From linnaean species to mendelian factors: Elements of Hybridism, 1751-1870

Annals of Science

Volumn 64, Issue 2, 2007, Pages 171-215

  Muller Wille S ^a   Orel, V ^b  

^a UNIVERSITY OF EXETER   (United Kingdom)

^b Mendelianum Brno ^*  (Czech Republic)

Author keywords

[No Author keywords available]

Indexed keywords

BIODIVERSITY; GENETIC ENGINEERING; RESEARCH AND DEVELOPMENT MANAGEMENT;

HYBRIDISTS; RECLASSIFICATIONS;

LIVING SYSTEMS STUDIES;

EID: 33947672977     PISSN: 00033790     EISSN: 1464505X     Source Type: Journal    
DOI: 10.1080/00033790601111567     Document Type: Article

References (197)

1. Michel Foucault, L'ordre de discours. Leçon inaugurale au collège de France prononcée le 2 décembre 1970 (Paris, 1971), 35-37;
2. see also Georges Canguilhem, 'L'objet de l'histoire des sciences', In Études d'histoire et de philosophie des sciences concernant les vivants et la vie, 7th edition (Paris, 1994; originally published 1968), 9-23.
3. On the various interpretations that historians of science and scientists have provided on Mendel, see Jan Sapp, 'The Nine Lives of Gregor Mendel', MendelWeb (1990). Available online at: http:// www.mendelweb.org/MWsapp. html (accessed 28 September 2006).
4. Vitezslav Orel, Gregor Mendel: The First Geneticist (Oxford, 1996). On Mendel and contemporary biology,
5. see Margaret Campbell, 'Mendel's Theory: Its Context and Plausibility', Centaurus 26 (1983), 38-69,
6. Sander Gliboff, 'Gregor Mendel and the Laws of Evolution', History of Science 37 (1999), pp. 217-35,
7. and Roger Wood and Vitezslav Orel, 'Scientific Breeding in Central Europe During the Early Nineteenth Century: Background to Mendel's Later Work', Journal of the History of Biology 38 (2005), 239-72.
8. Robert C. Olby, Origins of Menclelism, 2nd edition (Chicago, 1985);
9. Vitezslav Orel, 'Constant Hybrids in Mendel's Research', History and Philosophy of the Life Sciences 20 (1998), 291-99.
10. François Jacob, La logique du vivant (Paris, 1970), 201-4. Carl Correns emphasized that Mendel's choice of pea varieties was especially at odds with Carl Nägeli's idea that such varieties were not viable in nature;
11. see Carl Correns, 'Gregor Mendels Briefe an Carl Nägeli 1866-1873. Ein Nachtrag zu den veröffentlichten Bastardisierungsversuchen Mendels', in Gesammelte Abhandlungen zur Vererbungs-wissenschaft aus periodischen Schriften 1899-1924 (Berlin, 1924, originally published 1905), 1233-97 (1234).
12. See also Bentley Glass, 'Heredity and Variation in the Eighteenth Century Concept of the Species', in Forerunners of Darwin, 1745-1859, edited by Bentley Glass, Oswei Temkin, and William L. Strauss (Baltimore, MD, 1959), 144-72;
13. Hans Stubbe, Kurze Geschichte der Genetik bis zur Wiederentdeckung der Vererbungsregeln Gregor Mendels, Genetik: Grundlagen, Ergebnisse und Probleme in Einzeldarstellungen 1 (Jena, 1965), 108;
14. Leslie C. Dunn, A Short History of Genetics: The Development of Some of the Main Lines of Thought: 1864-1939 (New York, 1965), 18 (quoting Correns);
15. Peter J. Bowler, The Mendelian Revolution; The Emergence of Hereditarian Concepts in Modern Science ana Society (Baltimore, MD, 1989), 99;
16. Robert C. Olby, 'Mendel, Mendelism, and Genetics', MendelWeb (1990). Available online at: http://www.mendelweb.org/MWolby.html (accessed 23 October 2006).
17. Gregor Mendel, 'Experiments in Plant Hybridisation', in Gregor Mendel. Experiments in Plant Hybridisation, with commentary and assessment by the late Ronald A. Fisher, edited by James H. Bennett (Edinburgh, 1965), 7-51 (10). This edition of Mendel's 1866 paper is a reprint of the translation of C. T. Druery, commissioned by the Royal Horticultural Society of Great Britain, and published in its Journal in 1901.
18. The president of the Society at the time was William Bateson, and he reprinted the Druery's translation in his Mendel's Principles of Heredity: A Defense (Cambridge, MA, 1902), pp. 40-95. We will therefore refer to this translation as Bateson's in the following.
19. Augustine Brannigan, The Reification of Mendel', Social Studies of Science 9 (1979), 423-54 (424);
20. Robert C. Olby, 'Mendel no Mendelian?', History of Science 17 (1979), 53-72 (67);
21. L.A. Callender, 'Gregor Mendel: An Opponent of Descent with Modification', History of Science 26 (1988), 41-75 (72);
22. Floyd V. Monaghan and Alain F. Corcos, The Real Objective of Mendel's Paper', Biology and Philosophy 5 (1990), 267-92 (268).
23. For a critical discussion of these papers, see Vitezslav Orel and Daniel L. Hartl, 'Controversies in the Interpretation of Mendels Discovery', History and Philosophy of Life Sciences 16 (1994), 436-55.
24. Olby (note 7), 67.
25. Mendel (note 6), 18; cf. Gregor Mendel, 'Versuche über Pflanzen-Hybriden', Verhandlungen des Naturforschenden Vereines in Brunn 4 (1865, published 1866), 3-47 (14). For explanations, see note 142 below.
26. On this distinction, see Staffan Müller-Wille and Hans-Jörg Rheinberger, 'Heredity: The Production of an Epistemic Space', in Heredity Produced; At the Crossroad of Biology, Politics, and Culture, 1500-1870, edited by Staffan Müller-Wille and Hans-Jörg Rheinberger (Cambridge, MA, in press), 3-34 (13).
27. Mendel (note 6), 10.
28. James L. Larson, Interpreting Nature: The Science of Living Form from Linnaeus to Kant (Baltimore, MD, 1996), 61-65.
29. Carl Linnaeus, Generaplantarum eorumque characteres naturates [...] (Leyden, 1737), 'Ratio opens', par. 5 (unpaginated). The criterion of constancy had already been used by John Ray (1627-1605) in 1686 to distinguish specific differences from individual variations;
30. see John Ray, Historia plantarum species hactenus editas aliasque insuper multas noviter inventas & descriptas cotnplectens [. . .], 3 vols (London, 1686), I, 40.
31. Ernst Mayr's claim, that Mendel 'had little idea what a species was' must be seen against the background that for Mayr there was one true species concept only: his own biological species concept; see Ernst Mayr, The Growth of Biological Thought: Diversity, Evolution and Inheritance (Cambridge, MA, 1982), 712-13.
32. See Wood and Orel (note 3) for a recent overview over the discussions in the Sheep Breeders' Association and how it influenced Mendel.
33. Olby (note 4), 205-7.
34. Gliboff (note 3). 219.
35. Mendel (note 6), 10.
36. Alongside Art, Mendel (note 9) (36-37, 39) also used the expression Species to refer to what he otherwise called 'good species' (gute Arten), that is to plant kinds distinguished from each other by a whole set of traits.
37. This is especially palpable in the translation edited by Curt Stern and Eva R. Sherwood, where 'Art' is almost consistently rendered as 'variety', 'strain' etc., unless it is clear from the context that Mendel was indeed intending 'good' species; see Gregor Mendel, 'Experiments on Plant Hybrids', in The Origin of Genetics: A Mendel Source Book, edited by Curt Stern and Eva R. Sherwood (San Francisco, 1966). We are therefore using, with one exception only (note 94), R. A. Fisher's re-edition of Bateson's 1901 translation, which is truer to the original in this respect.
38. Carl Linnaeus, 'Disquisitio de sexu plantarum', in Caroli Linnaei Ammoenitates academicae, seu Dissertationen varias Physicae, Mediate, Botanicae antehac seorsim editae, 2nd edition, edited by Johann Christian Daniel Schreber, 10 vols (Erlangen, 1788-1790), X (1790, essay originally published 1760), 100-31 (126-27);
39. for a detailed reconstruction of this experiment, see Staffan Müller-Wille, '"Varietäten auf ihre Arten zurückfü hren": Zu Carl von Linnés Stellung in der Vorgeschichte der Genetik', Theory in Biosciences 117 (1998), 346-76.
40. Olby mentions that Linnaeus produced a second hybrid, Veronica maritima × Verbena officinalis, by manual cross-fertilization, and calls this combination 'extremely unlikely' (Olby 1985 [note 4], 4). The source he offers in support for this, a passage translated from Linnaeus' essay Generatio ambigena (1759), does not provide evidence, however, that this cross was indeed artificially produced (ibid., 142). As a matter of fact, in his first report on the Veronica × Verbena hybrid, Linnaeus only mentions that he had 'picked it up (lecta)' not far from the garden beds where the two parent species grew; see Carl Linnaeus, 'Plantae Hybridae', in Caroli Linnaei Ammoenitates academicae, seu Dissertationes variae Physicae, Medicae, Botanicae antehac seorsim editae, 1 volumes (Stockholm, 1749-1769), III (1764, essay originally published 1751), 28-62 (35).
41. Linnaeus (1751) 1764 (note 21), 32-33.
42. Linnaeus (1751) 1764 (note 21), 30.
43. Carl Linnaeus, 'Metamorphoses plantarum', in Caroli Linnaei Amoenitates academicae, seu Dissertationes varias Physicae, Medicae, Botunicae antehac seorsim editas, 2nd edition, edited by Johann Christian Daniel Schreber, 10 volumes (Erlangen, 1785-1789), VI (1788, essay originally published 1755), 367-86 (380).
44. For instance, in the 'bigeneric' case of Veronica spuria (speedwell), allegedly a hybrid of V. maritima and Verbena offlcinalis, and a few others; see Linnaeus (note 21), nos 1, 3, 6, 4, 18, 23, 29.
45. Ibid., 50.
46. For a more detailed account of Linnaeus's classification of hybrids see Müller-Wille (note 21), 366-68.
47. Joseph Gottlieb Koelreuter, Dritte Fortsetzung der Vorläufigen Nachricht von einigen das Geschlecht der Pflanzen betreffenden Versuchen und Beobachtungen (Leipzig, 1766), 37; see also Olby 1985 (note 4), 4.
48. Conway Zirkle, The Beginnings of Plant Hybridization, Morris Arboretum Monographs, 1 (Philadelphia, 1935), 7-41.
49. For a more detailed analysis see Staffan Müller-Wille, 'Nature as a marketplace: The political economy of Linnaean botany', in Oeconomies in the Age of Newton, edited by Neu De Marchi and Margaret Schabas, History of Political Economy Supplement 35 (Durham, NC, 2003), 154-72.
50. Joseph Gottlieb Koelreuter, Vorläufige Nachricht von einigen das Geschlecht der Pflanzen betreffenden Versuchen und Beobachtungen (Leipzig, 1761), 38.
51. Koelreuter must have heard of Linnaeus hybids much earlier already. He had studied natural history with Johan Georg Gmelin at the University of Tübingen, and Gmelin was a close correspondent of Linnaeus, and had devoted his inaugural address in 1749 to the topic of hybridization; see Ernst Mayr, 'Joseph Gottlieb Koelreuter's contributions to biology', Osiris, 2nd Series, 2 (1986), 135-76 (136-37). Gmelin, on the other hand, had been inspired to write his essay by Linnaeus, who in 1744 had reported peloric mutations from a wild population of Linaria and had speculated about their hybrid origin. As Linnaeus was unable to reproduce these plants in his garden, they did not play a role in his later writings on hybridization, until 1760, when he finally succeeded in propagating them asexually; see Müller-Wille (note 21) on this.
52. Olby (note 4), 9; according to Mayr (note 32), 135, Koelreuters experiments started in 1760 only.
53. Koelreuter (note 31), 41-42. In his prize essay of 1760, Linnaeus maintained that he had 'abraded the pollen [of the seed plant] by hand' (Linnaeus [1760] 1790 [note 21], 126), while the account from the essay Generatio ambigena, published a year earlier, reported that Linnaeus 'strongly blew into' the flowers of the seed plant to remove the pollen; see Carl Linnaeus, 'Generatio ambigena', in Caroli Linnaei Amoenitates academicae, seu Dissertationen varias Physicae, Medicae, Botanicae antehuc seorsim editas, 7 volumes (Stockholm, 1749-1769), vol. VI (1763, essay originally published 1759), 1-16 (12). If this is true - which is likely, as the essay was composed as a dissertation pro exerdtio by Linnaeus' student Christian Ludwig Ramström who probably witnessed the act - it would have been a ludicrously unreliable method to avoid self-pollination. It is also probable, however, that Koelreuter's suspicion was based on the fact that the plants growing from the seed sent by Linnaeus segregated.
54. Joseph Gottlieb Koelreuter Fortsetzung der Vorläufigen Nachricht von einigen das Geschlecht der Pflanzen betreffenden Versuchen und Beobachtungen (Leipzig, 1763), 9.
55. Cf. Olby"(note 4), 10-18.
56. Koelreuter (note 35), 60.
57. Koelreuter (note 28), 37.
58. Koelreuter (note 35), 47-48.
59. Ibid., 58.
60. The term Gattung designated species in eighteenth-century German, other than in modern German, where it designates genus. For 'genus', Geschlecht was used, while Art, corresponding to 'species' in modern German, was used to refer to kinds in general; see Christoph Girtanner, Über das kantische Prinzip für die Naturgeschichte (Göttingen, 1796), 6-10.
61. Koelreuter (note 31), 44; 'some' in all probability indicates Linnaeus.
62. Koelreuter (note 28), 35.
63. Koelreuter (note 35), 46; (note 28), 35.
64. Koelreuter (note 35), 44.
65. Koelreuter (note 28), 85-86; translation partly based on Mayr (note 32), 170.
66. Mayr (note 32), 170.
67. See ibid., 154-72, and Larson (note 12), 71-76, for detailed accounts of Koelreuter's 'alchemical' theory of inheritance.
68. Koelreuter employed the same strategy when dealing with an experimental result that formed the inverse of that just discussed. In crossing two kinds of Cucurbita (cucumber), which he had expected to behave like 'natural' species and thus to prove sterile, Koelreuter acquired perfectly fertile offspring. He accounted for this by declaring that the parental forms were 'essentially as little different as a lap-dog and an English mastiff'; see Koelreuter (note 28), 119.
69. Koelreuter (note 35), 62.
70. Olby (note 4), 25.
71. Zirkle (note 29), ch. 3.
72. Olby (note 4), 23-26.
73. For overviews, see Herbert F. Roberts, Plant Hybridization before Mendel (Princeton, NJ, 1929); Conway Zirkle, Gregor Mendel and his predecessors, Isis 42 (1951), 97-104; Stubbe (note 5); and Larson (note 12), ch. 3. It should be emphasized, that a large amount of observations on hybridizations from the late eighteenth and nineteenth century, in continuing the Linnaean tradition, was probably made in taxonomic and biogeographical literature.
74. Augustin Sageret, 1826, 'Considérations sur la production des hybrides, des variantes et des variétés en général, et sur celles des Cucurbitacées en particulier', Annales des Sciences Naturelles, 1. Ser., 8 (1826), 294-314 (300-302).
75. William Herbert, Amaryllidaceae: Preceded by an Attempt to Arrange the Monocotyledonous Orders, and Followed by a Treatise on Cross Bred Vegetables (London, 1837), 17;
76. on Herbert, see Abigail Lustig, 'Cultivating Knowledge in Nineteenth-Century English Gardens', Science in Context 13 (2000), 155-81.
77. Carl Friedrich Gärtner, Versuche und Beobachtungen über die Bastarderzeugung im Pflanzenreich (Stuttgart, 1849), x.
78. Ibid., 151.
79. Ibid., 163; translation partly based on Olby (note 4), 33.
80. Gärtner (note 57), 166.
81. Ibid, 168.
82. Ibid, 186.
83. ibid.,, 502-17.
84. ibid.,, 518.
85. ibid.,, 405.
86. ibid.,, 539.
87. ibid.,, 577.
88. ibid.,, 576-77.
89. ibid.,, 578-79.
90. See Olby (note 4), 26-31.
91. Gärtner (note 57), 581.
92. This was also the research programme of Charles Naudin (1815-1899) and Max Ernst Wichura (1817-1866); see Olby (note 4).
93. Larson (note 12), 97.
94. Gärtner (note 57), 234-35.
95. Olby (note 4), 33.
96. Gärtner (note 57), 254.
97. Olby (note 7), 67.
98. Callender (note 7), 41; with varying emphasis, these verdicts have been restated more recently by Monaghan and Corcos (note 7) and B.E. Bishop, 'Mendel's Opposition to Evolution and to Darwin'. Journal of Heredity 87 (1996), 205-13.
99. Mendel (note 6), 7; translation slightly altered. The term 'Entwicklung' in early nineteenth-century German covered both ontogeny and phylogeny; see Gliboff 1999 (note 3), pp. 225-26, who suggests to translate it as evolution, in line with English usage at Mendel's time.
100. In contrast to what Monaghan and Corcos (note 7), 271, have claimed.
101. Gärtner (note 57), 428.
102. Ibid., 446.
103. It should be noted, however, that this step was made possible in the context of a large-scale change in the notion of 'generation', which included Gärtner, and comprehended early-nineteenth century biological and social sciences in their entirety. While generation before the nineteenth century signified the procreation of individuals, it came to acquire the subsidary meaning of a population of individuals living at about the same time; see Ohad Fames, 'On the shoulders of generations: The new epistemology of heredity in the nineteenth century', in Heredity Produced: At the Crossroad of Biology, Politics, and Culture, 1500-1870, edited by Staffan Müller-Wille and Hans-Jörg Rheinberger (Cambridge, MA, in press).
104. 2 today), resulting from the selfing of hybrids, corresponds to Gärtner's 'mediated hybrids'. And Mendel's backcrosses would fall under Gärtner's 'mixed' and 'combined' hybrids, respectively.
105. Gärtner himself pointed this out explicitly; see Gärtner (note 57), 516-17.
106. Mendel (note 6), 8; translation slightly altered. The term 'Probe' (Mendel [note 9], 4) is translated misleadingly in both English translations: as 'results' in Bateson's translation (Mendel [note 6], 8) and as 'attempt' in Stern's and Sherwood's translation (Mendel [note 20], 2).
107. Mendel (note 6), 10.
108. Mendel (note 6), 11; translation slightly altered.
109. Again, this contrasts with what Monaghan and Corcos 1990 (note 7), 271, have claimed with respect to the quoted passage; cf. Raphael Falk and Sahotra Sarkar, 'The Real Objective of Mendel's Paper: a Response to Monaghan and Corcos'. Biology and Philosophy 6 (1991), 447-51 (449).
110. See especially the discussion of the dihybrid series in Mendel (note 9).
111. Mendel (note 6), 21.
112. Ibid., 19; Mendel's emphasis.
113. Correns (note 5), 1238. In a similar vein, Mendel emphasized 'that as an empirical worker I cannot understand anything else by becoming constant as the retention of characters during the period of observation' (ibid., 1242). Falk and Sarkar (note 88), 449, observe that Mendel neglected properties that are typical of hybrids, namely luxuriance and variegation (heterosis), although he noticed them. In a speculative reconstruction of Mendel's experiments, in the main undertaken to explain why Mendel's results appeared to be 'too good to be true' to a number of modern geneticists - a claim going back to Ronald A. Fisher, 'Has Mendel's Work Been Rediscovered?' Annals of Science 1 (1936), 115-37 - Frederico di Trocchio put forward the hypothesis, that Mendel made his monohybrid crosses 'just on paper', extracting the data from experiments that actually involved polyhybrid crosses throughout;
114. see Frederico di Trocchio, 'Mendel's Experiments: A Reinterpretation', Journal of the History of Biology 24 (1991), 485-519. This procedure would have presupposed that Mendel systematically focused on character pairs and neglected all other peculiarities that the individual hybrid plants may or may not have exhibited.
115. Mendel (note 6), 22-23.
116. Mendel (note 20), 15; we have chosen the translation offered by Curt Stern and Eva R. Sherwood in this instance, because William Bateson's translation corrupts the German original. It supposes that it is the seeds that possess 'one or other of the differing characters' (Mendel [note 6], 21), while the German original clearly speaks of 'Hybriden je zweier differierender Merkmale', that is, in literal translation, 'hybrids of two differing characters severally'.
117. Callender (note 7), 52. In fact, Callender maintained that Mendel distinguished two laws, '(a) "the law of simple combination of characters", and (b) "the law of combination of different characters'", the first presumably applying to monohybrid, the latter to polyhybrid crosses. However, Mendel mentions 'the law of simple combination of characters (Gesetze der einfachen Combiniermg der Merkmale)' with reference to an example of a polyhybrid cross (Mendel [note 9], 32). And the 'law of combination of different characters' is a mistranslation in the Bateson translation (on which Callender relied) for 'law of combination of differing (differierenden) characters' (Mendel [note 6], 37; cf. Mendel [note 9], 32; Mendel [note 20], 32, provides a correct translation). All in all, Mendel used the term 'law (Gesetz)' 15 times in his paper, in one instance qualifying it as a 'law of development (Entwicklungs-Gesetz)' (Mendel [note 9], 32). ...
118. See Gliboff (note 3), pp. 225-28, on Mendel's notion of a 'law of development'.
119. Cf. Monaghan and Corcos (note 7); Orel and Hartl (note 7).
120. Mendel (note 6), 21-22. Curt Stern considered this as the first instance in which 'a problem not only of genetics of individuals and their progeny, but also of a whole population' was attacked; see Curt Stern, 'Mendel and Human Genetics', in G Mendel Memorial Symposium 1865-1965: Proceedings of a Symposium Held in Brno in August 4-7 1965, edited by M. Sosna (Prague, 1965), 199-218 (204).
121. Mendel (note 6), 22-23 (translation slightly altered). We use the technical terms 'monohybrid' and 'polyhybrid' for the sake of convenience. They were not used by Mendel who referred to mono- and polyhbrid crosses in descriptive terms like the ones contained in the quote.
122. Cf. Orel (note 3), 162-63.
123. Cf. Falk and Sarkar (note 88), 450.
124. Mendel (note 6), 27; translation slightly altered.
125. This is also the gist of Ivan Fedorovitch Schmalhausen's (1849-1894) summary of Mendel's findings in the formers 1874 dissertation on plant hybrids; see Vitezslav Orel, 'Response to Mendel's Pisum Experiments in Brno since 1865', Folia Mendelianu 8 (1973), 199-209. An excerpt from Schmalhausens dissertation is translated in Olby (note 4), 225-26.
126. Mendel (note 6), 27-28; Mendel's emphasis.
127. Ibid., 29 (translation slightly altered).
128. Bateson ([note 6], 67) added a footnote to the quoted paragraphs stating: 'This and the preceding paragraph contain the essence of the Mendelian principles of heredity'.
129. 1) already.
130. This does not mean that egg and pollen cells must be identical; they are 'of like kind' in so far as their union, all else being equal, invariably brings about the same kind of organism.
131. Cf. Olby (note 7), 67.
132. Cf. J. Heimans, 'Mendel's Ideas on the Nature of Hereditary Characters', Folia Mendetiana 6 (1971), 91-98 (97); Olby (note 7), 58; Orel (note 3), 171-72.
133. Mendel (note 6), 31; translation slightly altered. Bateson translation has 'proving' for 'prüfen', which is simply wrong.
134. 1 do produce 'constant' forms in F2, and hybridity is therefore not a property that is inherited.
135. Callender ([note 7], 53) has maintained that Mendel distorted the views of Koelreuter and Gärtner. While Mendel ascribed to them the statement that 'hybrids are inclined to revert to the parental forms', Gärtner and Koelreuter 'stated categorically that Reversion [sic] was both necessary and inevitable'. Mendel was, however, quoting Gärtner and Koelreuter correctly; see, e.g., Joseph Gottlieb Koelreuter, Zweyte Fortsetzung der Vorläufigen Nachricht von einigen das Geschlecht der Pflanzen betreffenden Versuchen und Beobachtungen (Leipzig, 1764), 82; Gärtner (note 57), 446. The emphasis in speaking about a tendency or inclination to revert was not on denying the necessity of that process, but on stressing its gradual nature, dependent as it were, according to Koelreuter and Gärtner, on the interaction of reproductive forces. We will come back to this point later on.
136. Mendel (note 6), 35; Mendel's emphasis.
137. Cf. Heimans (note 109); Olby (note 7), 59-62; Orel (note 3), 177.
138. Orel and Hartl (note 7), 435-43.
139. Mendel (note 6), 43.
140. In fact, such semi-quantitative statements abounded in the hybridization literature before Mendel; see Roberts (note 54) and Stubbe (note 5) for useful overviews.
141. Mendel (note 6), 43.
142. Ibid., 43; Mendel's emphasis.
143. Ibid, 44-45.
144. As pointed out above already, the term for 'composition' used by Mendel in the German original - Beschaffenheit - is neutral about what it is exactly that determines this or that 'composition'. One should not conclude, therefore, that Mendel thought in terms of particulate inheritance. The reproductive cells might differ by overall qualities rather than constituents, for example. This is as true for them as it is for the plants that grow from them.
145. Ibid., 48; cf. Gärtner ([note 57], 463) for the original quote.
146. Gärtner (note 57), 251; for a translation see Olby (note 4), 157.
147. Gärtner (note 57), 458.
148. Mendel (note 6), 49; translation slightly altered.
149. Cf. Gärtner (note 57), 14-15, 249-50.
150. Mendel (note 6), 51. Mendel mentions 'Aquilegia atropupureu and cunadensis, Diunthus Caryophyllus, chinensis andjaponicus, Ntcotiana rustica smdpaniculata'.
151. Mendel (note 6). 41-42; Mendel's emphases.
152. Ibid., 25.
153. In 1886, C. W. Eichung, a horticulturalist, visited Mendel and asked him how peas growing in the latter's garden had been 'reshaped in height as well as in type of fruit'. Mendel simply answered: 'It is just a little trick, but there is a long story connected with it which it would take too long to tell' (quoted according to Olby [note 4], 90).
154. Cf. Callender (note 7), 54-55. To conclude from that, as Callender does (ibid., 72), that Mendel 'stated clearly that he accepted the general fixity of species' goes to far, however.
155. Mendel (note 6), 45.
156. Ihid., 45; Mendel's emphasis.
157. Ibid., 45. Callender ([note 7], 57) maintained that Mendel was again misrepresenting Gärtner's views here, because the latter added a remark to his list of 'constant' hybrids according to which 'there was always a steady loss of fertility and a general breaking up of the species' (cf. Gärtner [note 57], 421-22). Callender was wrong, however, to ascribe to Gärtner the general view that 'species hybrids were invariably sterile'. We demonstrated above that Gärtner was cautious to make such far-reaching empirical claims, and Mendel was therefore right to take the list as Gärtner announced it, namely as a list of 'exceptionally fertile hybrid plants propagatpng] themselves with no change of type, like pure species'. The German word that Callender translates with 'steady', moreover, is 'allmählig', meaning 'slowly' or 'gradually', rather than steady.
158. Mendel (note 6), 47.
159. Ibid., 45-46. Mendel had become acquainted with contemporary cell theory through the teaching of Franz Unger, who relied on the work of Matthias Jacob Schleiden (1804-1881) and Nägeli; see Vitezslav Orel, 'The teachings of J. M. Schleiden and its possible influence on G. Mendel', Janus 66 (1979), 33-47, and Olby (note 4), 199-207.
160. Mendel (note 6), 46.
161. Md., 47; translation slightly altered.
162. Ibid., 46-47; cf. Orel (note 3), 170. Olby ([note 7], 66), has argued that it is in these remarks that Mendel shows himself to be 'in conflict with classical Mendelian genetics'. We will come back to this point in our conclusions.
163. Mendel ([note 6], 46); Mendel's emphasis.
164. 2 (Mendel's 'first hybrid generation'). It is only this peculiar behaviour of a trait over three generations that indicates its 'hybrid character', and not, as in the instances of discoloration discussed by Mendel, the appearance of trait differences within one, or even two generations. Now, the verb vererben in German has a curious etymology, which is only insufficiently translated by the English 'inherit'. In its original legal sense, it was restricted to situations in which one person received from another something that the latter person had itself acquired by inheritance only. It was Immanuel Kant, in his writings on human races, who first exploited this legal terminology for biological purposes; see Peter McLaughlin, 'Kant on Heredity and Adaptation', in Heredity Produced: At the Crossroad of Biology, Politics, and Culture, 1500-1870, edited by Staffan Müller-Wille and Hans-Jörg Rheinberger (Cambridge, MA, in press).
165. Gregor Mendel, 'Über einige aus künstlicher Befruchtung gewonnene Hieraciumbastarde'. Verhandlungen des Naturforschenden Vereines in Brünn 8 (1869, published in 1870), 26-31 (28).
166. Cf. Callender (note 7), 58-59.
167. 1), not to members of the parental generation (P).
168. Mendel (note 142), 365. Some handwritten notes from later experiments indicate, that Mendel in the end succeeded to account for the polymorphism of Hieracium hybrids as a result of segregation of multifactorial traits; see Orel (note 3), 187.
169. Ibid., 364.
170. Ibid., 367.
171. Cf. J. Heimans, 'Mendel studying the theorem of the golden section', Folia Mendeliana 13 (1978), 235-42.
172. Mendel (note 6), 47; translation slightly altered.
173. Ibid., 47.
174. Ibid., 48.
175. Correns (note 5), 1270.
176. Olby (note 4), 254.
177. A similar view was expressed by R. A. Fisher in his introduction to Mendel (note 6), 4.
178. In a more recent re-assessment of his 1979 paper, Olby distinguishes between methodological and theoretical aspects of Mendel's achievements: 'By introducing the statistical approach and combinatorial mathematics into the subject of hybridization alongside his concept of the character-pair Mendel surely earned his place'.
179. We doubt, therefore, Gliboff's tentative conclusion that 'to Mendel himself, the goal of the research and the principal message of the paper was the law, not the mechanism', although we agree with Gliboff that this appeared so to his 'Austro-Ungerian reader[s]'; see Gliboff (note 3), p. 228.
180. Jean Gayon, 'From Measurement to Organization: A Philosophical Scheme for the History of the Concept of Heredity', in The Concept of the Gene in Development and Evolution: Historical and Epistemological Perspectives, edited by Peter Beurton, Raphael Falk and Hans-Jörg Rheinberger (Cambridge, MA, 2000), 69-90 (77-78).
181. On the nineteenth-century conception of heredity as a force, which was especially prevalent among breeders, see Jean Gayon and Doris T. Zallen, 'The role of the Vilmorin Company in the Promotion and Diffusion of the Experimental Science of Heredity in France, 1840-1920', Journal of the History of Biology 31 (1998), 241-62.
182. See, for example, Onno G. Meijer, 'Hugo de Vries no Mendelian?', Annals of Science 42 (1985), 189-232, on de Vries, and Olby (note 4) on William Bateson.
183. Frederic L. Holmes, Reconceiving the Gene: Seymour Benzer's Adventures in Page Genetics (New Haven, CT, 2006), ch. 1. Arguably, the various efforts to make sense out of Mendel's paper played a major role in the early history of this struggle. 'In the achievement of that "progress"', sociologists of science Barry Barnes, David Bloor, and John Henry argue, 'the misinterpretations of "Mendel's theory" were part of that creative use of the realist mode of speech which engendered adaptation and elaboration of
184. see Barry Barnes, John Henry, David Bloor, Scientific Knowledge: A Sociological Analysis (Chicago, 1996), 96.
185. On our reading, then, Mendel's concept of 'elements' or 'factors' came close to what Lenny Moss has called the preformationist notion of the gene ('gene-p'), without, however, conflating it with the developmental notion of the gene (gene-d), as many twentieth-century geneticists did; see Lenny Moss, What Genes Can't Do (Cambridge, MA, 2003), ch. 1. On genes as difference makers,
186. see C. Kenneth Waters, 'What was classical genetics?', Studies in History and Philosophy of Science 35 (2004), 83-89.
187. James Griesemer has argued on this basis that neither Mendel nor his twentieth-century followers ever separated development from heredity as many commentators - most prominently Iris Sandler and Laurence Sandler, A conceptual ambiguity that contributed to the neglect of Mendel's paper', History and Philosophy of the Life Sciences 7 (1985), 3-70 - have maintained. 'Genetic theories are theories of development', Griesemer says, 'but they are theories expressed in terms of developmental invariants';
188. see James Griesemer, 'Reproduction and the Reduction of Genetics', in The Concept of the Gene in Development and Evolution: Historical and Epistemological Perspectives, edited by Peter Beurton, Raphael Falk and Hans-Jörg Rheinberger (Cambridge, MA, 2000), 240-85 (261).
189. Gliboff (note 3), p. 223.
190. On the diversity of agendas of Mendel's 'rediscoverers' Bateson, Correns, de Vries, and Tschermak, see Ilse Jahn, 'Zur Geschichte der Wiederentdeckung der Mendelschen Gesetze', Wissenschaftliche Zeitschrift der Friedrich-Schiller Universität Jena, Mathematisch-naturwissenschaftliche Reihe 7 (1958), 215-27, and Olby (note 4), eh. 6.
191. Staffan Müller-Wille, 'Early Mendelism and the Subversion of Taxonomy: Epistemological Obstacles as Institutions', Studies in History and Philosophy of Biological and Biomedical Sciences 36 (2005), 465-87 (475-77).
192. Ibid., 70. Callender does not refer to Kuhn. His interpretation of Mendel seems more inspired by viewing history as an eternal struggle between theological and secular world views.
193. Olby ([note 7], 55) does not discuss Kuhn's work, but uses the expression 'paradigm' to designate the 'canonical ideas' invoked by twentieth-century interpretations of Mendel's work. Brannigan ([note 7], 449-51) discusses Kuhn in a long footnote. Brannigan's conclusion that 'Mendel's work figured as normal science in the hybridist tradition, while in 1900 the revival of Mendel's discovery of segregation constituted a relatively revolutionary achievement' (ibid, 424) comes very close to our own, with the important difference, however, that we regard the 'revolutionary achievement' of Mendel as a product of the 'normal science' he was engaged in.
194. Thomas S. Kuhn, 'Second Thoughts on Paradigms', in The Essential Tension: Selected Studies in Scientific Tradition and Change (Chicago, 1977), 293-319.
195. Cf. Hans-Jörg Rheinberger and Staffan Müller-Wille, 'Gene', in Stanford Encyclopedia of Philosophy, Winter 2004 edition, edited by E.N. Zalta. Available online at: http://plato.stanford.edu/entries/gene/ (accessed 28 September 2006).
196. See, for example, Thomas S. Kuhn, 'Logic of Discovery or Psychology of Research', in The Essential Tension: Selected Studies in Scientific Tradition and Change (Chicago, 1977), 266-92 (272).
197. For an interpretation of Kuhn that emphasizes this aspect, see Barry Barnes, T. S. Kuhn and Social Science (New York, 1982).