David Sloan Wilson tells the BBC that the evolution of altruism in humans is “solved”: it’s group selection (of course)

Reader Tony from the UK called my attention to yesterday’s “Start the Week” program on BBC Radio 4, which featured the evolutionary biologist David Sloan Wilson discussing the evolution of human altruism with host Tom Sutcliffe. (Click the screenshot below to go to the page, then press the arrow at lower left on that page. Alternatively, the mp3 is here and the podcast page is here.) Wilson was promoting his new book: Does Altruism Exist?: Culture, Genes, and the Welfare of Others, which came out in January.

If you know Wilson’s work, you’ll also know that he tends to explain nearly everything as a result of group selection: differential reproduction of groups rather than of individuals. And altruism is no exception. As he explains to Sutcliffe: “Groups of altruists do very well compared to groups without altruism.” That’s a pure group-selection explanation. He asserts that the group-selection explanation “is now becoming widely accepted.”

Have a listen (the Wilson segment begins at 1:05 and lasts until about 8:30. The other participants continue to discuss Wilson’s theory (with him chiming in from time to time), but the telling part comprises the first nine minutes.

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Sutcliffe asks Wilson a good question: how can you be so sure the problem of altruism has really been solved? Wilson simply responds that although there are a few holdouts who remain “unconverted,” like Richard Dawkins,  “it is still the case that this problem has been solved and still appears obvious in retrospect.” As Wilson affirms once again, “We now know it is so.”

That is a totally unwarranted degree of assurance about the issue, and in fact Wilson’s confidence angers me greatly.  The fact is that human “altruism” is a mixture of diverse and complex behaviors, only one of which corresponds to the real evolutionary issue of altruism: reproductive self-sacrifice by people that benefits unrelated people who give nothing back. And we simply haven’t the slightest idea whether that form of altruism evolved, or even if it has a genetic basis: i.e., that we have specific genes promoting such reproductive sacrifice. “True” biological altruism in humans appears rare, and when it does it appears to hijack behaviors that evolved, probably by individual or kin selection, for other reasons. Finally, there are formidable problems with explaining altruism and self-sacrificial cooperation by group selection compared to individual selection (see Pinker reference below)—problems that make the group selection explanation less parsimonious.

Wilson is in fact an enthusiast about group selection, and in the BBC show above his zealotry is deeply misleading.  If you were to ask me whether human altruism evolved by group selection, my answer would be “It’s theoretically possible, but we have no idea whether true biological altruism, if it indeed has a genetic and evolutionary basis, involved group selection. Given the problems with group selection, I think it unlikely.” That is what I see as a scientifically responsible answer. Wilson is scientifically irresponsible in his BBC presentation, leaving the untutored listener with the idea that science has finally answered the burning question of altruism. That’s simply wrong, and Wilson was wrong to leave that impression. It’s a lousy way to teach science to the public.

Here are a few problems with Wilson’s assured explanation. First of all, there is a panoply of behaviors we term “altruistic” that aren’t altruism in the true biological sense: an organism sacrificing all or part of its net reproductive fitness to benefit an organism to which it’s unrelated. Giving to charity or helping an old person cross the street are both considered “altruistic” acts in normal language, but don’t involve sacrificing one’s genetic output to benefit someone else. Rather, they’re nice gestures that involve financial or physical but not reproductive sacrifice. Such behavior doesn’t require a direct evolutionary explanation: that is, we needn’t posit a complex genetic scenario to explain it. There are plenty of alternative scenarios, involving a mixture of individual selection and cultural evolution, for how such behaviors came to be (see Peter Singer’s The Expanding Circle for one explanation).

Simple “helping” behaviors that likely evolved in our ancestors, in which individuals benefit those who aren’t especially closely related, could have evolved by individual selection, via a “tit-for-tat” strategy, also called “I’ll scratch your back; you scratch mine”). In these scenarios, individuals remember and recognize each other so that help given to a group-member will eventually be repaid. In other words, the “sacrifice” is only temporary and illusory since it’s repaid. If altruism like that—which isn’t true altruism in the sense that you don’t lose net reproductive fitness—evolved by individual selection, we’d expect to see it evolve in smallish groups in which individuals remember and recognize each other so that generous acts can be repaid to the right people. These are in fact precisely the conditions under which most of human evolution took place. Those kinds of groups aren’t necessary for altruism to evolve via group selection, which makes the group-selection explanation less necessary—and attractive.

Finally, do humans have genes for true biological altruism, in which we sacrifice our own lives and offspring production for others who are unrelated? Such acts do occur—among, for instance, volunteer firemen and soldiers who save their buddies by throwing themselves on grenades—but they are rare, not common. There is no indication that we have been selected to behave in such ways, and, in fact, most people don’t act like that. Further, even those kinds of behaviors can hijack evolved behaviors in a maladaptive way. There’s a reason why soldiers call each other “brother”; and “grenade-covering” behavior may involve hijacking of kin selection or small-group cooperation that evolved by individual selection in our ancestors. Adoption, which resembles true altruism since you’re rearing genetically unrelated individuals, is most likely a hijacking of parental instincts evolved by kin selection, and usually occurs in adults who can’t produce their own genetically related children. Finally, as Michael Price has pointed out, most altruistic-seeming behaviors really show the earmarks of having evolved for the actor’s benefit. As I say in Faith versus Fact, where I’m addressing Francis Collins’s claim that altruism couldn’t have evolved at all but must have been vouchsafed us by God:

In fact, many aspects of cooperation and altruism are precisely those we’d expect if their rudiments had evolved [by individual selection]. Altruism toward others is reciprocated most often when many people know about it, but often isn’t when you can get away with free riding. Humans have sensitive antennae for detecting violations of reciprocity, they choose to cooperate with more generous individuals, and they cooperate more when it enhances their reputation. These are signs not of a pure, God-given altruism, but of a form of cooperation that would evolve in small bands of human ancestors.

But for the most cogent critique of why human cooperation and altruism are unlikely to have evolved by group selection, see Steve Pinker’s Edge essay, “The false allure of group selection.” I won’t repeat his many arguments, but if you’re interested in the evolution of traits that seem bad for the individual but good for the group, it’s a must-read. One of his most telling arguments is that the traits that lead one group to dominate others are in fact not altruistic: they’re things like coercion, slavery, contempt for weakness, and so on. Groups that we see as really altruistic, like the Amish and San, don’t seem to have done well in inter-group competition.

In the end, Wilson is simply wrong in asserting that the evolutionary problem of altruism has been solved—and here I mean the existence of true biological altruism in humans. We don’t have any idea if such altruism is even based on “altruism” genes. (And if we all have such genes, why do so few of us display true biological altruism?) Wilson distorts the situation when claiming, erroneously, that nearly all evolutionists subscribe to Wilson’s own theory based on group selection. That is scientific self-promotion at its most self-serving.

101 Comments

  1. dorsaamir
    Posted April 15, 2015 at 10:04 am | Permalink

    This seems like something Gould would do: confidently declare an answer most scientists don’t agree with to the media.

    • Wayne Tyson
      Posted April 21, 2015 at 4:28 pm | Permalink

      Would you kindly cite a few cases in which Gould did that?

      WT

  2. kieran
    Posted April 15, 2015 at 10:13 am | Permalink

    Even when you’re really sure of something, you can still be wrong. Always use language to reflect that. There is nothing worse than a scientist speaking with absolutisms rather couched language. While it is always an argument for creationists that science changes, it is a great strength and should be encouraged when in public discourse.

  3. Paul S.
    Posted April 15, 2015 at 10:25 am | Permalink

    Not sure I understand what biological altruism is. In the example of a soldier jumping on a grenade, that seems like learned behavior, not an evolved trait. If it was altruism based in our genes, wouldn’t that prevent a soldier from killing a combatant? After all, aren’t most wars are based on ideological differences, not biological ones. Seems to me that Wilson is using evolution to explain something that may not be based on evolution, or more likely, I don’t understand biological altruism.
    As they say, when you only have a hammer, everything looks like a nail.

    • quiscalus
      Posted April 15, 2015 at 12:22 pm | Permalink

      I was thinking the same thing, that the soldier’s behavior is learned, or maybe more to the point, it’s a bit of brainwashing, as some in the military are fond of saying that they break you down in order to build you up. this then is enculturation, not genetics, and probably one more reason why they prefer such young recruits, beyond the purely physical aspects.

      as for what wars are based on, well that’ll open a whole new can of worms. In undergrad anthropology courses Marvin Harris’ book “Cows, Pigs, Wars, and Witches” is popular fodder for fighting over the causes of war. I believe it was Napoleon Chagnon that he would fight with, whether it was women, food, land or just “natural bellicosity” that cause the Yanomamo people to fight so frequently. I know we have at least one anthropologist reader in residence, perhaps they will chime in as well.

      • darrelle
        Posted April 15, 2015 at 1:44 pm | Permalink

        It is typical for militaries to foster team work and camaraderie to a very high degree. Soldiers are trained that they must be reliable, that their fellow soldiers must be able to rely on them. And that the one thing they can rely on in return is their fellow soldiers.

        This is further reinforced when soldiers actually experience combat. It has been described by soldiers since ancient times. You can find the same descriptions from ancient Greece (earlier even) to right now. When the shit has well and truly hit the fan soldiers don’t continue on because of dedication to king or country, or ideological ideals. They continue on because of their personal relationships with each other. They are as close, or closer, as true siblings.

        It is a reasonable hypothesis that this emotional bond results in altruistic behaviors that kin selection has been used to explain pretty well being co-opted.

        • Wayne Tyson
          Posted April 21, 2015 at 5:05 pm | Permalink

          In the context of battle, the mind is highly concentrated–it has to be, because the (artificially constructed) context is so narrow and the mind is in a state of confusion. Ironic, isn’t it, that concentration and confusion are, in this case, coexisting?

          Evolution is a much sloppier process, and we may have to explore beyond our imaginations and the “evidence” to come closer to whatever the truth is. It seems evident that we are not “there” yet, and that we have a long way to go . . .

          WT

    • Posted April 15, 2015 at 1:09 pm | Permalink

      Just because it “seems like” learned behavior doesn’t mean it is. That’s Prof CC’s main point. We don’t know. You’re probably right that it’s learned. But we don’t know. To claim, not only that we know it’s genetic, but that it arose by group selection, AND that this is commonly accepted, is preposterous.

    • Sastra
      Posted April 15, 2015 at 5:41 pm | Permalink

      It seems to me that the regular insistence that soldiers need to believe in God or they will be weak comrades in battle militates against the idea that altruism is innate.

    • Posted April 16, 2015 at 12:41 pm | Permalink

      I am not qualified to argue one way or the other. But it may be of value to say that from my experience in training soldiers and fighting in battles, it’s surprisingly (at least to me) easy to make them act against the seemingly basic personal survival instinct.
      At least in the IDF, the idea of “breaking the recruits and building a soldier from the ruins” has long been abandoned, and there is no convincing evidence that new soldiers are less “courageous”.

  4. John Matthews
    Posted April 15, 2015 at 10:28 am | Permalink

    I’m really glad.you covered this. I listened the other day and thought he was going way over the top with how ”solved’ all this was.

  5. bonetired
    Posted April 15, 2015 at 10:30 am | Permalink

    Apart from that, I gather you liked it! ( 🙂 )

  6. Dominic
    Posted April 15, 2015 at 10:33 am | Permalink

    There is a nice article that may be of interest that says-
    “Males with low socio-economic status were more likely to rescue in all the contexts (fire, drowning, violence and traffic accidents).”
    Lyons, Minna, Who are the heroes? Characteristics of people who rescue others. Journal of Cultural and Evolutionary Psychology, 3(2005)3– 4, 239–248

    • Kevin
      Posted April 15, 2015 at 10:43 am | Permalink

      I saw that article. I thought it was an interesting read. I have no solutions that I would put forward as science, but am certain of this: my altruism is driven by selfishness. If it makes me happier to save a person’s life and endanger my own then that’s what I do…because it makes me happier…a circle of motivation, bootstrapped from itself.

      • Dominic
        Posted April 16, 2015 at 10:04 am | Permalink

        I came across it from noting a number of deaf people who saved people from drowning…

  7. Jonathan Wallace
    Posted April 15, 2015 at 10:34 am | Permalink

    Wilson’s slogan: “Selfishness beats altruism within groups. Altruistic groups beat selfish groups. Everything else is commentary” neatly sums up the problem with group selection as an explanation for altruism (assuming, per Prof Ceiling Cat’s comments, above,that it is even an evolved trait). If selfishness trumps altruism within the group then it will keep on relentlessly undermining the between-group advantage conferred by altruism.

    It’s analogous to the ‘tragedy of the commons’ in a way, where a population would benefit from its members showing individual restraint in the exploitation of a common resource but each individual’s short term interest leads him to use it without restraint until the resource is depleted and everyone suffers the consequences.

    • Delphin
      Posted April 15, 2015 at 11:01 am | Permalink

      Group selection is faddish in some circles because people imagine it produces niceness. All the strong counters PCC or you or Linker cite will prove less convincing than pointing out it leads to the evolution of ingrained xenophobia and racism.

      • Posted April 15, 2015 at 1:18 pm | Permalink

        Do you really think group selection is real and produces xenophobia and racism?

        • rickflick
          Posted April 15, 2015 at 1:36 pm | Permalink

          Of course not. Xenophobia and racism don’t require group selection. Individuals in groups do it just fine.

        • Sastra
          Posted April 15, 2015 at 5:53 pm | Permalink

          I think Delphin’s point (please correct me if not) was that the proponents of group selection are secretly more swayed by what they think are the moral aspects of the belief than they are by the scientific case. Arguing that group selection leads to racism is therefore a bit like arguing that “Ancient Astronaut” theory is racist. Erik Von Daniken thought indigenous people in the past were too stupid to construct impressive monuments without help from extraterrestrials.

          No, that’s not really the main problem, is it? But if you want the general fans of the ‘theory’ to reject it (or maybe just be prepared to listen more favorably to the other side) it might be a tempting strategy.

          • Posted April 15, 2015 at 7:43 pm | Permalink

            Got it. The wording threw me. I should have read it as, “If group selection were real, it would lead to racism, etc.” Thanks.

            • Delphin
              Posted April 21, 2015 at 5:39 pm | Permalink

              Yes, exactly.

        • Delphin
          Posted April 21, 2015 at 5:38 pm | Permalink

          Group selection should produce a strong desire to draw group boundaries. Name two stronger ones. And the boundaries should be around the group’s that were selected. They should be hard to eradicate.

    • Tony Hoffman
      Posted April 15, 2015 at 11:55 am | Permalink

      I think that defense of evolutionary selection for altruism relies on group selection and Simpson’s paradox. I’m surprised nobdy’s mentioned that yet.

      http://www.zoology.ubc.ca/~doebeli/reprints/simon_etal2012.pdf

  8. Daniel McCoy
    Posted April 15, 2015 at 10:36 am | Permalink

    We are a social species, so it does seem likely that our groupishness would be important to our evolution in some way. But natural selection involves both a unit of selection and an environment of selection. Saying that our human groups have been a unit of selection seems really nonsensical to me. Our groups are just too fuzzy and fluid. It makes much more sense for human groups to be involved as the environment of selection. True genocide is rare; more commonly when one human group dominates another, the result is a blending, with cultural and genetic results more akin to mating than selection. Examples abound (the English language). One human group dominating another is not anything like natural selection; there are just no concrete examples of selection of whole groups, while the area of human adaptation to human social groups is a rich one.

    • Mark Sturtevant
      Posted April 15, 2015 at 12:59 pm | Permalink

      I think this captures a real problem with group selection — that groups are too fluid to stay as one ‘group’ from the beginning –>middle–> end of a process where a group winds up taking over.
      One can try to bring it down to a smaller unit than the individual, and that is the gene. It is the gene, or rather, an allele of a gene that has a singular origin in a single individual. It is not just the descendants of that individual that take over a population, it is the descendants of the gene carried in later generations of individuals, some of whom are not necessarily closely related to the individual that 1st had the allele.

    • reasonshark
      Posted April 19, 2015 at 5:41 pm | Permalink

      I think one helpful way of seeing the problem is applying the thought experiment to individual selection. Really apply it, I mean, because the gene-centric view, strictly speaking, doesn’t vindicate that either.

      Suppose you wanted a second level of selection “above” the gene, and you picked individuals. That makes sense because that’s how evolution by natural selection is often framed: kangaroos evolved longer legs and a hop because that enabled individual kangaroos to outpace their slower kin, say, from equally quick predators.

      Here’s the problem: Joey the kangaroo does not replicate himself when he reproduces with Sheila the kangaroo. Their offspring are not copies, but products of a copying process at the genetic level, so it just collapses to the genetic level. Even if you remove sexual reproduction and just have hopping aphid clones, it’s obvious Aphid generation one “copies” Aphid generation two because they both come from genetic copying.

      Well OK, but what about long legs? They get passed on like genes, so trait-level selection could be another hit, couldn’t it? Unfortunately, traits suffer the same problem: they’re end products of genetic level evolution. If it looks like they’re copying, it’s because they also emerge from copies. If an aphid in the line of generations suddenly lost a back leg, its trait isn’t passed on to its offspring. Its offspring emerge with the full set, showing that the clones are products of a copying process rather than a replication system in and of themselves.

      My point isn’t that there can never be a natural-selection-worthy replicator other than a gene. My point is that it’s devilish enough to try and apply that to a superficially replicator-like process like individual reproduction. Applying it to an even more nebulous concept like a bunch of organisms is only going to magnify the corresponding problems.

      • Daniel McCoy
        Posted April 19, 2015 at 6:56 pm | Permalink

        There is a difference between the unit of inheritance and the unit of selection. All of the genes in an individual have their survival fates locked together; they all live or die together. No human gene can survive unless the individual human carrying that gene survives. No human gene can make it to another generation unless the individual who carries it lives and reproduces (ignoring for the moment modern reproductive intervention).

        The unit of selection has switched levels in the past, for single-celled organisms the unit of selection is the cell. Then a transition began where some kinds of cells adapted to staying close to one another and the environment of their selection became dominated by other cells of the same kind. Over time this led to an adaptive state where survival of individual cells was nearly irrelevant to the survival of multi-celled organism which carried the genes. Eventually the adaption reached a point were reproduction is delegated to just a few of the cells, with the others merely providing an environment for the cells which actually pass on the shared genes. Human cells are adapted to live in a human organism where their fates have been tied together, the genes live, die, or reproduce as one organism, so we have group selection of human cells, where the group is the human individual. Your cells die all the time and you hardly notice. The replication and inheritance happens on the genes, but the selection happens on the organism.

        This transition to group selection has arguably already happened at the next level up with the social insects, with survival of individual worker ants or bees being only moderately related to the survival of the hive and all of the reproduction being delegated to a few individuals. Natural selection is operating on whole nests and hives rather than individual insects. The transition here from multi-celled organism to multi-organism hive went through a similar stage of individual insects staying near each other until the group became a dominate force in the environment experienced by the individual insects.

        One problem is that this transition from separate organism to multi-individual organism is a gradual one, as is usual in evolution. There is a lot of grey area in between, and social animals like humans are into the grey area where we are definitely adapted to living in human societies, not as solitary individuals, even if our social groups do not live and die as a whole.

        It’s a problem similar to speciation. The definition of “species” is a convenience for us to talk about nature, but the reality is that it is all grey area. There is no one generation you can point to in human ancestry where the parent is not human but the offspring is human; it’s gradual and we define where the line is. Similarly it would be hard to point to a generation where the dominate selection unit switched from the cell to the multi-celled organism.

        I think saying that humans have crossed through the transition to the point where traits are being selected at the group level is hard to line up with the way human groups work. There are analogies between human groups and hives but human groups are just not tight enough, they don’t live or die as a whole. Individuals (and their genes) from defeated or less successful groups just get absorbed into dominate groups.

        I don’t think the term “group selection” is useful until whole groups live or die together. That switch may happen before reproduction has been delegated to a few individuals, as it has with mammalian cells or hive bees, I’m not sure exactly where to draw the line, but humans just do not seem to be past it.

        We humans are definitely in the transition area though, where the environment of our fellow humans has as much or more to do with our survival than the rest of wild nature does. But it really seems that our groups are just too fluid and mutually interpenetrating to be subject to selection pressure above the level of the individual.

        It would take something more compelling than mathematical models based on sweeping assumptions (“spherical cows”) to convince me otherwise.

        • Gregory Kusnick
          Posted April 19, 2015 at 7:27 pm | Permalink

          No human gene can make it to another generation unless the individual who carries it lives and reproduces (ignoring for the moment modern reproductive intervention).

          You also seem to be ignoring kin selection. It is not the case that all my genes perish with me when I die, since copies of many of those genes exist in my nieces and nephews. Helping them survive and reproduce enhances my own fitness, even if I have no kids of my own.

          Kin selection isn’t the only counterexample. If a left-handed billionaire sets up a trust fund for left-handed orphans, then genes for left-handedness will benefit, even if the billionaire dies childless.

          So to me it makes sense to talk about genes as the unit of selection, since genes are both the causes and the ultimate beneficiaries of selectable phenotypic variation.

          • Daniel McCoy
            Posted April 19, 2015 at 7:44 pm | Permalink

            The comment was long enough without trying to describe how kin selection fits into the grey area between individual selection and group selection, but kin selection is clearly part of that along with more general niche construction. Kin selection is not group selection. Your left-handed philanthropist is a pretty contrived modern example, but actual examples of social behavior along those lines might fit into the niche construction realm.

            In either case it is still the individual who lives or dies. Kin and philanthropist are both part of the selection environment which decides whether the individual lives or dies.

            • Gregory Kusnick
              Posted April 19, 2015 at 8:30 pm | Permalink

              The point is that natural selection isn’t measured in individual deaths. Cases where we can blame a single death on specific genes are rare. Natural selection is by definition a statistical process, measured in changing gene frequencies. If a gene’s effect tends, on average, to increase the frequency of that gene in the next generation, then we say that natural selection has favored that gene. Conversely, selection against a gene doesn’t require that it kill anybody; all that’s needed is that it produce fewer copies than competing alleles.

              • Daniel McCoy
                Posted April 20, 2015 at 12:05 am | Permalink

                “The point is that natural selection isn’t measured in individual deaths.”

                What do you think the statistical process you mention is counting?
                The statistical process operates on the differential between the survival or death of large numbers of individuals,
                many coin flips deciding death or survival. The difference is in the granularity of the coin flips. For single cell organisms, the coin is flipped per individual cell, for multi-celled organisms, the coin is flipped per multi-celled organism. For a bee hive, the coin just might be flipped per hive.
                There is no evidence that the coin is flipped per group of humans.

              • Gregory Kusnick
                Posted April 20, 2015 at 12:50 am | Permalink

                As I already said, I think selection counts changes in gene frequencies, not deaths per se. Some deaths (e.g. death from old age) don’t have any material effect on gene frequencies. Nor is death required to alter gene frequencies; small changes in fertility, nutrition, sexual attractiveness, or any number of other factors can do it by modulating the birth rate rather than the death rate.

                So the idea that selection counts individual deaths, and nothing but individual deaths, is just too simplistic, and misses a big part of what natural selection is about.

                That does not, however, mean that I’m arguing for group selection. I said a few posts back that I think gene-level selection makes the most sense.

              • Daniel McCoy
                Posted April 20, 2015 at 9:46 am | Permalink

                The difference between the unit of inheritance and the unit of selection is the difference between the genotype and the phenotype. This concept is all straight out of The Selfish Gene. Read the wikipedia link I sent. The phenotype is the vehicle genes use to get from one generation to the next.
                Of course it’s not deaths, it’s survival to reproduce, death is just the inverse of survival.

        • reasonshark
          Posted April 20, 2015 at 4:40 am | Permalink

          There is a difference between the unit of inheritance and the unit of selection.

          This seems a needless way of bisecting what is really a single issue. If you want to find a unit of selection, then you want a replicator because natural selection “selects” among variants of this replicator, conditional on how its techniques to keep itself alive fare in the population. Its descendants – copies – then inherit that technique, precisely because they are copies. The replicator is where the issue of inheritance and selection are unified into a single explanatory framework.

          More to the point, it should be apparent that “units” of selection must either be replicators in their own right or simply vehicles for pre-existing replicators. You could make a case for cellular structures replicating, though as I understand it, that’s simply by dint of genetic manipulation (give or take whether you class RNA as a replicator in its own right or as a vehicle of DNA’s replication), but not individuals. Sexually reproducing organisms are mishmashes of genes who inherit features from two parents: they certainly aren’t selected individually. Even cloning individuals aren’t really replicators, because any traits that persist and are evolutionarily linked are byproducts of the genes.

          The unit of selection has switched levels in the past, for single-celled organisms the unit of selection is the cell. Then a transition began where some kinds of cells adapted to staying close to one another and the environment of their selection became dominated by other cells of the same kind.

          Here, I think you’re needlessly blurring two issues. That vehicles have increased in scale from cells to multicellular organisms to whole social collections is one thing. But the unit of selection hasn’t actually changed – the transition from RNA to DNA notwithstanding – because no genuine replicators have arisen in that time-frame. The fact that genes now live and die via multicellular organisms (you might think of them as highly coherent bee-like colonies with somatic and gametic members) is simply a change in vehicular complexity. It doesn’t entail “individual” selection merely because individual advantages and genetic propagation are so often one and the same. If anything, genetic selection predicts more phenomena, such as fatal semelparity (or big bang reproduction that kills the host) and inclusive fitness.

          So calling an individual a unit of selection is simply a happy convenience, made intuitively appealing because they’re the most obvious things we notice. It’s not as wrong-headed as group selection, but it’s still technically wrong.

          • Daniel McCoy
            Posted April 20, 2015 at 10:21 am | Permalink

            It’s the phenotype that carries the genotype from one generation to the next. The phenotype of a bacteria is a single cell. The phenotype of a human is an individual. One could describe the phenotype (or extended phenotype) of bees as the hive.
            There was a point in our evolutionary history where our phenotype jumped levels from cell to multi-celled organism.
            Sure, there is a whole process of natural selection happening to the genes, but the environment genes are interacting with is within the phenotype itself, all while the phenotype is being subject to selection pressures from the environment outside of the phenotype. Both levels of selection pressure matter.
            Saying that the the only useful way to talk about evolution is at the gene level is a bit like saying chemists should only ever speak in the language of physics. Sure all chemistry should be reducible to physics, but there are insights to be gained by speaking of chemical reactions in molecules rather than electrons. The action of the genes is a chemical one, should biologists always have to break it down to the electrons and protons when they talk about biochemical processes?
            (“Consilience” was one of E.O.Wilson’s better books.)
            Selection pressure at the level of the phenotype (or even extended phenotype) is a useful concept which can result in insights.
            I find definitions of “altruism” to be tricky business, but that’s the word in use.
            So-called “altruistic” traits seem to me most likely to be caused by selection pressure on the human phenotype (the individual) from the environment presented by human societies. Of course that pressure travels down to the genes the same way a chemical reaction is about electrons and protons. If the selection process is coherent enough then it is useful to talk about it without reducing it down to electrons every time. I’m just not convinced that human societies are coherent enough to be directly experiencing natural selection the way that beehives do.

            • Gregory Kusnick
              Posted April 20, 2015 at 12:47 pm | Permalink

              Selection pressure at the level of the phenotype (or even extended phenotype) is a useful concept which can result in insights.

              That I can agree with. But saying that phenotypes feel selection pressure is not to say that phenotypes are the units being selected, since phenotypes are ephemeral. Only replicators (i.e. genes) pass through the sieve of selection into the next generation.

    • Delphin
      Posted April 21, 2015 at 5:40 pm | Permalink

      Perfect.

  9. Delphin
    Posted April 15, 2015 at 10:55 am | Permalink

    First, thanks again.
    Ironic that he should be peddling this just after the evisceration of the other Wilson’s Nature article.

  10. Peter B
    Posted April 15, 2015 at 10:56 am | Permalink

    Here’s another review of Sloan Wilson:

    H. Allen Orr: The Biology of Being Good to Others. New York Review of Books, March 19 2015
    Review of Does Altruism Exist?: Culture, Genes, and the Welfare of Others by David Sloan Wilson
    Yale University Press/Templeton Press, 180 pp.
    http://www.nybooks.com/articles/archives/2015/mar/19/biology-being-good-others/

  11. Posted April 15, 2015 at 11:08 am | Permalink

    Yes, a very unattractive display by DSL. Disappointing.

    • Posted April 15, 2015 at 3:03 pm | Permalink

      DSW not DSL. (I need to get a cable modem!)

  12. MAUCH
    Posted April 15, 2015 at 11:16 am | Permalink

    Unless my reading of popular science articles suffers from confirmation bias I do not get the impression that there is resurgence among scientists accepting group selection as a major agent driving evolution. I had the impression that since the seminal work by W. D. Hamilton and over the years the accumulated work of scores of other scientists the scientific majority accept that group selection can not be considered the major agent driving evolution. If I misunderstand what the scientific consensuses is please let my know.

    • reasonshark
      Posted April 19, 2015 at 5:28 pm | Permalink

      I don’t think you do. An article appears every now and again from a few researchers on the subject, but quite frankly, the numbers involved in this resurgence strike me as deeply unimpressive. If I went back and trawled through the papers I’ve been sent to by such people, collecting names as I go, I’d still end up with a list of group selectionists shorter than the one for climate-change denying climatologists.

  13. Randy Schenck
    Posted April 15, 2015 at 11:30 am | Permalink

    The other question some of us might have here is a why. Why would an otherwise apparent educated person in this field go out in a public way like this? Is it for money and his 15 minutes of fame or is there something else that causes this kind of going off the reservation. Maybe just proves there can be sad cases at every level.

    • quiscalus
      Posted April 15, 2015 at 12:35 pm | Permalink

      I imagine that part of it is to be known as the one (or one of the few) who overturned a seemingly strong scientific theory and lead us to the correct one. Fame, pride, legacy, maybe, or just the strong desire to feel like you are “correcting” science for the ages. There certainly are cases in which scientific “certainty” was challenged and shown to be in error, so perhaps this is just one of those attempts, misguided as it appears to be, along with Gould’s punctuated equilibrium, and one I’ve recently heard attempting to return sans religion, that actually geological uniformitarianism is false and catastrophism is how the geological world really works, mostly by people not understanding biblical basis of catastrophism and conflating it with the idea that well, clearly volcanoes and earthquakes are, well, catastrophic.

      At least that’s what I assume is behind it, fame, of a sort.

    • reasonshark
      Posted April 19, 2015 at 5:25 pm | Permalink

      The other question some of us might have here is a why. Why would an otherwise apparent educated person in this field go out in a public way like this?

      I’ve been reading Bad Science by Ben Goldacre recently, and one of the things he points out is that a lot of “scientists” making interesting (read: unscientific) claims and research tend to go directly to the public with their views (and their credentials, however ropey) rather than to academic journals for peer review and criticism. That’s one of the more conspicuous things about how both Wilsons have gone about their group selection idea.

      I’m not sure if it’s because they are, subconsciously or not, aware of their need to be evasiveness, or if they’re convinced something’s biased about the “establishment” (or they just don’t get out much), but it’s notable that the only group selection pieces I know of that get put in genuinely peer-reviewed literature are mathematical models. As Pinker metaphorically puts it in the Edge article: “Imagine a perfectly spherical cow…”

      • Peter Nonacs
        Posted April 19, 2015 at 6:43 pm | Permalink

        I wasn’t going to weigh in on this anymore. But, oh really? Just models???

        Ecological causes of multilevel covariance between size and first-year survival in a wild bird population
        By: Bouwhuis, Sandra; Vedder, Oscar; Garroway, Colin J.; et al.
        JOURNAL OF ANIMAL ECOLOGY Volume: 84 Issue: 1 Pages: 208-218 Published: JAN 2015

        Site-specific group selection drives locally adapted group compositions
        By: Pruitt, Jonathan N.; Goodnight, Charles J.
        NATURE Volume: 514 Issue: 7522 Pages: 359-+ Published: OCT 16 2014

        Species-specific influence of group composition on collective behaviors in ants
        By: Modlmeier, Andreas P.; Keiser, Carl N.; Shearer, Taylor A.; et al.
        BEHAVIORAL ECOLOGY AND SOCIOBIOLOGY Volume: 68 Issue: 12 Pages: 1929-1937 Published: DEC 2014

        Multilevel selection and effects of keystone hyperaggressive males on mating success and behavior in stream water striders
        By: Chang, Ann T.; Sih, Andrew
        BEHAVIORAL ECOLOGY Volume: 24 Issue: 5 Pages: 1166-1176 Published: SEP-OCT 2013

        Cultural evolution of cooperation: The interplay between forms of social learning and group selection
        By: Molleman, Lucas; Quinones, Andres E.; Weissing, Franz J.
        EVOLUTION AND HUMAN BEHAVIOR Volume: 34 Issue: 5 Pages: 342-349 Published: SEP 2013

        Multilevel selection analysis of a microbial social trait
        By: Roditi, Laura de Vargas; Boyle, Kerry E.; Xavier, Joao B.
        MOLECULAR SYSTEMS BIOLOGY Volume: 9 Article Number: 684 Published: AUG 2013

        MULTILEVEL SELECTION WITH KIN AND NON-KIN GROUPS, EXPERIMENTAL RESULTS WITH JAPANESE QUAIL (COTURNIX JAPONICA)
        By: Muir, William. M.; Bijma, P.; Schinckel, A.
        EVOLUTION Volume: 67 Issue: 6 Pages: 1598-1606 Published: JUN 2013

        Multilevel Selection and Neighbourhood Effects from Individual to Metapopulation in a Wild Passerine
        By: Laiolo, Paola; Ramon Obeso, Jose
        PLOS ONE Volume: 7 Issue: 6 Article Number: e38526 Published: JUN 20 2012

        PHENOTYPIC ASSORTMENT MEDIATES THE EFFECT OF SOCIAL SELECTION IN A WILD BEETLE POPULATION
        By: Formica, Vincent A.; McGlothlin, Joel W.; Wood, Corlett W.; et al.
        EVOLUTION Volume: 65 Issue: 10 Pages: 2771-2781 Published: OCT 2011

        NATURAL GENETIC VARIATION IN SOCIAL ENVIRONMENT CHOICE: CONTEXT-DEPENDENT GENE-ENVIRONMENT CORRELATION IN DROSOPHILA MELANOGASTER
        By: Saltz, Julia B.
        EVOLUTION Volume: 65 Issue: 8 Pages: 2325-2334 Published: AUG 2011

        Evolution in structured populations: beyond the kin versus group debate
        By: Lion, Sebastien; Jansen, Vincent A. A.; Day, Troy
        TRENDS IN ECOLOGY & EVOLUTION Volume: 26 Issue: 4 Pages: 193-201 Published: APR 2011

        Transmission coupling mechanisms: cultural group selection
        By: Boyd, Robert; Richerson, Peter J.
        PHILOSOPHICAL TRANSACTIONS OF THE ROYAL SOCIETY B-BIOLOGICAL SCIENCES Volume: 365 Issue:1559 Pages: 3787-3795 Published: DEC 12 2010

        The Group-Level Consequences of Sexual Conflict in Multigroup Populations
        By: Eldakar, Omar Tonsi; Gallup, Andrew C.
        PLOS ONE Volume: 6 Issue: 10 Article Number: e26451 Published: OCT 24 2011

        Population structure influences sexual conflict in wild populations of water striders
        By: Eldakar, Omar Tonsi; Dlugos, Michael J.; Holt, Galen P.; et al.
        BEHAVIOUR Volume: 147 Issue: 12 Special Issue: SI Pages: 1615-1631 Published: NOV 2010

        THE ROLE OF MULTILEVEL SELECTION IN THE EVOLUTION OF SEXUAL CONFLICT IN THE WATER STRIDER AQUARIUS REMIGIS
        By: Eldakar, Omar Tonsi; Wilson, David Sloan; Dlugos, Michael J.; et al.
        EVOLUTION Volume: 64 Issue: 11 Pages: 3183-3189 Published: NOV 2010

        Group selection and social evolution in domesticated animals
        By: Wade, Michael J.; Bijma, Piter; Ellen, Esther D.; et al.
        EVOLUTIONARY APPLICATIONS Volume: 3 Issue: 5-6 Pages: 453-465 Published: SEP 2010

        All peer-reviewed mind you. Just a quick search over last 5 years… Let’s see what the next 20 bring!

        • reasonshark
          Posted April 20, 2015 at 4:19 am | Permalink

          I wasn’t going to weigh in on this anymore. But, oh really? Just models???

          Ah, good point. Correction: “mathematical models and needless misapplications crowbarred onto observational research.”

          For instance, your first paper shows how environmental factors and even the experience of individual birds feed back onto patterns of behaviour across populations and subpopulations (groups). Heck, one of the examples of the advantage of “multilevel analysis of selection” is of water striders Aquarius remigis with aggressive individuals gaining the advantage in a “group”, but females moving to less aggressive “groups” to turn it into a disadvantage.

          Really? This is the vaunted application of multilevel selection that will usher in 20 years of exciting new work? There’s no conceivable way “individual” selection doesn’t already account for this, such as ESS, extended phenotypes in the context of group living, frequency-dependent selection, and so on? If a gene “for” aggression drives itself extinct because it drives away potential mates from its catchment area, how is that any different from a cheater strategy driving its own population extinct even as it is prospering better than other strategies in an ESS model (see page 186 of The Selfish Gene)? Never mind all the issues not touched upon – like whether a “group” contains relatives, is actually a propagule or vehicle or replicator, whether what can be described is actually a copying process required for the application of natural selection to “higher” “levels” to work, etc.

          If the rest of the papers are like that, then I’m surprised the movement’s lasted five years.

  14. davidintoronto
    Posted April 15, 2015 at 11:39 am | Permalink

    …see Steve Pinker’s Edge essay, “The false allure of group selection.”

    **

    Not that it needed to be said but… hokey smoke, Pinker writes well.

  15. Geoff Benson
    Posted April 15, 2015 at 11:43 am | Permalink

    I listened to this broadcast the other night. As far a evolution is concerned, although I’m still reading WEIT, I’m little more than ordinary Joe Public on the matter, and I assumed Wilson was taking the scientific party line. It would seem not. I’m a bit disappointed with the BBC.

  16. rickflick
    Posted April 15, 2015 at 12:26 pm | Permalink

    I was struck by this: “Francis Collins’s claim that altruism couldn’t have evolved at all but must have been vouchsafed us by God:”

    This makes me ask why does Collins believe that? What occurs to me is he might be a misanthrope at heart and is desperately looking for a way to feel comfortable with himself. If he has no real empathy he may feel a need to find a justification for a sense of belonging and for behaviors deemed appropriate, like altruism. Fables about deities and what they expect of him would provide the rational.

  17. Peter Nonacs
    Posted April 15, 2015 at 12:29 pm | Permalink

    I agree that neither the “problem” of altruism is solved, nor that humans are proof positive for the dominance of group selection. However, I almost always see this debate framed in terms of supposed traits that are favored within groups, but disfavored when groups compete. Instead, the more common situation may be that a particular trait or behavior is favored within the group, and also makes groups, as a whole, better competitors. So yes, being nice to others can be an individually selected trait because the others are kin that have your selfsame genes, or that there is direct payback or positive benefits associated with having a good reputation. But Wilson’s point that all-nice groups are simply likely to function better and be more productive than not-nice groups seems rather obvious, too. What I think is overlooked is the selection strength at each level. It is certainly possible that being nice within a group results in a very small relative fitness gain versus being not nice, while at the same time nice groups may do very much better than not nice ones. Thus, the spread of “niceness” in a population is more actuated by across-group selection than by within-group (although both processes favor the same end result). Or the vice versa situation (which I’ll agree has more evidence for it). I find that too often in this debate, the evolutionary potential of synergisms across levels of selection is ignored. If a trait can be shown to have any individual or kin level benefit, then group level processes are conveniently dismissed. Or alternatively, if there is a demonstrable group-level benefit then group selection must have overwhelmed the individual level. Both viewpoints, if d*gmatically applied, are too narrow and may miss a significant part of the evolutionary story!

    • Gregory Kusnick
      Posted April 15, 2015 at 1:42 pm | Permalink

      Maybe I’m missing something, but it seems to me that synergistic group selection of the sort you’re describing isn’t much different from niche construction or Dawkins’ notion of the extended phenotype. The cooperative social environment of the group is the niche constructed by individual cooperators acting together, and the benefits of living in such a group favor the genes that drove the individual behavior. I don’t see what “group selection” adds to this picture.

      • Peter Nonacs
        Posted April 16, 2015 at 4:00 pm | Permalink

        The importance is that population structure and distribution of individuals matters in affecting how rapidly or whether a trait spreads at all. The DS Wilson structured deme models of group selection are theoretically robust and potentially widely applicable. Groups don’t have to be isolated entities. They can be local neighborhoods within a continuous population. If, however, you measure selection only in a single group/neighborhood or even just in the ‘average’ group, you may not see what is actually happening in the population. A good example is Jonathan Pruitt’s work with social spiders and the distribution of docility/aggression personalities within groups. The best predictor of fitness is whether or not the frequencies of the types within the group match the local conditions. And there is not much evidence that within-group frequencies will evolve to match those conditions. http://www.wired.com/2014/10/social-spider-group-evolution/

    • reasonshark
      Posted April 17, 2015 at 5:28 am | Permalink

      Instead, the more common situation may be that a particular trait or behavior is favored within the group, and also makes groups, as a whole, better competitors.

      In which case, any kind of multilevel selection idea, not just group selection, becomes a pointless idea because then we’d have the already-successful selfish gene theory doing the explanatory legwork to begin with. The thing that makes group selection stand out is its claim that it can explain traits that are disfavoured within groups but favoured when groups compete, and hence propagate in spite of the benefits of, say, individual selfishness. And it’s obvious why: as an explanation, group selection would be practically useless in any other scenario, because if you can’t tease out the signal from the noise, there’s no justification for claiming there’s a signal.

      If a trait can be shown to have any individual or kin level benefit

      Actually, there is no kin-level benefit (I assume you meant kin selection, but despite the use of the word “selection” in there, it’s not actually the kin-equivalent of group selection). There isn’t even, strictly speaking, an individual-level benefit, because there are adaptations (such as fatal semelparity) that sacrifice individuals when they’ve served their use. The actual units of selection are replicators, usually the genes in DNA (though other contenders, such as RNA, are plausible). This is most obvious in viruses and unicellular organisms such as bacteria, where horizontal gene transfer is possible.

      The closest individual selection could get would be perfect asexual clones, and even then they wouldn’t be genuine copies because if an aphid lost a leg before budding, its descendants would not inherit the lost leg but would be grown from scratch through genetically directed embryological processes. And if individual selection is so badly limited, then group of organisms standing in as a unit of selection? Fat chance.

      I’ve seen the evolution of social traits in animals, a herd of fast deer and a fast herd of deer, studies on how animals behave in the presence of other animals, animal social dynamics, biological altruism and parasitism, cultural inheritance, human social and moral psychology, and even memetic theory. In my experience, invoking group selection or other multilevel selection theories tends to do nothing positive for any of these fields, and if anything makes it worse, as it is almost always an invitation for confusion. I’m certainly not convinced by a Wired piece that grossly misunderstands evolutionary biology in the third paragraph, much less that assumes behavioural patterns in subpopulations of egg-laying chickens automatically demonstrates some new evolutionary principle.

      • Peter Nonacs
        Posted April 17, 2015 at 2:49 pm | Permalink

        I don’t know why advocates (and critics) of multilevel selection seem so fixated on whether or not it is some new evolutionary principle that overthrows a gene-centric view of adaptation. It simply is not, but that doesn’t mean it is valueless. Evolution by natural selection on genetic traits ALWAYS boils down to changes in allele frequencies, summing the processes that occur within the genome, at the level of the genome, and across a spatially distributed population. If, however, we only consider the level of the individual, we won’t recognize processes like meiotic drive that occur within individuals to bias representation in gametes. Group-level selection incorporates the importance of the spatial distribution of traits, and the variance in that distribution. Again, consider a trait for being nice that imposes no cost on the individual doing it, and the benefits are equally shared by all (nice and not so nice alike). If niceness were uniformly distributed in the population, it would not spread (except by random drift). If one compared the fitness of nice and not-nice individuals within any single “group” in the population, it would be equal, and you would conclude there is no selective advantage to being nice. Toss some variance into the population, however, such that some neighborhoods are nicer than others, and niceness can rapidly spread through the population as long as there is a positive correlation between neighborhood niceness and neighborhood productivity. You can dismiss this example as merely recapitulating an obvious truism – that nice spreads because its global relative fitness is higher. But it is also true how often people forget that spatial distribution (groupings!) can critically determine global fitness. Thus, if all that multilevel selection models do is remind us that spatial distribution matters, that’s still quite valuable.

        • reasonshark
          Posted April 19, 2015 at 3:11 am | Permalink

          I don’t know why advocates (and critics) of multilevel selection seem so fixated on whether or not it is some new evolutionary principle that overthrows a gene-centric view of adaptation. It simply is not, but that doesn’t mean it is valueless.

          I don’t think you grasp what the issue actually is. Group selection is not about behaviour pattern distributions. It’s not even some innocuous add-on to gene-centric evolution. It was an explicit way of answering the question of whose benefit was all this evolving and all these adaptations, especially when it came to answering questions about adaptations that don’t obviously seem to benefit their individual possessors. And on that score, the history of biology in the 20th and 21st century has simply not been kind to group selectionism.

          Your example with the spatial distribution of traits and fitness doesn’t even hold water. There are dozens of ways of measuring behavioural strategies compared with reproductive and survival advantages, one of the better ones being the ESS and Arms Race concepts, and none of them involve reviving a dead horse idea that has a long history of confusion and misdirection. Dawkins goes into depth about these sorts of issues in The Extended Phenotype, particularly in relation to his studies of digger wasp populations.

          Don’t take my word for it: the last few decades of biology come out against group selection as an explanatory mechanism, especially in contrast with kin selection theory:

          http://onlinelibrary.wiley.com/doi/10.1111/j.1420-9101.2007.01458.x/full

          “At one level, kin selection and group selection are just different ways of doing the maths or conceptualizing the evolutionary process. However, from a practical point of view, it could not be clearer that the kin selection approach is the more broadly applicable tool that we can use to understand the natural world. This is because kin selection methodologies are usually easier to use, allow the construction of models that can be better linked to specific biological examples, lend themselves to empirical testing and allow the construction of a general conceptual overview. In addition, the group selection approach is not only less useful, but also appears to frequently have negative consequences by fostering confusion that leads to wasted effort (see Errors 2 and 3 and West et al., 2007b; pp. 420–421 and 424–425). It is for these reasons that: (a) the arguments about group selection are only continued by a limited number of theoreticians, on the basis of simplified models that can be difficult to apply to real organisms (see Error 3); (b) theoretical models which make testable predictions tend to be made with kin selection theory (Tables 1 and 2); (c) empirical biologists interested in social evolution measure the kin selection coefficient of relatedness rather than the corresponding group selection parameters (Queller & Goodnight, 1989). It is best to think of group selection as a potentially useful, albeit informal, way of conceptualizing some issues, rather than a general evolutionary approach in its own right.”

          Note that this isn’t a popular magazine misreading of non-peer-reviewed work. This is the conclusions of a systematic review of the work done by the experts themselves.

          In conclusion, the experts (not including the tiny handful of partisans such as E.O. Wilson and D.S. Wilson) agree – and have the history to suggest that – you’re incorrect on the value of group and multilevel selection theories. They are valueless, both because we have much better models with which both to conceptualize and to study living things, and because there literally is nothing in favour of the concepts.

          • Peter Nonacs
            Posted April 19, 2015 at 12:46 pm | Permalink

            Well said, and I would have agreed with you 100%. About 20 years ago. But ideas also evolve and where multilevel selection is now, and where it is going differs from yesterday. The paper you cite is from the West group, who are the far end of the “It’s always just kin selection” spectrum. I could respond by pulling quotes from a number of eminent mathematical biologists who are at the opposite, “Forget about kin selection” end of the spectrum. What is really cool, however, is that more and more people are rejecting this either/or dichotomy and are simply asking what model(s) best explains their data. As an example, I cited the Wired article, but feel free to read the original Pruitt paper (if you can get past the paywall at Nature): http://www.nature.com/nature/journal/v514/n7522/full/nature13811.html
            Anyway, time will tell. Come back to me in 20 years and tell me again that multilevel selection ideas have been valueless. In the meantime, some of us are finding them quite useful and enlightening indeed.

            • reasonshark
              Posted April 19, 2015 at 4:50 pm | Permalink

              Well said, and I would have agreed with you 100%. About 20 years ago. But ideas also evolve

              That’s a pretty limp way of dressing up the fact that group selection has been redefined a bunch of times since its original inception. It’s a magnet for confusion. Just see Pinker’s Edge article Jerry linked to in the OP for examples of the muddying that ensues when the term is bandied around. It’s precisely how he describes it: an intellectual dust bunny.

              I could respond by pulling quotes from a number of eminent mathematical biologists

              You could indeed. And 9/11-truthers could point out their mass of “evidence”, for the sake of balance. That doesn’t mean what really happened on that date is in a “really cool” place somewhere in-between because “more and more people are rejecting this either/or dichotomy”. Unfortunately, the proportion of biologists who subscribe to group selection is as impressive as the proportion of climatologists who don’t think climate change is happening. The fact that a good chunk of group selection supporters are mathematical biologists, if anything, is what you’d expect when the empirical evidence is thin on the ground. There’s even a whole section on it in the OP’s linked Edge article.

              As an example, I cited the Wired article, but feel free to read the original Pruitt paper (if you can get past the paywall at Nature): http://www.nature.com/nature/journal/v514/n7522/full/nature13811.html

              Unfortunately, I can’t get past the paywall, but the abstract itself doesn’t look that promising. The definition alone doesn’t assure me the authors know what would be required to apply natural selection to “groups” (whatever a group is). And then it’s applied to what anyone else would recognize as an evolutionarily stable state of behavioural strategies in a polymorphic population.

              More specifically, the paper you cite has to answer a lot of questions, but I’ll just focus on the obvious issue: confusing the observation with the hypothesis. The survivorship or extinction of a ratio of polymorphic behavioural strategies, and the ratio adjusting to work better in certain backdrops, but not when pushed far enough out, are one basket of findings. Taking this as a vindication of natural selection applied to “groups” (whatever a group is) is a hypothesis, and not really a good one to explain the data.

              How do the “regulatory mechanisms” they cite work? They glibly talk about colonies as if they were agents, but there’s no suggestion that the adjustment or responses occur – for instance – because of individual decisions. How are the ratios copied (if they are) from group to group (spiders of this species form colonies of related individuals)? If you cut a chunk out of a spider population – akin to pulling a cloning aphid’s leg off – do the budding or fissioning descendants inherit the leftover structure (a Lamarkian evolutionary response) or build from scratch (which would be more likely to suggest genes are involved)? If ratios change over more than one generation, is this due to a developmentally stable strategy? What of the ethology of spiders: how do they pay attention to and respond to their environments? My point is not that you are burdened with answering these questions. My point is that nothing in the abstract suggests to me they were ever asked.

              Include the fact that one of the authors (Goodnight) appeared on here a way back and pretty much let rip in a couple of paragraphs with a revised history of group selection containing a high number of howlers and bluster, and I can’t say I see where your confidence in the next 20 years is coming from, other than the Dunning-Kruger effect.

  18. peepuk
    Posted April 15, 2015 at 12:34 pm | Permalink

    When I hear self-sacrifice, the first thing that comes to my mind is suicide attack.

    People are suckers for reputation, that’s a major factor for top quality reproductive success. And it makes it a lot easier to manipulate and being manipulated. So it has a very important function in our society.

    The good part is that it protects us somewhat from self-serving behavior,it makes us behave rather nicely. We look indeed good when compared to our closest relative the chimpanzee.

    Especial the BBC seems lately a sucker for reputation, after the altruistic activities of Robby Savage in children’s Hospitals.

  19. Christopher
    Posted April 15, 2015 at 1:08 pm | Permalink

    I listened to that radio show, and at the time my response was “eh?, that’s news to me!”

  20. NewEnglandBob
    Posted April 15, 2015 at 1:42 pm | Permalink

    DSW actually said “genetic evolution can happen quickly but cultural evolution happens slowly”. Is he out of his mind? Oh, yeah, he is DSW.

  21. Posted April 15, 2015 at 1:47 pm | Permalink

    Seems obvious to me that human culture evolves in ways that superficially resemble Darwinian evolution, and that human behavior is hugely influenced by culture.

    I’d suggest that “group selection” likely is a good way of understanding the evolution of culture…but to extend that to the biological level without showing how there’s a close tie between cultural and biological units…?

    If these “group selection” theories were true, then you’d expect an adopted infant to grow up socially more like its biological origins than its cultural upbringing, but that’s a laughable suggestion.

    Unless group selection proponents have a way to clear even such a low hurdle, I’m astonished that such otherwise-intelligent people could think it has any merit.

    b&

    • Mark Sturtevant
      Posted April 15, 2015 at 2:46 pm | Permalink

      Group selection of cultures is likely meant to mean selection of one culture, in toto over another culture. But like genomes, when cultures meet and compete one generally winds up assimilating parts of the other even as a losing culture fades away. So the ‘winning’ culture is no longer the same culture.

      • Posted April 15, 2015 at 2:51 pm | Permalink

        …and that’s even assuming you can draw clear boundaries between cultures. Save for populations that have been isolated fo many generations, I’ve no clue how that’s supposed to work. Is curry part of Indian culture, or Pakistani, or Korean, or Japanese, or British…?

        b&

        • Posted April 15, 2015 at 3:11 pm | Permalink

          [Curry] … or Thai, Malay, or Indonesian or Dutch or Jamaican …

          • Posted April 15, 2015 at 3:15 pm | Permalink

            Or, on further thought, the sandwich.

            Every culture has some variation on the theme of bread stuffed with something savory. Pirogi, baos, tacos, pasties, pitas…

            …hmmm…come to think of it, it may well be lunchtime….

            b&

            • Posted April 15, 2015 at 5:36 pm | Permalink

              Everyone knows the sandwich comes from the Sandwich Islands. Where native Hawaiians invented it by placing one egg and a slice of Spam between two halves of Portugese Sausage.

              • Posted April 15, 2015 at 9:13 pm | Permalink

                Wasn’t Earl who invented it? And who credited the inspiration to a Wicca priestess he met on the beach…?

                b&

              • darrelle
                Posted April 16, 2015 at 7:25 am | Permalink

                An Earl who wanted to be able to eat while playing cards, and without having to put down his hand.

                The Earl’s line continues to this day. And his currently extant descendant owns a chain of, wait for it . . . . sandwich shops!

                I ate at one 2 or 3 years ago. Pretty good. I wouldn’t rave about it, but not bad.

              • Posted April 16, 2015 at 11:19 am | Permalink

                Strangely enough…it turns out that one of their franchises is just up the road from me, at the corner of Fifth and Mill in downtown Tempe. I’ll have to keep that in mind.

                b&

              • darrelle
                Posted April 16, 2015 at 11:41 am | Permalink

                Ben,

                Have you ever eaten at the Wild Thaiger? I recently saw this restaurant on a foodie show on which Beau MacMillan, executive chef for Sanctuary on Camelback Mountain, and Guy Fieri were saying very nice things about it.

                I’ve got to say, there sure seems to be a lot of good food in your area. I’d have to spend a week or two there just to try all the food I’ve taken note of ’round your stomping grounds.

              • Posted April 16, 2015 at 4:59 pm | Permalink

                I actually have eaten there, some years ago. Good food, as I recall. I was doing graphic design production in the marketing department of The Arizona Republic at the time, and we would sometimes go to interesting places for lunch.

                I almost never eat out any more…takes a lot of time and money — and I just don’t want to eat like that all the time. It’s wonderful for special occasions, but not for an everyday sort of thing.

                Might also help to understand that the city is quite spread out…from my place to uptown, where Wild Thaiger is, would be almost half an hour drive…and could easily be double that when traffic gets bad.

                b&

  22. Steve Pollard
    Posted April 15, 2015 at 3:06 pm | Permalink

    Once again, many thanks to Jerry for his elegant and incisive analysis of this particular issue.

    But I am slightly amazed that Wilson thinks he can lead with the chin like this and get away with it. He may think he has the answer, but does he really believe that the question is settled and only a few standouts like RD dare to challenge it? Such hubris!

  23. Posted April 15, 2015 at 4:45 pm | Permalink

    Isn’t it fascinating how nearly every scientist who holds a minority position is convinced that it is “now becoming widely accepted”?

    In my area, proponents of paraphyletic taxa and panbiogeographers claim the same.

    • John Scanlon, FCD
      Posted April 21, 2015 at 12:07 pm | Permalink

      …where “now becoming widely accepted” is correctly understood to mean “still occasionally twitching” or “not yet fully extinct”.

    • Wayne Tyson
      Posted April 21, 2015 at 4:34 pm | Permalink

      A lot of “minority positions” in history are now “widely accepted.”

      WT

  24. Marella
    Posted April 15, 2015 at 5:39 pm | Permalink

    “Groups of altruists do very well compared to groups without altruism.”

    Where are these groups without altruism he talks about? I’ve never heard of a group of humans living entirely without altruism and I don’t believe it exists. He is pretending that the experiment has been done and it hasn’t. I am sure a group of humans without altruism would do badly but that isn’t science, it’s just an assertion.

    • Posted April 15, 2015 at 7:48 pm | Permalink

      I think he’s talking about models run on computers.

  25. Posted April 15, 2015 at 5:57 pm | Permalink

    The subject has a fascinating history. Robert Ardrey was the single most effective and influential individual in smashing the Blank Slate orthodoxy. The professionals and academics could never forgive him for being right when most of them were wrong, so they’ve made him an unperson. Their excuse? He wrote nice things about group selection. Instead, E. O. Wilson, a member of the tribe, became the knight in shining armor who “really” slew the Blank Slate dragon. The only problem – he just came out of the closet as a full fledged supporter of group selection. No doubt the “history” of the affair will have to be revised again.

  26. Sastra
    Posted April 15, 2015 at 6:13 pm | Permalink

    Here’s another potential problem with Wilson’s explanation: it seems to clash with his views on the evolution of religion.

    If altruism means “self-sacrifice with no expectation of future benefit” then wouldn’t a belief in God or Karma throw that off? “What goes around comes around.” If the Cosmos notices and cares about the difference between Good and Evil — and rewards the former and punishes the latter — then all bets are off when it comes to whether or not we’re dealing with genuine altruism. A future heaven or better reincarnation doesn’t have to be real to be part of a psychological motivation of tit for tat.

    Wilson believes that religion evolved “because it helped make groups of humans comparatively more cohesive, more cooperative and more fraternal, and thus able to present a formidable front against bands of less organized or unified adversaries.” But even if this were true how could it make any difference genetically? Instead, it seems that being altruistic is dependent on the cultural details of spirituality — and that can and does go anywhere.

  27. Posted April 15, 2015 at 9:41 pm | Permalink

    The group selectionists seem to base much of their hubris on the mathematical models of Martin Nowak. However, compared to the 3-D hydrodynamics codes used to model fusion implosions at the National Ignition Facility, they are kindergarten stuff. Those 3-D models inspired a similar confidence in the physicists at Livermore that they would achieve fusion ignition. In the so-called National Ignition Program, which lasted two years, they failed. Not only did they fail, but they didn’t even come close (which, of course, doesn’t mean they never will, using some other approach.) Nowak’s models can’t even be fact checked like fusion targets. They are a very frail basis for anything as extravagant as Wilson’s faith in group selection.

  28. Polymeron
    Posted April 15, 2015 at 10:20 pm | Permalink

    I’m surprised at no mention of the handicap principle – another individual-selection mechanism that can be hijacked to cause this sort of behavior. That also means that by this article’s standards this isn’t “true” altruism either, since its roots are in a net benefit to the individual.

    I’m uncomfortable with the word “true” being used to describe this extremely narrow definition of altruism. If the difference between a true behavior and a false one is in ultimate causes terms “it invalidates the centrality of natural selection”, then of course you wouldn’t find it among things that science observes.

    I think it’s clear that what Wilson is referring to in “altruism” is the actual phenomenon – which is incredibly common. People around the world take risks to their lives to help unrelated individuals all the time, often with dubious or very unlikely (read: no) return.

    So if you have an explanation for the ultimate cause of the behavior, it doesn’t stop being true – a better usage would be that it’s not true altruism only if the *proximate* cause is selfish. That happens a lot too, of course, but in nearly all cases? I think definitely not.

    But that’s nitpicking. The crux of the article is correct: While we have good ideas is to what caused this behavior to exist, the question is far from solved. To say that it is, with such desiciveness is wrong, and to state that the accepted solution is one that isn’t currently supported by a plurality of the researchers in the field is *very* wrong.

    Group selection is an alluring explanation in that it looks like an obvious extension of the natural selection process; so much so, in fact, that I developed it myself in my teen years, before I had technical understanding of it. When you examine the actual numbers, group selection doesn’t work as a primary driver.

    • darrelle
      Posted April 16, 2015 at 9:20 am | Permalink

      But you are forgetting one fairly important word that Jerry used. True biological altruism, is the term Jerry used. And he clearly stated the distinction he was making.

      “The fact is that human “altruism” is a mixture of diverse and complex behaviors, only one of which corresponds to the real evolutionary issue of altruism: reproductive self-sacrifice by people that benefits unrelated people who give nothing back. And we simply haven’t the slightest idea whether that form of altruism evolved, or even if it has a genetic basis: i.e., that we have specific genes promoting such reproductive sacrifice.”[my bold]

      “First of all, there is a panoply of behaviors we term “altruistic” that aren’t altruism in the true biological sense: an organism sacrificing all or part of its net reproductive fitness to benefit an organism to which it’s unrelated.“[my bold]

      He is distinguishing between a behavior that is directly selected for, an organism sacrificing itself for the benefit of another organism that is not directly genetically related to it, and other behaviors that are often considered to be altruistic. And he gave examples of the types of altruistic behaviors he was distinguishing “true biological altruism” from.

      “Giving to charity or helping an old person cross the street are both considered “altruistic” acts in normal language, but don’t involve sacrificing one’s genetic output to benefit someone else.”

      And he went on to explain why the distinction is relevant to the issue of the evolutionary origins of altruistic behavior.

  29. Daniel Engblom
    Posted April 16, 2015 at 1:53 pm | Permalink

    I have very little to contribute, I liked the article!
    Oh, with social primates like humans, status and reputation is also a thing individuals seek for selfish reasons, but what can change your status or reputation in a group can be very “selfless”, so another thing that could explain apparent altruistic behaviour in the standard neodarwinian framework with little to no extra assumptions.

  30. Posted April 16, 2015 at 4:34 pm | Permalink

    Very interesting post. Thank you.

  31. Keith Cook or more
    Posted April 16, 2015 at 8:21 pm | Permalink

    In twin studies it has been shown that monozygotic twins (identical) are more on average, cooperative than dizygotic twins (fraternal) what if anything, does that say about a genetic bases for altruism?
    First up, I wish I knew, it points towards some gene input to cooperative behaviours but not to altruism as such.
    It would not be difficult to think it could under the right conditions, if one twin was not so risk adverse as the other and willing to take a chance on surviving some ordeal.
    But this could also apply to non related individuals who are risk takers as to more conservative natured persons.
    Another example, a quick analytical or intuitive brain may access a dangerous situation and act to take advantage, with no outcome in mind but to act quickly, to be sure these are brave individuals and it smells of altruism, but is it?
    What about a testosterone fuelled individual with no regard to self preservation? is that in part a gene based altruism dictated by an belligerent environment?
    Certainly to my mind, when an outcome is not a given as to survival, it looks like anything but group selection and more like an individual’s gene package plus environment.

  32. Danny
    Posted April 17, 2015 at 2:36 am | Permalink

    Very interesting and informative as, always.

    As a lowly computer scientist, I don’t have anything to contribute other than general appreciation and admiration for ‘real’ science 🙂

  33. Posted April 17, 2015 at 1:42 pm | Permalink

    Perhaps part of the difficulty in comprehending the origins of a characteristic such as altruism is that, as many of you above have insinuated, it comes from a combination of evolutionary, cultural, and social factors. Both individual- and group-selection may be involved (Wilson’s insistence on one or ther other is overconfident, I think) along with learned cultural and social attributes (we think of altruism is good). This is too complex a process to be whittled down into one etiology.

    • reasonshark
      Posted April 19, 2015 at 5:04 pm | Permalink

      Nobody’s suggesting it be “whittled down into one etiology”. The point is that we have little to no reason to think that group selection is one of those factors, whether involved in informal altruism or involved in the biologically precise definition of altruism.

  34. Ken Kukec
    Posted April 17, 2015 at 2:32 pm | Permalink

    I’m no expert, and I’m never comfortable with what non-experts find “obvious” in scientific matters, but (as is invariably the case when such apophatic phraseology is followed by the b-word coordinating conjunction) I’ll take the plunge here:

    If “true altruism” means “reproductive self-sacrifice by people that benefits unrelated people who give nothing back,” then by definition non-altruistic free-riders enjoy a reproductive advantage in getting their genes into the next generation. As a result, those free-rider genes would propagate throughout the group, overwhelming the genes of the truly altruistic (since some of them, at least, have sacrificed themselves), thereby negating the opportunity for altruism to develop by way of group selection. (A hard-bought humility keeps me from adding “QED” here.)

  35. Wayne Tyson
    Posted April 19, 2015 at 2:06 am | Permalink

    I join the apparent minority, expressed by the following two posts by mystery people:

    kieran
    Posted April 15, 2015 at 10:13 am | Permalink
    Even when you’re really sure of something, you can still be wrong. Always use language to reflect that. There is nothing worse than a scientist speaking with absolutisms rather [than] couched language. While it is always an argument for creationists that science changes, it is a great strength and should be encouraged when in public discourse.

    septimus
    Posted April 17, 2015 at 1:42 pm | Permalink
    Perhaps part of the difficulty in comprehending the origins of a characteristic such as altruism is that, as many of you above have insinuated, it comes from a combination of evolutionary, cultural, and social factors. Both individual- and group-selection may be involved (Wilson’s insistence on one or ther other is overconfident, I think) along with learned cultural and social attributes (we think of altruism is good). This is too complex a process to be whittled down into one etiology.

    I haven’t much to add, in fact, but another way of putting it (those quoted please correct me as necessary; I take full responsibility for my own remarks and am not attempting to embellish my remarks by association), might be to say that is that much of the commentary seems to be in a bit of an intellectual snarl, thus losing focus (at least for me), although many posts contain statements that seem both relevant and useful. I will quote one of my greatest mentors once again, as I see it as relevant to the root of the snarl, to wit, “The suspension of judgment is the highest exercise in intellectual discipline.”

    Difficult or simple though the “answer” to this “question” may turn out to be, the arguments seem to be, in the main, polarizing. One of the reasons that I keep hanging on is that I am trying to “finish” a work I have been calling “Culture Against Society.” Needless to say, such a work will never be finished, but if it is a pebble in the pond, it will sink or skip on its own merits. And you’re right—I’m not up to the job. Part of it is to say that none of us are—Nature will always know more than we. My central guess-thesis is that we most likely were, in our “primitive” state, more social that we are now. More “altruistic” if you like. Pinker won’t like this, but after all, he is “higher above me than Heaven is Hell.” He has nothing to fear. One problem lies within the persistence of Hobbes’ insistence that “primitives” were somehow lesser beings than we. The only evidence was/is his/our insistence that we were/are superior to the “uncultured savages” observable then and now. All organisms do what they can, where they can, when they can. If “altruism” (cooperation) works for the moment, do it. If “competition” works, do that.

    Yet, the farther one proceeds and the higher one piles up knowledge or wisdom (whatever that is), the farther one can see into the great endless gulf of ignorance before one. (For some reason I am reminded of the time when I saw a lone cow in a feed-lot yard, standing upon a mountain of manure.) As a boy growing up on a stock-farm, I observed what I call the “barnyard truce,” under which familiar individuals and groups rarely molested each other, and actually enjoyed each other’s company; however, strangers were chased away or killed. Territorial imperative? I don’t know.

    • Gregory Kusnick
      Posted April 29, 2015 at 1:41 pm | Permalink

      This reply seems rather short on substance. As I read it, it pretty much boils down to “I’m right and Coyne’s wrong, and the proof that I’m right is that other group-selectionists agree with me. For details, buy my book.”


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