E. O. Wilson mistakenly touts group selection (again) as a key factor in human evolution

As most of you know, Edward O. Wilson is one of the world’s most famous and accomplished biologists.  He was the founder of evolutionary psychology (known as “sociobiology” back then), author of two Pulitzer-Prize-winning books, one of the world’s great experts on ants, an ardent advocate for biological conservation, and a great natural historian. His legacy in the field is secure.

So it’s sad to see him, at the end of his career, repeatedly flogging a discredited theory (“group selection”: evolution via the differential propagation and extinction of groups rather than genes or individuals) as the most important process of evolutionary change in humans and other social species. Let me back up: group selection is not “discredited,” exactly; rather, it’s not thought to be an important force in evolution.  There’s very little evidence that any trait (in fact, I can’t think of one, including cooperation) has evolved via the differential proliferation of groups.

In contrast, there is a ton of evidence for an alternative explanation for cooperation: kin selection, the selection of genes based on how they affect not just the fitness of the individual, but the fitness of relatives that share its genes.  Features like parental behavior, parent-offspring conflict, sibling rivalry, and preferential dispensing of favor to relatives, as well as features like sex ratios in insects—all of these are all easily explained by kin selection.  And many aspects of cooperation can easily be explained by individual selection: individuals that live in small groups, especially those in which one can recognize group members, can evolve cooperation as an individual good based on reciprocity: the “I scratch your back, you scratch mine” hypothesis.  And, as I’ve discussed before, the cooperative and “altruistic” behavior seen in our own species shows many features suggesting that it evolved via individual or kin selection and not group selection.

I’ve covered this issue many times (e.g., here, here, here, here, and here), so I won’t go over the arguments again. Wilson’s “theory” that group selection is more important than kin selection in the evolution of social behavior (published in Nature with Martin Nowak and Corina Tarnita) was criticized strongly by 156 scientists—including virtually every luminary in social evolution—in five letters to the editor, and sentiment about the importance of group selection has, if anything, decreased since Wilson’s been pushing it.

But Wilson persists, to the detriment of his reputation. In a new piece at the New York Times “Opinionator” site, “The riddle of the human species,” Wilson continues to make the same argument that group (or “multilevel”) selection was a key force in making humans (and social insects) the socially complicated species they are.  Since his arguments are virtually identical to those published in NYT Opinionator piece last June, and in his book The Social Conquest of Earth (see part of my review here), I won’t dissect them in detail. I just want to highlight three points that I think make Wilson’s argument for group selection—and against kin selection—deeply misleading. I wouldn’t spend my time writing time-consuming critiques like this were Wilson not famous, influential, and given a big public forum in the New York Times. Someone has to address his arguments!

Here are Wilson’s errors (quotes indented), and my responses:

1. Wilson: Humans are a “eusocial species”:

. . the known eusocial species arose very late in the history of life. It appears to have occurred not at all during the great Paleozoic diversification of insects, 350 to 250 million years before the present, during which the variety of insects approached that of today. Nor is there as yet any evidence of eusocial species during the Mesozoic Era until the appearance of the earliest termites and ants between 200 and 150 million years ago. Humans at the Homo level appeared only very recently, following tens of millions of years of evolution among the primates.

My response:  “Eusociality” as defined by Wilson and every other evolutionist is the condition in which a species has a reproductive and social division of labor: eusocial species have “castes” that do different tasks, with a special reproductive caste (“queens”) that do all the progeny producing, and “worker castes” that are genetically sterile and do the tending of the colony. Such species include Hymenoptera (ants, wasps and bees, though not all species are eusocial), termites, naked mole rats, and some other insects.

But humans don’t have reproductive castes, nor genetically determined worker castes.  Wilson is going against biological terminology, lumping humans with ants as “eusocial,” so he can apply his own theories of “altruism” in social insects (i.e., workers “unselfishly” help their mothers produce offspring while refraining themselves from reproducing), to humans. But human cooperation and altruism are very different from the behavior of ants, most notably in our absence of genetic castes and genetically-based sterility associated with helping others reproduce. Human females aren’t sterile, and don’t usually refrain from reproduction just to help other women have babies.  My guess is that Wilson lumps humans with insects as “eusocial” because he wants to subsume them both under a Grand Theory of Social Evolution.

2. Wilson: Kin selection doesn’t work, ergo it certainly couldn’t have played a role in the evolution of eusociality and human cooperation.

Still, to recognize the rare coming together of cooperating primates is not enough to account for the full potential of modern humans that brain capacity provides. Evolutionary biologists have searched for the grandmaster of advanced social evolution, the combination of forces and environmental circumstances that bestowed greater longevity and more successful reproduction on the possession of high social intelligence. At present there are two competing theories of the principal force. The first is kin selection: individuals favor collateral kin (relatives other than offspring) making it easier for altruism to evolve among members of the same group. Altruism in turn engenders complex social organization, and, in the one case that involves big mammals, human-level intelligence.

The second, more recently argued theory (full disclosure: I am one of the modern version’s authors), the grandmaster is multilevel selection. This formulation recognizes two levels at which natural selection operates: individual selection based on competition and cooperation among members of the same group, and group selection, which arises from competition and cooperation between groups. Multilevel selection is gaining in favor among evolutionary biologists because of a recent mathematical proof that kin selection can arise only under special conditions that demonstrably do not exist, and the better fit of multilevel selection to all of the two dozen known animal cases of eusocial evolution.

My response:  There is so much fail here I don’t know where to start.  The first paragraph is basically correct except that Wilson omits “individual selection” along with “kin selection” as an accepted evolutionary process that can promote the evolution of cooperation. As I mentioned, selection on individuals in small groups can allow the evolution of cooperation without any need to invoke the unparsimonious process of differential group survival based on genes.

Wilson’s claim that the “special conditions of kin selection” demonstrably do not exist is an egregious and (I think) willful misstatement.  Kin selection can cause evolution whenever the genes in an individual benefit relatives that share copies of that individual’s genes, and can do so whenever the benefit of that behavior to the recipients, devalued by their degree of relatedness to the donor (a figure usually ranging between 0 and 1, but which can be related if an individual helps another less related to it than the average member of the population) is greater than the reproductive cost to the donor.  (“Hamilton’s rule”: rb > c.) That is known to obtain in many cases, and explains things like parental care, parent-offspring conflict, sex ratios in insects, and many other features (see the five letters in Nature mentioned above, which list some features of social behavior that clearly evolved by kin rather than group selection).

The mathematical “proof” given by Nowak et al. does not show that group selection is a better explanation than kin selection for social behavior in insects, for their “proof” does not vary the level of kinship, as it must if it could allow that conclusion.

The second egregious and false claim in this paragraph (a paragraph that’s the highlight of the piece) is that “multilevel selection is gaining in favor among evolutionary biologists” because of the Nowak et al. paper. That’s simply not true.  The form of multilevel selection adumbrated in that paper is, to my knowledge, embraced by exactly four people: the three authors of the paper and David Sloan Wilson. There is, and has been, no increase in acceptance of group or multilevel selection in the past ten years. The Nowak et al. paper has sunk without a stone, except to incite criticism by other biologists and excitement by an uncomprehending press.

3. Wilson: Eusociality in insects arose not via kin selection, but via the initial construction of a defended nest site.

The history of eusociality raises a question: given the enormous advantage it confers, why was this advanced form of social behavior so rare and long delayed? The answer appears to be the special sequence of preliminary evolutionary changes that must occur before the final step to eusociality can be taken. In all of the eusocial species analyzed to date, the final step before eusociality is the construction of a protected nest, from which foraging trips begin and within which the young are raised to maturity. The original nest builders can be a lone female, a mated pair, or a small and weakly organized group. When this final preliminary step is attained, all that is needed to create a eusocial colony is for the parents and offspring to stay at the nest and cooperate in raising additional generations of young. Such primitive assemblages then divide easily into risk-prone foragers and risk-averse parents and nurses.

My response:  Phylogenetic studies show that eusociality in Hymenoptera always originated in species whose females mated only once: this is a statistically significant result.  And that alone militates for kin selection as an important factor in eusociality: if a female founds a colony consisting only of full siblings (as is the case when she mated only once), they are more related to each other than if she had mated multiply. In the later case, colonies would consist of half-sisters or even more distant relatives, making kin selection less efficient.

Further, relatedness is high in virtually every species of eusocial insect with the exception of a few highly derived species of ants that have many queens.  The connection between relatedness and eusociality is exactly what we expect if kin selection is important in social evolution, and is not expected if Wilson’s nest-based group selection was important. The model of Nowak et al., which starts with the construction of such nests by single females who stay in the nests with their offspring, produces precisely the condition in which relatedness can promote the evolution of sterility and cooperation.  They argue that this relatedness is a consequence of their model and not a cause of eusocial evolution, but that’s unconvincing, for they do not vary the level of initial relatedness in their model.

*****

Wilson’s claim, the theme of his newest book, is that humans are both angels and devils: we are both selfish and cooperative species, and this combination of good and bad is what makes our species unique. (That’s not true, of course, because many species show that mixture of behavior. Lions, for instance, cooperate when hunting, but when males take over a pride they immediately kill all the female’s cubs, which are unrelated to them. And that, by the way, is due to kin selection, because those cub-killing males replace the cubs with new cubs containing their own genes, including the genes for killing cubs. Cub-killing could have evolved only by individual selection and not group selection, for while killing another male’s cubs is good for an individual, it’s bad for the group, forcing females to waste reproductive energy.)

Yes, we have both selfish and cooperative behaviors, though most of our “cooperative” behaviors that didn’t arise through culture arose through forms of selection that involve maximizing our reproductive output—individual and kin selection.  There is not a scintilla of evidence, in humans or any other species, that group selection has been responsible for the evolution of any adaptation.  In contrast, individual and kin selection have productively explained the evolution of “problematic” traits like altruism and cooperation. They have been tested and work.

Why does Wilson keep writing article and article, and book after book, promoting group selection? I’m not a psychologist, so I don’t know the answer. What I do know, though, is that his seeming monomaniacal concentration on a weakly-supported form of evolution can serve only to erode his reputation.  His theories have not gained traction in the scientific community. That doesn’t mean that they’re wrong, for, in the end, scientific truth is decided by experiment and observation, not by the numbers of people initially on each side of an issue. But the facts of science already show that Wilson is unlikely to be correct. What is sad is that, as a great natural historian, he doesn’t recognize this.

Wilson’s reputation is secure. It’s sad to see it tarnished by ill-founded arguments for an unsubstantiated evolutionary process.

h/t: Phil Ward, Laurence Hurst

70 Comments

  1. Alice Wonder
    Posted February 26, 2013 at 9:45 am | Permalink

    Well this post is way above my head. I’ve got some reading to do.

  2. DiscoveredJoys
    Posted February 26, 2013 at 9:57 am | Permalink

    I’m not keen on the idea of group selection because I see no heritable mechanism for it to work.

    On the other hand I think you could make a case for (mostly) sterile human females helping to raise the young of others – they’re called grandmothers. A sort of timebanded eusociality, but still a case of kin selection.

    • Gregory Kusnick
      Posted February 26, 2013 at 10:49 am | Permalink

      Grandmothers have not delegated their reproduction to collateral kin, as eusocial insects do; on the contrary, they’re helping to raise their own direct lineal descendants.

  3. Sunny
    Posted February 26, 2013 at 10:13 am | Permalink

    Spiegel has an interview with him:

    Interview with Edward O. Wilson: The Origin of Morals

    http://www.spiegel.de/international/spiegel/spiegel-interview-with-edward-wilson-on-the-formation-of-morals-a-884767.html

  4. gbjames
    Posted February 26, 2013 at 10:13 am | Permalink

    E.O. Wilson was a hero of mine back in the old days when “Sociobiology” was published. This is sad.

  5. Posted February 26, 2013 at 10:30 am | Permalink

    Thanks for taking the time to write this post, Jerry. It’s fantastic.

  6. Posted February 26, 2013 at 10:45 am | Permalink

    Hear, hear!!

  7. Aaron S.
    Posted February 26, 2013 at 10:47 am | Permalink

    Good explanation of how group selectionists went wrong: LessWrong – The Tragedy of Group Selectionism

    • gbjames
      Posted February 26, 2013 at 11:20 am | Permalink

      Thanks for that.

      • Aaron S.
        Posted February 26, 2013 at 11:30 am | Permalink

        You’re welcome. LessWrong is a very fun site to get lost in. My mind was blown by the “Quantum Physics Sequence.” It now seems so obvious that Many Worlds is the correct interpretation.

  8. Wayne Tyson
    Posted February 26, 2013 at 10:57 am | Permalink

    MS WT Scientific discourse WEIT 130226

    The saddest thing may be the tendency to engage in attaching significance to personalities rather than discussing the relative merits of different interpretations or meanings of observed phenomena. Tangled webs of convoluted speculations are red meat for academic posturing, but good old-fashioned intellectual discipline can be lost as one wanders off into the weeds too far.

    “Reputations” are irrelevant, but our obsession with them is all too common. “Competition” is a cultural concept, projected upon other organisms because “we” have not yet developed a language up to the task of accurately describing and communicating and distinguishing the crucial distinctions between what is really happening from what we believe is happening (“science” as religion). Driven by our egocentric tendencies to obtain the recognition of others as “King of the Mountain,” we hasten to diminish others to create the illusion of superiority.

    I, for example, may question Wilson and Coyne’s interpretations of natural phenomena, but frankly m’dears I don’t give a damn who said what–all I care about is The Issue. That neatly proscribes the inflammation that occurs when an observation carries the label of its author. Yeah, when pigs fly. Egocentrism reigns. It trumps absolutely.

    Worse, the labeling of interpretations with authors’ names sets up limitless opportunities for digression, always a safer haven than sticking to the issue and responding to the points made and sticking to the subject or ignoring relevant elements of statements previously made in the course of discourse. Or ignoring a statement absolutely. In the mind of the “ignorer,” avoiding the subject is “a win.”

    With respect to the issue, what ultimately drives evolution beyond survival and reproduction and context? Please replace this statement with the correct one.

    • whyevolutionistrue
      Posted February 26, 2013 at 11:48 am | Permalink

      Thank you for your concern, which is duly noted. I categorically reject your argument that the post above is more about personalities than issues, but the sociology of science is of concern to some.

      • Wayne Tyson
        Posted February 27, 2013 at 8:42 am | Permalink

        I did not say “that the post above is MORE about personalities than issues,” as you obviously DID discuss the issues in the post. But the post did emphasize Wilson’s being wrong in a public forum (where he may or may not be present to defend himself and his interpretations–or care to). This reminded me of what I believe is a widespread defect in the way much discourse is conducted about issues, particularly scientific ones, which leaves the issues raised unresolved.

        I would like to see more discussion about the issues raised, and I asked a question directed at, presumably, those with knowledge superior to mine, to wit,

        “With respect to the issue, what ultimately drives evolution beyond survival and reproduction and context? Please replace this statement with the correct one.” (Or confirm it?) I’m here to learn about evolution from the experts.

        • Diane G.
          Posted February 27, 2013 at 11:26 am | Permalink

          Google is your friend.

        • Posted February 27, 2013 at 6:47 pm | Permalink

          I think probably no one answered your question because it doesn’t really make any sense. Perhaps you could re-phrase it?

          • gbjames
            Posted February 28, 2013 at 5:22 am | Permalink

            I’m sure this is the case.

          • Wayne Tyson
            Posted March 5, 2013 at 9:33 am | Permalink

            What is it about the question that is confusing?

            • gbjames
              Posted March 5, 2013 at 9:37 am | Permalink

              The words don’t make it clear what you are asking.

              • Diane G.
                Posted March 5, 2013 at 8:43 pm | Permalink

                ROFL!!

        • Wayne Tyson
          Posted March 6, 2013 at 12:00 am | Permalink

          With respect to the inquiries concerning the following question:

          ““With respect to the issue, what ultimately drives evolution beyond survival and reproduction and context? Please replace this statement with the correct one.” (Or confirm it?) I’m here to learn about evolution from the experts.”

          I’m sure that everyone understands what survival and reproduction mean, so I will hazard a guess that the problem lies in the word “context.” By context I mean what might be called “environmental” context, but my use of the word is an attempt to hazard a guess that “environmental” might lead to a misunderstanding because of the semantic baggage that word has acquired in other, well, contexts. Perhaps “biological” context would be better, but that, too, might lead to misunderstanding because I see the context in which an organism is able to maintain a viable population as one that includes both biotic and “abiotic” factors. Ecological context might be better, but that word also has acquired considerable semantic baggage. I appeal to my betters to suggest better terminology.

          I am asking whether there are factors other than the “environment” that an organism exists within in viable populations or that cause it to cease to exist or decline that drive evolution.

          I hope I am guessing correctly, but in the absence of references to the specific faults of my question I have been forced to go into detail that I had presumed, incorrectly, would be understood without further elaboration. I have trouble reading minds, especially via generalizations.

          I look forward to any specific suggestions regarding just how my language should be rephrased (as initially requested), but I also am willing to explain myself further when notified of a specific word or phrase that renders the question confusing.

          • gbjames
            Posted March 6, 2013 at 5:20 am | Permalink

            I think you are asking if there is an environment (aka “factors”) in which creatures exist that isn’t part of the environment in which they exist.

            I have no idea how to respond to that idea. It sounds like something from theology to me.

            If I still don’t understand then you’ll need to take another stab at it. But I’m guessing that we are having trouble connecting because the idea itself is amorphous.

            • Wayne Tyson
              Posted March 7, 2013 at 2:28 pm | Permalink

              That’s not what I’m asking.

              • gbjames
                Posted March 7, 2013 at 8:19 pm | Permalink

                Then you need to try again. Fewer words, perhaps. And more focus.

          • Posted March 6, 2013 at 5:34 am | Permalink

            “what ultimately drives evolution beyond survival and reproduction and context environment?”

            What makes you think there should be anything beyond survival and reproduction and environment?

            So… maybe the answer is: Nothing.

            /@

            • Wayne Tyson
              Posted March 7, 2013 at 2:32 pm | Permalink

              You may be right. If so, how does a trait that does not interfere with reproduction or one (or the lack of one) that causes death before reproduction, drive evolution?

              • Posted March 7, 2013 at 3:11 pm | Permalink

                Not all traits are adaptive and evolution can occur in part by genetic drift.

                /@

              • Diane G.
                Posted March 7, 2013 at 3:11 pm | Permalink

                Whoa, you really don’t know the first thing about evolution. There’s a great book called Why Evolution is True

              • Posted March 7, 2013 at 3:14 pm | Permalink

                Ah, Diane. I was saving that for my next reply! ;-)

                /@

              • Diane G.
                Posted March 7, 2013 at 3:15 pm | Permalink

                Yay! It’s rare to beat Ant Allan to the punch. :)

              • Wayne Tyson
                Posted March 7, 2013 at 8:43 pm | Permalink

                “Not all traits are adaptive and evolution can occur in part by genetic drift.”

                Yes, of course. But mustn’t the traits that drift do so because an organism lived to reproduce, taking them along for the ride, as it were?

                Should we be thinking in terms of traits that prevent success rather than promote it? I’ve heard it said that “evolution is what’s left over after everything else is gone.”

                If this isn’t true can anyone explain why not?

          • Wayne Tyson
            Posted March 7, 2013 at 9:11 pm | Permalink

            Here is the statement considered to be faulty, modified as one person [thank you] suggested (“environment” meaning the entire complex of factors that act upon an organism):

            “With respect to the issue, what ultimately drives evolution beyond survival and reproduction and environment? Please replace this statement with the correct one. (Or confirm it?) I’m here to learn about evolution from the experts.”

            [Please note the last sentence in the preceding paragraph.]

            Another question. Do all of you [evolutionary biologists] believe that organisms require an assemblage of traits that match the conditions of environment (environmental context) closely enough to continue to exist as a viable species? Is environment the ultimate arbiter in extinction and survival?

            • gbjames
              Posted March 8, 2013 at 5:41 am | Permalink

              I find your “questions” disingenuous. You are fishing for something. It is unclear exactly what (but I have my suspicions).

              Strip out the useless cruft. What value is the phrase “With respect to the issue”? It means nothing. (What issue?). “What ultimately drives evolution beyond survival and reproduction and environment?” Well, you can look this up. Evolution is the differential survival of genes in populations over time. That’s definitional. What do you mean by “ultimately”? It is a confusing and useless word in this context.

              “Im here to learn from the experts.” That sure sounds like sarcastic snark. Similarly your phrase: “Do all of you [evolutionary biologists]“. Many (most) of us are not evolutionary biologists, at least not by formal training or trade. What is the value of the phrase, used as you do? The context of misleading questions sure seem to be searching for a crevice in which to insert a deity. Maybe Vishnu. Maybe Gaia. But I’ll put my money on Jebbus, given the geography.

              • Diane G.
                Posted March 8, 2013 at 11:59 am | Permalink

                Hear, hear!

            • Dan Styer
              Posted March 8, 2013 at 6:57 am | Permalink

              Wayne Tyson asks: “What drives evolution beyond survival and reproduction and environment?” where “‘environment’ means the ENTIRE complex of factors that act upon an organism” (my caps).

              The answer is right there within the statement of the question. If environment is defined as everything that acts upon an organism, then there the nothing that drives evolution beyond environment, by your own definition.

    • Marta
      Posted February 26, 2013 at 1:21 pm | Permalink

      Jeepers, Mr. Tyson.

      If you’re going to write a post about personalities vs. issues (that’s your point, yes?) it’s probably more effective if you’re not so pompously long-winded while you’re at it.

      • Wayne Tyson
        Posted February 27, 2013 at 8:43 am | Permalink

        I rest my case.

        • John Scanlon, FCD
          Posted February 27, 2013 at 9:49 am | Permalink

          Case dismissed.

  9. DV
    Posted February 26, 2013 at 11:20 am | Permalink

    “eusociality in Hymenoptera always originated in species whose females mated only once”

    Wow! Thanks for this gem. I did not know this. This would indeed be a very big clue that group selection is wrong (as if we need more clues when the principle of it doesn’t even make sense).

    • Posted February 27, 2013 at 2:08 am | Permalink

      Does the naked mole rat mate only once? Does she practice (if that is the word) sperm retention? Is there a “drone”/”stud” male? (She could mate more than once if it were always with the same male, always producing full siblings.)

  10. Posted February 26, 2013 at 11:46 am | Permalink

    It is often said that Wilson was an early supporter of kin selection, and that he has now changed his mind because of new evidence. But even in 1975 when he wrote Sociobiology, Wilson clearly did not UNDERSTAND kin selection. He treated it then as a form of group selection, which demonstrates completely muddled thinking. Kin selection is not something additional, added on to normal neo-Darwinism. Kin selection follows logically and inescapably from neo-Darwinism, although this wasn’t properly understood until Hamilton (1964). Hamilton’s Rule states that an altruistic trait will spread if rB>C. There are many specific instances where B (Benefit to receiver) and C (Cost to donor) are such that r (coefficient of relationship) cannot be used to predict altruism. But Hamilton’s Rule still applies.

    • Posted February 26, 2013 at 1:33 pm | Permalink

      Now that’s an insightful observation.

      Was there any contemporary criticism of this error in Sociobiology?

      /@

      • Daniel Engblom
        Posted February 26, 2013 at 1:49 pm | Permalink

        I have a vague recollection of Dawkins pointing out the errors in Wilson’s thoughts in at least the second edition of The Selfish Gene (the footnotes?).

      • neil
        Posted February 26, 2013 at 1:49 pm | Permalink

        Just curious. Would the observed spread of an altrusitic trait among population members for whom rB<C be considered evidence for group selection, or would this be indistinguishable from reciprocal altruism? What if the population did not have the mental processing power for r.a.?

        What I am struggling with here, and perhaps the reason we who are not evolution scientists are confused by this controversy, is whether there is a definitive test of the group selection hypothesis. I am assuming there is not, or this debate wouldn't drag on.

        • Posted February 26, 2013 at 4:08 pm | Permalink

          For straightforward models of group selection, the groups become different by genetic drift, and consist of individuals more closely related to each other (on average). The benefit to relatives is the increased survival probability of the whole group. Bookkeeping that, theoreticians such as George Price and also Aoki and Crow have shown that Hamilton’s condition r b > c is in fact also the condition for group selection to favor the altruistic allele. Hamilton’s equation applies to both kin selection and group selection. So we should not look for violation of that condition in group selection.

          • neil
            Posted February 26, 2013 at 5:16 pm | Permalink

            Thank you. That is very helpful.

  11. Gordon Munro
    Posted February 26, 2013 at 12:14 pm | Permalink

    Could this sad regretful persistence in flogging a discreted thesis be a function of a Wilsonian family kin-selected trait for obtuse senior-citizen ba(i)sed self-rightousness?

    • Diane G.
      Posted February 27, 2013 at 1:19 am | Permalink

      Any-age citizen can be self-righteous.

      • Gordon Munro
        Posted March 6, 2013 at 12:00 pm | Permalink

        Self-evident. See this and take your pick of comments above.

  12. gillt
    Posted February 26, 2013 at 5:17 pm | Permalink

    JAC:

    In contrast, individual and kin selection have productively explained the evolution of “problematic” traits like altruism and cooperation. They have been tested and work.

    Another “problematic” trait is cannibalism. You mentioned male lions killing cubs but what about intraspecific predation among individuals within breeding populations? Hamilton predicted that behaviors such as cannibalism should be selected against in absence of benefit to the individual and when it does benefit should move toward a single optimum.

    Lori Stevens’ work with cannibalistic flour beetles (Tribolium) in the 1990s concluded

    Thus, theoretical studies combined with my empirical studies on the evolution of social behavior suggest that when the fitness of an individual depends on the distribution of genotypes within the population, population structure and multilevel selection must be considered in order to understand evolution.

    • BillyJoe
      Posted February 27, 2013 at 1:28 pm | Permalink

      “Hamilton predicted that behaviors such as cannibalism should be selected against in absence of benefit to the individual”

      If he said this, he was wrong. But I suspect he did not say this. Traits that are not beneficial to the individual tend to fade because they are not selected FOR, but they are not selected AGAINST. Also, traits that are not beneficial to the individual may be a necessary consequence of another trait which IS beneficial and, therefore, will tend to persist.

  13. H.H.
    Posted February 26, 2013 at 9:59 pm | Permalink

    I’m struggling to understand this, but a lot of it is going over my head. I don’t understand exactly what it means to say some species was shaped either by kin selection or group selection.

    It seems intuitive to me to imagine an ant colony as both a single entity and a collection of distinct units. Each colony has it’s own character, you could say. It’s own collective strategy. Particular defense responses, foraging techniques, what have you. Differences which could mean either survival or extinction should the evolutionary pressures.

    These differences in colony behavior may not even be reflected in the morphology of its members, though. Individual ants from one colony might look identical to ants in another. They just behave completely differently as a group.

    So what is natural selection at this ant-colony level if not group selection? Is group selection something completely different?

    • Posted February 27, 2013 at 2:19 am | Permalink

      Hofstadter in Gödel, Escher, Bach treats of this in some detail, under “Aunt Hillary” pp 314-33.

      (The aunt-ant wordplay doesn’t work in my accent, just as I hear Sinatra seeming to sing about an aunt moving a rubber tree plant.)

    • illinoisjoe
      Posted February 27, 2013 at 10:31 am | Permalink

      Your questions are excellent and I understand the confusion. I’ll try to explain the distinction succinctly, though I have no doubt that the reading suggested by Shuggy is probably better if longer: Basically you are right in your thinking that a colony is clearly a group and is composed of distinct individuals, much as a human body is composed of individual cells. Both kin selection and group selection attempt to explain how evolution shaped these groups of individuals. Kin selections takes a “gene’s eye view” which I, Dr. Coyne, Richard Dawkins, and most other scientists who study this stuff happen to believe is a much more useful way to look at things. Group selection/multi-level selection focuses on the relative magnitude of selective forces on individuals within a group versus on the group itself. They argue that sometimes selection on the group is more powerful than that on the individuals within it, and that the appropriate way to understand this is via a group selection/multilevel selection perspective.

      Kin selection explains the success of groups based on individual costs and benefits. It correctly defines “success” as copies of genes produced for the next generation. That is why relatedness is such an important consideration for kin selection: it explains altruistic behaviors like those seen in eusocial societies by pointing out that even sterile individuals benefit by helping their mother produce more (fertile) daughters. It points out that evolutionary successful strategies must at least obey this basic rule: The costs of an altruistic act to the actor (C)must be less than the benefits to the recipient (B) times the relatedness (R) of that recipient to the actor. R is between 0 and 1 such that it is 1 between genetic clones, 0.5 between parents and offspring, 0.5 among siblings, etc. Whether or not kin selection is or was important for the evolution of eusociality hinges on a more specific argument that for some kind of proto-eusocial animal, it was a better deal (in terms of genetic copies) to stay put and help mom make more children than to venture out and start your own nest. Again, the emphasis here is on individuals maximizing copies of their genes, either by direct reproduction or by helping others who have identical copies.

      Group selection describes the same phenomenon, but focuses on costs and benefits accrued within versus among groups. They would explain the same phenomenon by saying “groups with more self-sacrificing individuals will out-compete groups that have more selfish individuals”, at least in some ecological/evolutionary scenarios. The basic argument is that there exist cases where selective pressures at the group level outweigh those at the individual level. In the old incarnation of group selection, this was conclusively argued to be logically flawed: individuals within groups are produced and die much faster than the groups they compose. Thus any kind of self-sacrifice must benefit the individual’s genes in some way, or by definition the self-sacrifice will die with the individual who practices it: true self-sacrifice has no chance to evolve, so doing something at personal cost to benefit the group cannot be an evolutionary selected trait.

      I am told that in its current mathematical incarnation, group selection is no longer fundamentally flawed and that the math works out to be equivalent to kin selection. My critiques of it, nonetheless, are that 1.) Said math is pretty complicated. 2.) Pragmatically, as a scientist, I don’t know of any examples of “multilevel” (group) selection that cannot be more simply explained by kin selection. I think it is lousy at generating testable predictions and has given us zero insight into classic puzzles like eusociality. All it does is confuse me. If the proponents of group selection claimed that someday down the road it might be a better way to explain complex evolutionary phenomena, I might believe them. I only hear them claiming it explains things we already have better explanations for, like the self-sacrifice of clonal bacteria (R = 1 !) which to me is like saying your skin cells are making a sacrifice by not being your germ-line cells.

      • Diane G.
        Posted February 27, 2013 at 11:29 am | Permalink

        Bravo!

  14. Diane G.
    Posted February 27, 2013 at 1:35 am | Permalink

    He was the founder of evolutionary psychology (known as “sociobiology” back then), author of two Pulitzer-Prize-winning books, one of the world’s great experts on ants, an ardent advocate for biological conservation, and a great natural historian.

    Not to mention island biogeography. :)

    As for now…Pauling his Vitamin C, Wilson his group selection…Ultimately science triumphs, and it is to be hoped that lasting reputations of those who’ve made undisputed contributions will be based on the entire arc of one’s career, and not skewed too much by late-in-life hiccups.

    (Which is not to say that rebuttals such as this one aren’t necessary!)

    Also–the NYT Opionator section seems like an odd place to be making a case for this hypothesis…

    • Dominic
      Posted February 27, 2013 at 7:31 am | Permalink

      The sociobiology book is visible on Jerry’s shelf in the last old boot photo!

      • gbjames
        Posted February 27, 2013 at 7:41 am | Permalink

        I’ve still got my copy but the book cover is long gone. I see Jerry still has the remnants of a cover on his!

      • Diane G.
        Posted February 27, 2013 at 11:23 am | Permalink

        I vividly remember reading that around 1980. Definitely a personal paradigm changer. Have cringed through some of the hypotheses it’s led to over the years (actually, I cringed at a few of Wilson’s at the time!), but there’s certainly no way of going back to a pre-evo-psych worldview.

  15. Posted February 27, 2013 at 6:08 am | Permalink

    I might be mistaken about the history of this debate, but if I’m not mistaken, some years back Stephen Jay Gould advanced some group selectionist arguments, and if I’m also not mistaken, Wilson was very critical of it at the time. I’m not sure if I remember this accurately, but if that’s true, what made Wilson change his mind? Nowack’s mathematical model, perhaps, as there was no similar model back in the 70s and 80s?

    • Wayne Tyson
      Posted February 27, 2013 at 8:50 am | Permalink

      Is there an article or a book that covers the Gould-Wilson debate? I, too, vaguely recall this, but did not follow it at the time.

      • Posted February 27, 2013 at 7:22 pm | Permalink

        In the book “Dawkins vs. Gould” by Kim Sterelny, there is some discussion of Gould supporting a bit of group selectionism, but really at the “group” level of species, so it’s really species selection.

  16. Dominic
    Posted February 27, 2013 at 7:30 am | Permalink

    In November I went to the Linnean Society to hear Elliott Sober talk about Darwin & he made a comment at some point about feeling a chill in the air whenever he visited England & mentioned group selection! Several people picked on that even though it was not essential to his talk.

    • BillyJoe
      Posted February 27, 2013 at 1:47 pm | Permalink

      There was always a difference in emphasis in thinking about evolution between America led by Gould and Britain led by Dawkins. Dawkins always had the edge though and always had the better of Gould in areas of disagreement. For example, Gould criticised gradualism as if its proponents meant that evolution occurred at a constant rate, which was not what they were saying or implying at all.

      • gbjames
        Posted February 27, 2013 at 1:49 pm | Permalink

        That was not a US vs. UK thing. Plenty of us here in the States agreed with Dawkins.

      • Wayne Tyson
        Posted March 6, 2013 at 7:54 pm | Permalink

        What is the consensus now?

        • gbjames
          Posted March 6, 2013 at 8:12 pm | Permalink

          The consensus is that group selection is a concept that doesn’t contribute much.

  17. Kevin L.
    Posted March 16, 2013 at 6:05 pm | Permalink

    Regarding the discussion of whether the definition of eusociality applies to humans: Wilson isn’t alone in considering that humans may be eusocial. Kin selectionists have also suggested that humans could meet the criteria, depending on which definition of eusociality you use (for example, see Foster and Ratnieks 2005 TREE). This follows from the Grandmother Hypothesis, which suggests menopausal women are a reproductively sterile “caste”. Of course, this is considering reproductive skew at a given moment in time, not lifetime reproductive success.

    On a separate note, “genetically based sterility” doesn’t play a role in human sociliaty, but it is also rare in the eusocial insects. In many species, if not most, reproductive caste determination is driven by larval environment (typically nutritional), not genetics. Genetic caste determination is definitely not a part of the definition of eusociality, reproductive division of labor is.

    Sorry to quibble about definitions, but they are important when trying to make sure everyone is talking about the same thing!

  18. Wayne Tyson
    Posted March 17, 2013 at 6:38 pm | Permalink

    Let’s DO “quibble” about definitions.

    Organisms do what they can, where they can, when they can.

    “Social” is derived from the Latin, “socius,” meaning “partner” or “cooperator.” (Let the Latin scholars descend upon and correct me.)

    Social animals are those which have evolved to practice a certain amount of cooperation because it worked well for them and they hung onto the idea, as it were. In the case of humans, some slings and arrows of outrageous mutations gave rise, in the realm of the “Great Success” of cooperation, to hierarchical culture, or control over others of their species. Culture led to a compounding of “success” in terms of a reproductive bubble and control-freaky hierarchical authoritarianism, outside the “walls” of “Eden” and into the “Hell” of dominant corporate monarchies. The merely social fraction (shall we say, “the 99%”?) of cultures is fed upon by the dominant figureheads (shall we say, “the 1%”?) who do nothing but blissfully gamble with their own futures and the welfare of other humans and the earth. Humans are not well-adapted to being solitary. Kadafi found this out as he was dragged out of a drainpipe and beaten to death by his “own” “people.” The 0.0001 (plus or minus?) percent given a tiny dose of his own medicine.

    “Culture” comes from the Latin, “cultus,” to cultivate (the land). Sodbusters. Farmers. Urbanites. Destroyers of “Eden.” The land once “cooperated” with the migratory social units which, “In the [true] Beginning” rarely took more from their support system than they needed, accumulating no wealth, only necessities.

    Some mutations lead to extinction.

    In order to understand the present and the future, is it not necessary to understand the beginning as well as we can?


5 Trackbacks/Pingbacks

  1. [...] See on whyevolutionistrue.wordpress.com [...]

  2. [...] For a counterpoint from Jerry Coyne, professor of biology at the University of Chicago, please see this post at Coyne’s Why Evolution is True website (the linked post also includes links to previous posts [...]

  3. [...] is True.  I certainly agree with his take on evolution, but I found some of the arguments in his response to Wilson’s latest [...]

  4. [...] Why Evolution is True “E. O. Wilson mistakenly touts group selection (again) as a key factor in human evolution“ [...]

  5. [...] Wilson’s is making people angry, which he has done many, many, many times before. His most recent bit of horrendous offensiveness takes the form of a post [...]

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