During the second half of the twentieth century, Williams emerged as one of the most influential thinkers in evolutionary biology, and helped to clarify a number of key issues in the field. His defining style was to tackle difficult questions that had stymied other evolutionary theorists, and his thirst for challenge led him to propose influential theories about aging and senescence, the origin of sex, life history traits, and Darwinian medicine. He may have been one of the last great evolutionary theorists who could effectively rely on verbal models alone, and many lesser theorists have made careers out of following the mathematical consequences of Williams’ prose-based theories.
My greatest appreciation for Williams is in an area with which I mostly disagree with him: the role of group selection in evolution. In the 1960’s Williams completely dismantled the naive group selectionist paradigm by carefully and mercilessly pointing out every one of its false assumptions. In this sense, those of us who still entertain the idea that selection at a level above the individual (or the gene!) matters in the process of evolution owe to Williams a great debt. By helping to clear the field of sloppy thinking, he also cleared the way for more meaningful work on how group selection might actually function.
I use readings from his Plan & Purpose in Nature at the very beginning of my Evolution of Cooperation class, and my non-major students always respond strongly to his clear, provocative descriptions of how evolution does (and does not) work. That Williams could write about such abstract ideas and bring them alive for readers with little science background is a testament to his absolute clarity of thought on core issues of evolutionary biology.
I had a small connection to Williams by virtue of being a graduate student at Stony Brook University. Along with Larry Slobodkin and Robert Sokal, George founded the Ecology and Evolution department at Stony Brook, one of the first of its kind in the United States. To these men I owe in part my own academic development. By the time that I arrived at Stony Brook in 2001, George had already retreated into emeritus status, but he was occasionally present at seminars and talks held by the department. Quiet, unassuming — perhaps even shy — Williams carried himself with none of the pompous bearing so often seen in “famous scientists”. I remember quite clearly my closest interaction with him, which occurred during a meeting with my advisor Lev Ginzburg. Lev was considering whether the destabilization of ecological systems might be a potential evolutionary force, and I had warned him that this sounded like a form of group selection, so we decided to consult the expert. After hearing Lev’s exposition of his idea, George paused for a pretty long time and then simply said “I will have to think about that for awhile”. After about a week, he came back to Lev with valuable feedback. Clearly George was a man who valued precise thought over any other efficiency.
It is ironic that Williams’ death was preceded by mental decline. It is easy to see in his own life history an allegory for antagonistic pleiotropy: did cerebral brilliance in earlier life trade off with dementia at the very end of his life? But I think to focus too closely on the somatic existence of George Williams (or even on his reproductive success, as he left behind four children and nine grandchildren) is to miss the bigger impact of his life. Despite the fact that so much of his exceptionally-clear thinking was about death, decline, and gene-centered selfishness, George demonstrated most brilliantly the immortal power of valuable ideas. The legacy left by his diverse writings has already embedded itself in the foundation of modern evolutionary biology. Though new ideas will be laid upon Williams’, I suspect that his contributions will remain a prominent part of the bedrock underlying our field.
If you are interested in reading more about Williams’ life and work, there are a number of great tributes to him that have already been published by Randolph Nesse, Laura Betzig, David Sloan Wilson, Michael Ruse, Carl Zimmer, Richard Dawkins, and the “Reality Club”. Both the New York Times and the local Long Island newspaper Newsday have well-written obituaries for Williams.
Adaptation, Biography, Evolution, Group Selection, Obituary, Senescence