Re: Hamilton's rule
- From: "Perplexed in Peoria" <jimmenegay@xxxxxxxxxxxxx>
- Date: Sat, 19 Nov 2005 23:22:58 -0500 (EST)
"Guy Hoelzer" <hoelzer@xxxxxxx> wrote in message news:dll6au$1qor$1@xxxxxxxxxxxxxxxxxxxxxx
> in article dliqtb$ohd$1@xxxxxxxxxxxxxxxxxxx, Perplexed in Peoria at
> jimmenegay@xxxxxxxxxxxxx wrote on 11/17/05 12:58 PM:
>
> > "Guy Hoelzer" <hoelzer@xxxxxxx> wrote in message
> > news:dlh6sj$274$1@xxxxxxxxxxxxxxxxxxxxxx
[snip]
> >> Under your graphical model, which I accept as a representation of Hamilton's
> >> model, the strength of kin selection diminishes monotonically with the
> >> frequency of the allele.
> >
> > Well, if you mean that the difference between the inclusive fitness of an
> > altruist and the inclusive fitness of a member of the general population
> > decreases monotonically, then yes.
>
> That's right. Selection is generated by fitness differences, and the
> strength (effectiveness) of selection is determined by the extent of fitness
> differences.
>
> >> I guess you have swayed me to the position that
> >> under Hamilton's model, as opposed to a more realistic version of the same
> >> model, there is not a frequency threshold where kin selection becomes
> >> irrelevant. Instead, it becomes irrelevant monotonically.
> >
> > In the same way that any beneficial trait becomes irrelevant as more and
> > more of the population has that trait.
>
> Well, you have been arguing that it happens in a different 'way' because the
> strength of selection changes in a sigmoid fashion relative to frequency for
> 'ordinary' beneficial traits. A simple way to see this is to use Joe's
> freeware program called PopG, which simulates changes in allele frequencies
> under specified models of evolution. If you start a simulation with a
> beneficial allele in low frequency, you will see the sigmoid trajectory of
> its frequency by tracing the 'infinite population size' line that eliminates
> the noise of drift. While some aspects of the trajectory's shape will
> depend on dominance/recessiveness of the allele, it is always sigmoidal.
I claim that the trajectory of the altruism trait is sigmoid in almost exactly
the same way that the trajectory of any other trait is sigmoid. Felsenstein
recently delivered himself of the same opinion.
> > But perhaps a better way to look at it is to compare the possessor of the
> > trait to (not the population average, rather to) an individual without the
> > trait. If you compare in this way, then I don't agree that the 'strength of
> > kin selection' decreases monotonically. Once the allele becomes fixed in the
> > population, there is still strong selection AGAINST mutations that change the
> > altruism allele to selfishness (i.e. ineffectiveness).
>
> Here I think you have contradicted yourself.
I don't think so, though apparently I have contradicted your understanding
of what I said.
> While you have been convincing
> me of the validity of your argument along the way, that same argument
> persuades me that your last point is false. A largely altruistic population
> would be quite vulnerable to invasion and persistence of low frequencies of
> alleles for selfish behavior. If what you meant was that the largely
> altruistic population is unlikely to be replaced by a regime of selfishness,
> then I would agree.
The easiest way to refute your latest claim is to note that Hamilton's
rule also applies to selfishness. Selfishness genes are contra-selected
if rb < c with b and c negative.
If rb>c for altruism, then rb<c for selfishness. A selfish population is
subject to invasion by altruism, but an altruistic population is not
subject to invasion by selfishness.
To repeat for the last time, Hamilton's rule, as an arbiter of what traits
are favored by selection, IS NOT FREQUENCY DEPENDENT.
.
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