Abstract
In addition to his experiments on selection in pure lines, Wilhelm Johannsen (1857–1927) performed less well-known hybridisation experiments with beans. This article describes these experiments and discusses Johannsen’s motivations and interpretations, in the context of developments in early genetics. I will show that Johannsen first presented the hybridisation experiments as an additional control for his selection experiments. The latter were dedicated to investigating heredity with respect to debates concerning the significance of natural selection of continuous variation for evolution. In the course of the establishment of a Mendelian research program after 1900, the study of heredity gained increasing independence from questions of evolution, and focused more on the modes and mechanisms of heredity. Further to their role as control experiments, Johannsen also saw his hybridisation experiments as contributing to the Mendelian program, by extending the scope of the principles of Mendelian inheritance to quantitative characters. Towards the end of the first decade of genetics, Johannsen revisited his experiments to illustrate the many–many relationship between genes and characters, at a time when that relationship appeared increasingly complex, and the unit-character concept, accordingly, became inadequate. For the philosophy of science, the example shows that experiments can have multiple roles in a research programme, and can be interpreted in the light of questions other than those that motivated the experiments in the first place.
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Notes
The impact of the suggestion is best illustrated by the fact that the proceedings of the conference were published under the name Report of the Third International Conference on Genetics (on the importance of “baptising” for discipline formation, see Powell et al. 2007).
The perceived significance of Johannsen’s pure-line experiments is illustrated by the fact that Johannsen was nominated for the Nobel Prize for his experimental work in 1920 and 1923 (see Nielsen and Nielsen 2001, and the respective entries in the Nomination Database—Physiology or Medicine, Nobel Media (2014)). Johannsen introduced the terms gene, genotype, phenotype (see Johannsen 1909 and 1911, as well as Cook 1912 for contemporary discussion, and Winge 1958 and Wanscher 1975 for historiographic accounts, emphasising the terminological innovations). Furthermore, it was at Johannsen’s suggestion that Bateson’s term allelomorph was shortened to allele (see Johannsen 1923, p. 139 and 1926, p. 435, and the discussion in Sect. 3.2 of the present article).
The neglect is all the more remarkable given that the 1907 publication is one of the few English language articles published by Johannsen, besides the influential pieces from 1911 and 1923. For translations of the 1903 monograph, see Gall and Putschar (1955) providing the introduction, the first series of experiments (on the size qua weight of bean seeds) and a summary, as well as Peters (1959), containing only the summary chapter.
An earlier and shorter Danish version of Johannsen’s book appeared in (1905). The third edition (Johannsen 1926) is not analysed in detail, because the present article focuses on a shift in Johannsen’s thought (reflected in a shift in genetics generally) in the first decade after the rediscovery of Mendel’s results.
These period boundaries are often met in historiographic accounts of genetics, where the rediscovery of Mendel’s work obviously marks a beginning. Johannsen’s conceptual clarifications in 1909 draw this period to a close, while Morgan’s discovery of the white eye mutant marks the beginning of another period (see e.g. Dunn 1965; Carlson 1966).
In the first edition of the Elemente, Johannsen reported results from 1902 until 1907, i.e. six generations for several lines (Johannsen 1909, lecture 10). In the second edition he presented the results of further selection experiments spanning from 1906 to 1910, with the exception of 1908 where for reasons of lack of space no experiments were conducted. In this further series of experiments he met another objection occasionally put forward, namely that not a single bean, but the whole plant (the mean of all seeds of one plant) should be the starting point for selection. Johannsen found that also under this assumption selection has no effect (Johannsen 1913, pp. 179–181).
Johannsen traced the view that hybridisation increases fluctuating variability back to Weismann (Johannsen 1913, p. 551). He also made an interesting remark concerning the prejudices that are inscribed in different traditions that meet in the discourse on general questions such as that of heredity, most notably the traditions of zoology and botany. Zoologists, Johannsen observed, tend to think of self-fertilization as an abnormality, not being aware of its commonality in the realm of plants. Therefore, a zoologist like Plate can think that the non-effectiveness of selection in Johannsen’s pure lines is an experimental by-product of extensive inbreeding (Johannsen 1907, p. 106).
There are also crosses of bean lines with long and short seeds, respectively, mentioned on page 122 and 372, the latter referencing back to page 122. However, since p. 372 mentions a different number of seeds in the F2 generation, it seems to be yet another cross.
The quantitative/qualitative distinction is relative. Colour can, of course, also appear as a quantitatively characterised property, but in most cases discussed in early genetics, it is treated as a qualitative character, because the alternatives are clearly separated. Likewise dimensions such as length can be treated as qualitative alternatives, if the gap between the variants is large enough to exclude transgressing fluctuation, such as in Mendel’s ([1866] 1902) cross of tall and short pea plants.
Nilsson-Ehle’s results did not feature in the first edition of the Elemente, because they were published in the years from 1908 to 1913 (on Nilsson-Ehle, see Müller-Wille 2005). Tine Tammes (1871–1947) simultaneously produced similar results supporting a multi-factorial view (Stamhuis 1995). Shortly afterwards, though apparently independently, Edward Murray East (1879–1938) and Charles Davenport (1866–1944) also came to similar conclusions (Mayr 1982, p. 791).
Johannsen mentioned the F2 and F3 generations of the cross in question in 1909, but reported there that the material was not yet sufficiently “worked through” (Johannsen 1909, p. 327). In 1909, Johannsen still thought that impure segregation of genes was possible, and stated that he probably had a case at hand himself (ibid., p. 412). Cases of cumulative factors of same effect and transgressive segregation must have puzzled him, and it may have been a willingness to consider impure segregation that prevented him from analysing the material in a purely Mendelian fashion, and thus from coming to the kind of interpretations that were then presented by Nilsson-Ehle. In the second edition of the Elemente, Johannsen called the idea of impure segregation a childhood disease of Mendelism, and referred to Nilsson-Ehle’s interpretation as a paradigm for solving phenomena that suggested impure segregation (Johannsen 1913, p. 613). The point here is not that Johannsen was close to making these discoveries attributed to by Nilsson-Ehle. The point is rather, that the phenomena in question appeared so many times in the differential reproduction of the Mendelian experimental system, and that, combined with the corresponding practices of interpretation, they would have inevitably been resolved sooner or later by any of the early Mendelians (see also the simultaneous resolution of such cases by Tammes, East and Davenport mentioned in footnote 11).
Johannsen referred to work of Edith Rebecca Saunders (1865–1945), a collaborator of Bateson (see also Richmond 2001).
In every edition of Elemente Johannsen complained that the study of heredity became a servant to the theory of descent (see e.g. Johannsen 1909, p. 4). In the last edition he clearly stated that any discussion of the evolutionary implications of results from the study of heredity was tantamount to taking an excursion into another field. Johannsen claimed that he avoided such excursions in his lectures in order to establish the independence of a theory of heredity from evolutionary speculation (Johannsen 1926, pp. 248–249).
This seems to be true also for other early geneticists concerned with quantitative characters, such as Nilsson-Ehle or East.
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Acknowledgments
Research for the present article was conducted while I was as a fellow in the research group “Twentieth Century Histories of Knowledge About Human Variation” (Veronika Lipphardt), at the Max Planck Institute for the History of Science in Berlin, January–June 2012. The article was in large part written while I was as a fellow at the Institute for Cultural Inquiry, Berlin, September 2012–July 2014. The article benefited strongly from many conversations on the topic with Staffan Müller-Wille. I would also like to thank the anonymous reviewers for helpful criticism. Georg, thanks for trying.
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Meunier, R. The many lives of experiments: Wilhelm Johannsen, selection, hybridization, and the complex relations of genes and characters. HPLS 38, 42–64 (2016). https://doi.org/10.1007/s40656-015-0093-7
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DOI: https://doi.org/10.1007/s40656-015-0093-7