Galton's ‘Law of Ancestral Heredity’: Its Influence on the Early Development of Human Genetics

Galton F. , “Regression towards mediocrity in hereditary stature”, Journal of the Anthropological Institute of Great Britain and Ireland, xv (1885) 246–63.

Pearson K. , “Mathematical contributions to the theory of evolution. On the law of ancestral heredity”, Proceedings of the Royal Society, lxii (1898) 386–412. A less common variant was “the law of ancestral inheritance”, Life, IIIA, 21.

Swinburne R. G. , “Galton's law—formulation and development”, Annals of science, xxi (1965) 15–31.

Froggatt P. Nevin N. C. , “The ‘law of ancestral heredity’ and the Mendelian-ancestrian controversy in England, 1889–1906”, Journal of medical genetics, viii (1971) 1–36.

Spencer H. , Principles of biology (London, 1864, 1867). Spencer's maxim that function is prior to structure and the results of organic use and disuse immediately incarnate in structural adaptations capable of inheritance brought him into direct controversy with Weismann and Galton.

de M. Lamarck J. B. P. A. , Histoire des animaux sans vertèbres, i (Paris, 1815) 199 ff. Lamarck postulated four ‘laws’ to explain the organisation of animals and formation of different organs. This was the fourth.

Darwin C. , The variation of animals and plants under domestication (London, 1868), chapter 27. He had sent the MS. to Huxley in 1865: See Olby R. C. , “Charles Darwin's manuscript of pangenesis”, British journal of the history of science, i (1963) 251–63.

Vorzimmer P. , “Charles Darwin and blending inheritance”, Isis, liv (1963) 371–90, and Olby R. C. , “The Mendel centenary”, British journal of the history of science, ii (1965) 343–9 give good brief accounts of the issues.

Wallace to Sir C. Lyell of 20 February 1868. See Wallace A. R. , My life: A record of events and opinions, i (London, 1905) 422.

Galton F. , Hereditary genius: An enquiry into its laws and consequences (London, 1869) 363ff.

Galton F. , “Experiments in pangenesis, by breeding from rabbits of a pure variety, into whose circulation blood taken from other varieties had previously been largely transfused”, Proceedings of the Royal Society, xix (1871) 393–410.

See reference 7 (Darwin), ii, 374, 379: Also Galton F. , “Pangenesis”, Nature, iv (1871) 5–6; and Darwin C. , “Pangenesis”, ibid., iii (1871) 502–3. Strangely, Darwin had not corrected Galton in their regular correspondence. See Life, II, 113, 156 ff.

“But it does not appear to me that Pangenesis has, as yet, received its death blow; though … its life is always in jeopardy”. See Darwin C. , “Pangenesis”, Nature, iii (1871) 502–3.

For the development of Galton's thinking and the possibility that he might have reached his theory earlier but for his high regard for Darwin, see Life, II, 114–5, 169 ff.

Galton F. , “On blood relationship”, Proceedings of the Royal Society, xx (1872) 394–402.

Galton F. , “A theory of heredity”, Contemporary review, xxvii (1875) 80–95.

Galton F. , “A theory of heredity”, Journal of the Anthropological Institute of Great Britain and Ireland, v (1876) 329–48.

Galton's ‘stirp’ is very similar to Weismann's ‘germ plasm’: In fact the latter devoted two pages of his book to asserting (unsuccessfully) his own originality. See Weismann A. , The germ plasm. A theory of heredity, trans. Parker W. Newton Rönnfeldt Harriet (London, 1893) 198–200. Galton was too courteous to argue priorities.

Weismann A. , Ueber die Vererbung (Jena, 1883).

de Vries H. , Intracelluläre Pangenesis (Jena, 1889).

de Vries H. , Die Mutationstheorie. Versuche und Beobachtungen über die Entstehung der Arten im Pflanzenreich, i: Die Entstehung der Arten durc (Leipzig, 1901).

de Vries H. , “Das Spaltungsgesetz der Bastarde (Vorläufige Mitteilung)”, Bericht der Deutschen botanischen Gesellschaft, xviii (1900) 83–90.

For a review see Hughes A. , A history of cytology (London, 1959), and Coleman W. , “Cell, nucleus, and inheritance; An historical study”, Proceedings of the American Philosophical Society, cix (1965) 124–58.

Gasking E. B. , “Why was Mendel's work ignored?”, Journal of the history of ideas, xx (1959) 60–84; also Wilkie J. S. , “Some reasons for the rediscovery and appreciation of Mendel's work in the first years of the present century”, British journal of the history of science, i (1963) 5–17.

Boveri Th. , “Ueber mehrpolige Mitosen als Mittel zur Analyse des Zellkerns”, Verhandlungen der Physikalisch-medizinischen Gesellshaft zu Würzburg, xxxv (1902) 67–90.

Sutton W. S. , “On the morphology of the chromosome group of Brachystola magna”, Biological bulletin, Marine Biological Laboratory, Wood's Hole, Mass., iv (1902) 24–39; also his article “The chromosomes in heredity”, ibid., iv (1903) 231–51.

A description of particularly Sutton's work is given by McKusick V. A. , “Walter S. Sutton and the physical basis of Mendelism”, Bulletin of the history of medicine, xxxiv (1960) 487–97.

Life, II, 70.

Galton F. , “Hereditary talent and character”, Macmillan's magazine, xii (1865) 157–66, 318–27.

Undated MS (Life, II, 70). The quotation continues: “I am sure I assimilated it with far more readiness than most people, absorbing it almost at once, and my afterthoughts were permanently tinged by it.” Galton demonstrated this influence time and again.

Life, II, 84. Galton later corrected it (see reference 1).

Galton F. , Memories of my life (London, 1908) 309.

Galton F. , “The average contribution of each several ancestor to the total heritage of the offspring”, Proceedings of the Royal Society, lxi (1897) 401–13. See especially the paragraph “Again a wide … assurance a priori” on p. 403.

See reference 29, particularly the passage “We shall … point of interlacement” on p. 322.

Life, IIIA, 6 ff.

Ibid, 11.

Galton F. , “Family likeness in stature”, Proceedings of the Royal Society, xl (1886) 42–69. See also reference 1 and Life, IIIA, Chapter 14.

Galton F. , “Family likeness in eye-colour”, ibid., xl (1886) 402–16.

Life, II, 357 ff. The data were published separately: See Galton's Record of family faculties (London, 1884) and Life history album (London, 1884).

Galton F. , Natural inheritance (London, 1889).

Galton F. , “Co-relations and their measurement, chiefly from anthropometric data”, Proceedings of the Royal Society, xlv (1888) 135–45. ‘Co-relation’ was replaced by ‘correlation’ the following year (Life, IIIA, 57 footnote).

Life, IIIA, 22 ff.

See reference 40, pp. 134 ff.

Life, IIIA, 23.

See reference 38. Pearson has discussed invalidities in the method (Life, IIIA, 34–40).

Life, IIIA, 45–50.

Reference 32, p. 308.

Galton F. , “A new law of heredity”, Nature, lvi (1897) 235–7. Also reference 33.

About this time Pearson wrote, “… with all due reservations, it seems to me that the law of ancestral heredity … is highly probably … the simple descriptive statement which bring into a single focus all the complex lines of hereditary influence” (Reference 2—Pearson).

Galton F. , “Hereditary colour in horses”, Nature, lvi (1897) 598–9.

Life, IIIA, 21. Pearson considered this to be also Galton's view (ibid., p. 60).

Pearson K. , “Mathematical contributions to the theory of evolution. III: Regression, heredity, and panmixia”, Philosophical transactions(A), clxxxvii (1896) 253–318.

Galton initially failed to grasp the concept of multiple regression and adduced the ancestral law ambiguously in Natural inheritance (reference 40, pp. 132 ff.)

Galton wrote: “It delights me beyond measure to find that you are harmonising what seemed disjointed and cutting out and replacing the rotten planks of my propositions” (F.G. to K.P. of 4 January 1898, in Life, IIIB, 504).

Cited (p. 203) in: Pearson E. S. , “Karl Pearson: An appreciation of some aspects of his life and work. Part I: 1857–1906”, Biometrika, xxviii (1936) 193–257.

Pearson K. Lee Alice , “Mathematical contributions to the theory of evolution. VIII: On the inheritance of characters not capable of exact quantitative measurement”, Philosophical transactions, cxcv (1900) 79–150. See also Life, IIIA, 34 ff.

Pearson K. , “Mathematical contributions to the theory of evolution. VII: On the correlation of characters not quantitatively measurable”, ibid., 1–47.

Pearson K. , “Mathematical contributions to the theory of evolution. On the law of reversion”, Proceedings of the Royal Society, lxvi (1900) 140–64. This was sub-headed “A New Year's greeting to Francis Galton—January 1st, 1900.”

Pearson K. , The grammar of science, 2nd ed. (London, 1900) 494–5.

Reference 56 (Pearson and Lee) p. 121.

Pearson's three articles (references 56, 57, 58) were submitted to the R. S. on August 5, 1899 as a single memoir. Pearson later withdrew it for emendations and expansion and re-submitted the material as three articles between January and March 1900.

Short biographies and references to longer ones are in reference 4.

Pearson E. S. , “Karl Pearson. An appreciation of some aspects of his life and work, Part II: 1906–1936”, Biometrika, xxix (1938) 161–248.

Weldon W. F. R. , “On certain correlated variations in Carcinus moenas”, Proceedings of the Royal Society, liv (1893) 318–29; Pearson K. , “Contributions to the mathematical theory of evolution”, Philosophical transactions (A), clxxxv (1894) 71–110.

Members were to be Francis Darwin, Galton, A. Macalister, R. Meldola, E. P. Poulton and Weldon, and the petition stated that “it may afterwards be desirable to add a statistician”. See Pearson K. , “Walter Frank Raphael Weldon, 1860–1906”, Biometrika, v (1906–7) 1–52.

Facts from: Life, IIIA, 126 ff; reference 65; and correspondence W.F.R.W. to F.G. of 4 December 1893 and F.G. to K.P. of 16 July 1906, in Life, IIIA, 128–9.

Weldon W. F. R. , “Report of the committee … for conducting statistical enquiries into the measurable characteristics of plants and animals. Part I: An attempt to measure the death-rate due to the selective destruction of Carcinus moenas with respect to a particular dimension; Part II: Remarks on variation in animals and plants”, Proceedings of the Royal Society, lvii (1895) 360–79, 379–82.

Reference 65, p. 25.

Ibid., p. 26 footnote.

Bateson W. , Materials for the study of variation treated with especial regard to discontinuity in the origin of species (London, 1894).

Weldon W. F. R. , “The study of animal variation. Review of Materials for the study of variation, etc., by W. Bateson”, Nature, 1 (1894) 25–6.

The orthodox view, put by Weldon, Botting and Thistleton-Dyer, was that cultivated forms of Cineraria came from a wild species through the gradual accumulation of small variations induced by the horticulturist; Bateson held that they came through hybridization between several recognised species (see reference 4).

“Until the time—about 16 years ago [1890]—when his mind began to embitter itself against me, I was more intimate with him than I have ever been with anyone but you.” (W.B. to B.B. of 16 April 1906, in Beatrice Bateson, William Bateson, F.R.S. Naturalist. His essays and addresses together with a short account of his life (Cambridge, 1928) 102.) Pearson dated the start of the enmity more exactly: “Bateson's attacks did not start until Weldon had reviewed his book [Reference 71] in 1894 … and then they became incessant and ceased only with the death of Weldon.” (K. P. to F. G. of 11 July 1906, in Life, IIIA, 287–8.)

Reference 65, p. 26.

K. P. to F. G. of 14 July 1906 (Life, IIIA, 289–90).

Reference 10, 2nd ed., xviii-xix. Later Pearson (Life, IIIA, 80) defended this ‘heresy’ and wrote: “it is quite certain that Galton in 1892 supported evolution by mutations owing to an error in interpretation [in regression in the ancestral law]” (our italics). Galton distinguished between mutations (‘divergent variations’) and minor variations (‘transilient variations’) and ascribed each a different role in evolution. His writings on this are difficult to explain concisely: Details are in Life, passim; reference 3; and Wilkie J. S. , “Galton's contribution to the theory of evolution with special reference to his use of models and metaphors”, Annals of science, xi (1955) 194–205.

Galton F. , “Discontinuity in evolution”, Mind, iii (1894) 362–72.

Life, IIIA, 126; F. G. to W. F. R. W. of 17 November 1896 (Life, IIIA, 127); F. G. to K. P. of 15 February 1897 (Life, IIIB, 501) and 16 July 1906 (Life, IIIA, 290–1); and reference 73 (Beatrice Bateson), p. 60.

K. P. to F. G. of 28 June 1906 (Life, IIIA, 285–6).

Reference 55, p. 223.

Pearson K. Lee A. Warren E. Fry A. Fawcett C. D. , “Mathematical contributions to the theory of evolution. IX: On the principle of homotyposis and its relation to heredity, to the variability of the individual and to that of the race. Part I: Homotyposis in the vegetable kingdom”, Philosophical transactions, cxcvii (1901) 285–379. “Part II: Homotyposis in the animal kingdom”, was announced ten years later but never published.

Abstract of idem., Proceedings of the Royal Society, lxviii (1900) 1–5.

Quotation from Pearson K. , “On the fundamental conceptions of biology”, Biometrika, i (1902) 320–44.

Bateson W. , “Heredity, differentiation, and other concepts of biology: A consideration of Professor Karl Pearson's paper ‘On the principle of homotyposis”’, Proceedings of the Royal Society, lix (1901) 193–205. This paper was very temperate in tone, unlike Pearson's rejoinder (reference 83) and Bateson's own later attacks.

“… if the R. S. people send my papers to Bateson, one cannot hope to get them printed. It is a practical notice to quit. This notice applies not only to my work, but to most work on similar statistical lines” (K. P. to F. G. of 13 December 1900, in Life, IIIA, 241).

Reference 65, p. 35.

Life, IIIA, 241 ff.

Life, IIIA, 100.

The editors were discerning in their acceptance of papers, rejecting Bateson's reply to Pearson's rejoinder (reference 83) of Bateson's own criticism (reference 84) of Pearson's ‘homotyposis’ paper (reference 81). Bateson had it printed privately (in July 1903) by the same press and to the identical format of Biometrika! See Bateson W. , Variation and differentiation in parts and brethren (Cambridge, 1903).

Reference 65, p. 36.

To the Royal Horticultural Society on 8 May 1900. See Bateson W. , “Problems of heredity as a subject for horticultural investigation”, Journal of the Royal Horticultural Society, xxv (1900) 54–61.

By Druery C. T. Mendel Gregor , “Experiments in plant hybridisation”, ibid., xxvi (1901) 4–32.

One of the series from the Biometric Laboratory (written mostly by Pearson) is entitled Questions of the day and of the fray. It dealt polemically with controversial issues, including social ones. This sense of intellectual combat, the ‘fray’, was Pearson's main emotional outlet after he dropped his literary, philosophical, and political writing.

Fisher R. A. , “Has Mendel's work been rediscovered?”, Annals of science, i (1936) 115–37.

Weldon W. F. R. , “Mendel's law of alternative inheritance in peas”, Biometrika, i (1902) 228–53.

Phenotype differences between homozygote and heterozygote were becoming well-established, e.g. Correns C. , “G. Mendel's Regeln über das Verhalten der Nachkommenschaft der Rassenbastarde”, Bericht der Deutschen botanischen Gesellschaft, xviii (1900) 158–68; and “Ueber Bastarde zwischen Rassen von Zea Mays”, ibid., xix (1900) 211–20.

Bateson W. , Mendel's principles of heredity: A defence (Cambridge, 1902), 98. Ibid, p. vi.

Udny Yule described Bateson's method and style as those of “the religious revivalist [and] are illsuited to scientific controversy … A writer who indulges himself in displays of this kind loses his right to be treated either as an impartial critic or as a sober speculator”. See Yule G. U. , “Mendel's laws and their probable relations to intra-racial heredity”, New phytologist, i (1902) 193–207, 222–37. Bateson's wife referred to “the outspoken hardness and clarity of the ‘Defence»’ (reference 73, p. 74). Bateson considered himself as a “knocker-up … to rouse others from their slumber … and if I have knocked a trifle loud, it is because there is need” (reference 97, p. xii).

Weldon W. F. R. , “On the ambiguity of Mendel's categories”, Biometrika, ii (1902) 44–55. This was a temperate reply: Weldon was never acrimonious.

Bateson W. Saunders E. R. , Experimental studies in the physiology of heredity, Reports of the Evolution Committee of the Royal Society, i (London, 1902).

See particularly Weldon's review of the first volume of de Vries's Die Mutationstheorie (reference 21). See Weldon W. F. R. , “Professor de Vries on the origin of species”, Biometrika, i (1902) 365–74.

Until Pearson allied himself with Weldon, Bateson held him in high regard. Only five days after he had read Weldon's article refuting Mendelism (reference 95) and before Pearson's allegiance was known, Bateson wrote: “I respect you as an honest man, and perhaps the ablest and hardest worker I have met, and I am determined not to take up a quarrel with you if I can help it … you are probably the only Englishman I know at this moment whose first thought is to get at the truth of these [inheritance] problems … “(W. B. to K. P. of 13 February 1902, in reference 55, p. 204 footnote).

Reference 99 (Yule), p. 207.

Pearson K. , “Mathematical contributions to the theory of evolution. XII: On a generalised theory of alternative inheritance, with special reference to Mendel's laws”, Philosophical transactions (A), cciii (1904) 53–86. Weldon refused the joint authorship his contribution merited because “he had taken no part in certain portions of the more complicated algebraic analyses” (references 65, p. 43).

Fisher R. A. , “The correlation between relatives on the supposition of Mendelian inheritance”, Transactions of the Royal Society of Edinburgh, lii (1918) 399–433; republished, with critical commentary, by Moran P. A. P. Smith C. A. B. as Eugenics Laboratory Memoirs, xli (Cambridge, 1966).

Reference 99. Yule's conclusions dealt with the ancestral hypothesis and specifically excluded “with regret … the special laws as to the operation of Ancestral Heredity which were formulated by Galton and Pearson … [because] the fixity of the numerical constants involved, which they imply, has not stood the test of time.” Characteristically, Pearson did not let this pass without (adverse) comment! See Pearson K. , “Note on a paper by G. U. Yule”, Biometrika, ii (1903) 228–9.

Reference 65, p. 39.

Pearson K. , “The law of ancestral heredity”, Biometrika, ii (1903) 211–28.

Reference 107 (Pearson), p. 228.

Weldon was also a prime motivator (references 65, p. 43; 105, p. 53 footnote). Pearson nowhere mentions Yule's work in his article (which was written during the summer of 1903) though he had commented on it as early as January 1903 (reference 107). Yule had left Pearson's department in 1900 and relations between them after this were less than cordial.

Pearson gave values of 0.33 to 0.41 for ‘family size’ 8 to ∞, rightly accepting the higher value. He did not recognise that on Mendel's hypothesis no restriction is imposed by family size (reference 106, p. 410 footnote).

Cited from Pearson K. , “On the ancestral gametic correlations of a Mendelian population mating at random”, Proceedings of the Royal Society (B), lxxxi (1909) 225–9.

Reference 105, p. 86.

Ibid., p. 54.

See, for example, Pearson K. , “On a criterion which may serve to test various theories of inheritance”, Proceedings of the Royal Society, lxxiii (1904) 262–80; and reference 113.

Yule G. U. , “On the theory of inheritance of quantitatively compound characters on the basis of Mendel's laws”, Report of Third International Conference on Genetics (Royal Horticultural Society, London, 1906). See also Pearson's remarks on the above in Biometrika, v (1907) 481–2. Richardson, in October 1902, may have been the first to suggest that incomplete dominance could be the means to make the ancestral hypothesis and Mendelism compatible. See Richardson H. , “Theories of heredity”, Nature, lxvi (1902) 630–1.

Reference 65, p. 44.

Weldon hypothesised the integrity of the chromosomes up to meiosis and then random separation, into two moieties, of the ‘unit determinants’ in the paternal and maternal chromosomes. He considered, correctly, that this could provide the biological synthesis of Mendelian and ancestrian ideas. After Weldon's death Pearson published this hypothesis from Weldon's notes and papers. See Pearson K. , “On the mathematical theory of determinantal inheritance from suggestions and notes of the late W. F. R. Weldon”, Biometrika, vi (1908) 80–93.

von Guita G. , “Versuche mit Kreuzungen von verschiedenen Rassen der Hausmaus”, Bericht der Naturforschenden Gesellschaft zu Freiburg i. Br., x (1898) 317–22; xi (1900) 131–8.

Darbishire A. D. , “Note on the results of crossing Japanese waltzing mice with European albino races”, Biometrika. ii (1902) 101–5.

Darbishire A. D. , “Second report on the result of crossing Japanese waltzing mice with European albino races”, Biometrika, ii (1903) 165–71.

Bateson W. , “Mendel's principles of heredity in mice”, Nature, lxvii (1903) 462–3.

Weldon W. F. R. , ibid., p. 512.

Bateson W. , ibid., pp. 585–6.

Letters in ibid., p. 610; lxviii (1903) 33–4.

Reference 73 (Beatrice Bateson), pp. 464–5.

Weldon W. F. R. , “Mr. Bateson's revisions of Mendel's theory of heredity”, Biometrika, ii (1903) 286–98.

Bateson W. , “The present state of knowledge of colour-heredity in mice and rats”, Proceedings of the Zoological Society of London, ii (1903) 71–99.

Darbishire A. D. , “Third report on hybrids between waltzing mice and albino races. On the result of crossing Japanese waltzing mice with ‘extracted» recessive albinos’, Biometrika, ii (1903) 282–5.

Darbishire A. D. , “On the result of crossing Japanese waltzing mice with albino mice”, Biometrika, iii (1904) 1–51. For contrary interpretations by Pearson and by the Mendelians, Bateson and Punnett, see reference 4.

Bateson W. , “Presidential address to the zoological section, British Association”, Reports of the British Association for the Advancement of Science, (1904) 574–89.

For reports of papers, see ibid., 590–3; and Nature, lxx (1904) 538–9.

Punnett R. C. , “Early days of genetics”, Heredity, iv (1950) 1–10.

K. P. to F. G. of 4 July 1909 (Life, IIIA, 387–8); and reference 118.

Hurst C. C. , “On the inheritance of coat-colour in horses”, Proceedings of the Royal Society (B), lxxvii (1906) 388–94.

W. F. R. W. to K. P. of November 1905 (reference 65, p. 47).

Weldon W. F. R. , “Note on the offspring of thoroughbred chestnut mares”, Proceedings of the Royal Society (B), lxxvii (1906) 394–8.

F. G. to K. P. of 16 April 1906 (Life, IIIA, 280).

Reference 73 (Beatrice Bateson), p. 103.

F. G. to ‘Milly’ of 21 April 1906 (Life, IIIB, 569).

K. P. to Mrs. W. F. R. W. of April 1906 (reference 63, p. 163).

K. P. to F. G. of June 1906 (Life, IIIA, 285).

K. P. to F. G. of 29 April 1906 (ibid., 282).

Reference 55, p. 241.

It is said that when R. A. Fisher submitted his classic synthesis (reference 106) to the Royal Society it was refereed by Pearson and Bateson. Both recommended rejection! See “The centenary of Mendel's discovery”, British medical journal for 1965, i, 327–8.

See our forthcoming article on the fate of the ancestral law, in Journal of medical genetics.