Re: Hamilton's Nonsense
From: John Edser (edser_at_tpg.com.au)
Date: 01/06/05
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Date: Thu, 6 Jan 2005 06:11:42 +0000 (UTC)
"Perplexed in Peoria" <jimmenegay@sbcglobal.net>
> > > JM:-
> > > In fact, Hamilton's rule ("The gene (or trait) will increase
> > > in frequency if rb>c") is valid regardless of the frequency
> > > of the gene (or trait) in the population.
> > JE:-
> > The gene or trait in the population cannot
> > even RELATIVELY increase (an increase
> > measured as just a comparison between incomplete
> > totals of the altruistic allele and the
> > the wildtype allele) if the mutated allele
> > exists as dominant to its competitive wildtype
> > non altruistic allele and 100% kin selects.
> JM
> A relative increase is what I am talking about.
JE:-
Unles a relative increase refers either directly or
indirectly to an absolute (total) fitness count then
the idea is NOT scientific in any objective sense.
Popper was the first to underatnd such a _basic_
because without any absolute (total) fitness supposition
the model and the theory it was simplified/
oversimplifed from remain irrefuatble. Any
non refuitable idea can only be measured to
be verifified or non verified allowing "anything
goes" in the name of the sciences. Why the
mental block? Why do you flatly _refuse_ to name
the refutable theory from which Hamilton's non
refutable model was derived by the Neo Darwinian
process of modelling simplification/over simplification?
> JM:-
> And sure it can increase in frequency even if
> it is dominant and the behavior is expressed 100%
> of the time. Why not?
> I suspect that you are making
> the assumption that the donor is sacrificing ALL
> of its fitness here. But that is not necessarily
> the case. It is only sacrificing some of its fitness
> (i.e. in your world view, some of its children) to
> somewhat increase its relative's fitness (i.e. it
> is gaining lots of nephews).
JE:-
Because resources remain finite. If you
are forced to employ 100% of your resources
to reproduce by proxy, then even though you
remain fertile, you naturally reproduce just
a zero number of only infertile forms.
> Hamilton's rule even explains WHY "altruism" is not
> favored by selection in the extreme case in which the donor
> sacrifices its life.
> In this case, as you would point
> out, the recipient is likely to have no fitness of its
> own, because it is doomed to sacrifice its own life in
> a foolish act of "altruism". But since the recipient
> has no potential for children, doing it a favor cannot
> increase its fitness. So "b" is zero and Hamilton's
> rule correctly predicts that the gene will not increase
> in frequency.
JE:-
The actor can die because of his altruistic act
and b remain > 0. As long as the donar transfers
enough b resources before it "sacrifices its life"
and rb > c then according to Hamilton's rule the
donars death can be selected _for_.
> > JE:-
> > Jim, you are REQUIRED to mention EXACTLY what
> > can and CANNOT be measured by the rule and
> > list all over simplifications that Hamilton et al
> > stipulated.
> JM:-
> Surely I am not REQUIRED to mention all of this stuff
> every time I talk about the rule. That stuff is in the
> textbooks, as I keep trying to tell McGinn.
JE:-
In any BASIC discussion this reuqirement remains
absolute.
> > JE:-
> > You must also provide a REFUTABLE
> > THEORY with at least one point of refutation
> > that exists within it, from which
> > Hamilton's non refutable over simplified
> > model was simplified.
> JM:-
> No, I don't think that points of refutation are needed
> for models. Only for theories. And Hamilton's rule
> is a model, not a theory.
JE:-
Jim, I find this reply dissapointing. I reduces
your honest broker approach.
> > Logically you cannot
> > make modelling simplifications/oversimplifications
> > from just nothing at all! This question
> > ALWAYS remains ignored because Hamilton et
> > al cannot provide any answer to it.
> >
> > 1) What can be measured by Hamilton's Rule.
> > Just a 100% relative comparison of
> > the freq. of only one genomic allele at
> > just one locus. The rule cannot measure
> > the total fitness count of any allele because
> > it defines fitness as always ongoing so that
> > no total, i.e. finite fitness now exists.
>
> Again you are using "measure" in a sense different
> than I do. If you had titled this paragraph "what can
> be *predicted* by Hamilton's rule", then I would agree
> with your claim that the frequency prediction is a
> relative one. (That is what frequency MEANS in pop gen.)
> I'm not sure what you are saying, though in the sentence
> about "always ongoing".
>
> > 2) Simplifications:
> >
> > i) Random mating.
>
> Yep, as regards the 1964 "IBD" version. The 1970 "regression"
> version of the rule eliminates the random mating assumption at
> the cost of complicating the notion of relatedness.
>
> > ii) Zero epistasis including epistatic gene
> > fitnesses.
>
> Disagree.
>
> > iii) Dominance ignored.
>
> Strongly disagree. Much of the complexity of the 1964 paper
> exists because Hamilton did not simplify by assuming haploid
> genetics. He deals with the full diploid case and handles
> all of the complexities of partial and complete dominance.
> IIRC, he even discusses the complexities introduced if you
> have overdominance or underdominance.
>
> > iv) Gene counts only over organism generations
> > and not gene generation making Hamilton's fitness
> > count non logically self consistent. For the count
> > to become self consistent ALL gene replications from
> > each gene parent, including mitotic replications,
> > must be counted.
>
> Bullshirt. You keep saying this, but it is total nonsense.
> Hamilton's logic is completely at the organism level - not
> the gene level. So of course he is counting over organism
> generations. It was only later that someone (perhaps Dawkins)
> came up with heuristic justifications of the rule based on
> gene-level selection. And, in any case, if you really want to
> do gene-level modeling, you should calculated your fitness
> counts based on surviving descendents after a fixed time period
> (a year, say) rather than after a generation.
>
> > iv) All fitness totals remain ongoing and
> > therefore incomplete, by definition.
>
> If you are saying here that "b" and "c" represent fitness
> increments and decrements, rather than total fitnesses, then
> you are correct. I consider this a strength of the model,
> rather than a weakness. Altruistic behaviors are, after all,
> only one of the things that contribute to fitness. There are
> many other things going on simultaneously. But we don't want
> to investigate those other things, we want to investigate
> social behavior. So the rule talks about only those aspects
> of fitness that are impacted by social behavior.
>
>
> > 3) Over Simplifications:
> >
> > The total Darwinian fitness of the actor
> > remains deleted from the rule.
>
> As it should be.
>
> > 4) The theory from which the model was derived.
>
> The model is not derived from a theory, it is derived from a
> previous model - Fisher's. And it is a complexification, not
> a simplification, of the parent model.
>
> > [an excerpt from my book in preparation "Happy
> > Selecton's In Nature" (all rights reserved)
> > now follows:
> >
> > "Darwinian theory is the only refutable evolutionary
> > theory that exists, to this very day. This theory
> > states that each parent must maximise the total
> > number of fertile forms it reproduces into one
> > population because those that do not do so for
> > _any_ reason are selected against. This means that
> > the total Darwinian fitness of any Darwinian selectee
> > cannot be _selected_ to be reduced.
>
> So, if a hypothetical experiment measures that the fitness
> has been reduced, how can you tell whether it was _selected_
> to be reduced or whether it was reduced by something other
> than selection?
>
> > A measure of
> > Darwinian fitness is the total number of fertile
> > forms reproduced into one population by each parent.
> > This Darwinian fitness maximand is the only refutable
> > maximand that exists within biology. It can be tested
> > to refutation via a simple reversal of its selective logic.
> > This logic states that every parental total fitness is
> > compared to every other by simple default within one
> > population. Via this comparison parent/parents with the
> > largest total are selected for. In fact every form is
> > selected for if it trends to increase and not decrease
> > its total Darwinian fitness. This means that even though
> > a form is selected against using total fitness comparisons
> > as long as it exists within an expanding sub population
> > it has a future. The converse also applies: if a form is
> > selected for but is trending to an absolute Darwinian
> > fitness decrease it is heading for extinction. This is
> > only possible if a form increases its relative fitness but
> > decreases its absolute fitness. In economics this is known
> > as a relative gain for an absolute cost. Only Darwinism
> > explicitly prohibits such irrational action to be selected
> > for.
> >
> > If the total Darwinian fitness of every member of one
> > population can be artificially maintained as equal
> > then all evolution by natural selection must now cease
> > within that population because every maximand fitness that
> > is compared by simple default remains the same for all
> > selectees. If all selection is not halted then total
> > Darwinian fitness stands refuted. Only rhe total
> > Darwinian fitness proposition can force all evolution
> > within a natural population to be halted. No other
> > defined fitness, including Hamilton's can do so.
> > This is why only Darwinian total fitness can be
> > tested to refutation and only it constitutes a rational
> > fitness concept within biology.
> >
> > Hamilton et al established a 2nd but only heuristic and
> > therefore non refutable level of selection that was
> > incorrectly allowed to contest and win against Darwin's
> > refutable single fertile form level of selection forcing
> > organism fitness altruism via selfish geneism. A model
> > gene level of selection was argued to force a lowering of
> > Darwinian organism fitness via just one genomic gene maximising
> > its own (selfish) fitness independent of the Darwinian fertile
> > organism level of selection and all others genomic gene
> > fitnesses. To achieve this revolution Hamilton's gene must
> > be allocated an _independent_ fitness. Selfish geneism
> > cannot function if Hamilton's gene fitness remains epistatic
> > in fitness to any other gene within the same genome because
> > these genes are now being forced to compete against each other
> > and not just cooperate to provide a Darwinian fitness benefit.
> > To allow this hypothetical state all complex non linear fitness
> > (gene fitness epistasis) had to be deleted along with all other
> > non linear (epistatic) information. Thus Hamilton et al firstly
> > simplified Darwinian theory via his deletion of all epistasis.
> > This also required random mating to avoid the geometric costs
> > of epistasis.
>
> I don't see how random mating avoids "the geometric costs".
> As I see it, the only way to avoid those costs is to avoid
> mating altogether and become parthenogenic. Since this method
> of avoiding the costs is so rarely used, it is probably safe
> to conclude that the "costs" are not very severe.
>
> [remainder of Happy Selecton excerpt brutally snipped without
> having been read]
>
>
>
>
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