by Oren Harman
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 fr 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.
35. Samir Okasha, “Why Won’t the Group Selection Controversy Go Away?” British Journal of the Philosophy of Science 52 (2001), 25–50; Ayelet Shavit, “Shifting Values Partly Explain the Debate Over Group Selection,” Studies in History and Philosophy of Biological and Biomedical Sciences 35 (2004), 697–720, and One for All? Facts and Values in the Debates Over Group Selection (Jerusalem: Magnes Press, 2008) (in Hebrew).
36. David Sloan Wilson and Edward O. Wilson, “Rethinking the Theoretical Foundations of Sociobiology,” Quarterly Review of Biology 82 (2007), 327–48. See also Bert Hlldobler and E.O. Wilson, The Superorganism: The Beauty, Elegance, and Strangeness of Insect Societies (New York: W. W. Norton, 2009). Wilson, in fact, was never as opposed to group selection as either Dawkins or Williams—see Segerstrle, Defenders of the Truth, 37–38. Hlldobler, on the other hand, is not as convinced as Wilson of the importance of group selection. Dawkins, too, has yet to be won over, and the debate remains acrimonious. See his “The Group Delusion,” from January 10, 2009, at RichardDawkins.net. Also, see Ayelet Shavit and Roberta L. Millstein, “Group Selection is Dead! Long Live Group Selection?” BioScience 58 (2008), 574–75. For an interesting argument for group selection based on the plasticity of circadian clocks in bees, see Guy Bloch, “Plasticity in the Circadian Clock and the Temporal Organization of Insect Societies,” in Organization of Insect Societies, ed. Jrgen Gadau and Jenn
ifer Fewell (Cambridge, MA: Harvard University Press, 2009), 402–31.
37. Each differs in perspective. See S. A. Frank, “George Price’s Contributions to Evolutionary Genetics,” Journal of Theoretical Biology 175 (1995), 373–88, and “The Price Equation, Fisher’s Fundamental Theorem, Kin Selection, and Causal Analysis,” Evolution 51 (1997), 1712–29; Okasha, “Why Won’t the Group Selection Controversy Go Away?” A. Grafen, “Developments of the Price Equation and Natural Selection Under Uncertainty,” Proceedings of the Royal Society London B, 267 (2000), 1223–27, and “The First Formal Link Between the Price Equation and an Optimization Program,” Journal of Theoretical Biology 217 (2002), 75–91; but also see I. L. Heisler and J. Damuth, “A Method for Analyzing Selection in Hierarchically Structured Populations,” American Naturalist 130 (1987), 582–602, which provides a “contextualized” alternative to the Price equation.
38. Benjamin Kerr and Peter Godfrey-Smith, “Generalization of the Price Equation for Evolutionary Change,” Evolution 63, no. 2 (2009), 531–36.
39. The Price equation assumes fixed heritability, and is entirely devoid of mechanism. Though Frank decomposes it neatly, others are concerned that this precludes the interactivity inherent in nature, rendering it good exclusively for border cases. See Okasha, “Why Won’t the Group Selection Controversy Go Away?” for a thorough discussion of the relative merits of the equation. Also Massimo Pigliucci and Jonathan Kaplan, Making Sense of Evolution: The Conceptual Foundations of Evolutionary Biology (Chicago: University of Chicago Press, 2004), especially chapter 4, in which they discuss the relationship between statistical formalism and causal analysis. I thank Jim Griesemer for discussions on this point.
40. William B. Langdon, “Evolution of GP Populations: Price’s Selection and Covariance Theorem,” in Genetic Programming and Data Structures, (Norwell, MA: Kluwer Academic Press, 1998) 167–208; M. Van Veelen, “On the Use of the Price Equation,” Journal of Theoretical Biology 237 (2005), 412–26; T. Day, “Insights from Price’s Equation into Evolutionary Epidemiology,” DIMACS Series in Discrete Mathematics and Theoretical Computer Science 71 (2006), 23–43; J. W. Fox, “Using the Price Equation to Partition the Effects of Biodiversity Loss on Ecosystem Function,” Ecology 87 (2006), 2687–96; Stephen C. Stearns, “Are We Stalled Part Way Through a Major Evolutionary Transition from Individual to Group?” Evolution 61 (2007), 2275–80.
41. Darwin, The Origin of Species, 395.
42. Robert Trivers, Social Evolution (Menlo Park, CA: Benjamin/Cummings, 1985). For a defining volume on what became known as “evolutionary psychology” see Jerome H. Barkow, Leda Cosmides, and John Tooby, eds., The Adapted Mind: Evolutionary Psychology and the Generation of Culture (New York: Oxford University Press, 1992); Sarah F. Brosnan and Frans B. M. de Waal, “Monkeys Reject Unequal Pay,” Nature 425 (2003), 297–99. For a recent wide treatment of animal morality see Mark Bekoff and Jessica Pierce, Wild Justice: The Moral Lives of Animals (Chicago: University of Chicago Press, 2009).
43. R. Boyd and P. J. Richerson, “Punishment Allows the Evolution of Cooperation (or Anything Else) in Sizable Groups,” Ethology and Sociobiology 13 (1992), 171–95; E. Fehr and S. Gachter, “Altruistic Punishment in Humans,” Nature 415 (2002), 137–40; R. Boyd, H. Gintis, S. Bowles, and P. J. Richerson, “The Evolution of Altruistic Punishment,” Proceedings of the National Academy of Sciences of the United States of America 100 (2003), 3531–35; E. Fehr and U. Fischbacher, “The Nature of Human Altruism,” Nature 425 (2003), 785–91, are challenged by A. Dreber, D. G. Rand, D. Fudenberg, and M. A. Nowak, “Winners Don’t Punish,” Nature 452 (2008), 348–51.
44. See George Loewenstein, Scott Rick, and Jonathan D. Cohen, “Neuroeconomics,” Annual Review of Psychology 59 (2007), 647–72 for examples. Also, for a window into “positive evolutionary psychology,” see Dacher Keltner’s Born to Be Good: The Science of a Meaningful Life (New York: W. W. Norton, 2009), as well as new findings on cooperative infant behavior in Michael Tomassello, Why We Cooperate (Cambridge, MA: MIT Press, 2009).
45. Robert H. Frank, Passions Without Reason: The Strategic Role of the Emotions (New York: W. W. Norton, 1988). For an update of this argument from the field, see Frans B. M. de Waal, “Putting the Altruism Back into Altruism: The Evolution of Empathy,” Annual Review of Psychology 59 (2008), 279–300. For an update from neurophysiology see the work of Joshua D. Green et al., “An fMRI Investigation of Emotional Engagement in Moral Judgment,” Science 293 (2001), 2105–8, and “The Neural Bases of Cognitive Conflict and Control in Moral Judgment,” Neuron 44 (2004), 389–400, showing how emotion and moral reasoning relate in the brain. For a recent study on the importance of imitation as an empathy-building mechanism see A. Paukner, S. J. Suomi, E. Visalberghi, and P. F. Ferrari, “Capuchin Monkeys Display Affiliation Toward Humans Who Imitate Them,” Science 325 (2009), 880–83.
46. Robert Trivers, “The Elements of a Scientific Theory of Self-Deception,” Proceedings of the New York Academy of Science 187 (1999), 111–26.
47. C. Daniel Batson and Laura L. Shaw, “Evidence for Altruism: Toward a Pluralism of Prosocial Motives,” Psychology Inquiry 2 (1991), 107–22; Lise Wallach and Michael A. Wallach, “Why Altruism, Even Though It Exists, Cannot Be Demonstrated by Social Psychological Experiments,” Psychological Inquiry 2 (1991), 153–55; Sober and Wilson, Unto Others, presents the case for pluralism.
48. Charles Darwin, The Descent of Man, and Selection in Relation to Sex, selections and commentary by Carl Zimmer (New York: Plume, 2007), 157.
49. Peter J. Richerson and Robert Boyd, Not By Genes Alone: How Culture Transformed Human Evolution (Chicago: University of Chicago Press, 2005).
50. Trivers did not mean this as a criticism, but rather as a reflection of how others had viewed it. See Hamilton, Narrow Roads, 316; for an update of Hamilton’s argument, see Samuel Bowles, “Group Competition, Reproductive Leveling, and the Evolution of Human Altruism,” Science 314 (2006), 1569. Also J. Heirich, “Cultural Group Selection, Co-evolutionary Processes and Large Scale Cooperation,” Journal of Economic Behavior and Organization 53 (2004), 3–35.
51. Claudia Rutte and Michael Taborsky, “General Reciprocity in Rats,” PloS Biology 5 (2007), 196.
52. However, see David Sloan Wilson, Darwin’s Cathedral: Evolution, Religion, and the Nature of Society (Chicago: University of Chicago Press, 2002), for an argument showing religion functioning as a positive unifying system. From a neuropsychological angle, see also Jonathan Haidt, The Happiness Hypothesis: Finding the Truth in Ancient Wisdom (New York: Basic Books, 2005).
53. Peter Singer, The Expanding Circle: Ethics and Sociobiology (New York: Farrar, Straus & Giroux, 1981); Christopher Boehm, Hierarchy in the Forest: The Evolution of Egalitarian Behavior (Cambridge, MA: Harvard University Press, 1999), argues that we are laden with two conflicting evolutionary substrates: the foundational animal substrate reflecting hierarchy and status, and the overlaying cultural substrate allowing for genuine altruism. He finds this deeply embedded ambiguity in human nature both worrisome and hopeful. See also Stearns, “Are We Stalled Part Way Through a Major Evolutionary Transition from Individual to Group?” for an argument that humanity is stalled between individual and group selection.
54. J. Moll, F. Krueger, R. Zahn, M. Pardini, R. de Oliveira-Souza, and J. Grafman, “Human Fronto-mesolimbic Networks Guide Decisions About Charitable Donation,” Proceedings of the National Academy of Sciences 103 (2006), 15623–28.
55. Marc Hauser, Moral Minds: How Nature Designed Our Universal Sense of Right and Wrong (New York: Ecco, 2006); S. Anderson, H, Damasio, D. Tranel, and A. R. Damasio, “Impairment of Social and Moral Behavior Related to Early Damage in Human Prefrontal Cortex,” Nature Neuroscience 2 (1999), 1032–37. See also Marco Iacoboni, “Imitation, Empathy and Mirror Neurons,” Annual Review of Psychology 60 (2009), 653–70 for a presentation of the ways in which recent discoveries on the action of “mirror-neurons” complement cognitive models of imitation and social psychology studies
on empathy.
56. Zoe R. Donaldson and Larry J. Young, “Oxytocin, Vasopressin, and the Neurogenetics of Sociality,” Science 332 (2008), 900–904; S. Israel, E. Lerer, I. Shalev, F. Uzefovsky, M. Riebold, E. Laiba, R. Bachner-Melman, A. Maril, G. Bornstein, A. Knafo, and R. Ebstein, “The Oxytocin Receptor (OXTR) Contributes to Prosocial Fund Allocations in the Dictator Game and the Social Value Orientations Task,” PloS ONE 4 (2009), e5535.
57. “Academic Appeals to Oxbridge Students for an Egg Donor,” Daily Telegraph, December 4, 2008.