I review E. O. Wilson’s new book in the TLS

January 31, 2013 • 11:41 am

Most of the contents of the Times Literary Supplement are behind a paywall (a few pieces are free in each issue), but you can at least see the latest Table of Contents here. In that issue I’ve reviewed E. O. Wilson’s new book The Social Conquest of Earth, in which he claims that major features of human behavior evolved by group (rather than individual) selection.  Sadly, my own piece, “Genes first,” is also behind a paywall, but judicious inquiry might yield you a pdf file.

Here are two excerpts:

Although Wilson pushes this view [group selection] hard –it’s the book’s centrepiece – he is probably wrong. Most biologists have rejected group selection for two reasons: it doesn’t work well in principle, and, more important, there’s no evidence that it has been of any significance in evolution. For an obvious reason, selection among groups is far less efficient than selection among genes: genes replicate and replace other genes much faster than groups of individuals divide and replace other groups. Evolving all the social traits on Wilson’s list via group selection requires a slow and unrealistic sequence of episodes in which human populations replaced each other, each replacement based on one or a few behaviours. Further, once group selection fixes a disadvantageous trait like altruism within our species, individual selection proceeds to undo it within populations (“selfish individuals beat altruistic individuals”). In other words, altruism evolved by group selection is unstable and should disappear.

If the better angels of our nature really are based on genes that evolved – rather than on non-genetic aspects of culture – then they are much more likely to have done so by individual than by group selection. This becomes even more plausible with a more detailed look at how human altruism really works. It is preferentially directed towards friends and relatives, there is much concern with reciprocity and one’s own reputation, and psychological studies show that we dislike cheaters who hurt us personally much more strongly than cheaters who hurt our group. Such behaviours are precisely what you’d expect if altruism evolved by individual rather than group selection. But Wilson ignores these problems.

It is not even clear that altruistic groups of humans would beat non-altruists. Steven Pinker has noted that success of one group over another in the real world is based not on higher frequencies of altruistic individuals, but on matters like harsher discipline, better technology, and more brutal ideology. Indeed, altruistic groups may be more easily defeated because their empathy for the weak makes them susceptible to domination. But the most important problem is this: I know of not one evolved behaviour in any species that is harmful to individuals and their genes but good for their social groups. In the end, Wilson’s invocation of group selection is superfluous.

Needless to say after that, I don’t recommend buying this misguided book, which is a sad departure from Wilson’s usual high standard of thinking and writing.

61 thoughts on “I review E. O. Wilson’s new book in the TLS

    1. Or another choice is to buy a single electronic issue of the TLS via your appropriate Amazon site. Just search for The Times Literary Supplement [Kindle Edition] at your Amazon. For me in the UK it’s only £2.49

      It can be read on any PC, Mac, iPad, Android, iPhone, iPod touch or Windows Phone 7 which has the free Kindle software installed

      [I first tried buying the epaper version from TLS directly, but gave up due to their complicated subscription system ~ like most publishers they still have a physical print mindset]

  1. Are there not many examples of individuals behaving “sub-optimally” for the greater good of the group? The individual is not “harmed” but is constrained from eating the entire carcass by the number of his/her fellow feeders and the total kilograms of the catch. After an appropriate period for digestion and napping, they can indulge their carnal instincts and pass on their alleles, n’est-ce pas?

    1. Well, if I read you correctly, then in that case the individual is not behaving sub-optimally by not being allowed to eat the entire carcass but is being externally constrained by the social behaviors of its peers.

      1. Perhaps it is not “suboptimal” to share, with the result that others share with you, and then you don’t risk starving whenever you are unlucky in the hunt.

  2. I still remember reading the full length article in NYT on Wilson’s new book published last year. Though the article was quite effusive in its praise towards Wilson but it also included a couple of lines from your side.

    “ ‘Sociobiology’ is still a very great book, and now he’s trashing it all,”

    “It’s crazy.”

    Well both of your lines do say a lot !!

    The book which i like on Human Cooperation is the one written by Bowles and Gintis. Actually, the group selectionist view point is quite a hit with normal public.”Family is the universal human grouping” as said by the famed Yale anthropologist, George Peter Murdock. For the public, in general group selection gives a scientific explanation or a sort of justification for the formation of human families, i guess.




    P.S- Why don’t you write a review on “A Cooperative Species” ? I would be happy to know your views on it.

  3. I find it hard to get my head around some of these arguments, not least because some group selection arguments just appear to be kin selection expressed in a different way.

    Also there are many levels between a single gene centric view and group selection at the level of individual phenotypes, such as say selection for groups of cooperating genes that interact to contribute to some particular phenotypic effect.

    Not sure that this is territory that non specialists can venture into easily and there seems to be some confusion among some of the professionals as to the correct interpretation.

    1. “not least because some group selection arguments just appear to be kin selection expressed in a different way”

      This point has been made a number of times in the academic literature, as well, and by both sides. AFAICT, the Nowak, Tarnita, & Wilson paper (which Jerry has criticized several times on this site) is correct on this point–kin selection is a subset of group selection. It might be that this subset makes up most, or (although I do not think this is likely) even all, of the cases of group selection in nature, however.

      Consequently, I view any claim to the effect of “It’s not group selection, it’s kin selection!” with wariness. That doesn’t not appear to be one of the options. If it’s kin selection, it’s also group selection… and vice versa (at least a lot of the time). You may as well say: “That’s not an ape, it’s a human!”

      1. “If it’s kin selection, it’s also group selection”
        Errr….no… this can’t really be the case. Kin selection is measured by inclusive fitness… a weighing of the costs and benefits by genetic relationship factors. As there is no kin relatedness within wider members of a non-kin group the inclusive element is zero: fitness goes back to an individuals own specific fitness. At this point everything is reduced to that individuals own gain or loss and the possibility of reciprocity, if that’s a factor which is in play. There are numerous examples in nature of different altruistic behaviours by kin within a larger group, compared to the non-kin within the same group.

  4. What bothers me about this topic is that it is assumed that altruism is passed down from one generation to the next like its a bad painting or old chest of drawers. We then try and explain why anyone would want to keep an old oil painting hanging around. Of course genes don’t work that way. We inherit alleles and sometime these alleles are quisecent unless we inherit two copies. The allele for altruistic tendencies might be a perfectly beneficial allele to possess if it were incompletely dominant or incompletely expressed. In this fashion I am not overtly altruistic (if I have only one copy) but perhaps some of those around me are. In this case I benefit from those around me who are altruistic but I don’t suffer by being selfless. The allele would be exploiting itself.
    One might argue that I would lessen my chance of passing on the allele if I have a bunch of truly altruistic offspring. But so long as more of my offspring survive because these selfless tendencies are present in the gene pool, than in populations without, then the allele would thrive. This arguement seems to work for sickle cell disease so why not altruism?

    1. However, in this case an individual with 2 “selfish” copies would still outcompete the heterozygote carrier (unless cheating is punishable through a mechanism such as individual recognition) because they would pay NO cost of altruistic behavior while gaining the benefits from altruistic or partially altruistic individuals.

      It does not work like the sickle cell case b/c when the risk of malaria is high those carrying a single copy of the sickle cell mutation have greater fitness than either homozygote. In your scenario, where altruism is partially dominant it would more take longer for a cheater allele to reach fixation however.

    2. In the case of sickle cell it’s advantageous for an individual to have one allele, but *not* both of them and that’s why it can be selected for. In your scenario it’s not clear that there is a benefit of *you* having one allele and unless there is one it won’t be selected for given a standard gene centric approach. It doesn’t make any difference what the frequency is in the population outside of you.

    3. I very much doubt that altruism is a discrete trait like monogenic sickle cell, where a single mutation means you have it. Plus as someone already mentioned, sickle cell is under balancing selection, as opposed to purifying selection.

  5. “Further, once group selection fixes a disadvantageous trait like altruism within our species, individual selection proceeds to undo it within populations (“selfish individuals beat altruistic individuals”)”
    Wouldn’t that depend on the specifics? One altruistic person commonly saves or benefits many more lives than a selfish one who claws his way to the top of the pile – and usually thanks to the space provided by the altruistic one.

    I’m also curious as to how altruism gets to be a “disadvantageous trait”.

    And is it supposedly disadvantageous to the individual or the group?

    And what about looking at it from the meme point of view where a meme would be an altruistic value like “unity is strength” or “Do unto others as you would have them do unto you.” I’m almost inclined to think that in some ways a meme has more power than a gene. Consider some altruistic people who were also good speakers or teachers such as Martin Luther King Jr. and Mahatma Gandhi, they might not have lived to great age (though they lived long enough to have children) but the memes they passed on gave them, in a sense, more children than any passing on of the genes ever could.

    (Sorry, lots of questions here.)

    1. The argument is that selection at the individual level will be a faster process that selection between groups, and that altruistic behavior by definition is costly to the individual. So selfish individuals would outcompete altruistic ones within a group by benefitting from others altruism while not paying the cost.

      To me, an idea like “do unto others” is closer to reciprocal altruism, implying repricocity if it is going to work. Memes will not evolve in the same way as genes and I agree that many altruistic behaviors found within societies are more likely a result of cultural evolution and are probably more often examples of reciprocal altruism than pure altruism.

      1. That raises way too many questions for me to accept it. Outcompete in which respect? Are you suggesting that selfish people tend to have more children than altruistic ones? Surely not. And wouldn’t selfish people tend to have less success with raising their children? Here, in South Afica, the children of selfish fathers tend to end up in Aids orphanages – unless they’re lucky enough to be adopted by a white Christian couple from a progressive church.

        I’m also probably going to disagree with you about what altruism is. It’s not the proverbial “doormat” attitude that comes from the common misreading of Jesus’ “turn the other cheek” teaching. Altruism is a virtue, and every virtue is moderated by the others. Eg: it’s not patience if you don’t have the discernment to know when to give up waiting. Altruism is easily undone by gullibility or the absence of foresight. There is no such thing as pure altruism. Like any virtue, it has to be a successful mix with other virtues or it’s worthless.

        1. I believe the issue here is that we are coming from different perspectives. I am not discussing altruism in the colloquial definition. In evolutionary biology/behavioral ecology, altruism is specifically defined as a behavior that benefits other organisms at a cost to the actor. Because altruistic behavior is by definition costly to the actor it should be selected against. There are several hypothese proposed to explain how seemingly altruistic behaviors would evolve and be maintained despite this individual cost:

          The most common are 1. reciprocal altruism, where altruistic behaviors are expected to be reciprocated, and cheaters punished. 2. kin selection, where altruistic behaviors are directed towards kin, and 3. the hypothesis that Dr. Coyne is arguing against here, that altruistic groups would outcompete and replace non-altruistic groups

            1. For my part I am not very up to date on the theories of how the complex altruistic-like behaviors came about and are maintained in human societies through cultural/societal forces.

          1. But then, if one unpacks the concept of behaviour, aren’t you left talking in the same terms as me? It’s all very well to talk in general and abstract terms about benefit and cost but as soon as you give an example of a behaviour, and specify a cost and a benefit you’re back with vices and virtues. Yes, a generalisation like “when food is short, actors who starve themselves so others can live are selected against,” would seem true on the surface of it but when you get down to the specifics it doesn’t follow. For example, starving so others can live wouldn’t be seen as good in young ones, ones who have not had the chance to procreate. For an old person to do it is a different matter entirely.

            I don’t understand how anyone can define altruism so vaguely, so generally as to make it indifferent to a distinction like this. By the biologist’s definition, both the old and the young would both be behaving altruistically yet the outcome is very different. Truth is the behaviour is actually different, that is why the outcome is different. The one is genuinely altruistic the other is not.

            1. Yes, you do have to very carefully identify the specific costs and benefits of the altruistic behaviors to learn how they might have evolved.

              Your specific example touches on other concepts in evolutionary biology that would suggest that altruistic behaviors expressed in young individuals will evolve differently than those expressed in post-reproductive individuals.

              First, natural selection will generally not be very effective at eliminating maladaptive traits when acting on cohorts that have already survived to reproduce and have passed their reproductive period. In your example, the cost of the altruistic behavior is very low for older individuals since they are giving up little or no chance chance to increase their direct fitness.

              However, they could be indirectly increasing their indirect fitness if they preferentially share their food with younger relatives. Indeed, if we were to study instances of individuals starving themselves when resources are scarce I would predict that such behavior would preferentially occur in older individuals, past their reproductive age, and that this behavior would be much more likely to occur with old individuals forgoing food so that their younger relatives (rather than unrelated individuals) could eat.

  6. “and psychological studies show that we dislike cheaters who hurt us personally much more strongly than cheaters who hurt our group.”

    But aren’t cheaters who hurt the group punished through the group structures? The individuals may feel less intense anger or pain because of a few degrees of removal from the cheating, but the group structures are likely to take more severe action than an emotional individual could.

    1. Business fraud is often treated more tolerantly than theft that directly affects individuals. In fact what amounts to fraud is often considered to be good business practice. Perhaps that’s something that needs addressing if we are aiming to reach a more egalitarian society.

      1. Sheesh! You’re damn right about that!
        And corrupt government officials are another example of the scum rising to the top.

        So would you say that selfish individuals who cream it off the top in a more general way are likely to be more successful than those who do their thing on a one-to-one basis?

    2. If someone is a rapist, you want him to go to jail. If he rapes your daughter, you want to kill him. I think that pretty clearly demonstrates Jerry’s point.

      Keep in mind that in the first case we are applying an abstract concept of justice, in the latter we are having a visceral emotional reaction.

      Its pretty safe to say that much of our visceral emotional reactions had evolved prior to anything resembling civilization with laws and formal group institutions.

      As our brain got larger we were able to engage in a parallel kind of evolution usually called cultural evolution. In cultural evolution we inherit by learning, not by genes.

      I only have a superficial familiarity with evolutionary psychology, so I’m guessing here and you can take what I say with a grain of salt. But we’ve probably had an enormous amount of cultural evolution and a relatively tiny amount of biological evolution in the last 50,000 years. And during that time most of the group structures you are thinking of evolved culturally, not biologically. Our culture is a sort of byproduct of having a larger brain capable of language, abstraction, metaphor, recursion, etc. This larger brain probably evolved for other reasons, but at some point some kind of intelligence threshold or tipping point was reached that enabled explosive mental and cultural evolution not seen in other species. This cultural evolution is completely uncoupled from biological evolution, though it depends on our prior biological evolution. But trying to untangle what is biologically evolved and culturally evolved in human behavior is tough, especially when it comes to morality and group dynamics. There is a lot of interplay between our emotions and our reason, and I figure our reason is heavily parameterized or influenced by learned ideas that are culturally evolved. Our emotions are often powerful enough to totally override our reason though, creating situations where we kick ourselves in the rear wondering why we did something stupid. This is I think the war between our culture and our biology that plays out in our brain.

      With altruism it seems conceivable that a biological form of reciprocal altruism formed a starting point for a more abstract cultural form of human altruistic goodness that is not genetically founded at all. It may be that certain genetically determined emotional dispositions make a person more receptive to cultural systems or social pressures in favor of altruistic decisions. Which is not at all the same thing as a genetic altruism.

      1. Well, Jeff, I’ll throw my salt over my shoulder if you take what I’m saying with a gulp of Nitrous Oxide because I know next to nothing about the biology. I’m coming with a philosophy background and trying very hard to bend my brain around the concepts.

        There are still too many confounding factors here for me. In your example of the rapist, sure, there is a difference between knowing something and feeling it, and the victim’s father would want to kill the rapist but he, being an individual with little resources at his disposal and probably being older and weaker is less likely to be able to track down and punish the rapist in ANY way. The group or the state, on the other hand, has many pairs of eyes, and whole long-term structures devoted to tracking down and punishing the rapist.

        Also, you may distinguish between applying an abstract concept of justice vs following a visceral emotion but I don’t think one can assume either that the members of the state doing their job are unfeeling or that the father would retain the same level of feeling over the whole period of pursuit. In either case, combining the feeling with the knowledge gives the feeling an extended trajectory, extended both through time and through social structure.

        I’m clueless here, but could one not rather say that individuals who are able to extend their power through creating and using group structures are evolutionarily advantaged?

        I’d also like to mention that altruism and all the other group skills sound to me like inventions or discoveries rather than advantageous accidents.

        The way we talk about the biology all sounds too bloody reductionist and category error -ridden to me. That remains my biggest problem with evolution as Prof Coyne teaches it. What you say about a decoupling of cultural evolution from biological evolution is very interesting and may be a way forward for me. I’ll be reading up about that. Thanks.

        1. You wrote, “I’m clueless here, but could one not rather say that individuals who are able to extend their power through creating and using group structures are evolutionarily advantaged?”

          I’m not 100% sure what you mean, but if an individual is “extending [his/her] power” in any way, that sounds like individual selection, not group selection. But what does “extending power” mean with regard to allele frequencies in future generations.

          I think you are getting confused by mixing scientific definitions with common usage for many different terms, including “group selection” and “altruism.” To compound the confusion, you are also mixing cultural evolution with genetic evolution.

          I’d recommend searching this website for the words “group selection” and then reading everything. Then google Steven Pinker and group selection, Richard Dawkins and group selection, etc. Pinker’s article “The False Allure of Group Selection” is especially good because he replies to his critics with a follow up article.

      2. Irrelevant but interesting: what you say about “the war between our culture and our biology” sounds an awful lot like what the apostle Paul had to say about the war between the “spirit” and the “flesh”! Bear in mind that the Spirit was not understood to be some kind of animated gas that defied the laws of nature but was most commonly identified with the concept of the Word. The “Word” can be construed as a kind of personified cultural medium in which we “live and move and have our being.” The “fruits of the Spirit” as he called them, were all virtues which are skills of relating, or group skills, altruism being one.

        I don’t understand why we talk here almost exclusively about altruism, since all the virtues are skills of relating in a group, and involve some kind of deferred gratification for the greater good.

        1. What I was talking about is really flesh against flesh, or perhaps information against information. On the one hand you have the information contained in the genes that imprints on how our brain is structured and how it learns, on the other hand you have the environmental information, much of which we have created as part of our culture, which imprints on our brain as it develops and we learn. Because of the complexity of our brain involving many parallel impulses in competition, internal struggle is inevitable.

          Paul, unfortunately for him, lived in a time of relative ignorance, so he had little knowledge of why things are as they are. Observing interior struggle is pretty intuitively obvious, even to children. Paul in his ignorance was under the influence of the idea that some invisible thing he called spirit was acting upon him.

          The way our mind understands the world involves using conceptual categories that we impose on the world. The categories don’t necessarily exist. A word like society for example, is not an object with independent existence in the world, but rather a conceptual way the human mind groups together a large number of people in order to think and talk about their collective actions and effects.

          Flesh and spirit are natural conceptual categories to invent. There are obvious differences between the limbs, torso, skin, organs, that are weak and vulnerable to disease and injury, and sensitive to pain or pleasure, while our mind seems to be invisible, insubstantial, and not subject to gross physical limitations. But our deeper scientific understanding today suggests that these differences are illusory, and that this ancient duality is really an imaginary human creation. The human is all flesh, an integrated whole composed of various organs, including the brain. The conscious mind is the effect of the brain’s actions, wholly dependent on the proper functioning of all the organs in concert.

          So the battle between flesh and spirit is merely a poetic metaphor Paul used for what later became known as “jihad” in that part of the world. Today we see it as a result of the fact that the brain has many parallel and contrary processes competing for control of our actions and decisions.

        2. I don’t understand why we talk here almost exclusively about altruism…

          We are talking about it in this way because it is a concept in evolutionary biology that is well studied. In this context it is not viewed as a virtue (and not not a virtue, either). Altruism in the world of biology is found in social species of all sorts and in this context it is only confusing to mix in talk of “spirit and flesh”, etc.

  7. I picture the situation like this:

    Let’s assume there is a parasite that attaches itself between the shoulder blades of a monkey, leading to their death after a few days. The such afflicted cannot remove the monkey all by itself. A fortunate combination of mutated genes bring into the world another monkey that somehow has the habit of picking parasites of other monkey’s backs. It has no advantage itself from this behavior, but isn’t handicapped from it either. However, it makes a lot difference for the other monkeys in the group. This “new” monkey passes down his genes, making the behavior more common in the next generation. Over time, this group outperforms other groups, but there is no “group selection” whatsoever. The genes act entirely “selfishly” . Exact copies riding within other group members create an invisible bond, where “scratching some other’s back” is, from a genetic perspective, much like “scratching one’s own back”, i.e. reciprocal altruistic behavior and selfish genes are really two sides of the same coin.

    Is this a workable (layman’s) way of putting it?

    1. Reciprocal ‘altruism’ can work as symbioses between individuals of different species that don’t share alleles, or trade relationships between nations or corporations that don’t reproduce genetically, so it doesn’t depend on there being selfish genes for the behaviour.

    2. The problem with your example is that theoretically, picking off the parasites should “cost” the altruistic monkey. He’s wasting time and energy helping others with no benefit to himself. Over time, the genes for “picking off parasites” should be selected against since he’s putting himself at a disadvantage compared to monkeys who don’t waste energy helping others in this way. If the other monkeys notice and are nicer to him for helping them, then it’s not true altruism, by definition. If his group becomes stronger and healthier than other groups and outcompetes them, and soon his group takes over the monkey world – that would be an example of group selection; but once that happens individual selection within the group would eliminate the genes/behavior for the reasons described above.

    3. You assume the parasite picking behavior requires genetic transmission. That means this behavior must improve the chances of that individual reproducing, or at least not hurt those chances, in order for the behavior to propagate.

      Depending on the behavior of other group members, this parasite picking might enhance that individual’s status and increase mating opportunities, or it could anger some more dominant member of the group, leading to violence, ostracism, and less chance of reproducing. So the fact that the group is benefiting could, but doesn’t necessarily, help that individual to mate.

      For this particular example, in primates a more likely reason that such a behavior would spread rapidly is that primate intelligence enables others in the group, and other groups, to observe and learn this behavior. So the more advanced brain and learned culture enable useful social behaviors to be adopted and spread far more rapidly than genetic mutation and natural selection.

      The various aspects of intelligence could have evolved for entirely different reasons, for example successful hunting and foraging, ability to function in more diverse environments and range over wider areas, ability to use and invent tools, ability to flexibly devise more strategies for dealing with scarcity and harsh seasons.

      This example shows how intelligence can give rise to behaviors that are not products of genetic selection, but are rather side effects of evolved abilities. Language and learned culture serve to transmit and preserve such side effects far more rapidly than biological evolution ever could. Culture seems to provide a fairly natural explanation for how the idea of pure altruism might have developed from the more biologically selfish version of reciprocal altruism.

      It’s questionable whether pure altruism really exists at all. Even the most stunning acts of self sacrifice have indirect benefits. The ideals of noble purpose, of charity, protection of life, and of honor and patriotism confer admiration and elevated social status, and hence interior feelings of satisfaction and pleasure, with possible future material benefits that accrue to heroes. Individual acts viewed in isolation of the larger social context can appear superficially to be altruistic, but future rewards are almost certain to follow, even if they are indirect and abstracted from the immediate act.

      This shows how the idea of heaven could not have evolved biologically, but how as a cultural artifact it could have been cleverly devised as the ultimate logical extension of the idea of indirect future reward for acts of goodness. And this idea is the ultimate in deceptive self-interest because the infinite postponement of reward enables people, especially leaders, the ability to escape responsibility for any rewards in reality, while the never fulfilled promise of imaginary future rewards provides cover for the deception. This makes heaven one of the cleverest and most selfish lies ever told, the Ponzi scheme of all Ponzi schemes.

      1. I doubt that “future rewards” certainly follow for a soldier who throws himself in a grenade. Of course there is pure altruism in humans.

        1. Perhaps it exists, but it seems very rare. I’m still not sure though.

          How many cases of the ultimate sacrifice were aided by the false promise of future reward offered by religion?

          So the hurling one’s self onto the grenade could have a similar level of altruism as a suicide bomber in some cases; it’s done with false expectation of great reward.

          And many other sacrifices resulting in death happen because the hero still assumes he will not die.

          Human psychology is baffling. Self-hatred, masochism, and suicide exist. Some acts of heroism could have such “abnormal” psychological causes.

          So the rational psychologically healthy atheist could still dive on the grenade. But still this person must have dedicated himself so totally to the goal of winning the battle or the war, that no other consideration matters or enters into decision making. In this case, in the heat of battle perhaps, a kind of zen-like band of brothers state of mind might take over, so that one’s own life seems no more or even less important than those you are fighting with.

          This individual sacrifice to save many is a purely selfless rational calculation, yet it is unlikely that any calculating precedes such an act. It is more likely an adrenaline fueled purely emotional and impulsive act. It shouldn’t surprise us to learn, if we had some way of knowing, that the last thought going through the mind of the guy on the grenade is “what the hell am I doing?”. I suppose another popular last thought might be “my god unto thee I commend my spirit.”

          I don’t know how common such actions are. But it does exist, and appears to be pure altruism. But this is a case involving extreme psychological stress, possible suicidal tendencies, possible religious false hopes of future reward, and other complications. These cases are not typical of human behavior. So at best it seems to be suggestive of human quirks at the margins rather than suggesting pure altruism is a general human pattern.

          1. It would be a far more clever strategy for me to convince you to jump on the grenade yourself Jeff, than to do it myself, thus enhancing my own chances of reproductive success. Unfortunately my being a cowardly defector introduces even more defector genes into the gene pool. No….there is hardly any escape from the influence of individual “selfish” genes to explain this extreme form of altruism. The only reason that ants get away with it is that inclusive fitness makes it a better reproductive strategy to save the queen than to die oneself. Termite sacrifice is a bit more problematic. Perhaps, as you say, the reason that grenade jumping altruism is so highly discussed by us humans is that it is both extremely rare and pretty damn nonsensical. In any case, the complexity of human behaviour is so great that ANY Evolutionary Psychology explanations are plausible, hence all are useless.

            1. I wouldn’t be surprised if some clever person had at least once convinced someone else to jump on the grenade. Especially if that clever fellow were a chaplain. Of course there is no time for persuasion once the grenade hits, so it would have to be accomplished in advance as an insurance policy, possibly even using veiled suggestions to plant the idea.

              I think you are going too far in saying that ALL evolutionary psychology is useless. I wouldn’t want to be in the position of debating Steven Pinker on that question if I were you. He’s written pretty persuasively that evo psych should be open to criticism, and a lot of work under the field deserves criticism, but that the field has merits and when done well can contribute to our understanding of humans. I think this is similar to the position Jerry took on the field in an old post: there is plenty to ridicule, but some of the work is good.

              I’m not quite sure exactly what point you are making relative to my discussion of altruism. I can summarize what I was trying to say, that it seems if altruism exists at all, it is an idea or ideal that is purely a product of human cultural evolution, probably based on extrapolations from observations of reciprocal altruism, and isn’t really a biologically evolved feature of human behavior. I think this is in agreement with your statement that there is no escapt from the selfish genes. I believe that reciprical altruism is biologically evolved, and I think this view is widely accepted.

              1. “I believe that reciprical altruism is biologically evolved, and I think this view is widely accepted.”
                I totally agree with you, but don’t think that falling on grenades, in particular, is reciprocal altruism.

                While we are on the general subject of falling onto grenades Jeff, I’m going to tell you a rather rambling story that may shed some light on the issue. I apologise for the rambling nature of it in advance. I once attended a lecture given by Dr. John Polkingham, an ex-Quantum Physicist turned Anglican minister. Polkingham asserted that falling on grenades was a characteristic of what he classified as “extreme altruism” for which no satisfactory biological or evolutionary explanation was possible. Therefore this phenomenon of falling on grenades was a proof of the existence of God. He went on to explain this was not just any God, but that the God in particularly had to be the Christian God as defined by the Anglican Church. When I asked at question how he made the great mental leap from noting the existence of a particular form of altruistic human behaviour to the certainty that the explanation was the byproduct of a particular desert dwelling carpenter, back several thousand years ago, and later interpreted through the agency of Henry VIII, he answered (at length) that it was self evident. The question that struck me(and still does) is how can a person with Polkinghams great intelligence, education and scientifically honed mind (substitute Francis Collins or Martin Nowak if you wish etc. etc.) possibly holds such a preposterous idea in their mind and acts on this idea, as if it were valid. If I could figure this behaviour out, maybe I could figure out why people do such a stupid thing as to fall on a grenade.
                An example of suicidal sacrifice I recall in nature is that of an ant, climbing a blade of grass so that it will increase his chance of it being consumed by grazing cattle (Dennet gives this example). The reason is not that it benefits the ant at all; it is certainly totally counterproductive to the ant’s reproductive strategy. But the ant’s brain has been inflected by a lancet fluke which “takes over” the ants brain, and redirects the ants action to aid in the lancet fluke’s life cycle strategy of being ingested into the gut of grazing animals
                Which takes me to the subject of memes, evolution and Polkingham. We are unique among animals in that we not only have innate heritable behaviours; we have culturally transmitted ones too. Our large brain (evolved toward social efficiency – e.g. language, problem solving, generation-to-generation transmission of survival strategy) is in itself an environment for reproduction – the reproduction of competing cultural entities – memes– ideas, symbols and behavioural strategies . They can become the” lancet fluke” in our behaviour system. To fall on a grenade for example, or to believe preposterous things, as in the reasoning of John Polkingham.
                It’s a lifetimes work to sceptically edit through the memes getting into our own minds. This creates another rather interesting aspect of the arguments on freewill…. but that’s for another thread.

                BTW – I don’t believe grenade falling has anything to do with group selection.

              2. @ Howard Kornstein

                Thought-provoking and intuitively convincing.

                I wanted to point out that we are not unique in having culturally transmitted behaviors, though.

                Scientists have begun to rethink their ideas on culture within monkey society in a large part because of the Japanese macaques. It has been observed that the macaques invent new behaviors and pass them on by immitation.


                (Not the only example nor the most scientific reference; just the first that sprung to mind in a quick effort to find one supporting example.)

      2. This makes heaven one of the cleverest and most selfish lies ever told, the Ponzi scheme of all Ponzi schemes.

        I love that!

        1. “(Not the only example nor the most scientific reference; just the first that sprung to mind in a quick effort to find one supporting example.)”
          Thanks Diana – fascinating. It’s impressive to see such examples of the capability of an animal group to be able to learn a new strategy in one generation and then to use it in successive generations as part of a new adapted strategy of survival. This is a sort of Lamarckian form of evolution, which we can say is happening at a “cultural” level. The new strategies that are passed on from generation to generation utilise genetically acquired cognitive/ learning CAPABILITIES but they are certainly not genetic in their own right.
          It is rather amusing to consider that if an engineer were the one designing the mechanism of Evolution he would certainly make it Lamarckian, it is so much faster and so much less clumsy and wasteful than any taking place at the genetic level. (so much for “Intelligent Design” in the way evolution normally works)
          I suppose you could consider these learned strategies as being equivalent to memes. Furthermore you could say that if one group, with one set of memes, is competing for resources with another group with a different set of memes, it is thereby competing at the level of “group selection”. But this sort of meme based group selection doesn’t negate either kin or individual levels of selection, as Wilson himself attempts to do. There again, it could never apply to ants – they are incapable of such cognitive skills. Indirect altruism does require increased cognitive skills, as the various players need to track one anothers “reputation” as well as to recognise them.
          BTW…lovely to see how fond and protective the Japanese are of these Maacque monkeys. Pity their level of their compassion and concern for beautiful sentient creatures doesn’t extend to whales.

  8. Something about this entire discussion seems to omit important aspects of behavior.
    JC uses altruism as an example of an “disadvantageous trait”. Commenter Ryan offers a definition of altruism as, “altruism is specifically defined as a behavior that benefits other organisms at a cost to the actor.”

    Evolution is not my strong suit, but I have spent far too long studying human behavior. The definition of altruism as applied, seems grossly oversimplified, and its characterization I believe erroneous, as it is applied to Homo sapiens. Behavior does not occur in a vacuum. Altruistic behavior is almost certain to garner reinforcing consequences, most likely in the form of social reinforcement, and probably reciprocal altruism which yields more fundamental reinforcers (e.g., food, shelter, protection, etc.) This is certainly beneficial to an individual behaving altruistically. In regards to altruism B.F. Skinner said, “I think you would have to show that there must be reinforcing consequences or the behavior wouldn’t occur.” I believe this to be true in most instances. Other altruistic behavior that may appear spontaneous and self-sacrificing can easily be explained by an individuals learning history or acculturation.

    There is nothing I can see that confirms altruism is disadvantageous when discussing it in the context of human behavior, nor is the definition applicable because it is so narrow. I would very much like to see a discussion informed by biologists and behaviorists.
    I guess I need to become better acquainted with the topic at hand. It seems there is much work that needs to be done.

    Lastly, I really don’t see how a trait like altruism, or any human behavior for that matter, can be discussed in terms of Mendelian genetics (comments above). It can only be clear that we are dealing with polygenetic traits, which is much more difficult to understand and explain.

    1. All that makes a lot of sense to me. Thanks.

      I’d go further and say that to explain human behaviour in terms of genetics is to commit the category error and I always wonder why no one is trying, at least some of the time, to explain the the genetics in terms of the behaviour, ie, how our gene pool provides adaptable capacities to achieve the improvements we want?

      Re what B.F. Skinner said, “I think you would have to show that there must be reinforcing consequences or the behavior wouldn’t occur.” I can’t help observing how many individuals have argued and fought and given their lives for group advantages without much hope of seeing it in their lifetime.

      1. I think I’ve had more than my fair share to say – and ask – on this one so I’ll shut my trap now.

  9. An excellent review.

    I really do wonder at E O Wilson’s plunging into such a lopsided and irrational dismissal of major concepts in theoretical Biology – including both kin selection and the effect of selection made at the level of the individual. He has done this in the face of a mass of mathematical modelling which underpins these effects, combined with a numerous studies confirming their existence within animal behaviour. He seems in the thrall of some of the recent mathematical models created by Martin Nowak (not that Wilson professes to have the mathematical wherewithal to fully follow those models himself). Wilson and Nowak publish jointly on these ideas. There can be little question of Nowak’s capability as a mathematician, but what is questionable is why he has stricken off in the direction he has – creating a model requiring such a very complex and highly tuned sequence of stages to arrive at any sort of a plausible mathematical justification of altruism via the route of group selection in social animals. If this wasn’t enough he uses this frail model to dismiss a multitude of much more simple and more robust models of the same effects. As Maynard Smith himself said, a model is of no relevance at all unless it has strong substantiation in experimental observation. This level of verification is certainly the state of affairs with kin selection, but not with group selection. I cannot help but wonder if this thrust of Nowak’s models are not somehow influenced by his strongly professed religious beliefs and the massive amount of funding he has received from the Templeton Foundation ($10 million I believe). After all, kin and individual selection reek of base materialism and determinism, while group selection can be envisaged as reflecting the “hand of god”. Or am I being a very naughty boy to even consider this possibility?

  10. MR. Coyne,

    I was just thinking about something like this over the last few days.

    in your words, “It is not even clear that altruistic groups of humans would beat non-altruists. Steven Pinker has noted that success of one group over another in the real world is based not on higher frequencies of altruistic individuals, but on matters like harsher discipline, better technology, and more brutal ideology”

    Where do you think the tipping point is with respect to fitness, when you consider the human population? We are definitely genetically loading ourselves and our species is becoming less fit in physicality, and we rely so much on better technology to allow extension of life and reproductive capacity of less than ideal individuals. Will our lack of fitness kill us off in the end, or will our selfish, warring behavior? or will we just go “poof” in a cataclysmic event? your thoughts?

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