George Williams, the man whose name will forever be associated with the rejection of group selection in the 1960’s, passed away on September 8. He previously suffered from Alzheimer’s disease, so his voice has not been heard for a number of years. He was my intellectual opponent but also a good friend.
I first met George when I was a graduate student in 1974. I had just written my first paper on group selection and was driving to Harvard to convince E.O. Wilson to sponsor it for publication in the Proceedings of the National Academy Of Sciences. I decided to visit George at Stony Brook along the way and strode into his office with the words “I’m going to convince you about group selection.” His response was to offer me a postdoctoral position on the spot.
I didn’t accept the position but I did take the opportunity to visit with George and his wonderful wife Doris whenever I could and they would often stop at our house in Binghamton on their way to or from their summer home in Canada. George had a slow way of talking and a droll sense of humor. Once, when introducing me at a seminar, he recounted our first meeting, paused for a long time, and continued “I’m sure that he has been disappointed before and will be disappointed again.” During the same visit, he marked his house so that I could find it by parking his lawnmower by the entrance with a sign taped on that read “Super-organisms welcome here.” The sign has been gracing the door of my laboratory ever since.
When my wife Anne and I discovered that George had been diagnosed with Alzheimer’s, she observed that someone should record his story and thoughts while he was still able to relate them. The perfect person to do this was Will Provine, the great historian of evolution at Cornell University. Will, Anne and I bundled ourselves into a car and drove down to Stony Brook, where we stayed with George and Doris and interviewed them for two days. It was one of the best decisions that we ever made.
George always accepted multi-level selection as a theoretical framework. In Adaptation and Natural Selection, his most influential book, he affirmed the basic fact that altruism and other traits that are “for the good of the group” are selectively disadvantageous within groups and require between-group selection to evolve. His claim was that lower-level selection is invariably stronger than higher-level selection so “group selection does not, in fact. exist.”
George did change his mind about group selection, at least for some specific traits. In the 1990’s he acknowledged that female-biased sex ratios count as an example of group selection and his interest in Darwinian medicine caused him to become downright enthusiastic about reduced virulence in parasites as an example of group selection. In a letter he acknowledged that group selection is probably important in human evolution, causing me to shake my head in wonder, knowing how many members of the Human Behavior and Evolution Society are staunch anti-group selectionists and treat George as a god. In general, however, George retained his worldview and I didn’t convince him about group selection.
Ironically, on the day after George passed away, Elliott Sober and I were notified that a paper we submitted to the Journal of Evolutionary Biology titled “Adaptation and Natural Selection Revisited” was accepted for publication. In this paper, we argue in favor of George’s central position about the relationship between adaptation and natural selection, which we call Williams’ principle, against some modern-day individualists who think they are following in his footsteps. I cannot think of a more fitting tribute to a man who represents the best of scientific discourse, so much that he treated his intellectual adversary as a friend.