jimmenegay{at}sbcglobal.net (Jim Menegay) wrote in
news:c4qqe1$30q0$1{at}darwin.ediacara.org: 

> wlhunt{at}earthlink.net (William L Hunt) wrote in message
> news:... 

>>  If anyone is interested, here is a paper that discusses the
"spite"
>> aspect of Hamilton's rule with a few examples (most notably the green
>> beard gene, Gp-9, in the red fire ant).
>>  Spite: Hamilton's unproven theory - Foster, Wenseleers, and Ratnieks
>> 2001
>> http://www.shef.ac.uk/uni/projects/taplab/pdf/fwrannzoolfenn2001.pdf

> Thanks for the link.  Several things about this paper are worth 
> comment.

I also thank William Hunt for the link. The paper definitely helped me 
understand the topic better.


> First, the three examples of spite all fit into a category I would
> call enforcement actions in defense of a larger pattern of mutualism.
> I believe that "spiteful behaviors" of this kind may actually be
> fairly common, and I hope to defend this opinion on the thread
> "Mutualism" that I recently opened.
> Second, it is amusing that one of the examples is "Green beard spite".
> I recently argued that green beard altruism is unlikely (due to
> epistasis) but my arguments don't apply to green beard spite -
> particularly if the spite is enforcement.

> Third, I notice that several possible examples of spite, by Bill
> Morse's definition of spite, are ruled to actually be examples of
> selfishness. That is, the authors believe that reducing the
> competition should not be counted as spiteful.  

Let me just note that the authors distinguish between Wilsonian spite and 
Hamiltonian spite. I was in fact alluding to Wilsonian spite (which does 
not require a negative r) but which could also be considered to be simply 
a subset of altruism - reducing the competition really just boils down to  
benefiting relatives. I agree that if reducing the competition results in 
a net reproductive benefit to the one doing the reducing, then there is 
no real cost and we are back to simple selfish behavior. I think we also 
agree that in the context of intelligent social animals, some behavior 
that may be spiteful in the short run in fact serves a policing function 
that helps longer term reciprocal altruism.

> Hamilton's 1970 paper
> seems to support this interpretation.  However, in Chapter 6 of
> "Narrow Roads", Hamilton says some remarkable (to me) things that seem
> to indicate that he wishes he had adopted a more Morse-like
> interpretation back in 1970-1971.  To set the stage, let me first
> include what I wrote against Morse. 
 
>> >Negative r is, of course, a theoretical possibility, but not a
>> >realistic one IMO in models of human spitefulness (xenophobia). 
>> >Hamilton's rule cannot validly be used directly to explain the kinds
>> >of active hostility to other tribes that Tim Tyler is fascinated
>> >with (and receives so much grief for).  It may well be involved
>> >indirectly in such an explanation, mediated by reciprocity, sexual
>> >selection, or whatever, but it cannot be directly explanatory. 
>> >There is just no way for the genes to know that a neighboring tribe
>> >has a small negative r with respect to this tribe, rather than a
>> >small positive one. 
 
> Hamilton wrote:  "One of the most surprising conceptual points in the
> (1971) paper ... may seem even contradictory to altruism being
> correlated with relatedness.  This is in my verbal caution that low
> dispersal by itself (population "viscosity") as a way of reaching high
> relatedness has snags.  The point is that, to be effective, altruism
> must put offspring into competition with non-altruists, not bunch them
> in a wasteful competition with their own kind.  Recently this point
> has been treated explicitly and quantitatively with some quite
> surprising conclusions." [References here to 1992-1994 papers by
> Queller and others]. 

(snip)
 
> Still, Hamilton made a big advance by noticing that, panmixis or not,
> everyone is not equally related to everyone else, and this fact can be
> captured in the factor r.  Morse now points out that everyone is not 
> equally in competition with everyone else. Perhaps this should be
> captured in r as well?  Of course, we will have to switch to absolute
> fitnesses in computing b and c, to avoid double counting, but this
> should be no problem. We can keep "rb>c", but now the
definition of r
> is different. 
> 
> We have the defining equation:
>   r = g - e
> where
>   g = genetic relatedness (this is the old r)
>   e = ecological competitiveness. 
> 
> If we do this, we will notice the effect that gave Hamilton so much
> pain - in a viscous population, g and e tend to move together -
> cancelling each other out.  Hence, there may be no net tendency toward
> altruism. 
> 
> However, when you look in more detail, you see that e doesn't always
> have to cancel out g.  Wright, in "Non Zero" makes the case that e is
> frequently negative for H. sap.  And, if you are a zebra migrating
> across a savanna with plenty of fodder and a finite number of lions,
> each with finite appetite, a certain gregariousness seems called for.
> 
> Mr. Morse, I'm not sure how much of this theory is yours and how much
> is mine, but I think I kind of like it!
 

I will cheerfully give the credit to you. I had not intended to fold the 
ecological competitiveness into the equation for r, but that may in fact 
be easier than the "double counting" method. While the cited paper (and a 
nice explanation you give in a separate follow) helped clarify why r is 
calculated as a comparison to the average relatedness, and why a negative 
r is possible, I don't really like the convention. My thought had been to 
have an absolute r and absolute b's. Then, if i represents members of a 
population of size N, you would calculate the sum over i from 1 to N of 
rsubi times bsubi in order to arrive at a number to compare to c.

But in practice this is extremely cumbersome, and it misses the 
generality of your equation above. I will have to think more about this, 
but at first glance one of the nice features of r = g -e is that it can 
encompass interactions with other species, allowing the equation to be 
generalized to symbiosis. Good work, Jim!

Yours,

Bill Morse
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