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Articles

Cambridge geneticists and the chromosome theory of inheritance: William Bateson, Leonard Doncaster and Reginald Punnett 1879–1940

Pages 468-496 | Received 25 Mar 2022, Accepted 08 Aug 2022, Published online: 17 Aug 2022
 

ABSTRACT

Early in the 20th century Bateson, Doncaster and Punnett formed a cooperative collective to share research findings on the chromosome theory of heredity (CTH). They cross-bred plants and animals to correlate behaviour of chromosomes and heredity of individual traits. Doncaster was the most enthusiastic proponent of the new theory and worked for years to convince Bateson and Punnett on its relevance to their own research. The two younger biologists collaborated with Bateson, the preeminent geneticist in England. As their own reputations developed, their research findings allied with the consensus on the importance of the CTH by the broader scientific community. After Doncaster’s tragic death in 1920, major objections to the theory had been resolved; Bateson and Punnett then utilized the CTH to construct chromosome maps detailing locations of specific genes on particular chromosomes in several different species. The marriage of heredity and cytology enhanced confidence that the theory was an accurate mechanism to explain inheritance in both plants and animals.

Disclosure statement

No potential conflict of interest was reported by the author(s).

Notes

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10 Bateson, Materials, p. 574.

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19 Clarence E. McClung, ‘Notes on the Accessory Chromosome’, Anatomischer Anzeiger, 20 (1901), 220–26; Leonard Doncaster, ‘Letter to William Bateson 18 March c1902’.

20 J. A. Venn, Alumni Cantabrigienses (Cambridge: Cambridge University Press, 1944), p. 318.

21 William Bateson, ‘Obituary of Leonard Doncaster’, Proceedings of the Royal Society, 92B (1921), xl–xlvi (xlii).

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23 William Bateson, Mendel’s Principles of Heredity: A Defence (Cambridge: Cambridge University Press, 1902), p. 30.

24 William Bateson, ‘Discussion’, Memoirs of the Horticultural Society of New York, 1 (1902), 123.

25 H. Lyster Jameson, ‘Zoological Station at Naples’, Report of the Meeting of the British Association for the Advancement of Science, 69 (1899), 433; Reginald C. Punnett, ‘On the Formation of the Pelvic Plexus, in Especial Reference to the Nevus Collector in the Genus Mustelus’, Philosophical Transactions, 192B (1900), 331–51.

26 Reginald C. Punnett, ‘On some Artic Nemerteans’, Proceedings of the Zoological Society of London, 2 (1901), 90–107; F. A. E. Crew, ‘R. C. Punnett’, Biographical Memoirs of Fellows of the Royal Society, 13 (1967), 309–26.

27 Reginald C. Punnett, ‘Merism and Sex in Spinax niger’, Biometrika, 3 (1904), 313–62.

28 William Bateson, ‘Letter to Leonard Doncaster’, 18 December 1903. MS Bateson Collection BA 193; William Bateson, ‘Letter to Leonard Doncaster’, 21 December 1903. MS Bateson Collection BA 194; Leonard Doncaster, ‘Letter to William Bateson’, 20 December 1903. MS Bateson Collection BA 191.

29 William Bateson, ‘Letter to Reginald C. Punnett’, 25 December 1903. MS Bateson Collection BA 193; Reginald C. Punnett, ‘Letter to William Bateson’, 30 December 1903. MS Bateson Collection BA 196; William Bateson, ‘Letter to Beatrice Bateson’, 10 January 1904. MS Bateson Collection BA 197; Reginald C. Punnett, ‘Early Days of Genetics’, Heredity, 4 (1950), 1–10 (4–5).

30 ‘Balfour Studentship’, University of Cambridge Calendar, (1904), 616–17; William Bateson, Edith R. Saunders, and Reginald C. Punnett, ‘Experiments in the Physiology of Heredity’, Reports to the Evolution Committee of the Royal Society, 2 (1905), 1–160 (89).

31 Stewart A. McDowall, ‘A Preliminary Note on the Maiotic Phenomena in the Eggs of the Hermaphrodite Angiostomum nigrovenosum’, Proceedings of the Cambridge Philosophical Society, 13 (1904), 309–11.

32 William Bateson, Edith R. Saunders, and Reginald C. Punnett, ‘Sweet Peas’, Reports to the Evolution Committee of the Royal Society, 2 (1904), 80–99 (80).

33 ‘University and Educational Intelligence’, Nature 69 (1903), 69.

34 Leonard Doncaster, ‘Letter to Reginald C. Punnett’, 13 April c1905. MS Bateson Collection BA 1464.

35 Leonard Doncaster, ‘On the Inheritance of Tortoiseshell and Related Colours in Cats’, Proceedings of the Cambridge Philosophical Society, 13 (1904), 35–8.

36 Leonard Doncaster, ‘On the Maturation and Early Development of the Unfertilised Egg in Certain Sawflies’, Proceedings of the Cambridge Philosophical Society, 13 (1904), 103–05.

37 Leonard Doncaster, ‘On the Maturation of the Germ-Cells in the Sawfly, Nematus ribesii’, Proceedings of the Cambridge Philosophical Society, 14 (1905), 22–3.

38 Yearbook of the Royal Society (London: Harrison and Sons, 1905), p. 87.

39 R. P. Gregory, ‘Letter to William Bateson’, 19 December 1903. MS Bateson Collection BA 2566; R. P. Gregory, ‘The Reduction Division in Ferns’, Proceedings of the Royal Society, 73B (1904), 86–92; R. P. Gregory, ‘Some Observations upon the Determination of Sex in Plants’, Proceedings of the Cambridge Philosophical Society, 12 (1904), 430–40.

40 E. B. Wilson, ‘Mendel’s Principles of Heredity and the Maturation of Germ Cells’, Science, 16 (1902), 991–93.

41 Beatrice Bateson, pp. 242–43.

42 W. E. Agar, ‘Letter to E. R. Babcock 21 December 1947’, quoted in Coleman, p. 262.

43 E. B. Wilson, ‘Letter to William Bateson’, 12 October 1905. MS Bateson Collection BA 835.

44 Leonard Doncaster and G. H. Raynor, ‘On Breeding Experiments with Lepidoptera’, Proceedings of the Zoological Society of London, (1906), 125–33 (126–28).

45 Sidney F. Harmer, ‘Letter to William Bateson’, 23 December 1906. MS Bateson Collection BA 148; William Bateson, ‘Letter to Sidney F. Harmer’, 27 December 1906. MS Bateson Collection BA 199.

46 Reginald C. Punnett, Mendelism (London: Macmillan and Bowen, 1905).

47 Reginald C. Punnett, Mendelism, 2nd edn (Cambridge: Bowes and Bowes, 1907), p. 64.

48 Reginald C. Punnett, ‘Mendelism’, Encyclopedia Britannica (1910–1911), xviii pp. 115–20 (p. 119).

49 Reginald C. Punnett, ‘Mendelism in Relation to Disease’, Proceedings of the Royal Society of Medicine, 1 Epidemiology Section (1908), 135–68 (153).

50 ‘Lecturers’, University of Cambridge Calendar, (1908), p. 881.

51 Beatrice Bateson, pp. 106–07.

52 ‘Professors’, University of Cambridge Calendar, (1908), p. lxiv; William Bateson, ‘The Methods and Scope of Genetics 1908’, in Beatrice Bateson, pp. 317–25.

53 William Bateson, ‘Facts Limiting the Theory of Heredity’, Science, 26 (1907), 649–60 (651–52); William Bateson, ‘Letter to C. B. Davenport’, 23 August 1907. MS Bateson Collection BA 4320.

54 Eric Ives, ‘Mason College: Staff and Students’, in The First Civic University: Birmingham 18801980, ed. by E. Ives, D. Drummond and L. Schwarz (Birmingham: University of Birmingham Press, 2000), pp. 48–69; Leonard Doncaster, ‘Inheritance and Sex in Abraxas grossulariata’, Nature, 76 (1907), 248; Leonard Doncaster, ‘Gametogenesis and Fertilisation in Mematus ribesii’, Quarterly Journal of Microscopical Science, 51 (1907), 101–14.

55 Leonard Doncaster, ‘Letter to William Bateson’, 14 January 1908. MS Bateson Collection BA 1433; Leonard Doncaster, ‘Letter to William Bateson’, 20 March c1908. MS Bateson Collection BA 1439.

56 Leonard Doncaster, ‘Letter to William Bateson’, 2 August 1908. MS Bateson Collection BA G70-21; Leonard Doncaster, ‘On Sex Inheritance in the Moth Abraxas grossulariata and its var. Lacticolor’, Reports to the Evolution Committee of the Royal Society, 4 (1908), 53–7.

57 Leonard Doncaster, ‘Gametogenesis of the Gall Fly Neuroterus lenticularis’, Proceedings of the Royal Society, 82B (1909): 88–113.

58 Leonard Doncaster, Heredity in the Light of Recent Research (Cambridge: Cambridge University Press, 1910), pp. 132–34.

59 E. B. Wilson, ‘Secondary Chromosome Coupling and the Sexual Relations in Abraxas’, Science, 29 (1909), 704–06.

60 E. B. Wilson, ‘Recent Researches on the Determination and Heredity of Sex’, Science, 29 (1909), 53–70 (68–9).

61 E. B. Wilson, ‘Letter to William Bateson’, 2 April 1909. MS Bateson Collection BA 3135.

62 Ernst Mayr, The Growth of Biologic Thought (Cambridge: Harvard University, 1982) p. 734.

63 Cock and Forsdyke, pp. 381–83; Robert Olby, ‘Scientists and Bureaucrats in the Establishment of the John Innes Horticultural Institution under William Bateson’, Annals of Science, 46 (1989), 497–510.

64 Coleman, p. 252.

65 Olby, ‘Scientists and Bureaucrats’, p. 507.

66 William Bateson, ‘Letter to Beatrice Bateson’, 1910. MS Bateson Collection Add.8634/A16:9.2v.

67 Crew, p. 316.

68 Cock and Forsdyke, p. 386; A. W. F. Edwards, ‘Reginald Crundall Punnett: First Arthur Balfour Professor Genetics, Cambridge, 1912’, Genetics, 192 (2012), 3–13; ‘Professors’, University of Cambridge Calendar, (1913), p. lxxix.

69 Cambridge University Calendar (1914–1915), pp. 610 and 874.

70 Leonard Doncaster, ‘Note on the Spermatogenesis of Abraxas grossulariata’, Proceedings of the Cambridge Philosophical Society, 16 (1911), 44–5; Leonard Doncaster, ‘Letter to William Bateson’, 17 February 1912. Bateson Collection BA 1445; Leonard Doncaster, ‘Letter to William Bateson’, 29 December 1912. Bateson Collection BA 1450.

71 Leonard Doncaster, ‘Letter to William Bateson’, 26 December 1912. Bateson Collection BA 1449.

72 Leonard Doncaster, ‘Letter to William Bateson’, 5 January 1913. Bateson Collection BA 1451.

73 Garland Allen, Thomas Hunt Morgan (Princeton NJ: Princeton University Press, 1978).

74 Mayr, pp. 752–61; Rafael Falk, ‘Linkage: From Particulate to Interactive Genetics’, Journal of the History of Biology, 36 (2003), 87–117 (101).

75 T. H. Morgan, ‘Chromosomes and Heredity’, The American Naturalist, 44 (1910), 449–96 (465).

76 Calvin B. Bridges, ‘Direct Proof through Non-Disjunction that the Sex-Linked Genes of Drosophila are Borne by the X-chromosome’, Science, 40 (1914), 107–09.

77 Mayr, pp. 769–72.

78 T. H. Morgan, ‘An Attempt to Analyze the Constitution of the Chromosomes on the Basis of Sex-Limited Inheritance in Drosophila’, Journal of Experimental Zoology, 11 (1911), 403–09 (409).

79 A. H. Sturtevant, ‘The Linear Arrangement of Six Sex-Linked Factors in Drosophila, as Shown by their Mode of Association’, Journal of Experimental Zoology, 14 (1913), 43–59.

80 T. H. Morgan, Heredity and Sex (New York: Columbia University Press, 1913), pp. 92–3.

81 T. H. Morgan, ‘Sex-Limited Inheritance in Drosophila’, Science, 32 (1910), 120–22.

82 C. D. Darlington, ‘Genetics and Plant Breeding 1910–1980’, Philosophical Transactions, 292B (1981), 401–06 (401).

83 Leonard Doncaster, ‘Chromosomes, Heredity and Sex: A Review of the Present State of the Evidence with Regard to the Material Basis of Hereditary Transmission and Sex-Determination’, Quarterly Journal of Microscopical Science, 59 (1914), 487–521 (511–12).

84 Leonard Doncaster, Determination of Sex (Cambridge: Cambridge University Press, 1914), pp. 35–68.

85 Doncaster, Determination of Sex, p. vii.

86 Ibid., p. 56.

87 William Bateson, ‘Review of The Mechanism of Mendelian Heredity’, Science, 44 (1916), 536–43 (542).

88 William Bateson, ‘Letter to Leonard Doncaster’, 27 August 1917. Bateson Collection BA 1463; Martins, pp. 327–67.

89 Leonard Doncaster, ‘Letter to William Bateson’, 21 July 1916. Bateson Collection BA 3915.

90 Leonard Doncaster, ‘Letter to William Bateson’, 2 November 1916. Bateson Collection BA 1461.

91 Robert H. Lock and Leonard Doncaster, Recent Progress in the Study of Variation, Heredity and Evolution, 4th edn (London: John Murray, 1916) pp. 265–66.

92 William Bateson, ‘Letter to Leonard Doncaster’, 27 August 1917. Bateson Collection BA 1463.

93 Reginald C. Punnett, Mendelism, 3rd edn (London: Macmillan, 1911), p. vi

94 William Bateson, Mendel’s Principles of Heredity (Cambridge: Cambridge University Press, 1909), pp. 157–61.

95 William Bateson, ‘Letter to Erwin Baur’, 13 April 1911. Bateson Collection BA 967.

96 William Bateson, ‘Letter to Erwin Baur’, 29 April 1911. Bateson Collection BA G9C-25.

97 William Bateson and Reginald C. Punnett, ‘A Gametic Series Showing Reduplication’, Journal of Genetics, 1 (1911), 293–302; William Bateson and Reginald C. Punnett, ‘On the Inter-Relations of Genetic Factors’, Proceedings of the Royal Society, 84B (1911), 3–8.

98 Cock, p. 47.

99 Reginald C. Punnett, ‘Reduplication Series in Sweet Peas’, Journal of Genetics, 3 (1913), 77–103.

100 A. H. Trow, ‘Forms of Reduplication—Primary and Secondary’, Journal of Genetics, 2 (1913), 313–24.

101 Calvin B. Bridges, ‘The Chromosome Hypothesis of Linkage Applied to Cases of Sweet Peas and Primula’, The American Naturalist, 48 (1914), 524–34.

102 A. H. Sturtevant, ‘The Reduplication Hypothesis as Applied to Drosophila’, The American Naturalist, 48 (1914), 535–49 (548).

103 Morgan, The Mechanism of Mendelian Heredity, p. 77.

104 Reginald C. Punnett, ‘Reduplication Series in Sweet Peas. II’, Journal of Genetics, 6 (1917), 185–93 (190).

105 J. B. S. Haldane, ‘The Combination of Linkage Values and the Calculation of Distances between the Loci of Linked Factors’, Journal of Genetics, 8 (1919), 299–309.

106 Reginald C. Punnett, ‘Letter to William Bateson’, 28 August 1916. Bateson Collection BA 4044.

107 Reginald C. Punnett, ‘Letter to William Bateson’, 17 February 1917. Bateson Collection BA 950.

108 Reginald C. Punnett, ‘Letter to William Bateson’, 31 October 1918. Bateson Collection BA 1530.

109 Reginald C. Punnett, ‘Letter to William Bateson’, 26 May 1921. Bateson Collection BA 955.

110 William Bateson, ‘Root-Cuttings, Chimaeras and “Sports”’, Journal of Genetics, 6 (1916), 75–80.

111 William Bateson, ‘Studies in Variegation’, Journal of Genetics, 8 (1919), 93–9; William Bateson and I. Sutton, ‘Double Flowers and Sex-Linkage in Begonia’, Journal of Genetics, 8 (1919), 199–207.

112 Leonard Doncaster, An Introduction to the Study of Cytology (Cambridge: Cambridge University Press, 1920), p. 229.

113 Leonard Doncaster, ‘Biology since Darwin’, in Recent Developments in European Thought, ed. by F. S. Marvin (London: Oxford University Press, 1920), pp. 229–46.

114 Leonard Doncaster, ‘Genetic Studies of Drosophila’, Nature, 105 (1920), 405–06.

115 Thomas Kelley and R. F. Whelan, For the Advancement of Learning: The University of Liverpool, 18811981 (Liverpool: University of Liverpool Press, 1981), p. 238; W. A. Herdman, ‘Leonard Doncaster, FRS’, Sphinx, 27 (1920), 4–5.

116 Bateson, ‘Obituary of Leonard Doncaster’, pp. xl–xlvi; William Bateson, ‘Obituary of Leonard Doncaster’, Nature, 105 (1920), 461–62; William Bateson, ‘Letter to Mrs. Leonard Doncaster 29 May 1920’, quoted in Cock, 54.

117 Herdman, p. 5.

118 Bateson, ‘Root-Cuttings’.

119 William Bateson and I. Sutton, ‘Double Flowers and Sex-Linkage in Begonia’, Journal of Genetics, 8 (1919), 199–207.

120 William Bateson, ‘Progress of Mendelism’, Nature, 104 (1919), 214–17 (215); William Bateson and Caroline Pellew, ‘The Genetics of “Rogues” among Culinary Peas, Pisum sativum’, Proceedings of the Royal Society, 91B (1920), 186–95 (194).

121 William Bateson and A. E. Gairdner, ‘Male Sterility in Flax, Subject to Two Types of Segregation’, Journal of Genetics, 11 (1921), 269–75 (272).

122 William Bateson, ‘Gametic Segregation’, Proceedings of the Royal Society, 91B (1920), 358–68.

123 William Bateson, ‘Letter to H. Chisholm’, 8 November 1920. Bateson Collection BA G5-L8.

124 Darden, pp. 121–37.

125 Coleman, p. 260.

126 William Bateson, ‘Letter to Caroline Pellew’, 21 December 1921. Bateson Collection BA 1152.

127 William Bateson, ‘Letter to Beatrice Bateson’, 24 December 1921. Bateson Collection BA G8G-01J.

128 William Bateson, ‘Letter to Beatrice Bateson’, 26 December 1921. Bateson Collection BA G8-01K.

129 William Bateson, ‘Evolutionary Faith and Modern Doubts’, Nature, 109 (1922), 553–56 (553).

130 Alan G. Cock, ‘Bateson’s two Toronto Addresses: 1. Chromosomal Skepticism’, Journal of Heredity, 80 (1989), 91–5; William Bateson, ‘Lecture Notes “The Outlook of Genetics”’, 1922. Bateson Collection BA 1213.

131 Helen Gwynne-Vaughan, ‘Letter to William Bateson’, 28 June 1922. Bateson Collection BA 2278; William Bateson, ‘Note regarding W. C. F. Newton’, c1922. Bateson Collection BA 2279.

132 Bateson and Pellew, pp. 186–95.

133 R. R. Gates, ‘A Preliminary Account of the Meiotic Phenomena in the Pollen Mother-Cells and Tapetum of Lettuce’, Proceedings of the Royal Society, 91B (1920), 216–23.

134 R. P. Gregory, D. de Winton, and William Bateson, ‘Genetics of Primula sinensis’, Journal of Genetics, 13 (1923), 219–53.

135 W. C. F. Newton and Caroline Pellew, ‘Primula kewensis and its Derivatives’, Journal of Genetics, 20 (1929), 405–67.

136 G. Jorgensen, ‘Letter to William Bateson’, 14 November 1924. Bateson Collection BA 1090; G. Jorgensen, ‘Letter to William Bateson’, 17 January 1925. Bateson Collection BA 1095.

137 W. C. F. Newton, ‘Studies on Somatic Chromosomes I. Pairing and Segmentation’, Annals of Botany, 38 (1924), 199–206; W. C. F. Newton, ‘Chromosome Studies in Tulipa and some Related Genera’, Journal of the Linnean Society of London, Botany, 47 (1926), 339–54; C. D. Darlington, ‘Chromosome Studies in the Scilleae’, Journal of Genetics, 16 (1926), 237–51.

138 W. C. F. Newton, ‘The Inheritance of Flower Colour in Papaver rhoeas and Related Forms’, Journal of Genetics, 21 (1929), 389–404; Oren S. Harman, The Man Who Invented the Chromosome: A Life of Cyril Darlington (Cambridge: Harvard University Press, 2004), pp. 31–5.

139 William Bateson, ‘Notes for University of London Lecture’, 3 November 1921. Bateson Collection BA G8C-08.

140 William Bateson, ‘Notes for Leicester Lecture’, 20 November 1922. Bateson Collection BA G8C-10.

141 William Bateson, ‘Notes for Birmingham Lecture’, 26 November 1923. Bateson Collection BA G8C-11; William Bateson, ‘Notes on Primula Linkage’, February 1924. Bateson Collection BA G8C-13.

142 Harman, p. 33.

143 C. D. Darlington, ‘Comments by William Bateson 1923–24’, quoted in Coleman, p. 262.

144 Clifford Dobell, ‘Letter to William Bateson’, 25 May 1924. Bateson Collection BA1065.

145 William Bateson, ‘Letter to Clifford Dobell’, 22 May 1924. Bateson Collection BA 1046.

146 Reginald C. Punnett, Mendelism, 5th edn (London: Macmillan, 1922), p. v.

147 Punnett, Mendelism, 5th edn, pp. 137–46.

148 Cock, William Bateson’s Recognition’, pp. 54–5.

149 Reginald C. Punnett, ‘Linkage in Sweet Peas’, Journal of Genetics, 13 (1923), 101–23.

150 Reginald C. Punnett, ‘Genes and Linkage Groups in Genetics’, Nature, 117 (1926), 514–15, (515).

151 Reginald C. Punnett, ‘As a Biologist Sees It’, The Nineteenth Century and After, 97 (1925), 697–706 (697).

152 Reginald C. Punnett, Mendelism, 7th edn (London: Macmillan, 1927), p. v.

153 Punnett, Mendelism, 7th edn, p. 135.

154 Ibid., p. 149.

155 Hutchinson, p. 99.

156 Reginald C. Punnett, ‘Linkage Groups and Chromosome Number in Lathyrus’, Proceedings of the Royal Society, 102B (1927), 236–38.

157 Reginald C. Punnett, ‘Further Studies of Linkage in the Sweet Pea’, Journal of Genetics, 26 (1932), 97–112; Reginald C. Punnett, ‘Notes on the D-Chromosome of the Sweet Pea’, Journal of Genetics, 39 (1940), 301–08.

158 Reginald C. Punnett, ‘Genetic Studies in Fowl X. Linkage Data for the Sex Chromosome’, Journal of Genetics, 39 (1940), 335–42.

159 Punnett, Mendelism, 2nd edn, p. 64.

160 Derek J. DeSolla Price and Donald DeB. Beaver, ‘Collaboration in an Invisible College’, American Psychologist, 21 (1966), 1011–18 (1011); Margery Purver, The Royal Society: Concept and Creativity (New York: Routledge, 2009) pp. 194–95.

161 J. S. Katz and B. R. Martin, ‘What is Research Collaboration?’ Research Policy, 26 (1997), 1–18.

162 D. DeB. Beaver and Richard Rosen, ‘Studies in Scientific Collaboration III. Professionalization and the Natural History of Scientific Co-Authorship’, Scientometrics, 1 (1979), 231–45.

163 Donald DeB. Beaver, ‘Reflections on Scientific Collaboration (and its Study): Past, Present and Future’, Proceedings of the Second Berlin Workshop on Scientometrics and Informetrics, (2000), 29–40.

164 Coleman, pp. 294–98.

165 Donald DeB. Beaver and Richard Rosen, ‘Studies in Scientific Collaboration I. The Professional Origins of Scientific Co-Authorship’, Scientometrics, 1 (1978), 65–84.

166 Darlington, ‘Genetics and Plant Breeding’, p. 401.

167 Rushton, ‘William Bateson and the Chromosome Theory’, p. 154.

168 Ibid., p. 153.

169 Darden, pp. 121–30.

170 James B. Farmer and J. E. S. Moore, ‘On the Essential Similarities Existing between the Heterotype Nuclear Division in Animals and Plants’, Anatomischer Anzeiger, 11 (1895), 71–80.

171 Cock, ‘William Bateson’s Recognition’, pp. 48–50.

172 Wei Lu et al., ‘Scientific Collaboration and Career Stages: An Ego-Centric Perspective’, Journal of Informetrics, (2022), https://doi.org/10/j.joi2021.101207.

173 Cock, ‘William Bateson’s Recognition’, p. 53.

174 Brush, p. 489.

175 Bateson, ‘Obituary of Leonard Doncaster’, p. 462.

176 Darlington, ‘Genetics and Plant Breeding’, p. 401.

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