The evolutionary biologist George Williams died on Wednesday at the age of 84. He was one of the most important evolutionary biologists of the twentieth century, although he's not a familiar name beyond scientific circles. Gently, persistently, he reformulated how we think about natural selection and its many effects. As Richard Dawkins noted today, "He was one of the great evolutionary thinkers of my lifetime." In 2004, I wrote a profile of Williams for Science. The occasion was a meeting that was held in Williams's honor, in which one scientist after another stood up to talk about the influence he had had on their work. They investigated everything from human behavior to the mating of fish to disorders of pregnancy. How on Earth could he have so much influence in so many different directions? Permit me to self-plagiarize:
In the 1950s, when Williams was doing his graduate work at the University of California, Los Angeles, the science of evolutionary biology had just gone through two decades of spectacular advances. Ronald Fisher and Theodosius Dobzhansky, among others, had used the new science of genetics to work out some of the molecular underpinnings of evolution. Natural selection was now recognized as a change in the frequency of genes in a population. Yet one important part still hadn't been nailed down: the nature of adaptations. It was clear that adaptations evolved, but few biologists had given serious thought to the rules that govern the process.