Re: The status of Infertile forms ( was: What is the evolutionary



Guy A Hoelzer hoelzer@xxxxxxx wrote:-


JE:-
But this remains the case on just logical grounds (see below).

My hesitation in agreement was based on examples
like mules, where the offspring have nothing to do with the parents'
reproductive strategies beyond possibly being a failed attempt to
create a
reproductive offspring. Sometimes infertile offspring do not remain an
integrated part of a fertile form, which is why I did not fully agree
with
your statement that they "must" remain so.

JE:
As previously indicated my "must" remains a proposition of logic which
confines choices to just two alternatives but does not force either choice.
If a sterile organism such as a mule cannot be selected to assist a fertile
form (preferably its own parents) for _whatever reason_, then its own
independent fitness must remain zero. There are only two choices: be
selected to be reduced to just a fitness sub part (here the sterile form
assumes a dependent nested fitness within a fertile form) or remain a
fitness nothing as far as nature is concerned. I agree that a sterile form
such as mule may not be able to be selected to assist a fertile form (if it
could do so the infertile form obtains for itself a dependent nested fitness
which is better than just a zero independent fitness). However, this does
not change these two choices: remain independent with only a zero fitness or
be reduced to a dependent part of some other fertile form. It seems the only
possible way nature can select for a nested fitness is via hormones
requiring an enclosed space, e.g. sterile eusocials.

JE:-
The "must" I used is only based on logic: genes within an infertile form
can
only possess a zero fitness relative to an infertile body or any meta
level
above this except for a fertile body no matter how well adapted they
allow a
meta infertile form to become and irrespective of how much longer these
genes increase the lifespan of an infertile form.

This is IMHO badly flawed logic. It seems to be based on the false
assumption that every organism must have or contribute to reproductive
fitness in some way. You conclude that if they aren't going to reproduce
themselves, they must be play a role in another's (the parents) fitness.
This is simply not true. I already gave you a concrete example that
demonstrates the falsehood of your logic. Infertile mules rarely play any
role in the reproductive success of their parents.

JE:-
The fact "infertile mules rarely play any role in the reproductive success
of their parents: does not exclude the logic that I have detailed. Please
review the argument presented above. To refute it all that is required is
the provision of a 3rd alternative for the sterile form. To recap: I reason
that only two logical alternatives can empirically exist: a dependent nested
fitness within a fertile form or fitness independency which only provides a
zero fitness.

You could explicitly
assume that they do for the sake of a model, but that doesn't make it so.

JE:-
I was not assuming a simplified/oversimplified model. What I carefully
articulated above represented an empirically refutable theory contingent on
my definition of fitness which can empirically and objectively separate
fitness dependency from fitness independency providing the required rigor to
underwrite the basic organism concept on which the science of biology
remains based.


snip<

JE:-
Why did you exclude selection OF the group in which each group
fitness is
just the sum of the fitness of each fertile individual which
comprises any
one group?


Because under this description there is no evidence that the group
actually
exists as a possible unit of selection.

JE:-
But you failed in the past and have failed here to explain WHY this is the
case. Your agreement that "under this description there is no evidence that
the group actually exists as a possible unit of selection" without providing
any explanation as to why this is the case only confirms a belief. Surely
you realize that since you rejected my argument as to why, the validity of
your agreement as to why remains contingent on you providing an alternative
theory with worked examples (which you have not provided).

The theory and the worked examples as to why that that I have provided here
(which you reject out of hand) remain empirically testable: ANY additive
fitness _proves fitness independence_ thereby refuting fitness dependence
(and vice versa). Here is my explanation as to why: addition only employs a
reversible set intersection whereas a NON additive fitness must assume a non
reversible set nesting which whenever it becomes reversed provides a
required refutation.

JE:-
But I have been posting the same argument to sbe for nearly 5 years
now.
Previously you rejected it (I think the closest was about 3 months
ago).

I have always argued that fitnesses at different levels of selection
must be
independent, so I'm sure that I never posted anything consistent with
the
claim that group selection can occur when group fitness was merely the
sum of
its individual fitnesses. I have consistently argued against this idea
over
the years on sbe and I challenge you to find anything I have posted
contrary
to this.

Example:
To: Sci.bio.evolution moderation account <sbe@xxxxxxxxxxxxxxxxxxx>
Date: Jan 29 2006:

------------------------------ repost ---------------------------------

Guy Hoelzer hoelzer@xxxxxxx wrote Jan 29 2006

I agree that every level of selection, such as the individual
organisms,
must have a phenotype (including the trait of fitness) that is a non-
additive result of single gene influences.

JE:-
Do you agree that "the trait of fitness" as measured at the gene level
is
always dependent on another level and is therefore not an independent
fitness?

No. I have tried to convey to you that I do not see
dependence/independence
as a black and white issue.


JE:-
You did _convey_ this. The problem was is it made no sense to me. The
reason
why is this: If you allow "dependence/independence" to be complimentary
and
not contradictory (it cannot be both) it becomes impossible to
distinguish
mono-centricity from poly-centricity.

Thank you for acknowledging that I did convey this and have been
consistent.

JE:-
Your consistency only appears to me to be that you prefer sitting on the
fence resolving nothing.

I do not feel any obligation to pretend that dependence/independence is
anything other than a continuum, nor do I see any value in doing so.

JE:-
If fitness dependence cannot be empirically differentiated from fitness
independence via SOME testable evolutionary theory then "anything goes"
simply because what remains the one same system and what constitutes a
Darwinian competing system to it, cannot be empirically differentiated.


snip<

IMHO, no two things or processes in the
universe are 100% absolutely independent.

------------------------------------ end repost ------------------------
----

JE:-
Your final comment in the above repost: "no two things or processes in
the
universe are 100% absolutely independent" is a get out of jail free
card. It
allows you to only subjectively dictate what differentiates group
fitness
interdependence from group fitness dependence without having to be
empirically accountable.

I disagree, and I think I have explained my reasons in sbe dialogues with
you before.

JE:-
You have not provided:

1) WHY you have rejected my argument out of hand.

2) An alternative argument which can be tested empirically.

3) How a science of evolutionary theory can possibly exist if researchers
remain free to define fitness independence and fitness independence in any
way that they like. Surely it is obvious that whenever fitness independent
forms are not able to be objectively differentiated from fitness dependent
forms then an evolutionary theory which remains testable against nature does
not exist. In such an unhappy situation only a bunch of beliefs exist which
remain subject to nothing which can be described as an empirical test
against nature allowing bias and inevitable political misuse.


The only possible reason why "under this description there is no
evidence
that the group actually exists as a possible unit of selection" is
because
additive fitness associations can only produce _fitness independent_
associations. Associations between what? You haven't said what you
are
referring to.

JE:-
Just, any group of biological entities, e.g. any group of organisms, any
group of genes, any group of cells....

I am talking about independence of fitness at different levels of
organization. For example, individuals in a group could have high
fitnesses,
but the group could have low fitness, and vice versa.

JE:-
This remains empirically possible if the fitness of a group is not just
the
simple sum of the fitness of each part. If most individuals in such a
group
have a higher fitness then any additive group must also have a higher
fitness. However, if any group fitness remains NON additive then it can
become geared to become either much higher or much lower than the mean
individual fitness. The cost: a loss of fitness independence.

Every time I try to make the discussion more clear by emphasizing that
group
fitness may not be a function of individual fitnesses at all you seem to
steer it back to the additive/non-additive issue. Can we try to avoid
the
semantics about "dependence" and focus on a complete absence of a
functional
relationship?

JE:-
I will continue to steer discussion towards and not away from *ANY*
empirical test simply because the epistemology of science requires me to do
so. I am happy to consider other empirically based alternatives as to WHY an
additive in fitness group cannot constitute one selectee. So far you have
not provided one.

to reiterate: without being able to _empirically_, i.e. objectively
differentiate between fitness dependency and fitness independency (which
includes fitness interdependency) using anybodies unambiguous theory, no
basis exists for the organism concept within Darwinism. Without this concept
the science of biology cannot exist. If you argue that it can, what are you
replacing it with?


This means that selection between additive in fitness parts must be
occurring before selection between groups possibly can.

This strikes me as rubbish, but then I may not understand your
argument.

JE:-
My hypothesis which I have often repeated here remains precise and
empirically refutable, i.e. remains rigorous. I define any additive in
fitness group to consist entirely of independent in fitness wholes which
_intersect_ their fitnesses allowing the largest fitness to reduce all
the
other fitnesses to intersecting (reversible) subsets of itself. This
represents a testable logic of natural selection. This is the reason why
I
state that selection between additive in fitness parts (which actually
constitute wholes) must be occurring before selection between groups of
them
can. The facts of this matter is that any additive in fitness parts of a
population are not fitness dependent parts of one selectee but
empirically
fitness independent wholes. Reversing this proposition is empirically
unsustainable.

As I think about levels of selection, it is clearly possible to have
selection at the group level without any selection occurring at the
individual level.

JE:-
Only when supposed as competition between non additive in fitness wholes
which empirically represent an individual. When employing selection via
competing additive in fitness wholes within an additive meta population,
selection has already operated on these "parts" (competing Darwinian
individuals) before it could possibly occur between populations of them.
The
first additive in fitness level is just the fertile organism level. It
dominates every other level simply because of this fact. If you can
provide
a lower additive in fitness level which is empirically based then please
provide one.

If selection remains equal for each Darwinian selectee then it has
operated
before selection at the additive group level has, e.g. extinguishing one
additive in fitness population in favor of another group. This is
routinely
but incorrectly employed to represent an act of group selection.

Selection at different levels are independent processes that need not
influence one another, although they may often influence one another in
nature.

JE:-
But this "influence" can employ two _contradictory_ pathways:

1) A non mutualised influence directed from above (using nested sets of
fitness): Here an enforced fitness loss at a lower nested set level
provides
a fitness gain for the largest fitness set it remains within. A
schematic
example: if A is entirely nested within B (not the reverse) then A CAN
be
selected to sacrifice fitness for B simply because selection only occurs
as
a contest between B's, i.e. competition remains entirely at the B level
of
selection. Note: the opposing nested set fitness configuration provides
an
empirically based refutation.


2) A mutualised exchange influence not imposed from above (using
intersected
sets of fitness): Fitnesses reversibly intersects within an additive in
fitness proposition allowing those receiving a bad fitness deal to
simply
leave and find a better one. A schematic example: If A only reversibly
intersects with B then selection between A and B operates before
selection
between additive in fitness populations of either do. Selection between
groups constituting additive in fitness selectees can produce a grouped
selective force but not a grouped selectee. These groups can only act as
one
(of many) selective forces on Darwinian individuals e.g. the Baldwin
effect.
Note that V. C. Wynne-Edwards confirmed this in his final book on this
subject which occupied most of his life (this last book remains
ignored).
Wynne-Edwards recanted only allowing group selection via groups but not
on
groups (which of course is not group selection). These groups can act as
a
selective force but not as a selectee.

I predict that an additive fitness association can never allow the
selection
of a fitness loss in order to sustain a fitness gain at a meta additive
level. A non additive set nesting association represents a different
kettle
of fish because any inner nested set remains stuck within the largest
one so
it can only be selected via the fitness of the larger set. OTOH additive
fitness associations remain entirely reversible allowing those suffering
a
reduced fitness to move somewhere else.

To which of the above do you refer?

I am referring to neither one. I am trying to steer the discussion to the
case of no influence.

JE:-
No influence? How is such a thing possible? It seems to me that everything
affects everything else given chaos theory where just one butterfly flutter
can trigger the weather to change. What is required from is an empirical way
to separate two entirely _contradictory_ effects as far as _fitness_ is
concerned: dependent and independent (which includes interdependent).

Here is a riddle for you. How can a human
religious
group that prohibits reproduction and advocates suicide for its group
members have high fitness at the group level? [Remember - group fitness
refers to the propensity for persistence and spawning of daughter groups.]

JE:-
At best it can only have an entirely illusionary "high fitness at the group
level". At worst it remains obvious that such groups have a lower fitness
compared to others. I know of no human group which "prohibits reproduction".
What human groups do is regulate this in some way. It seems clear to me that
group selection is not required here given the fact that mean total
productivity per individual increases as group size increases but at a cost
which has to be paid.

Groups can evolve "suicide" but as a rare last resort mutualised group
defense mechanism without any group selection required, if and only if, the
costs on average remain less than the gains where both may be units of risk
(like an insurance company). I am happy to expand on this argument which
appears to me to remain neglected within evolutionary theory. Suicide as a
way of life cannot evolve for the same reason that cannibalism as a way of
life cannot evolve: the individual costs are greater than the individual
gains.

Your definition of group fitness as "the propensity for persistence and
spawning of daughter groups" remains a forever ongoing proposition and
therefore non testable. Group persistence can always be increased at the
expense of group reproduction and vice versa allowing a reduction of one to
be _explained away_ as some ongoing future increase for the other. For your
argument to make rational sense at some stage you will have to define
criteria which require either "propensity for persistence" to become more
important than a "spawning of daughter groups" or vice versa. It should be
self evident that group reproduction cannot be selected to become reduced in
order to increase group persistence simply because selection for increased
group persistence (group survival) cannot win against group reproduction.
What serves group selection the best is the same logic as to what serves
individual selection the best: lean and mean survival strategies which free
up limited resources in order to maximize reproduction.


This in turn means group selection must always remain subject to
individual
selection disallowing the evolution of organism fitness altruism in
nature
via selection between additive in fitness groups.

JE:-
The above is deductive from the inductive inference that additive groups
of
fitness cannot constitute one selectee, just the one selector (the
contradictory opposite proposition).



It suggests that the existence of the group as a potential target of
selection is a figment of our imagination.

JE:-
I argue that group selection between additive in fitness groups
remains 100%
subject to individual selection, i.e. it can act as an amplifier for
Darwinian selection acting at the fertile organism level. The common
argument that it can contest and win against the Darwinian fertile
organism
level was and remains _entirely_ false.

It seems to me that what you are talking about has nothing to do with
multilevel selection. You are making a straw man by defining groups as
the
sort of thing that selection does not grab hold of, then pointing out
that
selection doesn't grab hold of them.

JE:-
No. I stated clearly and unambiguously that an additive in fitness group
cannot possibly represent a single selectee but it can constitute one
(of
many) selectors. On the other hand a non additive in fitness group can
and
does constitute one selectee, e.g. one group of somatic cells
empirically
represents a single organism fitness. The empirical test: provide a
verification of just one organism which is the simple sum of the fitness
of
any of its parts. None exist. Not a single additive somatic gene or
somatic
cell fitness can be found. No additive organ fitness exists. No body
part
fitness can be just added to another to provide an empirically based
additive fitness. The very first additive in fitness level is the
organism
level which must become fertile to reproduce itself. This is why the
organism induction remains essential to the science of biology. I cannot
see
how it is possible to make this critical but basic distinction any
clearer.

Clarity may not be the problem. I just don't accept your premises. I
have
tried to be clear about the problems I have seen.

JE:-
But my premises remain empirically based. What are you replacing them with
and why are you rejecting them out of hand?


You should try considering the sort of groups that selection can get
traction
with, such as groups with fitnesses that are either non-linear
functions of
their individual's fitnesses or groups with fitnesses that are not
functions
of their individual's fitnesses at all.

JE:-
The only empirical example that I know of groups which have fitnesses which
remain non-linear functions of their individual's fitnesses are organism
parts grouped within one fertile organism. To my knowledge all organism
group fitnesses remain linear. I you can provide documented examples which
refute this then please do so.

JE:-
Please provide an example. In this case each group can only constitute a
single selectee and not a group of selectee's.

Following on my riddle above, the fitness of a religious group could be
independent of the fitnesses of its component individuals in the sense
that
their may be no link between the two.

JE:-
I can see no way that "the fitness of a religious group" can be
"independent of the fitnesses of its component individuals" simply because
the very existence of these groups remains dependent on just the existence
of the individuals concerned. The only valid question here is the nature of
their fitness relationship. For some reason it is this which remains evaded.


snip<


I prefer to think of group selection acting when there is absolutely
no
correlation between individual fitnesses and group fitnesses.

JE:-
If the fitness of the group is NOT just the simple sum of the fitness
of the
parts then that group must constitute one Darwinian selectee.

Then maybe you are a multilevel selectionist after all.

JE:-
No, simply because selection at the very _first additive and independent
fitness level_ determines the final fitness. All so called higher
multi-levels of fitness that remain additive evolve to become fitness
mutualistic to just this first additive in fitness level. Nested higher
levels of fitness remain lethally contradictory forcing a resolution as
to
who is the boss: e.g. the transformation of fitness dependent to fitness
independent cancer cells. These newly fitness independent cells have no
other choice but to contest the fitness of the body they are within.
Healthy
cells constitute a dependent nested (not independent intersected)
fitness.
Once these cells transform to have an independent fitness a selectee a
war
inevitably results which destroys both the cancer and the body.

IOW what we are looking at here is one organism and any infertile
offspring
as just a single unit of selection. Quite clearly the fitness of each
cell
or each organ does not simply add up to provide the fitness of one
organism
no matter how you define fitness. I might add neither does the fitness
of
the genomic genes. Therefore it is not possible to argue that any of
these
parts have an independent fitness.

We are having a semantic problem here. If the fitnesses of higher
units was
constrained to be merely the sum (or average) of the fitnesses of
their
components, then I would describe the fitnesses of these different
levels in
the hierarchy as dependent.

JE:-
But the semantics I use remain empirically refutable. How can it be
claimed
to be empirically true that "the sum (or average) of the fitnesses of
their
components"... "would describe the fitnesses of these different levels
in the hierarchy as dependent" when empirically these can be proven to
remain fitness interdependent (just reversible intersecting sets
fitness)
and therefore fitness _independent_?


They are independent if they have no influence on one another.

JE:-
But this is empirically impossible (see above).

They are
independent to a degree if the fitness at one level is only one of several
factors influencing fitness at the other level.

JE:-
I put it to you that factors which appear to you "at one level" as "only one
of several factors influencing fitness at the other level" are actually
contained within a fitness. IOW, the "several factors influencing fitness"
at a higher level must all be contained within a fitness at the lower level.

If the fitnesses at different levels were not so constrained, I would
call
them independent. You seem to be using "dependence" in the opposite
way.

JE:-
My argument is that you have these critical propositions in reverse
where my
claim can be empirically substantiated.

Put another way, any such argument constitutes an oversimplification
of
empirical reality which must be corrected before it can be validly
applied.

In my opinion fitnesses at nested levels of organization are often
intertwined and only semi-independent, but it is important to
recognize
that being semi-independent is not the same as being entirely
dependent. It
is autonomy by degree.

JE:-
Fitness "autonomy by degree" means nothing to evolutionary theory if you
cannot/refuse to define what does and what does not constitute an
autonomous
fitness. My definition: any additive fitness (no matter how you define
fitness) represents an autonomous fitness. Therefore, any non autonomous
fitness is any non additive fitness which can only be represented as a
nested fitness.

Please supply your definitions.

Done above.

JE:-
I can see no definition/definitions that you have provided above which
enables me or anybody else here to be able to empirically discriminate
between independent and dependent fitnesses!

Regards,

John Edser
Independent Researcher

edser@xxxxxxxxxxxxxx



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