Hamilton - fitness mutualisation

"John Edser" <[email hidden]> wrote in message news:<[email hidden]>...

Quoted message said:

Quoted message said:

JM:- Immature forms are seen by John as acting on behalf of the parents.

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JE:- What else could they possibly do?

[snip]

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JM:- There are some other curious aspects to this. What a Hamiltonian would call parent-child
conflict must be viewed as a conflict between the two parents in John's accounting. As a
consequence, we might expect the children of strictly monogamous parents to be better
behaved. :-}

Quoted message said:

JE:- Please elaborate.

JM:- Trivers, I believe, has written about parent/child conflict - for example, the child does not
appreciate being weaned. Trivers interprets this as a conflict of interest between child and
mother. However, Edser believes

Quoted message said:

maturity. Therefore, weaning conflict is a conflict between the interests of the mother and the
interests of the father (in whose interests the child is also acting). The father wants this child
to be well fed. The mother, on the other hand, thinks that the child is ready to start eating
solid food, so that she can get pregnant again (perhaps with a different father). However, in a
monogamous species, the interests of father and mother are the same, so the child is expected to
align with those common interests.

JE:-

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because they have evolved opposed_ reproductive strategies.

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at all, then these opposed reproductive strategies must be resolved. This resolution requires
a ritual,
i.e. a way of winning and losing using "a game". I will elaborate on this further down.

What is being maximally reproduced within Darwinism are not genes, cells, organs or even
organisms, just _fertile_ organisms. It is the mutualisation of _opposed_ strategies that provides
an absolute but not necessarily equal larger fitness gain to each mutualised parent. This is what
the process of fitness mutualisation is all about: setting up games that provide absolute gains
for all. Everyone is a winner, but everyone has to pay a cost (e.g. losing a game) which is much
less than the mutualised gain (everybody wins) but more than cost incurred by just going it alone
(losing the war). Also, because mutualised gains are not necessarily equal, natural selection
still operates when each parents absolute fitness is compared to every other, within one Darwinian
population. As far as inheritance is concerned the assumption is that parental absolute fitness is
a complex heritable trait.

Neo Darwinian game theory does not include any measure for absolute parental fitness. Like
Hamilton's rule, these games only compare a relative with yet another relative so they can only
produce an _arbitrary_ winner allowing selection for cheating. In a mutualised game, cheating
provides a relative fitness gain at an absolute fitness loss to all players including the cheater,
so it is selected against. Because no absolute measure of each player's fitness exists within Neo
Darwinism, the absolute cost of winning by cheating cannot be measured so it remains invisible
allowing cheating. The only thing that is visible to Neo Darwinism is the winner in just
_relative_ terms.

Quoted message said:

Males take more risks and mostly die earlier compared to females but are amply compensated by
having a potentially higher absolute fitness using polygamy, if and only if, these children are
raised to fertile adulthood. For females their fecundity (the reproduction of immature forms) is
determined by how many babies they can have at birth

Quoted message said:

it is for males. Like the males, only the number raised to fertile adulthood count as valid
fitness units. IMHO females are better served by serial monogamy (exchange poorer males for better
ones in serial order) and males by polygamy (cram as many females as you can on one territory).
Thus the chess board is set and the game begins: males are black (more aggressive) and females
white (more cooperative) and they must play to win. The resolution of the game provides a relative
winner and a loser where however, both win, _absolutely_. Those that do not play the mutualised
game forgo HIGHER gains. Their costs are lower but their gains are lower so they are selected
against. Those that cheat to win a mutualised game win relatively but lose _absolutely_, so
cheating is selected against. The problem within a mutualised game is to find a point of
mutualisation using conventionalised competition. This is something like a "market price" where
the seller selfishly maximises profits and buyer selfishly minimises costs. Obviously the extreme
conditions where the buyer has a zero cost and the seller makes an infinite profit are not an ESS
(evolutionary stable strategy) and is the simple reason why selfishness is not the same as
mutualism. Mutualists balance seller and buyer so both make an absolute gain. For the seller,
relative profits are measured every trade but absolute profits are a _total_ of trades made over a
defined time period. Thus the seller can only maximise his profits when profit*turnover within a
defined time period INCREASES. Just increasing relative profits, i.e. the profit per trade, can
REDUCE abolute profits because selling at a higher price may reduce the numbers sold over a
defined time period. In This instance it pays the seller to REDUCE his price in order to INCREASE
his absolute profits. For the buyer, pushing the price below a mutualised price sends sellers to
the wall so the goods and services just disappear. Thus for seller and buyer finding the
mutualised price is for everybody's benefit. Here capital is funnelled into goods and services
demanded by consumers at an optimal rate. The capitalist system only works when sellers and buyers
are free to find a mutualised price. Parents use the same logic to mutualise their absolute
fitness counts.

JM:- John's "accounting rule" that attributes the child's actions to the parents creates a more
complicated conceptual structure than does Hamilton's method of accounting which has every
organism (even immature and infertile ones) acting in its own interest. On the other hand,
Hamilton's idea of "inclusive fitness" is much more complicated than John's. Hamilton needs his
fitness to be inclusive, otherwise an infertile form would have no fitness.

JE:- OK.

JM:- Those who are familiar with the philosophical foundations of economics and game theory may
worry about how a child can be said to balance the interests of both parents, giving equal weight
to each. In critiques of the ethical doctrine of utilitarianism, for example, theoretical
economists object to the idea of "equal weight". Those objections may well apply in economics,
where utility functions are freely chosen. The objections do not seem to apply in evolutionary
theory, because Nature is assigning the utility function. (PS. If this paragraph makes no sense to
you, don't worry about it. It is addressed only to people who understand the distinction between
cardinal utility theory and ordinal utility theory.)

JE:- The above is important. Please explain it in a more simple way so that sbe readers can more
easily understand it and identify what your explaining within evolutionary theory

JM:- I'm not sure exactly what you are looking for, since I am not sure that it does apply to
evolutionary theory, but I will try.

Economists talk about a quantity called "utility" that has some similarity to the quantity that
evolutionary theorists call "fitness". John finds, picks, and eats some strawberries. Economists
would say that John has incrementally added to his utility. Evolutionists, noting the nutritional
value of the strawberries, would say that John has incrementally added to his fitness. John got
scratched up while picking the strawberries. Economists would say there was a decrement to John's
utility from the scratches. Evolutionists would say that there was a decrement to John's fitness
from the scratches.

Which was greater, the increment from the eating, or the decrement from the scratches? An economist
considers this to be an objective question. He answers by observing John's behavior. If John knew in
advance that he would get scratched, and fetched the strawberries anyways, then the utility (to
John) of the strawberries was greater than the disutility of the scratch. If Jim refuses to fetch
strawberries under these conditions, then the disutility must be greater (for Jim) than the utility.
John and Jim appear to disagree, but actually both are right. There are two different utility
"scales" involved - JimUtility and JohnUtility. John is the sole arbiter of JohnUtility and Jim is
the sole arbiter of JimUtility. The economic theorist has no basis for judging between the two.

An evolutionary theorist looks at it differently. Eating the strawberries increased the expected
number of offspring John succeeds in producing. But getting scratched decreased it. It is still
an objective question which is greater, but John's preferences have nothing to do with it.
John's instincts could be wrong. If the scratching does more harm than the eating does good,
then an evolutionary theorist would say that John had lowered his fitness (produced fewer
offspring) by fetching and eating the strawberries. The economist still thinks John was right to
fetch - if offspring are not important to John, then so be it. The evolutionist might say that
John made a mistake.

Now, it turns out that economists have no way of comparing JohnUtility units to JimUtility units.
That is, they cannot come up with an exchange rate between the two kinds of utility. It is possible
that John and Jim can come up with a mutually agreeable exchange rate through trade and
reciprocity, but the economists cannot compute and impose one. JimUtility is strictly incomparable
to JohnUtility. Hence, any attempt by Jim and John to ally themselves according to Bentham's
principles and both try to maximize the sum of JimUtility and JohnUtility is hopeless unless Jim
and John first agree on some kind of exchange rate. The sum cannot be computed. We are trying to
add apples and oranges.

Evolutionists, on the other hand, believe that JohnFitness and JimFitness are measured in the same
units. John gains exactly as much by raising one

JohnFitness units from JimFitness units - we can just talk about fitness - plain and simple.

All of this is irrelevant for most purposes in evolutionary theory. But, there are some cases where
the fitnesses of one individual must be compared with that of another individual. For example, if
Jim is an altruistic donor of fitness, and John is the recipient, then Hamilton's rule (rb>c)
involves comparing "c" which is measured in JimFitness, with "b" which is measured in JohnFitness.
Not a problem, in evolutionary theory, as long as the units are the same - and they are. (Or at
least they are if the theorist is careful. I believe that Hamilton [1964] was careful.)

Now, back to the child that is supposed to be acting as a "somatic extension" of its parents. How is
it supposed to behave? It is not acting to maximize it's mother's fitness exactly, nor is it acting
to maximize its father's fitness exactly. John must say that it is maximizing a 50-50 blend of the
two kinds of fitness. And, that is a well-defined concept in fitness theory. It would not be a well-
defined concept in utility theory.

But now, here is a weakness of John's model. The child's genome is not really a 50-50 blend of the
two parents. There is first a random selection of half of the genes out of each parent to form
zygotes. The child is a blend of two zygotes, not of two parents. It is possible that the child's
"vector" for maximizing its own fitness is not a blend of the two parents. It may point in some
other direction altogether. John's theory allows for this after the child reaches adulthood, but it
is an artificiality of John's theory that the interests of the child suddenly veer from a blend

maturity.

gives reasonable answers in most cases. But I think that a Fisher-style model of fitness that weighs
each offspring by its "reproductive value" is really the best model to use.

I also am guessing that John doesn't really understand how neo-Darwinists count fitness. For
example, in a social insect with infertile castes, the fitness of a queen is not counted by
including all offspring - sterile forms and fertile forms together - the "establishment" is not
that stupid.

Quoted message said:

JM:- Those who are familiar with the philosophical foundations of economics and game theory may
worry about how a child can be said to balance the interests of both parents, giving equal weight
to each. In critiques of the ethical doctrine of utilitarianism, for example, theoretical
economists object to the idea of "equal weight". Those objections may well apply in economics,
where utility functions are freely chosen. The objections do not seem to apply in evolutionary
theory, because Nature is assigning the utility function. (PS. If this paragraph makes no sense to
you, don't worry about it. It is addressed only to people who understand the distinction between
cardinal utility theory and ordinal utility theory.)

Quoted message said:

JE:- The above is important. Please explain it in a more simple way so that sbe readers can more
easily understand it and identify what your explaining within evolutionary theory

JM:- I'm not sure exactly what you are looking for, since I am not sure that it does apply to
evolutionary theory, but I will try. Economists talk about a quantity called "utility" that has some
similarity to the quantity that evolutionary theorists call "fitness". John finds, picks, and eats
some strawberries. Economists would say that John has incrementally added to his utility.
Evolutionists, noting the nutritional value of the strawberries, would say that John has
incrementally added to his fitness. John got scratched up while picking the strawberries. Economists
would say there was a decrement to John's utility from the scratches. Evolutionists would say that
there was a decrement to John's fitness from the scratches. Which was greater, the increment from
the eating, or the decrement from the scratches? An economist considers this to be an objective
question. He answers by observing John's behavior. If John knew in advance that he would get
scratched, and fetched the strawberries anyways, then the utility (to John) of the strawberries was
greater than the disutility of the scratch. If Jim refuses to fetch strawberries under these
conditions, then the disutility must be greater (for Jim) than the utility. John and Jim appear to
disagree, but actually both are right. There are two different utility "scales" involved -
JimUtility and JohnUtility. John is the sole arbiter of JohnUtility and Jim is the sole arbiter of
JimUtility. The economic theorist has no basis for judging between the two.

JE:- If john and Jim trade then the economic theorist does have “a basis for judging between the
two” because their interests have become mutual. If John picks strawberries for Jim because Jim
provides something that JohnUtility says no to, then both John and Jim are joint arbiter’s to each
other’s utililty. If the gains both make increase in _absolute_ terms above the gains either would
have made on their own, then John has a vested interest in looking out for Jim and vice versa. This
is the basis of mutualising selection where zero OFA exists.

Quoted message said:

snip<

JM:- It is possible that John and Jim can come up with a mutually agreeable exchange rate through
trade and reciprocity, but the economists cannot compute and impose one. JimUtility is strictly
incomparable to JohnUtility. Hence, any attempt by Jim and John to ally themselves according to
Bentham's principles and both try to maximize the sum of JimUtility and JohnUtility is hopeless
unless Jim and John first agree on some kind of exchange rate. The sum cannot be computed. We are
trying to add apples and oranges.

Evolutionists, on the other hand, believe that JohnFitness and JimFitness are measured in the same
units. John gains exactly as much by raising one

JohnFitness units from JimFitness units - we can just talk about fitness - plain and simple.

JE:- Yes fitness units are eqivalent.

JM:- All of this is irrelevant for most purposes in evolutionary theory.

JE:- It is not “irrelevant” it is of extreme importance. This is because organism fitness
mutualism (OFM) has been routinely misrepresented as organism fitness altruism (OFA) within Neo
Darwinian theory.

JM:- But, there are some cases where the fitnesses of one individual must be compared with that of
another individual.

JE:- “Some cases?” EVERY case of Darwinian evolution by natural selection requires such a
comparison.

JM:- For example, if Jim is an altruistic donor of fitness, and John is the recipient, then
Hamilton's rule (rb>c) involves comparing "c" which is measured in JimFitness, with "b" which is
measured in JohnFitness. Not a problem, ..

JE:- A big problem...

Each, independent finess within b must be compared to each other and the donor, _before_ one b total
can even be compared to c (which is only being done for the benefit of Hamilton’s_hypothetical_
selfish gene). Note that b is a _proven_ group of _independent_ Darwinian organism selectees that
_are_ competing against each other at the Darwinian level of selection.

Only when the donor is innitiating the action it is “atruistic”. Here the donor’s that give nothing
away are always selected over those who give anything away. On the other side of the fence, selfish
recipients that accept the donation are selected over those that do not. When the recipient
innitiates the action, it becomes “selfish” because b has been stolen and not donated. Here
recipients who steal are favoured compared to those that don’t steal. On the other side of the
fence, donors that allow recipients to steal from them are selected against. In all cases it is,
war. For Van Valen his Red Queen theory means nature must work harder and harder just to stand
still. At the Mad Hatter’s Tea Party the Red Queen just shouts “off with his head”. OFM removes this
absurd Red Queen. The logic for “selfishness” and “altruism” is the same except the pigs have become
the farmer within Orwell’s “Animal Farm”. Selfishness and altruism are just relative sides of the
SAME logic
i.e. you cannot have one without the other. The only thing that is an absolute opposite to _both_ is
the process of mutualisation yet it is the misuse of Hamilton’s rule that has caused
researchers to all but ignore, OFM.

JM:- in evolutionary theory, as long as the units are the same - and they are. (Or at least they are
if the theorist is careful. I believe that Hamilton [1964] was careful.)

JE:- The units of fitness counted must be self consistent and eqivalent. If gene fitnesses are
supposed to contest organism fitnesses and win, then one fitness has to be converted into the
other, just to find out. Total fitness is the number of fitness units reproduced over a defined
time period by a parent selectee. To convert an organism fitnesses into a valid gene fitnesses, all
the genes replicated within an organism by mitosis and between organism generations by meisosis
must be counted and then compared. Fitness now becomes not just how many Darwinian organisms you
reproduced but also, how fat you are! One fat individual can be selected over many thin ones
reproduced if the total number of genes in the fat one is more than in all the thin ones. Of course
such a situation is just Mad Hatter nonsense. However, if Hamilton wants a real gene level of
selection then this is it...

When Hamilton counts genes over organism generations he is NOT counting genes he is only using genes
as a handy index for an organism count. This is why gene fitness is entirely dependent on the
Darwinian organism level and cannot compete against it as Hamilton stipulated. Note that b is NOT an
organism level it is a group of organisms, i.e. it is an organism _group_ level of selection.
Inclusive fitness has been misrepresented as an organism level of selection because it is a group
level of selection.

JM:- Now, back to the child that is supposed to be acting as a "somatic extension" of its parents.
How is it supposed to behave? It is not acting to maximize it's mother's fitness exactly, nor is it
acting to maximize its father's fitness exactly.

JE:- It is acting to maximise their mutual fitness where this fitness provides an absolute fitness
gain to both, i.e. compared to going it alone, they are BOTH better off but NECESSARILY EQUALLY.
Here lies the rub.

JM:- John must say that it is maximizing a 50-50 blend of the two kinds of fitness.

JE:-

independent but has become mutualised. Thus the 50-50 does NOT have to be equal in absolute terms
and each parent is free to break the association and make another (note the mayhem this causes!).

JM:- And, that is a well-defined concept in fitness theory. It would not be a well-defined concept
in utility theory. But now, here is a weakness of John's model. The child's genome is not really a
50-50 blend of the two parents.

JE:- This does not matter in the least because the mutualised fitness is way above the fitness
either parent could have achieved by just going it alone (reproducing without

Quoted message said:

snip<

JM:-

gives reasonable answers in most cases. But I think that a Fisher-style model of fitness that weighs
each offspring by its "reproductive value" is really the best model to use.

JE:- No, because it deletes genetic epistasis. This has been fudged by redefining epistasis as
additive epistasis which is zero real epistasis.

JM:- I also am guessing that John doesn't really understand how neo-Darwinists count fitness. For
example, in a social insect with infertile castes, the fitness of a queen is not counted by
including all offspring - sterile forms and fertile forms together - the "establishment" is not
that stupid.

JE:- It was that stupid. Hamilton ENDS UP, (no matter what lip service he pays to raising forms to
fertile adulthood) by just counting genes in the very next organism generation; fertile or
infertile. The group fitness count, b, is just the number of extra immature organisms that each
recipient is now able to produce using the donation and NOT how many of them are raised to fertile
adulthood.

The only reason everybody was overjoyed at Hamilton’s rule was because it was supposed to provide a
cause for the evolution of _entirely_ sterile eusocial casts by allocating an altruistic fitness to
them. THEY were most certainly NOT raised to fertile adulthood, were they! Allowing a totally
sterile cast a fitness to give away was absurd. These casts have zero INDEPENDENT fitness. Like my
washing machine, its fitness contribution was ALWAYS to the owner and never to itself... but look
out for the nano machines ;-)

Best Wishes,

John Edser Independent Researcher

PO Box 266 Church Pt NSW 2105 Australia

[email hidden]

"John Edser" <[email hidden]> wrote in message news:<[email hidden]>...

Quoted message said:

JM:- For example, if Jim is an altruistic donor of fitness, and John is the recipient, then
Hamilton's rule (rb>c) involves comparing "c" which is measured in JimFitness, with "b" which is
measured in JohnFitness. Not a problem, ..

JE:- A big problem...

Each, independent finess within b must be compared to each other and the donor, _before_ one b
total can even be compared to c (which is only being done for the benefit of
Hamilton?s_hypothetical_ selfish gene). Note that b is a _proven_ group of _independent_ Darwinian
organism selectees that _are_ competing against each other at the Darwinian level of selection.

JM:- Quibble: "b" is not a group of organisms. "b" is the fitness received by one recipient (who
is related to the donor by r). There is a total quantity - call it Btot - which can be computed by
adding rb for each recipient. The group or set of recipients certainly exists, and it may be
desirable to talk about it and give it a name. But please don't call it "b". Request for
clarification: Why does the fact that the recipients and the donor are all competing organisms
make a difference. If you are worried about the fact that there will be some mortality after the
act of altruism, but before the actors have raised their families, then this fact can be taken
into account by decreasing the estimate of b and c to take into account that some of the altruism
was futile. Whoops - maybe you need to decrease b, but you don't decrease c. Hmmm. Is this what
has you bothered?

Quoted message said:

Only when the donor is innitiating the action it is ?atruistic?. Here the donor?s that give
nothing away are always selected over those who give anything away. On the other side of the
fence, selfish recipients that accept the donation are selected over those that do not. When the
recipient innitiates the action, it becomes ?selfish? because b has been stolen and not donated.
Here recipients who steal are favoured compared to those that don?t steal. On the other side of
the fence, donors that allow recipients to steal from them are selected against. In all cases it
is, war. For Van Valen his Red Queen theory means nature must work harder and harder just to stand
still. At the Mad Hatter?s Tea Party the Red Queen just shouts ?off with his head?. OFM removes
this absurd Red Queen. The logic for ?selfishness? and ?altruism? is the same except the pigs have
become the farmer within Orwell?s ?Animal Farm?. Selfishness and altruism are just relative sides
of the SAME logic
i.e. you cannot have one without the other. The only thing that is an absolute opposite to _both_
is the process of mutualisation yet it is the misuse of Hamilton?s rule that has caused
researchers to all but ignore, OFM.

JM:- in evolutionary theory, as long as the units are the same - and they are. (Or at least they
are if the theorist is careful. I believe that Hamilton [1964] was careful.)

JE:- The units of fitness counted must be self consistent and eqivalent. If gene fitnesses are
supposed to contest organism fitnesses and win, then one fitness has to be converted into the
other, just to find out. Total fitness is the number of fitness units reproduced over a defined
time period by a parent selectee. To convert an organism fitnesses into a valid gene fitnesses,
all the genes replicated within an organism by mitosis and between organism generations by
meisosis must be counted and then compared. Fitness now becomes not just how many Darwinian
organisms you reproduced but also, how fat you are! One fat individual can be selected over many
thin ones reproduced if the total number of genes in the fat one is more than in all the thin
ones. Of course such a situation is just Mad Hatter nonsense. However, if Hamilton wants a real
gene level of selection then this is it...

When Hamilton counts genes over organism generations he is NOT counting genes he is only using
genes as a handy index for an organism count. This is why gene fitness is entirely dependent on
the Darwinian organism level and cannot compete against it as Hamilton stipulated. Note that b is
NOT an organism level it is a group of organisms, i.e. it is an organism _group_ level of
selection. Inclusive fitness has been misrepresented as an organism level of selection because it
is a group level of selection.

JM:- Yes, there is a "group" of recipients, but that does not make this "group-level" selection. As
I understand it, we have group level selection only if there is mortality and reproduction of groups
- for example, if a group of lazy ands is wiped out by an ant-eater or if a group of busy-bees gives
birth to many swarms.

I continue to be mystified why you think that Hamilton has anything to do with group selection. My
understanding is that Hamilton was welcomed by the "establishment" because it could explain altruism
without invoking a Wynn-Edwards-style group selection. Hamilton is the opposite of group selection.

Quoted message said:

Quoted message said:

snip<

JM:-

Quoted message said:

gives reasonable answers in most cases. But I think that a Fisher-style model of fitness that
weighs each offspring by its "reproductive value" is really the best model to use.

JE:- No, because it deletes genetic epistasis. This has been fudged by redefining epistasis as
additive epistasis which is zero real epistasis.

JM:- I suspect we are talking about two different aspects of Fisher. Fisher definitely did not model
real epitasis in his assignment of fitnesses to genes and in his analysis of how gene frequencies
change in response to organism-level selection. But "reproductive value" is, as far as I can tell,
independent. Here, Fisher is not looking at genes - he is simply turning organism-level fitness into
something that changes during an organism's lifetime - reaching a maximum around the onset of
fertility.

Quoted message said:

JM:- I also am guessing that John doesn't really understand how neo-Darwinists count fitness. For
example, in a social insect with infertile castes, the fitness of a queen is not counted by
including all offspring - sterile forms and fertile forms together - the "establishment" is not
that stupid.

JE:- It was that stupid. Hamilton ENDS UP, (no matter what lip service he pays to raising forms to
fertile adulthood) by just counting genes in the very next organism generation; fertile or
infertile. The group fitness count, b, is just the number of extra immature organisms that each
recipient is now able to produce using the donation and NOT how many of them are raised to fertile
adulthood.

The only reason everybody was overjoyed at Hamilton?s rule was because it was supposed to provide
a cause for the evolution of _entirely_ sterile eusocial casts by allocating an altruistic fitness
to them. THEY were most certainly NOT raised to fertile adulthood, were they! Allowing a totally
sterile cast a fitness to give away was absurd. These casts have zero INDEPENDENT fitness. Like my
washing machine, its fitness contribution was ALWAYS to the owner and never to itself... but look
out for the nano machines ;-)

JM:- Well, John, I have to admit that I have only read Fisher and Hamilton [1964]. I have not yet
read any of the later papers in which the transition is made to counting fitnesses from birth-to-
birth and to modeling parental care as just another kind of altruism to close relatives. So, maybe
you are right that they ARE stupid. I will have to find out when I read the papers. But I am
skeptical that all offspring are counted as equal, unless they are saying that the sterility is "self-
selected".

Incidentally, if there are any experts out there following this thread, I would appreciate exact
references to later Hamilton papers that I should read. I will need to provide the reference to my
neighborhood library branch, and they will send away for photocopies, so I need title, journal name,
date, and page numbers.

On Wed, 11 Feb 2004 00:46:17 +0000 (UTC), [email hidden] (Jim Menegay) wrote: ... [much
snipped] ...

Quoted message said:

I continue to be mystified why you (Edser) think that Hamilton has anything to do with group
selection. My understanding is that Hamilton was welcomed by the "establishment" because it could
explain altruism without invoking a Wynn-Edwards-style group selection. Hamilton is the opposite of
group selection.

Quoted message said:

Quoted message said:

snip<

JM:-

WH:- Maybe this is a quibble because what you say is true. But by 1972 Hamilton had corresponded
with Price and was using Price's formulation of group selection (the Price equation). Hamilton
(1975) re-derived his expression using the logic of selection within and between groups. I think if
Hamilton were still here he would now see kin selection as just a strong form of group selection
although not a Wynn-Edwards-style group selection. William L Hunt

... [much snipped] ...

Quoted message said:

I continue to be mystified why you (Edser) think that Hamilton has anything to do with group
selection. My understanding is that Hamilton was welcomed by the "establishment" because it could
explain altruism without invoking a Wynn-Edwards-style group selection. Hamilton is the opposite of
group selection.

WH:- Maybe this is a quibble because what you say is true.

JE:- It is simply a proven contradiction to suggest that "Hamilton is the opposite of group
selection".

WH:- But by 1972 Hamilton had corresponded with Price and was using Price's formulation of group
selection (the Price equation). Hamilton (1975) re-derived his expression using the logic of
selection within and between groups. I think if Hamilton were still here he would now see kin
selection as just a strong form of group selection although not a Wynn-Edwards-style group
selection.

JE:- Inclusive fitness, rb is the group fitness of a number of independent Darwinian selectee's
i.e. rb is ONE group fitness where inclusive fitness is ONE rb value that is being compared to a NON
group fitness, c.

Best Wishes,

John Edser Independent Researcher

PO Box 266 Church Pt NSW 2105 Australia

[email hidden]

"John Edser" <[email hidden]> wrote in message news:<[email hidden]>...

Quoted message said:

JM:-

Quoted message said:

I continue to be mystified why you (Edser) think that Hamilton has anything to do with group
selection. My understanding is that Hamilton was welcomed by the "establishment" because it could
explain altruism without invoking a Wynn-Edwards-style group selection. Hamilton is the opposite
of group selection.

WH:- Maybe this is a quibble because what you say is true.

JE:- It is simply a proven contradiction to suggest that "Hamilton is the opposite of group
selection".

WH:- But by 1972 Hamilton had corresponded with Price and was using Price's formulation of group
selection (the Price equation). Hamilton (1975) re-derived his expression using the logic of
selection within and between groups. I think if Hamilton were still here he would now see kin
selection as just a strong form of group selection although not a Wynn-Edwards-style group
selection.

JE:- Inclusive fitness, rb is the group fitness of a number of independent Darwinian selectee's
i.e. rb is ONE group fitness where inclusive fitness is ONE rb value that is being compared to a
NON group fitness, c.

JM:- My thanks to WH for pointing this out, and my apologies to JE for not having appreciated the
germ of truth in his position. It appears that I am going to have to dig into the literature to
understand the group selection theories of Price, and the use that was made of this in the later
publications of Hamilton. If anyone can give me some pointers into this literature, I would
appreciate it. I have only sporadic use of a good medical-school library for research. My serious
reading, so far, is limited to Hamilton's 1964 paper.

In 1964, Hamilton did not write down the formula rb>c. Nonetheless, from what he did write down, it
is easy to derive this formula. In fact, there are two different meanings that can be given to this
formula depending on how you interpret r and b.

1. You can take r and b as scalars and interpret the formula as applying to behaviors with a single
donor and a single recipient. In this case, b is the benefit to the (single) recipient and r is
the Wright coefficient of relationship between donor and recipient.

2. You can take r and b as vectors, with the multiplication being an inner product and the index
ranging over individuals within the deme (excluding the donor). One applies this formula to
behaviors of a single individual which create a benefit for many individuals. As in the first
case, each scalar r within the vector is a Wright coefficient, and each scalar b is the benefit
to a single individual.

To my mind, this cannot be called group selection, because there is no identification of a group and
there is no "group fitness". All fitnesses being considered are individual fitnesses. To be sure, in
the second part of the paper, Hamilton suggested some applications in which the demic structure of
the population might result in typical r values being higher than in an unstructured population, and
you might stretch things to call this group selection. Nonetheless, I stand by my claim that the
1964 paper of Hamilton does not involve group selection, in the strict sense.

Now, one can imagine a third possible interpretation of rb>c in which b is a scalar "group fitness"
delta and in which r is some measure of the relationship between a typical individual and the group
- a number that also somehow serves as a conversion factor between group fitnesses and individual
fitnesses. If it turns out that this is what rb>c became in modern evolutionary biology, then I
suspect that I am going to approach this literature with a skepticism of only slightly less than
Edserian proportions. I hope that my mind will remain open, though, until I see the details.

Quoted message said:

JM:- For example, if Jim is an altruistic donor of fitness, and John is the recipient, then
Hamilton's rule (rb>c) involves comparing "c" which is measured in JimFitness, with "b" which is
measured in JohnFitness. Not a problem, ..

Quoted message said:

JE:- A big problem... Each, independent finess within b must be compared to each other and the
donor, _before_ one b total can even be compared to c (which is only being done for the benefit of
Hamilton?s_hypothetical_ selfish gene). Note that b is a _proven_ group of _independent_ Darwinian
organism selectees that _are_ competing against each other at the Darwinian level of selection.

JM:- Quibble: "b" is not a group of organisms. "b" is the fitness received by one recipient (who is
related to the donor by r).

JE:- All of Hamilton's sins are hidden within b. Mostly, b is the GROUP fitness total of two or more
kin selected recipients. I can kin select two individuals related 0.5 or 10 individuals related 0.1
or just kin select myself related 1. According to Hamilton they are all "equal" inclusive fitness
counts. Only if I kin select myself is group selection avoided.

JM:- There is a total quantity - call it Btot - which can be computed by adding rb for each
recipient. The group or set of recipients certainly exists, and it may be desirable to talk about it
and give it a name. But please don't call it "b".

JE:- Yes Btot is HIDDEN within b. Selection occurs between the Darwinian selectee's within Btot
BEFORE b can be concluded and compared with c.

JM:- Request for clarification: Why does the fact that the recipients and the donor are all
competing organisms make a difference.

JE:- Because Darwinian organism fitness is a comparison of fitness at EACH AND EVERY Darwinian
organism level. If this is the first independent fitness level, which it is, then Hamilton's
_dependent_ gene level cannot be selected BEFORE Darwin does his selecting, only afterwards, making
OFA a logical impossibility. Only the 1st _independent_ fitness count can cause natural selection
for obvious logical reasons. Any 2nd independent fitness count, e.g. at the Darwinian organism group
level, can only be constituted AFTER Darwinian selection has already operated at the 1st independent
level. Thus group selection can go with but not against, organism selection. Hamilton's b value is a
group of independent fitness values so it is group selective. Like all group selection, it can only
go with and not against selection at the Darwinian organism level since if it does so they BOTH
lose. Thus OFA cannot be selected using Hamilton's logic.

Quoted message said:

snip<

Quoted message said:

Only when the donor is innitiating the action it is "atruistic:. Here the donors that give nothing
away are always selected over those who give anything away. On the other side of the fence,
selfish recipients that accept the donation are selected over those that do not. When the
recipient innitiates the action, it becomes "selfish" because b has been stolen and not donated.
Here recipients who steal are favoured compared to those that don't steal. On the other side of
the fence, donors that allow recipients to steal from them are selected against. In all cases it
is, war. For Van Valen his Red Queen theory means nature must work harder and harder just to stand
still. At the Mad Hatter's Tea Party the Red Queen just shouts "off with his head". OFM removes
this absurd Red Queen. The logic for "selfishness" and "altruism" is the same except the pigs have
become the farmer within Orwell's "Animal Farm". Selfishness and altruism are just relative sides
of the SAME logic
i.e. you cannot have one without the other. The only thing that is an absolute opposite to _both_
is the process of mutualisation yet it is the misuse of Hamilton's rule that has caused
researchers to all but ignore, OFM.

Quoted message said:

JM:- in evolutionary theory, as long as the units are the same - and they are. (Or at least they
are if the theorist is careful. I believe that Hamilton [1964] was careful.)

Quoted message said:

JE:- The units of fitness counted must be self consistent and equivalent. If gene fitnesses are
supposed to contest organism fitnesses and win, then one fitness has to be converted into the
other, just to find out. Total fitness is the number of fitness units reproduced over a defined
time period by a parent selectee. To convert an organism fitnesses into a valid gene fitnesses,
all the genes replicated within an organism by mitosis and between organism generations by meiosis
must be counted and then compared. Fitness now becomes not just how many Darwinian organisms you
reproduced but also, how fat you are! One fat individual can be selected over many thin ones
reproduced if the total number of genes in the fat one is more than in all the thin ones. Of
course such a situation is just Mad Hatter nonsense. However, if Hamilton wants a real gene level
of selection then this is it... When Hamilton counts genes over organism generations he is NOT
counting genes he is only using genes as a handy index for an organism count. This is why gene
fitness is entirely dependent on the Darwinian organism level and cannot compete against it as
Hamilton stipulated. Note that b is NOT an organism level it is a group of organisms, i.e. it is
an organism _group_ level of selection. Inclusive fitness has been misrepresented as an organism
level of selection because it is a group level of selection.

JM:- Yes, there is a "group" of recipients, but that does not make this "group-level" selection. As
I understand it, we have group level selection only if there is mortality and reproduction of groups
- for example, if a group of lazy ands is wiped out by an ant-eater or if a group of busy-bees gives
birth to many swarms.

JE:- Strictly speaking one group fitness is the number of groups that a group selectee reproduces.
However, classical group selection never defined it that way. Neo Darwinians just compared group
sizes or group mortalities and defined this as group selection. In your example, differential
mortality of groups is just classical group selection. The irony is that this type of group
selection is actually selection at the Darwinian organism level and NOT group selection, at all. Which-ever-
way you define group selection, it fails to compete and win against the unit that is grouped, if and
only if, each unit within remains an independent selectee. Independence is allowed by the term
"additive",
i.e. independent fitness counts are "additive" but dependent fitness counts are "non additive". I
will add this is is because additive events only employ standard set processing (intersection
and set union) but multiplicative events can be expressed as more complex, nested sets.

JM:- I continue to be mystified why you think that Hamilton has anything to do with group selection.
My understanding is that Hamilton was welcomed by the "establishment" because it could explain
altruism without invoking a Wynn-Edwards-style group selection. Hamilton is the opposite of group
selection.

JE:- Yes, Hamilton was supposed to be "the opposite of group selection". Neo Darwinians wanted
another way to explain OFA other than group selection. They thought that selection at the lower gene
level instead at the higher organism group level could do it. As it turned out, selection at the
lower gene level had to employ selection at the higher organism group level, to work. All that
happened was Hamilton's group selection remained cleverly hidden within b for over 50 years.

Quoted message said:

Quoted message said:

snip<

Quoted message said:

JM:-

Quoted message said:

gives reasonable answers in most cases. But I think that a Fisher-style model of fitness that
weighs each offspring by its "reproductive value" is really the best model to use.

Quoted message said:

JE:- No, because it deletes genetic epistasis. This has been fudged by redefining epistasis as
additive epistasis which is zero real epistasis.

JM:- I suspect we are talking about two different aspects of Fisher. Fisher definitely did not model
real epitasis in his assignment of fitnesses to genes and in his analysis of how gene frequencies
change in response to organism-level selection.

JE:- You have to remove ALL real gene fitness epistasis to allow genes to be independently
selectable so Fisher just deleted all of it within his model.

JM:- But "reproductive value" is, as far as I can tell, independent. Here, Fisher is not looking at
genes - he is simply turning organism-level fitness into something that changes during an organism's
lifetime - reaching a maximum around the onset of fertility.

JE:- Yes, but the stage was set for Hamilton to reverse cause an affect and explicitly say, "hey its
selection at the gene level that was happening within Fisher", because it WAS. When real gene
fitness epistasis is deleted then YES, selection can happen at an independent gene level. Hamilton
just "forgot" the Fisher's view was only an oversimplified model so he misused it. Everybody since
has compounded more and more misuse.

Quoted message said:

JM:- I also am guessing that John doesn't really understand how neo-Darwinists count fitness. For
example, in a social insect with infertile castes, the fitness of a queen is not counted by
including all offspring - sterile

forms

Quoted message said:

and fertile forms together - the "establishment" is not that stupid.

Quoted message said:

JE:- It was that stupid. Hamilton ENDS UP, (no matter what lip service he pays to raising forms to
fertile adulthood) by just counting genes in the very next organism generation; fertile or
infertile. The group fitness count, b, is just the number of extra immature organisms that each
recipient is now able to produce using the donation and NOT how many of them are raised to fertile
adulthood. The only reason everybody was overjoyed at Hamilton's rule was because it was supposed
to provide a cause for the evolution of _entirely_ sterile eusocial casts by allocating an
altruistic fitness to them. THEY were most certainly NOT raised to fertile adulthood, were they!
Allowing a totally sterile cast a fitness to give away was absurd. These casts have zero
INDEPENDENT fitness. Like my washing machine, its fitness contribution was ALWAYS to the owner and
never to itself... but look out for the nano machines ;-)

JM:- Well, John, I have to admit that I have only read Fisher and Hamilton [1964]. I have not yet
read any of the later papers in which the transition is made to counting fitnesses from birth-to-
birth and to modelling parental care as just another kind of altruism to close relatives. So, maybe
you are right that they ARE stupid. I will have to find out when I read the papers. But I am
skeptical that all offspring are counted as equal, unless they are saying that the sterility is "self-
selected".

JE:- Hamilton misused Fisher's model and everybody was happy to jump onto the bandwagon of model
misuse. Now the chickens have come home to roost.

Best Wishes,

John Edser Independent Researcher

PO Box 266 Church Pt NSW 2105

[email hidden]

Quoted message said:

JM:-

Quoted message said:

I continue to be mystified why you (Edser) think that Hamilton has

anything

Quoted message said:

Quoted message said:

to do with group selection. My understanding is that Hamilton was welcomed by the "establishment"
because it could explain altruism without invoking a Wynn-Edwards-style group selection. Hamilton
is the opposite of group selection.

Quoted message said:

WH:- Maybe this is a quibble because what you say is true.

Quoted message said:

JE:- It is simply a proven contradiction to suggest that "Hamilton is the opposite of group
selection".

Quoted message said:

WH:- But by 1972 Hamilton had corresponded with Price and was using Price's formulation of group
selection (the Price equation). Hamilton (1975) re-derived his expression using the logic of
selection within and between groups. I think if Hamilton were still here he would now see kin
selection as just a strong form of group selection although not a Wynn-Edwards-style group
selection.

Quoted message said:

JE:- Inclusive fitness, rb is the group fitness of a number of independent Darwinian selectee's
i.e. rb is ONE group fitness where inclusive fitness is ONE rb value that is being compared to a
NON group fitness, c.

JM:- My thanks to WH for pointing this out, and my apologies to JE for not having appreciated the
germ of truth in his position. It appears that I am going to have to dig into the literature to
understand the group selection theories of Price, and the use that was made of this in the later
publications of Hamilton. If anyone can give me some pointers into this literature, I would
appreciate it. I have only sporadic use of a good medical-school library for research. My serious
reading, so far, is limited to Hamilton's 1964 paper.

In 1964, Hamilton did not write down the formula rb>c. Nonetheless, from what he did write down, it
is easy to derive this formula. In fact, there are two different meanings that can be given to this
formula depending on how you interpret r and b.

1. You can take r and b as scalars and interpret the formula as applying to behaviors with a single
donor and a single recipient. In this case, b is the benefit to the (single) recipient and r is
the Wright coefficient of relationship between donor and recipient.

2. You can take r and b as vectors, with the multiplication being an inner product and the index
ranging over individuals within the deme (excluding the donor). One applies this formula to
behaviors of a single individual which create a benefit for many individuals. As in the first
case, each scalar r within the vector is a Wright coefficient, and each scalar b is the benefit
to a single individual.

To my mind, this cannot be called group selection, because there is no identification of a group and
there is no "group fitness". All fitnesses being considered are individual fitnesses. To be sure, in
the second part of the paper, Hamilton suggested some applications in which the demic structure of
the population might result in typical r values being higher than in an unstructured population, and
you might stretch things to call this group selection. Nonetheless, I stand by my claim that the
1964 paper of Hamilton does not involve group selection, in the strict sense.

Now, one can imagine a third possible interpretation of rb>c in which b is a scalar "group fitness"
delta and in which r is some measure of the relationship between a typical individual and the group
- a number that also somehow serves as a conversion factor between group fitnesses and individual
fitnesses. If it turns out that this is what rb>c became in modern evolutionary biology, then I
suspect that I am going to approach this literature with a skepticism of only slightly less than
Edserian proportions. I hope that my mind will remain open, though, until I see the details.

JE:- It seems to me that b is EITHER a set of independent Darwinian individuals or it isn't, it
can't be both! Naturalists I have spoken with who mostly ignore Neo Darwinism because it is
"unintelligible" and then blame themselves because "they don't really understand the maths" scratch
their heads in disbelief at how a gene that _lowers_ the total fitness of each Donor parent by just
giving it way to a _group_ of Darwinian competitors because they are related in some way, can ever
be selected _for_. They are right, it can't! Hamilton's rule is just a misused model that deleted
the total fitness of the donor from the model allowing ALL FITNESSES to become JUST relative. This
means the goal posts for selection can be shifted to anywhere you like. Prof. Felsenstein is correct
when he suggested all Neo Darwinian models are "non testable". Hamilton's model is non testable
because all fitnesses remain relative. The "everything is relative" school of epistemology is called
Post Modernism. Note that Darwinism is NOT post modern. Implied within it is each parents total
fitness within one Darwinian population.

______________________________________________
Until a full and frank discussion re: the proper relationship between models and the theories they
are just a simplification from eventuates, the problem of simplified model misuse will continue to
destroy evolutionary theory from the inside out, like a cancer.
_______________________________________________

Best Wishes,

John Edser Independent Researcher

PO Box 266 Church Pt NSW 2105 Australia

[email hidden]

"Jim Menegay" <[email hidden]> wrote in message
"]news:[email hidden]...

Quoted message said:

"John Edser" <[email hidden]> wrote in message

news:<[email hidden]>...

Although you are superbly analytical (and very subtly so), and admirably patient!

Here is my take on how to cut through the core of what is being considered:

Given a population that across generations undergo somewhat incremental but sometimes phase
transitioning (significantly phenotype-altering) genomic changes:

Failure / Gene ->Behaviour-> Reproductive Benefit? \

Success

\

Gene->Behaviour->Reproductive Benefit? (a.s.f)

In the above diagram the only really interesting component is the "?".

?=Any actual (and relevant) outcome of the "Evolutionary Pressure Totality".

You have not embarked on a 'scientifically (or philosophically) remaining-to-take' -- i.e., not
already thoroughly tried, tested/contested, and sufficiently collectively conceptually crystalized
-- non-knit-picking general evolution theoretical approach to explaining and understanding how we
came to be how we are.

What you are doing is to re-analyse evolution theoretical concepts inorder to understand how
someone's psyche has quirks to the effect of that he refuses to absorb and accept the fact that
altruistic acts of individuals need not be "mutualistic" to have become selected-in (and so to
become an as if 'genetically conserved behavioural potential' within a population).

In reality this is a subject best to be theoretically tackled (analyses) from various angels of
Psychobiology (including with evolutionary psychology type thinking).

In contrast to your effort, a relatively new or unexplored and untapped possibility to explain how
we are and evolved is one that takes the tack of defining and categorizing amongst different
phylogenetically significant selective or evolutionary pressures (against a backround the E.P.T.).

Whilst being as generalizing (in parts) as it is, this complementary analytical approach is of
course rather more individual-centered than most (other 'evolution philosophical/theoretical'
approaches).

As a complementary approach or aspect of evolution theory, "E.P.T. thinking" does of course
accommodate the possibility that, amongst several individual "candidates for reproductive success"
within a population, the _least_ flexibly or well adapted of these may yet be fit enough to
reproduce if it is selected _not_ by competition with other candidates but because pure happenstance
caused all other candidates to 'crumble'.

This aspects of reality is easily accommodated because such happenstances do not generate a
genophenotyping "strange attraction" within the phylogenetic patterning process.

Recognitions of such "attractors" in the process of phylogeny is what EPT is about.

The EPT approach within anthropobiology is to consider how we are against a back-drop of 'EPTly'
theorized and defined types of phylogenetic (evolutionary) pressures on (and in different respects
also *within*) "individual candidates for reproductive success". (Including pressures on
"prematures"; and even on infertiles to the extent these behave altruistically because of genes
shared with more or less close relatives - the survivial of which is altruistically promoted.)

The schematic scenario of "selective pressures" operating on individuals within phylogeny is however
especially relevant to the phylogeny of fauna, because the EPT approach is, ultimately, most of all
a philanthropically oriented overview.

[BTW, EPT is a SEPTIC-humored approach at seriously understanding What Is going on.]

P

"Peter F." <[email hidden]> wrote in message news:<[email hidden]>...

Quoted message said:

[snip] You have not embarked on a 'scientifically (or philosophically) remaining-to-take' -- i.e.,
not already thoroughly tried, tested/contested, and sufficiently collectively conceptually
crystalized -- non-knit-picking general evolution theoretical approach to explaining and
understanding how we came to be how we are.

What you are doing is to re-analyse evolution theoretical concepts inorder to understand how
someone's psyche has quirks to the effect of that he refuses to absorb and accept the fact that
altruistic acts of individuals need not be "mutualistic" to have become selected-in (and so to
become an as if 'genetically conserved behavioural potential' within a population).

In reality this is a subject best to be theoretically tackled (analyses) from various angels of
Psychobiology (including with evolutionary psychology type thinking).

In contrast to your effort, a relatively new or unexplored and untapped possibility to explain
how we are and evolved is one that takes the tack of defining and categorizing amongst different
phylogenetically significant selective or evolutionary pressures (against a backround the
E.P.T.). [snip]

As always, Peter, I am not sure I understand you. But it appears that you are wrong about what I am
doing. I am definitely NOT seeking to "understand how someone's psyche has quirks" nor am I
interested in "explaining and understanding how we came to be how we are." Human psychology is your
area of interest - not mine.

What am I interested in? Theoretical concepts. Other people have them - I want them. I only learn
and use enough psychology to let me communicate efficiently - I am not particularly interested in
why communicating efficiently is more difficult than it "ought" to be. Your interests are different
than mine. Vive le difference.

"John Edser" <[email hidden]> wrote in message news:<[email hidden]>...

Quoted message said:

Naturalists I have spoken with who mostly ignore Neo Darwinism because it is "unintelligible" and
then blame themselves because "they don't really understand the maths" scratch their heads in
disbelief at how a gene that _lowers_ the total fitness of each Donor parent by just giving it way
to a _group_ of Darwinian competitors because they are related in some way, can ever be selected
_for_. They are right, it can't!

Let me attempt an explanation of Hamilton's [1964] logic without math. The hypothetical successful
gene for altruism lowers the total fitness of its carrier (the donor) in the following sense: A
carrier of the gene (an altruist) has lower fitness than a non-carrier (a non-altruist) who lives in
the same deme. On the other hand, the hypothetical gene INCREASES the fitness of its carrier in the
following sense: A typical carrier of the gene has higher fitness than the typical non-carrier
within the population. And, solely for this reason (higher average fitness within the population)
the gene will prosper.

How is this possible? Well, it is possible if the typical carrier of the gene (within the whole
population) is a recipient more often than is a typical non-carrier (within the whole population).
Why are carriers "luckier" than non-carriers? Because the carrier probably inherited the gene, and
this indicates that there are probably a higher-than-average number of other altruists in his family
and in his neighborhood. The gene brings with it a burden of being altruistic, but it also brings
the blessing of helpful relatives and neighbors. The gene doesn't "cause" the blessing, but it does
tend to come with the blessing.

Consider four kinds of individuals -
(A) non-altruists that live in mostly non-altruistic demes,
(B) non-altruists that live in mostly altruistic demes,
(C) altruists that live in mostly non-altruistic demes,
(D) altruists that live in mostly altruistic demes.

Of these four kinds of individuals, the most fit are (B), next is (D), next is (A), and last is (C).
Altruists really are less fit than similarly situated non-altruists; the cost of altruism is real.
But, it turns out that
(E) individuals are more common than (C) and (A) is more common than (B). So, the "average" altruist
in the population turns out to be more fit (!) than the "average" non-altruist. Natural
selection - at the population level - causes the altruistic gene to thrive.

Now, when Hamilton's [1964] argument is expressed in this way, it is easy to see how someone (i.e.
you) might react in horror with cries of "Group selection! Group selection! Oh, the horror!!" But I
think that this horror is misguided, in this case. Only if (D) individuals compete for resources
more rigorously against (B) individuals in their own deme than they do against individuals in other
demes, will there be problems.

Now, you can claim that if the higher-than-population-average fitness of the typical altruist is
true, and especially since it results from higher than average receipts of altruism, then "altruism"
is the wrong work and it should be called "mutualism". I suppose that you are free to use words
however you wish, except:
1. If you use a word in a non-standard way, you may fail to communicate.
2. You cannot logically accuse Hamilton of error if you redefine the words he used so that what he
said no longer makes sense under your new meanings.

Hamilton [1964] never claimed that altruism could spread in the face of Darwin's natural selection.
What he did was simply to point out that in some social situations, your fitness must include
aspects not only of "what you do", but also of "who you are related to". Altruists tend to be
related to some very helpful relatives. And, this is a heritable characteristic.

There is another curious aspect to all this. It is a standard analytical technique in evolutionary
thinking to ask "Would it be to this organism's advantage to acquire this gene by mutation?" But
this technique turns out to be invalid in social situations. The fitness of an individual who
acquires a gene by mutation may be very different from the fitness of an individual who acquires the
gene by inheritance. If you acquire one of Hamilton's genes for altruism by mutation, then all you
acquire are some costs. But, if you acquire the gene by inheritance, then along with the cost come a
bunch of very helpful relatives that can increase your fitness by more than the cost. Hamilton's
logic only work if most organisms acquire their genes by inheritance, rather than by mutation. My
understanding of the biology is that this assumption is valid.

One final point: You may ask why I claim that (D) individuals are more fit than (A). Good question.
It is not true for an arbitrary altruistic gene, but it is true for the ones that satisfy "rb>c".
But proving that would take math.

"John Edser" <[email hidden]> wrote in message
news:<[email hidden]>...

Quoted message said:

Naturalists I have spoken with who mostly ignore Neo Darwinism because it is "unintelligible" and
then blame themselves because "they don't really understand the maths" scratch their heads in
disbelief at how a gene that _lowers_ the total fitness of each Donor parent by just giving it way
to a _group_ of Darwinian competitors because they are related in some way, can ever be selected
_for_. They are right, it can't!

JM:- Let me attempt an explanation of Hamilton's [1964] logic without math. The hypothetical
successful gene for altruism lowers the total fitness of its carrier (the donor) in the following
sense: A carrier of the gene (an altruist) has lower fitness than a non-carrier (a non-altruist) who
lives in the same deme.

JE:- This is where the argument ends and not starts ;-) Hamilton starts his argument at the end and
ends it at the start; a neat trick. Of course this was made possible because of the reversal of
cause and affect within his massively simplified model where cmax was deleted.

JM:- On the other hand, the hypothetical gene INCREASES the fitness of its carrier in the following
sense: A typical carrier of the gene has higher fitness than the typical non-carrier within the
population. And, solely for this reason (higher average fitness within the population) the gene
will prosper.

JE:- If the fitness of the donor absolutely decreases then it does not matter if it appears to
relatively increase as just a mean value. A relative measure can indicate the gene may spread
relative to a non altruist gene but the absolute value loss means that it cannot be selected to do
so. Thus it is an argument over what must win logically? The absolute donor fitness decrease that is
not selectable or Hamilton's hypothetical relative mean fitness increase for the donor?

JM:- How is this possible? Well, it is possible if the typical carrier of the gene (within the whole
population) is a recipient more often than is a typical non-carrier (within the whole population).

JE:- But donor loss exactly equals recipient gain because altruism can only transfer fitness, it
cannot create any of it.

JM:-

Why are carriers "luckier" than non-carriers? Because the carrier probably inherited the gene, and
this indicates that there are probably a higher-than-average number of other altruists in his family
and in his neighborhood. The gene brings with it a burden of being altruistic, but it also brings
the blessing of helpful relatives and neighbors. The gene doesn't "cause" the blessing, but it does
tend to come with the blessing.

Consider four kinds of individuals -
(A) non-altruists that live in mostly non-altruistic demes,
(B) non-altruists that live in mostly altruistic demes,
(C) altruists that live in mostly non-altruistic demes,
(D) altruists that live in mostly altruistic demes.

Of these four kinds of individuals, the most fit are (B), next is (D), next is (A), and last is (C).
Altruists really are less fit than similarly situated non-altruists; the cost of altruism is real.
But, it turns out that
(E) individuals are more common than (C) and (A) is more common than (B). So, the "average" altruist
in the population turns out to be more fit (!) than the "average" non-altruist. Natural
selection - at the population level - causes the altruistic gene to thrive.

JE:- Hamilton's mythical increase in relative mean fitness for the carrier is ye olde group
selection. Hamilton saved altruism from group selection by only hiding it away within
F. Nobody noticed so who cares?

JM:- Now, when Hamilton's [1964] argument is expressed in this way, it is easy to see how someone
(i.e. you) might react in horror with cries of "Group selection! Group selection! Oh, the horror!!"
But I think that this horror is misguided, in this case. Only if (D) individuals compete for
resources more rigorously against (B) individuals in their own deme than they do against individuals
in other demes, will there be problems.

JE:- Either b is a group of independent selectees competing against one selectee c costs, or it
isn't! If it is, then rb is just classically group selective and fails because selection at the
lower individual level used in c must occur, _firstly_.

JM:- Now, you can claim that if the higher-than-population-average fitness of the typical altruist
is true, and especially since it results from higher than average receipts of altruism, then
"altruism" is the wrong work and it should be called "mutualism".

JE:- No, I claim that altruism is a selected reduction in absolute donor fitness but mutualism is a
selected increase. The mean value tells you nothing about the absolute fitness of each independent
parental absolute fitness component used to calculate it. Before such a mean can even be calculated,
selection between these diverse absolute fitness components is over and done with. This is why any
mean fitness is a group selective measure of any independent fitnesses that are being averaged when
used to form a competing fitness.

Quoted message said:

snip<

JM:- Hamilton [1964] never claimed that altruism could spread in the face of Darwin's natural
selection.

JE:- He simply failed to include any discussion of total parental fitness so he just evaded this
most _basic_ of issues.

JM:- What he did was simply to point out that in some social situations, your fitness must include
aspects not only of "what you do", but also of "who you are related to". Altruists tend to be
related to some very helpful relatives. And, this is a heritable characteristic.

JE:- Hamilton failed to discriminate between helping relatives as an investment cost for a _gain_ in
absolute donor fitness (OFM) or the exact opposite (OFA), because he deleted any reference to total
parental fitness. Hamilton's skeleton key that could open _any_ level of fitness was just the
smashing of Darwin's single lock that allowed Darwin's view to remain testable: total parental
fitness. Hamilton's free "unlocked" view remains arbitrary and thus, non testable.

JM:- There is another curious aspect to all this. It is a standard analytical technique in
evolutionary thinking to ask "Would it be to this organism's advantage to acquire this gene by
mutation?" But this technique turns out to be invalid in social situations. The fitness of an
individual who acquires a gene by mutation may be very different from the fitness of an individual
who acquires the gene by inheritance. If you acquire one of Hamilton's genes for altruism by
mutation, then all you acquire are some costs. But, if you acquire the gene by inheritance, then
along with the cost come a bunch of very helpful relatives that can increase your fitness by more
than the cost. Hamilton's logic only work if most organisms acquire their genes by inheritance,
rather than by mutation. My understanding of the biology is that this assumption is valid.

JE:- Yes, Hamilton's relative gain at an absolute cost cannot even start.

JM:- One final point: You may ask why I claim that (D) individuals are more fit than (A). Good
question. It is not true for an arbitrary altruistic gene, but it is true for the ones that satisfy
"rb>c". But proving that would take math.

JE:- None of them can start. What is needed to start any of them is OFM. OFA and OFS can only exist
when OFM is greater than either of them.

Respectfully,

John Edser Independent Researcher

PO Box 266 Church Pt NSW 2105 Australia

[email hidden]

Jim Menegay wrote: <snip>

Quoted message said:

JM:- My thanks to WH for pointing this out, and my apologies to JE for not having appreciated the
germ of truth in his position. It appears that I am going to have to dig into the literature to
understand the group selection theories of Price, and the use that was made of this in the later
publications of Hamilton. If anyone can give me some pointers into this literature, I would
appreciate it. I have only sporadic use of a good medical-school library for research. My serious
reading, so far, is limited to Hamilton's 1964 paper.

For the science, the volume edited by Laurent Keller (1999) "Levels of Selection in Evolution", is a
good overview of the current state of the art in this area. If you want to know more of the history
of these disputes, then "Defenders of the Truth: The Sociobiology Debate" by Ullica Segerstr†le is a
good read.

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: rni.helsinki.fi~bohOpen ↗ Journal
of Negative Results - EEB: jnr-eeb.orgjnr-eeb.orgOpen ↗

"Jim Menegay" <[email hidden]> wrote in message
"]news:[email hidden]...

Quoted message said:

As always, Peter, I am not sure I understand you. But it appears that you are wrong about what I
am doing. I am definitely NOT seeking to "understand how someone's psyche has quirks"

I tried to point out that - whilst you might well have great fun and clearly show me and others that
you are analytically clever and intelligent - there is ultimately not much substantial to be
understood about Edser's theoretical concepts than the very important and substantial
psychobiological causes behind them.

It is perfectly fine - and you express it perfectly fine too - that you don't have any interest in,
or inclination, to absorb anything of what I am *trying* to communicate. The reality of that it is
so is something entirely for me to deal with.

Celebrating our differences is, I have to say, a lot more easy for me to do when someone who is so
different to me, as you are, is also such an apparently decent and high-quality a person.

If that implies 'a boomerangs turning back towards my own head', then I suppose I just have
to duck. ;->

P