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echo: evolution
to: All
from: William Morse
date: 2004-01-21 11:04:00
subject: Re: : Re: Hamilton`s rule

"John Edser"  wrote in
news:bu7blf$1r78$1{at}darwin.ediacara.org: 

I have left in most of the previous argument, even though our esteemed 
moderator may object, so we know what 1) and 2) refer to.


 
>> JE:-
>> Spencer only use the word "survival" to mean 
>> how long any organism lived. Thus:
>> Those that live longer are fitter, because
>> fitter forms must live longer.
>> If you define fitness in genetic survival terms then
>> Spencer's jingle equates "survival" with both maximal 
>> phylogenetic survival (genes over organism generations)
>> AND maximal organism reproduction, thus allowing the 
>> word to be ambiguous, i.e. it can mean two CONTRADICTORY 
>> causations:
>> 1) Survival = Maximal organism reproduction.
>> 2) Survival = Maximal gene replication. 
>> Since 2) is empirically fitness dependent on 1) 
>> and not vice versa, then only 1 is a valid _causation_.
>> Nobody here will admit that 2) is entirely 
>> dependent on 1). Hamilton argues that 1)
>> is entirely dependent on 2) against ALL
>> THE EVIDENCE using just an invalid proposition,
>> Hamilton's rule.
> 
> 
> WM:-
> Well we finally see the root of your objection to Hamilton, and it is
> - (surprise, surprise) - "gene centrism"! 
> But your problem seems to be only in your own mind. It is obvious that
> naked genes will not survive outside organisms - so in order for genes
> to reproduce the organisms that carry them must reproduce. Hamilton's 
> argument is in fact based on that proposition. Hamilton DOES NOT argue
> that 1) is dependent on 2).
> 
> JE:-
> Then you do agree that 2) is dependent on 1)?

Yes. Genes do not reproduce outside of organisms (other than in the lab).
 
> WM:-
> What Hamilton does argue is that maximal 
> organism reproduction does not mean maximal reproduction of only one's
> own offspring, but must include reproduction of any offspring that
> share one's genes. 

> JE:-
> Unfortunately for Hamilton, all adult 
> (fertile) forms within the same population 
> must compete against each other using Darwinian 
> reasoning BEFORE fitness at Hamiton's gene
> level can even start. Even adult 
> relatives must compete against each
> other. This being the case, any reduction 
> of fitness at the donor organism level via 
> Hamilton's gain at just a supposed gene 
> level of selection, is always selected against 
> at the actual Darwinian organism level of the 
> donor.


This is simply not true. There are several examples of eusociality in the 
animal kingdom. I believe you have argued against haplo-diploids such as 
bees as being a true counter example to your logical objection, since the 
workers are not capable of reproduction. But there are also examples of 
animals that are capable of reproduction forgoing their own to help 
relatives. Perhaps the best example is the naked mole rat. The conditions 
that could give rise to a mammal  exhibiting eusocial behavior was 
predicted by Richard Alexander based on Hamilton's rule before it was 
known that naked mole rats were eusocial, and before Alexander even knew 
that naked mole rats existed! 

Even without eusociality, there are numerous examples of animals, such as 
the meerkat and the Florida scrub jay, where conspecifics help out others 
in raising young at a cost to themselves. You would  do well to review 
the article on page 634 of the October 24, 2003 issue of Science. You 
might also try doing a search on the web for some of these species. Yet 
another interesting example is slime molds (although I have no idea if 
the slime molds that combine to form a fruiting body are related). Slime 
molds are aggregations of individuals, some of whom do not reproduce 
while others get to become reproductive spores. 

To put it very simply, there are in the real world tests of your 
statement above, and they prove it to be incorrect. Since you love 
testable theories, how do you respond to that?


 
> It does not matter that organism fitness
> altruism allows a hypothetical increase in 
> fitness at just a supposed gene fitness level if
> this costs an absolute decrease in donor organism
> fitness. Hamilton's organism fitness altruism
> caused by his selfish geneism is selected 
> against before it can even begin. This alone,
> is logically consistent with the fact that
> 2) is dependent on 1). If a gene fitness
> was actually independent of organism fitness 
> then Hamilton's argument may be sustained. Not 
> a single independent genomic gene fitness has 
> ever been documented within nature. Hamilton
> was living in a population genetics fairy land. 

Let me try to parse the above. The first sentence is clearly false - if 
"organism fitness altruism" gives an increase in "gene
fitness", then it 
must give an increase in the total number of organisms carrying the gene 
This follows from your previous statement (with which I agreed) that an 
increase in gene fitness can only be achieved by and increase in 
organisms carrying the gene. . In this case even if the "donor" organism 
has a decrease in fitness (which is what Hamilton supposed) there is 
still an overall increase in organism fitness, i.e. the number of 
organisms carrying the gene. Since this is clearly true, your second 
sentence is clearly false. Your third sentence is clearly false, since 
(as I have just shown) a "Hamiltonian" increase in fitness is logically 
consistent with 2) being dependent on 1). Your fourth and fifth sentences 
are moot, since nobody is arguing that gene fitness is independent of 
organism fitness. In your sixth sentence you mysteriously misspelled John 
Edser ;-)     




(snip) 

> JE:-
> Darwin was correct. However, what nearly everybody has 
> neglected to understand is that only epistatic gene
> information and not just "genes" are important
> "in determining survival". Genes are only mere
> letters of the alphabet. Heritable information 
> is coded in words, sentences, paragraphs pages, 
> chapters and books.

So far I am in agreement, except that very many have explicitly stated 
that they understand this, including your arch nemesis Mr. Dawkins.

> Hamilton deleted all gene
> epistasis within has rule as well as any 
> representation of absolute fitness. You may
> have noticed that I did bother to include 
> epistasis within:
> 
>           r^eb>c
> 
> What chance can a selfish letter have battling 
> an entire encyclopaedia of information?

Hamilton's rule includes epistatic effects by definition, since they 
affect the value of b. And of course he represents absolute fitness, in 
fact he extends it to be an accurate measure of absolute fitness instead 
of confining it to just the fitness of one organism.

Having said all the above, there is in fact considerable discussion in 
the real world of evolutionary biology over whether the observed 
"altruistic" care for relatives in a number of animal species is due to 
kin selection according to Hamilton's rule or is due to other effects. 
Perhaps, John, you could discuss real results of real research. My 
prediction is that you won't.

Yours,

Bill Morse
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