John Edser wrote:
>>>>JE:-
>>>>If -c is mutualism and +c is altruism
>>>>but c is arbitrary within the rule then
>>>>the rule cannot discriminate between them.
>>>
> 
>>>BOH:-
>>>Yes, I totally agree.
>>
> 
>>>JE:-
>>>Do you agree that the
>>>sign of c remains arbitrary
>>>within the rule?
>>
>  
> 
>>BOH:-
>>Yes.
> 
> 
>>JE:-
>>Do you now conclude that Hamilton's
>>rule has been misused to explain apparent
>>OFM in nature?
> 
> 
> BOH:-
> No.
> 
> JE:-
> Please explain why you are not just contradicting
> yourself. IF you agree that OFM and OFA can
> only be indicated within the rule via the
> sign of c but the sign of c remains _arbitrary_
> THEN, OFA cannot be separated from OFM can it? 

Of course it can.  By examining the sign of c.

> Thus Hamilton cannot separate OFA and OFM
> within nature, can he.
> 
Of course he can.  By examining the sign of c.

> 
> 
>>>JE:-
>>>Do you agree that Hamilton's rule only
>>>measures differences in relative fitnesses?
>>
>  
> 
>>BOH:-
>>Yes.  The fitnesses associated with carrying out the behaviour relative
>>to the fitness associated with not carrying out the behaviour.
> 
> 
>>JE:-
>>Then you agree that it is possible for the altruistic
>>gene to relatively spread as the absolute fitness of both
>>genes, i.e. Hamilton's hypothetical altruistic genes and the
>>wildtype non altruistic gene it is contesting becomes
>>absolutely reduced?
> 
> 
> BOH:-
> Yes.
> 
> JE:-
> So you agree that as Hamilton's OFA
> gene spreads, Hamilton's rule ignores the
> possibility that both genes may be paying
> the price of extinction to allow this?

Yes,  The concept is called evolutionary suicide.

> Do you agree that OFM entirely 
> prohibits such an event from being
> _selected_  but OFA allows it?
> 
I'm not sure, because I haven't thought about this in great detail.  My 
guess is that it would be possible to find a counter-example (at least a 
theoretical one).

To be honest, I have more important things to be doing at the moment, so 
  if I do work out an answer, it'll be on the bus home...

> 
> 
>>JE:-
>>..
>>All sterile forms come from non sterile
>>forms and not vice versa. The effect sterile
>>forms  can have is only selectable at the fertile
>>level of selection and not at a sterile
>>level of selection. Do you agree or disagree?
> 
> 
> BOH:-
> Yes, I agree.
> 
> JE:-
> Given your agreement do you now conclude
> that sterile eusocial casts must have a
> zero fitness, 

No.  They have zero direct fitness (assuming that a sterile individual 
is always sterile - if it has the option of becoming fertile, then it's 
fitness - or in your terms its expected fitness - is non-zero), but 
their indirect fitness does not have to be zero.

> 
>>snip<
> 
> 
>>JE:-
>>Isn't this an evolutionary discussion
>>group where the point of discussing OFA
>>here, is only about whether this "behaviour can
>>evolve"? Isn't it a fact that nothing we
>>have discussed allows OFM to evolve including
>>Hamilton's rule, i.e. Hamilton's rule has
>>been utterly misused as an evolutionary
>>mechanism that can allow OFA to evolve?
> 
> 
> BOH:-
> No, as I've been trying to point out to you, Hamilton's rule is still 
> applicable for mutualiism.  Far less than not allowing OFM to evolve, it 
> give the conditions under which is will evolve.
> 
> JE:-
> The above contradicts your agreement that the 
> sign of c is arbitrary! 

Rubbish.  Hamilton's rule is rb>c.  It is possible for this to be true 
whether c is positive or negative.  In that sense the sign of c does not 
matter - we can still use Hamilton's rule whether c is positive or negative.

> 
> 
>>>BOH:-
>>>(as long as the
>>>inequality in Hamilton's rule holds, then the prediction is that the
>>>behaviour can invade the population)
>>
> 
>>JE:-
>>No that is not the prediction! The prediction
>>is that a behaviour that may be EITHER OFM or OFA
>>can only increase _relatively_ in a population
> 
> 
> BOH:-
> This is, in essence, a re-statement of what I wrote.  Therefore we seem 
> to agree.
> 
> JE:-
> Then you agree that the reason
> this "behaviour can invade the population"
> was either via OFM or OFA so that Hamilton cannot
> validly claim that _only_ OFA allowed so called
> "altruistic" behaviour because no way exists
> using Hamilton's rule to prove the behaviour
> was "altruistic"! At all times the behaviour
> was EITHER altruistic or mutualistic, period.
> 
Again, we agree.  Hamilton's rule does not tell us whether a behaviour 
is atruism or mutualism, onlt whether the genes coding for the trait 
will increase in frequency.


>>JE:-
>>Do you agree that DARWINIAN competition
>>between donors must fail to let Hamilton's
>>hypothetical altruistic gene pass otherwise
>>Darwin stands refuted?
> 
> 
> BOH:-
> I have to admit that I'm not aware of Darwin's own position on the 
> levels of selection debate, and to be honest it onlt seems of historical 
> relevance.  Even if Darwin is refuted on this point, it makes little 
> difference.  His contribution to biology as a science was immense, and I 
> am more than happy to forgive him a few errors.
> 
> JE:-
> "Forgive him a few errors"?
> Your arrogance astounds me!
> It is the gene centric Neo Darwinian 
> establishment that keep making all the 
> errors because they have entirely
> and consistently deleted 
> gene fitness epistasis.
> 
*SIGH*  Fisher explicitly included epistasis in his fundamental theory. 
  And this is a separate issue to the levels of selection debate.

> The proposition was VERY SIMPLE:
> EITHER Darwin's fertile organism level
> allows or disallows the selection of
> Hamilton's OFA gene. Does it allow it
> or does it disallow it?
> 
I'm not sure that "Darwin's fertile organism level" allows or disallows 
anything.  It hardly seems to have the necessary capabilities to make 
these sorts of decisions.


>>>JE:-
>>>___________________________
>>>Do you understand
>>>the difference between
>>>"absolute parental fitness"
>>>and "population growth"?
>>>___________________________
>>
> 
>>BOH:-
>>Simply the difference between the
>>individual and the population.
> 
> 
> 
>>What aspect of "the individual"
>>does "absolute parental fitness"
>>represent and why is it _not_ a
>>"population" aspect?
> 
> 
> BOH:-
> Growth rate.  And because they may differ between individuals.
> 
> JE:-
> Exactly.
> Is it not a fact that "because they may differ between individuals"
> Darwinian selection must select there and then against any donor
> with a lower reproduction of fertile forms no matter if this
> event does not suit Hamilton's "selfish gene"?
>  
No.  Because it may lower the absolute fitness of EVERY individual in 
the population.  Growth rate goes down, but unless there is selection at 
the species level, this is not important.

> BOH:-
> Population growth rate is the average of the individual absolute 
> fitnesses.  Hence, if you only reduce individual absolute fitnesses, you 
> must also reduce the population growth rate.
> 
> JE:-
> Thus is it not a fact that within just
> "the average of the individual absolute 
> fitnesses" higher "individual absolute 
> fitnesses" ARE being selected over the
> lower?
> 
Yes.

Bob

-- 
Bob O'Hara

Dept. of Mathematics and Statistics
P.O. Box 4 (Yliopistonkatu 5)
FIN-00014 University of Helsinki
Finland
Telephone: +358-9-191 23743
Mobile: +358 50 599 0540
Fax:  +358-9-191 22 779
WWW:  http://www.RNI.Helsinki.FI/~boh/
Journal of Negative Results - EEB: http://www.jnr-eeb.org
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