The Price of Altruism (59 page)
Read The Price of Altruism Online
Authors: Oren Harman
7.
Alexander M. Carr-Saunders,
The Population Problem: A Study in Human Evolution
(Oxford: Clarendon Press, 1922).
8.
Wynne-Edwards’s descriptions of nature in the Arctic are very similar to those of Kropotkin in
Mutual Aid
: “Except perhaps among carnivorous predators, competition between individuals for space and nourishment seems commonly to be reduced to a low level among members of the Arctic flora and fauna; they live somewhat like weeds, the secret of whose success lies in their ability to exploit transient conditions while they last, in the absence of serious competition. In the Arctic the struggle for existence is overwhelmingly against the physical world, now sufficiently benign, now below the threshold for successful reproduction, and now so violent that life is swept away, after which recolonization alone can restore it.” V. C. Wynne-Edwards, “Zoology of the Baird Expedition,”
The Auk
69, no. 4 (1952), 353–91, quote on 384.
9.
V. C. Wynne-Edwards, “Intermittent Breeding of the Fulmar (
Fulmarus glacialis
) with Some General Observations on Non-Breeding in Sea-Birds,”
Proceedings of the Zoological Society of London
A109 (1939), 127–32.
10.
There are a number of known density-dependent reproduction-control mechanisms, such as reducing the rate of ovulation via a change in hormone output, or resorption of the embryos in the uterus as a result of stress. Wynne-Edwards, however, was talking about voluntary restraint accomplished via ritualized convention, as in territoriality displays, migration, and abstention.
11.
This had been the original insight of Georgii Gause, which he called the “principle of competitive exclusion.” See Sharon E. Kingsland,
Modeling Nature: Episodes in the History of Population Ecology
(Chicago: University of Chicago Press, 1985), 146–62.
12.
David L. Lack,
The Natural Regulation of Animal Numbers
(Oxford: Clarendon Press, 1954). See also Lack, “My Life as an Amateur Ornithologist,”
Ibis
115 (1973), 421–31,
The Life of the Robin
(London: Witherby, 1943),
Swifts in a Tower
(London, Methuen, 1956), and
Darwin’s Finches: An Essay on the General Biological Theory of Evolution
(London: Methuen, 1973).
13.
V. C. Wynne-Edwards,
Animal Dispersion in Relation to Social Behavior
(Edinburgh: Oliver and Boyd, 1962), 20. The fishing analogy first appeared in V. C. Wynne-Edwards, “The Control of Population Density Through Social Behaviour: A Hypothesis,”
Ibis
101 (1959), 436–41, quote on 437.
14.
Wynne-Edwards called the kind of behavior necessary for such convention “epideictic.” His main study of such behavior was on the red grouse,
Lagopus lagopus scotica
, in the vicinity of Aberdeen. Olavi Kalela, too, developed this idea. See his “ber die Funktion der Mandibeln bei den Soldaten von
Neocapritermes opacus
(Hagen),”
Zoologischer Anzeiger
152 (1954), 228–34.
15.
Charles Darwin,
The Origin of Species
, 139, emphasis added.
16.
Quoted in Kohn,
A Reason for Everything
, 227. For a study of Wynne-Edwards’ place in the history of the debate on group selections see Mark Borrello,
Evolutionary Restraints: The Contentious History of Group Selection Theory
(Chicago: University of Chicago Press, 2010).
17.
Ibid., 224.
18.
David Lack,
Population Studies of Birds
(Oxford: Oxford University Press, 1966); Charles Elton, “Self-Regulation of Animal Populations,”
Nature
197 (1963), 634.
19.
V. C. Wynne-Edwards, “Population Control in Animals,”
Scientific American
211 (1964), 68–74.
20.
“The Nature of Social Life,”
Times Literary Supplement,
December, 14, 1962, 967. More reviews are discussed in Borrello,
Evolutionary Restraints
. I thank Mark Borrello kindly for sharing with me a prepublication version of his manuscript.
21.
David Cort, “The Glossy Rats: A Review of Animal Dispersion in Relation to Social Behavior,”
The Nation
, November 16, 1963, 326–29, quote on 327. Wynne-Edwards himself often made the comparison to man.
22.
August Weismann,
Ueber die Dauer des Lebens
[On the Duration of Life] (Jena, 1882);
Ueber Leben und Tod
[On Life and Death] (Jena, 1884). Both appear in English in Weismann,
On Heredity
(Oxford: Clarendon Press, 1891).
23.
See John Maynard Smith, “The Causes of Ageing,” in Review Lectures on Senescence,
Proceedings of the Royal Society of London B
157 (1962), 115–27. Peter Medawar and George C. Williams had already been thinking along these “pleiotropic” lines. See Medawar,
An Unsolved Problem of Biology
(London: H.K. Lewis, 1952), and Williams, “Pleiotropy, Natural Selection, and the Evolution of Senescence,”
Evolution
11 (1957), 398–411.
24.
This formulation belongs to Richard Dawkins,
The Selfish Gene
(Oxford: Oxford University Press, 1976; reprint, 1989), 175.
25.
The haplodiploid hypothesis turned out to be flawed: many haplodiploid species have not evolved eusociality, whereas other, diplodiploid species have. Still, the “spirit” of the hypothesis survived in the notion that there is genetic conflict between sisters’ interest in producing more female reproductives than males versus the queen’s interest in an equal investment in the sexes. See R. L. Trivers and H. Hare, “Haplodiploidy and the Evolution of the Social Insects,”
Science
91 (1976), 249–63, and more recently, N. J. Mehdiabadi, H. K. Reeve, and U. G. Mueller, “Queens Versus Workers: Sex Ration Conflict in Eusocial Hymenoptera,”
Trends in Ecology and Evolution
18, no. 2 (2003), 88–93. For a definitive presentation of current knowledge on social insects see Bert Hlldobler and E. O. Wilson,
The Superorganism: The Beauty, Elegance, and Strangeness of Insect Societies
(New York: W. W. Norton, 2009).
26.
For a full-length biography of Hamilton see Ullica Segerstrle,
Nature’s Oracle: A Life of W. D. Hamilton
(Oxford: Oxford University Press, 2010). There are many shorter appreciations of Hamilton’s life and science. See Alan Grafen, “William Donald Hamilton, 1936–2000,”
Biographical Memoirs of Fellows of the Royal Society
50 (2004), 109–32.
27.
This was the writer Alisdair Gray’s remark, quoted in Kohn,
A Reason for Everything
, 259; 260; interview with Janet Hamilton (Bill’s sister), October 24, 2007.
28.
“I realized that I had little talent in mathematics and even less training for it,” he later wrote, “so that my efforts to teach myself what was necessary to understand even the merely standard theoretical population genetics of the day were tedious in the extreme.”
29.
On the jacket of the 1999 edition of Fisher’s book.
30.
W. D. Hamilton,
Narrow Roads of Gene Land
,
Volume 1: Evolution of Social Behavior
(Oxford: Spektrum, 1996), 21; the quote on 22 is from Vincent Wiggles-worth,
The Life of Insects
(London: Weidenfeld and Nicolson, 1966).
31.
Letters to Mary quoted in Kohn,
A Reason for Everything
, 265, 266.
32.
Charles Darwin,
On the Origin of Species
, facsimile of the first edition with an introduction by Ernst Mayr (Cambridge, MA: Harvard University Press), 189.
33.
Hamilton,
Narrow Roads
, 22.
34.
On the decline of eugenics including Penrose’s role see Daniel J. Kevles,
In the Name of Eugenics: Genetics and the Uses of Human Heredity
(Cambridge, MA: Harvard University Press, 1995), especially 148–222.
35.
Hamilton,
Narrow Roads
, 14.
36.
Ibid., 4.
37.
Ibid., 12–13.
38.
Ibid., 15, 5.
39.
In fact Hamilton’s supervisor Cedric Smith had once introduced the two, but Maynard Smith couldn’t remember the encounter. Hamilton was shy and very unassuming, and Maynard Smith, like everyone else at the time, wasn’t interested in the genetics of altruism.
40.
Ullica Segerstr
Defenders of the Truth: The Sociobiology Debate
(Oxford: Oxford University Press, 2000), 64.