RIP, G.C. Williams

The American biologist George C. Williams has died at the age of 84. Williams worked and taught for a long time at SUNY Stony Brook, and is best known for his book Adaptation and Natural Selection, published in 1966, where he criticized the idea of 'group selection', which can be caricatured as the idea that traits evolve 'for the good of the species'. He emphasized how natural selection can only act on individuals; his book was one of a clutch in the 1960s that led to the formulation of the gene's-eye view of natural selection. This change of perspective would bring evolutionary thinking back to the basics of selectionism and population-thinking, as first proposed by Darwin.

These are topics that are standard fare in any biology or evolution course today, so it is with some difficulty that I first appreciated its importance when reading his book. It's a problem that my roommate, a big movie buff, complained about in a different setting: when he watched a classic Western for the very first time, he thought that it was cliched and boring, using all the tropes and tricks found in every Western outlaw film. Why did people say that it was such a great movie, then? Because it was the first to do these things, and all the other films in the genre since then have been imitating it. A similar situation awaits the modern reader of Adaptation - we've been steeped so thoroughly in the gene's-eye view by popular science writers such as Richard Dawkins, so much so that we might think that Williams's book is just stating the obvious. In the early 1960s, though, biologists could still seriously speak of groups or whole populations as having adaptive traits, i.e. that natural selection could work at the level of the group. The demolition of this so-called 'group selection' was so complete that even today the term still has a whiff of taboo. The anthology Group Selection published in 1971, and edited by Williams, gives a good overview of the transformations in biological thinking during this period, through selections from the original literature.

Williams is also known for his theory of aging, or senescence. Growing old is a problem, not just for people, but for science. Why should an organism age? Why not be immortal? Wouldn't a gene for immortality be selected for? He explained it in evolutionary terms as a trade off - suppose a given trait allowed the individual to be more fecund earlier in life, but at a cost to it when it grew older. That trait would be selectively favored, because in a natural setting, the probability of reaching old age is low given the pressures of predation and the harsh environment. If you knew that you were going to die young, wouldn't you empty out your bank balance, too? This observation is borne out in humans, as modern medicine has allowed us to increase our lifespans way beyond what we would expect to live in the wild. As a result, so-called 'diseases of aging' are becoming ever more prevalent. In the mid-1990s, Williams and a medical doctor, Randolph Nesse, collaborated on a book that explored just this topic for a general audience, Why we get sick: The new science of Darwinian medicine. It was indeed a new science, which even today is still experiencing growth, but it traced its beginnings to Williams's theory of aging which he first proposed in the 1950s.

Aside from his original writings, Williams also served editorially at the Quarterly Review of Biology - the main book review for the biological sciences - for over forty years. He was not a flashy popularizer in the mold of Dawkins or Stephen Jay Gould, but his books remain readable and very relevant today. They are still among the best introductions for general audiences or biology students to gene-centered thinking, which has become populated by modelers and theoreticians to the extent that the basic biology can be obscured. In science, primary sources and original writings often become obsolete - because important ideas are repackaged and worked over and sometimes even overturned. It then becomes unnecessary to go back to the original publication because it doesn't necessarily improve your understanding. To understand newtonian mechanics, for example, one doesn't read the Principia but instead the latest version of some college textbook which (hopefully) incorporates refinements in our understanding and pedagogy that have been accumulated in the centuries since Newton. For biology, though, and with Williams in particular, the philosophical nature of these changes in theoretical framework, and their frequent subtlety, make the original texts worthwhile to read, and certainly even today can still stimulate renewed debate and appreciation.

Obituaries:
Chronicle of Higher Education
Edge Magazine
Wikipeida article