By JERRY COYNE - WEIT
Added: Mon, 30 Aug 2010 18:24:38 UTC

I don’t know what’s gotten into E. O. Wilson. He’s certainly the world’s most famous evolutionary biologist, and has gone from strength to strength over the years, winning two Pulitzer Prizes, writing great general books on not only ants but conservation and social behavior. And he’s kept his hands in the ant work, producing any number of technical papers and monographs. Frankly, I don’t know how he does it. I haven’t always agreed with what he says—I think he overreached with the sociobiology stuff, for instance—but you have to admire the guy’s diligence, breadth, dedication to conservation, and sheer workaholism.

But now Wilson, along with some collaborators like David Sloan Wilson and Martin Nowak, is definitely heading off on the wrong track. They’re attacking kin selection, not only maintaining that it has nothing to do with the evolution of social insects, but is a misguided way to look at evolution in general. And they’re wrong—dead wrong.

Read the rest of Jerry's excellent article here

And here are my own notes on the infamous Nowak/Wilson paper, written a few weeks ago when I first read it:-

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 1979). Kin selection is not a subset of group selection, it is a logical consequence of gene selection. And gene selection is (everything that Nowak et al ought to mean by) 'standard natural selection' theory: has been ever since the neo-Darwinian synthesis of the 1930s. 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 synthesis ever since Fisher and Haldane, but had been largely overlooked because people (with the exception of those two geniuses) didn't think about collateral kin.

Another way of expressing what was logically inherent in the synthesis is Hamilton's rule, rB>C: a gene for altruism will spread if the cost to the actor, 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. Haplodiploidy always was a side issue, so nothing has changed there.

Finally, Nowak et al do Darwin an injustice, in discussing his theory of the evolution of sterile worker adaptations. 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 pure inclusive fitness theory, nineteenth century version. Alex Kacelnik points out to me that kin selection is the only way in which worker adaptations such as soldier jaws and honeypot abdomens – phenotypes that are never expressed in reproductive individuals – could have evolved. 'Colony selection' and 'superorganisms' don't do the trick. You have to talk about shared genes in individuals, with conditional phenotypic expression.

Richard Dawkins

TAGGED: BIOLOGY, EVOLUTION, SCIENCE