Selection of and in groups

By Razib Khan | July 9, 2012 9:45 pm

Eric Michael Johnson has an excellent round-up of and commentary upon recent debates about “group selection” (also, a decent primer on the basics). If there is one major issue I might take with the narrative outlined, it is the idea that E. O. Wilson has made a recent about-face in regards to group selection, going from a skeptic to a believer. In Defenders of the Truth: The Sociobiology Debate Ullica Segerstrale pointed out that this was obviously not the case even in Wilson’s grand book Sociobiology. If you don’t accept this, David Sloan Wilson seems to implicitly confirm Segerstrale’s position in his semi-autobiographical book Evolution for Everyone. Finally, I’ve heard it from acquaintances at Harvard that it was an open secret that E. O. Wilson was skeptical of the “Hamiltonian orthodoxy” ascendant in evolutionary biology and ecology. The controversy over the notorious paper, The evolution of eusociality, was years in the making. From what I gather many of Wilson’s colleagues at Harvard were skeptical that he comprehended the depths of the mathematics which he promoted to support his more intuitive empirically informed skepticism of the power of inclusive fitness. It isn’t an unheard of sin for an eminent empiricist to go shopping for a theoretician to add some luster to his logic, but it does frame the reason that some of Wilson’s colleagues seem so irritable about the whole affair.

I think this is at the heart of the acidity in Richard Dawkins’ review of Wilson’s latest book, The Social Conquest of Earth. If I, a nobody, could hear through the grapevine that Wilson is lending his name to ideas which he doesn’t have a deep understanding of in terms of the mathematical superstructure, then Dawkins surely knows as well. This is what I believe is at the heart of this section of Dawkins’ review: “But one can make a good case that the 2010 paper would never have been published in Nature had it been submitted anonymously and subjected to ordinary peer-review, bereft of the massively authoritative name of Edward O Wilson. If it was authority that got the paper published, there is poetic justice in deploying authority in reply.” Wilson has a great deal of authority as an entomologist and a synthetic thinker, but not as a formal theoretician. Obviously there are many great partnerships between theoreticians and empiricists in science, but it strikes me that many feel that the relationship between Wilson and his two mathematically oriented co-workers was too convenient by a half. In his gut Wilson long knew that inclusive fitness was not all there was, and here were these biologically naive mathematicians willing to stand by his side. It makes fascinating sociology of science.

Finally, you should also check out Steven Pinker in Edge, The False Allure of Group Selection. Pinker’s essay is fine as far as it goes, but the responses are also excellent. As it happens I agree with the group selection skeptics that the framework is often incoherent, muddled, and lacking in robust empirical illustration. On the other hand, do read the responses of Joe Heinrich, Robert Boyd, and Peter Richerson. They make cogent defenses of the multi-level framework, but to be frank I think that E. O. Wilson has kind of lost it at this point. He claims too much for multi-level selection, and I do wonder if his fellow travelers are starting to wonder if he’s a liability. On the other hand I do think that Pinker sells group level dynamics short for human cultural modeling. I will be frank, I think inclusive fitness and reciprocal altruism are not sufficient to explain human cultural complexity. This does not mean that I accept inter-group selection as “the answer.” But even if it’s a difficult and dodgy framework, it is an alternative.

One last thing. My personal experience is that this is an issue where discipline and “who you know” matters in terms of what you think as to the science. I’ve encountered eminent behavioral ecologists who think Wilson is in the right. On the other hand, there are many others, especially in genetics, who are absolutely skeptical. Overall I suspect that the apologists for multi-level selection, especially in terms of biological process, are in the minority. But probably not a tiny minority.

Addendum: Readers may be curious that W. D. Hamilton reported that his paper, Innate Social Aptitudes of Man: An Approach from Evolutionary Genetics, which Eric Michael Johnson references as evidence of Hamilton’s rapprochement with group selection, was referred to as the ‘fascist paper.’ For more in that vein see Narrow Roads of Gene Land: The Collected Papers of W. D. Hamilton Volume 1: Evolution of Social Behaviour.

MORE ABOUT: Social Evolution
  • Michael Watts

    I have one mostly-undeveloped thought on group selection:

    In (I think) The Selfish Gene, there’s a discussion to the effect that a group of monkeys might end up in a no-grooming equilibrium (bad for the group) or in a reciprocal-grooming equilibrium (good for it), and that you can’t predict which way will occur because the monkeys don’t choose what to do based on the good of the group. My thought is: granting that, can I still say that, if I look in on the planet at various points in time, I’m more likely to see reciprocal-grooming groups (of whatever species happen to exist at the time) than no-grooming groups, because a group in no-grooming equlibrium is more likely to die out than one that practices reciprocal grooming?

    I guess my intuition is that over time, groups with “good group behavior” might be more prevalent in the world than groups with “bad group behavior”, since although they might evolve more or less by coincidence, they exist for longer spans of time.

    Is there someone making a case for some formalism of that intuition? Am I hopelessly confused?

  • Nathan Taylor

    Eric Michael Johnson makes a great comment that “multilevel selection is little more than a rebranding of Hamilton’s inclusive fitness (albeit the “enhanced” 1975 version).” The albeit is the key here. Dawkins and some of the others so up in arms are using inclusive fitness in the narrow and original version of rB>C from 1964. Not the Price equation co-variance model from 1975 which allows non-relation selection. In fact it’s not quite clear if you using the 1964 version as well when you say “inclusive fitness and reciprocal altruism are not sufficient to explain human cultural complexity”. The “enhanced” 1975 version of inclusive fitness would include the possibility that humans evolved from group on group conflict without necessarily having a narrow kin selection relatedness dynamic.

  • TJ

    It is not a secret that Wilson doesn’t understand the math that he thinks supports his argument. He told Jonah Lehrer as much in interviews for the hagiography that ran in the March 5, 2012, issue of the New Yorker.

  • Grundle

    Group selection won’t be obvious to geneticists, but it’s already obvious to anyone well-versed in epigenetics.

  • BDoyle

    @ #1: I think most people would call mutual grooming “reciprocal altruism.” Both participants get a boost to their individual fitness, and it’s not really controversial to think that the behavior could result from selection pressure on individuals. It’s only when trying to explain behavior that is clearly bad for the individual that some people resort to multi-level selection.

  • Chad

    How is it obvious to anyone well-versed in epigenetics? This sounds like the typical abuse of Epigenetics, which has come to serve as a refuge of all kinds of bad ideas. Epigenetics is a part of Genetics. Its simply a regulatory mechanism.

  • Razib Khan

    #6, ditto. but perhaps we are in the darkness…


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About Razib Khan

I have degrees in biology and biochemistry, a passion for genetics, history, and philosophy, and shrimp is my favorite food. In relation to nationality I'm a American Northwesterner, in politics I'm a reactionary, and as for religion I have none (I'm an atheist). If you want to know more, see the links at


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