Levels of selection: controversies no one cares about?

Bora made two quick references to "group selection" today. I don't have much time...and shouldn't be blogging, but I want to make a few quick points before this topic goes down the memory hole (I know, unnecessary caveat, but I am driven by personal guilt in expressing it, not public shame). For those "not in the know" (e.g., most readers), Bora and I have a history.

Update: Robert Skipper's ruminations are worth a read, as always. And of course I was just making shit up about his political views and draft....

My problem with Bora comes down to assertions like this:

And I have realized that opposition to multi-level selection is now reduced to a few die-hards who have either invested too much of their careers opposing it to be able to now back down, or people like Razib who oppose it for ideological reasons.

First, let us constrain the field here, "levels of selection" isn't really of interest to many (most) areas of biology. This is an issue that crops up in evolution and genetics, and it is not trivial, but if you aren't the type of person who enjoys books like Defenders of the Truth or some of Elisabeth Lloyd's papers you probably won't get worked up over this. But here is a concrete reason why I get worked up over this: I was discussing some of Robin Dunbar's ideas about the evolution of cranial capacity with a graduate student in anthropology and he mentioned that "selection happens to groups." I was like, "wait up here, selection doesn't happen to groups," and he was like, "Yes it does, evolution selects on groups," and I was like, "Nope, the dominant view is that selection occurs mostly on individuals, or, at least lower levels than groups, though groups are part of the extended context in which selection occurs." Long story short, I know this guy well enough to know that he isn't really conversant in the details of the levels of selection debate, and wouldn't be able to tell you who David Sloan Wilson is. But, he was expressing an intuition that many individuals possess that it is groups which are being selected up by natural selection. The most common manifestation of this is the "for the good of the species" type talk which peaked with VC Wynne-Edwards work in the 1960s.

At this time there was a "revolution" in evolutionary biology and W.D. Hamilton and George C. Williams shifted the focus and methodological bias from group to individual level selection. Roughly speaking these workers simply contended that there were all sorts of problems with "for the survival of the species" style arguments, the most simple one being that it seems that such behaviors are vulnerable to being exploited by cheaters who invade a social system. A more technical objection is that intergroup variance is simply too low when compared to intragroup variance on a trait for between group level variation to be competitive with within group variation over the evolutionary long haul.

But these are details and nuances you can explore if you have an interest in the field, I want to speak to two of Bora's points which I believe are disputable and/or bizarre.

First, I do not believe that "opposition to multi-level selection is now reduced to a few die-hards" as a matter of fact. I haven't seen a survey, and I'm a scientific small-fry so perhaps the big boys just aren't telling me what's going down (it isn't like philosophical debates about levels of selection are part of typical lab-chat), but just to check if the sky wasn't purple I did email a few friends who do graduate work in evolutionary biology and they don't think multi-level selection is the dominant paradigm either. This does not mean that Richard Dawkins style gene-level selection is universally accepted, and even Dawkins himself admits that higher levels of selection do occur. And I tend to concur, the point is whether scientifically this is a fruitful field of study with a vibrant empirical and theoretical research program. My impression is that it isn't vibrant, and when I read Unto Others I recall Wilson even admitting that unfortunately there was simply too little experimental work being done in this area (he was writing in the late 1990s). So, the issue here is not (for me) whether multi-level selection is correct or not, it is simply whether it is the dominant conceptual paradigm in evolutionary biology (I wouldn't bet on this), and whether it is even a vigorous one (I am sketchier here, but my impression is no, it isn't that vigorous). An important bigger issue, but one I won't address here, is whether generalizations like "most selection is on the level of the gene" or "selection is pluralistic" are really important aside from selling semi-popular press books and giving rise to a reasonable literature in philosophy of science (I do think that Dawkins style "gene selectionist thinking" is pretty weird and revolutionary to many people, and it is worth understanding and internalizing even if it isn't as revolutionary or ubiquitous as Dawkins might have you believe).

The second issue is Bora's attempts to make connections between these particular scientific ideas and a particular politics. Myself, I was a libertarian before I became interested in evolutionary biology, and I remain a libertarian despite my varied levels of emphasis on a "Fisherian" world-view over time (that is, individual level selection upon additive genetic variation as the prime driver of biological evolution). In fact, if you wanted to plot "libertarianism" and "individual selectionist" on the same graph as level of personal intensity the former would probably show a steady drop since circa 1997, while the latter would have peaked circa 2003. In terms of the bigger picture, consider the adaptionist evolutionary biologists who I believe Bora has in his sights, the school which began with R.A. Fisher, an Anglican Tory eugenicist, J.B.S. Haldane, an atheist Communist for most of his life (the Lysenko affair soured him on active participation) and W.D. Hamilton, an agnostic Thatcherite. Finally, there is the great popularizer, Richard Dawkins, who has taken gene level selectionism to the people, but is a confirmed Laborite militant atheist. In contrast, both Richard Lewontin and Stephen Jay Gould would be in the multi-level selection camp, and would be considered men of the Left, though I do not know where David Sloan Wilson would be slotted politically, and he has been the genuine evangelist on this issue for the past generation. I have pointed out that one Wilson's "followers" in extending group level selection is Kevin MacDonald, a noted intellectual anti-Semite who promotes the idea that Jews have been shaped by selection as a group over the past 2,000 years, and that European gentiles are vulnerable to parasitism by selfish groups because of their excessive altruism due to weak intergroup selection. The point which that bizarre digression speaks to is that correspondences between political and scientific orientations can be a fun parlor game, but it must be done judiciously, and in many esoteric areas (and levels of selection is one) there just isn't much there to work with because the topic is at too far a remove from public policy and political concerns.

Addendum: Note that one of those posts is rather old, so I don't know if Bora holds all those positions in the details at this point. But, I will say that I am tempted to make some argument about how Robert Skipper's line of reasoning about "genetic draft" is ideologically driven by beliefs as a conservative Democrat :)

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My take on this, as someone who during his grad school years thought far too much about this issue, is that while most biologists are willing to consider multi-level selection, the phenomenon doesn't seem to be that prevalent. Ultimately, this is an empirical, system-by-system question in the same way that the selectionist-neutralist debates were (although there are still diehards there too).

system-by-system question in the same way that the selectionist-neutralist debates were

i can buy that. also, taxa by taxa....

Trivers was, um, not so impressed w/ *Unto Others*: http://www.metanexus.net/metanexus_online/show_article2.asp?id=3041
And we know his political background (white Black Panther). Maynard Smith took the same view of group selection, and he was an ex-Communist. Here's a good quote that gets at the basic idea of reciprocal altruism / expulsion of cheaters, from one of the intellectual grandfathers of left anarchism, Peter Kropotkin in _Mutual Aid: A Factor in Evolution_:

'Let us take a group of volunteers, combining for some particular enterprise. Having its success at heart, they all work with a will, save one of the associates, who is frequently absent from his post. Must they on his account dissolve the group, elect a president to impose fines, or maybe distribute markers for work done, as is customary in the Academy? It is evident that neither the one nor the other will be done, but that some day the comrade who imperils their enterprise will be told: "Friend, we should like to work with you; but as you are often absent from your post, and you do your work negligently, we must part. Go and find other comrades who will put up with your indifference!" '

He didn't do any of the math, and wasn't so precise as later figures, but most of the book documents how members of the same species help each other out, rather than ruthlessly compete exclusively. This was in response to Social Darwinism, and again, he didn't do the math or formalize in any way, but this is like the discoveries in evolutionary game theory, where in realistic cases, everyone pursuing a hawk strategy won't be stable, thus will be invadable by doves. Now, Kropotkin thought he saw almost no "tooth and claw" behavior, so he undoubtedly down-played reality rather than present the nuances of frequency-dependent selection.

I don't want to give him too much credit, but the point is that, while mostly verbal, this book on evolution & cooperation by a seminal Left figure isn't out-and-out group selectionist, and shows a hint of what the formal models would later show. And as long as we're including ourselves, I take the standard view, and you can guess my rough poli / econ orientation...

Also, Chomsky prides himself on being a "sociobiologist" and many consider him a "genetic determinist" -- both facts dovetail perfectly w/ his conservative, hierarchical poli-econ orientation. Wait a sec...

Ah, Razib, that is so ancient...

First, I have modified my views several times since that old post and pretty much gave up on the whole political blogging endeavor - though I may repost some of the better oldies just for kicks (and with caveats) - don't need to react all over again.

Second, you and I do not define political ideology the same way or along the same axes. I do not use party affiliation, policy choices, or even results of online quizzes to define political ideology - I am looking more for psychological underpinnings of a worldview that may )or may not) lead one to adopt a particular political ideology, which in turn may (or may not) lead one to accept or reject evolution in the first place, which may lead the accepters of evolution to favor or disfavor hierarchical view of selection vs. one-level view. Still, none of this matters if you are a professional philosopher of evolutionary biology: you can be a communist or a member of KKK, if you study the issue in detail you just HAVE to accept that selection CAN act at different levels, then we can all go into the field and try to figure out how much it actually acts at which level in the real world (in which group selection is unlikely to be all that common for reasons you know very well and mention in this post).

Third, I do not think that "group selection" is a dominant paradigm in all biology, but that is is held by most of the people who have studied this phenomenon in particular detail. In a way, what I believe is that it is 'becoming' a dominant paradigm over time, as more and more philosophers of biology and evolutionary theorists get to understand how it works, and this may require a new generation of evolutionary biologists to replace the old school and finally start teaching the most modern evolutionary theory to undergraduates. It is a slow process.

As for non-professionals, if they accept evolution at all, they are most likely to fall for some kind of Desmond Morris crap because it is sexy. That is better than Creationism, but the battle is soooo uphill, I'd rather not waste time on lamenting that they do not understand Sober, Brandon or Hamilton - leave that to professionals.

On the other hand, as professionals are also a conduit of evolution to the popular press, we should do better than keep confirming the crude genocentric view. Neither Dawkins nor Razib are crude genocentrists, but there is quite a lot of that in the popular consciousness.

Here is an example of a die-hard in a prominent place - Clutton-Brock. He refuses to even discuss group selection and dismisses it with a magic wave of hand. And he has data of his own (on meerkats) that he says he does not know how to explain, yet a 1/10th second glance at the graph makes you scream "group selection!" - he will not accept it. If oyu mention it his face gets dark and he will not talk to you again.

I'm not going to get into the biological debate since my knowledge of biology begins and ends with knowing the evidence for evolution. But I do know enough about psychology, sociology, and political science to see that there isn't much (or any) evidence for Bora's definition of ideology. In a nutshell:

1. Spillover of bias into science correlates with traditional political ideology better than with sociological or psychological theories of bias. For example, all proponents of Cultural Theory agree that environmentalism is an example of Egalitarianism, but those who analogize it to religious cults tend to be predictably conservatives or libertarians more than Individualists or Hierarchists. For another example, Lewontin and Wilson are both Egalitarians, but their views of evolution are opposite and correlate better with their politics (Marxist vs. liberal).

2. Serious theories of bias and ideology have an element of Kuhnian surprise, whereby overwhelming evidence can shift a person from one bias to another. As a corollary, theories based exclusively on models of upbringing cannot work.

3. The main basis for Bora's theory is extraordinarily shaky. First, Lakoff's theory of frames isn't particularly mainstream. Second, Lakoff unfortunately engages in excessive introspection and explicitly eschews empiricism, something that's a lot more acceptable in linguistics than in social sciences. Third, the connection between Lakoff's cognitive theory and his political theory is weak at best.

What about selection for genetic architecture? Seems to me that's only relevant at the group level.

I'm thinking, for example, of the fact that the system of sexual reproduction in mammals is such that it is very hard for a mutant to suddenly reproduce asexually. Does that account for the success of this architecture? Or is it just chance?

I'm a big fan of Bill Hamilton, but not his rather silly ideas on human eugenics. He seemed to be worried that we are headed to a 'hospital' society while we allow mutants to survive. He vaguely suggested a 'voucher' system for good breeders. My leftish anti-authoritarian views immediately ask 'who dispenses the vouchers?'
S J Gould had politics more sympathetic to me, but I note his fat book has just one line on Bill H. even though Bill was called 'the greatest theoretical biologist since Darwin' or some such, by someone big (?)

man I was gonna read the fat book until I read that about one line on Hamilton.

bah!

also, screw group selection, and screw the lack of formal evolutionary theory teaching in bio undergrad.

Maybe someone can explain to me why when it comes to 'multi-level selection' biology gets hijacked by these troupe of philosophers (?) whose only achievement is to obscure and confuse arguments that have LONG been settled with rock-hard solid evidence (which of course is a completely alien concept to philosophers)???
By the way Sloan Wilson is a BIG TIME charlatan who wants to put himself at the level of Williams, Trivers and Hamilton and doesn't even play in the same league.