The original robots
We are haunted by Hamilton. William D. Hamilton specifically, an evolutionary biologist who died before his time in 2000. We are haunted because debates about his ideas are still roiling the intellectual world over a decade after his passing. Last summer there was an enormous controversy over a paper which purported to refute the relevance of standard kin selection theory. You can find out more about the debate in this Boston Globe article, Where does good come from? If you peruse the blogosphere you’ll get a more one-sided treatment. So fair warning (I probably agree more with the loud side which dominates the blogosphere for what it’s worth on the science).
What was Hamilton’s big idea? In short he proposed to tackle the problem of altruism in social organisms. The biographical back story here is very rich. You can hear that story from the “horse’s mouth” in the autobiographical sketches which Hamilton wrote up for his series of books of collected papers, Narrow Roads of Gene Land: Evolution of Social Behaviour and Narrow Roads of Gene Land: Evolution of Sex. For the purposes of the issue at hand the first volume is obviously more important, but the second volume has an enormous amount of personally illuminating material because of Hamilton’s untimely passing in 2000 before it could be edited. In Ullica Segerstrale’s Defenders of the Truth and Oren Harman’s The Price of Altruism Hamilton looms large as a major secondary character in the narrative. The Altruism Equation, A Reason for Everything, and The Darwin Wars, all give him extensive treatment, both his scientific ideas and relevant biographical context. Hamilton’s scientific influence on Richard Dawkins was enormous. There are nearly fifty references to him in both The Selfish Gene and The Extended Phenotype. In writing his obituary Dawkins began: “W. D. Hamilton is a good candidate for the title of most distinguished Darwinian since Darwin.”
In terms of the details of his science, Hamilton proposed that genetic relatedness between individuals can explain altruism within groups. In this way Hamilton reduced a phenomenon which had often been explained as a group-level one (e.g., “for the good of the species”) to an individual-level one (e.g., “for the good of the individual/gene”). According to Hamilton when he was a young scientist in the early 1960s most people did not perceive this problem to be a problem at all, and he had difficulty finding support for this line of research, and was in fact warned off it by his superiors. The end culmination of those early years of lonely introspection were two dense, abstruse, and difficult papers (in part due to their peculiar notation), The genetical evolution of social behaviour – I and The genetical evolution of social behaviour – II. But the basic heuristic at the heart of these papers was condensed earlier in a short essay in The American Naturalist as Hamilton’s Rule:
rB > C or rB – C > 0
There is a new paper in Nature which is a full frontal attack on the utility of William D. Hamilton’s inclusive fitness framework in explaining eusociality. Martin A. Nowak, Corina E. Tarnita, & Edward O. Wilson are the authors. Wilson is famous in large part for his authorship of Sociobiology: The New Synthesis, and is arguably the doyen of American organismic biology. He is both an active scientist, and, a premier public intellectual. So with that in mind, I notice that Dienekes Pontikos alludes to “E.O. Wilson’s change of mind about group selection.” This is conventional wisdom, but it is I think wrong (though from what I can tell Wilson has not done much to disabuse the press of the notion). In Defenders of the Truth Ullica Segerstrale notes that Wilson did not expunge group selection thinking even in Sociobiology. In Evolution for Everyone David Sloan Wilson recounts that it was in fact E. O. Wilson who pointed out a group selective interpretation of data he was presenting at a conference, helping to push him early on in a rather unfashionable direction. From what I have heard Wilson always believed that the empirical data was not adequately explained by a pure inclusive fitness model, and simply waited until things shook out before pushing back with more theoretically trained colleagues who had the same skepticism.
From page 30 of Sociobiology: