The Price of Altruism (65 page)

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.

16.
Amotz Zehavi, “Mate Selection—a Selection for a Handicap,”
Journal of Theoretical Biology
, 53, 1975, 205–14, and Amotz Zahavi and Avishag Zahavi,
The Handicap Principle: A Missing Piece of Darwin’s Theory
(Oxford: Oxford University Press, 1997). See also Joseph Laporte, “Selection for Handicaps,”
Biology and Philosophy
, 16, 2001, 239–49.

17.
Michael Ghiselin,
The Economy of Nature and the Evolution of Sex
(Berkeley: University of California Press, 1974), 274.

18.
Richard Dawkins, “Twelve Misunderstandings of Kin Selection,”
Zeitschrift f
r Tierpsychologie
51 (1979), 184–200, quote on 190.

19.
Helena Cronin,
The Ant and the Peacock
(Cambridge: Cambridge University Press, 1991), 265.

20.
For a counterargument based on the rejection of sexual selection in favor of social selection, see Joan Roughgarden,
The Genial Gene: Deconstructing Darwinian Selfishness
(Los Angeles: University of California Press, 2009).

21.
Dawkins made a point of arguing in the 1989 edition of
The Selfish Gene
that his language was not merely metaphorical. In his next book,
The Extended Phenotype
(New York: Oxford University Press, 1982), he even went so far as to argue for the boundlessness of the individual organism as due to the all-importance of the gene. For a counterargument see Eva Jablonka, “From Replicators to Heritably Varying Phenotypic Traits: The Extended Phenotype Revisited,”
Biology and Philosophy
19 (2004), 353–75.

22.
Sober and Wilson,
Unto Others
; Samir Okasha,
Evolution and the Levels of Selection
(Oxford: Oxford University Press, 2006); David Sloan Wilson and Kevin M. Kniffin, “Altruism from an Evolutionary Perspective,” in
Research on Altruism and Love: An Annotated Bibliography of Major Studies in Psychology, Sociology, Evolutionary Biology, and Theology
, ed. Stephen G. Post, Byron Johnson, Michael E. McCullough, and Jeffrey P. Schloss (Philadelphia: Templeton Foundation Press, 2003), 117–36. It’s important to add that the exclusive role of genetic inheritance in evolution has been seriously challenged in the last decade. See in particular Eytan Avital and Eva Jablonka,
Animal Traditions: Behavioural Inheritance in Evolution
(Cambridge: Cambridge University Press, 2000), Eva Jablonka and Marion Lamb,
Evolution in Four Dimensions: Genetic, Epigenetic, Behavioral, and Symbolic Variation in the History of Life
(Cambridge, MA: MIT Press, 2005), and Scott F. Gilbert and David Epel,
Ecological Developmental Biology: Integrating Epigenetics, Medicine, and Evolution
(Sunderland: Sinauer Associates, 2008).

23.
Sober and Wilson,
Unto Others
, 332.

24.
Actually, the preferred term is “interactors,” since “vehicles” implies being under the control of the “replicators.” For a clear explanation of the interactor concept see David Hull, “Individuality and Selection,”
Annual Review of Ecology and Systematics
11 (1980), 311–32.

25.
For a recent illustration of this principle see Jeffrey A. Fletcher and Michael Doebeli, “A Simple and General Explanation for the Evolution of Altruism,”
Proceedings of the Royal Society of London B
276 (2009), 13–19.

26.
For a clear explanation of the errors in Maynard Smith’s “Haystack Model,” see Sober and Wilson,
Unto Others
, 67–71.

27.
W. D. Hamilton, “Innate Social Aptitudes in Man, An Approach from Evolutionary Genetics,” in
Biological Anthropology
, ed. Robin Fox (London: Malaby Press, 1975), 133–53.

28.
This is a game in which a pair constitutes a group, but N-person evolutionary game theory can handle groups comprised of any number. The advantage of modeling an N-person game, as opposed to a two-person game, is that it more closely resembles what happens in nature: Whereas in a two-person game altruism always incurs a cost to the altruist, in a many-player situation where many altruists are acting, altruism can benefit the group, including the altruists, which in turn means that it can more easily evolve.

29.
For a firsthand description of the vagaries of group selection, see a series of online, ongoing articles at the
Huffington Post
by David Sloan Wilson titled, “Truth and Reconciliation for Group Selection,” beginning December 27, 2008.

30.
This is the case whether or not one sees it as a general assault on “methodological individualism,” both from within biology and from other disciplines like psychology and sociology.

31.
R. K. Colwell, “Group Selection Is Implicated in the Evolution of Female-Biased Sex Ratio,”
Nature
190 (1981), 401–4.

32.
R. C. Lewontin, “The Units of Selection,”
Annual Review of Ecology and Systematics
1 (1970), 1–18, quote on 14–15. Lewontin, however, stopped short of implicating group selection directly.

33.
G. C. Williams and R. M. Nesse, “The Dawn of Darwinian Medicine,”
Quarterly Review of Biology
66 (1991), 1–22; Randolph M. Nesse and George C. Williams,
Why We Get Sick: The New Science of Darwinian Medicine
(New York: Times Books, 1994). For more recent appreciations see Wenda R. Trevathan, E. O. Smith, and James McKenna,
Evolutionary Medicine and Health: New Perspectives
(New York: Oxford University Press, 2007), and Stephen C. Stearns and Jacob C. Koella,
Evolution in Health and Disease
, 2nd ed. (New York: Oxford University Press, 2008).

34.
J. L. Brown, “Types of Group Selection,”
Nature
211 (1966), 870–71; M. J. Wade, “Group Selection Among Laboratory Populations of
Tribolium
,”
Proceedings of the National Academy of Science
73 (1976), 4604–7, and “An Experimental Study of Group Selection,”
Evolution
31 (1977), 134–53; D. S. Wilson, “The Group Selection Controversy: History and Current Status,”
Annual Review of Ecology and Systematics
14 (1983), 159–87; S. W. Rissing, G. B. Pollock, M. R. Higgins, R. H. Hagen, and D. R. Smith, “Foraging Specialization Without Relatedness or Dominance Among Co-Founding Ant Queens,”
Nature
338 (1989), 420–22; D. C. Queller, “Quantitative Genetics, Kin Selection, and Group Selection,”
American Naturalist
139 (1992), 540–58; Leticia Avilés, “Interdemic Selection and the Sex-Ration: A Social Spider Perspective,”
American Naturalist
142 (1993), 320–45; W. M. Muir, “Group Selection for Adaptation to Multiple-Hen Cages: Selection Program and Direct Responses,”
Poultry Science
75 (1995), 447–58; C. J. Goodnight and L. Stevens, “Experimental Studies of Group Selection: What Do They Tell Us About Group Selection in Nature?”
American Naturalist
150 (1997), 59–79; P. B. Rainey and K. Rainey, “Evolution of Cooperation and Conflict in Experimental Bacterial Populations,”
Nature
425 (2003), 72–74; P. J. Werfel and Y. Bar-Yam, “The Evolution of Reproductive Restraint Through Social Communication,”
Proceedings of the National Academy of Sciences of the United States of America
101 (2004), 11019–20; Benjamin Kerr, Claudia Neuhauser, Brendan Bohannan, and Antony Dean, “Local Migration Promotes Competitive Restraint in a Host–Pathogen ‘Tragedy of the Commons,’”
Nature
442 (2006), 75–78.