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Paul Kammerer and the Inheritance of Acquired Characteristics

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Paul Kammerer and the Inheritance of Acquired Characteristics Contributions to Zoology, 85 (4) 457-470 (2016) Paul Kammerer and the inheritance of acquired characteristics Jacques J.M. van Alphen1, 2, 3*, Jan W. Arntzen1* 1 Naturalis Biodiversity Center, P.O. Box 9517, 2300 RA Leiden, the Netherlands 2 Institute for Biodiversity and Ecosystem Dynamics, University of Amsterdam, P.O. Box 94248, 1090 GE Amsterdam, the Netherlands 3 E-mail: [email protected] * Both authors contributed equally to this work Key words: amphibia, Arthur Koestler, Darwinism, epigenetics, evolution, Lamarckism Abstract Introduction The Austrian biologist Paul Kammerer (1880-1926) would by On August 7, 1926 the American herpetologist Noble now be long forgotten if Arthur Koestler had not published published in Nature evidence of fraud with scientific ‘The case of the Midwife toad’, in which he depicted Kammerer as a victim of the paradigm battle between neo-Darwinists and results of Paul Kammerer (Noble, 1926). Six weeks Lamarckists. Kammerer is still on the scientific agenda, with at later Kammerer committed suicide. Kammerer was a least 10 publications since 2005. The question is still out if biologist in Vienna, Austria. He studied toads, sala- Kammerer fabricated his scientific results or not. In this paper manders and lizards and Noble had discovered that the we provide the evidence that Kammerer consistently faked ex- front legs of his last preserved specimen of the Midwife perimental results. We show (1) that the design of his experi- toad (Alytes obstetricans (Laurenti, 1768)) had been in- ments could never have produced the results that he claimed, (2) that the assumptions he made about the developmental biol- jected with India ink to suggest the presence of nuptial ogy of the species he studied are falsified by recent research, pads. If nuptial pads really had been present, it would and (3) that the specimens he showed as proof for the success of possibly have been evidence for an acquired character his experiments came from nature. to become a hereditary trait. Midwife toads mate on land and have no nuptial pads. Kammerer claimed to have bred midwife toads in water over several genera- Content tions, and that the male descendants now developed nuptial pads in the mating season, like males of water Introduction ................................................................................... 457 breeding frogs and toads. Kammerer had been un- The Blind cave salamander ........................................................ 459 masked as a scientific fraud. Many saw his suicide as a Recent research on reproduction in the Blind cave salamander .............................................................................. 460 confession. Probably, Kammerer would by now be long Eye development ..................................................................... 460 forgotten if Arthur Koestler had not published ‘The Recent research ...................................................................... 461 case of the Midwife toad’ (Koestler, 1971). In that book The Fire salamander .................................................................... 461 he pictures Kammerer as a victim of the paradigm bat- Variation in colour patterns ................................................ 462 tle between neo-Darwinists and La marckists, and sug- The evidence ............................................................................ 463 gests that others had committed the fraud. The book Conclusion ............................................................................... 464 Viviparity in Salamandra salamandra ............................... 464 saved Kammerer not only from oblivion, but also Kammerer’s results ................................................................ 464 placed him back on the scientific agenda: since 2005 at Recent research on viviparity in S. salamandra .............. 465 least 10 publications (Gliboff, 2005, 2006, 2010; Coen, Geographical distribution of viviparity in 2006; Logan, 2007; Pennisi, 2009; Svardall, 2010; S. salamandra .......................................................................... 465 Wagner, 2009; Weisman, 2010; Liu, 2011) have ap- Developmental biology ......................................................... 466 Conclusions ............................................................................. 466 peared about him and his work, and it has even been General conclusions ..................................................................... 466 suggested that Kammerer is the founder of epigenetics Acknowledgements ...................................................................... 468 (Sermonti, 2000; Vargas, 2009). Although the events References ...................................................................................... 468 took place in the period 1903-1926, they continue to Downloaded from Brill.com09/27/2021 07:50:05PM via free access 458 Van Alphen & Arntzen – Paul Kammerer’s Lamarckism generate discussion and the question is still out if believed that inheritance of acquired characteristics Kammerer fabricated his scientific results or not. played an important role in evolution and his re- Several aspects in this case impede a clear view of search was aimed to demonstrate just that. Kam- the problem: merer presented his data as evidence that character- 1. The language in which Kammerer published. The istics acquired by exposure to experimental envi- original publications are in German, a language ronments became heritable. Lamarckism was un- that not all of the people who have participated in derstandably quite attractive to Communists, be- the discussion have mastered equally well, limiting cause of the plastic view of life that was consistent their access to literature. with the plastic view of human nature insisted upon 2. The quality of his publications. In his publications by Marxist-Leninist dogma (Zirkle, 1949, 1959). Kammerer is often unclear about the details of the The Russians even made a film inspired by the life research protocol followed, and - according to cur- and works of Kammerer (Anonymous, 1961). The rent standards - essential quantitative data and sta- film can be seen at YouTube (https://youtube/ tistical analyses are lacking. Where there is clarity 6Au0lh7BuK4) and two stills from the film are here about the procedures followed, alternative hypoth- shown as Fig. S1. Kammerer’s opponents were neo- eses are not excluded and other interpretations of Darwinians and argued fiercely against Lamarck on the results are often possible. the basis of Mendel’s and Weismann’s theories. 3. Phenotypic plasticity. Most of the experimental They demanded hard evidence from their oppo- work of Kammerer involved the breeding of am- nents for the inheritance of acquired characteristics. phibians under ambient conditions (humidity, light, Weismann discovered that the cells that will be temperature) that differ from those in the natural used for reproduction (the germ line) are already set habitat. Such experiments may reveal phenotypic apart during early development (Weismann, 1892). plasticity. In Kammerer’s time, phenotypic plasticity The other cells form the body (the soma). On the of individual organisms was widely known, but not basis of this separation Weisman reasoned that new how it related to genetics. One was not yet aware that genetic properties acquired by somatic cells cannot the expression of a gene could vary with the envi- be transmitted to the offspring. The controversy ronment and thus produce different phenotypes. also suggested that prejudice rather than scientific This also contributed to the confusion about the im- arguments played a key role in the debate on Kam- plications of his experiments. If a different pheno- merer’s claims. type occurred in a different environment some saw 6. No independent replication of Kammerer’s experi- this as an acquired characteristic. It would of course ments. A scientific solution of the disagreement still have to be shown that such an acquired charac- could have been that Kammerer’s opponents would teristic was also passed on to the next generation to have repeated his experiments, but Kammerer provide evidence for Lamarckian evolution. worked with amphibians with long generation times, 4. The role of selection. Another source of confusion some of which are difficult to breed in captivity. Re- is that the experiments did not exclude the occur- peating the experiments would have been an even rence of selection. As a rule Kammerer collected greater challenge, as this requires breeding under large numbers of animals, which he kept in terrari- conditions normally not experienced by the animals. ums. Maybe he already selected particular individ- For example, to breed midwife toads in the water is uals in the field. In addition, some mortality must an impossible task. A carrying male will routinely have occurred in his terrariums, but he did not pro- shed the egg string at slight disturbance. Many have vide data on this mortality. Then, animals were ex- tried to raise the embryos from such an abandoned posed to extreme environmental conditions. These clutch, with no success. Kammerer’s work on the must also have caused some mortality. Certainly, Sea squirt Ciona intestinalis (Linnaeus, 1767) has high mortality occurred among their offspring, and been repeated, but the results could not be duplicat- so it is likely that the specimens that finally sur- ed (Fox, 1923, 1924; Whittaker, 1975, 1985). vived represented genotypes that were more resist- 7. No preserved
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