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  Topic: multi-level selection models, examining observation and evidence< Next Oldest | Next Newest >  

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(Permalink) Posted: Jan. 05 2007,18:46   

I've been thinking about doing this for a while now, and considering I'm working on a new course curriculum for a course in evolutionary biology from a history of prediction standpoint, I thought it would be worthwhile to set up a thread as a repository of information and discussion on the history and current debate surrounding multi-level selection models.

Typically, this boils down to the debate over what level selection acts at:  gene, individual, group, population, species, or some combination of the above.

I'm going to keep the introduction to this topic short, as I really want the tread to become more of a repository of specific studies that examine the practical application of multi-level (or purely group) selection models, which those interested can discuss in detail at leisure.  So, I'm just going to very briefly touch on some of the history, and then go right into listing some references, and encourage others to post relevant references they run across, with commentary as they see fit.  As time permits, we can discuss specifics of some of the papers, and maybe come to some resolution as to the actual efficacy of some of the models that go beyond individual selection.  pardon the quick and dirty generalizations, and my leaving out players that are/were also important in the debate.

Because selection must act on phenotypes, and those phenotypes typically must be heritable for evolution to proceed via selection, it is natural that the early models developed by Fisher in the 1930's were based on the idea that selection operates at the level of the individual.

the idea of group selection in the "modern" era was basically introduced by Wynn-Edwards in the early 1960's, but the introduction of game theory and its application by Maynard Smith also in the 60's (think: cheaters) tended to invalidate the concept of selection acting on groups, as any group behavior would be subject to individual cheaters.  

Moreover, the explanation of specific behaviors such as those typically considered 'altruistic' also were considered hurdles to the idea of individual selection. However, with the introduction of the concept of inclusive fitness by Hamilton (60's, early 70's), much of altruistic behavior was seen to be readily explained and predicted by selection at the level of the individual.

Another shaping force around this time was Williams, who correctly pointed out that the notions of group selection postulated by Wynn-Edwards were simply unsupported by actual observational data, and essentially set the stage as one of the individual selection model being the most parsimonious based on observation.  Moreover, he created what is still considered the generally accepted rule for examining the veracity of any group-selection model:

"Our search must be specifically directed at finding adaptations that promote group survival but are clearly neutral or detrimental to individual reproductive survival in within-group competition." (Williams, 1966)

With that in mind the onus was placed on the shoulders of group selection supporters to eliminate any possible counter explanations from an individual perspective. This criterion makes logical sense in that one must disprove all other possible scenarios before proving the existence of group selection, however one could argue that it is too restrictive, and perhaps nigh impossible to experimentally achieve.

As we started more and more to apply the idea of selection to human behavior and morphology, we again saw another debate about the possibility of group selection raised by Gould and others in the 80's.  For example, Gould raised the possibility of group selection acting on race differences based on some of Wells earlier work.  There is some argument as to whether Gould was strictly supporting the idea of group selection as being worthy of pursuit in and of itself, or whether he really was trying to broaden the thinking of evolutionary biologists to gain a better understanding of higher levels of interactions between organisms and their subsequent influences on selection.

Regardless, the debate has continued, for the most part from a theoretical and philosophical level, ever since.  Empirical evidence to support group or multi-level selection models have typically been few and far between, and always controversial.  For example, several fairly recent reviews have concluded that long-term observations of birds, mammals, and other higher organisms have not encountered populations in which group selection seems to operate.  However, to counter that, other studies looking at social adaptations in microorganisms (some might remember a recent paper on slime molds that appeared on PT a few months back) have made the suggestion that group selection models do indeed best explain observations in these organisms.
Moreover, some have suggested that the observations of specific things like sex ratio bias also can only be explained with the use of multi-level selection models.

Having spent time embroiled in the middle of this controversy while a grad student at Berkeley in the late 80's/early 90's, I see the value of examining the evidence that actually supports or rejects the various models proposed by theoretical biologists and philosophers of biology.  So far, I tend to think that the application of multi-level or group only models are extremely limited at best, and not of significance from the overall standpoint of looking at evolution in general.

However, as all of us are typical busy, nobody can keep on top of all the literature in an area as large as this one, so the purpose of this thread is mainly to look at the practical application of multi-level and group-level models, to see just when and where (and IF), we should be considering the application in a more general sense than we currently are.

I hope that was short enough, but not too short.

I'm going to list some general and specific references to start, and keep adding to the list as time goes on, and anybody who has read any of the papers, or has their own they wish to contribute, is encouraged to do so and argue whether or not they think the data from the articles answers the long running debate:

Is group selection necessary?

As i find online, free versions of anything, I'll add a link to it.


WYNNE-EDWARDS, V.C. 1962. Animal Dispersion in Relation to Social Behaviour. Hafner Publishing Company. New York, NY. 653 pp.

online version of RA Fisher's original models of selection (er, and everything else about the man too):

Maynard-Smith J. (1964) Nature 201: 1145&#8722;1147

a site where you can check out Gould's work (including his commentaries on group selection):

(I'll add more later)

recent reviews of empirical evidence for selection models in large organisms:

Zahavi A & Zahavi A. (1997) The Handicap Principle Oxford University Press: New York.

...and in microorganisms:

Bassler BL. (2002) Cell 109: 421&#8722;424.

some specific studies of interest old and recent:

(note this is the list that I really want to focus on, and will be adding a lot more to this a bit later)

I know there are several others over on Pharyngula that wish to add their own references, so I'll open this up to them for now, and be back to add more later, and then maybe we can pick a paper or two to dissect and discuss.

for now, I'm happy to just keep this thread open as a repository, even if there is little interest in detailed discussion.


"And the sea will grant each man new hope..."



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(Permalink) Posted: Jan. 05 2007,19:24   

Hello all,

Title says it all:

- Goodnight et. al. (1995) Multlevel selection in natural populations of Imapiens Capensis Am. Nat. Vol. 145, No. 4 pp. 513-26

One such model suggested for multi-level selection is contextual analysis (which has a bit of lit. in epidemiology). I'm not sure whether it works or not (probably not) but it has some recent practitioners.
Examples of group selection w/ contextual analysis:

- Tsuji, K. (1995) Reproductive Conflicts in the Ant Pristomyrmex Rungens: Contextual Analysis and Partitioning of Covariance Am. Nat. Vol. 154 pp. 599-613.

Contextual analysis, from a sociologist.
Although this paper isn't about group selection, it is a first-rate paper about the uses and mis-uses of contextual analysis:

- Blalock, Hubert M. (1984) Contextual-Effects Models: Theoretical and Methodological Issues Ann. Rev. Sociol. 10:353-72.

And I know you'll probably complain about this link, but it is probably relevant: Biological Altruism

Also, Sober and D.S. Wilson's book here, may be worth looking at. If nothing else it has a really nice history talking about Price's eqn., Williams, Hamilton, etc...

have a good day

Bob O'H

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(Permalink) Posted: Jan. 06 2007,08:13   

I agree with curious that Sober & Wilson is useful, although one should always keep in mind that they're advocating one point of view.

For a overview of the sociology, have a look at Segerstråle's book Defenders of the Truth.

There's also Keller's edited volume "Levels of Selection".  The intention is obviously to move the debate on from the old discussions about group selection and trait group selection, and more towards JMS's evolutionary transitions viewpoint.  Because of this, the introduction is a hoot with its rather grouchy "oh, just shut up will you?" approach to the old debates.

Oh, and the three volumes of Bill Hamilton's Narrow Roads of Geneland have to be mentioned as well.


It is fun to dip into the various threads to watch cluelessness at work in the hands of the confident exponent. - Soapy Sam (so say we all)


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(Permalink) Posted: Jan. 06 2007,10:13   

Here's a good paper:

Author(s): QUELLER DC
Source: AMERICAN NATURALIST 139 (3): 540-558 MAR 1992

If you have Web of Science: check out the papers that have recently cited Queller. There's been a bit of a debate going on about the role of kin selection and group selection in the evolution of altruism. I don't have time to post all the links right now.

After much reflection I finally realized that the best way to describe the cause of the universe is: the great I AM.



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(Permalink) Posted: Jan. 06 2007,14:24   

thanks much, and just know I'll be keeping this thread open, occassionally popping in with commentary, and bumping it.

feel free to add specific commentary (positive or negative) wrt to any of the contributions in the thread.

I won't be able to get back to this until tuesday (busy writing).

"And the sea will grant each man new hope..."



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(Permalink) Posted: Jan. 06 2007,14:27   

I've been here for years, and even I'm still stunned occasionally by the expertise our little community has.


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(Permalink) Posted: Jan. 06 2007,18:17   

I hope you're just as stunned by the lack of expertise some of us have.

After all, in DaveWorld, the spread from lack of expertise to expertise isn't from bottom to top on the x axis, it's from side to side on the y, where the knowledge, detail, and sophisticated analysis of the expert are balanced--nay, trumped!--by the intuition, common sense, and simplicitas of the dunce.


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(Permalink) Posted: Jan. 07 2007,13:04   

Hello all,

As I said in some other threads, I think a good way to illustrate the problem is selfish DNA and genomic conflits :
- B chromosomes
B-chromosome evolution
Author(s): Camacho JPM, Sharbel TF, Beukeboom LW

- modile DNA (thousands of references)
- Segregation distorters
Author(s): LYTTLE TW
Source: ANNUAL REVIEW OF GENETICS 25: 511-557 1991

- conflict between nuclear and cytoplasmic DNA (including endosymbionts) causing
  * Cytoplasmic Male Sterility
Schnable PS, Wise RP
The molecular basis of cytoplasmic male sterility and fertility restoration
TRENDS IN PLANT SCIENCE 3 (5): 175-180 MAY 1998

  * Feminization, and other effects

When there is a conflict between units of selection (gene/individual/population), the gene always has the last word.
My position, like Dawkins', is that we can explain our perception of the living world more precisely if we consider that selection acts on genes.


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(Permalink) Posted: Jan. 07 2007,13:42   

I think you're setting youself up for some arguments from the evo-devo folks, but I especially appreciate this particular input in this thread, so thanks.

we now have both ends of the spectrum represented, and I expect eventually this will spark some heated debate.

I'd prefer to stick to specific papers rather than general discussion myself, so with that in mind, do you have a specific article you would like to champion, Jean?

"And the sea will grant each man new hope..."


Bob O'H

Posts: 2561
Joined: Oct. 2005

(Permalink) Posted: Jan. 07 2007,13:59   

A couple more theoretical papers from JEB:
Social semantics: altruism, cooperation, mutualism, strong reciprocity and group selection


The evolution of cooperation and altruism – a general framework and a classification of models

The latter has a bunch of commentaries after it too.  As I recall, they were quite entertaining.  I can't remember if they referenced any empirical work, though.


It is fun to dip into the various threads to watch cluelessness at work in the hands of the confident exponent. - Soapy Sam (so say we all)


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Joined: Jan. 2006

(Permalink) Posted: Jan. 07 2007,16:37   

Quote (Ichthyic @ Jan. 07 2007,13:42)
I think you're setting youself up for some arguments from the evo-devo folks, but I especially appreciate this particular input in this thread, so thanks.

we now have both ends of the spectrum represented, and I expect eventually this will spark some heated debate.

I'd prefer to stick to specific papers rather than general discussion myself, so with that in mind, do you have a specific article you would like to champion, Jean?

It's not evo-devo, as far as I can't tell, rather some situations found in nature that have been explain by selection acting on gene, in spite of their deleterious effects on population growth and the fitness of other genes.

I haven't read this papers actually. But maybe the review paper on segregation distorters is a good start.

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