000. It was also the subject of a workshop
titled "The Viennese Roots of Theoretical Biology: The Vivarium Centenary," organized by Manfred Laubichler, Gerd B. Muller, and Werner Callebaut at the Konrad
Lorenz Institute, Vienna and Altenberg, 2002, abstracts available from http://
www.kli.ac.at/workshops-a.html.
16Cook,
1'
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532
SANDER GLIBOFF
organ transplantation, and testing the advantages of purported adaptations. '8
Przibram spared no expense on equipment to provide the bestcontrolled conditions achievable for testing environmental effects on the
growth and development of greenhouse plants and terrestrial, freshwater, and marine organisms.'9 A deep cellar provided an environment
for cave animals and space for storing rainwater and the seawater that
was brought by train from Trieste. An electric pump circulated the
seawater through aquaria upstairs. Outdoors were experimental ponds
and open-air terraria. Indoors were more terraria and insectaria. For
the botanists there were extensive gardens and two greenhouses with
electric lighting. There were rooms with remotely controllable light
sources and heating systems, apparatus for manipulating air pressure,
and electric centrifuges and "clinostats" for simulating gravitational
forces. Different temperatures were maintained in different parts of the
building, using central steam heat, supplemented with stoves in the
".warmrooms." For especially cold temperatures there was the cellar, a
supply of ice, and rented cold rooms at a nearby warehouse.
The main hall of the Vivarium was lit by tall windows and was kept
as free as possible from dust and toxic chemicals. It had taps for fresh
water and seawater, aeration outlets from a central air-pumping system,
and it housed 15 work tables with electric sockets and lights and Bunsen
burners. Oddly, there was no mention of photographic equipment.
Przibram, a talented artist,20 preferred to do scientific illustrations by
hand, despite his evident fascination with modern technology.
Work was organized and funded on the table-system, modeled after
the Naples Zoological Station,2' and institutional sponsors or paying
researchers were sought to fund and occupy each table. Any experimental project was to be welcomed. There was to be no other laboratory
doctrine than experimentalism, and Przibram directed his department
with a light hand, encouraging diverse ideas to be tested on many different organisms.
Kammerer was free to explore the idea of "somatic induction,' which
had environmental effects being transmitted from the modified body
parts to the hereditary material or germplasm. At the same time, Przibram's second assistant, Franz Megusar was out to prove that variation
was generated internally in the germplasm, while Przibram inclined
towards "parallel induction," whereby the environment modified soma
18
19
20
21
Przibram, 1908-1909, 1959. On Steinach: Sengoopta, 1998.
The following description is based on Przibram, 1908-1909.
Coen, in prep.
Groeben, 1985.
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THE CASE OF PAUL KAMMERER: EVOLUTION AND EXPERIMENTATION
533
and germ simultaneously, causing the former to acquire characteristics
and the latter the means to perpetuate them, but with no communication
between the two. Przibram let all three views flourish while he tested
whether heat, light and other environmental factors could penetrate to
the reproductive organs as required for parallel induction.22 Results with
salamanders kept open the possibility, while the same experiments on
different species sometimes seemed to rule it out.23
Przibram wrote that the laboratory's "main purpose is not the
investigation of a narrow group of organisms.... Its activities should not
be limited to specific problems, instead it should draw all the big
questions of biology into its purview. Not specialization, but generalization from the experiences we gain is our goal: the favoring of a
certain methodology and technology alone sets our Institute apart from
earlier biological research facilities."24 To Przibram, "experimental
biology" meant research on whole organisms, kept alive under experimental conditions for an entire life-cycle, or at least for an extended
period. He felt that short-term, reductionistic experimentation was
already well established at physiological and medical laboratories and
need not be included in the Vivarium program. Similarly, research in
genetics was excluded, even though it might have fit Przibram's definition of experimental biology. He said genetics already had a foothold in
Vienna, thanks to Erich von Tschermak at the Agricultural College, and
so it was not necessary for the Vivarium to duplicate the effort.
Przibram's executive decisions and the Vivarium's methodological
mission and lavish facilities had formative influences on Kammerer's
work. He had to develop methods of rearing diverse animals and controlling their environments. His publications served the interests of the
laboratory by demonstrating and advertising the power of the approach
and the technology. In successive articles in Roux's Archiv one can see
him trying out the gamut of artificial environments and terrestrial,
aquatic, and marine organisms. Kammerer's high profile in the press
and on the lecture circuit was also beneficial, for the sake of attracting
researchers and sponsors to the tables.
Kammerer'sExperimental Pipeline
Kammerer's scientific career began with his 1904 dissertation on the
anatomy and reproductive adaptations of two salamanders, the lowland
22
23
24
Przibram, 1912.
Secerov, 1912a, b.
Przibram, 1908-1909, p. 234.
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SANDER GLIBOFF
Salamandra maculosa and the alpine S. atra. The former produces 40
eggs or more and releases them into the water when they are ready to
hatch into tadpoles. The latter carries its eggs longer, and they hatch
already metamorphosed into the terrestrial form, larger than S.
maculosa tadpoles, and usually only two in number. Kammerer found
that increasingly dry conditions induced S. maculosa to delay releasing
its eggs until it produced large, metamorphosed young like S. atra.
Simultaneously, the typical brood size decreased to two. A reciprocal
experiment transformed S. atra's life history into S. maculosa's. Both
changes were effected within a single generation. The same individuals
produced increasingly land- or water-adapted offspring in successive
breeding periods.
Kammerer concluded, based on these results and comparative
anatomy, that the two species were closely related, despite their contrasting life histories.25 Kammerer did not discuss mechanisms of evolution or plans to follow up the project with breeding experiments.
Indeed Kammerer's emphasis on the close relation between the two
species runs counter to his later claim to have induced a significant
evolutionary change in the laboratory. His professors, Hatschek and
Carl Grobben passed the dissertation with praise for the experimental
skills and the interesting results, but with doubts about Kammerer's
depth in theoretical matters and the relevance of the experiment to the
phylogenetic questions with which he had begun in Hatschek's laboratory.26
After receiving his doctorate, Kammerer continued to work at the
Vivarium as Przibram's assistant. He dropped the phylogenetic questions in favor of testing environmental effects and their heritability.
However, in contrast to the pattern being established elsewhere in the
study of heredity, Kammerer used long-lived organisms, with generation times of up to three and a half years in the case of the salamanders.
To deal with the long experimental durations, Kammerer got a pipeline
going, containing a staggered series of experiments, using different
species and laboratory environments each time, and aiming for evermore-striking transformations. Positive results began to emerge in 1908:
the salamanders with altered life histories had offspring with the same
traits, the water-breeding midwife toads not only bred true, but
acquired nuptial pads, spotted salamanders changed color or became
striped when raised on differently colored substrates, blind cave olms
regained their sight when raised under special lighting, sea squirts
25
26
Kammerer, 1904.
Hirschmiiller, 1991, p. 35.
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THE CASE OF PAUL KAMMERER: EVOLUTION AND EXPERIMENTATION
535
acquired long, droopy siphons following amputations.27 In the cases
where Kammerer bred the modified animals, the acquired characteristics reappeared in the next generation, even in the absence of the
environmental treatments.
These experiments do not exhaust Kammerer's program. He also
worked on regeneration and symbiosis, which, like the acquisition of
new characteristics, represented creative and constructive responses to
external forces. They bolstered Kammerer's case against variation being
random, internally generated, or predetermined in an isolated germplasm. There was also a sideline on the mechanism of heredity intended
to build bridges to Mendelism and the chromosome theory. Kammerer
back-crossed his midwife toads and salamanders to unmodified individuals and produced hybrids more or less in Mendelian ratios. By
Kammerer's reasoning, this indicated that the acquisition of characteristics caused Mendelian genes to form on the chromosomes. Where
the expected ratios did not appear, he argued that the genes were not
fully formed and stabilized and so did not Mendelize yet. He also
transplanted ovaries between modified and unmodified salamanders to
test the influence of the recipient body on the donated eggs. The results
were inconsistent, but Kammerer felt they created more problems for
the Weismannian side than for himself.28
Taking the Show on the Road
For his work on the inheritance of breeding behaviors in salamanders,
Kammerer received his Habilitation, the qualification to offer courses at
the university, initially as a Privatdozent [unsalaried instructor]. He
capitalized on the academic rank right away to promote himself as a
science writer and lecturer. In July 1910, still waiting for the last
bureaucratic hurdles to be cleared ("Missing only signatures from the
Hausmeister [concierge] and the minister [of education]," he joked to
Iltis), he already had a full summer schedule of "big show-lectures" in
Munich, Berlin, and Vienna.29 He was to travel a great deal, lecturing
before local scientific societies, adult-education programs, and chapters
of the Monist League, associated with the Haeckelian world-view.
Although some university scientists might have thought his public
27
Kammerer, 1908, 1906, 1913, 1911-12, 1915, pp. 273-277. In English: Kammerer,
1912, 1924.
28
Kammerer, 191Ic.
29
Kammerer to Iltis, 6 July 1910, Kammerer Papers.
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SANDER GLIBOFF
performances detracted from his credibility, as Koestler claims they did,
Kammerer was in good company on the lecture circuit. Public lectures
and adult education were important sources of income for many
scholars languishing at the rank of Privatdozent. They also served
modern artists and thinkers as a medium for disseminating controversial
new ideas.30
Kammerer's lectures were given wide exposure in newspapers,
magazines, and scientific journals. For example, he spoke before the
Society of Naturalists in Brno several times, and the texts were often
published or discussed by Iltis in local newspapers.:3 A talk at the
Scientific Society of Berlin in 1910 was first printed in Himmel und Erde
and ended up in translation in the Smithsonian Annual Report.32He got
the most mileage from a preliminary report on the salamander coloration experiments, which began as lectures at the Society of German
Naturalists and Physicians in Salzburg in 1909 and the International
Zoological Congress in Graz in 1910. Versions were printed in the
proceedings of the former conference, and by Natur (Leipzig), Umschau,
and the German Society for Breeding, before entering the specialized
literature in the leading German genetics journal - the Zeitschrift fur
induktive Abstammungs - und Vererbungslehre,edited by Erwin Baur
[Journal for the inductive study of evolution and heredity, hereinafter,
"Baur's Zeitschrift"]. Following the Vivarium's arrangement, the full
report appeared in Roux's Archiv, but not until 1913.33 Kammerer also
used some of the same material in general presentations on inheritance
of acquired characteristics, of which his lecture and slide show at the
Society of Naturalists in Brno was probably representative.34
Kammereron Evolutionary Theory
During the contentious period between the death of Darwin (1882) and
the Modern Synthesis of the 1930s and 1940s , even self-styled Darwinians differed on the relative importance of natural selection and
30
Felt, 1996; Stewart, 1998; Stifter, 2002; Hofer, 2002.
Kammerer, 191ic; Iltis, 1921, "Jungbrunnen der Wissenschaft," Rote Armee,
Czernowitz, and Iltis, 1922 [?],"Paul Kammerer," both newspaper clippings, Kammerer
Papers. Kammerer also mentions lecturing engagements in Brno, Prague, Vienna, and
Germany: Kammerer to Iltis, 17 January 1920 and 1 September 1919, Kammerer
Papers.
32
Kammerer, 1912.
33 Kammerer, 191lb, 1913. These contain references to the other published versions.
34 Kammerer, 191Ia.
31
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THE CASE OF PAUL KAMMERER: EVOLUTION AND EXPERIMENTATION
537
other agents of change, the nature and causes of variation, and the
mechanism of heredity. Peter J. Bowler provides the best-known analysis of the period, which he describes, following Julian Huxley, as the
"6eclipseof Darwinism," because so many rivals obscured Darwin's
light. Kammerer appears in his account as one of the shadier
Lamarckians.35
Bowler successfully challenges the idea of a decisive Darwinian revolution, but the eclipse metaphor can also be misleading. It seems to
imply that an essentially modern Darwinism was available, just waiting
for the cloud to pass and the synthesis with Mendelism to begin. But
among the theories that Kammerer encountered, which was the future
Darwinism, and who was eclipsing whom? Here I will try to place
Kammerer into a context where modern Darwinism is not being
eclipsed, but created and debated, and where he is not an obstacle, but a
participant. Even if his answers were soon to be rejected, Kammerer
raised important questions about the nature and causes of variation at a
time when the concepts of genes and "random" mutations were still
unsettled.
One of the most important 19th-century interpretations of Darwinism in Germany was the neo-Darwinism of August Weismann, which
rejected the inheritance of acquired characteristics and diminished the
role of the environment and made internal processes in the germplasm
the main source of variation. Weismann's main Darwinian opponent at
the close of the 19th century was Haeckel, who thought Weismann
sacrificed the best explanation of variation and adaptation, namely the
inheritance of environmental effects. Darwin had clearly endorsed it,
and it was also mechanistic, in the sense that it had discernable physical
causes, unrelated to mysterious inner workings of the germplasm, or to
will, purpose, or divine providence. Haeckel worried that Weismann
opened the door to theism, under the assumption that the course of
evolution and all future adaptations were preformed or foreordained in
a primeval germplasm.36
Kammerer supported Haeckel's old-school (as opposed to "neo-")
Darwinism.37 With Richard Semon and Ludwig Plate, he continued
Haeckel's campaign to brand Weismann's isolated germplasm as arbitrary and hypothetical. A friend of Semon's and an admirer of
Haeckel's, Kammerer identified personally and theoretically with the
3'
36
37
Bowler, 1983, 1988. Also Huxley, 1942, pp. 22-28.
Haeckel, 1893, pp. vii-xi.
On Kammerer's relationship to Haeckel: Gliboff, 2001.
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538
SANDER GLIBOFF
old school. Kammerer even described Semon (along with Przibram) as
his mentor,38 and in his early publications, Semon is the authority
Kammerer cites to counter objections to the inheritance of acquired
characteristics. This group is commonly labeled "Lamarckian," but
they rejected any innate tendency to progress and any role for the
perceived needs of the organism in mediating evolutionary change. They
described responses of organic material to environmental stimuli as
strictly mechanistic. Most important, they recognized that individuals
inevitably differ in their exposure and responses to the environment,
hence that they will vary in fitness. Natural selection therefore still had a
role to play in completing the process of adaptation, which only begins
with environmental effects.39
Mendelian genetics deepened the divisions among the Darwinians
after 1900. Whereas neo-Darwinians warmed to it early and identified
their germplasm subunits [Determinanten]with genes,40 the old-school
Darwinians were divided on the subject. Semon rejected Mendelism and
developed an alternative system of heredity and development, his
Mneme theory of organic memory.41 Plate accepted Mendelism, but
without letting it be the exclusive mode of hereditary transmission.42In
contrast, Kammerer was committed to the inheritance of acquired
characteristics within a Mendelian and chromosomal conception of
heredity.
Most of Kammerer's quarrels were with the neo-Darwinians and the
Mendelian saltationists. The latter included his nemesis Bateson and the
more tolerant Richard Goldschmidt, who thought that evolution proceeded by discrete jumps, or large-scale mutations, taking a lineage from
one stable form to another, and with little or no significant variation
and selection in between. On the periphery of the Kammerer controversies were also theories of orthogenesis, which held that variation was
determined by laws of development that did not respond much to the
environment.
Kammerer envisioned the origin of favorable variation as a two-step
process of somatic induction. First, the environment induces the
organism to acquire new bodily or behavioral characteristics, then the
innovation is communicated somehow to the chromosomes for hereditary transmission. Extending the metaphor of inheritance, Kammerer
38
39
Anonymous, 1912.
Haeckel, 1866, vol. 2, pp. 191-193.
40 Ziegler, 1910.
41 Semon, 1908,
42
Plate, 1913b.
building on Haeckel, 1876; see also Hering, 1895.
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THE CASE OF PAUL KAMMERER: EVOLUTION AND EXPERIMENTATION
539
likened the chromosomes to a bank account from which inherited
wealth flows - perhaps in the form of hormones - to invest in the growth
and physiology of the body. Profits from acquired characteristics flow
back to the chromosomal coffers.43
This was intended to bring Darwinism up to date with genetics and
cytology, without giving up the inheritance of environmental effects:
"Darwin himself never assumed anything but an eliminative and isolating effect of selection and always ascribed any positive effects to the
environment...."
"We return directly to the original Darwinian standpoint when we
assert: the struggle for existence can produce nothing new, not even with
the help of selection. And we go a step further when we assert in
addition: struggle as such cannot even conserve the old. Struggle can
neither create nor preserve, but only destroy...." [Emphasis original.]44
Especially important to Kammerer were characteristics that led to
cooperation and symbiosis. As the title of his programmatic 1909 article
indicates, he viewed "General symbiosis and struggle for existence as
equally valid driving forces of evolution."45 Struggle provided external
stress, and organismal responses ranged from competition to cooperation. Cooperative responses were the most productive, evolutionarily,
because they brought out behaviors that stimulated the acquisition of
more new characteristics. Kammerer also ascribed political importance
to them.
Kammerer described himself as a socialist, but with reservations
about accepting any political label.46There is a liberal streak in him as
well and a concern for balancing individual and collective interests.
Even in a symbiotic relationship, he felt, each partner strove to take
advantage of the other and continually threatened to turn cooperation
into parasitism.47 Progress, including progress towards cooperation,
depended on individuals improving and asserting themselves: "Sooner
or later [my research] must gain general recognition for that which so
far, on the basis of my experimental experience, has become an
unshakeable conviction to few other than me: The power of the milieu
and the high significance of the individual for the history of its race!"48
43
Kammerer, 1925, p. 80.
44 Kammerer, 1909a, on pp. 603-604.
41
Ibid.
Kammerer, 1918b, pp. 9-10. lltis says he was a socialist, but not a party member:
Iltis, 1951, "Paul Kammerer," Kammerer Papers.
47 Kammerer, 1909a.
48 Kammerer, 1913, p. 8.
46
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540
SANDER GLIBOFF
Scientific Reception
The only writer I can find who backed Kammerer on every point was Iltis.
A socialist, he considered the inheritance of acquired characteristics a
necessary antidote to racism and selectionist eugenics, and he valued
Kammerer's efforts to make it compatible with Mendelism.49 He was
secretary to the Society of Naturalists in Mendel's hometown of Brno,
gave Kammerer's show-lecture some exposure there, and included him in
the Mendel-Festschrift of 1911.50 He also edited and annotated the
German edition of R. C. Punnett's book on Mendelism, in which he cited
Kammerer's work as proof of the heritability of environmental effects.5'
Kammerer's natural allies should have been the old-school
Darwinians, but Haeckel was not a friend of experimentalism and did
not get involved in public, and Plate was reluctant to be associated with
Kammerer, out of mistrust and anti-Semitism, as discussed below.
Semon was supportive, but differed with Kammerer on Mendelism and
methodology. He argued that history and environmental effects made
every individual unique, so one could not generalize from an experimental result. Nonetheless, until his death in 1918, Semon defended
Kammerer and used his results to support his own ideas on heredity.52
He also put in a good word for Kammerer with Haeckel, who valued
Kammerer's lectures and writings on monism, if not his experiments.53
Another important supporter was Przibram, but he, too, disagreed
with Kammerer on specifics. Przibram accepted the Weismannian barrier between germ- and somatoplasm, favored the compromise theory of
parallel induction, and was undecided about whether the chromosomes
were the physical bearers of heredity.54
After the war, Kammerer had an ally in the British embryologist
Ernest W. MacBride, who is usually called a Lamarckian, but had much
in common with the old-school Darwinians. However, MacBride, too,
disagreed with Kammerer on specifics. He was a gradualist, and
Kammerer's single-generation laboratory transformations did not fit his
conception of the naturalprocess.55Other criticsalso questionedwhethera
49 litis, 1927.
50
Kammerer, 1911a 1911c; Iltis, 1922 [?], "Paul Kammerer," newspaper clipping,
Kammerer Papers; Iltis, 1921.
51
Punnett, 1910, p. 117.
52
Semon, 1908, 1911b, 1912.
53 Semon to Haeckel, 16 January 1911, Ernst-Haeckel-Haus, Jena, Best. A, Abt. 1.
54 Przibram to Iltis, 23 July 1923, Kammerer Papers.
55 Bowler, 1984.
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THE CASE OF PAUL KAMMERER: EVOLUTION AND EXPERIMENTATION
541
trait that changed so readily could count as hereditary or play a role in
evolution.56
Lamarckians of various sorts had various reactions to Kammerer,
mostly negative. Some agreed that use and disuse and direct environmental effects were the primary causes of change,57 but others
emphasized the modifying power of the organism's survival needs,
either with58 or without59 a mediating role for the will or psyche.
Lamarckian ideas on hereditary mechanisms also varied. None accepted Kammerer's complete package of somatic induction, Mendelism, chromosomes, and experimentalism.
In Britain, H. G. Cannon attacked somatic induction, genetics, and
"heredity" as the term is commonly understood. He said form resulted
from dynamic interactions of the parts of the organism with each
other and with the environment. In a stable environment, similar
characteristics are produced anew in succeeding generations, giving a
false impression that something is transmitted. He rejected Kammerer's assertion that an acquired characteristic could persist after the
organism was returned to its original environment.60 Another British
Lamarckian whom Kammerer could not win over was J. T. Cunningham, an experimentalist, whose studies of fish coloration and
flounders' eyes provided standard arguments for the inheritance of
acquired characteristics. Cunningham even accepted somatic induction, just not Kammerer's evidence. He called Kammerer a victim of
self-deception and lamented that, "Lamarckian doctrine has often
suffered more from the indiscretion of its advocates than from the
attacks of its enemies.",6'
Ironically, Kammerer's embrace of Mendelism fell afoul of the
Mendelians, his worst critics. Although they did not all agree on the
nature and causes of variation they did not think it arose as an adaptive
response to environmental change or use and disuse. Several questioned
whether any changes reported by Kammerer fit their definitions of
mutation, and whether they were even caused by Kammerer's treatments. Saltationists considered the continuous traits Kammerer studied
to be irrelevant to evolution.
56
Weismann, 1913, pp. 74-75; Megusar, 1914.
Cunningham, 1892.
58 Pauly, 1905.
59 Cannon, 1959.
60
Ibid., pp. 43-47.
61
Cunningham, 1923a, b, quote from the latter.
57
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542
SANDER GLIBOFF
Goldschmidt gave Kammerer the most neutral evaluation, but
suggested alternative explanations of his results. For example, in one
of the less well-known midwife-toad experiments, Kammerer prolonged
the larval stage. One larva out of a large batch survived the treatment
and became sexually mature without metamorphosing. Goldschmidt
said that such a unique and fortunate individual might have
had a mutation in the saltationist sense, rather than an acquired
characteristic.62
Bateson's marginal notes and underlining in his copies of Kammerer's articles suggest that, like Goldschmidt, he was most interested in
the experiments that yielded the fewest transformed individuals, as if he
were looking for a saltation. Bateson focused on the nuptial pads of
the midwife toad as the best candidate. Everything else he dismissed as
just "a matter of more or less,"63 not significant in evolution, and
probably not even heritable. One case that Bateson discussed with
Przibram was the transformation of spotted salamanders to striped.
That change might have risen to the level of a genetic mutation by
Bateson's criteria, because the striped S. maculosa taeniata was recognized as a separate subspecies. However, Bateson pointed out that there
were many intermediate variants, and he thought the burden was on
Kammerer to establish the genetic basis of the differences before
claiming to have changed anything.64
Baur was not committed to saltationism, but did not see mutations of
any magnitude in Kammerer's experiments. In his 1911 textbook, Baur
classified Kammerer's reported changes as non-heritable "modifications." They recurred in the offspring if sperm, eggs, or their precursors
were affected by the environment while still in the parental body. Both
generations were modified simultaneously; the trait did not pass from
one to the next. Baur was prepared to argue that the protoplasm of as
many as three generations, nested one within the other, could have been
modified by Kammerer's treatments.65But when Kammerer reported a
fourth generation of water-breeding midwife toads, Baur reached his
limit and dismissed Kammerer's account as vague and unreliable.66
Several opponents pointed out that Kammerer's results were compatible with other mechanisms besides somatic induction, and some
62
Goldschmidt, 1913, pp. 456-469.
Bateson to Quastel, 24 April 1923, Bateson Papers, American Philosophical Society
Library, Philadelphia, History of Science Collection, Microfilm no. 26, Reel E, Section
Balto #32, Index 5 [hereinafter, "Bateson Papers"].
64 Bateson to Przibram, 21 October 1920, Bateson Papers, Index 7.
65 Baur, 1911, pp. 37 & 260.
63
66
Baur, 1919, p. 55.
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THE CASE OF PAUL KAMMERER: EVOLUTION AND EXPERIMENTATION
543
demanded further information about the genetics and physiology of his
modified traits in order to narrow down the possibilities. Weismann and
other neo-Darwinians suggested that Kammerer had caused atavisms,
or reversions to ancestral forms, latent in the germplasm, which were
therefore old characteristics, not acquired ones. Either that or inadvertent selection was at work.67 A similar criticism from the Mendelian
side was that Kammerer failed to use highly inbred and genetically
uniform "pure lines" as defined by Wilhelm Johannsen. He might
merely have isolated pre-existing mutations from his starting populations. 68
In hindsight, Kammerer's accounts are transparent enough to reveal
where selection could have occurred. High mortality rates can be
inferred (e.g., in midwife-toad eggs laid in water), and variation was
available: Kammerer mentioned intermediates between S. maculosa and
S. atra; striped, spotted, and other patterns were known in S. maculosa;
there are long- and short-siphoned populations of the sea squirt (Ciona)
in the Mediterranean; and midwife toads with rudimentary nuptial pads
had been described.69The changes that Kammerer reported were within
the species' known ranges of variation (or phenotypic plasticity). There
is no need to invoke fakery to defend neo-Darwinism.
Kammerer considered the possibility of selection, but rejected it. In
the 1908 salamander paper, he wrote: "I did not in any way need to
select the offspring of those specimens that displayed the altered
reproductive habits most sharply. It was at most just a question of time
before the adaptations could be brought about, in an equal degree, in all
the specimens that were exposed to the relevant factor, insofar as they
could bear the changed conditions at all." [Emphasis added.]70There was
the problem: not all the animals could bear the changed conditions,
hence the likelihood of selection.7'
Kammerer responded to alternative interpretations of his results
impatiently. All together they seemed to add up to pure sophistry:
67
Plate agreed with the atavism objection at first: Plate, 1908; later changed his
opinion: Plate, 1913a, pp. ix & 468-478. Weismann and Ziegler suggested both atavism
and selection as explanations Weismann, 1913, pp. 74-75; Ziegler, 1910.
68
Hanstein, 1911.
6)
On nuptial pads in midwife toads: Kaindler, 1924. This information was apparently
still unknown to Kammerer and Przibram in 1926. On Ciona: Whittaker, 1985, but note
that he argues for fraud, not error.
70
Kammerer, 1908. p. 42.
71 Kammerer, 1915, pp. 268-276, adds that some treatments were too innocuous to be
selective, e.g., background color or cutting off the siphons of Ciona, and that some of
the acquired characteristics (e.g., lengthened siphons) were not adaptive.
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544
SANDER GLIBOFF
"Whatever is altered, one [critic] says it is an acquired characteristic and
therefore not heritable; the other says it is heritable and therefore not an
acquired characteristic."72In the case of salamander coloration, Baur
and Goldschmidt allowed that the trait was acquired, but Baur said it
was not transmitted, and Goldschmidt that it was transmitted but not
genetic. For Bateson it was a continuous trait and therefore uninteresting. Meanwhile, other geneticists claimed, with Weismann, that the
changes were hereditary, but selected and not acquired. Given the lack
of consensus on what constituted a hereditary trait or a mutation, or
what else might be going on in his experiments, perhaps Kammerer's
complaint was justified.
However, Kammerer's pipeline system did not allow him to clarify
matters, had he been inclined to do so. By the time an experiment was
published and criticized, similar ones, on other species, were in the
pipeline and they would fail similarly to answer the critics and eliminate
alternatives. Moreover, Kammerer's experimental designs dated from
the earliest days of genetics, before the pure-line concept and before
geneticists settled on model organisms and methodological standards.
From the geneticists' point of view, Kammerer was living and working
in the past. Kammerer could go so far as to agree that a pure line would
have been the ideal starting population, but he could not easily redo his
experiments, and breeding a pure line was unfeasible in animals with
such long generation times as his salamanders. So he argued that it was
an unreasonably high standard, that his starting populations were
uniform enough in the relevant traits, and that it was safe for him to
assume that he was working with simple Mendelian characters.73
When pressed, Kammerer acknowledged that he could not rule out
every alternative explanation, and was prepared to retreat to the following minimal claim: "...I maintain that no matter whencethe ability to
react purposefully derives, whether it is atavism or selection, one thing
remains unaffected: changes have arisen under the influence of external
factors, and those changes have appeared again in subsequent generations, despite the absence of these factors." [Emphasis original.]74 In
other words, a heritable change had occurred, and the environment had
caused it. Even if he could not elucidate the intermediate mechanisms,
the change was initiated by environmental influences and not by
internal, orthogenetic, teleological, or random processes.
72
73
74
Kammerer, 1920a, p. 32.
Kammerer, 1913, on pp. 30 & 163-164.
Kammerer, 1908, pp. 42-43.
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THE CASE OF PAUL KAMMERER: EVOLUTION AND EXPERIMENTATION
545
Growing Mistrust
Deficiencies in his designs and alternative explanations of his results
were only part of Kammerer's problems. As Przibram complained
to Iltis, "One must distinguish between the scientific attacks on K. and
those that are made not for scientific reasons. Here the gentlemen from
Germany are accomplishing great things [leisten die Herrn
ReichsdeutschenGrosses] and will embarrass themselves very much."75
Scuttlebutt about Kammerer's vanity, his penchant for self-promotion, and his Jewish background made some potential allies keep their
distance. For example, the cell biologist Theodor Boveri, who had some
sympathy for Lamarckian evolution,76 considered hiring him for the
planned Kaiser Wilhelm Institute, but decided against it: "...I have met
Kammerer briefly and seen his salamanders and cave olms. We really
must get a man who can work with animals that way, and it sure seems
like he could be gotten cheaply. But unless Braus turns us down, we
can't even think about it. And besides, there is of course the question of
whether his extraordinary characteristics are not connected with other,
unpleasant ones."77
Goldschmidt heard from a friend in Vienna that the Vivarium was
full of over-inflated egos like Kammerer's that just begged to be popped.
Kammerer was "personally repulsive," and the attention he got from
the Jewish press aroused jealousy and gave him and Przibram reputations as publicity seekers. Moreover, the laboratory as a whole was
resented and mistrusted because it was viewed as an exclusively
"Semitic" institution.78
Plate vacillated in his support of Kammerer. Skeptical at first,79he
accepted Kammerer's counterarguments for a time,80 then turned
against him because of the innuendo,8' particularly Megusar's accusations, discussed below. Later still, after the scandal, Plate added antiSemitic invective: "All these reports by Kammerer the Jew must be
taken with extreme skepticism....The emphasis on race seem to me
75
76
Przibram to lltis, 23 July 1923, Kammerer Papers.
Boveri, 1906.
77
Boveri to Goldschmidt, 16 January 1913, Richard Goldschmidt Papers, Bancroft
Library, University of California, Berkeley, Box 1, Boveri folder [hereinafter, "Goldschmidt Papers"].
78 Eugen Neresheimer to Goldschmidt, 1 June 1909 and Neresheimer to Goldschmidt,
30 December 1909, Goldschmidt Papers, Box 3, Neresheimer folder.
79 Plate, 1908. Here he ascribed the changes in the midwife
toads to atavism.
80
Kammerer, 1908-9; Kammerer, 1909b, p. 524; Plate, 1913a, pp. ix & 468-478.
81 Plate, 1922-1924, p. 9; Plate, 1925, p. 119.
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SANDER GLIBOFF
appropriate here, because the Kammerer case is a good example of the
unscrupulousness of the Jews."82
Weismann evidently heard a thing or two about Kammerer from his
former student Richard Woltereck, who worked for a time in Lower
Austria. He wrote in a notebook: "Kammerer (Vienna) is a little, miserable, sticky Jew, who has proven himself on earlier occasions to be a
quite unreliable worker (Woltereck). 25/3/10.,,83 It is no wonder that he
later declared Kammerer's results to be "hardly credible."84
Kammerer was badly served by his own practice of trumpeting
conclusions in the popular press and the show-lecture before writing up
the formal research paper. For example, early accounts of his salamander color-change experiments85 left out crucial details about
rearing- and recording methods, environmental controls, and the
appearance of the modified animals. They provoked suspicious and
acerbic responses which put Kammerer on the defensive before he wrote
the full report for Roux's Archiv.
The herpetologist Edward Boulenger accused Kammerer of procedural error, based on those incomplete previews. He suggested that
Kammerer had not kept his salamander colonies strictly separated and
might have let his experimental lines mix. He also called attention to the
fact that Kammerer did not mark his salamanders. Kammerer raised
them in groups of 40 to a terrarium and boasted that his practiced eye
could recognize every individual - this despite the fact that the goal of
the experiment was to make the salamanders' appearance change.
Boulenger doubted that Kammerer could follow the progress of individual salamanders.86
Some critics wrote to Kammerer for more information, asked to
borrow specimens for examination, or visited the Vivarium. Bateson
was most persistent in pursuing information on the midwife toads. He
doubted they had acquired all the features of nuptial pads at once, in the
saltational manner. Kammerer had written little more in 1909 than that
there were "black-gray discolored calluses on the upper side of
the thumb and on the ball of the thumb,"87 which was ambiguous,
82
Plate, 1933, vol. 2, p. 1175.
Weismann, notebook "1909: 11. 9 Aug. 1909 bis 26 Juli 1910," p. 100, NachlaB
August Weismann, Universitatsbibliothek Freiburg, Freiburg im Breisgau, box labeled
"Tagebucher/Notizbucher, 1909-1914" [hereinafter, "Weismann Papers"]. On Woltereck, Harwood, 1996.
84 Weismann, notebook "1911. 3," p. 50, Weismann Papers.
85
See p. 33, above.
86
Boulenger, 1911, p. 346. This was a response to Kammerer, 1910.
87
Kammerer, 1909b, p. 516.
83
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THE CASE OF PAUL KAMMERER: EVOLUTION AND EXPERIMENTATION
547
especially since the ball of the thumb is not the normal location in frogs.
Bateson and the herpetologist G. A. Boulenger (father of E. Boulenger,
above), whom he consulted frequently, felt they still needed to know the
precise location of the pad, whether it was thickened and rough in
addition to being dark, and whether the roughness was due to characteristic asperities (horny spines and spicules). Bateson could not get
Kammerer to send him a specimen or to show him one when he visited
the Vivarium in September 1910, apparently because he had not preserved any, and the live toads only exhibited the pads in the spring.
Koestler, following Kammerer, portrays Bateson's requests as
unreasonable, on the grounds that the midwife toads were not a central
part of Kammerer's overall argument. He says that Bateson should have
focused on the sea-squirt experiments, which provided much more
decisive evidence. However, in Kammerer's pre-war writings the salamanders and midwife toads really were the mainstays of his case for the
inheritance of acquired characteristics. Kammerer downplayed them
only after criticism mounted. Even then, he never published a formal
research paper on sea squirts, so he could hardly rest his case on those
results. Besides, siphon length was a continuous characteristic, which
Bateson would not accept as evidence in any case.
The problem highlights Kammerer's inattention to documentation.
He did not preserve modified individuals in a routine manner and never
found a satisfactory balance between producing results and documenting them. Typically, his experiments yielded very few modified
individuals - sometimes just a single one - and the more he killed and
preserved, the fewer he had left for breeding and follow-up experiments.
He sacrificed all of his modified cave olms for display, for example. He
regretted the decision,88 and seems never again to have erred on the side
of documentation. He sometimes referred readers to his living specimens, but they were not always suitable for showing as evidence,
because their appearance was not constant. Salamanders continued to
change their spots after the experiment, the midwife toad did not exhibit
nuptial pads when Bateson came calling, and none of modified stocks
lived through the war.
Kammerer's published illustrations did not compensate for the lack
of preserved specimens. He appears not to have been able to draw very
well, and the Vivarium was not well equipped for photography. His
photos of whole animals were few and of the very lowest quality. Most
were made at studios in town, which were unaccustomed to having
glistening, wet amphibians sit for portraits. They are almost always
88
Kammerer, 1920a.
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548
retouched or out of focus, or they simply do not show the details
described in the text. They might have been good enough for the showlectures, but they did not stand up to scientific scrutiny.
Conflict with Erwin Baur
Kammerer called attention to his own documentation problems by
picking a fight with Baur, whose Zeitschrift had published one of
Kammerer's preliminary reports on salamander coloration in 1911.89
Kammerer was frustrated by the criticism it drew, and when he finally
published the full-length research paper in Roux's Archiv in 1913, he
lashed out at Baur, E. Boulenger, and others, and accused Baur of
reproducing his photos poorly and misplacing some of the captions.90
However, he did not provide improved photographs in the 1913 article.
He now drew the changing spots by hand on a series of schematic
salamander outlines. Only a few photographs were included as "supplementary documentation," but they were colored over so heavyhandedly in black and yellow that they look as if renderedentirelyby hand.
Baur responded angrily in Roux's Archiv. He said Kammerer's
originals had all the shortcomings Kammerer was blaming on his
Zeitschrift: "They are all very bad, extensively retouched photographs.
The photo laboratory got as much out of them as could be done, that is
to say, they reproduced the originals faithfully. [Emphasis original].91
Baur recognized some of Kammerer's Archivphotos as versions of those
from the Zeitschrift, with the same added spots, but without any indication that spots had been added: "...Whereas the figures in the 'Zeitschrift' reproduce exactly what can be seen on the photographs, the
photographs...in the 'Archiv,' supposedly only 'colorized,' are redrawn
to such an extent as to lack any value as 'documents."' Note, however,
that Baur did not object to retouching in itself. After all, he had
accepted Kammerer's photos for the Zeitschrift article. The problem
was with saying they were ";only colorized" and claiming they were
better documentation than schematic drawings: "No one will be able to
deny that a photograph that is supposed to serve as a piece of
documentary evidence may not be redrawn in the way it was done.. .in
the 'Archiv."'92
89
90
91
92
Kammerer, 191lb.
Kammerer, 1913, p. 6n.
Baur, 1914, p. 682.
Ibid., pp. 682-683.
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Kammerer replied that the plates were honest likenesses, even with
the hand coloring. They "...were portraits, produced under multiple
controls. Since they are not autochromatic images, but hand-colorized,
any well-intentioned judge knows that the term 'supplementary documentation' cannot be meant to exclude subjectivity." The important
thing was not to exclude all the subjectivity, but to include all the spots:
"...The glare from the skin gave the impression of spots where none was
present, and spots that were present were washed out in the glare. With
my preserved specimens I can demonstrate the correspondence of the
pictures with the objects any time. From this results the lack of correspondence with the figures in the 'Zeitschr. f. indukt. Abst.- u.
Vererbungsl., which for this reason had to be improved and published
again in the 'Archiv."'93Kammerer assumed that readers would take his
word for the likenesses, but he did not have everyone's trust.
Conflict with Franz Megusar
Later in 1913 came embarrassing accusations from Megusar. In a paper
presented to the German Society of Naturalists and Physicians, Przibram's second assistant cast doubt upon Kammerer's care in maintaining environmental conditions as well as the veracity of his
publications. The two had fundamental differences on theory. Concerning blind cave animals such as Kammerer's olms or the crustaceans
he worked on himself, Megusar argued that development of the eyes in
response to light spoke against the inheritance of acquired characteristics. He argued that changes effected in a single generation could not
play a role in evolution, because any trait that changed so readily could
not have been truly hereditary in the first place: "...If long-lasting fix-
ation of externally induced changes were possible, then nature would
have accomplished it, given the immeasurable time periods and the
radicalism of the influences. The cave animals should have retained the
characters they had acquired in the darkness (pale coloring, rudimentary organs) for at least one generation in the light.",94 Although
Kammerer later called him a Weismannian,95because of his rejection of
the inheritance of acquired characteristics, Megusar was more of an
orthogenecist. He said that first and foremost "inner causes,"
93
94
95
Kammerer, 1914a.
Megusar. 1914, p. 718.
Kammerer,1920.
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SANDER GLIBOFF
rather than environmental influences, drove evolutionary change and
accounted for the diversity of forms.
Megusar's work on salamander coloration, although similar to
Kammerer's, supported his own view that the environmental effects
were unimportant. Background colors had no effect on his salamanders'
spots, and he attributed the discrepancy to sloppiness and dishonesty on
Kammerer's part: "The processes that Kammerer reports I could not
confirm, neither in his experiments nor in my own, even though I have
been monitoring his extremely imprecisely executed experiments constantly for nearly 10 years. I checked up on the laboratory workers as
they maintained his experiments, and I supervised them personally for
several years. Kammerer's representations contain crude untruths and
falsifications of the actual circumstances."96
According to Iltis, personal animosity motivated the attack. He
recalled a visit to the Vivarium: "Megusar came to the door. When I
told him that I would like to see Dr. K. he left me standing in the door
after giving me a scornful look, and walking away with an impolite
growling. When I told K. about this incident, he smiled and told me that
he knows that Megusar would poison him if he had a chance." Iltis does
not explain the origin of the hostility, but it was bad enough to make
him suspect Megusar of doctoring the midwife toad. However, Iltis
changed his mind upon learning that Megusar had fallen in the First
World War, too early (presumably) to have framed Kammerer.97
Przibram seems to have been caught entirely off guard by Megusar's
paper, which was extremely embarrassing, especially because it was
given at a meeting in Vienna and Przibram was trying to show off his
laboratory to visiting scientists. Although Przibram vouched for the
authenticity of Kammerer's results, he never fully distanced himself
from Megusar, at least not in public.98
Kammerer'sPostwar Career
As the exchanges with Baur and Megusar show, Kammerer's reputation
was already suffering before the war shut off his experimental pipeline.
Przibram had the foresight to donate the Vivarium to the Academy of
Sciences, who kept the laboratory afloat through the war and the
96
Megusar, 1914, p. 719.
Iltis, 1951, "Paul Kammerer," Kammerer Papers.
98
Later and privately he said Megusar was mentally ill: Hirschmuller, 1991, pp. 42-43;
Przibram, 1918; and Przibram to Iltis, 9 February 1930, Kammerer Papers.
97
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postwar hyperinflation that consumed the Przibram fortune.99 The
Vivarium carried on with a skeleton crew that included Kammerer's
wife, but Kammerer had to work for the military censor. His stocks of
modified animals died of disease and neglect, but he found a new outlet
for his ideas in pacifist essays that made evolutionary arguments against
war and for cooperation between individuals, races, and countries.'00
He also began work on a big book about coincidences, which came out
in 1919 as Das Gesetz der Serie [The Law of Series].10' It attempted to
make coincidences amenable to scientific study, but it smacked of
mysticism and pseudo-science, and it cost him a promotion to professor,
for which he had applied in 1918.
He thought he had a good case for promotion, because he had been a
Privatdozent for 8 years, had published a great deal, and because he was
only asking for the rank (i.e., of auJ3erordentlicherProfessor), not a
position on the payroll. He also had good backing. Haeckel wrote in on
Kammerer's behalf,'02 and Hatschek and Przibram, as members of the
promotion committee, were able to downplay the clashes with Baur and
Megusar.'03 After months of deliberation, the committee was leaning
Kammerer's way, but then Gesetz der Serie turned the tide against
him.'04
Kammerer did little new research after the war, but wrote up a
backlog of results, including a 1919 follow-up on the midwife toad. It
provided photographs of Kammerer's last specimen (preserved and
photographed in 1913) and of microscopal cross sections from one of its
nuptial pads, which the text described as follows: "The first finger, on its
upper and outer sides, as well as on the ball, was discolored black-gray
and at the same time was somewhat thickened.'105 But the paper only
provoked new criticism.
Bateson thought that Kammerer had never established the dark
patches as nuptial pads in the first place, and that he now contradicted
the 1909 paper by moving them from the "thumb" to a different side of
a different finger. G. Boulenger told Bateson he could not see asperities
99 Anonymous, 1914.
'?? Kammerer, 1914c, 1918a, 1919b.
""1 Kammerer, 1919a.
102 At Kammerer's request, Haeckel wrote to Hatschek and possibly Grobben: Kammerer to Haeckel, 27 March 1918 and 14 April 1918, Haeckel Papers, Ernst-HaeckelHaus, Jena, Best. A, Abt. 1, Nr. 3 165/1-7; see also KrauBe, 1998.
103
For details, see Hirschmiiller, 1991.
104
Przibram, 1926b, p. 403.
?5 Kammerer, 1919c, p. 327.
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SANDER GLIBOFF
in the photo and that the photo might even prove they did not exist.'06
Acrimonious exchanges in Nature ensued.
E. Boulenger examined the specimen at the Vivarium in 1922 and
reported that: "Nearly the whole hand is colored black, but I could not
detect any obvious rugosities. The black color might have been merely
pigment." However, because spines and spicules were visible in the
cross-sections, Boulenger seemed willing to believe the pad originally
did have a rough surface: "I had a long talk with Przibram who not only
takes full responsibility for all K's experiments, but states that he was
present when the sections were cut which prove that that the black color
is due to horny substance and not pigment. If we still disbelieve we must
assume that Przibram is a dishonest person."'07
Disillusioned with the University and with scientific research,
Kammerer quit the Vivarium in 1921 and lived from writing and lecturing, which took him to Britain and the U. S. in 1923 and 1924. The
show-lectures changed in character, as did Kammerer's attitude toward
evolutionary progress. He no longer considered steady progress to be
the natural and inevitable outcome of evolution. He now called on
humanity to take charge of its own biological and cultural future.
Recent findings from the Vivarium would provide the tools for a new
"organic technology" [Organik,as opposed to Technik],a more effective
and humane alternative to selectionist eugenics.'08
Selectionist eugenics, Kammerer argued, could only preserve desirable traits that happened to be present in the population. In contrast,
organic technology would create new ones. Steinach's work on glandular secretions and their effects on body and mind suggested how new
traits might be induced, and Kammerer's research showed that induced
changes could become hereditary. The postwar show-lectures blended
the two lines of work and touted the possibilities for improving
humanity. Thyroid, thymus, pituitary, and adrenal preparations could
adjust bodily proportions, intelligence, virtues and vices, likes and dislikes; testicle implants could cure homosexuality; and X-ray treatments
might produce the same effects by stimulating the growth of glandular
tissue.
On tour in 1923, Kammerer also displayed his midwife toad and
other specimens in Cambridge, where he gave a lecture and tried to
106
Bateson, 1919; Boulenger to Bateson, 23 June 1919, Bateson Papers, Index 3.
Actually Kammerer's two descriptions are of two different specimens.
107
Boulenger to Bateson, 24 January 1923, Bateson Papers, Index 3.
108
Kammerer, 1920, "Entwicklungsmechanik der Seele." Der freie Gedanke, Prague,
and Kammerer, [1921?], "Organische und soziale Technik," both newspaper clippings,
Kammerer Papers; Kammerer, [1920].
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settle his dispute with Bateson, but Bateson would not take more than a
cursory look at the nuptial pad. He conceded nothing: "The question
remains, what is the real nature of the swellings in the animal exhibited?
That on the palm did not look like a nuptial pad."'09 Indeed, the palm is
the wrong side for a functional nuptial pad, because it faces away from
the female.
Kammerer was better received in the Soviet Union. His ideas resonated in doctrinaire Marxist circles, who shared his commitment to the
inheritance of acquired characteristics and human improvability. On a
1926 visit, the Communist Academy offered him the directorship of a
new laboratory to be built under its auspices in Moscow.'10 Kammerer
was unconvinced that the laboratory would materialize, and unhappy
about leaving Vienna, but initially he accepted the offer.
What must have made the Russian offer attractive was the brewing
scandal. It had not yet been publicized, but G. Kingsley Noble, a herpetologist from the American Museum of Natural History in New York
had come to Vienna in January or February 1926 to examine the
midwife-toad specimen. Kammerer was informed of the visit and
assented to it, and he went to Moscow knowing that Noble found the
India ink. Like Bateson, Noble had doubted whether the reported
modifications were truly nuptial pads, but to him the problem was not
their position, color, or texture, but the skin glands in Kammerer's
cross-sections. They were not greatly enlarged as they would be in
frogs.'11 However, there is no reason to think Noble suspected fraud
before he saw the specimen.
The Midwife-toad Scandal
Noble reported not only the presence of ink, but the absence of
asperities, and concluded that there was no trace of a nuptial pad on the
I
specimen. 112 He now rejected the evidence of the cross sections, suspecting that they were from a frog - this despite his previous judgment
that they were not frog-like enough, because of the skin glands. There
were also three photographs of the specimen, dating to when it might
still have had asperities, but Noble rejected this evidence as well. No
109 Bateson, 1923a; see also Bateson,
1923b, c.
'o
Hirschmuller, 1991; Gaissinovitch, 1980.
111 He first made this argument in an unpublished talk before the British Association
for the Advancement of Science in 1925, and he repeats it in Noble, 1926a.
112
Noble, 1926b.
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SANDER GLIBOFF
asperities were visible in the two photos available to him, from 1913113
and 1923.114 A third, from 1922,115 was said to exhibit spines, but Noble
inferred from verbal descriptions that these were bigger than the spines
in the cross sections and must have been foreign objects. Finally, Noble
dismissed the eyewitness accounts from Cambridge. He said the rough
texture some of them reported must have been due to the uneven distribution of India ink granules, not to genuine asperities. Noble could
not imagine a qualified herpetologist being fooled."6
Przibram agreed that the midwife-toad had been tampered with and
could no longer be presented as evidence for Kammerer's claims.
However, he insisted that it did have nuptial pads once but they had
sloughed off over the years and on its travels. He still had the evidence
of the cross-sections, which were distinguishable from any taken from
frogs. He had a list of scientists who had looked at the specimen in
Cambridge in 1923 and seen nuptial pads. And there was the third
photo, which in his opinion had two or three distinct spines at the edge
of the "hand." He obtained letters from Cambridge biologists attesting
to the authenticity of the photo." 7
Przibram reported that Kammerer was "greatly astonished" and
could not account for Noble's findings."8 Kammerer's only published
remarks on the scandal were in his last letter to the Communist
Academy, dated 22 September 1926. The Academy published it, and an
abridged translation appeared in Science. Kammerer acknowledged
Noble's findings and predicted that the apparent forgery, "...as far as
one can expect will be blamed on me alone." He declared his innocence,
but felt that under such a shadow he could no longer accept the job in
Moscow and go on with his research, or even with his life. The next day
he shot himself on a mountain trail south of Vienna."19
Did Kammerer really did inject his midwife-toad specimen with India
ink? And if so, then what is the answer to his rhetorical question to Iltis?
How could an intelligent scientist do something so blatant and foolish,
especially if, as I have suggested, he had really produced nuptial pads?
Taken at the Vivarium by a visiting researcher; published in Kammerer, 1919c.
Taken on Kammerer's visit to Cambridge; published in Kammerer, 1924.
115
Taken at a studio in Vienna; published in Kammerer, 1925; detail reproduced in
Koestler, 1971.
116 Noble, 1926b.
"3
114
117
Przibram, 1926a.
Przibram cites a letter from Kammerer, dated 18 February 1926, so Noble must
have made his discovery earlier that month or late January.
"'9 For the full text: Hirschmuller, 1991, p. 62. See also Kammerer, 1926.
118
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I think the Baur incident suggests that he might have been just foolish
enough back in 1913 to touch up his evidence.
A letter to Nature in 1927 suggested that someone might have wanted
"to bring out some of the natural markings [of the midwife toad] with
ink so that they would reproduce better in print."'20 Although Przibram
dismissed the idea,'2' bringing out skin markings had been a problem
for Kammerer before, and in 1913 he did not think that retouching
amounted to falsification of evidence. That was a defensible position,
too. Photography was not yet the norm in Baur's Zeitschrift or Roux's
Archiv, and standards of presentation were in flux. Baur had not
objected to the retouching, but to the labeling. Kammerer need not have
used photos at all, but could have rendered his evidence entirely by
hand. After all, when a midwife toad with nuptial pads was collected in
the field in 1924, the report was illustrated by hand, and no one questioned its veracity.122
Perhaps Kammerer took just a little more artistic license with the
midwife toads than with the salamanders. Instead of touching up the
photos, he used ink on the toad to reduce the glare. He could easily have
justified it, at least to himself, using the same logic with which he
responded to Baur - especially if he did it in 1913, when the earliest
photo was taken, and before his run-ins with Baur and Megusar. After
those embarrassing experiences, he would have had to be "a Dummkopf
or an idiot" to touch anything up, but beforehand he might have been
naive enough to try it.
Przibram dated the injection to 1921 because neither of Kammerer's
midwife-toad articles (1909 and 1919) mentions darkening of the palm,
but darkening is visible in the 1922 photo.'23 This dating saves face for
the Vivarium, since it lets all the work Kammerer did while still a staffmember precede the tampering, and it directs suspicion away from
Megusar, who died in 1916. However, the 1909 paper does not describe
the specimen in question, and the 1919 description is very brief. Its
failure to mention darkening of the palm does not mean that none was
present.
Koestler's dating to 1925 is also weak. He argues that the Cambridge
biologists would have recognized the ink in 1923 if it had been there,
and that even if they had missed it, it would have dissolved and faded by
120
121
122
Kiplinger, 1927.
Przibram, 1927.
Kandler, 1924.
123
Przibram, 1927. Later he suggests cryptically to Iltis that something could have
happened in Cambridge in 1923: Przibram to Iltis, 9 February 1930, Kammerer
Papers.
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SANDER GLIBOFF
1926. The latter claim is based on unsuccessful attempts by the biologist
Holger Hyden to replicate the forgery in 1970.124This dating leaves all
of Kammerer's research and all three photos above suspicion, and
suggests that the injecting was done just before Noble's visit, the perfect
time for an enemy to set Kammerer up. However, Hyden did not know
what materials Kammerer had used. His failed replication only shows
that some inks dissolve and fade in some preservatives. As for the testimony of the Cambridge biologists - who among them looked carefully, given the setting at Kammerer's lecture, and who was expert
enough to know what to look for? The only thing they all agreed on was
the dark coloration. Only a few reported seeing a roughened surface,
and none specifically described characteristic asperities. The best qualified observer, E. Boulenger had specifically noted their absence in 1922.
The Cambridge people probably saw the same thing in 1923 as Noble
saw in 1926: a patch of skin made dark and uneven by India ink
granules.
None of these arguments is conclusive, but one offers an answer to
Iltis: the earliest possible dating, to 1913, when Kammerer was still
unworried about accusations of dishonesty, and still could refer skeptics
to his stocks of live toads. Later, once the stocks were lost, it would have
been difficult for him to discard the inked specimen. He did not have
any other midwife toads to display, and the nuptial pads were among
his standard examples in the show-lectures and popular writings, which
became his main source of income in the 1920s. He might have thought
it was worth the risk if only he could keep his show on the road.
Conclusion
It takes more than one person to cause a scandal. Someone has to do the
scandalous deed, others must become suspicious and uncover it, and
still more people must care enough to be scandalized by it. Thus, even
granting that Kammerer tampered with the toad, the scandal is not fully
explained by Kammerer's actions. We must also consider the reasons
why his work attracted so much attention and scrutiny at the time.
For Kammerer's opponents, the scientific dispute was not simply
about whether acquired characteristics could be inherited by means of
somatic induction, but about how else to explain Kammerer's results.
What were the nature and causes of variation? The varied responses to
his work reveal the disunity of geneticists and evolutionists on that
124
Koestler, 1971, pp. 104-115.
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557
subject. Kammerer was able to keep them engaged in debate by
persistently raising a difficult question and offering "Lamarckian" answers that could not easily be refuted.
The other proposed answers bring out the wide range of theories that
flourished during the so-called eclipse of Darwinism. They
included spontaneous saltations, atavisms to ancestral traits stored in an
immortal germplasm, environmental "modifications" that could persist
for two or three generations, organic memory, and parallel induction.
Within this context, it is clear that Kammerer did not stand simply for
"Lamarckism" and against "Darwinism" in a two-sided debate. His
was one of many competing positions, as he aimed for a new synthesis
of natural selection and the inheritance of acquired characteristics with
the latest developments in genetics, chromosomal inheritance, and
experimentalism, and with a vision for evolutionary progress in the arts,
education, and culture.
In the history of experimentalism, Kammerer, and indeed the whole
Vivarium enterprise, broaden our picture of the experimental turn in
whole-organism research. They show that there were diverse motives for
rejecting the old morphology and that the development of experimental
institutions and approaches were influenced by local needs and resources
as well as by international trends. The Vivarium benefited from Viennese
industrial capital, middle-class engagement in the arts and sciences, and a
population of Jewish Privatdozenten, forced to seek their scientific fortunes outside the universities. Kammerer adapted his research practices
to his situation in Vienna and at the Vivarium. His experiments showed
off his creative skills and provided tales of animal transformations with
which to regale and educate show-lecture audiences, while also displaying the potential of the experimental method and the laboratory's
technological capabilities. In return, Przibram and the Vivarium provided Kammerer with an opportunity for a research career outside the
university, where he was free to follow his own scientific inclinations and
pursue his musical, cultural, and popularizing side-lines.
However, what made sense in Vienna and at the Vivarium, did
not always meet the expectations of experimentalists elsewhere. The
Vivarium environment encouraged him to work with a wide variety of
organisms, mostly long-lived, slow to produce results, and little known
to geneticists. Greater interest in genetics at the Vivarium might have
prompted Kammerer to respond to the critics by investigating the
genetic basis of the traits he worked with, beyond making a small
number of backcrosses of modified toads and salamanders. Most fatefully, the Vivarium did not cultivate the art of photography or impose a
system of preserving and presenting results.
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SANDER GLIBOFF
Finally, there is the story of an individual playing two professional
roles, as a researcher and a public intellectual in fin-de-siecle Vienna.
His popular writings and lectures were more than just a source of
income. Kammerer needed his message to reach artists, musicians,
teachers, laymen, and students at all levels. His choices of organisms
and methods, and even his display of the midwife toad, were aimed
towards demonstrating the transforming power of the environment to
these varied audiences, rather than elucidating mechanisms of heredity
to the satisfaction of academics.
Acknowledgements
This research was supported by a grant from the National Science
Foundation. The article has been through many incarnations as talks
and drafts. My thanks to all who listened, read, and commented. For
a critique of the last draft I am indebted to Melinda Fagan and participants in the Biology Studies Reading Group at Indiana University.
References
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