George Price was born a Jewish half-breed to parents who kept his Semitic side a secret; lived much of his life an aggressive atheist and skeptic of the supernatural; and died a Christian, twice converted, albeit, to his mind, a defeated one. Several years before he abandoned his career in a mission to shelter and comfort homeless alcoholics, he made a number of extraordinary contributions to evolutionary biology, a field in which he had no training. Educated as a chemist, Price had worked previously for the Manhattan Project on uranium enrichment, helped develop radiation therapy for cancer, invented computer-aided design with IBM and dabbled in journalism.
Shortly after Christmas 1974, Price slashed his carotid artery with a pair of tailor’s scissors in his room in a London squat. John Maynard Smith, with whom Price published a paper that applied game theory to natural selection, was one of the few people, along with some of those homeless alcoholics, to attend his funeral. Also present was William Hamilton, the father of kin selection, which proposed that self-sacrificing behavior was able to evolve between related organisms because of the advantages conferred to their shared genes. Price used Hamilton’s ideas about kin selection to derive his own equation, one that could explain selection at multiple levels of organization—the genetic level, as well as among individuals in kin groups and populations of unrelated others. The equation marked a breakthrough in the field: Price had provided a working mathematical model for the emergence of altruism in a theory of the world that took dogmatic self-interest as its first principle.
Richard Dawkins, who considers Hamilton, Price’s closest collaborator, a "good candidate for the title of most distinguished Darwinian since Darwin," based his gene’s-eye-view theory of natural selection on Hamilton’s work in kin selection. This paradigm, which Dawkins popularized with The Selfish Gene in 1976, has dominated evolutionary biology and its progeny, sociobiology and evolutionary psychology, ever since. Maynard Smith and Hamilton are now dead, as are, presumably, those homeless alcoholics, while Dawkins has become one of the most strident and significant of the voices that blare on in what is known as "the God debate."
Price has remained, for some three decades, a footnote in the history of science. This is in part the doing of Price himself, the cost, perhaps, referred to in the title of The Price of Altruism, although I suspect that his biographer, Oren Harman, is alluding more to the misery and suicide that followed his subject’s failed stab at altruism than to the marginalization of his scientific work. Hamilton, who deemed the Price equation "a far better tool for all forms of selection acting at one level or at many" than he had ever before had at his disposal, implored Price to keep working on the equation and to publish his results. By then, Price was deep into his mission ministering to wayward souls, eventually giving up his home and few possessions in an attempt to live as Jesus would. "Oh yes," Price told Hamilton not long before he died. "But I have so many other things to do…population genetics is not my main work, as you know."
* * *
If the main work of evolutionary theory has been to explain and substantiate the process of evolution through natural selection, the main obstacle to that work has been, since Darwin’s time, the problem of altruism. In 1838, two years after his return from his journey on the HMS Beagle, Charles Darwin read An Essay on the Principle of Population by Thomas Malthus. Its proposition that population growth always exceeds food supply astonished Darwin, and offered him, at last, "a theory by which to work." "As more individuals are produced than can possibly survive, there must in every case be a struggle for existence," Darwin wrote some twenty years later in the Origin of Species. "It is the doctrine of Malthus applied with manifold force to the whole animal and vegetable kingdoms." Yet despite the myriad predators and prey who bore out this theory, there were obvious outliers: sterile worker insects, such as ants, who labor for the good of the nest rather than their own reproductive success. Darwin was perplexed by this "most serious special difficulty" facing his theory, but he accounted for it with the supposition that "division of labour" must be as beneficial for an insect community as it was "useful to civilised man." "It may not be a logical deduction," Darwin wrote, but nevertheless, such self-sacrificing behavior showed that natural selection was even more efficient than he had imagined.