Truth and Reconciliation for Group Selection XI: Dawkins Protests (Too Much)

Richard Dawkins did not invent naïve gene selectionism (see T&R X) but he spread it far and wide with the publication of The Selfish Gene. Let's follow his logic, beginning on page 6 of the 1989 paperback edition:

This book will show how both individual selfishness and individual altruism are explained by the fundamental law that I am calling gene selfishness. But first I must deal with a particular erroneous explanation for altruism, because it is widely known, and even widely taught in schools. This explanation is based on the misconception that I have already mentioned, that living creatures evolve to do things 'for the good of the species' or 'for the good of the group'.

There's the caution against naïve group selectionism. Good for you, Richard! Now for the explanation of why it is naïve:

[A] group, such as a species or a population within a species, whose individual members are prepared to sacrifice themselves for the welfare of the group, may be less likely to go extinct than a rival group whose individual members place their own selfish interests first. Therefore the world becomes populated mainly by groups consisting of self-sacrificing individuals. This is the theory of 'group selection', long assumed to be true by biologists.

Right! This explains how "for the good of the group" traits might evolve. Now for the explanation of why they might not evolve.

The quick answer of the 'individual selectionist' to the argument just put might go something like this. Even in the group of altruists, there will almost certainly be a dissenting minority who refuse to make any sacrifice. If there is just one selfish rebel, prepared to exploit the altruism of the rest, then he, by definition, is more likely than they are to survive and have children. Each of these children will tend to inherit his selfish traits. After several generations of this natural selection, the 'altruistic group' will be over-run by selfish individuals, and will be indistinguishable from the selfish group. Even if we grant the improbable chance existence initially of pure altruistic groups without any rebels, it is very difficult to see what is to stop selfish individuals migrating in from neighboring selfish groups, and, by inter-marriage, contaminating the purity of the altruistic groups.

Done. Allow me to make three observations about these passages.

1) Dawkins' portrayal of within- and between-group selection is utterly standard, and that's a good thing. Throughout the T&R series, I have stressed the simplicity of the group selection controversy. From Darwin to Dawkins, it's all about what I have called the original problem (see T&R II).

2) Dawkins implies that between-group selection (described in the second passage) is no match for within-group selection (described in the third passage), but he provides no proof. Models (such as the modified haystack model), experiments, and field studies are required to make this determination, not rhetorical flourishes.

3) The status of genes as replicators -- what Dawkins calls the fundamental law of gene selfishness -- is utterly beside the point. Genes are the replicators regardless of whether altruism wins or loses in Dawkins' own scenario (also see T&R X). Using the replicator concept to argue against group selection is arguably the greatest case of comparing apples with oranges in the annals of evolutionary thought!

I do not believe the cynical adage "science progresses -- funeral by funeral" but I worry that it might be the case for Richard Dawkins on the subject of group selection. In my dreams, I imagine him reading my modified haystack model and saying "Well done, David! I have been wrong all these years. It turns out that a gene coding for altruism can evolve on the strength of between-group selection, even when it is selectively disadvantageous within groups. I do think it is important to keep in mind, however, that when altruism evolves by group selection, it is still an example of gene selfishness because the gene for altruism is more fit than the gene for selfishness, all things considered."

Scientists would clap their hands red at such an act of nobility, but it hasn't happened yet. Instead, Dawkins has behaved like a cowboy fighting off the Indians in an old western movie. When one gun runs out of ammo, he grabs another and another. Here are three guns that have run out of ammo.

The selfish gene gun. As we have seen, this gun didn't have any bullets to begin with.

The vehicle gun. Recall that if individual organisms don't qualify as replicators, they must quality as something else to be so manifestly well adapted. That "something" is variously called "interactors", "targets", or "vehicles" (Dawkins' term) of selection. In one of Dawkins' famous metaphors, genes in individuals are like rowers in a rowing crew. Since they are literally "all in the same boat", they must "pull together" to win the race. As soon as Dawkins threw away the selfish gene gun, he started to claim that groups fail as vehicles because individuals in groups are not completely bound together in a common fate in the same way as genes in individuals. But this was never a requirement for group selection to occur! The groups in the haystack model and in Dawkins' own portrayal of group selection quoted above aren't like individuals in this respect, but they suffice for altruism to evolve despite its selective disadvantage within groups.

The extended phenotype gun. When the vehicle gun didn't work, Dawkins decided to bury the vehicle concept altogether by describing genes as having extended phenotypes. Two examples of extended phenotypes are a bird's nest and a beaver's dam. The first is an individual-level adaptation in conventional terms; birds that build better nests raise more offspring than birds in the same group that build worse nests. The second is a group-level adaptation in conventional terms; beavers that build better dams are providing a public good for all of the beavers in the pond at their own expense. The fact that the genes result in alterations of the physical environment in both cases is irrelevant. In short, the concept of extended phenotypes doesn't address the original problem and certainly doesn't provide a novel solution.

With the Indians closing in, Dawkins has now started to throw chairs, bite, and kick. Consider his response to my recent article with Edward O. Wilson in American Scientist magazine titled "Evolution 'For the Good of the Group'"

Genes Still Central: David Sloan Wilson's lifelong quest to redefine "group selection" in such a way as to sow maximum confusion--and even to confuse the normally wise and sensible Edward O. Wilson into joining him--is of no more scientific interest than semantic double talk ever is. What goes beyond semantics, however, is his statement (it is safe to assume that E.O. Wilson is blameless) that "Both Williams and Dawkins eventually acknowledged their error [that the replicator concept provides an argument against group selection]...I cannot speak for George Williams but, as far as I am concerned, the statement is false: not a semantic confusion; not an exaggeration of a half-truth; not a distortion of a quarter truth; but a total, unmitigated, barefaced lie. Like many scientists, I am delighted to acknowledge occasions when I have changed my mind, but this is not one of them. D.S. Wilson should apologize. E.O. Wilson, being the gentleman that he is, probably will.

Gracious! What a hierarchical guy! Dawkins acts as if he is the No. 2 monkey, kowtowing to the No. 1 monkey (Ed) while dishing it out to the No. 3 monkey (me)! As Ed commented to me after reading Dawkins' comment, "What does he think--that you slipped me a Mickey?"

If you still have the patience, let me make a few observations about this gem of a tantrum. First, why on earth would Dawkins title his response Genes Still Central ? Isn't he ever going to get over the fact that selfish genes have no bearing whatsoever on the group selection controversy?

Second, I trust that I have provided ample evidence that the original problem has provided the basis for defining group selection for everyone, including Dawkins and myself. There has been no redefining.

Third, when it comes to semantic confusion, you can't beat selfish gene theory. Genes are "the fundamental unit of selection" but this has no bearing on the "levels of selection" controversy. A gene might be selfish because it is selectively advantageous within groups, or it might be selfish because it evolves in the total population all things considered. Individuals might be perfected "vehicles" of selection now, but we also need to use the term "vehicle" to explain how such perfection evolved. This is how the simplicity of the original problem has turned into a terminological quagmire.

Dawkins continued his tantrum on his website after Ed and I quoted the passage in The Extended Phenotype where he abandons the replicator argument and takes up the vehicle argument:

The Wilson quotation from The Extended Phenotype is a ludicrous attempt to justify their lying statement that I "eventually" acknowledged an earlier error. For one thing, The Extended Phenotype was published way back in 1982, which makes nonsense of Wilson's "eventually". But more important, the point I was making in 1982 (and would make again now) was a general one about the important distinction between replicators and vehicles...I was explaining that those models of group selection that had been proposed were vehicle models not replicator models. I was not for a moment suggesting that I accepted those models as valid. They were (and are) invalid vehicle models, as opposed to invalid replicator models.

There you have it from Dawkins himself. The word "eventually" is appropriate for the six-year period between 1976 and 1982, regardless of how much time has elapsed since then. Poor Richard is still trying to fend off the Indians with the butt end of his replicator gun ("genes still central") and vehicle gun ("invalid vehicle models"). If only Bill Hamilton was still alive to fight alongside and defend his brilliant ideas!

Fortunately, Bill Hamilton wrote plenty while he was still alive, showing that Richard Dawkins fights on alone.

To be continued.

Comments

[whitecap]

Thank you for this series of posts on the state of play in the multilevel selection debate Dr. Wilson. I have to confess that I haven’t attended overly much to articles, interviews, etc, featuring Richard Dawkins over the years, so my understanding of his ideas is sometimes second hand. But even as a non-believer myself, his militantly intolerant stance toward religious phenonema has long made me rather suspicious about his capacity for objectivity in that area, and after encountering your work and that of many other scholars (e.g. Peter Turchin, Peter Richerson, Robert Boyd, Joseph Henrich, Jonathon Gottschall, Brian Boyd) who make use of multilevel selection theory, it seems clear that it is by far the more promising paradigm for understanding the observed facts of human prosociality (inclination toward altruism and altruistic punishment, capacity for large-scale cooperation, etc), human cultural evolution, psychology, history – everything about us in fact!

[whitecap]

Somewhat off-topic question, but after going “paleo” this past year and eliminating grain and sugar from my diet as best I can (thoroughly recommended!), I've been interested in the changes to human life since the advent of the Holocene, in particular the shift toward increased inequality and hierarchy that accompanied the rise of agriculture and civilization and how that relates to our innate psychology.

A recent article by Kameda et al. contends that “although income inequality sharply divides industrialized societies all over the world, material egalitarianism may still operate as a fundamental principle affecting social sharing and exchange under uncertainty in many domains of human activity”, nonetheless, “human minds may be structured fundamentally as egalitarian sharers and that merit-based ideologies which pervade our modern lives may be seen as an adjustment on top of such a psychological foundation”.

I was wondering, do you see the rise of inequality during the Holocene as representing in part, at least, the “re-emergence” of ancient primate dominance hierarchy, as for example Polly Wiessner suggests, or are we fundamentally egalitarian? Have the “supra-tribal social systems” of the past 5,000 years been ultimately unstable because they fail to satisfy our tribal social instincts, at least until more inherently stable forms of organization such as representative democracy (we hope!) emerged to mimic the “reverse dominance” social structure of our ancient tribal past?

[whitecap]

Sorry for the multiple postings (that 250 word limit can be tough), but I wanted to provide the details for the references cited in my previous post:

Kameda, T., Takezawa, M., Ohtsubo, Y., & Hastie, R. (in press). Are our minds fundamentally egalitarian? Adaptive bases of different socio-cultural models about distributive justice. Retrieved 12 June 2008, from http://lynx.let.hokudai.ac.jp/~kameda/UBC_Erlbaum_Draft_Kameda_revised.pdf

Wiessner, P. (2002). The vines of complexity: Egalitarian structures and the institutio­nalization of inequality among the Enga. Current Anthropology, 43(2), 233-269. Retrieved 1 May 2009, from http://www.santafe.edu/files/gems/bowles/Wiessner6752614.pdf

[David Sloan Wilson - Huffpost Blogger]

Following Christopher Boehm (read Hierarchy in the Forest), I think that egalitarianism is deeply built into human nature, but it is built upon typical animal dominance hierarchies. Thus, at least some individuals will try to dominate, but they are usually successfully suppressed. Human egalitarianism is GUARDED egalitarianism. When social control mechanisms break down, then dominance quickly reasserts itself. Social control can be exceptionally difficult in large-scale society, but it is still possible. Finally, two kinds of social hierarchy need to be distinguished--the exploitative kind, in which dominants benefit at the expense of the subordinates, and the kind that benefits everyone. All societies must differentiate as they increase in scale, and people will tolerate hierarchy when it is "for the good of the group". Victor Turner's "The Ritual Process" shows how rituals are used to keep powerful people humble and to preserve the egalitarian sense of "communitas" even when society has "structure".

[David Sloan Wilson - Huffpost Blogger]

Thanks very much--I am often portrayed as the Lone Ranger of group selection, but in fact there is an active and growing community, including the folks that you list. I encourage readers to visit their websites by googling their names, for access to their publications and lists of their books.

[Jay Farar]

Professor D.S. Wilson,

Please consider moving your blog to a more appropriate venue. These are extremely good and important science post you have been offering. Having them on Huffpo in way unfairly diminishes their respectability just by virtue of being here. In the other words, the quality of the science, writing and presentation far exceeds the ability of Huffpo to offer the type of exposure called for with your blog postings. This site is primarily political with no science section (as you know) and very questionable science offered as popular postings, usually by celebrities who are not scientist (and since the site has an obvious political bias that also taints the objectivity of science).

I'm sure you've considered what I am talking about, so you may have thoughts on the matter already. I may suggestion a situation similar to what Massimo Pigliuuci has, or perhaps going over the ScienceBlogs (there are other options of course, including for science). At this point I think its getting the right amount and type of exposure (and if I may suggest - while avoiding taking funds from organizations which may not have the best interest of science in mind).

Finally, if you do decide to move, which I am really hoping for, please take this series of blog postings to present wherever you land. The confusion Dawkins has caused and the general misunderstanding of the science (stemming mainly from areas you have so eloquently detailed) make it important these post follow you.

Thanks,
Jay

[David Sloan Wilson - Huffpost Blogger]

Thanks for your comments and suggestion that I take my blog elsewhere. It's true the these blogs appear only briefly on the HuffPost, which does not showcase science as much as it should (see my previous blog on that topic). Fortunately, HuffPost blogs are picked up by many other sites, so they develop permanent presence on the internet. I receive feedback from all over the world about them, in addition to the posted comments. Finally, I intend the "Truth and Reconciliation" series to have a permanent impact on the field. The blog format should not mask my serious intent.

I'll think about better blogging options and check with my colleagues such as Massimo.

[brjohn9]

It does seem that Dawkins has always insisted that genes as replicators must mean more than it could possibly mean. His language suggests that even individuals are not subject to selection, let alone groups. Darwin would be baffled.

[David Sloan Wilson - Huffpost Blogger]

Let me stress that the "gene's eye view" is a popularization of the concept of "average effects" in population genetics theory and can be very insightful, as long as it isn't used as an argument against group selection. In my book "Evolution for Everyone", in the chapter titled "The Case of the Mad Monkey" I use the gene's eye view to show how a gene can evolve that causes (some) males to become psychopathic, based on positive effects expressed in females. The benefits must outweigh the costs for the GENE to evolve, but the benefits can be concentrated in some individuals and the costs in others. The tallying of costs and benefits takes place at the gene level, not the individual level. This is what George Williams meant by bookkeeping.

You're right that the replicator concept acquired a life of its own. I look forward to what historians will say about this period of evolutionary thought. Why did it become necessary to stress that genetic evolution takes place at the gene level, as if there is an alternative???

[John T]

I posted David's blog on rd.net, perhaps we'll see some spill over since they seem to move in packs on the internet when directed (while often patting themselves on the head for being calm and rational).

Jrogger, if you haven't seen, Richard posted a pretty funny one today:
http://richarddawkins.net/article,3767,Truckling-to-the-Faithful-A-Spoonful-of-Jesus-Helps-Darwin-Go-Down,Jerry-Coyne#368197

It's strange in a way to watch Richard dismantle himself so completely. Watching him lately on an Origins panel and a speech at the AA, he just looked even more desperate than he has in recent years. It appears he now seems the need to advocate even greater ridicule and ask: "Or should we continue to go along with the appeasers and be all nice and cuddly, like Eugenie and the National Academy?" The nearly bizarre cult of personality that has risen around him is equally strange and odd to witness again in my lifetime happening within a skeptical movement. It's usually defended by dismissing that charge altogether with claims of "he's just a brilliant man that I admire" or something like that. I guess we can't look forward to Richard and his group to "go along with the appeasers" anymore, heh.

[David Sloan Wilson - Huffpost Blogger]

No response from Richard yet, and I'm not holding my breath. There is a forum on multilevel selection being organized in London in July (still tentative) and a symposium at the European evolutionary biology meetings in Turin, italy, in August (certain), but Richard will not be at either one.

I should add that Richard has not contributed to the scientific literature on this subject for many years. He might be a public icon, but when I and my colleagues debate multilevel selection, no one feels the need to mention Richard.

[royniles]

Dawkins' attitude is a bit surprising, but may have something to do with problems in both of your positions. One thing stands out from reading the American Science article cited. You note that human hunter gatherer societies differed from the supposed "cheater" advantage dynamic in that they were fiercely egalitarian. You propose this was due to selection for teamwork which acted to suppress cheating activities.
What I think both your side and Dawkins' fail to consider is the effect of punishment on the degree of cheating that prevails. And punishment differs when the resources are scarce and the environment is too dangerous not to require the utmost in cooperation. And differs according to the ability of the group to detect cheaters and not only apply punishment but predict the consequences for not applying it severely. Killing our cheaters may be almost uniquely human.

[royniles]

Plus humans can predict such consequences for the long term as well as short term where other life forms cannot. Which will tend to make successful cheating a well thought out process and even at times an acceptable cooperative strategy where prospective inter group competition is concerned.
Cheaters in peace becoming assets in times of war. Altruistic cannon fodder perhaps.
Kind of throws the whole multi-level analysis for a bit of a loop?

[royniles]

And if this is about the original problem of understanding how and why first level groups existed in forms that were essentially dysfunctional, with the answer being that second level groupings provided functionality, this invites the paradox of reciprocal causation. Because what is the overall point here - that if it wasn't for the second level group selection, the first group hierarchy wouldn't have been formed? But if there had been no such hierarchy preceding it, there would be no second group. Did the second level formation act to create the first level hierarchy when at the same time the first level hierarchy enabled the second level to function?

Doesn't it seem more likely that the initial grouping had to have been successful in order to enable the next group to capitalize on that success? And if we start to look at the picture after the second group is already functional, might we have missed seeing that the first group had changed because the emphasis now switched to that of maintaing the effectiveness of the second level operation? And thus the so-called cheaters were more likely to be losers in the game without which there would have been no team of players to begin with?

[royniles]

Further, I have to ask, why are these "cheaters" here at all of they aren't in some sense fulfilling a need, perhaps for some sort of corrective function - perhaps they give a group flexibility of purpose, without which a group should eventually self-destruct. Do away with this diversity and you may well have some biological variety of fanaticism in its place.

And in a way, there's a strong inference here that if what these evolutionary biologists label "cheating" leads to greater relative fitness of cheaters, then it's not exactly cheating. Not if fitness is a desirable goal and "cheating" is the best strategy for obtaining it. So-called cheaters might be better labeled as outsmarters, innovators, inventors, creators.

In sum, these alleged cheaters may represent the potential for dissent in any biological grouping, without which a group would not have the flexibility or capacity to adapt when necessary. From this perspective, "cheating" by any measure seems at the very least to be a necessary evil for survival.

[David Sloan Wilson - Huffpost Blogger]

I and my colleagues DO emphasize the importance of punishment, even if it was not appropriately showcased in this particular blog. There is no egalitarianism without social control, which begins with low-cost mechanisms such as gossip and escalates to execution when necessary. Christopher Boehm made this point in a 1993 Current Anthropology article and his 1999 book Hierarchy in the Forest. The academic literature on "altruistic punishment" is all about this. The punishment is altruistic because imposing social control benefits everyone in the group at a private cost (however small) to the punisher.

[royniles]

This explanation of why punishment is altruistic simply exposes the weakness in the altruistic punishment concept. It's saying that if there is any cost at all to the punisher, no matter how small, this fits with the acceptable definition of altruism. But how often is the punisher buying a larger reward by expending this initial effort? In some way, every role in a social group hierarchy involves the same sort of ultimately profitable trade-off.

[Jrogger]

I have to say, David, its hard to take Dawkins seriously anymore. His rants against religion have done him no favour and he now looks more and more like every other hack on a tirade. His comments on your work with Ed Wilson is sad; sad that a man once respectable for his contributions to evolutionary thought now cannot tolerate any dissent. I think the excessive reverence that falls his way within the evolutionary field (and I suspect primarily within the EP field) has finally corrupted his sense of self-worth.

[David Sloan Wilson - Huffpost Blogger]

I don't want to beat up on Richard TOO much. He has turned thousands of people on to evolution, for example. However, I agree with you that science is all about being held accountable for what you say, and that Richard has become unaccountable, in part by becoming a public icon. That disqualifies him as a spokesperson for science.