This post apparently didn't make it to Ediacara.  I'm reposting
because I reference this post elsewhere:

"Perplexed in Peoria"  wrote in
message news:...
>
> "John Edser"  wrote in message
news:cc1bkm$2lp8$1{at}darwin.ediacara.org...
> >
> >
> > Multi-level selection
> > Sean Rice
> > Dept. of Ecology & Evolutionary Biology
> > Yale University :
> >
> > http://pantheon.yale.edu/~sean/group.html
> >
> I am inserting here a portion of Rice's essay that
> John clipped.  It is important, though John probably
> did not realize this.
>
> [Rice writes:]
> I suggest that a useful test of whether selection is
> acting at some level is to ask the following question:
>
> In order for selection to act, is it necessary and sufficient
> for there to be multiple (>1) units at that level and for
> there to be variation between these?
>
> If the answer to this question is "yes", then we say that
> selection is acting at the level under consideration.
>
> We can see how this test works by applying it to the case
> of kin selection. Consider a population composed of a single
> family of full siblings. Now imagine that some of these
> offspring carry an allele for altruism and others do not.
> In this case there are multiple individuals and variation
> in the trait (altruism) that influences fitness. Individuals
> are also related to one another and interact. Thus, if kin
> selection is a case of selection acting on individuals or
> alleles, it should happen here.
>
> Since r = 0.5 for every pair of siblings in this scenario,
> [here, John begins his quotation from Rice with the remainder
> of this sentence]
>
> > "Hamilton's rule predicts that altruism should
> > increase in frequency if the benefit to the
> > recipient of an altruistic act is greater than
> > twice the cost of that act to the altruist. In
> > fact, altruism will always DECREASE in this
> > scenario, regardless of the costs and benefits".
> > [JE's capitalisation of DECREASE]
>
> I, and I think most other defenders of Hamilton in this
> group, agree that in Rice's scenario, altruism will
> decrease.  We agree, John, and we have always agreed.
>
> However, Rice is making two mistakes.  The first and
> obvious one is that he left the word "heritable" out
> as a modifier of "variation" in his proposed criterion.
>
> The second is less obvious, and may be debatable.  In
> fact, I have just completed a debate with Dr. Hoelzer
> on this subject:
>
> Rice, I claim, is wrong to state that r = 0.5 for every
> pair of siblings in this scenario.  Actually, in this
> highly artificial scenario, r = 0.  Hamilton's rule, if
> properly understood, does NOT predict that altruism will
> increase in this scenario.
>
> What Rice has done is similar to what Hoelzer did.  He
> has constructed a scenario in which IBD is no longer a
> good estimator of r.
>
> For details, refer to my debate with Hoelzer on the thread
> "Kin Selection Contradiction?"
>
> Hamilton's model is sometimes described as involving gene
> level selection, sometimes as involving individual level
> selection, and sometimes as involving group level selection.
> Rice's conclusion from his toy model seems to be that group
> selection is the best characterization of Hamilton.  The
> odd thing is that Rice is a proponent of group selection,
> and of the kin selection model, yet Edser welcomes his
> analysis because Edser believes in neither group selection
> nor kin selection.
>
> To repeat my own view on this, I think that all three
> viewpoints are defensible, but that the individual
> selection viewpoint is best.  Calling it gene level
> selection simply invites the standard objection to gene
> level models - namely epistasis.  It turns out that
> epistasis isn't troublesome to Hamilton, but this takes
> some effort to see.  Similarly, calling it group level
> selection invites various standard objections too.  These
> can be answered in the Hamilton case, but only with effort.
>
> This is not to say that the individual level viewpoint is
> without conceptual problems of its own.  Hamilton's model
> turns causality on its head in that the success of the
> altruistic gene derives not from what it causes in the
> donor (the altruism), but rather from what it is merely
> correlated with in the donor (his status as a disproportunate
> RECIPIENT of altruism).  And this is a subtlety that some
> people have trouble apprehending, let alone comprehending.
>
>
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