The Price of Altruism

by Oren Harman

  19. Lewontin would summarize his finding in the classic book The Genetic Basis of Evolutionary Change (New York: Columbia University Press, 1974).

  20. Charles Darwin, The Origin of Species, 84; Motoo Kimura original paper was “Evolutionary Rate at the Molecular Level,” Nature 217 (1968), 624–26. His major findings were later summarized in his book The Neutral Theory of Molecular Evolution (Cambridge: Cambridge University Press, 1983). There is much literature on the history of the neutralism debate, see in particular Michael Dietrich, “The Origins of the Neutral Theory of Molecular Evolution,” Journal of the History of Biology 27 (1994), 21–59, and James F. Crow, “Motoo Kimura and the Rise of Neutralism,” in Rebels, Mavericks, and Heretics in Biology, ed. Oren Harman and Michael Dietrich (New Haven: Yale University Press, 2008), 265–81.

  21. George viewed the genetic polymorphism work as a “digression” from the “Sex and Rapid Evolution” paper he was working on at the time but undertook it nevertheless since he felt it would help him better understand a point involved in the sex paper: George Price letter to John Maynard Smith, April 20, 1972; “Antlers” File, John Maynard Smith Papers, British Library (BLJMSC).

  22. Harry Harris, “Polymorphism and Protein Evolution: The Neural Mutation–Random Drift Hypothesis,” Journal of Medical Genetics 8, no. 4 (December 1971) 444–52; “A Theoretical Investigation into Genetic Polymorphism,” MRC application for a project grant, signed by Cedric A. B. Smith, January 31, 1973, BL:KPX1_10.2.

  23. George Price letter to Paul Samuelson, August 19, 1972, GPP. Samuelson had become interested in population genetics and written a short article on “The Hardy-Who Law of Genetics?” in March 1971, which he now sent to George.

  24. George Price letter to John Maynard Smith, April 20, 1972.

  25. The model used the following probabilities: Probability of serious injury from a single D play = 0. 10. Probability that a “Prober-Retaliator” will probe on the opening move or after opponent has played C = 0. 05. Probability that “Retaliator” or “Prober-Retaliator” will retaliate against a probe (if not injured) by opponent = 1. 0. Payoffs were calculated as follows: Payoff for winning = +60. Payoff for receiving serious injury = -100. Payoff for each D received that does not cause serious injury (a “scratch”) = -2. Payoff for saving time and energy (awarded to each contestant not seriously injured) varied from 0 for a contest of maximum length to +20 for a very short contest.

  26. George Price letter to Kathleen Price, August 5, 1972, GPP.

  27. George Price letter to John Maynard Smith, October 19, 1972, BLJMSC.

  28. Ibid.

  29. John Maynard Smith letter to George Price, October 24, 1972, BLJMSC; Bill Hamilton letter to George Price, November 22, 1972, BL:KPX1_4.14.

  30. George Price letter to Frieda (last name not known), May 29, 1971, GPP; Interview with Sam Berry, May 5, 2008.

  31. George Price letter to Dr. Webb, December 21, 1972, GPP; George Price letter to the Home Office, Immigration and Nationality Department, December 28, 1972, GPP.

  32. George Price letter to Annamarie Price, January 23, 1973, GPP.

  33. The Latin is: “Amicabilia quae sunt ad alterum vererunt amicabilibus quae sunt ad seipsum.” Aristotle, Nichomachean Ethics, book IX, chapter 4.

  34. Thomas Aquinas, The Summa Theologica of St. Thomas Aquinas 5 vols., trans. Fathers of the English Dominican Province (Westminster, MD: Christian Classics, 1981); David Hume, A Treatise of Human Nature (1739), ed. L. A. Selby-Bigge and P. H. Nidditch (Oxford: Clarendon Press, 1978), 521.

  35. Adam Smith, An Inquiry into the Nature and Causes of the Wealth of Nations, 2 vols., ed. R. H. Campbell and A. S. Skinner (Indianapolis: Liberty Fund, 1981), I.ii.2, 26–27; Bernard Mandeville, The Fable of the Bees: or, Private Vices, Publick Benefits, 4th ed. (London: J. Tonson, 1725), vol. 1, 9.

  36. Andrew Brown, “The Kindness of Strangers,” The Guardian, August 27, 2005; Roger Bingham, “Trivers in America,” Science 80 (March–April 1980), 56–67; communication with Robert Trivers, May 27, 2008.

  37. Robert Trivers, “The Evolution of Reciprocal Altruism,” Quarterly Review of Biology 46 (1971), 35–57. For a description of writing this paper see Trivers, Natural Selection and Social Theory (Oxford: Oxford University Press, 2002), 3–18.

  38. “A Theoretical Investigation,” MRC application for a project grant.

  39. Ibid.

  40. George Price letter to Bill Hamilton, May 3, 1971, BLGPC, BL:KPX1_5.9.

  41. George Price letter to Bill Hamilton, May 30, 1973, BLGPC, KPX1_2.4.

  42. George Price letter to Annamarie Price, March 3, 1973, GPP.

  43. George Price letter to Rosemarie Hudson, February 23, 1973, GPP; Henry Noel letter to George Price, January 30, 1973, GPP.

  44. George Price letter to Kathleen Price, March 24, 1973, GPP.

  45. George Price letter to Julia Price, March 10, 1973, GPP.

  46. George Price letter to Julia Price, March 24, 1973, GPP.

  47. Jack London, The People of the Abyss (London: Macmillan, 1903).

  48. See Exploring 20th Century London—Homelessness, Museum of London, www.20thcenturylondon.org.uk/server.php?show=conInformationRecord.77, for a well-researched, colorful exhibition. For a classic study of homelessness see Christopher Jencks, The Homeless (Cambridge, MA: Harvard University Press, 1994).

  49. George Price letter to Jack (surname unknown—a functionary at Saint Mark’s Church), August 18, 1973, GPP.

  50. John Price letter to George Price, April 5, 1973, GPP; Henry Noel letter to George Price, April 7, 1973, GPP; Tatiana letter to George Price, August 17, 1973, GPP; Edison Price letter to George Price, April 16, 1973, GPP.

  51. George Price to Jack, August 18, 1973.

  52. The Reverend R. F. H. Howarth letter to George Price, July 16, 1973, GPP; George Price letter to Henry Noel, July 22, 1973, GPP.

  53. George Price letter to Kathleen Price, March 19, 1973, GPP.

  CHAPTER 12: RECKONINGS

  1. UCL letter to George Price (signed by the assistant secretary of personnel), April 26, 1973, GPP; George Price letter to Julia Price, May 6, 1973, GPP.

  2. George Price letter to Mr. Norman Ingram-Smith, May 27, 1973, GPP.

  3. Ibid.

  4. George Price letter to Jack, August 18, 1973.

  5. Communication from Ursula Mittwoch, May 4, 2008; George Price letter to Dr. Gilfillan, July 3, 1973, GPP.

  6. Trevor Russell (Smoky) to George Price, July 3, 1973, GPP.

  7. Trevor Russell letter to George Price, July 9, 1973, GPP.

  8. Muriel Challenger letter to George Price, September 3, 1973, GPP.

  9. Julia Price letter to George Price, July 30, 1973, GPP.

  10. George Price letter to Smoky, September 14, 1973, GPP.

  11. Ibid.

  12. George Price letter to John Maynard Smith, October 19, 1973, BLJMSC, “Conflict Draft” folder.

  13. George Price letter to John Maynard Smith, February 12, 1973, BLJMSC, “Conflict Draft” folder.

  14. John Maynard Smith letter to George Price, October 24, 1972, BLJMSC, “Conflict Draft” folder. In fact Maynard Smith had been more than scrupulous and entirely generous both to have initially contacted George to get his permission to cite his unpublished antlers paper, and then in offering joint authorship. As for the 1964 Hamilton affair, Maynard Smith always had a conscience about it and did his best to set the record straight. He often remarked that he had a knack, blessed or cursed, for getting ideas from papers he reviewed.

  15. J. Maynard Smith and G. R. Price, “The Logic of Animal Conflict,” Nature 246 (1973), 15–18, quote on p. 15. Half a year later, in “On Fighting Strategies in Animal Combat,” Nature 250 (1974), 354, Valerius Geist of the University of Calgary accused John and George of not acknowledging his prior explication of the retaliation principle in V. Geist, “On the Evolution of Hornlike Organs,” Behaviour 27 (1966), 175. It had been an oversight that George and John were sorry about, and George wrote a letter of apology. See
George Price letter to V. Geist, March 24, 1974, BLJMSC, “Antlers File.”

  16. Al Somit letter to George Price, September 27, 1973, GPP.

  17. George Price letter to Edison Price, September 26, 1973, GPP.

  18. Cedric Smith note to George Price, November 7, 1973, GPP; Paul Garvey letter to George Price, November 11, 1973, GPP.

  19. George Price letter to Joan Jenkins, November 15, 1973. I thank Jim Schwartz for providing me with copies of the correspondence between Joan and George.

  20. “Monthly Message No. 202,” June 1974, London Healing Mission, GPP.

  21. George Price letter to Kathleen Price, December 6, 1973, GPP.

  22. George Price letter to Morris Goodman, June 7, 1974, GPP; phone interview with Professor Sam Berry, May 5, 2008.

  23. Interviews with Al Somit, December 6, 2007 and April 16, 2008, and communication on May 6, 2009.

  24. Al Somit letter to George Price, November 27, 1973, GPP.

  25. On the speculating bonanzas in London in the sixties see Oliver Marriot, The Property Boom (London: Hamish Hamilton, 1967); Nick Wates, The Battle for Tolmers Square (London: Routledge and Kegan Paul, 1976), 41.

  26. Wates, The Battle for Tolmers Square, 23, 24.

  27. Only 4 percent of the properties in the neighborhood were occupied by their owners, whereas 80 percent, three-quarters of which unfurnished, were being rented from a private landlord. Wates; ibid., 8, 9.

  28. Private Eye, February 9, 1973; Wates, The Battle for Tolmers Square, 43–73, quotes on 48, 68, 71, 43.

  29. Communication from Paul Nicholson, February 4, 2008; Wates, The Battle for Tolmers Square, 120.

  30. Wates, The Battle for Tolmers Square, 129; communication with Paul Nicholson, May 10, 2009.

  31. Communications with Ches Chesney, January 14 and 15, 2008.

  32. Paul Nicholson, “Room At the Top,” unpublished short story.

  33. Wates, The Battle for Tolmers Square, 162. Also see Nick Wates and Christian Wolmer, eds., Squatting: The Real Story (London: Bay Leaf Books, 1980).

  34. Communication with Paul Nicholson, February 4, 2008.

  35. Paul Nicholson, “Room At the Top.”

  36. Ibid.

  37. George Price letter to Dr. P. J. D. Heaf, December 16, 1973.

  38. George Price letter to the official solicitor, Royal Courts of Justice, January 30, 1974, GPP.

  39. James Schwartz, “Death of an Altruist,” Lingua Franca 10, no. 5 (July/August 2000), 51–61, quote on 59,

  40. Beginning in February, George had embarked on a project involving polymorphism analysis of macaque data from Morris Goodman, a professor of anatomy at Wayne State University, via Harris’s collaborator Dr. Nigel Barnicot, an anthropologist at UCL.

  41. Bill Hamilton letter to Dr. Kelly, undated (circa December 10, 1974), BLWHC, Z1X102_1.1.21; Hamilton, Narrow Roads, 174.

  42. George Price letter to Bill Hamilton, March 4, 1974, BLGPC, KPX1_2.5; George Price letter to Annamarie Price, April 27, 1974, GPP.

  43. George Price letter to Bill Hamilton, March 4, 1974, op. cit.

  44. Bill Hamilton letter to George Price, undated (circa March 6, 1974) BLGPC, BL:KPX1_4.12; Bill Hamilton letter To Whom It May Concern, on Imperial College letterhead, March 14, 1974, BLGPC KPX1_6 arrow KPX1_10.7.2.

  45. George Price letter to Bill Hamilton, May 18, 1974, BLGPC, KPX1_2.5.

  46. I thank Jim Schwartz for kindly sharing with me his notes from his meeting and conversations with Joan Jenkins from 2000. Joan Jenkins died shortly after. For her work on estrogen replacement therapy she was granted an OBE.

  47. George Price letter to Kathleen Price, January 25, 1974, GPP; George Price letter to Joan Jenkins, May 19, 1974; Schwartz notes.

  48. George Price letters to Joan Jenkins, May 20, June 6, June 9, 1974. Often the liaison between the two was Ruth Lang, CABS’s secretary at the Galton.

  49. George Price letter to Kathleen Price, January 25, 1974, GPP; interview with Kathleen Price, April 12, 2008.

  50. George Price letter to Kathleen Price, May 31, 1974, GPP.

  51. Bill Hamilton letter to George Price, June 11, 1974, BLGPC, BL:KPX1_4.6.; George Price letter to Bill Hamilton, August 21, 1974, BLGPC, KPX1_2.5.

  CHAPTER 13: ALTRUISM

  1. Immanuel Kant, Critique of Practical Reason and Other Works on the Theory of Ethics, trans. Thomas Kingsmill Abbott, 4th rev. ed. (London: Kongmans, Green and Co., 1889), 260.

  2. Williams, Adaptation and Natural Selection, 93.

  3. Richard Dawkins, The Selfish Gene (Oxford: Oxford University Press, 1996), 201, 254.

  4. Gene-centrism became wedded to “adaptationism,” and huddled together within a worldview called sociobiology, both were furiously attacked from different quarters. In particular see the anthropologist Marshall Sahlins, The Use and Abuse of Biology: An Anthropological Critique of Sociobiology (Ann Arbor: University of Michigan Press, 1976); the philosopher Mary Midgley, “Gene-Juggling,” Philosophy 54 (1979), 439–58; and the biologists Steven J. Gould and Richard R. Lewontin, “The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme,” Proceedings of the Royal Society of London B 205 (1979), 581–98.

  5. The notion of altruism as a mistake (due to imperfect design) is strengthened by the observation that some species of host birds have developed anticuckoo behavior. See A. Moksnes, E. Roskaft, A. T. Braa, L. Korsnes, H. M. Lampe, and H. C. Pedersen, “Behavioral Responses of Potential Hosts Towards Artificial Cuckoo Eggs and Dummies,” Behaviour 116 (1990), 64–89.

  6. P. W. Sherman, J. Jarvis, and R. Alexander, eds., The Biology of the Naked Mole-Rat (Princeton: Princeton University Press, 1991); D. W. Pfennig, H. K. Reeve, and P. W. Sherman, “Kin Recognition and Cannibalism in Spadefoot Toads,” Animal Behavior 46 (1993), 87–94; Robert B. Payne, Michael D. Sorenson, and Karen Klitz, The Cuckoos: Cuculidae (Oxford: Oxford University Press, 2005).

  7. John Tyler Bonner, The Social Amoebae: The Biology of Cellular Slime Molds (Prince ton: Princeton University Press, 2009).

  8. For a nontechnical description of this research program, see “Altruism among Amoebas,” Joan E. Strassman and David C. Queller, Natural History 116 (September 2007), 24–29.

  9. Martin Daly and Margo Wilson, The Truth about Cinderella: A Darwinian View of Parental Love (New Haven: Yale University Press, 1999). There has been, of course, much criticism of Daly and Wilson’s claims. For a study presenting contradictory data see H. Temrin, S. Buchmayer, and M. Enquist, “Step-Parents and Infanticide: New Data Contradict Evolutionary Predictions,” Proceedings of the Royal Society of London 267 (2000), 943–45.

  10. J. S. Gale and L. J. Evans showed that Retaliator was not, in fact, an ESS. See their “Logic of Animal Conflict,” Nature 254 (1975), 463–64.

  11. The name “Mouse” appeared only once in the literature—in George and Maynard Smith’s 1973 paper. From then on it was “Dove.”

  12. G. A. Parker and E. A. Thompson, “Dung Fly Struggles: A Test of the War of Attrition,” Behavioral Ecology and Sociobiology 7 (1980), 37–44; John Maynard Smith laid down the logic of games in evolution in his classic book Evolution and the Theory of Games (Cambridge: Cambridge University Press, 1982); for a good explanation of the limits and advantages of game theory for understanding natural evolution see G. A. Parker and J. Maynard Smith, “Optimality Theory in Evolutionary Biology,” Nature 348 (1990), 27–33.

  13. Amos Twersky, Daniel Kahneman, and Paul Slovic, eds., Judgment under Uncertainty: Heuristics and Biases (Cambridge: Cambridge University Press, 1980); Larry Samuelson, Evolutionary Games and Equilibrium Selection (Cambridge, MA: MIT Press, 1997), provides a technical account of how theoretical economists have adopted evolutionary ideas; Ran Shpigler, “The Invisible Hand and Natural Selection,” Odyssey 2 (2009), 45–51 (in Hebrew).

  14. Trivers, “The Evolution of Reciprocal Altruism,” and Trivers, Natural Selection and Social Theory, 3–18, 51–55; Robert Axelrod and William Hamilton, “The Evolution
of Cooperation,” Science 211 (1981), 1390–96.

  15. C. Packer, “Reciprocal Altruism in Papio anubis,” Nature 265 (1977), 441–43; G. S. Wilkinson, “Reciprocal Food Sharing in Vampire Bats,” Nature 309 (1984), 181–84; M. Melinski, “Tit for Tat in Sticklebacks and the Evolution of Cooperation,” Nature 325 (1987), 433–35; H. Godfray, “The Evolution of Forgiveness,” Nature 355 (1992), 206–7; A. H. Harcourt and F. B. M. de Waal, eds., Coalitions and Alliances in Humans and Other Animals (New York: Oxford University Press, 1992); S. Frank, “Mutual Policing and Repression of Competition in the Evolution of Cooperative Groups,” Nature 377 (1995), 520–22; T. H. Clotton-Brock and G. Parker, “Punishment in Animal Societies,” Nature 373 (1995), 209–16; Lee Alan Dugatkin, Cooperation Among Animals: An Evolutionary Perspective (New York: Oxford University Press, 1997); M. Nowak and K. Sigmund, “Evolution of Indirect Reciprocity by Image Scoring,” Nature 393 (1998), 573–77 and “The Dynamics of Indirect Reciprocity,” Journal of Theoretical Biology, 194 (1998), 561–74; G. Roberts and T. Sherratt, “Development of Cooperative Relationships Through Increasing Investment,” Nature 394 (1999), 175–79; J. L. Sachs, “The Evolution of Cooperation,” Quarterly Review of Biology 79 (2004), 135–60; Martin Nowak, “Five Rules for the Evolution of Cooperation,” Science 314 (2006), 1560–63; S. A. West, A. S. Griffin, and A. Gardner, “Social Semantics: Altruism, Cooperation, Mutualism, Strong Reciprocity and Group Selection,” Journal of Evolutionary Biology 20 (2007), 415–32. The above are choice citations.