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?
> 
No.


>>BOH:-
>>Depends on which definition of fitness you want to use, because it
>>doesn't matter.
> 
> 
>>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?
> 
Yes.
> 
> 
> 
>>>>>>>JE:-
>>>>>>>1) No sterile form can have any fitness
>>>>>>>at all, i.e. it cannot be altruistic
>>>>>>>because it has no fitness to give away.
> 
> 
>>>>>>BOH:-
>>>>>>Are you saying that a sterile form cannot
>>>>>>act  to help a related individual?
> 
> 
>>>>>>JE:-
>>>>>>Obviously not.
> 
> 
>>>>>BOH:-
>>>>>So are you saying that any help provided by a sterile
individual cannot
>>>>>change the fitness of the related individual?
> 
> 
>>>>>JE:-
>>>>>The answer IS IN WHAT YOU SNIPPED.
>>>>>I explicitly wrote that a sterile
>>>>>form cannot have an INDEPENDENT
>>>>>FITNESS.
> 
> 
>>>>BOH:-
>>>>But that wasn't what I was asking.  I was asking whether it
is possible
>>>>for a sterile individual to act in such a way as to change
the fitness
>>>>of another, related, individual.
> 
> 
>>>>JE:-
>>>>ANYTHING can, the wind, a tin can,
>>>>wearing a hat with monkey on it...
>>>>So what?
> 
> 
>>>BOH:-
>>>Well, if a sterile individual can have an effect on a related
>>>individual's fitness, then it seems to use this fact when modelling the
>>>fitness of a group of sterile and non-sterile individuals.  Especially
>>>when looking at individuals who can switch between sterility and
>>
>>fertility.
> 
> 
>>>JE:-
>>>A sterile individual, like a hat with a monkey
>>>on it, can have an effect on related or non
>>>related individuals. So what?
> 
> 
>>BOH;-
>>Well, if a sterile individual can have an effect on a related
>>individual's fitness, then it seems to use this fact when modelling the
>>fitness of a group of sterile and non-sterile individuals.  Especially
>>when looking at individuals who can switch between sterility and
> 
> fertility.
> 
> 

> 
> JE:-
> OK.
> 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?
> 
Yes, I agree.

>>>>BOH:-
>>>>Hamilton's rule isn't intended to be used as a
discriminator of altruism
>>>>and mutualism.  Why should it?  It applies to both cases, and can be
>>>>used to predict whether a behaviour can be selected for.
> 
> 
>>>>JE:-
>>>>Then you admit that because the
>>>>sign of c within Hamilton's is
>>>>arbitrary then Hamilton's rule
>>>>has been consistently misused to say
>>>>when OFA or OFM is operating within
>>>>the rule? Please answer, yes or
>>>>no.
> 
> 
>>>BOH:-
>>>No.  One can say if a behaviour is altruism or mutualism by estimating
>>>the signs of c (oh, and b - positive c, negative b would be parasitism)
> 
> 
>>>JE:-
>>>Here we go around the mulberry bush...
>>>You have proven that the sign of c is
>>>arbitrary.
> 
> 
>>BOH;-
>>For deciding whether a behaviour can evolve, yes..
> 
> 
> 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?
> 
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.

> 
> 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

This is, in essence, a re-statement of what I wrote.  Therefore we seem 
to agree.

> where this relative increase can require an absolute
> reduction of both the altruistic and non altruistic
> genes.
> 
The relative increase "CAN require an absolute reduction", but this is 
not necessary.  Again, we're in agreement.


>>JE:-
>>Hamilton's gene can only invade the population
>>if it can pass the fertile organism
>>level of selection. Yes or no?
> 
> 
> BOH:-
> Again, this question makes no sense.
> 
> JE:-
> Well it can only make "no sense" if you
> do not understand the logic of Darwinian
> selection at the organism level. Since
> all the selective competition when rb>c
> is supposed to explain OFA
> is between donors and not between recipients
> (if it was between recipients then OFS would
> be the behaviour under examination and not OFA)
> then the donors who give nothing away are
> always selected at the organism level over
> those that give something away simply
> because donors are Darwinian organisms that
> must compare their _organism_ fitness results
> and giving any of it away reduces their fitness.

John, this demonstrates that you simply haven't understood Hamilton's 
central point - if you are going to look at the evolution of a behaviour 
that affects not just the actor (i.e. the donor), but also other 
individuals (i.e. the recipients), then you need to consider the 
fitnesses of all of the individuals, and the probabilities of them 
carrying the alleles that code for the behavioural phenotype.

> Do you agree that DARWINIAN competition
> between donors must fail to let Hamilton's
> hypothetical altruistic gene pass otherwise
> Darwin stands refuted?
> 
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:-
>>>How can you say if a behaviour
>>>is altruism OR mutualism! ALWAYS it
>>>is altruism AND mutualism  and you
>>>can never know which!
> 
> 
>>BOH:-
>>No.  It is altruism if c is positive, and mutualism if c is negative.
>>These are mutually exclusive events, so you cannot have both.
> 
> 
>>JE:-
>>Since you have proven that the sign of c is
>>arbitrary within Hamilton's rule then
>>it is _impossible_ to know when c is
>>positive or negative so it is impossible
>>to verify EITHER OFA or OFM within
>>Hamilton's rule.
> 
> 
> BOH:-
> No.  You go into the real world and measure it.
> 
> JE:-
> We are attempting to INTERPRET measured observations
> of the real world , 


> Unless you have clear theories of OFA and OFM
> on the table as  testable views of nature, it becomes
> impossible to test between OFA and OFM. 

As I wrote earlier:  "It is altruism if c is positive, and mutualism if 
c is negative."  That seems clear enough to me.


>>>>>>>>>JE;-
>>>>>>>>>__________________________________
>>>>>>>>>please explain how OFM can allow
>>>>>>>>>the SELECTION of a REDUCTION in
>>>>>>>>>absolute fitness.
>>>>>>>>>__________________________________
> 
> 
> 
>>>>>>>>BOH:-
>>>>>>>>By poisoning the environment with a toxin
for which you, and your
>>>>>>>>relatives, have limited immunity.  It goes
on in bacteria, where it's
>>>>>>>>mediated by a plasmid which as both the
genes for production of the
>>>>>>>>toxin, and a gene for resistance to it. 
All you need is a cost of
>>>>>>>>resistance, and you have a clear example.
> 

> 
>>>>>>>>JE:-
>>>>>>>>Here a SELECTION for an
>>>>>>>>absolute reduction in fitness
>>>>>>>>is not occurring. We all agree
>>>>>>>>that absolute fitness reduction
>>>>>>>>can and does, occur but it is
>>>>>>>>not chosen by selection.
>>>>>>>>snip<
>>>>>>>>The population is predicted to deal
>>>>>>>>with these toxins by curtailing
>>>>>>>>population growth,
> 
> 
>>>>>>>BOH:-
>>>>>>>What is "curtailing population
growth" if not reducing absolute
>>>>>>>fitness?
>>>>>>>And at what level is this curtailment acting?
>>JE:-
>>This is evasive pedantry.
>>There is such a thing as
>>"a curtailment".
>>___________________________
>>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?
> 
Growth rate.  And because they may differ between individuals.

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.

Bob

-- 
Bob O'Hara
Department 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: www.jnr-eeb.org
---
þ RIMEGate(tm)/RGXPost V1.14 at BBSWORLD * Info{at}bbsworld.com

---
 * RIMEGate(tm)V10.2áÿ* RelayNet(tm) NNTP Gateway * MoonDog BBS
 * RgateImp.MoonDog.BBS at 2/23/04 3:00:47 PM