by Oren Harman
36. Ibid., 4.
37. Ibid., 12–13.
38. Ibid., 15, 5.
39. In fact Hamilton’s supervisor Cedric Smith had once introduced the two, but Maynard Smith couldn’t remember the encounter. Hamilton was shy and very unassuming, and Maynard Smith, like everyone else at the time, wasn’t interested in the genetics of altruism.
40. Ullica Segerstrle, Defenders of the Truth: The Sociobiology Debate (Oxford: Oxford University Press, 2000), 64.
41. The other two Oxford men at the meeting were the ethologist Niko Tinbergen and the geneticist Arthur Cain. Maynard Smith recalled that it was Cain who came up with the term “kin selection” though Cain himself had no such recollection.
42. John Maynard Smith, “Kin Selection and Group Selection,” Nature 201 (1964), 1145–47. Maynard Smith’s model became known as the “haystack model” and has excited a steady amount of discussion and criticism ever since. See Sober and Wilson, Unto Others, 67–71. For Wynne-Edwards’s own reply see Nature 201 (1964), 1147.
43. William Hamilton, “The Evolution of Altruistic Behavior,” American Naturalist 97 (1963), 354–56. Here Hamilton collapsed cost and benefit into one variable, k, and so the equation was even simpler: k > 1/r. This was the only math that appeared in the article.
44. Hamilton, Narrow Roads, 29–30.
45. Sober and Wilson, Unto Others, 35–36.
46. David Lack, Population Studies of Birds (Oxford: Clarendon Press, 1966).
47. G. C. Williams and D. C. Williams, “Natural Selection of Individually Harmful Social Adaptations among Sibs with Special Reference to Social Insects,” Evolution 11 (1957), 32–39.
48. Fisher, The Genetical Theory of Natural Selection, 158.
49. This formulation is taken from Sober and Wilson, Unto Others, 38–39.
50. George C. Williams, Adaptation and Natural Selection (Princeton: Princeton University Press, 1966), 273.
51. W. D. Hamilton, “Extraordinary Sex Ratios,” Science (156) 1967, 477–88.
52. Hamilton, Narrow Roads, 131–42.
53. Hamilton, “Extraordinary Sex Ratios,” 477.
54. The first to point this out was Robert Colwell, “Group Selection Is Implicated in the Evolution of Female-Biased Sex Ratios,” Nature 190 (1981), 401–4.
55. Hamilton, “Extraordinary Sex Ratios” see the section on “sex ratios and polygyny.” Hamilton did, however, make a mathematically imprecise allusion to the effect that certain optimal strategies may reflect a compromise between conflicting individual and group-based selection pressures. He buried it, however, in footnote 43.
56. Hamilton, Narrow Roads, 186. I thank Peter Henderson for a wonderful description of his time with Hamilton in the Amazon.
CHAPTER 8: NO EASY WAY
1. George Price letter to Hubert Humphrey, May 2, 1957, GPP.
2. George Price correspondence with Senator Hubert Humphrey, February 1956–December 1959, GPP; Donald S. Harrington, “To Deal with China Crisis; Adjudication by International Court of Justice Proposed,” New York Times, September 19, 1958.
3. George R. Price articles in Popular Science Monthly and THINK, March 1959–November 1960; George Price letters to Claude Shannon, October 11 and December 8, 1960; George Price–H. J. Muller correspondence, January–February 1960, GPP.
4. On Skinner see Daniel W. Bjork, B. F. Skinner: A Life (Washington, DC: American Psychological Association, 1997); William T. O’Donohue and Kyle E. Ferguson, The Psychology of B .F. Skinner (Thousand Oaks, CA: Sage, 2001); Daniel N. Weiner, B. F. Skinner: Benign Anarchist (Boston: Allyn and Bacon, 1996).
5. “Skinner’s Utopia: Panacea, or Path to Hell?” Time, September 20, 1971, 47–53, found in GPP. See also Philip J. Pauly, Controlling Life: Jacques Loeb and the Engineering Ideal in Biology (New York: Oxford University Press, 1987).
6. George R. Price, “The Teaching Machine,” THINK, March 1959, 10–14, quote on 10.
7. George Price letter to B. F. Skinner, October 27, 1957, GPP.
8. Alexis de Tocqueville, Democracy in America (1840), quoted in drafts of No Easy Way, chapter 6, GPP. Also George Price, “The Warning from the Distant Past,” draft, probably of chapter 10, GPP.
9. Cass Canfield letter to George Price, March 3, 1958, GPP; A. Boardman letter to George Price, August 11, 1966, GPP.
10. Ross Wetzsteon, Republic of Dreams: Greenwich Village, the American Bohemia, 1910–1960 (New York, Simon & Schuster, 2002); George Price passport, GPP.
11. George Price letter to Hubert Humphrey, December 8, 1959, GPP; George Price–Richard Winslow Correspondence, 1959–1961, GPP; No Easy Way draft, circa 1960, GPP.
12. George Price letter to Dr. Nathan S. Kline, October 28, 1958, GPP.
13. George Price letter to Richard Winslow, February 29, 1960, GPP.
14. George Price letter to Robert Latimer, February 13, 1961, GPP; Joan letter to George Price, November 12, 1961, GPP; George Price letter to Henry Noel, February 10, 1961, GPP; George Price letter to Al Somit, October 12, 1961, GPP; Fred Schneider letter to George Price, April 10, 1960, BLGPC, KPX1_1.5.
15. Donald Ferguson letter to George Price, September 21, 1960, GPP.
16. George R. Price, drafts of “Fallacies of Random Neural Networks and Self-Organization” and “A Theory of the Function of the Hymen,” GPP.
17. IBM Annual Report (1961), 7.
18. Ibid., 6; George Price letter to Donald Ferguson, October 28, 1960, GPP; George Price letter to Richard Winslow, October 15, 1962, GPP; George Price letter to Al Somit, June 19, 1962, GPP.
19. Graham DuShane (editor) letter to George Price, July 19, 1962, GPP; George Price letter to Ralph Graves, September 5, 1970, GPP.
20. Alice Avery Price letters to George Price, November 1962–November 1963, GPP.
21. B. F. Skinner, “The Science of Learning and the Art of Teaching,” Harvard Educational Review 24 (1954), 86–97.
22. George R. Price, “Some Suggestions About Programmed Instruction,” Market Requirements Memorandum, IBM, January 31, 1963, GPP.
23. George Price letter to Richard Winslow, December 17, 1963, GPP.
24. George Price letter to Emanuel Piore, March 17, 1969, GPP; George Price letter to John Isaacson, October 28, 1964, GPP; George Price letter to Stephen (last name not known), January 30, 1972, GPP.
25. George R. Price, “The Climate of Invention,” THINK, June 1959, 24.
26. George Price–Paul Samuelson correspondence, December 17, 1965–September 14, 1966, GPP; George R. Price, “Report on Marginal Optimization,” draft, IBM-DSD Poughkeepsie, January 1964, GPP.
27. George Price letter to Paul A. Samuelson, December 17, 1965, GPP. Bob Solow and Andreas Papandreou both became prominent economists.
28. Paul A. Samuelson letter to George Price, January 13, 1966, GPP.
29. George Price letters to Fred Brooks, October 12 and 21, 1964, GPP; George Price letter to Tatiana, May 31, 1964, GPP; George Price letter to Fairfield, August 6, 1964, GPP.
30. John C. Eccles letter to George Price, January 20, 1965, BL:KPX1_3.2, BLGPC.
31. Donald Ferguson letter to George Price, September 21, 1960, GPP; George Price letter to Howard Klevens, October 31, 1967, GPP.
32. George Price letter to Donald Ferguson, March 12, 1966, GPP.
33. George Price letter to Donald Ferguson, September 4, 1966, GPP; Donald Ferguson letter to George Price, January 2, 1967, GPP.
34. St. Hilda’s and St. Hugh’s “Reservation Agreement,” September 6, 1966, GPP; George Price letter to Annamarie Price, October 12, 1966, GPP; George Price–A. Boardman correspondence, August 26, 1966-June 12, 1967, GPP; Edison Price letter to George Price, September 11, 1970, GPP.
35. George Price letter to W. A. Brocker, July 11, 1967, GPP; George Price letter to Heinrich Kluver, July 8, 1967, BL:KPX1_1.4, BLGPC; George Price letter to Annamarie and Kathleen Price, November 13, 1967, GPP; George Price letter to Howard Klevens, October 31, 1967, GPP.
CHAPTER 9: LONDON
1. It w
ouldn’t last long, though. The boutique closed eight months after opening, a victim of local business interests and shoplifting. The Fool’s mural was removed by civic order, but not before Paul McCartney graffitied the windows with the name of the Beatles’ upcoming new hit, “Hey Jude.”
2. Andrew Loog Oldham, Stoned: A Memoir of London in the 1960s (London: St. Martin’s Press, 2001).
3. George Price letters to Tatiana, November 11 and December 12, 1967, GPP; George Price letter to Thomas Meyer, December 18, 1967, GPP.
4. Ibid.; George Price to Howard Klevens, October 31, 1967, GPP.
5. George Price letter to A. Boardman, December 25, 1967, GPP.
6. Ibid.
7. On the New Left in postwar England see Michael Kenny, The First New Left: British Intellectuals After Stalin (London: Lawrence and Wishart, 1995). Descriptions of the clashes at Grosvenor Square can be found in Ian Bone, Bash the Rich: True-Life Confessions of an Anarchist in the UK (London: Naked Guide Ltd., 2006) and in “Mick Farren—the Battle of Grosvenor Square London 1968” on YouTube.
8. Tatiana letter to George Price, December 30, 1967, GPP; George Price letter to Thomas Meyer, December 18, 1967, GPP; Edison Price letter to George Price, August 1, 1946, GPP; George Price letter to Alive Avery Price, February 27, 1968, GPP.
9. George Price letters to UCL librarian, December 12, 1967, and May 10, 1968, GPP; George Price letter to UCL Medical School Librarian, January 15, 1968, GPP; George Price letter to Thomas Meyer, December 18, 1967, GPP.
10. George Price letter to Bill Hamilton, March 5, 1968, GPP/BLGPC.
11. George Price letter to Thomas Meyer, December 18, 1967, and January 29, 1968, GPP; George Price letter to Kathleen Price, April 11, 1968, GPP.
12. Bill Hamilton letter to George Price, March 26, 1968, BL:KPX1_4.5.5.
13. Hamilton describes his own perception of George’s depressing revelation in Narrow Roads, 320.
14. Draft of an unsent letter from George Price to Bill Hamilton, March 26, 1968, BL:KPX1_4.5. 8.
15. The letter is addressed to “W.D. Hamilton, ? ?, Brazil.”
16. George Price letter to Kathleen, April 11, 1968, GPP.
17. Charles Darwin, The Descent of Man, chapter 17. Antlers are effective shields against other branching antlers, Darwin wrote, but would not protect against unbranched antlers projecting forward.
18. G. Stonehouse, “Thermoregulatory Function of Growing Antlers,” Nature 218 (1968), 870–72.
19. Draft of George Price, “Antlers, Intraspecific Combat, and Altruism,” 1–32, BLGPC.
20. Ibid., 16.
21. The precise description was this: “Basically, get even behaviour requires that animal A in conflict with B should hold in memory a measure, SB, of the seriousness of recent Category II (all-out combat) acts by B, and a measure, SA, of its own recent category II acts against B. Then the tendency, RAB, for A to retaliate against B is given by RAB = SB – SA. The strategy for A against a roughly equal opponent, B, can be stated as: ‘Fight as hard as possible at Category I level (deescalated). If SB > SA, retaliate at level II. If SB SA, fight at level I; except that when combat has been at level I for a long time, try a probe’” Ibid., 17.
22. Ibid., 19.
23. George did not consider that punishing exacts a price, and that free riders who fail to punish would therefore hold an advantage over those who did. This kind of free-riding game-theoretic thinking would occupy modelers in later years.
24. George Price letter to Bill Hamilton, August 3, 1968, GPP.
25. George Price letter to Thomas Meyer, October 14, 1968, GPP; John Orr declaration on Flat 3, 1a, Little Titchfield, July 30, 1968, GPP.
26. Bill Hamilton letter to George Price, August 29, 1968, GPP.
27. The condition was that there should be statistical association between the genotypes of donor and receiver; this is what is meant by “altruists finding one another”: organisms that share those genes that contribute to their behaving altruistically coming together in groups.
28. Strictly speaking, George had not shown how the covariance equation could translate into Hamilton’s kin-selection equation, replacing relatedness with association, but he understood immediately that this was now possible. For a formal derivation that does this, see Appendix 1: Covariance and Kin Selection.
29. George Price letter to Alice Avery Price, September 21, 1968, GPP.
30. George Price letter to Alice Avery Price, September 24, 1968, GPP.
31. George Price letter to Annamarie Price, October 2, 1968, GPP; George Price letter to Howard Klevens, October 9, 1968, GPP.
32. Kohn, A Reason for Everything, 221.
33. Steve Jones, “View from the Lab: A Mastermind for a Number of Reasons,” UK Telegraph, January 23, 2002.
34. Ibid.; Newton E. Morton, “Professor Cedric B. Smith, pioneer in statistical genetics: Died January 10, 2002, at the age of 84,” Genetic Epidemiology 22, no. 4 (2002), 283–84.
35. George Price letter to Annamarie Price, October 2, 1968; interview with Annamarie Price, April 13, 2008.
36. George Price, “Supplementary Details of Intended Research,” draft proposal to the Science Research Council, BLGPC, KPX_5.4.
37. A corollary of this system would be for males to take advantage of any mating opportunity outside of marriage, but not to provide for any children born. Thus, two distinct categories of female sexual partners would become recognized, with dichotomous treatment of each.
38. George Price letter to Ludwig Luft, January 29, 1969, GPP. George asked Luft, a German-born Jewish émigré to America, for the lyrics from Schumann’s Dichterliebe for a paper he was preparing on the evolution of love.
39. George Price letter to Edison Price, February 3, 1969, GPP; Edison Price letters to George, February 22 and March 2, 1969, GPP; George Price letter to Kathleen Price, March 5, 1969, GPP.
40. George Price letter to Kathleen Price, May 1, 1969, GPP.
41. Interview with Kathleen Price, April 13, 2008; George Price letter to Ludwig Luft, July 22, 1969, GPP.
42. George Price letter to Tatiana, May 11, 1969, GPP; George Price letter to Kathleen Price, March 5, 1969, GPP. In truth Edison had paid for George’s airfare from London and expenses in New York, but George was sour over an unpaid loan and little help with the apartment.
43. George Price letter to Ludwig Luft, July 22, 1969, GPP; George Price letter to Tatiana, May 11, 1969, GPP.
44. Garrett Hardin, “The Tragedy of the Commons,” Science 162 (1968), 1243–48.
45. The amateur mathematician was William Forster Lloyd (1794–1852) and the source, W. F. Lloyd, Two Lectures on the Checks to Population (Oxford: Oxford University Press, 1833), reprinted (in part) in Population, Evolution, and Birth Control, ed. Garret Hardin (San Francisco: Freeman, 1964), 37.
46. Hardin, “The Tragedy of the Commons,” 1244.
47. Sewall Wright letter to George Price, May 22, 1968, GPP.
48. George Price, “Supplementary Details of Intended Research,” draft proposal to the Science Research Council, BLGPC, KPX_5. 4. These observations would later be challenged in the ethological literature.
49. SRC grant proposal, written by Cedric Smith, March 31, 1969, and signed by the head of the department, Harry Harris, BLGPC, KPX1_10.1.
50. George Price letter to Tatiana, May 11, 1969, GPP.
51. John S. Price, “The Ritualization of Agonistic Behaviour as a Determinant of Variation along the Neuroticism/Stability Dimension of Personality,” Proceedings of the Royal Society of Medicine 62 (1969), 37–40.
52. Ibid., 39.
53. See S. A. Frank and M. Slatkin, “The Distribution of Allelic Effects under Mutation and Selection,” Genetics Research 55 (1990), 111–17.
54. See Appendix 2: The Full Price Equation and Levels of Selection for a derivation of the full Price equation and an explanation of how it can be partitioned and expanded.
55. Hamilton, Narrow Roads, 173.
56. W. D. Hamilton, “Selection of Selfish a
nd Altruistic Behaviour in Some Extreme Models,” in Man and Beast: Comparative Social Behavior, ed. J. F. Eisenberg and W. S. Dillon (Washington, DC: Smithsonian Press, 1971), 57–91.
57. For Hamilton’s entertaining description of the conference see Narrow Roads, 185–97; quotes on 187, 189. Hamilton’s reply, following his hero Fisher before him, was eugenics. In the long run the best humans would have to be consciously selected.
58. Ibid., 173.
59. George Price letter to Bill Hamilton, July 28, 1969, BLGPC, BL:KPX1_4. 4. The precise correction George alluded to was that in his 1964 paper Hamilton had failed to state that his kin-selection model only held for an indefinitely large population where the average relatedness between individuals was zero, whereas in realistic finite populations that wasn’t quite the case, and it was there that spite could arise.