"John Edser" <[email hidden]> wrote in message news:<[email hidden]>...
Quoted message said:JM:- Fisher's "fundamental theorem" says that the
genetical component of fitness is non-decreasing.JE:- This is consistent with the Darwinian proposition
that each parents absolute fitness (the _total_ number of
_fertile_ forms reproduced by _each_ parent into _one_
population) cannot be selected to be reduced, i.e. at all
times parents must maximise the total number of fertile
forms they reproduce into one population.JM:- But he avoids having continuously increasing
populations by claiming that the environment is
continually deteriorating. Only recently have theorists
realized that Fisher was including the average fitness of
competing conspecifics as a component of the environment.
So, we have a "Red Queen" race - fitness continually
increases, but number of offspring do not.JE:- Most populations have been observed to expand and
then become more stable. Populations that only tend to
decrease are on the road to extinction.
JM:- A minor quibble: extinction is inevitable only if the
trend of population decrease persists. But populations have
also sometimes been observed to contract and then become
more stable. This frequently happens after an environmental
change. At the risk of transforming the minor quibble into a
major one, it could also conceivably happen after a genetic
innovation inside the species.
For example, a large population of small shrubs could
transform itself into a smaller population of taller,
broader quasi-trees. (Why has so little been written about
the sociobiology of plants? In theory, they should play a
mean game of iterated prisoner's dilemma - they are stuck in
long term relationships with neighbors and should have no
trouble remembering who defected. And they can retaliate, if
necessary by strangling encroaching branches.)
Quoted message said:
JE:- If each parent must maximise the total number of
fertile forms it reproduces into one population (_not_
just one genomic genes count which can be indicated by the
presence of that gene in _either_ immature sterile forms
or fertile forms) populations should keep expanding until
they reach a stable fluctuation. These are indicated by r
and K points respectively, observable within most
population curves. Thus r selection operates during a
geometric increase well below the carrying capacity of the
environment but K selection must operate at an
environmental ceiling capacity. It is here that a
population can degrade its environment, and it is here
that all the trouble starts. Can _individuals_ be selected
to limit their numbers at this point to avoid destruction
of their environment? Is such an event an altruistic act?
Is group selection or Hamiltonian logic required to
maintain an environment?
JM:- At this point in your analysis, I would like to repeat
my recommendation of the following paper: "Fisher's
fundamental theorem of natural selection", Steve Frank and
M. Slatkin, 1992 stevefrank.org92TREEOpen ↗
R.html They list two different K-selection strategies
available at this point. (They overlook the third one
that you are going to suggest.) Their two strategies are
(1) grab a bigger piece of the pie, thus reducing K for
everyone else and reducing the population, or (2) use
your piece of the pie more efficiently, increasing your
fitness without hurting anyone else - until the next
generation when the population creeps up and everyone has
to accept a smaller piece.
Quoted message said:JE:- I contend that organism fitness mutualism (OFM) can
select Darwinian individuals to mutualise their
reproductive output in concert with all the other life
forms that form its environment requiring no altruism and
no group selection.Van Valen's Red Queen hypothesis supposes that a war
exists within nature between an expanding population and
all the other living forms that constitute its
environment. This produces the effect that all the species
concerned have to "run harder and harder just to stand
still", i.e. in order for every parent to produce an
obligatory absolute fitness increase it has to face
_increasing_ offence/defence costs. Such a solution is not
sustainable. OFM enters this situation to decrease these
costs substantially but not necessarily equally, for all
concerned. This will include a measure of "the average
fitness of competing conspecifics as a component" but such
an average hides _all_ the selective events that are
occurring at the Darwinian fertile organism level of
selection. Van Valen's view is organism fitness selfish
(OFS). Logically such a view is the same as organism
fitness altruistic arguments (OFA) because what is taken
by OFS produces an OFA deficit. What is being ignored is
organism fitness mutualism (OFM) which stands in
contradiction to both OFA and OFS. It is OFM that
dominates nature and it is the deceptively simple logic of
OFM that has been neglected within Neo Darwinian reasoning
because OFA has dominated for over 50 years.
JM:- I pretty much agree with this, except that I would add
the following points:
1. True OFM is a relationship between two, or a few,
organisms - not a relationship between an organism and
its species.
2. Mutualistic associations protect themselves against
deadbeats by enforcement behaviors; at minimum the
deadbeat is thrown out of the association.
3. True OFM relationships can exist between organisms in
different species, but the restriction to direct organism-to-
organism relationships still holds.
4. Insect eusociality is not true OFM, nor is a behavior
that benefits self and just "accidentally" happens to
benefit other members of the species (c<0, b>0). True OFM
requires reciprocity.
5. Examples of true OFM are pair bonding, "friendship",
hunting bands, lichens, mitochondria and their hosts, and
even ritualized territorial displays between songbirds.
(Sometimes, the best that OFM can do with a finite pie is
to avoid wasting time dividing it!)
6. A hypothesis of true OFM is refuted if repeated defection
does not elicit enforcement behaviors. In that case, we
have to consider the hypothesis that that sucker really
IS altruistic.