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Robert Trivers, Richard Dawkins, Daniel C. Dennett, Niles Eldredge, Randoph Nesse, M.D., Carl Zimmer

EDITOR NOTE: A strong case can be made that in the past 50 years, the two most influential evolutionary theorists are George C. Williams and Robert Trivers. Here, Trivers remembers Williams ...


In memory of George C. Williams

The last time I spoke with George Williams was in 2002 when I called about something and he told me he had pre-Alzheimer's. There were simple memory tests now that were diagnostic, he said. In the background I could hear his wife Doris saying something and George said, "Doris always tells me not to tell people" and continued by saying that what he first noticed is that all words starting with capital letters were disappearing from his mind — arbitrary words for cities, buildings, people and so on.

A few months later, I sent my Selected Papers book but I never heard from him. He was gone. The person I felt for was Doris, a beautiful woman about half his size, and a very welcome compliment to him. It is those closest to someone with Alzheimer's who often suffer the most but George had a sweet disposition that, I hear, greatly reduced the cost to those closest to him.

We last saw each other at the William Hamilton memorial session at Amherst in 2000 during the meetings of the Human Behavior and Evolution Society, at which both of us spoke. He was sitting behind me while others preceded us and I could hear Doris saying, "Now, George, don't do what you are thinking of. Just tell the stories you have about Bill, don't do it." So I was full of anticipation when George got up because I knew he was surely going to do exactly what his wife thought was a bad idea. George gets up and says "I wish Bill were here today, because I have a bone to pick with him".

And then he went and picked that bone for the entire talk. It had to do with the evolution of sex and patterns of evidence that George had pointed out years ago that contradicted (so George said) aspects of Bill's parasite approach. I thought it was wonderful. There were those that said it was inappropriate and why didn't he tell stories, but I thought it was perfect for the occasion, both vintage George Williams — no wasted motion with that organism! — and a tribute to the enduring importance of Bill's ideas.

My first contact with George was when as a graduate student. I sent him my chapter then in press on "Parental investment and sexual selection". When I wrote the paper I had completely forgotten that a key portion of the argumentation came right out of George's 1966 book, Adaptation and Natural Selection. I had only relearned this when I reread his book in preparation for teaching my first course on social evolution. There was "sex role reversed" species (as well as female choice for genes and investment) and the relevant pages were full of underlining and marginal comments by me. None of this was acknowledged in the chapter I was sending him, so I pointed this out and said I would try to put some in before the book was printed. I was therefore feeling a little nervous when a letter came from George Williams. I braced myself for an unpleasant experience.

Instead, I found one of the warmest and most generous letters I have ever received. Among other things, he said my paper had rendered obsolete a chapter in his own forthcoming book on "Sex and Evolution", namely the one on differential mortality by sex, which chapter he enclosed. He said nothing about not being properly cited but dealt only with scientific content. His chapter had my essential insight regarding male mortality — that higher variance in male reproductive success would often select for traits more costly in survival. The larger book was the first to systematically explore the consequences of seeing that sex usually had an immediate 50% cost in every generation (compared to asexuality) which cost had to be overcome in any successful model.

I invited him to Harvard in 1974 and he lectured on his ideas on sex. I do not say he was shy so much as reserved, but with a warm smile and sense of humor. A classic I use often occurred while telling me about the joys of grand-fatherhood. "If I could have figured out how to have grandchildren without having children first, I would have done so." I knew just what he meant — high relatedness, no work! Or as Melvin Newton (Huey's brother) puts it, "You can serve them ice cream for breakfast, what do you care?"

Having started with the evolution of senescence in 1957 in later life he tackled Darwinian Medicine, memorably saying that he did not think there was any compound — arsenic included — that was not beneficial if given in sufficiently small doses. This was almost surely an overstatement but a bracing and useful one. His knowledge of biology was so deep that he is the only person I know of to have predicted in advance the existence of an entire category of selfish genetic elements — androgenesis in which paternal genes eject maternal ones and take over the genome of an organism, a system now known from three very different groups of organisms.

He was a beautiful man, with a very simple and clear style of thinking, in a warm and humble personality.


George Williams was one of the great evolutionary thinkers of my lifetime. Universally respected, he was a tall, shy, diffident scholar, rather like W D Hamilton in character, his legendary wisdom seeming somehow enhanced by a physical resemblance to Abraham Lincoln. A wonderful scientist and a great gentleman, sadly missed.


I find that I really can't improve on what I said before in The Third Culture (on Edge). I think I nailed it then. That someone so incisive, such a corrosive corrector of sloppy thinking, could be so gentle and generous is something of a marvel. He was a lover of truth, not a lover of his own perspective and reputation.


[Dennett's 1995 comments:]

As other people have said, George Williams is the Abraham Lincoln of his field. He has a wonderful, laconic, pithy way of talking, and he seems to be an amazingly astute and clearheaded thinker. Reading George Williams showed me for the first time how hard it is to be a good evolutionary thinker, and how easy it is to make simple mistakes. Again and again, Williams issues his pithy little correctives to otherwise superficially good ideas and just calmly, firmly, wipes them out. Then you realize that this is a harder game to play than any of us realize, and George plays it better than anybody else in the world.

His main contribution, of course, was blowing the whistle loud and clear on the idea of "good for the species." In his 1966 book, he saw that Wynne-Edwards' — and others' — ideas, which were very familiar fare in the textbooks and popular treatments of evolution, had to be wrong. This was a wake-up call. Williams pointed out that it's not "What's good for the species is good for the organism (or vice versa)"; it's "What's good for the gene is good for the gene." Usually, other things being equal, what's good for the gene is good for the organism — and thus, you might say, for the species. But the gene is in the driver's seat.


Though he edited a major intellectual journal, for much of his career George really did not know much about the publishing world outside of the domain of university presses. He was incredulous when I told him, during a casual breakfast conversation in Sweden many years ago, that publishers do in fact offer advances on books! Indeed, I presume you got him one.

I never had the chance to tell him this, but many years later, while working on the Darwin exhibition, I found out that Darwin was one of the first authors known to have obtained an advance —  for the Origin of Species —  so upset was he that he apparently never received a farthing for his immensely popular Journal of Researches (aka Voyage of the Beagle)first published 20 years earlier. I can imagine George grinning at that little piece of news.


It is wonderful that you recall our first meeting. I remember your skepticism at first about our book, and George's skepticism about anyone in a fancy suit, and how, when George left the room, you said, "Someone told me George IS somebody. Is that true?" . You were by no means alone in not knowing his accomplishments. He never advertised himself. He just did his work. It is a great shame that is over, but a select few live on in their work. George is one of them.

Here are my extended comments on George:

George C. Williams died on September 8, 2010 at the age of 84. One of the most important biologists of the 20th century, his influence came not from big grants, flashy talks, magazine articles, or scores of graduate students. Instead, he pursued methodical thinking about important questions and distilled his conclusions into crystal clear prose. His approach was consistent. He would be struck by some apparent contradiction between fact and evolutionary theory, and work on it until a resolution emerged. Senescence, sex, menopause, and vulnerability to disease, all are hard to explain in evolutionary terms. In each case, he thought and thought, eventually coming up with a major contribution.

My tribute to his influence on the field and my life is in: Nesse, RM: Maladaptation and natural selection. Quarterly Review of Biology 80(1):62-70, 2005. In that article, I noted that he had a cognitive anomaly, I called it "Williams vision," that made it very difficult for him to blind himself to evidence that contradicted his ideas. Instead of marshaling all the evidence for his theory, and disparaging the rest, his work is a model of objectivity. In that article I also argued that he was neither an adaptationist nor an anti-adaptationist; if anything, he was a "maladaptationist." We shared a fascination with aspects of bodies and minds that just didn't seem to make sense in evolutionary terms. Gradually, we figured out ways of providing evolutionary explanations for many such traits. Now, scores of researchers are expanding the field of evolutionary medicine. In typical fashion, he insisted on naming the field "Darwinian medicine," because that term was slightly more accurate, too bad about readers who might better accept a blander designation. He is, and will always be, the father of Darwinian medicine.

Many other commentators will confirm the magnitude of his influence. Here, I want to offer a more personal perspective. His 1957 paper on senescence and antagonistic pleiotropy changed my career. I started trying to figure out senescence in 1968, as a sophomore at Carelton College. It is highly heritable and highly deleterious, so why didn't natural selection reduce its influence? Days spent paging through volume after paper volume of Index Medicus turned up all kinds of great ideas about aging, but not George's article. So, I wrote a paper arguing that senescence might have evolved to speed up the evolution of a species. My professors, not having read Williams, 1966, loved it.

In 1983, after I had joined the medical faculty at Michigan, I was spending lots of time with evolutionary biologists. It was clear they had a scientific foundation for understanding behavior that was lacking in psychiatry. Finally I got up my nerve and told them my theory about senescence. They laughed and laughed. Hadn't I ever read Williams, 1957? Didn't I know about the problems with group selection? I realized then that a truly excellent medical education had left me fundamentally ignorant about why organisms are the way they are, and why diseases exist. Once I recognized that senescence had at least two potential evolutionary explanations, I wanted to try to explain everything else about the body that seemed suboptimal. Why is obesity so common? Why is the birth canal so narrow? Why hasn't natural selection eliminated the genes that cause schizophrenia?

I started looking for an evolutionary biologist who might want to collaborate on this project. It was my great good fortune that Michigan was at the center of new evolutionary thinking in the 1980s. Richard Alexander and his grad students meet weekly for riveting seminars, and Bill Hamilton and George Williams came by often. Colleagues at Michigan, some of the best and brightest evolutionary biologists you could hope to find, were remarkably patient with my often embarrassingly ignorant questions. When I finally met George, it turned out that he had been thinking along similar lines; he was looking for a physician collaborator. We spent months talking and exchanging notes, trying to find evolutionary explanations for cancer, schizophrenia, and Alzheimer's disease. The process was frustrating. Finally, in a simple but profound reframing, we recognized that diseases do not have evolutionary explanations. Instead, we turned our focus to aspects of the body that leave us vulnerable to disease. Then we made progress.

Our first major paper came out in 1991 with the grand title, "The Dawn of Darwinian Medicine." The bravado turned out to be somewhat justified. The field has grown exponentially. We wrote our 1994 book, Why We Get Sick: The New Science of Darwinian Medicine in an engaging style, hoping that people would read it just for fun. That has worked out well.

Hundreds of publications on aspects of evolution and medicine are now available. In 2009, second editions of major edited volumes and a new textbook were published. A web journal, The Evolution and Medicine Review, provides a central information source. The field is healthy and growing, with one exception: medical schools seem to have no way of incorporating yet another basic science into the curriculum, not even something as fundamental as evolutionary biology. This isn't really surprising. There are no evolutionary biologists on medical school faculties to explain what is missing, and few evolutionary biologists bring in the kind of big grants that make medical school deans eager to start new departments. With time, undergraduates now learning about evolution and medicine will become deans. In the meanwhile, things are developing nicely everywhere else.

George was a wonderful collaborator. We wrote the book by sending drafts of chapters back and forth on computer disks. One of us would write a first draft, the other would completely rewrite it. We agreed on a simple method for staying friends. The revising author could make whatever changes seemed sensible, but was not allowed to look back at his earlier draft. After half a dozen iterations for each chapter, we forgot about who came up with what. We argued at length about many issues, allergy, defenses, menopause, senescence, and morality. It was real back and forth, the kind of very productive careful listening and hard arguing that carves away vagueness.

His wife Doris was not only his steady and loving companion, but a vigorous early research collaborator, and my friend as well. Their joint paper in 1957 on the evolution of altruism came close to describing inclusive fitness, but they veered away and relied instead on a variation of group selection. In typical fashion, they explicitly avoided the engaging word "altruism" and instead advocated using the more neutral terms "social donors" and "social non-donors." As memes, these were non-starters; selfish genes displaced them completely.

Some have described George as a bit curmudgeonly. I loved his directness. He was never trying to manipulate, always trying to understand. If he thought someone was wrong, he felt it was his obligation to talk with them to figure out what was correct. He was always polite, but he didn't pretend to agree just to be congenial. Not everyone liked that.

There was also a private side of George that few know about. For instance, he devoured fiction and history. He wrote a long work of fiction, an Icelandic saga. Eventually biographers will tell us more, but he has made the task challenging. He had the habit of keeping all of his notes and Icelandic. Such a characteristic quirk.

It was Alzheimer's disease that took him. The irony is nearly unbearable. We spent hours discussing why such a devastating disease should be so extraordinarily common. The obvious answer, of course, is that selection is weak at advanced ages. The alternative hypothesis, while unlikely, also needed consideration. Perhaps some aspect of the mechanism that causes Alzheimer's disease offers some kind of advantage. For decades I have asked neurologist friends for their thoughts about this, without much luck. Then, just a few months ago, an article appeared about beta-amyloid (Soscia, et al., PLoS ONE, 2010) , the substance contained in plaques on the neurons of those with Alzheimer's disease. It turns out to be quite a potent antimicrobial.

How George would've loved that! We would have talked for hours about what this means. Are infectious processes involved in Alzheimer's disease? Or is beta-amyloid merely a response to inflammation? Or is it an epiphenomenon? We also would have talked about the failure of drugs intended to disrupt beta-amyloid formation. In publications just last month, they were shown to be worse than useless, they speed the progression of Alzheimer's disease. And yet, in all of reports of the failure of the drug, there was no mention about the new evidence that beta-amyloid might have some useful function. I wish I could talk with George about all of this, and what we can do about it. I do so miss him.


...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... [MORE]

Back to: GEORGE WILLIAMS (1926-2010)

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