Re: The Cost of Substitution [possible REPOST]





"Perplexed in Peoria" jimmenegay@xxxxxxxxxxxxx wrote:-

Wilson basically supports ReMine on the 'misrepresentation' question,
while remaining quite skeptical on whether the Dilemma constitutes
a problem for 'evolutionists'.

JE:-
The following problems

1) What is the cost of substituting one gene for another in a single
population?

2) What is significance of the many _different_ answers to the cost of
substitution to "Haldane's Dilemma"?

are related but remain _separate_ propositions. In this thread we are only
concerned with 1. If you wish to debate 2 (which depends entirely on a
resolution of 1) please start another thread.




Here is a copy of that posting:
--------------------------------------------------------------------------
---
In article <pan.2006.11.24.21.52.33.239921@xxxxxxxxxxxxx> on November 24th
in talk.origins Mark Isaak <eciton@xxxxxxxxxxxxx> wrote:
On Thu, 23 Nov 2006 22:01:57 +0000, Perplexed in Peoria wrote:
snip<
1. "Since humans and apes differ in 4.8 x 10^7 genes, ..."
This claim has already misled at least one talk.origins poster to
ridicule
ReMine for making such a silly blunder. But the blunder here is entirely
the
fault of whoever wrote CB121. ReMine does not make this claim in _The
Biotic
Message_ (or anywhere else as far as I know).

2. "Only 1,667 nucleotide substitutions in genes could have occurred if
their
divergence was ten million years ago."

JE:-
No, 1667 _fitness dependent_ gene sets (NOT separate single genes). Haldane
employed Fisher's oversimplified gene centric assumption which disallowed
heritable genetic epistasis. This preempted Haldane's false dilemma.


"The vast majority of differences would probably be due to genetic
drift,
not selection."

JE:-
Drift acts randomly so these genes were being randomly substituted long
before humans and chips even evolved and continued to do so afterwards. The
genes which have to be proven to evolve and substituted in 5-10 million
years were NON neutral genes carried by a supposed common ancestor to both.
This answer was provided by the human and chimp genome project: less than 1%
of 20-25,000 polypeptide coding genes are different between man and chimp
(which well within Haldane's 1667).

The only significant question to be answered is WHY did Haldane et al reason
that 1667 was much too small a number for a significant selectable gene
difference? The short answer is because they assumed genetic epistasis was
inherited but remained non heritable and therefore non selectable (after
Fisher's oversimplification). The number 1667 remains too little to code for
just heritable (additive) differences between man and chimp but sufficient
to code for any heritable NON ADDITIVE differences, IOW heritable epistatic
traits. Key example: ANY gene fitness! Not one, single, additive gene
fitness has ever been documented in nature, no matter how you define
fitness.


snip<

The criticisms which CB121 makes of Haldane's "Cost of Natural Selection"
are
also erroneous in my opinion:
1. "Haldane's "cost of natural selection" stemmed from an invalid
simplifying assumption in his calculations. He divided by a fitness
constant
in a way that invalidated his assumption of constant population size, and
his cost of selection is an artifact of the changed population size."

JE:-
The only valid constant (at least one constant algebraic term is always
required to supply a necessary Galilean frame of reference allowing a
proposition to remain rational and not just logical) is the proposed size of
the original population to be substituted. This CANNOT be changed "ad hoc"
(Felsenstein reduced it "ad hoc") so as provide a relative cost to just
nothing defined by anybody. If you increase the size of the original
population "ad hoc" the cost of substitution can NEVER be paid. If you can
reduce it "ad hoc" you can substitute the population "for free" via an
illegal concealed payment.


2. "He also assumed that two mutations would take twice as long to reach
fixation as one, but because of sexual recombination, the two can be
selected simultaneously and both reach fixation sooner."

This is nonsense. Haldane did _not_ _assume_ this. He _derived_ it from
his other assumptions. The derivation is given at the bottom of page
20 of _The Cost of Selection_". It does _not_ assume that the mutations
only become fixed one at a time, and it _does_ take full account of the
role of sexual recombination. Contrary to the specious conclusions which
at least one talk.origins poster has tried to draw from a calculation
given
on ReMine's web-site, _he_ does not assume this either. It would
appear that he has read Haldane's paper a little more carefully than
some of his critics, since he says explicitly on page 216 of _The Biotic
Message_:

"This does not mean these substitutions occur sequentially, one
by one. Several genes can undergo substitution simultaneously
at various speeds. If you average all these speeds, then the
total rate can be one per 300 generations."

And this is an accurate description of what Haldane showed (given his
assumptions).

Haldane's arguments, and hence his conclusions, are, however, limited to
sets of genes which have _independent effects on fitness_.

JE:-
The above constitutes the key as to why Haldane's gene centric dilemma was
always just a gene centric _misrepresentation_. Not one, single, additive
gene fitness has ever been documented within NATURE (including meiotic gene
fitnesses). At least one is required to verify an independent fitness
"effect" (no matter how you define fitness).


All genomic genes (please note: I include meiotic drive gene fitnesses here
and remain happy to debate this point) were and remain fitness DEPENDENT on
just the one, same, _genome_ fitness. The only refutable fitness that
actually exists is TDF (Total Darwinian Fitness). This is defined as: the
total number of _fertile_ forms reproduced per parent per population. It is
this fitness constant per parent per population which represents a refutable
Darwinian maximand allowing a refutable Galilean frame of reference to exist
for ALL of evolutionary theory.

snip<
In the 1950s Haldane calculated the maximum rate at which genes could
be replaced through differential reproduction and survival. He
concluded that it posed a serious problem for evolution. His conclusion
was that the long-term average rate at which beneficial alleles become
fixed in a population can be no greater than one every 300 generations.
This means that over 10 million years of alleged human evolution from an
ape-like ancestor, with an average generation time of 20 years, at most
1,667 nucleotides could be selectively replaced in the population. This
is
insufficient to account for the difference between modern humans and any
such alleged ancestor.

JE:-
The above assumption predicted on a misuse of Fisher's oversimplified
assumption of what is and what is not heritable in nature was refuted
empirically via the human and chimp genome projects.

What remains evaded via the (quiet) dumping of Haldane's false Dilemma is:
why Haldane and Fisher predicated this false dilemma on empirically false
premises and even more critically, have these false premises been corrected
today? Shamefully, the short answer to is NO, they haven't been corrected.
They remain the basis of today's population genetics.


snip<

Regards,

John Edser
Independent Researcher

edser@xxxxxxxxxxxxxx



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Relevant Pages

  • Re: The uncorrected simplifications/oversimplifications of
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  • Re: Haldanes Dilemma - again, again, again, ...
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  • Re: Ernst Mayr: Where Are We (1976)
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  • Re: Haldanes Dilemma
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    (sci.bio.evolution)