It bothers me that so little is being said about George Williams, and it seems to me that more of his intellectual debtors should step up.
Back in the day, there were three big names in evolutionary biology: Bob Trivers, Bill Hamilton and George Williams. Trivers, the youngest, made his reputation in the early ‘70s with papers on reciprocal altruism, parental investment and sexual selection, parent-offspring conflict, and sex ratios. Every one of those papers applied evolutionary theory, explicitly, to H sapiens. “One human being leaping into water at some danger to himself, to save another distantly related human from drowning may be said to display altruistic behavior,” is the proposition that starts the paper on reciprocal altruism; data on paternity uncertainty in Kalahari Bushmen starts the paper on parental investment and sexual selection; the paper on parent-offspring conflict—which reads like an autobiography, to me—ends with a wonderful discussion of the priesthood; the paper on sex ratios ends with an “application of the model to humans,” and preliminary data sex ratios and SES in the US. Trivers started out studying history, and has always worked closely with people in anthropology. His work, more than the others’, bridges the gap to us.
Bill Hamilton, the second youngest, died ten years ago. Of the big three, Hamilton was most in love with “maths;” and he was most at home in the field. He made a number of trips to the Amazon, to look at wasps; and in the months before he died, he was on an expedition to the Congo, looking for the origins of HIV. He was probably most interested in insects and pathogens—but always intensely curious about humans. The short list of Hamilton’s classics—on kin selection, senescence, sex ratios, selfish herds, tit-for-tat, parasites and sex—run the gamut of social interaction; and the longer list isn’t shy of titles like “Selfishness re-examined: no man is an island” and “Innate social aptitudes of man.” Hamilton started out studying biology at Cambridge; but he got his advanced degree in “Human Demography” at LSE. He read Ronald Fisher with Franz Kafka; and the autobiographical essays attached to his collected scientific papers, Narrow Roads of Gene Land—named for the Japanese poet, Matsuo Basho—are good English literature.
But the first, and the oldest, of the big three was George Christopher Williams. Unlike the other two, George got both of his degrees in biology, from Berkeley and UCLA, where he studied fish. He was less of a fieldworker than Hamilton, and less of an anthropologist than Trivers. But more than the others, he was a great reader, and a great writer, of natural history. Hamilton and Trivers wrote papers; George Williams wrote books. He did start out, in 1957, with a pair of hugely influential articles in Evolution, on kinship and death—the first, on “Pleiotropy natural selection, and the evolution of senescence;” and the second, written with his wife Doris, on “Natural selection of individually harmful adaptations among sibs with special reference to social insects.” But afterwards, he elaborated on those ideas, and others, at greater length. There was the 1994 collaboration with Randy Nesse that launched Darwinian medicine, called Evolution and Healing in London and Why We Get Sick in the US; there was the 1992 clarification on adaptation, called Natural Selection: Domains, Levels, and Challenges; there was the 1975 Sex and Evolution, on the most important “question and paradox” in natural selection; and there was Williams’ great, 1966 book, Adaptation and Natural Selection: A Critique of some Current Evolutionary Thought. Those books are full of arctic foxes and phalaropes, rotifers and oysters. But the lessons he drew from them—on everything from the levels of selection, to sexual selection—are fundamental to our understanding of Homo sapiens.
When Bob Trivers wrote about sexual selection—reprised in his own collected essays, Natural Selection and Social Theory—he had to admit that “the importance of sex role-reversed species was first suggested by George Williams (1966), and I certainly read it there, but I forgot where I had learned it and imagined that I had made this extension myself.” And when Bill Hamilton wrote about inclusive fitness, or senescence, or selfish herds, or sex—reprinted in Narrow Roads of Gene Land—he gave credit to George Williams, whose “conclusions are doubtless correct,” who “seemed to have understood the matter right through,” who “as usual had anticipated me in studying this egotistic interpretation,” who had a “characteristic knack of seeing a problem from a totally fresh point of view.”
Our theoretical debt doesn’t stop there, of course. As everybody from Richard Dawkins to E O Wilson continues to think through the fundamentals of the levels of selection and adaptation, their first debt is to George Williams, who defined and clarified the problem.
Our empirical debt is enormous. When Sarah Blaffer Hrdy studied infanticide in Hanuman langurs, she was looking for evidence of selection at the individual level—as were others who studied infanticide in other species, and in us. When Mary Jane West Eberhard studied dominance interactions in paper wasps, she was looking for individual level adaptations—as were others who studied dominance in other oragnisms, and in us. When Kristen Hawkes and her friends study grandmothering in humans, they’re following George Williams on parental care as an individual level adaptation. And when Linda Partridge and her fellow gerontological researchers look into human aging, they’re following George Williams on senescence as an effect of individual level selection.
Undergraduates in Richard Alexander’s courses on social behavior at Michigan pored over Adaptation and Natural Selection. My copy is marked up with several inks, and the binding long ago fell apart. I love the part about phalaropes—which solved the problem of sexual relations for me then, and has since solved it for another generation. “The evidence strongly supports the conclusion that promiscuity, active courtship, and belligerence toward rivals are not inherent aspects of maleness. They will be developed in whichever sex cannot effectively increase its production of offspring merely by increasing its material contribution” (p. 186). I love the constant insistence on parsimony in the search for adaptations to social life. “We should not invoke biological principles where statistics suffices” (p. 257). And I love the part in the introduction that explains why it’s so hard for us to admit the importance of natural selection. “It is difficult to image that the blind play of the genes could produce man” (p. 4).
I last saw George Williams at the end of the last century, on a trip to New York. My career had just fizzled, a little: I’d passed up a job out east, in order to stay home and raise my children in the midwest. But I invited George to dinner anyway, and he came. He met me, and John Hartung, in Helen Fisher’s apartment west of Central Park—in spite of the fact that he’d been up all the previous night, attending to the birth of one of his grandchildren on Long Island. He gave each of us a signed copy of The Pony Fish’s Glow, his last book. I was so honored. We all had a really good time.
At the end of that year, or the next year, Paul and I got a holiday letter. It came with the most wonderful picture of George and Doris’ nine beautiful grandchildren—dark headed, tow headed, redheaded—tumbling down the stairs in their grandparents’ house. It makes me happy that George’s own statistical contributions to the next two generations were good ones. I’ve held onto that card.
Most students of human behavior don’t get bogged down in biology courses. They study psychology, sociology or anthropology. If they become interested in what’s variously known as sociobiology, evolutionary anthropology or evolutionary psychology, they learn George Williams’ name, and occasionally they cite him. Probably very few take the time to read his books. But we’re all enormously indebted to him. My words can’t convey how much. But I can’t help pointing it out.