Harman Oren Solomon. The Price of Altruism: George Price and the Search for the Origins of Kindness

18. Since the Price equation is a statistical, not a causal, decomposition of

selection, there is an interesting reflection of this problem in the very bones of the

math itself. In the end the Price equation can tell you that a trait evolved, but why

it evolved remains elusive.

19. Bill Hamilton letter to Kathleen Price, June 25, 1997, BLWHC,

BL:Z1X101_1.2.2.3.

20. “Love is the Greatest!” found in George’s scattered papers, GPP.

APPENDIX 1: COVARIANCE AND KIN SELECTION

1. Here I follow S. A. Frank, “George Price’s Contributions to Evolutionary

Genetics,” Journal of Theoretical Biology 175 (1995), 374–75.

2. D. C. Queller, “Kinship, Reciprocity and Synergism in the Evolution of Social

Behaviour,” Nature 318 (1985), 366–67. See also his “A General Model for Kin

Selection,” Evolution 46 (1992), 376–80.

3. W. D. Hamilton, “Selfish and Spiteful Behaviour in an Evolutionary Model,”

Nature 228 (1970), 1218–20.

APPENDIX 2: THE FULL PRICE EQUATION AND LEVELS OF

SELECTION

1. Once again, I follow Frank, “George Price’s Contributions to Evolutionary

Genetics,” 375–79. Samir Okasha, Evolution and the Levels of Selection (Oxford:

Oxford University Press, 2006), provides a full, fleshed-out interpretation for

those interested.

2. A. Robertson, “A Mathematical Model for the Culling Process in Dairy Cattle,”

Animal Production 8 (1966), 95–108; C. C. Li, “Fundamental Theorem of Natural

Selection,” Nature 214 (1967), 505–6.