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Rediscovery revised the cooperation of Erich and Armin von Tschermak-Seysenegg in the context of the rediscovery of Mendel s laws in

Plant Biology ISSN REVIEW ARTICLE Rediscovery revised the cooperation of Erich and Armin von Tschermak-Seysenegg in the context of the rediscovery of Mendel s laws in M. Simunek 1,
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Plant Biology ISSN REVIEW ARTICLE Rediscovery revised the cooperation of Erich and Armin von Tschermak-Seysenegg in the context of the rediscovery of Mendel s laws in M. Simunek 1, U. Hoßfeld 2 & V. Wissemann 3 1 Academy of Sciences of the Czech Republic, Centre for the History of Sciences and Humanities, Prague, Czech Republic 2 Friedrich-Schiller-University, AG Didactics of Biology (Bienenhaus), Jena, Germany 3 Justus-Liebig-University Giessen, Spezielle Botanik, Giessen, Germany Keywords Genetics; heredity; history of science; hybridisation; Mendelism. Correspondence M. Simunek, Academy of Sciences of the Czech Republic, Centre for the History of Sciences and Humanities, Puškinovo nám. 9, Prague, Czech Republic. Editor M. Van Kleunen Received: 8 December 2010; Accepted: 19 May 2011 doi: /j x ABSTRACT The rediscovery of Mendel s laws in 1900 is seen as a turning point in modern research on heredity and genetics. In the first half of the 20th century it was generally held that the rediscovery was made several times, independently, and in a parallel fashion by three European botanists (Carl Correns, Hugo de Vries and Erich von Tschermak-Seysenegg). Since the 1950s, however, serious questions have arisen concerning both the chronology and the specific conceptual contribution of the scientists involved. Not only the independence but also parallelism was analysed in the context of individual research programmes of all three of these scholars. The youngest of them, Austrian botanist Erich von Tschermak-Seysenegg, was excluded from the rank of rediscoverers. It is the aim of this paper to use new archival evidence and add important facts both to the chronology and conceptual framework of Erich von Tschermak-Seysenegg s work. An entirely new aspect is added by identifying his older brother, the physiologist Armin von Tschermak-Seysenegg ( ), as a significant spiritus movens of the events of 1900 and A selected part of their correspondence, covering the period from 13 March 1898 until 19 November 1901, is made available in transcriptions. INTRODUCTION Almost nothing about the event was simple or straightforward (...)Zirkle 1964: 67 According to a traditional view adopted and fostered by almost all textbooks of genetics, Mendel s laws were presented in the first half of annus mirabilis 1900 simultaneously and independently by three European botanists: Hugo de Vries ( ) of the Netherlands, Carl Correns ( ) of Germany and Erich von Tschermak-Seysenegg ( ; hereafter E.T.S.) of Austria Hungary (Johannsen 1923; Iltis 1924; Roberts 1929; Grant 1956; Dunn 1965; Stubbe 1965; Johansson 1979; Keller 2000). The importance of William Bateson ( ), a leading British botanist, in this process is stressed as well (Olby 1985; Falk 2009). In connection with these events, the term rediscovery is often used. Having laid the foundations of further development in the 20th century research of heredity, they continue to attract considerable interest. There are two interrelated issues that are essential for reconstructing this process. The first one concerns the independence of research and discovery (Platt 1959), that is, if through their own practical research, results were obtained that were interpreted in the same sense as Mendel but without knowing the paper of Mendel. Or, if after reading Mendel s paper, existing results could be explained on the basis of 1 This paper is published as part of the DFG Research Project Ho this paper. In relation to each protagonist, two questions arise: (i) whether they discovered the laws of J.G. Mendel on their own or used Mendel s papers to interpret previously obtained results; and (ii) whether each of the scholars made the discovery independently of each other. Especially the latter question touches upon the parallelism of the discovery, and, eo ipso, also later claims of priority. Depending on the answers and the level of analysis and singularity, events of 1900 might be seen not as a rediscovery but rather as a new discovery (Jahn 1957), delayed discovery (Zirkle 1964) or complex of multiple discoveries (Brannigan et al. 1981). Issues pertaining to independence, originality and parallelism are thus reflected in various interpretations of the rediscovery (Roberts 1929; Stomps 1954; Jahn ; Dunn 1965; Stubbe 1965; Sturtevant 1964; Zirkle 1968; Olby 1985; Bowler 1989). Moreover, given the evolving nature of experimental research programmes of the protagonists, it remains unclear what should be the extent of the period covered by the term rediscoverery (Jahn 1957: 215). Doubts regarding the general claims of the youngest of the three, E.T.S., appeared relatively early. Nonetheless, he was seen as one of the rediscoverers for a long time (Johannsen 1923; Nilsson-Ehle 1924; Roemer 1941; Stubbe 1941; Reinöhl 1950; Gasking 1959; Heinisch & Rudorf 1961). Some contemporaries even saw him as...the very first Austrian scientist after Mendel who understood the rules of heredity (Hänsel 1962: 13). It may also be relevant that E.T.S. lived until the Plant Biology ª 2011 German Botanical Society and The Royal Botanical Society of the Netherlands 1 The rediscovery of Mendel s laws, Simunek, Hoßfeld & Wissemann early 1960s. He supported the position of triple parallel (simultaneous) rediscovery already in his first papers (Tschermak- Seysenegg 1900a: 239,b: 555), and consistently promoted this view also in his later writings (Tschermak-Seysenegg 1901a: 642,c: 1029, 1908, 1928, 1956, 1958). Implying the identity of discoveries in 1865 and 1900, he even mentioned...a 35 years long interval when comparing, in 1931, the original publication of Mendel s paper and events of 1900 (Tschermak-Seysenegg 1931: 1). Very important was also his contribution to the commemoration and celebration of Mendel s personality and work, which started immediately after 1900 and took the form of, for example, re-publication of Mendel s paper in 1901 (Vol. 121 in Ostwald s Classics of the Exact Sciences Series). Last, but not least, he saw himself as a leading contributor in applying the new Mendelian knowledge in the field of plant breeding and botany (Wunderlich 1951; Harwood 1997, 2000). In the mid-1960s, however, it was shown that his original understanding of Mendel s explanations in the context of 1900 had limited validity, both in conceptual and in terminological terms (Dunn 1965: 76; Allen 1975: 48). He was dropped from the list of the rediscoverers (Stern & Sherwood 1966, 1978; Bowler 2000) and viewed rather example of a biologist who interpreted what we now regard as Mendelian phenomena within a pre-mendelian concept of heredity (Olby 1985: 114), or an example of an scientist who was influenced by...preceding generations of the older hybridizers rather than the direct influence of Mendel himself (Roberts 1929: 356). Recently (e.g., Moore 2001), it has been argued that he failed because (i) he was not able to draw any generalisations from the data he obtained; and (ii) he did not attempt to explain the results he obtained by discovery within an appropriate theoretical framework. To a large extent, existing interpretations have been based on the contemporary statements and or memoirs. In the summer of 2009, a personal collection of E.T.S. held by the Archives of the Austrian Academy of Sciences in Vienna was catalogued and opened for historical research. At the same time, a significant part of the personal possessions of Armin Tschermak-Seysenegg ( ) (hereinafter A.T.S.) 2, his older brother and a prominent physiologist, was identified 2 Armin (Eduard Gustav) von Tschermak-Seysenegg (21 September 1870 in Vienna, Austria 9 October 1952 in Bad Wiessee, Germany) studied medicine at the Universities of Vienna, Austria, and Heidelberg, Germany. After completing his studies in Vienna in 1895, he moved to Leipzig. From 1899 until 1906, he worked at the University of Halle, where he was appointed adjunct professor in He allied himself to the school of E. Hering. After 1906, he returned to Vienna where he temporarily received full professorship in physiology and medical physics at the Veterinary College (Hochschule für Tiermedizin), a school he helped to establish. In , he served as its Rector. In 1913, he came to Prague, Bohemia, to fill a vacancy at the Institute of Physiology, a venerable institution founded in the 19th century by J. E. Purkinje. After receiving full professorship, he was appointed director of this Institute. He continued to lecture in Prague at the German (Charles) University until the end of WWII in His research focused mainly on general and special physiology (physiology of sight), anatomy and neurology. He coedited the Zeitschrift für Physiologie, the Zeitschrift für Sinnesphysiologie and the Archiv für Augenheilkunde. He was a member of several scientific societies and academies across Europe. In 1936, together with Th. H. Morgan, he became a member of the Academy of Sciences of the Holy See (Pontificia Academia Scientiarum) in the Vatican. He died on 9 October 1952 in Bad Wiessee in Bavaria. and described thanks to the generosity of Dr Armin Tschermak von Both collections contain correspondence between the two brothers, in all 87 items of letters, correspondence cards, postcards and telegrams. They cover the period from 1898 until 1951 (the year before the death of A.T.S.). The vast majority of the extant correspondence was written and sent from A.T.S. to E.T.S. (83 items); only four items conversely. There are 14 pieces of correspondence sent by A.T.S. to E.T.S. in the period from 13 March 1898, until 19 November Despite some very clear statements of E.T.S. (Tschermak- Seysenegg 1958: 6) and the assumption that the two brothers were very close (Harwood 2000: 1062), the extent, quality and significance of their relationship for the early research in genetics remained, until recently, completely unknown. 3 It is, however, certain that A.T.S. supported his brother in promoting the view of the triple simultaneous rediscovery from the very beginning (Tschermak-Seysenegg A. 1901, 1923). Although the mutual correspondence is clearly incomplete, it is certain that information therein contained can help provide much more detailed answers to questions pertaining to degrees of success in interpreting Mendel s works in the crucial period of The key issues are: (i) when were Mendel s papers read for the first time; (ii) when was the phenomenon of segregation understood in Mendelian terms; (iii) what was the explanation of the mechanical base of heredity; and (iv) how did this influence the future research of heredity? And finally-and this should be seen as an entirely new aspect of the traditional rediscovery story-how relevant was the contribution of A.T.S., whose role was until now completely underestimated? ROUTES TO REDISCOVERY : THE CHRONOLOGY In the spring of 1898, E.T.S. went to the Belgian city of Ghent to study botanical techniques. At the suggestion of Dr Julius MacLeod ( ), professor of botany at the University of Ghent and director of the local botanical garden, he soon started experimenting with self- and cross-pollination. Exploring the effects of both self- and cross-fertilisation in plants, he was also inspired by some remarks of Charles Darwin (Tschermak-Seysenegg 1900a: 232, 1958: 53; Roberts 1929: 343). For hybridisation experiments, E.T.S. chose several varieties of Pisum. In the same year, he visited H. de Vries in Amsterdam, with whom he then stayed in touch by correspondence (Tschermak-Seysenegg 1958: 47 8). In the autumn of 1898, E.T.S. returned to Austria hoping to obtain a position as an assistant at the University of Agriculture in Vienna with Dr Adolf von Liebenberg ( ). At Liebenberg s advice, however, he first accepted a practical position in Eßlingen, at the farm of the Habsburg imperial family, in October There he continued with hybridisation experiments using the plant material he brought from Ghent (Roberts 1929: 346; Tschermak-Seysenegg 1958: 53). The move back to Vienna a year later, in October 1899, gave him an opportunity to analyse the results of his experiments in the F2 generation, which he needed for his inaugural dis- 3 The only two persons who went through the correspondence in the 1980s and 1990s were Armin von Tschermak Seysenegg s son, Wolfgang Tschermak von Seysenegg, and his grandson, Dr Armin Tschermak von 2 Plant Biology ª 2011 German Botanical Society and The Royal Botanical Society of the Netherlands Simunek, Hoßfeld & Wissemann The rediscovery of Mendel s laws, sertation. In Vienna, he could also use the scientific literature in Liebenberg s private library. 4 According to E.T.S. s memoirs, it was here that he first became aware of the existence of G.J. Mendel and his 1850s and 1860s research on Pisum and Hieracium. He came across it in the 1881 work of Wilhelm O. Focke (Tschermak-Seysenegg 1958: 54). 5 Then he found a copy of Verhandlungen des Naturforschenden Vereines in Brünn in the library of the University of Vienna. While finishing his thesis, he studied Mendel s papers between 10 October and Christmas 1899, that is, he read them in 2.5 months. 6 He delivered his thesis Ueber kuenstliche Kreuzung bei Pisum sativum (Concerning Artificial Crossing in Pisum sativum) on 17 January Two months, later in March 1900, he made a second set of corrections. According to E.T.S., he received H. de Vries paper Sur la loi des disjonction des hybrides (Concerning the Law of Segregation of Hybrids; published in French) of 26 March 1900, at the beginning of April 1900 (Roberts 1929: 346; Tschermak- Seysenegg 1958: 54; Sturtevant 1964: 26). Later he received another paper by H. de Vries, Das Spaltungsgesetz der Bastarde (The Law of Segregation of Hybrids; written in German), which was submitted for publication in the March volume of Berichte der Deutschen Botanischen Gesellschaft (further Berichte) (Tschermak-Seysenegg 1958: 55). At this point, E.T.S. decided to publish his own results as soon as possible. One of the few options open to him was the official journal of the Imperial Austrian Ministry of Agriculture Zeitschrift für das landwirthschaftliche Versuchswesen in Österreich (further Zeitschrift) (Tschermak-Seysenegg 1900b). While going through the second round of corrections at the beginning of May 1900, E.T.S. received a reprint of C. Correns s paper written in German, Gregor Mendel s Regel über das Verhalten der Nachkommenschaft der Bastarde (Gregor Mendel s Law Concerning the Behaviour of Progeny of Varietal Hybrids) (Correns 1900; Rheinberger 2000a). It was published in volume 18 (3) for April 1900 of the Berichte (Tschermak-Seysenegg 1958: 55). 7 Within a few days, and certainly before 16 May 1900, E.T.S. prepared a short abstract which shared its title with his thesis. This paper was submitted to the Berichte on 2 June 1900, and published in volume 18 (5) for June 1900 (Tschermak-Seysenegg 1900a). At about the same time, he sent the pre-prints of his thesis (i.e., not the abstract submitted to the Berichte!) to both H. de Vries and C. Correns (Tschermak-Seysenegg 1958: 55 6). In the course of the following year (1901), E.T.S. published three more papers that he saw as a contribution to the same topic, based on an extension of his research (Iltis 1924: 223). The first one, Weitere Beiträge über Verschiedenwerthigkeit der 4 Manuscript called Geschichtliche Darstellung der Wiederentdeckung der Mendelschen Vererbungsgesetze von Seiten Erich v. Tschermak-Wien, Archiv der Österreichischen Akademie der Wissenschaften Wien, collection of Erich von-tschermak-seysenegg. 5 Ibid. 6 Ibid. 7 The only paper of E.T.S. to be translated in full into English was Concerning artificial crossing in Pisum sativum, in: Genetics 35 (suppl. to No. 5) 1950: 42 7 (translated by Aloha Hannah). Some parts of E.T.S. papers from 1900 and 1901 are available in English translations also in H. F. Roberts (1929), Plant Hybridization before Mendel, Princeton, Princeton University Press (346 57). Merkmale bei Kreuzung von Erbsen und Bohnen (Further Contributions to the Different Valuation of Traits in Crossing Peas and Beans) from 19 January 1901, was published in the February volume of Berichte (Tschermak-Seysenegg 1901a). The second one dealt with the same issues, and was basically a longer version of the same paper. It was published under the same title in the Zeitschrift (Tschermak-Seysenegg 1901b). The third was called Ueber Züchtung neuer Getreiderassen mittelst künstlicher Kreuzung (On the Breeding of New Varieties of Cereal Through Artificial Hybridisation) with the subtitle Kritisch-historische Betrachtungen (Critical Historical Remarks). It too was published in the Zeitschrift (Tschermak- Seysenegg 1901c). TOWARD THE CONCEPT: REGULAR DIFFERENTIAL VALUATION OF THE TRAITS After 1900, E.T.S. reviewed the research he started in 1898, which included experimental hybridisation, as a repetition and continuation (Wiederholung und Weiterführung), or perhaps a contribution to the new build-up (neuer Ausbau) of Mendel s teaching (Tschermak-Seysenegg 1901c: 1029). His research focused primarily on traits in hybrids, in particular on data he obtained in F1 and F2 generation by heteromorphic xenogamy (Tschermak-Seysenegg 1901a: 37). In Mendel s work, he appreciated the analysis of traits (Merkmalsanalyse) or (...) decomposition of the whole habitus into individual traits (Tschermak-Seysenegg 1901c: 1030). He also analysed the expression of various traits including the socalled vegetative traits (vegetative Merkmale). He concluded that some traits are realized in hybrids only in an alternative form (alternierend). This he saw as a proof of different values of the traits. Based on this observation, he introduced his own terms for recessive ( undervalued unterwertig) and dominant ( overvalued überwertig) traits. He did not, however, see the valuation of traits as something generally valid but rather as something pertaining only to vegetative traits (Tschermak-Seysenegg 1901a: 51, 1901c: 1032). In relation to both the breeding praxis and a racial history he was especially interested in an age of a trait (Tschermak-Seysenegg 1901c: 1058). He even implied that the dominant traits may be older than the recessive ones (Tschermak-Seysenegg 1901c: 1060). Interestingly, Mendel s factors did not play any significant part in his early analysis, i.e., in association with a pair of determiners that segregate to individual reproductive cells as they are formed. In papers published by E.T.S. in 1900 and 1901 there is just one detailed remark on Mendel s view, namely, that he even deduced some elements to be in the special organs of the cell or parts of the plasm (Tschermak- Seysenegg 1901c: 1030). 8 In several places, the so-called carriers of the traits (Träger der Merkmale) were used in connection with the ratio 3:1 (Tschermak-Seysenegg 1901a: 39, 37, 1901b: 644). It may be deduced that at this time E.T.S. failed to appreciate the distinction between phenotypic characters and genotypic factors. Trying to explain the different valuation of traits, he also allowed for further factors such as, for example, the sex of 8 In original text: (...) ja sogar an eine Zurückführung jener Elemente auf gesonderte Zellorgane oder Plasmatheile gedacht (...). Plant Biology ª 2011 German Botanical Society and The Royal Botanical Society of the Netherlands 3 The rediscovery of Mendel s laws, Simunek, Hoßfeld & Wissemann the transmitter (Tschermak-Seysenegg 1901a: 47), the influence of the race or a combination of the races (Tschermak-Seysenegg 1901a: 48; 1901c: 1059; Tschermak-Seysenegg A. 1901: 1428). Following an older tradition, he even mentioned a strengthening (or weakening) of paternal traits in the hybrids (Tschermak-Seysenegg 1901a: 49 50; Roberts 1929: 356; Olby 1987: 416), which he suggested could be used in valuation tables (Wertigkeitstabellen) for individual traits (Tschermak-Seysenegg 1901c: 1032). In 19
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