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Did Paul Kammerer Discover Epigenetic Inheritance? No and Why

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Did Paul Kammerer Discover Epigenetic Inheritance? No and Why COMMENTARY AND PERSPECTIVE Did Paul Kammerer Discover Epigenetic Inheritance? No and Why Not SANDER GLIBOFFÃ Department of History and Philosophy of Science, Indiana University, Bloomington, Indiana ABSTRACT In a recent article in this journal, Alexander Vargas presents a new, epigenetic explanation of Paul Kammerer’s controversial midwife toad experiments, but he has constructed his model without first reading Kammerer’s original articles. A look at the articles shows that Vargas is seriously misinformed about what Kammerer did and what the results even were. His model simply cannot explain the results as they were originally reported and it cannot easily be corrected. Similarly, Vargas’ historical inferences about the Kammerer affair, Kammerer’s priority for the discovery of parent-of-origin effects, and the negative reactions of geneticists to this purported discovery, are unsupported and do not stand up to scrutiny. J. Exp. Zool. (Mol. Dev. Evol.) 314B:616–624, 2010. J. Exp. Zool. & 2010 Wiley-Liss, Inc. (Mol. Dev. Evol.) 314B:616–624, How to cite this article: Gliboff S. 2010. Did Paul Kammerer discover epigenetic inheritance? No 2010 and why not. J. Exp. Zool. (Mol. Dev. Evol.) 314B:616–624. A great deal of ink has already been spilled in connection with innocence does not hinge on the details and assumptions of this the midwife toad scandal of 1926, most if it, initially, vilifying preliminary model’’ (Vargas, 2009; p 672), and in one sense I Paul Kammerer as a Lamarckian fraud. Later, Arthur Koestler (’71) agree with him: the case for Kammerer’s innocence does not promoted an alternative view, according to which Kammerer was hinge on anything in Vargas’ model, because that case has misunderstood and maligned by a neo-Darwinian establishment already been made in various ways by other authors, without the that was closed to his ideas about the modifying power of the aid of epigenetics. But, if Vargas wants to make a meaningful environment. Now, a recent article in this journal by Alexander contribution to these lines of argument, the details, assumptions, Vargas (2009) proposes a variation on Koestler’s theme, in and above all the fit of his model to Kammerer’s reported results which Kammerer is misunderstood and maligned not for his are crucial. Lamarckism, but for his discovery of epigenetic effects. Thanks to I am also aware that the stated purpose of his article is only the current knowledge of epigenetics, the argument goes, ‘‘to illustrate how, in the context of our modern knowledge, a Kammerer’s results now gain new credibility and significance. working hypothesis for a mechanistic explanation can be I write in response to Vargas because I am astonished that attempted’’ (Vargas, 2009; p 672), rather than to check that anyone would venture to publish a new analysis of Kammerer’s hypothesis against Kammerer’s published results. I admit that the results without first consulting Kammerer’s original articles. I am article meets this low expectation. But what is the point of afraid that Vargas has given readers the false impression that he rushing to publish a hypothesis that can so easily be shown to be has explained what Kammerer actually reported. false? Just as baseless are Vargas’ historical inferences. Before I also wish to make it clear that I am talking about a particular suggesting that Kammerer deserves credit for discovering epigenetic model, not the potential of epigenetics generally to epigenetic effects, or rather stumbling upon them, one really shed light on the Kammerer case. Who knows? Maybe some other ought to check the literature for prior reports. And, before making claims about what early 20th century geneticists ÃCorrespondence to: Sander Gliboff, Indiana University, Department of understood or failed to understand about environmental effects History and Philosophy of Science, 130 Goodbody Hall, 1011 E. 3rd St., and non-Mendelian heredity, one really ought to read some of Bloomington, IN 47405. E-mail: [email protected] their work, too. Received 3 March 2010; Revised 19 June 2010; Accepted 16 August 2010 Of course, I have read Vargas’ disclaimers. I know that he Published online 17 September 2010 in Wiley Online Library (wileyonline wants it ‘‘clearly understood that the case for Kammerer’s library.com). DOI: 10.1002/jez.b.21374 & 2010 WILEY-LISS, INC. KAMMERER AND EPIGENETICS 617 epigenetic model can be contrived to fit the data. But even then, Of these environmental manipulations, two are at issue here. what would that tell us about the Kammerer affair and the In the first, Kammerer took the eggs from an untreated male, who reception of his work? Vargas suggests that an epigenetic had been brooding them normally on his hind legs, and he put explanation would make fraud unlikely, on the grounds that the eggs into the water to complete their development. This is the Kammerer would not have concocted data that support a theory treatment that Kammerer later called a ‘‘controlling test.’’ of which he had no knowledge, and that argument might have The second is the better-known one, in which he used elevated some merit. But fraud is not the only available explanation of temperatures to induce the toads to spend more and more time in Kammerer’s results. There are others with which an epigenetic the water and gradually to give up their terrestrial mating and one would have to compete. brooding behavior. The behavioral changes occurred over the For instance, there are selectionist explanations, such as the course of several breeding seasons, but within a single ones outlined in responses to Koestler by Stephen J. Gould (’72) generation. Kammerer kept the offspring of each breeding season and C.H. Waddington (’71). Or one might allow that the design separate and used them to found several lines of water-breeding and execution of the experiments was well intended, but faulty or toads. Eventually, the parental toads were all mating in the water ambiguous. Maybe the results were reported only selectively. Or and abandoning their eggs there. Simultaneously, they began to maybe the reports were perfectly honest, but nonetheless lay the smaller and more numerous ‘‘water eggs.’’ Meanwhile, in incomplete or erroneous. Why should the availability of an the control terrarium, in an unheated room, the toads maintained epigenetic explanation rule out all these possibilities? Ought we their usual terrestrial breeding habits and continued to lay the to really declare the results to be valid, just because they are larger and less numerous ‘‘land eggs’’ (Kammerer, ’06). consistent with a favored hypothesis? At this point, Kammerer sampled eight water-breeding toads Finally, there are problems with the historical picture Vargas from the warm room for backcrossing to the untreated, land- paints, of Kammerer’s Mendelian critics having been totally breeding stock (Kammerer, ’10, ’11). Vargas’ main concern, and unprepared to understand his results, and suspecting fraud rather mine, is with these backcrosses, with which Kammerer aimed to than reconsider their ideas about the modifying power of the make the case for a synthesis of Lamarck with Mendel. environment. The picture is in line with Koestler’s view of the Kammerer’s idea was that acquired characteristics could become field and his desire to make Kammerer into something of a fixed in heredity in the form of Mendelian alleles, and he wanted martyr, but it is not well grounded in historical research. to show that the newly acquired water-breeding traits would Vargas claims that ‘‘Paul Kammerer could be the true indeed behave in the Mendelian manner. The modifying effects of discoverer of non-Mendelian, epigenetic inheritance’’ (Vargas, the environment, he would argue, must have been communicated 2009; p 671), but there were several notable cases that were somehow to the chromosomes, where they created a new allele. under discussion before 1910, which suggest epigenetic effects at The second generation of toads from the warm room also least as strongly as Kammerer’s. Even if he did encounter mated in the water and laid water-type eggs. As a test for the epigenetic effects, his report is very unlikely to have been a first. inheritance of these acquired characteristics, Kammerer trans- Vargas also ignores a variety of mechanisms that the early ferred some eggs to the more normal environment of an outdoor Mendelians proposed to account for various anomalous patterns terrarium, where they, too, grew up to be water breeders of heredity. He is not justified in assuming that they were unable (Kammerer, ’09). But the geneticist Erwin Baur criticized this to deal with anything that seemed to modify the expression or test, on the grounds that the modified toads were transferred to transmission of an allele. On the contrary, they did not shy away the ‘‘normal’’ environment only after they had already been from exploring the possibilities for the environment, physiology, exposed, however briefly, to the experimental conditions. Their sex, or interactions with other genes to influence heredity. water-breeding traits, therefore, were not necessarily inherited, The following is a corrected and expanded account of but could have been induced by exposure to the warmth and the Kammerer’s results and the context in which they were first water during fertilization and at least part of the egg stage. He interpreted, as well as a critique of Vargas’ epigenetic model. thought that the warmth might even have affected the eggs while still in the parental ovary (Baur, ’14; p 55–58). In response, Kammerer later reconceptualized and renamed the other experi- EXPERIMENTS IN BRIEF ment as his ‘‘controlling test’’ to use as an answer to Baur. The Kammerer’s long-running midwife toad project began by gist of his new interpretation was that it showed that exposure to demonstrating the power of the environment to modify the water at the egg stage was not the crucial modifying factor life-history and breeding behavior of the animals.
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