Do we need group selection to explain human cooperation?

January 31, 2012 • 11:15 am

Readers of this site will know that I’m not a big fan of group selection—the idea that adaptations in different species often result not from selection acting among individuals with different genetic constitutions, but from selection acting among groups, with some whole groups replacing others by virtue of their average genetic difference.

One supposedly group-selected trait is altruism, or any form of cooperation in which a “donor” of help receives little in return. In evolutionary terms, a genotype that sacrifices its fitness for a non-relative is at a disadvantage, and genes promoting such behaviors will be eliminated.  But if different groups had different proportions of altruists, then perhaps the altruist-rich groups would survive at the expense of groups containing more selfish members, and the altruistic trait could spread via this differential survival of groups.

Besides the lack of empirical evidence for such selection in nature, there are two theoretical problems. First, group proliferation and extinction is much slower than the reproduction of individuals within groups, so it’s hard to see how the former could outweigh the conflicting pressures of the latter.  Second, even if altruism is established via group selection, it’s vulnerable to the invasion of mutant individuals carrying non-altruistic genes: such “free riders” would be at an advantage within-all altruist groups. This would lead to the erosion of altruism, which would thus be unstable if it evolved through group selection.

Actually, pure altruism is exceedingly rare in nature, as we might expect if it’s subject to selection against individuals behaving that way. We do see it, though, in humans. Individuals regularly sacrifice their fitness (survival and reproduction) for unrelated individuals. Policeman and firemen, for example, do this, as do soldiers who risk their lives to save their comrades.

We don’t know whether such altruism in our own species has any genetic basis, but a new book review in Evolutionary Psychology by Michael Price of Brunel University, London, suggests that even if it does, one can account for it by selection on individuals rather than groups. Brunel is reviewing a new book by Samuel Bowles and Herbert Gintis, A Cooperative Species: Human Reciprocity and its Evolution (Princeton University Press, 2011), and it’s one of those rare reviews that makes its own contribution to science, bringing together a lot of evidence—in this case against the book’s thesis that human cooperation, which includes altruistic acts, evolved by group selection. (The pdf of the review, by the way, is available free at the links above and below.)

If you’re not interested in the technicalities, you can skip this paragraph from Price’s review, but I like it because it’s a mini-review of the evidence that human cooperation shows all the evolutionary signs of having evolved by individual rather than group selection.  There are, of course, many evolutionary scenarios showing how cooperation and reciprocal altruism can evolve by individual selection alone, but Price summarizes the evidence that they did:

The view that group selection is needed to explain most human cooperation seems inconsistent with the fact that over the past several decades, most successful research on this cooperation has theorized that it is produced by individual-level adaptations (Price, 2011; Price and Johnson, 2011). The most influential and predictive theories of group cooperation have assumed that people contribute to group efforts in order to acquire resources for themselves, and that the main obstacle to successful cooperation is that members often do better individually by contributing less, or by consuming more, than would be optimal for group success (Hardin, 1968; Olson, 1965). These theories have served as the basis for an immense body of research which has demonstrated their predictive power; a quick overview of this research follows. Individual group members tend to acquire return benefits via their cooperation, by engaging in behaviors that can be regarded as n-person reciprocity or conditional cooperation (Fischbacher, Gaechter, and Fehr, 2001; Tooby, Cosmides, and Price, 2006), competitive altruism (Hardy and Van Vugt, 2006; Roberts, 1998), and status-for-altruism transactions (Price, 2003, 2006); they free ride frequently, when they can get away with it (Fehr and Gaechter, 2000); they monitor other members’ contribution levels so that they can detect and punish free riding (Ostrom, 1990; Price, 2006), and they experience more punitive sentiment towards free riders when they are more individually vulnerable to being free ridden (Price, 2005; Price, Cosmides, and Tooby, 2002); they engage in partner choice, which allows highly cooperative individuals to assort positively and thus avoid being exploited by free riders (Barclay and Willer, 2007; Page, Putterman and Unel, 2005); and they engage in more cooperation and third party punishment when they can acquire more reputational benefits from doing so, or when they detect cues that their actions are being monitored (Bateson, Nettle, and Roberts, 2006; Kurzban, DeScioli, and O’Brien, 2007; Milinski, Semmann, and Krambeck, 2002).

Price notes that the book’s main argument against individual selection is based on games conducted by experimental psychologists showing that cooperation continues even when the participants get no individual-level benefits.  His response is that such games are not realistic models about how cooperation probably evolved in our ancestors:

The main problem with this suggestion has been pointed out repeatedly (Trivers, 2004; Burnham and Johnson, 2005; Hagen and Hammerstein, 2006; Price, 2008): Experimental economic games are not ecologically valid contexts from which to draw conclusions about how humans are adapted for one-shot, “anonymous” social activity. One-shot games are easy enough to orchestrate in experimental labs, but what would the analogue be in ancestral environments? Ancestrally, no experimenter was present to enforce the one-shot nature of an interaction, so social interactions were intrinsically iterative; for instance, if you cheated somebody, he might retaliate (Trivers, 2004). There’s no real reason, therefore, to expect the human mind to be adapted to a one-shot interaction context, or to process such experimental interactions as if they were truly one-shot. Further, for a behavior to be perceived as anonymous in the ancestral past, the actor would need to feel sure that no one else could consciously observe the act (e.g., she would need to be alone in the middle of the forest). This is nothing at all like the environment of an experimental lab, where you may be surrounded by other participants, you believe you are interacting with other conscious participants, and you know that your behavior is being recorded and scrutinized by researchers. Even if a participant consciously believes that his behavior is anonymous, his semi-autonomous adaptations producing his cooperative behavior may not act as if they believe this.

In sum, results from experimental economics games can be highly illuminating and useful for many purposes, but just like any kind of behavioral data, they have limitations. It is doubtful that experimental economic results actually reveal much of anything about how people are adapted for one-shot anonymous interactions, and they should not be regarded as evidence about the relevance of the individual as a vehicle of selection in ancestral environments.

Price gives the book a mixed review, praising it for its review of the literature on cooperation and of the models designed to explain it, but faults it for the unrealism of the formal models and its neglect of empirical data inimical to the book’s thesis. I haven’t yet read the book, but I found the review illuminating. So few book reviews these days do anything more than summarize the book’s contents and tack on a superficial positive judgment like, ‘This is a book that belongs on every graduate student’s shelf.”


Price, M. 2012.  Group selection theories are more sophisticated, but are they more predictive? (A review of Samuel Bowles and Herbert Gintis, A Cooperative Species: Human Reciprocity and its Evolution. Princeton University Press: Princeton, NJ). Evolutionary Psychology 10:45-49

71 thoughts on “Do we need group selection to explain human cooperation?

    1. In the U.S., there are volunteer firefighters — far more, I believe, than professionals. There are also many easier as well as safer jobs than being a firefighter or a police officer. And it’s not like they get paid only when they perform altruistic acts.

        1. But the other 30% (career firefighters) cover the bulk of the population, which is urban. These volunteer firefighters are found in small towns, where many of the town citizens are on “standby” in case of fire. It is not their full time job. There are clear-cut advantages for citizens in small towns doing this rather than paying taxes to support career firefighters. For one thing, it raises one’s status in the community. For another, there is a strong element of reciprocity. One who shirks his duty to volunteer might find his neighbors slow to respond in the event he should have a fire. No altruism need be invoked.

  1. Surely when people argue that one form of selection cannot explain certain altruistic behaviours, this is being implicitly reductionist. Certain altruistic acts may seem like a problem for individual selection, but only if we forget all the other ingredients that go into behaviour. Perhaps I’m missing something.

  2. I’m not a fan of Group Selection. It seems like a far weaker force than selection at the level of the individual/gene

    Reciprocity, kin-selection and reputation can account for human altruism.

    Do memes play a role? Has anyone written about a meme version of the Green Beard Effect?


    1. I agree, it may be a weaker force.

      That said, if one simply defines “group selection” as the covariance term of the Price Equation (George, not Michael incidentally), then the reciprocity and so forth merely become special cases of the more general phenomenon. Where the covariance is smaller than the expected, group selection is weaker than direct selection.

      The theorem does not preclude the covariance being zero, either in particular or in general (in which case, there would be “no group selection”); however, showing that experimentally would seem to be a remarkable result.

      Of course, I’m not a biologist; I’m just an eclectically read amateur.

  3. I was just reading a paper from last year in Science, where researchers did a review of kinship in a variety of hunter-gatherer societies, and found that, by and large, their bands were not heavily interrelated. Instead, society was founded in good part by sibling relationships, and by the granting of family status to the family members of one’s spouse, and, more generally, by joining groups of people that you were friendly with. Admittedly, this is just one paper, of course, but it seems like such studies of kinship would heavily weight against notions of ‘group selection’, since the groups themselves (namely, the individual bands) are not isolated genetically, and not necessarily at constant odds with one another.

    1. Probably this paper by Kim Hill “Co-Residence Patterns in Hunter-Gatherer Societies Show Unique Human Social Structure.”

      In the same issue Bernard Chapais comments on the article. He has a book called “Primeval Kinship” that you might want to have a look at.

      1. Exactly the one, thank you. But yeah, it is interesting how people postulate group selection among humans without actually looking to see if humans even form non-interbreeding groups.

  4. I find it highly unlikely that the group environment would have little effect on individual selection, in fact it doesn’t make sense not to.
    Why would groups begin to exist if they weren’t beneficial for individuals in the first place?
    Co-operation is not anywhere near the same as altruism, either, why link the two?
    Working for a boss is co-operation, watching out for each other for predators and dangers, hunting together, task delegation for exchanging specialized skills needed for the whole groups survival, etc., etc.

    I mean, groups started out small, and free riders would quickly deplete resources and extinguish the group, so groups containing unco-operative members would die out quickly. The groups themselves that survive would be necessarily more co-operative internally. If they didn’t isolate users and embraces ‘sharers’, no one would co-operate in the first place because the whole species would die out trying to indiscriminately support freeloaders.

    I must be missing something, this isn’t even an issue, as far as I know.

    1. Empirically speaking, groups containing unco-operative members don’t die out quickly. That is one of the points of the review: group selection would predict that free riders would disappear altogether, but they don’t. Instead, many people engage in free rider behavior when they can get away with it, and societies develop many mechanisms to try and actively prevent it.

      Its sort of like pointing to a very heavily policed, low violence society, and claiming they are non violent by nature. Well, no. If that were true, they wouldn’t be heavily policed. The observation that our societies spend a great deal of effort suppressing free-ridership indicates that groups containing un-cooperative members did not die out quickly: we are those groups.

  5. While I don’t have any particular dog in this fight, I do find it a bit annoying that the reviewers refers primarily to papers by himself in order to critique the book.

    Frankly, I’d be more convinced if the countervailing evidence didn’t come from the same source — the author of the review.

    Note to reviewers of books you disagree with: Don’t do this.

    1. The basic premise of the peer review process is for experts in a given field to review and critique the works of their peers. It seems counterproductive to ask expert reviewers to refrain from citing their own published works when they are relevant. Moreover, the resulting dialogue is the fulcrum which often lifts the overall level of knowledge in a given field.

    2. I hate the phrase “dog in this fight” and its loose use (it is almost as bad as “turn a blind eye” or “turn a deaf ear”). Please, just stop it.

  6. Readers of this web site that enjoy beating up group selection theory should be aware of some recent developments. The concept of group selection that Jerry (and most other researchers, actually) endorse is based on a “well known” result that says that group selection can be explained by “inclusive fitness theory”, i.e., kin selection, or individual-level selection. The result, which is sort of a “folk theorem” (everybody knows the result, but not a proof), is responsible for most, if not all, of the nonsensical arguments people have on the topic. Well, the folk theorem was never true, and hopefully it is now officially dead: Van Veelen, et. al. (2011), “Group selection and inclusive fitness are not equivalent: the Price equation vs. models and statistics”, J. Theor. Biol. I have some work appearing soon that will hopefully put another nail in the coffin. It’s silly to beat up on a straw-man version of group selection. With that said, I’m no big fan of the modeling work of Bowles and Gintis, although I have not seen their book yet. Their mathematical models are simplistic. and don’t get group-level events right.

    My own view on this topic is that once you understand that group selection and individual-level selection are apples and oranges, you see that they can work together (possibly synergistically), and this is probably how the evolution of cooperation typically occurs in the real world.

    1. I’m sorry, but lots of scientists say you’re wrong, and that the arguments for equivalency are not “nonsensical”. Look at the huge panoply of workers in the field who criticized the Nowak et al. paper that asserted that their form of “group selection” was not the same thing as kin selection. As far as we can see, it is.

      And if you’re going to tout your own expertise here, and the fact that you have an upcoming paper, then you must give your name. One has little credibility making such assertions unless we know who you are.

      1. I am extremely aware that lots of scientists think I’m wrong! But take a look at Van Veelen et. al. It is actually a very enjoyable read. I’m newer to the field, but I’m very proud of Simon, B. (2010), A dynamical model of two-level selection, Evol. Ecol. Res. It’s clear from that work how (I think) group selection should be viewed. I will be happy to share recent work that hasn’t officially appeared yet with people that contact me directly.

        1. So WEIT?

          Why did you attack the credibility of your opponent? Much less the mere mention of his name?

          That sounds like bravado before a boxing match. And it does not read as an intellectual use of logic.

          Either your opponent has support for his position, or does not.

          Either you have support for your position or you do not.


          I think your bravado revealed an attempt to disguise your fear of being wrong.

          BTW, my name is Wayne ….

          1. Re Van Veelen’s veracity, see Gardner et al. 2011. The genetical theory of kin selection. J. Evol. Biology 24:1021-1043 On pp. 1031-1033, the authors show that Van Veelen’s attack on kin selection and Hamilton’s rule is completely wrong and that, in fact, Van Veelen appears not to understand kin selection. Here’s a snippet:

            van Veelen (2009) has argued that, although the issue of synergy can be resolved for two-player games, Hamil- ton’s rule cannot be guaranteed to give a correct prediction of the action of natural selection in games of three or more players. This is based upon the idea that Hamilton’s rule contains fewer parameters than are needed to model such complexity. van Veelen (2009) illustrated this point with an example of a three-player ‘stag hunt’ game. Here, we provide a detailed reanalysis of this game, from a Hamilton’s-rule perspective. . .

            . . . This is exactly the result derived by van Veelen (2009). However, van Veelen (2009) claimed that this result cannot be formulated in terms of Hamilton’s rule (in particular, ‘cannot be given in a formula with costs, benefits and relatedness only’). He argued that the same is true of any model with three or more social partners interacting in a synergistic way. This conclusion follows from an argument that: (i) in such models, details of population structure in addition to relatedness (genetic similarity of pairs of individuals) must be known to predict the action of natural selection; and (ii) such details do not feature as terms in Hamilton’s rule, so this cannot generally provide correct predictions. For example, two model populations can be set up such that the relatedness between pairs of social partners is the same but other moments of population structure are different, such that cooperation is selected for in one model but selected against in the other. van Veelen (2009) asserted that the corresponding Hamilton’s rules for both models will be identical, and hence unable to account for the difference in selective outcome.
            However, as we have shown, this is based upon an incorrect understanding of kin selection and Hamilton’s rule. We have derived van Veelen’s (2009) condition for cooperation to be selectively favoured, using Hamilton’s rule. Irrespective of higher moments of population structure, Hamilton’s rule correctly predicts the action of natural selection. This is because the cost and benefit terms are also mediated by population structure, so that although higher moments of genetic structure are not captured in the relatedness coefficient, they may exert their effects through the cost and benefit terms of Hamilton’s rule. van Veelen (2009) has therefore con- fused the cost and benefit terms in Hamilton’s rule with arbitrary payoff parameters in an evolutionary game (another recent example of this occurs in Nowak et al., 2010b). In reality, the cost and benefit terms in Hamil- ton’s rule are the direct and indirect average fitness effects of a genetic predisposition for a character of interest, which are prone to be mediated by the genetic and ecological milieu.

            Now granted, this is van Veelen (2009) and not van Veelen (2011), but you can see why I don’t have a lot of confidence in his work on kin/group selection.

            1. Jerry, This is a different issue. The Van Veelen (2011) paper is a proof that the “folk theorem” (my name for it), which says in effect that every mathematical model of group selection can be reformulated as a model of kin selection (inclusive fitness), is false. In the acknowledgement paragraph at the end of the paper is a long list of major players in the debates, including Andy Gardner. It’s not clear that they all endorse the results, but apparently they were in on the conversation that lead to the paper. It’s a strange situation since a lot of these people have an awful lot at stake here.

              It will be interesting to see how this plays out over the next couple of years. I don’t mean to come off as a know-it-all.

              1. Just because somebody is listed in the acknowledgements, that doesn’t mean they endorse the paper, or even agree that it should be published. In most fields, it’s common practice to list anybody you’ve had a substantial interaction with regarding the issues in the paper – and doubly so if they disagree with you, and have critiqued the work.

    2. I don’t have anything riding on this, but I had to laugh at the description of how individual-level selection and putative other selection to be demonstrated “can work together … synergistically”.

      Perhaps, but it is at the very least an oxymoron.

      1. Thanks, that probably isn’t the best way to word it! Maybe I should just say they can both operate at the same time. They are sufficiently different that there’s no logical problem with it. And by the way, this has not yet been demonstrated (to my knowledge). Just a hunch.

      2. Funny, it’s the second time in a couple of days that someone on this website uses oxymoron to mean redundant instead of contradictory.
        When did this change started happening? I’ve never noticed this usage before.

  7. I can think of at least two possible meanings of “group selection.” One meaning is plain old Darwinian selection, as applied to groups rather than individuals. The other is selection by the group itself, and is more akin to sexual selection. We do practice the second kind, selection by the group, in our penal systems, which act against those who are anti-social. In earlier eras, that selection by the group was probably a lot more intensive than it is today.

  8. MikMik says, “Co-operation is not anywhere near the same as altruism, either, why link the two?”

    The common definition of altruism is to act for the benefit of another organism with no expectation of any compensation or benefits, either direct, or indirect. [paraphrase]

    But Trivers seems to use a “tit for tat” concept, which makes altruism more like cooperation.

    MIkMik asks an interesting question, and far far from my expertise. Narrowly defined altruism is, as JAC points out, quite rare, but is that what ethologists are referring to? (I don’t know.) Is altruism genetically related to cooperation, the latter really more related to one’s more direct self-interest, or are they different as MikMik opines?

    1. My understanding of the concepts, based on readings of the literature over the past year or three.

      An example of cooperation:
      Wildebeest encircle calves in order to protect them from the lion pride.

      An example of altruism:
      A lioness intentionally sacrificing herself to hyenas to save another lioness’ cub, knowing her own cub is safe.

      Obviously, the second scenario, in addition to being rarer (having the extra burden of being made up on the spot), imposes an additional burden of knowing intent. Altruistic behavior is one where the giver knowingly sacrifices itself on behalf of other individuals or the group — and most definitions add on top of that a burden that the ‘saved’ group not include family/close kin.

      Since the concept of knowing intent is pretty much found only in big-brained mammals (primates, maybe cetaceans), it’s kinda hard to demonstrate. Sacrifice for the group is one thing — intention to offer yourself as a sacrifice on behalf of a group that doesn’t include your progeny/close kin is another thing entirely.

      Of course, there are those who say altruism doesn’t exist in humans, either…ie, war heroes who “take the grenade” do so on behalf of surrogate family. Seems to me that in some circles, it might actually be considered a psychopathy.

      1. I think if someone continues to cooperate for the collective good of an unrelated group, when no one else in the group cooperates, that would be evidence of altruism. Or stupidity.

  9. “mutant individuals carrying non-altruistic genes” I have heard this statement from a lot of biologist. But nobody told me what is the name or the sequence of that gene that will change the neurological “wiring” of an individual to behave in a selfish way? this way of describing possible phenotypes has always puzzled me. No one gene or mutation causes anything without its genomic network. Anyway, I am not taking sides on this debate, because I don’t know all of the arguments for both sides yet. One thing I will say though that natural selection is not necessarily the answer to the question of altruism – or the only answer.

  10. The observation that our societies spend a great deal of effort suppressing free-ridership indicates that groups containing un-cooperative members did not die out quickly: we are those groups.

    It may be more accurate to say that groups that fail to effectively police/contain/suppress/restrain uncooperative behavior by its members (any individual member can behave either cooperatively or non-cooperatively depending on circumstance, so it is non-sensical really to talk about “cooperative” or “non-cooperative” members) tended to die out quickly.

  11. Arr fireman and police officers the best examples for altruism?

    Well, today’s police officers are something of a still short-term novelty.

    But we do have examples in nature of cooperative societies that deal with the threat of cheaters and free riders by having a subset of individuals tasked with “policing” cooperation, and these individuals are “rewarded” for their efforts by being allowed, themselves, to cheat a little bit, within limits.

    And historically, that is exactly how human societies have “paid” their police. By allowing them the exclusive privilege of cheating, (within prescribed limits.)

    Yep. Evo-psyche has produced another just-so story to explain police corruption….

    (Hell, even the best police of modern times are allowed to do things, like carry weapons, use force against individuals on their own judgment, break the speed limit, etc, within limits, that ordinary citizens are either forbidden from doing, or severely restricted in the doing of.)

    1. It could also be argued that police also gain benefit from their actions, especially in smaller communities like many rural American towns or in historic smaller communities.

      For example, a man who acted as sheriff in a small community during the California Gold Rush might have been putting himself at greater risk, but his actions would have had a direct influence on the safety his family of his family.

  12. I’ve yet to see any behavior that’s undeniably altruistic.

    Firemen and policemen certainly don’t qualify. Putting oneself at risk to help others is not inherently altruistic. No one becomes a policeman or fireman who hates the position – such a person gets a great deal of satisfaction from doing the job, or enjoys the power and/or prestige that comes with it.

    From a genetic standpoint, said power and/or prestige can certainly help with reproductive success. Our culture is saturated with the fact that men in uniform do better with the ladies.

    As for women in uniform, I can’t really say with any confidence what the impact is. They still get the power/prestige angle, but it seems plausible that their reproductive success is actually hurt by their choice of careers.

  13. I am going to chime in here – though I should point out that I am a molecular biologist and not an evolutionary biologist, so please forgive any naivety on my part in the later field.

    Altruism is an extremely complete trait – likely driven by numerous genes which no doubt also control many other traits. Is there any reason to believe that Altruism is not the result of selection for other human tendencies? For example we have relatively helpless young which have a long term of dependency on their parent or for that matter the larger social group. Is a person that makes a more attentive, caring parent (with a larger chance for offspring survival) not also likely to be more altruistic? Other genetically associated traits may be more obscure than this.

    Also, perhaps in social groups,in which we evolved, where no doubt altruism is reciprocal, doesn’t being altruism benefit the altruist in the long term because others are more likely to be altruistic to the altruist? Is it true to say then that altruism does not benefit the altruist, if they live in a group of altruists.

    In terms of free loaders – I was reading an interesting article recently ( on psychopathy which indicated that psychopathy (superficially charming, egocentric, insincere, manipulative, grandiose, exploitative, and lacking in empathy, not necessarily a serial killer but maybe a CEO on Wall Street) is a sound survival strategy – PROVIDED there is a significantly large proportion of the population to take advantage of (i.e. altruists). Psychopathy likely has a genetic component – though again is a complex trait – presumably retained in the human population at a rate of about 1% due certain aspects of the trait being advantageous during evolution.

  14. If group selection is real, then why don’t lions cease eating antelopes in order to preserve the mammals?

    No: group selection is illusory.
    The only things that select are atomic replicating units.
    Some candidates are:
    * genes
    * ideas
    * crystals
    But NOT ‘groups’ that have been arbitrarily generalised by human definition!

    1. The first part of the comment contains a fallacy. It does not follow that since lions eat antelopes then group selection does not happen.

      The rest is a profession of faith. You’d think this alone would attract some criticism here.

      1. Mmm…

        The first part of the comment contains a fallacy. It does not follow that since lions eat antelopes then group selection does not happen.

        The first part is a necessary (and what I hoped was a ‘snappy’ summary) contraction of a potentially book-length exposition.
        I had trusted that intelligent readers might grasp that implicit fact.
        But I was incorrect.
        For your edification, this was an OBVIOUS[1] paraphrasing of Richard Dawkins, so, should you have any quibbles with it, take it up with him.

        As for the ‘The rest is a profession of faith’, I stand incredulous!
        Oh, anonymous one, how on earth is my “some candidates are” in any way a”profession of faith”??

        You’d think this alone would attract some criticism here.

        Your wish is my command, oh vHF-thingy.

        I do despair at times, honestly.
        If you advocate for ‘group selection’, then you must first define it, and then defend it against the indomitable fact of gene selection.

        Try Again.
        [1] At least to those who are even vaguely familiar with the topic

        1. Wide of the mark on both counts. Your paraphrase was fallacious and I called it out. I wonder what you would make of the fact that lions do not eat lions? Also, it is simply not true that a vague familiarity with group selection — whether I possess it or not — requires familiarity with Richard Dawkins’ writings, let alone with specific passages in these.

          As to the second issue, you chose the phrase “some candidates are” rather conveniently to challenge my “profession of faith” characterisation. How about “group selection is illusory”, or “The only things that select [are selected? vHF] are atomic replicating units”? Got any evidence that’s strong enough to support such strong assertions?

          And I’m afraid you’ll have to deal with the fact that I don’t sign my real name here. I know our host doesn’t like this, but as far as I understand I am not (yet) in breach of his rules on this issue.

          1. Male lions killing the cubs of other male lions is a well documented fact. (Yes, I know you used the word “eat” but the killing part is the significant element here.)

  15. Can anyone suggest a good introduction to the mathematics of this area for people who are mathematically competent but unfamiliar with the literature on this particular topic?

    (Ideally available free on the web of course).


  16. In a manner of speaking, unicellular organisms evolved to apply group selection when they morphed into multicellular organisms.

    Animals which live and work in packs or colonies make group choices, too.

    So, why not group selection? In fact, the only way around it that I can imagine, in the case of humans, is well grounded education, critical reasoning skills, and intelligence enough to recognize when the group has been taken over by one whose egocentric goals put the group in danger.

    As if that would ever happen… LOL!!!!

  17. You wrote:

    “Readers of this site will know that I’m not a big fan of group selection—the idea that adaptations in different species often result not from selection acting among individuals with different genetic constitutions, but from selection acting among groups, with some whole groups replacing others by virtue of their average genetic difference.”

    As far as I understand the issue, that is the late 1960s conception of group selection. The “modern” conception is multilevel selection applied to a particular tire of the natural hierarchy (that is the tier were the particles are individuals and the collectives groups). Of course, there are semantic issues and researchers like West or Gardner prefer to not use the term “group selection” anymore.

    Multilevel selection theory (MLS) comes in two flavours MLS2 is like what you described and has been rejected for the individual-group tier. In MlS1, however, group structure only structures individual selection and a dynamics of group resolution and re-establishment prevents cheats from taking over altruistic groups.

    I prefer to call it MLS1, because the term “group selection” is so confused, but what’s the useof continuing that confusion by freezing a concept to a long since reject model.?

    1. P.S.: I haven’t seen the book in question. If it wants to reinstate an MLS2 scenario within (human) species (Wynne-Edwards’s style group selection), I’d also be skeptical.

  18. There used to be conflict between group selection and individual selection in population biology circles. The intellectual issue is settled, but people still enjoy throwing around heat instead of seeking light. This blog is a case in point. Why comment on a book before you have read it? This just produces exponentially growing inaccuracies.

    Biological selection can always be analyzed on the gene level, usually on the individual level, and sometimes a higher level. These levels never contradict each other; one can always reformulate an individual selection model as a gene-selection model, and reformulate a group selection model as an individual selection model. There is no conflict.

    The point of multi-level slection models is that the social organization of a species affects the fitness coefficients that enter into the individual fitness and gene-level analysis. That is all. This is a fundamental principle of sociobiology. This is also a fundamaental principle of niche construction in all species (if you don’t know what this means, Google it), and of gene-culture coevolution in humans (ditto).

    This blog is contributing to know-nothingism by not explaining niche construction and gene-culture coevolution, while throwing around the term “group selection” as though nothing has happened in sociobiology in the past two decades.

    Michael Price, as well as his colleagues John Tooby and Leda Cosmides have been critics of our work for many years. They can’t find any real criticism, so they label us as “group selectionists,” knowing that a lot of people will reject our argument without further inspections, as though we are idiots and/or ideologues. We are not. In particular, we are aware of every study mentioned in the paragraph from Price’s paper that you quote so approvingly, and they fit just fine with the models developed in our book.

    In particular, the fact that individuals sometimes behave altruistically does not mean that they ignore self-interest, or that they are unconditional altruists. Self-interest and reciprocal altruism are very important in human behavor, and human altruism is conditional, not absolute. It is conditional on the participation of others, without which cooperation unravels in most cases.

    Perhaps at some point you will have time to read our book and comment on it intelligently.

    1. I did not comment on the book at all, or imply that I’ve read it; I highlighted a BOOK REVIEW that makes independent criticisms of group selection. See the difference? And I don’t appreciate the snark of the last line, or the implication that I don’t know what niche construction or gene-culture coevolution is (I do, and discussed the latter in my last book).

      As for contributing to know-nothing-ism, you clearly haven’t read this site. I’ve gone into detail about papers like the Nowak et al. critique of kin selection; and there are many commenters, pro and con, who have discussed group selection. So spend some time here before you throw out baseless accusations.

      And again, note that I did not say anything about your book at all, but simply highlighted a review. If you don’t like the review, take it up with the reviewer.

      1. You cited the review approvingly without having read the book. This to my mind is not okay. At least you might have cited Peter Richerson’s review in Nature as well (Nature 476, 29–30 (03 August 2011), for some balance.

        You may have talked about niche construction and gene-culture coevolution in the past, but it would have been good to bring them up in regard to the current discussion of our book. We go into these issues in some detail in our book.

        You are making a mistake in discussing our book as though the issue of group selection is important. It is not. No model in our book is incompatible with individual level selection, or could operate “against” individual selection (whatever that might mean, which is probably nothing).

        1. Frankly, I don’t care what you think is “okay” for me to cite. I was concentrating, as you’ll see if you read the post, on the points that Price makes supporting the evolution of cooperation by individual selection. Yes, I approve of those citations, and I didn’t give either approval or disapproval of your book. I lauded Price’s review because it brought together a bunch of references of which I was unaware.

          And of course the issue of group selection is important. Maybe not in your book, but it is to many people, as the recent flap about the Nowak, Wilson, and Tarnita paper shows.

  19. Jerry,

    I realize, of course, that you have a very different perspective on this subject than do. I also acknowledge that my title may be (and probably is) overstated. Still, I wanted to let you know that this post of yours inspired me to write the following…

    2012: The Triumph of Multi-Level Selection Theory and a Renewed Appreciation of the Role of Cooperation in Human Cultural Evolution

    Thanks for so consistently articulating a skeptical position on this subject with such passion and clarity. I admit that I could be wrong. Time will tell whether or not, or to what degree, this year will be a pivotal one in this field. At the very least, however, it seems to me that the publication of these three books (Haidt, E.O Wilson, and Boehm) should generate lots of good discussion.

    I fully expect yours to one of the clearest and most thoughtful voices on the subject, even if I may (at least in part) disagree with you.


    ~ Michael

  20. I”m just a country doctor, but I think that kin selection might be a moving target, How do individuals in a community determine their kin? Does adoption of individuals or friends contribute to altruism? How does a cat adopting kittens from another litter enhance her survival? Maybe you could help explain this to me.

    1. Individuals don’t necessarily have to “determine” their kin. They only need to live in situations where kinship proximity correlates with the chances or interacting with one another. It is all a matter of statistics and heritability.

  21. If it is so hard to explain altruism as an adaptation, then maybe it isn’t an adaptation. There are two parties to any altruistic transaction, the donor and the donee. Perhaps what is evolving is the ability to manipulate people around you to be altruistic.

  22. I find it difficult to see altruism as an evolutionary force.

    If a man (or woman) moves into a new neighbor hood, and his child runs into the house and yells, “Across the street, the house is on fire.”

    The man yells to his wife, “Call the fire department.”

    And he runs into the burning home to see if he can help the people.

    This is perfectly acceptable for SOME. MOST would not risk their lives for someone they do not know.

    But, having responded to emergencies, I can tell you that not everyone responds.

    People can be trained to be more altruistic. And I believe family ties can make them stronger.

    Yet, I find it difficult to believe it would be purely evolutionary forces which grow altruism. Although, some altruism exists for animals across the animal kingdom.

    Birds sometimes raise other birds in their nests. Dolphins have been known to keep people alive who have fallen overboard. Yet, I have heard they have been known to kill people after a few days, as if they get tired after a few days.


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