If you’ve read the critiques of the Nowak et al. paper on kin selection that I highlighted this morning, you may have noticed a conspicuous absence among the authors: the name of Richard Dawkins. Why, as the most famous exponent of kin selection since W. D. Hamilton, didn’t he co-author one of the critiques?
It turns out that nobody asked him. I think this was simply an oversight, because all of us simply assumed that Richard would be penning his own criticism. He didn’t, but he did write a brief piece on Nowak et al. for New Scientist, which he decided not to publish. I’m posting it here with his permission, along with a link to his excellent paper about common misunderstandings of kin selection. I’m not sure whether Richard will answer comments from readers, but you can certainly pose them below. Here’s his take on Nowak et al.:
This is no surprise. Edward Wilson was misunderstanding kin selection as far back as Sociobiology, where he treated it as a subset of group selection (Misunderstanding Two of my ‘Twelve Misunderstandings of Kin Selection‘: Zeitschrift für Tierpsychologie 1969). Kin selection is not a subset of group selection, it is a logical consequence of gene selection. And gene selection is ‘standard natural selection’ theory. Inclusive fitness theory is not some kind of supernumerary excrescence, to be ‘resorted to’ only if ’standard natural selection theory’ is found wanting (Misunderstanding One). On the contrary, inclusive fitness theory is one way of expressing what was logically inherent in the neo-Darwinian synthesis ever since the 1930s but had been largely overlooked because people didn’t think hard enough about collateral kin. ‘Standard natural selection theory’ MINUS inclusive fitness would be like Euclidean geometry minus Pythagoras’ theorem.
Another way of expressing what was logicially inherent in the synthesis is Hamilton’s rule, rB>C: a gene for altruism will spread if the cost to the altruist, C is exceeded by the Benefit to the recipient, B, devalued by the coefficient of Relatedness, r. If you think, as Nowak et al. do, that ‘Hamilton’s rule almost never holds’, that simply means you haven’t been measuring B and C carefully enough. r is not the only term in Hamilton’s inequality. B and C matter too, and your game theoretic considerations are subsumed within them.
Perhaps most irritating is Nowak et al.’s concentration on haplodiploidy, which, in Hamilton’s original paper was a throwaway side-issue, interesting enough to pique the interest of generations of students, but not in any sense central to his paper. Of course Hamilton was well aware that eusociality is present in diplo-diploid animals, exactly as inclusive fitness theory would predict given appropriate B/C ratios. Indeed, Hamilton himself put forward an ingenious theory of the evolution of eusociality in termites, predating by seven years the version usually attributed to Bartz (attributed by Hamilton himself, indeed, with characteristically absent-minded generosity as I described in The Selfish Gene, second edition p 317).
Finally, Nowak et al. do Darwin an injustice, in discussing his theory of the evolution of worker sterility in social insects. They paraphrase Darwin’s ‘well-flavoured vegetable’ analogy. Let me quote it exactly: “Thus, a well-flavoured vegetable is cooked, and the individual is destroyed; but the horticulturalist sows seeds of the same stock, and confidently expects to get nearly the same variety . . . I do not doubt that a breed of cattle, always yielding oxen with extraordinarily long horns, could be slowly formed by carefully watching which individual bulls and cows, when matched, produced oxen with the longest horns; and yet no one ox could ever have propagated its kind. Thus I believe it has been with social insects . . .” It is true that Darwin goes on to phrase his idea in terms of benefit to the colony, but his analogy of the long-horned (castrated) oxen could not be clearer. No colony is involved. This is early inclusive fitness theory. It is entirely clear that, if Darwin had been alive to read Hamilton on social insects, he would have embraced inclusive fitness, not as an add-on to natural selection theory but as the logical way to express it in the age of the gene.