The Price of Altruism (55 page)

12.
Udney G. Yule had made the same claim back in 1902 but went unheeded. See his “Mendel’s Laws and Their Probable Relations to Intraracial Heredity,”
New Phytologist
1 (1902), 193–207, 222–38.

13.
Kohn,
A Reason for Everything,
142. See also Ronald W. Clark,
J. B. S.: The Life and Work of J. B. S. Haldane
(Oxford: Oxford University Press, 1968).

14.
See Martin Goodman,
Suffer and Survive: Gas Attacks, Miner’s Canaries, Spacesuits and the Bends: The Extreme Life of J. S. Haldane
(London: Pocket Books, 2007).

15.
On J. S. Haldane’s philosophy see S. Sturdy, “Biology as Social Theory: John Scott Haldane and Physiological Regulation,”
British Journal of the History of Science
21 (1988), 315–40. For J. S. Haldane’s own writings see
The Philosophical Basis of Biology
(Garden City, NY: Doubleday, 1931), and
The Philosophy of a Biologist
, 2nd ed. (Oxford: Clarendon Press, 1935).

16.
“I am enjoying life here very much,” he wrote to his father in February 1915. “I have got a most ripping job as a bomb officer.” To his mother he wrote: “I find this sort of fighting very enjoyable.”

17.
Haldane’s wartime paper showing the genetic linkage between albinism and pink eyes in mice was written with his sister Naomi and his friend A. D. Sprunt, who was killed in battle before it was published: J. B. S. Haldane, A. D. Sprunt, and N. H. Haldane, “Reduplication in Mice,”
Journal of Genetics
5 (1915), 133–35; J. S. Haldane is credited with inventing the gas mask that saved countless Allied lives in World War I.

18.
Kohn,
A Reason for Everything
, 149.

19.
“J.B.S. was against—against authority, and against the government, any authority and any government; if possible in the cause of reason; if not as a matter of principle,” Clark,
The Life and Work of J. B. S. Haldane
, 20.

20.
John Herschel,
Physical Geography
(Edinburgh: Adam and Charles Black, 1861), 12.

21.
For a perspective on Fisher’s statistical accomplishments and legacy see Anders Hald,
A History of Mathematical Statistics 1750 to 1930
(New York: Wiley, 1998).

22.
Edwards, “The Genetical Theory,” 1423.

23.
Interesting appreciations of
The Genetical Theory of Natural Selection
can be found in Edwards, “The Genetical Theory” James F. Crow, “R. A. Fisher, a Centennial View,”
Genetics
124 (1990), 207–11, and “Fisher’s Contributions to Genetics and Evolution,”
Journal of Theoretical Biology
38 (1990), 263–75; E. G. Leigh, Jr., “Ronald Fisher and the Development of Evolutionary Theory,” in
Oxford Surveys in Evolutionary Biology
, vol. 3, ed. Richard Dawkins and Mark Ridley (London: Oxford University Press, 1986), 187–223; Richard Lewontin, “Theoretical Population Genetics in the Evolutionary Synthesis,” in
The Evolutionary Synthesis: Perspectives on the Unification of Biology
, ed. Ernst Mayr and William B. Provine (Cambridge, MA: Harvard University Press, 1980), 58–68; and in the foreword to the Variorum Edition of
Fisher’s Genetical Theory of Natural Selection
, J. H. Bennett (Oxford: Oxford University Press, 1999). Quotation in Edwards, “The Genetical Theory,” 1422.

24.
Attempts to understand the fundamental theorem, and George Price’s solution, will be discussed in chapter 10.

25.
On Fisher’s notion of progress see Michael Ruse,
Monad to Man: The Concept of Progress in Evolutionary Biology
(Cambridge, MA: Harvard University Press, 1996), 295–303.

26.
Quoted in Kohn,
A Reason for Everything
, 96.

27.
Friedrich Nietzsche,
On the Genealogy of Morals
(1887), trans. Douglas Smith (Oxford: Oxford University Press, 1996); Charles Darwin,
M Notebook
, 1838.

28.
R. A. Fisher, “The Evolution of the Conscience in Civilized Communities,”
Eugenics Review
14 (1922), 190–93.

29.
R. A. Fisher, “The Renaissance of Darwinism,”
The Listener
37 (1947), 1001, 1009, quoted in Kohn,
A Reason for Everything
, 108; R. A. Fisher, “Indeterminism and Natural Selection,”
Philosophy of Science
1 (1934), 99–117.

30.
Cyril Darlington, “Recollections of Haldane” (draft), Darlington Papers (DP): C. 108: J. 86.

31.
Quoted in Clark,
The Life and Work of J. B. S. Haldane,
160.

32.
Ibid., 115.

33.
J. B. S. Haldane,
The Inequality of Man and Other Essays
(London: Chatto and Windus, 1932), quote on 121. On the culture and politics of British science at the time see William McGuken,
Scientists, Society, and State: The Social Relations of Science Movement in Great Britain 1931–1947
(Columbus: Ohio State University Press, 1984).

34.
J. B. S. Haldane,
Possible Worlds and Other Essays
(London: Harper and Brothers, 1928), 220–21. See also Charlotte Haldane’s
Truth Will Out
(London: Weidenfeld and Nicolson, 1949) for a colorful account of the couple’s life together.

35.
Haldane vacillated between attributing actual novel scientific discovery, as opposed to understanding, or knowing “what to look for,” as opposed to telling you what you “are going to find,” to the dialectical method. Compare “A Dialectical Account of Evolution,”
Science & Society
1 (1937), 473–86, to
The Marxist Philosophy and the Sciences
(London: Ayer 1939; reprint, Freeport, NY: Books for Libraries Press, 1969), 43. As for how he got to Marxism, Haldane cited English capitulation to fascism alongside “recent developments in physics and biology.”

36.
J. B. S. Haldane,
The Causes of Evolution
(London: Longmans, Green, 1932). The series of nine papers can be found in Mark Ridley,
Evolution,
CD-ROM (Oxford: Blackwell, 1996).

37.
Both Anthony Edwards and James Crow shared this view of Haldane as a mathematician: Kohn,
A Reason for Everything
, 145.

38.
To be fair, Haldane didn’t follow dialectical materialism blindly. Fisher argued that selection could produce modifier genes that would “negate” the effect of dominant deleterious mutations, a “beautifully dialectical theory” that Haldane rejected.

39.
For a critical rendition of the story of the
betularia
see Judith Hopper,
Of Moths and Men: The Untold Story of Science and the Peppered Moth
(New York: W. W. Norton, 2002).

40.
Scholars argue about the extent and manner in which dialectical materialism played a role in Haldane’s science. Arthur M. Shapiro shows that Haldane’s evolutionary papers in the 1920s became increasingly dialectical but puts it down to Hegel, via Uffer, rather than Marx. See his “Haldane, Marxism and the Conduct of Research,”
Quarterly Review of Biology
68 (1993), 69–77. Sahotra Sarkar argues that it was via his mechanistic and reductionist scientific work that Haldane became a Marxist, rather than his politics, or any form of philosophy, influencing his science. See his “Science, Philosophy, and Politics in the Work of J.B.S. Haldane, 1922–1937,”
Biology and Philosophy
7 (1992), 385–409.

41.
J. B. S. Haldane, “The Effect of Variation on Fitness,”
American Naturalist
71 (1937), 337–49, and “The Cost of Natural Selection,”
Journal of Genetics
55 (1957), 511–24. This line of thought later led to Motoo Kimura’s “neutral theory” of evolution, whereby natural selection is “blind” to much of the genetic mutation in a population, which therefore has little or no effect on population fitness.

42.
On Lysenko see David Joravsky,
The Lysenko Affair
(Cambridge, MA: Harvard University Press, 1970), Zhores Medvedev,
The Rise and Fall of T. D. Lysenko,
trans. Michael Lerner (New York: Columbia University Press, 1969), and Nils Roll-Hansen,
The Lysenko Affect: The Politics of Science
(New York: Humanity Books, 2004). A large literature is discussed in Oren Solomon Harman, “C. D. Darlington and the British and American Reaction to Lysenko and the Soviet Conception of Science,”
Journal of the History of Biology
36 (2003), 309–52. On Vavilov see Peter Pringle,
The Murder of Nikolai Vavilov: The Story of Stalin’s Persecution of One of the Great Scientists of the Twentieth Century
(New York: Simon & Schuster, 2008).

43.
Haldane,
The Inequality of Man
, 136; J. B. S. Haldane,
Heredity and Politics
(New York: W. W. Norton, 1938), 14.

44.
Boris Ephrussi said this of Haldane, quoted in Clark,
The Life and Work of J. B. S. Haldane
, 109.

45.
Ruse,
Monad
, 367.

46.
Sewall Wright, “The Genetical Theory of Natural Selection: A Review,”
Journal of Heredity
21 (1930), 340–56.

47.
See Sewall Wright, “Evolution in Mendelian Populations,”
Genetics
16 (1931), 97–159. Also, “The Roles of Mutation, Inbreeding, Crossbreeding and Selection in Evolution,”
Proceedings of the Sixth International Congress of Genetics
1 (1932), 356–66, and “Adaptation and Selection,” in Glenn L. Jepson, Ernst Mayr, and George Gaylord Simpson,
Genetics, Paleontology and Evolution
(Princeton: Princeton University Press, 1949), 365–89.

48.
Sewall Wright finally produced a book-size exposition of his complete evolutionary worldview late in life in
Evolution: Selected Papers
, ed. William B. Provine (Chicago: University of Chicago Press, 1986). There is also Wright,
Evolution and the Genetics of Populations
, 4 vols. (Chicago: University of Chicago Press, 1968).

49.
R. A. Fisher, “The Measurement of Selective Intensity,”
Proceedings of the Royal Society of London B
121 (1936), 58–62.

50.
It was T. H. Huxley’s grandson and Haldane’s old Eton friend Julian Huxley who gave the enterprise its name. See his
Evolution: The Modern Synthesis
(London: Allen and Unwin, 1942).

51.
The treatment of this problem came in the chapter “The Evolution of Distastefulness” in Fisher,
The Genetical Theory of Natural Selection
.

52.
Ibid., 159.

53.
Sewall Wright, “Coefficients of Inbreeding and Relationship,”
American Naturalist
56 (1922), 330–38.

54.
This exposition exists in Dugatkin,
The Altruism Equation
, 81–82.

55.
Sewall Wright, “Tempo and Mode in Evolution: A Critical Review,”
Ecology
26 (1945), 415–19.

56.
On group selection see Elliott Sober and David Sloan Wilson,
Unto Others: The Evolution and Psychology of Unselfish Behavior
(Cambridge, MA: Harvard University Press, 1998).