The altruism equation: Seven scientists search for the origins of goodness

The Altruism Equation: Seven Scientists Search for the Origins of Goodness

Volumn , Issue , 2011, Pages

  Dugatkin, Lee Alan ^a  

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References (436)

1. H. M. Fraser, History of Beekeeping in Britain (London: Bee Research Association, 1958).
2. R. Huish, A Treatise on the Natural Economy and Practical Management of Bees (London: Baldwin, Cradock and Joy, 1815).
3. F. R. Prete, "The Conundrum of the Honey Bees," Journal of the History of Biology 23 (1990): 272-90.
4. J. Browne, Charles Darwin: The Power of Place (New York: Knopf, 2002), 203.
5. C. Darwin, On the Origin of Species, 1st ed. (London: J. Murray, 1859), 95, 90, 109.
6. C. Darwin, The Voyage of the H.M.S. Beagle (London: J. Murray, 1845).
7. R. Dawkins, The Selfish Gene (Oxford: Oxford University Press, 1976).
8. W. Kirby and W. Spence, Introduction to Entomology (London: Longman, Hurst, Rees, Orme and Brown, 1818).
9. R. J. Richards, Darwin and the Emergence of Evolutionary Theories of Mind and Behavior (Chicago: University of Chicago Press, 1987), 144.
10. Questions and Experiments notebook 5[a], 9, in P. Barrett, P. J. Guatrye, S. Herbert, D. Kohn, and S. Smith, eds., Charles Darwin's Notebooks: 1836-1844 (Ithaca, N.Y.: Cornell University Press, 1987).
11. May, 1872, in R. B. Freeman, "Charles Darwin on the Routes of Male Humble Bees," Bulletin of the British Museum (Natural History), Historical Series 3: 182.
12. Browne, Charles Darwin, 203.
13. Darwin, Origin, 236.
14. R. Stauffer, ed., Charles Darwin's Natural Selection, Being the Second Part of His Big Species Book (Cambridge: Cambridge University Press, 1975), 366.
15. Darwin, Origin, 241.
16. Darwin manuscript, June 1848, Darwin's Archive at Cambridge University, as cited in Richards, Darwin and the Emergence of Evolutionary Theories, 145, 149.
17. Richards, Darwin and the Emergence of Evolutionary Theories, 147.
18. William Youatt, Cattle: Their Breeds, Management and Disease (London: Library of Useful Knowledge, 1834).
19. Darwin, Origin, 238.
20. In Stauffer, Charles Darwin's Natural Selection, 370.
21. Richards, Darwin and the Emergence of Evolutionary Theory
22. also see Prete, "Conundrum of the Honey Bees."
23. Darwin to Huxley, July 3, 1860, in F. Burkhardt, D. Porter, et al., eds., The Correspondence of Charles Darwin (Cambridge: Cambridge University Press, 1993).
24. P. Kropotkin, In Russian and French Prisons (London: Ward and Downey, 1887).
25. T. H. Huxley, "The Struggle for Existence: A Programme," Nineteenth Century 23 (1888): 163-65.
26. P. Kropotkin, Mutual Aid, 3rd ed. introduction by George Woodcock (1902; New York: Black Rose Books, 1989), 75.
27. R. Clark, The Huxleys (London: Cox, Wyman, 1968).
28. T. H. Huxley, Autobiography and Selected Essays (Boston, Houghton Mifflin, 1909), 3.
29. A. Desmond, Huxley: From Devil's Disciple to Evolution's High Priest (New York: Addison Wesley, 1994), 3.
30. T. H. Huxley, Autobiography, 4.
31. L. Huxley, Life and Letters of Thomas Henry Huxley (New York: D. Appleton, 1901), 6.
32. T. H. Huxley, Autobiography, 5.
33. Desmond, Huxley, 9.
34. L. Huxley, Life and Letters, 16.
35. Desmond, Huxley, 36.
36. Introduction to T. H. Huxley, Autobiography, vi.
37. T. H. Huxley, Autobiography, 13.
38. Huxley to Darwin, November 23, 1859, in L. Huxley, Life and Letters.
39. Quoted in Desmond, Huxley, 361.
40. Huxley to Hooker, December 19, 1860, in L. Huxley, Life and Letters.
41. T. H. Huxley, "Evolution and Ethics: The Romanes Lecture," in J. Huxley, Touchstone for Ethics, 1893-1943 (New York: Harper and Brothers, 1947), 92.
42. Desmond, Huxley, 598.
43. T. H. Huxley, "The Struggle for Existence: A Programme." Nineteenth Century 23 (1888): 167, 165, 168, 166.
44. Theologically, Huxley was an agnostic, believing that one simply cannot know whether God exists; in fact, Huxley coined the term "agnostic" (from the Greek "not known"). Even if he had believed in God with any certainty, he no doubt had become quite angry with the Almighty about the time he wrote the gladiator article that so enraged Kropotkin.
45. The official cause of death was pneumonia, but she died at Charcot's Salpetrier Hospital (France) while being treated for her severe mental problems.
46. Draft of Manchester address 1887, as cited in Desmond, Huxley, 558n.64.
47. Huxley to Foster, January 8, 1888, in L. Huxley, Life and Letters.
48. T. Malthus, An Essay on the Principle of Population, as It Effects the Future Improvement of Society (1798; reprint, London: William Pickering, 1986), 1:4. Malthus wrote a much revised second edition in 1803: An Essay on the Principle of Population, or a View of Its Past and the Present Effects on Human Happiness with an Inquiry into Our Prospects Respecting the Future Removal or Mitigation of the Evils Which It Occasions.
49. Malthus, Essay on the Principle of Population, 5. Malthus, though he recognized that his principle of population applied to animals as well as humans, was clearly concerned with the latter, as evidenced by the subtitle of the revised version of his essay (see note 28).
50. C. Darwin, Autobiography (1876), ed. N. Barlow (London: Collins, 1958), 120.
51. C. Darwin, On the Origin of Species, 1st ed. (London: J. Murray, 1859), 63.
52. T. H. Huxley, On the Origin of Species, or The Causes of the Phenomena of Organic Nature (New York: D. Appleton, 1873), 123-24.
53. Huxley, "The Struggle for Existence," 166.
54. D. Todes, Darwin without Malthus: The Struggle for Existence in Russian Evolutionary Thought (New York, Oxford University Press, 1989), 20.
55. N. Riasanovsky, A History of Russia, 2nd ed. (New York: Oxford University Press, 1969), 5.
56. Todes, Darwin without Malthus, 23.
57. Kovalevskii cited and translated in A. Vucinich, "Russia: Biological Sciences," in T. Glick, ed., The Comparative Reception of Darwin (Austin: University of Texas Press, 1972).
58. P. N. Tkachev, "Nauka v poeziia v nauke," Sochinenie v dvukh tomakh 1: 398 (as cited in Todes, Darwin without Malthus).
59. See Todes, Darwin without Malthus, 25, for more on this.
60. K. Timiriazev, "Zemledelie i fizologiia rasteniia," Sochinenie 3: 31 (as cited in Todes, Darwin without Malthus).
61. P. Kropotkin, "The Scientific Bases of Anarchy," Nineteenth Century 21 (1887): 248.
62. N. Danilevskii, ed., Rossiia i Evropa (1869; New York: Johnson Reprint Co.), 146 (as cited in Todes, Darwin without Malthus).
63. Todes, Darwin without Malthus, 29.
64. Letter from Marx to F. Engels, June 18, 1862, Marx-Engels Internet Archive, http://www.marxists.org/archive/marx.
65. See Todes, Darwin without Malthus.
66. L. Tolstoy, What to Do?, trans. I. Hapgood (New York: Crowell and Co., 1887), 172.
67. R. F. Christian, ed., Tolstoy's Letters, vol. 2: 1880-1910 (New York: Charles Scribner's Sons, 1978), 717.
68. Darwin, Origin, 52. While this latter claim is often paraded as an example that struggle need not be taken in the literal sense, it is very clear from the totality of Darwin's arguments that he generally believed the struggle was a real one, a direct struggle between individuals for survival in a world of limited resources. Overpopulation led to this competition, which in turn led to adaptations and even new species.
69. Petr Kropotkin in a letter to Marie Goldsmith, April 7, 1915, Boris Nicolaevsky Collection, Hoover Institution on War, Resolution and Peace, Stanford University, file 7.
70. P. Kropotkin, "The Direct Action of Environment and Evolution," Nineteenth Century and After 85 (1919): 89.
71. P. Kropotkin, Memoirs of a Revolutionist (1899; rpt., New York: Grove Press, 1968), 5, 8, 28.
72. Kropotkin, Memoirs, 98.
73. G. Woodcock and I. Avakumovic, The Anarchist Prince: A Biographical Study of Peter Kropotkin (London: T. V. Boardman and Co., 1950), 53.
74. Kropotkin, Mutual Aid, xxxvii.
75. Woodcock and Avakumovic, The Anarchist Prince, 61.
76. Kropotkin, Memoirs, 198.
77. Kropotkin, Mutual Aid, xxxvi-xxxvii.
78. There is certainly still debate over the exact definitions of altruism. For most people who study social behavior, game theory has become the most common modeling technique used to study altruism and cooperation. Within game theory, both cooperation and altruism are defined as behaviors that entail a cost to the actor and a benefit to others. In other contexts, cooperation is defined strictly in terms of benefiting all parties involved, and thus entails a benefit, not a cost, assigned to the payoff of the cooperator.
79. Kropotkin, Mutual Aid,10-11.
80. Kropotkin, Memoirs, 216.
81. Kropotkin, Mutual Aid, 186, 76.
82. Kropotkin, Memoirs, 217.
83. P. Kropotkin, "Charles Darwin," Le Revolte, April 29, 1882.
84. Kropotkin, Mutual Aid, 4.
85. Kropotkin, "The Scientific Bases of Anarchy," 238.
86. Kropotkin, Mutual Aid, 186.
87. Kropotkin, "The Scientific Bases of Anarchy," 238-39.
88. Kropotkin, Memoirs, 403.
89. S. J. Gould, "Kropotkin Was No Crackpot," Natural History 7 (1988): 12-21.
90. Kropotkin in a letter to Marie Goldsmith, August 15, 1909, as cited in Todes, Darwin without Malthus, 123, my italics.
91. J. Huxley, T. H. Huxley's Dairy of the Voyage of the H.M.S. Rattlesnake (1935; New York: Kraus Reprint, 1972).
92. Kropotkin was also the geologist who correctly determined the directional orientation of the major mountain ranges in Asia.
93. Encyclopedia Britannica, Thomas Henry Huxley.
94. Desmond, Huxley, xvii.
95. T. H. Huxley, Man's Place in Nature, introduction by Stephen Jay Gould (New York: Random House, 2001), x. Originally published in 1863.
96. P. Marler and D. R. Griffin, "The 1973 Nobel Prize for Physiology or Medicine," Science 182 (1973): 464-66
97. R. A. Hinde, "Nobel Recognition for Ethology," Nature 245 (1973): 346.
98. D. Dewsbury, Studying Animal Behavior: Autobiographies of the Founders (Chicago: University of Chicago Press, 1985)
99. R. Burkhardt, Patterns of Behavior: Konrad Lorenz, Niko Tinbergen and the Founding of Ethology (Chicago: University of Chicago Press, 2005). When some experimental work on animal behavior began to emerge at the start of the twentieth century, it was rough going, with one early investigator lamenting that to keep his work on doves and pigeons afloat, financial circumstances might force him to eat some of his subjects. The researcher was Wallace Craig (see Burkhardt, Patterns of Behavior, 15).
100. Kropotkin, Mutual Aid, 10.
101. Pronounced AL-lee.
102. G. Mitman, The State of Nature: Ecology, Community and American Social Thought, 1900-1950 (Chicago: University of Chicago Press, 1992), 48.
103. Warder Clyde Allee Papers (WCAP), Special Collections, University of Chicago, box 26, folder 7.
104. K. P. Schmidt, "Warder Clyde Allee," Proceedings of the National Academy of Sciences, Biographical Memoirs 30 (1957): 3-40.
105. Allee was primarily a summer school student at first, teaching high school during the course of the academic year.
106. Mitman, The State of Nature, 30.
107. Whitman is credited with giving one of the first formal lectures on animal behavior at the Marine Biological Laboratory in 1898.
108. Mitman, The State of Nature, 7.
109. Frank Lillie, "The Department of Biology in Relation to the New Organization," Daily Maroon, December 11, 1930.
110. W. C. Allee, "An Experimental Analysis of the Relation between Physiological States and Rheotaxis in Isopoda," Journal of Experimental Zoology 13 (1912): 270-344.
111. Obituary for Majorie Hill Allee, WCAP, box 25, folder 3.
112. M. H. Allee, Jane's Island (New York: Houghton Mifflin, 1931).
113. W. C. Allee and M. H. Allee, Jungle Island (New York: Rand McNally, 1925).
114. Schmidt, "Warder Clyde Allee," 7.
115. Mitman, The State of Nature, 46 n.103.
116. W. C. Allee, "Animal Aggregations," Quarterly Review of Biology 2 (1927): 367-98.
117. W. C. Allee, Animal Aggregations: A Study in General Sociology (Chicago: University of Chicago Press, 1927). A second edition appeared in 1931.
118. Allee, "Animal Aggregations," 379.
119. W. C. Allee, Cooperation among Animals with Human Implications (New York: Henry Schuman, 1951), 26.
120. A. Popovici-Baznosanu, "L'influence de quelques sur l'accroissement des Gastropodes d'eau douce," Archives de Zoologie Experimentale et Générale 60 (1921): 501-21.
121. Allee, Cooperation among Animals.
122. T. B. Robertson, "Experimental Studies on Cellular Reproduction: II. The Influence of Mutual Contiguity upon Reproductive Rate in Infusoria," Biochemical Journal 15 (1921): 612-19.
123. Allee, "Animal Aggregations," 385.
124. V. L. Kellogg, Headquarters Nights (Boston: Atlantic Monthly Press, 1917).
125. Watson to Allee, December 29, 1949, WCAP, box 21, folder 4; and Allee to Watson, December 31, 1949, WCAP, box 21, folder 4.
126. Springfield News-Record, February 9, 1917.
127. Quoted in E. Banks, "Warder Clyde Allee and the Chicago School of Animal Behavior," Journal of the History of Behavioral Science 21 (1985): 351.
128. Only the draft version of this article is in Allee's files, but it appears to have been aimed at a Quaker audience.
129. W. C. Allee, "Reexamination of One Fundamental Doctrine in the Light of Modern Knowledge," WCAP, box 26, folder 3.
130. W. C. Allee, "Concerning the Biology of War," June 1940, WCAP, box 2, folder 7.
131. Allee, "Animal Aggregations," 387, 379.
132. For background information on Sewall Wright, see the American Philosophical Society at http://www.amphilsoc.org/library/mole/w/wrights.htm.
133. W. Provine, The Origins of Theoretical Population Genetics (Chicago: University of Chicago Press, 1971)
134. Sewall Wright and Evolutionary Biology (Chicago: University of Chicago Press, 1986).
135. S. Wright, "Tempo and Mode in Evolution: A Critical Review," Ecology 26 (1945): 415-19.
136. D. S. Wilson, "The Group Selection Controversy: History and Current Status," Annual Review of Ecology & Systematics 14 (1983): 159-87.
137. D. Wilson, "Structured Demes and the Evolution of Group-Advantageous Traits," American Naturalist 111 (1977): 157-85
138. D. S. Wilson, The Natural Selection of Populations and Communities (Menlo Park, Calif.: Benjamin Cummings, 1980)
139. R. Dawkins, The Selfish Gene (Oxford: Oxford University Press, 1976)
140. R. Dawkins, "Twelve Misunderstandings of Kin Selection," Zeitscrift fur Tierpsychologie 51 (1979): 184-200
141. E. Sober and D. S. Wilson, Unto Others (Cambridge, Mass.: Harvard University Press, 1998).
142. D. S. Wilson, The Natural Selection of Populations and Communities (Menlo Park, Calif.: Benjamin Cummings, 1980)
143. L. A. Dugatkin and H. K. Reeve, "Behavioral Ecology and Levels of Selection: Dissolving the Group Selection Controversy," Advances in the Study of Behaviour 23 (1994): 101-33
144. D. C. Queller, "Group Selection and Kin Selection," Trends in Ecology & Evolution 6 (1991): 64.
145. S. Wright, "Size of Population and Breeding Structure in Relation to Evolution," Science 87 (1938): 430-31
146. Wright, "Tempo and Mode in Evolution."
147. Allee to Julian Huxley, April 4, 1939, WCAP, box 18, folder 15.
148. Mitman, The State of Nature, 13.
149. Allee, "Animal Aggregations."
150. Allee, Animal Aggregations, 338.
151. Allee, Cooperation among Animals, 28.
152. Allee, Animal Aggregations, 339-40.
153. Allee, "Animal Aggregations," 390.
154. Mitman, The State of Nature, 81-84. While most anthropologists saw their views on kinship and cooperation as antithetical to Allee's, there were exceptions, one of whom was his friend Ralph Linton, author of The Study of Man. Linton's theories seemed eerily reminiscent of Allee's ideas on family life and cooperation, just cast in anthropological terms. It is possible that Allee and Linton met at the Hyde Park gatherings of the Society of Friends, a Quaker group with which Allee was associated for decades.
155. R. Linton, The Study of Man: An Introduction (New York: Appleton, 1936)
156. A. Linton and C. Wagley, Ralph Linton (New York: Columbia University Press, 1971).
157. Funded indirectly by the Rockefeller Foundation.
158. W. C. Allee, "Where Angels Fear to Tread: A Contribution from General Sociology to Human Ethics," Science 97 (1943): 517-25.
159. W. C. Allee, "Cooperation among Animals," University of Chicago Magazine, June 1928, 424.
160. Allee, Animal Aggregations, 362.
161. Mitman, The State of Nature, 84.
162. Allee, "Where Angels Fear to Tread," 524.
163. Newman to Allee, November 17, 1953, WCAP, box 9, folder 4.
164. "Concerning Biology and Biologists" WCAP, box 9, folder 4.
165. Allee, Animal Aggregations, 355.
166. Allee, Cooperation among Animals, 213.
167. October 26, 1946, WCAP, box 2, folder 6.
168. Allee to Phillips, February 13, 1933, WCAP, box 21, folder 3.
169. Mitman, The State of Nature, 139.
170. Griggs to Allee, January 31 and March 31, 1947, WCAP, box 2, folder 6.
171. WCAP, box 26, folder 7.
172. J.B.S. Haldane and J. Huxley, Animal Biology (Oxford: Claredon, 1927).
173. J.B.S. Haldane, Possible Worlds (New York: Harper Brothers, 1928), 298.
174. For a fascinating history of Haldane's "inordinate fondness" quote, see S. J. Gould, "A Special Fondness for Beetles," Natural History, January 1993, 4-11. Two other published versions of this story have Haldane noting: "the creator would appear as endowed with a passion for stars, on the one hand, and beetles on the other, for the simple reason that there are nearly 300,000 species of beetles known"
175. J.B.S. Haldane, What Is Life? [New York: Boni and Gaer, 1947], 239 and "the creator, if he exists, has a special preference for beetles, and so we might be more likely to meet them than any other type of animal on a planet that would support life"
176. A. Clarke, "Haldane and Space," in Haldane and Modern Biology, ed. K. Dronamraju [Baltimore: Johns Hopkins University Press, 1968], 244.
177. K. Dronamraju, Haldane: The Life and Work of J.B.S. Haldane with Special Reference to India (Aberdeen: Aberdeen University Press, 1985), xv.
178. Clarke, "Haldane and Space," 248.
179. N. Pirie, "John Burdon Sanderson Haldane: 1892-1964," Biographical Memoirs of Fellows of the Royal Society 12 (1966): 219-49. Included in Haldane's popular writings were almost 350 articles he wrote during his tenure as science correspondent for the communist paper the Daily Worker.
180. Pirie, "John Burdon Sanderson Haldane," 219, quoted from J.B.S. Haldane
181. "The Scientific Work of J. S. Haldane," Penguin Science Survey 2 (1961): 11.
182. Haldane as quoted in R. Clark, JBS: The Life and Work of J.B.S. Haldane (London: Hodder and Stoughton, 1968), 13.
183. Haldane and Huxley, Animal Biology.
184. As quoted in Clark, JBS, 22.
185. For more on Goodrich, see M. Ruse, From Monads to Man (Cambridge, Mass.: Harvard University Press, 1996).
186. Haldane as quoted in Clark, JBS, 27.
187. Clark, JBS, 37.
188. Pirie, "John Burdon Sanderson Haldane," 219
189. J.B.S. quote from J.B.S. Haldane, The Inequality of Man and Other Essays (London: Chatto and Windus, 1941).
190. Haldane as quoted in Clark, JBS, 39.
191. J.B.S. Haldane, A. D. Sprunt, and M. N. Haldane, "Reduplication in Mice," Journal of Genetics 5 (1915): 133-35.
192. John Maynard Smith, Haldane, J.B.S. Science and Life: Essays of a Rationalist (London: Humanist Library, 1968), ix.
193. Haldane as quoted in Clark, JBS, 65, 63.
194. Bateson quoted in Clark, JBS, 79.
195. Clark, JBS, 80-82.
196. Much of the work in these papers is summarized for the lay reader in J.B.S. Haldane, The Causes of Evolution (London: Longmans Green, 1932).
197. Haldane, Causes of Evolution.
198. E. Mayr, "Haldane's Causes of Evolution after 60 Years," American Naturalist 67 (1992): 175-86
199. J. Maynard Smith, "In Haldane's Footsteps," 347-56 in Studying Animal Behavior: Autobiographies of the Founders ed. D. Dewsbury (Chicago: University of Chicago Press, 1985).
200. Haldane, "A Mathematical Theory of Natural and Artificial Selection, Part II" Proceedings of the Cambridge Philosophical Society 1 (1924): 158.
201. Haldane, Possible Worlds, 35.
202. For more on Haldane's view of the power of science, see Ruse, From Monads to Man, as well as Haldane, Possible Worlds
203. J.B.S. Haldane, Science Advances (New York: Macmillan, 1948); and Haldane, Science and Life.
204. Clarke, "Haldane and Space," 138.
205. J.B.S. Haldane, "Population Genetics," New Biology 18 (1955): 34-51.
206. Haldane, Causes of Evolution, 1.
207. In the appendix to Causes, Haldane does in fact go beyond a verbal model of altruism, providing the mathematically trained reader with some equations. The model, however, does not consider the effects of kinship per se, and rather generally demonstrates that altruism will be rare in large populations, and by implication, rare in nonkinbased populations.
208. Haldane, "Population Genetics," 44.
209. Haldane, Causes of Evolution, 71, 120.
210. R. A. Fisher, "On the Dominance Ratio," Proceedings of the Royal Society of Edinburgh 42 (1922): 321-22.
211. R. A. Fisher, The Genetical Theory of Natural Selection (Oxford: Claredon Press, 1930).
212. R. A. Fisher, "The Correlation between Relatives on the Supposition of Mendelian Inheritance," Transactions of the Royal Society of Edinburgh 52 (1918): 399-433.
213. His original table is actually a seven-by-five matrix with thirtyfive entries, but for ease of presentation, I have deleted one column, reducing this to a six-by-five matrix.
214. W. Provine, The Origins of Theoretical Population Genetics (Chicago: University of Chicago Press, 1971), 140.
215. T. Eisner and D. Aneshansley, "Spray aiming in the bombardier beetle: photographic evidence," Proceedings of the National Academy of Sciences 96 (1999): 9705-9.
216. Fisher, Genetical Theory of Natural Selection, 158.
217. S. Wright, "Systems of Mating, I: The Biometric Relation between Parent and Offspring," Genetics 6 (1921): 111-23.
218. S. Wright, "Coefficients of Inbreeding and Relationship," American Naturalist 56 (1922): 330-38.
219. This idea was suggested by an anonymous review of an early version of this book.
220. For example, Fisher, "Correlation between Relatives"; Wright, "Systems of Mating";
221. Haldane, "A Mathematical Theory of Natural and Artificial Selection, Part II," Transactions of the Cambridge Philosophical Society 23 (1924).
222. Author's interview with Thomas Seeley. Ithaca, New York, January 31, 2003. A similar opinion was expressed by Harvard's Naomi Pierce in an interview with me on February 24, 2003.
223. Author's interview with Kern Reeve. Ithaca, New York, January 31, 2003.
224. J. Maynard Smith, "J.B.S. Haldane," Oxford Surveys in Evolutionary Biology 4 (1987): 9.
225. This idea was also suggested by an anonymous review of an early version of this book.
226. Personal communication from Paul Ewald.
227. W. D. Hamilton, Narrow Roads of Gene Land: The Collected Papers of W. D. Hamilton, vol. 1 (Oxford: W. H. Freeman, 1996), viii-ix.
228. The Callender-Hamilton bridge design remains on the market to this day. For more on Archibald Hamilton's work, see A. M. Hamilton, Road through Kurdistan: The Narrative of an Engineer in Iraq (New York: AMS Press, 1937).
229. Author's interview with Mary Bliss, Bill Hamilton's sister, Sevenoaks, Kent, England, March 9, 2003.
230. A. Grafen, "William David Hamilton," Biographical Memoirs of the Fellows of the Royal Society of London 50 (2004): 109-32.
231. M. Bliss, "Professor William Donald Hamilton: Memoirs of My Brother's Youth," speech read at the memorial ceremony for W. D. Hamilton in the chapel of the New College, London, July 1, 2000.
232. Hamilton's lecture when accepting the prestigious Kyoto Prize, British Museum, Hamilton Archive.
233. For an excellent review of the life and work of Francis Galton, see M. G. Bulmer, Francis Galton: Pioneer of Heredity and Biometry (Baltimore: Johns Hopkins University Press, 2003).
234. Hamilton in The Secret of the Clouds videotape. British Library. Originally aired on "The Edge Show."
235. Kyoto Prize lecture.
236. Bliss, "Professor William Donald Hamilton."
237. E-mail message from Mary Bliss to author, April 18, 2003.
238. British Library, BL:z1X55 17.5, June 3 and 6, July 18, 1956. This journal runs from June 3, 1956, to May 27, 1957, and is 146 pages in length.
239. Hamilton, Narrow Roads, 1:21.
240. W. D. Hamilton, Narrow Roads of Gene Land: The Collected Papers of W. D. Hamilton, vol. 2 (Oxford: Oxford University Press, 2001), 543, 542.
241. From Hamilton's endorsement of the 1999 Oxford University edition of The Genetical Theory of Natural Selection.
242. Despite the fact that Fisher's book had a profound effect on him while he studied at Cambridge, Hamilton notes that he did not have Fisher, or for that matter, Haldane's verbal model for the evolution of altruism "consciously in mind" during the early development of his own ideas on this subject. "One reads and forgets," Hamilton recounted. "Hints not understood probably leave their traces, but one has to return to the topic in a better state of preparation and to re-read before such throwaway items become meaningful." Narrow Roads, 1:22.
243. Who would one day be knighted.
244. Some fifteen years later, Leach would become one of the most severe critics of E. O. Wilson's Sociobiology: The New Synthesis (Cambridge, Mass.: Harvard University Press, 1975).
245. Hamilton, Narrow Roads, 1:24.
246. Bill Hamilton to Mary Bliss, November 11, 1959, cited in Bliss, "Professor William David Hamilton."
247. In so doing, Hamilton was following in the footsteps of Ronald Fisher, who was a schoolteacher early in his career.
248. Hamilton, Narrow Roads, 1:24.
249. Bliss, "Professor William Donald Hamilton," citing a February 8, 1960, letter from Bill Hamilton.
250. Hamilton, Narrow Roads, 1:11.
251. In fact, during his early days in graduate school, Hamilton once again considered an alternative career path-this time, as a carpenter. In the end, though, his obsession with evolution, kinship, and altruism would overpower any ideas about other careers.
252. Hamilton, Narrow Roads, 2:543.
253. W. D. Hamilton, "The Evolution of Altruistic Behavior," American Naturalist 97 (1963): 354.
254. Although the argument can get technical, Hamilton later claimed that Wright's coefficient of relatedness was not the perfect variable to use in his model. He would come to recognize, with the help of Richard Michod, that his equation "requires, in principle, a regression coefficient of genotype of recipient on genotype of altruist, whereas Wright's coefficient is the corresponding correlation coefficient." R. E. Michod and W. D. Hamilton, "Coefficients of Relatedness in Sociobiology," Nature 288 (1980): 694-97.
255. In his 1963 paper, Hamilton folded the costs and benefits into a single variable, which he labeled k-where k is simply the benefits divided by the costs, or b/c.
256. I thank Oxford University's Alan Grafen for emphasizing the importance of economics in Hamilton's model.
257. This is a slightly different way of calculating things than Haldane adopted in the New Biology paper.
258. For instance, it applied only to haploid organisms, and to the case where the altruistic allele was rare.
259. Hamilton, Narrow Roads, 1:230.
260. J. Maynard Smith, "J.B.S. Haldane," Oxford Surveys in Evolutionary Biology 4 (1987): 9.
261. J. Maynard Smith, "Group Selection and Kin selection," Nature 201 (1964): 1,145-46.
262. V. C. Wynne-Edwards, Animal Dispersion in Relation to Social Behavior (Edinburgh: Oliver and Boyd, 1962).
263. Maynard Smith, "Group Selection," 1,145.
264. W. D. Hamilton, "The Genetical Evolution of Social Behaviour, I and II," Journal of Theoretical Biology 7 (1964): 1-52.
265. In "Death of an Altruist," Maynard Smith told Jim Schwartz, "I seem to have this fate of getting ideas from other people's manuscripts when I referee them." With respect to the Hamilton Journal of Theoretical Biology papers, Maynard Smith noted, "I wasn't trying to steal his idea, or I don't think I was, so it wasn't conscious." J. Schwartz, "Death of an Altruist," Lingua Franca 10 (2000): 51-61.
266. Author's interview with Naomi Pierce, Harvard University, Cambridge, Mass., February 25, 2003.
267. Hamilton, "The Genetical Evolution of Social Behaviour," 2.
268. In this paper, Hamilton used his address at University College rather than the London School of Economics: W. D. Hamilton, The Galton Laboratory, University College, London, WC2.
269. Hamilton, "The Genetical Evolution of Social Behaviour," 8.
270. It could hardly have been otherwise; recall that the American Naturalist paper (1963), though published before the Journal of Theoretical Biology papers (1964), was in fact written after Hamilton had derived the models in the 1964 paper.
271. Hamilton, "The Genetical Evolution of Social Behaviour," 19.
272. Work over the past forty years has generally supported Hamilton's claim that the donor and recipient in grooming bouts will often be related, but other factors, such as the degree of reciprocity between donor and recipient, are often in play as well.
273. Hamilton, "The Genetical Evolution of Social Behavior," 21.
274. R. Gadagkar, "Evolution of Eusociality: The Advantages of Assured Fitness Returns," Philosophical Transactions of the Royal Society of London 329 (1990): 17-25
275. D. C. Queller, "The Evolution of Eusociality: Reproductive Head Starts of Workers," Proceedings of the National Academy of Sciences 86 (1989): 3,224-26
276. E. O. Wilson and B. Holldobler, "The Rise of the Ants: A Phylogentic and Ecological Explanation," Proceedings of the National Academy of Sciences 102 (2005): 7,411-14.
277. These journals are currently housed in the British Library. The first (British Library: Z1X55 17.1) runs from September 12, 1963, through November 11, 1963. The second (BL: Z1X55 17.14) runs from November 12, 1963, through June 1, 1964. The third starts on June 1, 1964, and ends on July 26, 1964. The address on the journals is Sr. W. D. Hamilton, Faculdade de Filosofia Ciencias E Letra De Rio Claro, Rio Clare, Estado De São Paulo, Brasil.
278. Originally published in Japanese in W. D. Hamilton, "My Intended Burial and Why," Insectarium 28. Reprinted in English in W. D. Hamilton, My Intended Burial and Why," Ethology Ecology and Evolution 12 (1991): 111-22.
279. Hamilton, Narrow Roads, 1:85, 86.
280. Now known as the International Institute of Entomology.
281. W. D. Hamilton, "The Moulding of Senescence by Natural Selection," Journal of Theoretical Biology 12 (1966): 12-45.
282. Although one reviewer of the initial manuscript referred to the model as "verbal diarrhea," this was clearly a minority position, as the Hamilton model of aging is today one of the most important ever constructed on the subject. 5. W. D. Hamilton, "Extraordinary Sex Ratios," Science 156 (1967): 477-87.
283. W. D. Hamilton, "Geometry of the Selfish Herd," Journal of Theoretical Biology 31 (1971): 295-311.
284. W. D. Hamilton, "Altruism and Related Phenomena, Mainly in Social Insects," Annual Review of Ecology & Systematics 3 (1972): 192-232; W. D. Hamilton, "Selfish and Spiteful Behaviour in an Evolutionary Model," Nature 228 (1972): 1,218-19.
285. J. Schwartz, "Death of an Altruist," Linga Franca 10 (2000): 52.
286. For research on uranium analysis as part of the Manhattan Project. British Library, George Price file, BL21X 102_13.
287. G. R. Price, "Science and the Supernatural," Science 122 (1955): 359-67
288. G. R. Price, "Where Is the Definitive Experiment?" Science 123 (1956): 17-18. In these papers, Price argued that not a single instance of purported extrasensory perception meets even the most liberal definitions of a scientifically demonstrable phenomenon. This article was picked up by newspapers all over world, including a front-page story in the New York Times. British Library, George Price file, BL21X 102_13.
289. British Museum, George Price file, BL21X 102_13.
290. Price most likely came across Hamilton's model in the Senate House Library. Hamilton, Narrow Roads, 1:.
291. Hamilton, Narrow Roads, 1:173.
292. Because natural selection works on relative fitness, spite can, in principle, evolve, as the relative fitness of the spiteful individual, though decreased by its own action, is still greater than that of the recipient of the spiteful act. The problem that all models of spite come up against is that individuals who are not spiteful at all should have higher fitnesses than either the initiator or recipient in a spiteful encounter.
293. E. C. Waltz, "Reciprocal Altruism and Spite in Gulls: A comment," American Naturalist 118 (1981): 588-92.
294. R. Pierotti, "Spite, Altruism, and Semantics-A Reply to Waltz," American Naturalist 119 (1982): 116-20
295. K. R. Foster, T. Wenseleers, and F.L.W. Ratnieks, "Spite: Hamilton's Unproven Theory," Annales Zoologici Fennici 38 (2001): 229-38
296. L. Keller, M. Milinski, M. Frischknecht, N. Perrin, H. Richner, and F. Tripet, "Spiteful Animals Still to Be Discovered," Trends in Ecology & Evolution 9 (1994): 103
297. R. Gadagkar, "Spiteful Animals Still to Be Discovered-Reply," Trends in Ecology & Evolution 9 (1994): 103
298. R. Gadagkar, "Can animals Be Spiteful?" Trends in Ecology & Evolution 8 (1993): 232-34.
299. Hamilton, Narrow Roads, 1:175.
300. W. D. Hamilton, "Selfish and Spiteful Behaviour in an Evolutionary Model," Nature 228 (1970): 1,218-19
301. G. R. Price, "Selection and Covariance," Nature 227 (1970): 520-21.
302. Schwartz, "Death of an Altruist," 52, 56.
303. Hamilton, Narrow Roads, 1:322-23.
304. Schwartz, "Death of an Altruist," 56.
305. Indeed, Cedric Smith spent considerable time writing grant proposals whose function was to support George Price's work. British Library, George Price file, BL21X 102_13.
306. Interview with John Maynard Smith. This interview can be found in L. A. Dugatkin, Principles of Animal Behavior (New York: W. W. Norton, 2004), 502-3.
307. Schwartz, "Death of an Altruist," p. 58.
308. Price had apparently attempted suicide once before, but precisely when is not known.
309. Hamilton, Narrow Roads, 1:174, 323-24.
310. Author's interview with Richard Dawkins, Royal Society of London, London, March 10, 2003.
311. G. Williams, Adaptation and Natural Selection (Princeton, N.J.: Princeton University Press, 1966), 5.
312. Interview with Dawkins, March 10, 2003.
313. Dawkins, The Selfish Gene, 97. In a later, more detailed analysis, Dawkins found that it was not until 1974 that citations to Hamilton's inclusive fitness work began to grow. Thenceforth, "like a gathering epidemic," references to Hamilton's altruism models increased steadily.
314. For example, in 1997, Gould and Dawkins reviewed each other's books in back-to-back papers appearing in Evolution: S. J. Gould, "Selfhelp for a Hedgehog Stuck on a Molehill (A Review of Richard Dawkins's Climbing Mount Improbable)," Evolution 51 (1997): 1,020-23
315. R. Dawkins, "Human Chauvinism (A Review Stephen Jay Gould's Full House)," Evolution 51 (1997): 1,015-20.
316. B. Rensberger, "Updating Darwin on Behavior," New York Times, May 28, 1975, 85.
317. For more on the debate surrounding Sociobiology, see U. Segerstrale, Defenders of the Truth: The Battle for Science in the Sociobiology Debate and Beyond (New York: Oxford University Press, 2000)
318. J. Alcock, The Triumph of Sociobiology (New York: Oxford University Press, 2001).
319. Gould was senior author of a November 13, 1975, letter entitled "Against Sociobiology," New York Review of Books
320. E. O. Wilson, "For Sociobiology," New York Review of Books, December 11, 1975
321. S. J. Gould and R. Lewontin, "The Spandrels of San Marcos and the Panglossian Paradigm: A Critique of the Adaptationist Programme," Proceedings of the Royal Society of London (ser. B) 205 (1979): 581-98.
322. J. Schwartz, "Oh My Darwin," Linga Franca (November 1999).
323. E. O. Wilson, Naturalist (New York: Warner Books, 1994), 319, 320.
324. Although he agrees that when the benefit of altruistic behavior discounted by the degree of relationship exceeds the cost, then altruism evolves, he is also in the midst of developing a new model that suggests "a supervening force," which he calls "ecological selection," could "virtually erase any effect of kin selection" in social insects. Author's interview with E. O. Wilson, Harvard University, Cambridge, Mass., February 26, 2003.
325. E. O. Wilson and B. Holldobler, "The Rise of the Ants: A Phylogenetic and Ecological Explanation," Proceedings of the National Academy of Sciences 102 (2005): 7,411-14
326. E. O. Wilson and B. Holldobler, "Eusociality: Origin and Consequences," Proceedings of the National Academy of Sciences 102 (2005): 13,367-71.
327. Wilson, Sociobiology, 554.
328. R. L. Trivers, "Parent-Offspring Conflict," American Zoologist 14 (1974): 249-65.
329. Wilson, Sociobiology, 555.
330. R. Dawkins, The Selfish Gene, 2nd ed. (Oxford: Oxford University Press, 1989), 328.
331. J. L. Brown, Helping and Communal Breeding in Birds (Princeton, N.J.: Princeton University Press, 1987)
332. P. Stacey and W. Koenig, eds., Cooperative Breeding in Birds: Long-Term Studies of Ecology and Behavior (Cambridge: Cambridge University Press, 1990).
333. J. L. Brown, E. Brown, S. D. Brown, and D. D. Dow, "Helpers: Effects of Experimental Removal on Reproductive Success," Science 215 (1982): 421-22.
334. A queen has two sets of chromosomes and hence two sets of genes. Both male and female offspring each receive a single set of these genes, so the queen shares copies of half her genes with each offspring, regardless of sex.
335. Fisher, Genetical Theory of Natural Selection.
336. R. Trivers and H. Hare, "Haplodiploidy and the Evolution of the Social Insects," Science 191 (1976): 249-63.
337. When Alexander taught his first course in animal behavior, Emlen was his teaching assistant.
338. Author's interview with Stephen Emlen, Cornell University, Ithaca, N.Y., January 31, 2003.
339. E-mail message from Stephen Emlen, February 17, 2003.
340. Approximately half of all nests have helpers at one time or another. S. T. Emlen and P. H. Wrege, "A Test of Alternative Hypotheses for Helping Behavior in White-fronted Bee-eaters," Behavioral Ecology and Sociobiology 25 (1989): 705. The bee-eaters is one of the few bird species that is both a cooperative and colonial breeder.
341. S. T. Emlen and N. Demong, "Bee-eaters: An Alternative Route to Cooperative Breeding," paper presented at the Symposium on Altruism in Birds, 27th International Ornithological Congress, West Berlin, 1978.
342. Little did he know that he would be making such visits each spring for more than the next twenty years.
343. Emlen interview, January 31, 2003.
344. S. T. Emlen, "An Evolutionary Theory of the Family," Proceedings of the National Academy of Sciences 92 (1995): 8,092-99.
345. The CGSS surveyed 13,495 households.
346. J. Davis and M. Daly, "Evolutionary theory and the human family," Quarterly Review of Biology 72 (1997): 407-35.
347. Emlen, "An Evolutionary Theory of the Family."
348. S. Pruett-Jones and M. Lewis, "Sex Ratio and Habitat Limitation Promote Delayed Dispersal in Superb Fairy Wrens," Nature 348 (1990): 541-42.
349. Indeed, in some cases (males, age 15-24), married individuals living away from home were more likely to remain in contact with their parents than were singles.
350. Instead, Davis and Daly suggest that that human parents act as postreproductive helpers to their own offspring. This help to children and grandchildren may have selected for strong family bonds that do not easily dissolve when offspring get married.
351. Emlen, "An Evolutionary Theory of the Family."
352. J. L. Brown, "Alternate Routes to Sociality in Jays-with a Theory for the Evolution of Altruism and Communal Breeding," American Zoology 14 (1974): 63-80. Not only are the offspring that remain on high-quality territories receiving a benefit, but their parents are as well, since they pass down the best-quality territories to their blood kin.
353. P. B. Stacey and J. D. Ligon, "Territory Quality and Dispersal Options in the Acorn Woodpecker, and a Challenge to the Habitat- Saturation Model of Cooperative Breeding," American Naturalist 130 (1987): 654-76.
354. L. White, "Coresidence and Leaving Home: Young Adults and Their Parents," Annual Review of Sociology 20 (1994): 81-102. Reviewed by Davis and Daly and used to test the hypothesis.
355. Emlen, "An Evolutionary Theory of the Family."
356. The extreme version of this hypothesis predicts that, under certain conditions, natural selection might even favor a stepparent's killing its offspring. If the stepparent is a male, patricide may result in his mate's quickly returning to an estrous state.
357. There is growing evidence that avian stepparents also destroy clutches containing potential stepchildren.
358. G. Hausfater and S. Blaffer-Hrdy, Infanticide (New York: Aldine, 1984).
359. J. Bray, "Children's Development during Early Remarriage," in Impact of Divorce, Single Parenting and Stepparenting on Children, ed. E. Hetherton and J. Arasteh (Hillsdale, N.J.: Lawrence Earlbuam, 1988), 279-98
360. L. White, "Stepfamilies over the Life Course: Social Support," in Stepfamilies: Who Benefits? Who Does Not?, ed. A. Booth and J. Dunn (Hillsdale, N.J.: Lawrence Earlbaum, 1994), 109-37.
361. Daly and Wilson, The Truth about Cinderella, 7.
362. Daly and Wilson, Homicide, and The Truth about Cinderella
363. D. Kaplun and R. Reich, "The Murdered Child and His Killers," American Journal of Psychiatry 133 (1976): 809-13
364. M. Daly and M. Wilson, "Some Differential Attributes of Lethal Assaults on Small Children by Stepfathers versus Genetic Fathers," Ethology and Sociobiology 15 (1994): 207-17.
365. H. Temrin, S. Buchmayer, and M. Enquist, "Stepparents and Infanticide: New Data Contradict Evolutionary Predictions," Proceedings of the Royal Society of London 267 (2000): 943-45.
366. From an e-mail exchange with Margo Wilson, December 9, 2004.
367. Emlen interview, January 31, 2003.
368. Sherman's dissertation work was done at the University of Michigan. He first learned of the ground squirrel system from Martin Morton.
369. P. W. Sherman, "Nepotism and the Evolution of Alarm Calls," Science 197 (1977): 1,246-53.
370. This was the first of a series of such papers that Sherman would publish, including: "The Limits of Ground Squirrel Nepotism," in Sociobiology: Beyond Nature / Nurture?, ed. G. Barlow and J. Silverberg (Boulder, Colo.: Westview, 1980), 505-44
371. "The Meaning of Nepotism," American Naturalist 116 (1980): 604-6
372. "Kinship, Demography, and Belding's Ground Squirrel Nepotism," Behavioral Ecology and Sociobiology 8 (1981): 251-59
373. "Alarm Calls of Belding's Ground Squirrels to Aerial Predators: Nepotism or Self-preservation?" Behavioral Ecology Sociobiology 17 (1985): 313-23
374. P. W. Sherman and W. Holmes, "Kin Recognition: Issues and Evidence," Fortschritte der Zoologie 31 (1985): 437-60.
375. Author's interview with Paul Sherman, Cornell University, Ithaca, N.Y., January 31, 2003.
376. Sherman interview, January 31, 2003.
377. Sherman, "Alarm Calls of Belding's Ground Squirrels to Aerial Predators," 313.
378. Sherman interview, January 31, 2003.
379. Termites are part of the insect order Isoptera.
380. P. W. Sherman, J. Jarvis, and R. Alexander, eds. The Biology of the Naked Mole-Rat (Princeton, N.J.: Princeton University Press, 1991), ix.
381. Sherman interview, January 31, 2003.
382. Compared with other species, naked mole-rats lead extraordinarily long lives. P. Sherman and J. Jarvis, "Extraordinary Life Spans of Naked Mole-rats (Heterocephalus glaber)," Journal of the Zoological Society of London 258 (2002): 307-11.
383. E. A. Lacey, R. D. Alexander, S. H. Braude, P. W. Sherman, and J. Jarvis, "An Ethogram for the Naked Mole-rat: Non-vocal Behaviors," in The Biology of the Naked Mole-Rat, 209-42.
384. E. A. Lacey and P. W. Sherman, "Social Organization of Naked Mole-rats: Evidence for Divisions of Labor," in The Biology of the Naked Mole-Rat, 275-336.
385. H. K. Reeve and P. W. Sherman, "Intracolonial Aggression and Nepotism by the Breeding Female Naked Mole-rat," in The Biology of the Naked Mole-Rat, 337-57.
386. J.U.M. Jarvis, "Eusociality in a Mammal: Cooperative Breeding in Naked Mole-rat Colonies," Science 212 (1981): 571-73.
387. H. K. Reeve, D. F. Westneat, W. A. Noon, P. W. Sherman, and C. F. Aquadro, "DNA 'Fingerprinting' Reveals High Levels of Inbreeding in Colonies of the Eusocial Naked Mole-rat," Proceedings of the National Acadamy of Sciences 87 (1990): 2,496-2,500.
388. Reeve et al., "DNA 'Fingerprinting,'" 2,499. Why the deleterious effects typically associated with inbreeding (for example, an increase in diseases caused by recessive alleles) are absent in naked mole-rats remains a mystery.
389. H. K. Reeve, "Queen Activation of Lazy Workers in Colonies of the Eusocial Naked Mole-rat," Nature 358 (1992): 147-49.
390. When a queen is removed from a colony, it is the workers that are least related to her that are most likely to decrease their workload.
391. D. Fletcher and C. Michener, Kin Recognition in Animals (New York: Wiley, 1987)
392. P. G. Hepper, Kin Recognition (Cambridge: Cambridge University Press, 1991)
393. D. W. Pfennig and P. W. Sherman, "Kin Recognition," Scientific American, June 1995, 98-103
394. P. Hepper and J. Cleland, "Developmental Aspects of Kin Recognition," Genetica 104 (1998): 199-205.
395. B. Waldman, "Mechanisms of Kin Recognition," Journal of Theoretical Biology 128 (1987): 159-85
396. H. K. Reeve, "The Evolution of Conspecific Acceptance Thresholds," American Naturalist 133 (1989): 407-35.
397. The internal template of an individual may be generated by either some sort of genetically based rule or via learning. E. Boyse, G. Beauchamp, K. Yamazaki, and J. Bard, "Genetic Components of Kin Recognition in Mammals," in Kin Recognition, 162-219
398. R. Crozier, "Genetic Aspects of Kin Recognition: Concepts, Models and Synthesis," in Kin Recognition in Animals, 55-74
399. S. Robinson and W. Smotherman, "Fetal Learning: Implications for the Development of Kin," in Kin Recognition, 308-34
400. R. D. Alexander, Darwinism and Human Affairs (Seattle: University of Washington Press, 1979)
401. R. D. Alexander, "Social Learning and Kin Recognition," Ethology and Sociobiology 12 (1991): 387-99.
402. D. W. Pfennig, H. K. Reeve, and P. W. Sherman, "Kin Recognition and Cannibalism in Spadefoot Toads,"Animal Behavior 46 (1993): 87-94
403. M. Elgar and B. Crespi, Cannibalism: Ecology and Evolution among Diverse Taxa (Oxford: Oxford University Press, 1992).
404. W. Holmes and P. W. Sherman, "Kin Recognition in Animals," American Scientist 71 (1983): 46-55
405. P. W. Sherman and W. Holmes, "Kin Recognition: Issues and Evidence," Fortschritte Zoologie 31 (1985): 437-60
406. A. Blaustein, "Kin Recognition Mechanisms: Phenotype Matching or Recognition Alleles," American Naturalist 121 (1983): 749-54
407. A. Blaustein, M. Bekoff, J. Byers, and T. Daniels, "Kin Recognition in Vertebrates: What Do We Really Know about Adaptive Values?" Animal Behavior 41 (1991): 1,079-83.
408. M. D. Beecher, I. Beecher, and S. Lumpkin, "Parent-Offspring Recognition in Bank Swallows: I. Natural History," Animal Behavior 29 (1981): 86-94
409. M. D. Beecher, I. Beecher, and S. Hahn, "Parentoffspring recognition in bank swallows: II. Development and acoustic basis," Animal Behavior 29 (1981): 95-101
410. M. D. Beecher, B. Medvin, P. Stoddard, and P. Loesch, "Acoustic Adaptations for Parent-Offspring Recognition in Swallows," Experimental Biology 45 (1986): 179-93.
411. Hamilton, Narrow Roads, 1:486.
412. Afaculty member at Michigan's Institute for Public Policy Studies (now called the School of Public Policy).
413. Hamilton, Narrow Roads, 2:119.
414. Hamilton had some experience with game theory, reciprocity, and the prisoner's dilemma. W. D. Hamilton, "Extraordinary Sex Ratios," Science 156 (1967): 477-87
415. W. D. Hamilton, "Altruism and Related Phenomena, Mainly in Social Insects," Annual Review of Ecological Systems 3 (1972): 192-232
416. W. D. Hamilton, "Selection of Selfish and Altruistic Behaviour in Some Extreme Models," in Man and Beast: Comparative Social Behavior, ed. J. F. Eisenberg and W. S. Dillon (Washington, D.C.: Smithsonian Institution Press, 1971), 57-91.
417. Hamilton, Narrow Roads, 2:120.
418. W. Poundstone, Prisoner's Dilemma: Jon Von Neuman, Game Theory and the Puzzle of the Bomb (New York: Doubleday, 1992).
419. More precisely, they were searching for the evolutionarily stable strategy (ESS) to this game. An ESS is a strategy that at high frequencies cannot be "invaded" and replaced by any other strategy.
420. In fact, Axelrod and Hamilton ran a second computer tournament with sixty-two strategies. Results were similar to those in the initial tournament.
421. Submitted by Anatol Rapoport of the Institute of Advanced Study in Vienna.
422. It is also the case that a strategy called "always defect" also does well in the iterated prisoner's dilemma game.
423. Since the original models of tft appeared, dozens of variants of this strategy have been examined, but most share the essential characteristics
424. L. A. Dugatkin, Cooperation among Animals: An Evolutionary Perspective (New York: Oxford University Press, 1997) for more on this.
425. Hamilton, Narrow Roads, 2:124.
426. http://www.royalsoc.ac.uk/funding/ResProf/CondTenRP1A.pdf, condition 5.
427. Hamilton, Narrow Roads, 2:303.
428. Richard Dawkins in a forward to Hamilton, Narrow Roads, 2:xvii.
429. Hamilton, Narrow Roads, 2:100.
430. Hamilton was still able to recognize friends and family.
431. E. Hooper, The River: A Journey Back to the Source of HIV and AIDS (New York: Little, Brown, 1999).
432. Alan Grafen, Guardian, March 9, 2000.
433. London Times, March 9, 2000.
434. Natalie Angier, New York Times, March 10, 2000.
435. Richard Dawkins, Independent (London), March 10, 2000.
436. Hamilton, Narrow Roads, 1:93.