Thanks for your response. First off, I agree from the standpoint of Fisher’s actual theorem, it doesn’t seem to be reflected explicitly in development neo-Darwin/moder synthesis theory.

I read some of the references by Edwards (whom you recommended). Edwards said in:

*R.A. Fisher’s gene-centred view of evolution and the Fundamental Theorem of Natural
Selection*

As an aside we may note that, just as Fisher defined the technical meaning of the word variance in the 1918 paper, so the first recorded occurrence of the word covariance is in The Genetical Theory (1930b, p. 195). Surprisingly, the above theorem did not appear explicitly in the literature until Robertson (1966). A more general form making allowance for changes in the character values themselves was given by Price (1970, 1972a).

It seems that only during the writing of The Genetical Theory did Fisher then realise that fitness itself could be considered as the character under selection, so that, since fitness is perfectly correlated with itself, the Fundamental Theorem emerges: the rate of change in fitness ascribable to

gene-frequency change is equal to the genic variance.

But I think a charitable reading of Basener Sanford (2017) would permit the claim Fisher was first to link Darwinism and Mendelism based on Fisher’s 1918 paper:

https://en.wikipedia.org/wiki/The_Correlation_between_Relatives_on_the_Supposition_of_Mendelian_Inheritance

The not-so fundamental Fundamental Theorem came later in the 1930 paper. I scoured the earlier edition your book Theoretical Evolutionary Genetics to actually find a formula stating Fisher’s theorem, and that’s why I noticed it’s complete absence from earlier editions of your book, so I know from that, it is definitely your view Sewall Wright’s formula was foundational, not Fisher’s.

It is understandable one might think Basener Sanford 2017 claim Fisher’s formula was foundational, but I can attest I appraised them in 2016 and thereafter of our discussion in December 2015 where you said Fisher’s theorem was not so fundamental:

http://theskepticalzone.com/wp/absolute-fitness-in-theoretical-evolutionary-genetics/#comment-99127

So understandably I read the meaning of the paper differently than you would! But their choice of words and the meaning of what they are saying is worth clarifying. I leave that discussion between you Michael Lynch and Bill and John….

But, backing up a bit, my understanding is that according to Queller 2017, there is a move (including Michael Lynch and Walsh) to create the hierarchy depicted below.

Also, while backing up a bit, there are three formulas. One is what I call the Bonkers Formula which Gruar apparently used to argue “If ENCODE is right, evolution is wrong.” This applies to recombining diploid populations like humans.

I derived it here:

http://www.creationevolutionuniversity.com/science/?p=22

The way I interpret the Bonkers Formula is that it is relatively independent of the mutation/selection balance formula. This formula would take precedence over the mutation/selection balance formulas since hypothetically, from a medical standpoint, we can have harmful traits that have neutral to “beneficial” selection coefficients. Thus the ratio of “deleterious” to “beneficial” is moot if the absolute number of deleterious traits is high enough. It doesn’t make sense that a population is getting better if for every increase of IQ points and memory we add a kidney defect…..thus there is the never ending clash of “fitness” in the pop gen sense vs. fitness in the medical sense. So as far as Michael Lynch’s studies on “compensatory mutations”, they are of little comfort to those suffering heritable diseases, since having 13 babies like Octomom doesn’t necessarily translate into more personal well being….

I call the above formula the Bonkers Formula because Graur claimed ENCODE was “bonkers” because of that formula (though I think he punched some numbers in his calculator wrong). It also seems to accord with what you said on page 157-158 of your book for recombining diploid population in relation to ENCODE:

Clearly an organism with as much DNA as we have would be in severe trouble. Yet in humans well

over 98% of all newborns survive to adulthood in most industrial countries.

WHY WE AREN’T ALL DEAD.There are several possible resolutions of the dilemma. If

much of the DNA is simply “spacer” DNA whose sequence is irrelevant, then there will

be a far smaller mutational load.

….

The mutational load calculation continues to be relevant to understanding whether

most eukaryotic DNA has any function that is visible to natural selection. Recent announcements

(Encode Project Consortium, 2012) that 80% of human DNA is “functional”, based on finding some transcription or binding of transcription factors in it, are very misleading. Junk DNA is still junk DNA, however often its demise has been

announced.

So my reading of what you say is that independent of Fisher’s theorem, there is a point where enough bad mutation will lead to decline. My understanding (which could be wrong) is that mutational load may or may not be independently derivable from Fisher’s theorem? Is that right?

Then there are the mutation selection formulas

https://en.wikipedia.org/wiki/Mutation%E2%80%93selection_balance

haploid:

diploid:

These aren’t derived from Fisher’s formula, as far as I can tell. Aren’t they for the infinite population size case, and aren’t they assuming s is some fixed value rather than s being a mean? Do those formulas apply in the finite population case???

Thanks in advance.

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