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Fisher’s Fundamental Theorem of Natural Selection: the death sentence for Darwinism

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Consider the following claims:

Darwinism requires that the Fundamental Theorem [of Natural Selection] does not apply most of the time.

Walter ReMine
Biotic Message

and

a relative lack of natural selection may be the prerequisite for major evolutionary advance

Mae Wan Ho
Beyond neo-Darwinism

and

Concerning this theory [Darwinian evolution], I believe that we might question (or at least note) the following:
….
(10) The internal contradiction in its major theoretical cornerstone — Fisher’s fundamental theorem

Stanley Salthe
Analysis and critique of the concept of Natural Selection

and

many genomic features could not have emerged without a near-complete disengagement of the power of natural selection

Michael Lynch
opening, The Origins of Genome Architecture

Distinguished evolutionary biologist Michael Lynch recently published a much anticipated book, The Origins of Genome Architecture. Curiously, in this magnificent 494-page book, only an obligatory mention of the name of Charles Darwin was made. Darwin was mentioned passingly on 3 pages in the chapter entitled “GenomFart”.

It was also in this book Lynch demonstrated his great irritation with the advocates of Natural Selection (like Richard Dawkins). So great was his irritation that he gave the hard core Darwinists the ultimate insult, he likened them to ID proponents!

the uncritical acceptance of natural selection as an explanatory force for all aspects of biodiversity (without any direct evidence) is not much different than invoking an intelligent designer

Michael Lynch
The Origins of Genome Architecture, p 368

Why the disdain for Natural Selection? It follows beautifully from Fisher’s Fundamental Theorem of Natural Selection.

Lynch is one of the world’s foremost experts in population genetics. Population genetics is the sort of anomalous discipline in biology that has a rich tradition of mathematics, and one that commands both respect and disdain. Lynch himself points this out, “It is well known that most biologists abhor all things mathematical…”

What sort of things do population geneticists do? They mathematically describe the evolution of populations. They quantify the amount of natural selection going on in a population. If natural selection is like the notion of force in physics, then it only makes sense to attempt to quantify how much or how little of this “force” must be in operation.

A central figure creating the ability to quantify and measure the amount of selection acting in a population is R.A. Fisher. Fisher was a first rate mathematician, and his work is celebrated in ID circles as it led to the formalization of the Explanatory Filter and other means of complexity analysis.

But in addition to the groundwork Fisher provided for design detection, he has provided much ammunition for ID proponents in the field of population genetics. Make no mistake, Fisher is a Darwinist, hailed by Stephen Gould as a “patron saint”, so the irony is that he continues to be an unwitting hero for the ID hypothesis.

If his Fundamental Theorem of Natural Selection gave population geneticists the tools to measure the amount of natural selection in a population, what would happen if these sophisticated techniques demonstrated Natural Selection had to be next to non-existent for evolution to advance? Answer: Darwin’s theory would be formally disproved! And in fact that is the case, and it is only quietly acknowledged in the literature (as hinted by the quotations above).

How can we measure natural selection? Without going into the deep details, we conventionally assert selection exists if there has been a reduction in diversity in a population. For example, let’s say we have a population of fruit flies and apply a pesticide to the population such that only 1 in 1000 fruit flies survives. The diversity of the population is severely reduced, and we can assert the pesticide applies a very strong selection force on the population.

It’s actually a bit clumsy to use the fundamental theorem of natural selection to describe the strength of natural selection and the reduction of diversity. With some degree of gyration one can probably do it using Fisher’s Fundamental theorem. But the bottom line, according to Fisher’s fundamental theorem, as “fitness” increases, diversity must decrease! [I’ll save the technicalities in the notes below and comment section.]

As an aside, Fisher’s theorem was controversial and misunderstood until the Creationist George R. Price reformulated it. The new version of Fisher’s Fundamental Theorem by creationist George Price can be found at Wiki here.

The Wiki biography comments:

[Price] thus clarified Fisher’s fundamental theorem of natural selection, which had caused some controversy and misunderstanding. He believed that this equation had been a gift from God, a miracle after a religious experience.

And given that Fisher’s theorem essentially destroys any hope of Darwinism being true, I would have to agree with George Price, that Fisher’s Theorem and Price’s reformulation was a gift from God. After Price’s renounced Darwinism and became a creationist, he managed to publish his ideas in the prestigious scientific journal Nature in the article, and The Journal of Theoretical Biology. It is noteworthy he published in the prestigious journal Science on Science and the Supernatural.

In Death of Altruist we read:

Price made his final revisions to “The Logic of Animal Conflict” the following February. In a cover letter, he explained to Maynard Smith that he had made a few changes to accommodate his newfound belief in creationism. “I think I found wordings that you won’t object to, and that won’t shock Nature’s readers by making them suspect what I believe,”

Ah the irony of it all!

But let me return to the issue of Fisher’s Fundamental Theorem, and particularly Lynch and Salthe’s observations. Recall Lynch wrote:

the uncritical acceptance of natural selection as an explanatory force for all aspects of biodiversity (without any direct evidence) is not much different than invoking an intelligent designer

I highlighted the word “diversity” for a reason. How does fitness improve according to Fisher’s Fundamental Theorem or Price’s Equation? Answer: by reducing diversity.

If we presume that all life descended from a single species and diversified, how can we logically argue that diversification happens through a process of removing diversification! Some may invoke things like allopatric (geographic) speciation or sympatric speciation where mutant forms are isolated somehow from the parent population, but is this not essentially a means of protecting new species from the culling effects of natural selection? It’s surprising the illogic of Darwinian claims has not been readily apparent!

Consider for example the problems of evolving a 3-chambered heart to a 4-chambered heart. The IDEACenter website has a wonderful treatment of the problem. See: The Vertebrate Animal Heart: Unevolvable, whether Primitive or Complex.

If we have species with 3-chambered hearts, how can species with 4-chambered hearts evolve? Let’s say a creature with a 3-chambered heart gives birth to a creature with a 4-chambered heart. This of course would be a saltational miracle in light of the fact a LOT of associated plumbing and developmental pathways have to be in place to make this possible. If the plumbing is hooked up in the wrong way, death results, no reproduction, no evolution.

But even granting this miracle set of mutations can happen, what role can selection possibly play? It has to allow both changes to exist simultaneously. If selection for the 4-chambered heart overtakes the population, then the 3-chambered hearts are eliminated (which is definitely not the case since reptilian 3-chambered hearts still exist). If selection for the 3-chambered heart takes place, then 4-chambered hearts are eliminated (which is clearly not the case since mammalian 4-chambered hearts exist).

Of course, one will argue that the two kinds of heart architectures might fall in separate niche’s (either geographically or via other means) and thus we prevent competition between the 3-chambered and 4-chambered heart. But competition is at the heart of natural selection. Thus I’ve demonstrated that in order for natural selection work, we have to prevent natural selection from working!

One of course would object and say that such saltations from 3 to 4 chambers violates Darwinian gradualism. Fair enough, but one is still confronted with the same problem. Let’s say one part of the population has pre-cursors to 4-chambered hearts and the other part of the population does not. For the 4 chambered heart to evolve, selection pressure has to be non-existent on the pre-cursors, exactly in the spirit of what Mae Wan Ho asserted: “a relative lack of natural selection may be the prerequisite for major evolutionary advance”. And if one thinks of it, this is actually a more forceful statement of the problem posed by irreducible complexity. But I save development of that brainstorm for another day…:-)

This contradiction between Fisher’s Fundamental Theorem and Darwinism has not been lost upon those I quoted above, and it has not been lost on those who research molecular evolution. In the 1960’s and 70’s, a problem emerged in trying to reconcile the existence protein polymorphisms (another word for “diversity”) and the “fact” of natural selection. Heated debates ensued, but the molecular quants like Jukes, King, and Kimura prevailed in putting forward the idea of non-Darwinian evolution for molecular evolution. See: Non-Darwinian Evolution and Neutral Theory of Molecular Evolution.

This fine tradition of “non-Darwinian” evolution has been quietly handed down and slowly extended to other domains of evolution, not just molecular. Lynch’s latest book is rich with Kimura’s math, and if one wishes to see the ID perspective on population genetics and Kimura’s math, I highly recommend Genetic Entropy by renowned Cornell geneticist John Sanford or Biotic Message by Walter ReMine.

And I close with this thought. Fisher’s Fundamental Theorem and its logical consequences cannot be properly taught in the School District of Dover Pennsylvania since Judge Jones ruled it unconstitutional to critically study the ideas of Charles Darwin. I’m grateful that the internet still provides a means for spreading the truth.

NOTES:

1. Walter ReMine has a wonderful exposition of Fisher’s fundamental theorem in his book Biotic Message. He explains Fisher’s Theorem by likening genotypes to bank accounts. Consider you have a portfolio of 2 bank accounts, one account starts with 10,000 earning 10% interest and the other starts with 10,000 at 5%.

The composite starting value of the portfolio is $20,000 with an instantaneous mean interest rate of 7.5% [ 7.5% = (10% + 5%) / 2].

However the mean interest rate of the portfolio will not be 7.5% forever, but will eventually move asymptotically toward 10% over time. The portfolio will thus become over-weighted and less diversified toward the more “fit” account bearing 10%. Fisher-Price describes the evolution toward an over-weighted portfolio.

How are interest rates related to the notion of selective fitness? This follows from Fisher’s Malthusian notion of fitness as is readily seen in this Wiki treatment Fitness

Wabs = N_after/N_before

Wabs = absolute fitness
N_after = number of individuals or money after selection
N_before = number of individuals before

For example, using the money analogy with an interest rate of 10% we can say a generational cycle is 1 year. Let

N_after = $11,000
N_before = $10,000

Wabs = 1.1 or a 10% increase

It can be see from the theorem it is a bit clumsy to actually apply it in terms of trying to analyze something like heart evolution, but I believe it is still correct, and I hope it suffices to at least get the discussion of these issues going.

2. I provided my take on Massimo Pigliucci’s review of Michael Lynch’s book in Michael Lynch: Darwinism is a caricature of evolutionary biology

3. To the formalists out there, I concede that reduction of diversity is formally only necessary, but not sufficient condition to assert the existence of selection since it is possible survival might be owing only to luck and not “fitness” as Raup pointed out in his book Bad Genes or Bad Luck! But Lewontin essentially pointed out (in Santa Fe Winter 2003) enforcement of this formalism would effectively discredit the concept of fitness and natural selection altogether! Kimura also shows the problem of distinguishing the effect of random walks from the effects of selection. Good luck often trumps good genes!!!!

4. A thriller movie actually W delta Z came out this year with Price’s equation as a central theme.

5. NAS member Masatoshi Nei is extending Kimura’s ideas to domains outside of molecular evolution. See: Prominent NAS member trashes neo-Darwinism

Comments
Greetings, Sal: Throughout this discussion, I think it is very important to keep in mind that Darwin's algorithm (Daniel Dennett's term) for natural selection has three prerequisites: 1) variety (that is, substantive differences between phenotypes in populations); 2) heredity (that is, vertical inheritance of phenotypic characters); and 3) fecundity (that is, reproduction that can, but need not necessarily, exceed replacement). Given these prerequisites, the following outcome is virtually inevitable: 4) unequal non-random survival and reproduction (i.e. proliferation of certain phenotypes relative to others). As I have pointed out repeatedly, the key to understanding evolution is to focus on the "engines of variation"; that is, those mechanisms (there are literally hundreds, perhaps thousands) by which new phenotypic characters are produced. These characters, produced by the various mechanisms that I list at my blog: (http://evolutionlist.blogspot.com/2007/10/rm-ns-creationist-and-id-strawman.html) provide the raw material upon which natural selection, sexual selection, genetic drift, meiotic drive, and all other sources of linkage disequilibrium operate. Fisher's Fundamental Theorem, concisely stated, says that the rate of natural selection is a direct function of the amount of genetic variance in a population. That is, the greater the variation, the more raw material natural selection has to work on, and therefore the greater the degree of genetic change over time (note the emphasis on genetic change; as I have already pointed out in other threads, this is a serious limitation of Fisher's approach, a limitation that also applies to the "modern evolutionary synthesis" as it existed in 1959). But, according to Fisher's mathematical models, natural selection can only result in a decrease in overall genetic variance. This is because the effect of natural selection is to remove some variants while preserving others. The variants that are removed represent a decrease in genetic variance, hence Sal's contention is essentially correct. While it is true that disruptive selection (also called "diversifying selection") has the effect of producing two (or more) peaks in mean character states, rather than one (as in the case of both directional and stabilizing selection), it is not true that disruptive selection produces a net increase in genetic variation by itself. Only if the "engines of variation" produce the requisite new genetic material to allow for the shift in mean character state(s) will this happen. Ergo, it is not natural selection that "produces" new variation at all. On the contrary, natural selection merely channels the immense variety produced by the "engines of variation" into particular adaptive zones. Those zones exist, not because of selection, but because of the tremendous capacity for producing new variations that exist in the sources of variation that I have listed on my blog. So, in the interests of fair play, I hereby suggest to the ID supporters reading this post that opposing natural selection is a pointless exercise. Current evolutionary theory has already moved far beyond the limited theoretical models that formed the basis for the "modern evolutionary synthesis". You are literally attacking a dead horse. Rather, the real focus of everyone's attention should be on the "engines of variation". That is, on those mechanisms by which variation in phenotypic characters is produced. If I were an ID supporter (I'm not, BTW, but I have good friends who are), I would strongly recommend investigating the dynamics of the various mechanisms by which phenotypic variation is produced (it is maintained, of course, by selection and drift, but that's not the point). If the processes by which variation are produced can be shown convincingly to demonstrate foresight (i.e. that they produce characters that, when produced, are already necessarily adaptive), then ID may be a reasonable and useful component of any hypothesis for the origin and proliferation of adaptive characters in populations. If, however, the processes by which variation are produced can be shown convincingly to not demonstrate foresight (i.e. that they produce characters that, when produced, are not necessarily immediately adaptive, but become so following a change in environmental conditions), then ID is unnecessary as an hypothesis for the origin and proliferation of adaptive characters in populations. And please: in the interests of intellectual integrity, please do not quote parts of the foregoing analysis out of context. In particular, it would be grossly intellectually dishonest to characterize the foregoing as asserting only that “ID may be a reasonable and useful component of any hypothesis for the origin and proliferation of adaptive characters in populations”, without also including the second half of that assertion: that without a convincing demonstration that new variations are immediately adaptive, "ID is unnecessary as an hypothesis for the origin and proliferation of adaptive characters in populations." Thank you for a very stimulating discussion!Allen_MacNeill
April 4, 2008
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Good analogy, Gerry. In theory, natural selection is always interested in what provides the greatest gain. However, given that natural selection only acts in the present (and only with one particular living organism at a time), it is more accurate to say that it is only interested in immediate gain. As you point out, this could well be very different from what is best in the long run (this is another example, really, of the foresight problem irreducible complexity attacks). Frankly, natural selection is only interested in culling the organisms that don't make the grade today. Here. Right now. It has no foresight; creates nothing; produces nothing; generates nothing. It only culls. Thus, natural selection, by definition, can only eliminate, not create. Yet the irony is that people often confuse the issue and think that natural selection is somehow a driving force for creating new kinds of things. The concept of natural selection may have some (exceedingly limited, in my view) value in terms of describing the survival of the fittest, but has absolutely nothing to say about the arrival of the fittest. I think Sal's post is interesting because it highlights that not only is natural selection unable to create anything, but also that natural selection -- the culling force that it is -- often needs to be suspended in order for any real creative work to take place. I'll have to think about Sal's point a bit more, but I think this is a very significant additional nuance.Eric Anderson
April 4, 2008
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Seems to me, since future conditions are unknown and may favor either this or that, the most fit species is the one that is most diversified. Any population culled for fitness to a particular set of circumstances (by natural selection or whatever) must therefore be less diverse and less fit to face a foreboding and indeterminate future. A portfolio maximized for short-term gains is not likely to be a portfolio suited to weather the vicissitudes of an unknown future economy. Is this this what you're trying to say, Sal?Gerry Rzeppa
April 4, 2008
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[...] UD on fundamental theorem of NS the uncritical acceptance of natural selection as an explanatory force for all aspects of biodiversity (without any direct evidence) is not much different than invoking an intelligent designer [...]» Fisher and natural selection
April 4, 2008
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Note to the reader, the interest rate calculation I made was in a 1 year discrete one shot iteration, not continuous compounding over infinitesimally small (as in less than pico seconds), hence you'll find the final interest rate I provided above is slightly off. If one uses a daily compounded formula, the result is more accurate where the initial effective annual interest rates are: 5.1267% and 10.5156% for a starting mean of 7.8212% and a variance of .0726%. The end result after year in light of this refinement of daily compounding is 7.8212% + .0726% = 7.8938% which is better. Further refinement to nano-second compounding would probably yield better results. Let the reader understand, Bob accuses me of not fundamentally understanding the theorem. I've laid out the case in terms of an interest rate bearing portfolio. He can at least argue that I don't understand Fisher's theorem in terms of the math in this context. I welcome his analysis.scordova
April 4, 2008
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As to his initial statement: "Secondly, selection can also act to increase diversity, for example if selection is disruptive." Bob O'H adds this clarification: "There will be a reduction in the frequency of genotypes after selection, but then there will be mating and recombination, which (unless there is assortative mating) will regenerate these genotypes." ------------ Thus we can see that the first statement was indeed an inaccurate description of the particular process in mind. Notice that recombination is not a selective event. It may indeed result in diversity, but it did not result from natural selection. If what Bob is saying is that selection (by eliminating some genotype) makes way for others, then that is probably true as far as it goes (and indeed, may be so obvious as to avoid the need for significant discussion). However, selection does not itself produce the new "others." I think this is an example of ascribing powers to selection that properly should be ascribed to other events (in this case recombination). Regardless of the nuances of Fisher's theorem, are we actually suggesting that selection is not, by definition, a culling process? Any way you cut it, it seems difficult to sustain the idea that selection produces anything. That is certainly not how Darwin used the term, and it is not the typical understanding today.Eric Anderson
April 4, 2008
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Let GR be the average growth rate. Rate of increase of GR = Variance of growth rates Suppose there are N groups, each with its own Malthusian growth rate. let p_i be the fraction of the total group that is in group i. Let m_i represent the growth rate of group i. Then GR is the growth rate averaged over m_i and the distribution of p_i. The mean is the sum of m_i p_i as i goes from 1 to N. Variance is sum of [p_i (m_i^2-GR)] as i goes from 1 to N. Recall, the growth rate of each group is correlated to Fisher's Malthusian notion of fitness. In the above interest rate case, the mean is .075 the variance is 0.000625. Thus the average Malthusian growth rate is .075 and the rate of increase in that growth rate will be 0.000625. At the end of one year the average growth rate will be .075 + 0.000625. Is that calculation materially incorrect? Unless the individual groups have identical absolute growth values, I don't see that there wil be any thing but a reduction in the variance of the growth rates. Is that correct, Bob. And if they all have identical fitness values, it's hard to argue natural selection will work at all!scordova
April 4, 2008
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The mean fitness increases (i.e. like the mean interest rate increases in my example form 7.5% asymptotically to 10%) in relation to the genic variance in fitness.
That would be marked incorrect, but not totally. Firstly, "in relation to" is vague. Are you a field biologist? Secondly, it's not genic fitness that's important, but additive genetic fitness. For the purpose of the discussions here you got one thing right. Nothing is said about a reduction in the variance or diversity. IOW the whole post is based on a total mis-understanding of the theorem.Bob O'H
April 4, 2008
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Do you actually know what And Fisher’s Fundamental Theorem’s states? It certainly does not say that diversity is reduced.
I believe I do. The mean fitness increases (i.e. like the mean interest rate increases in my example form 7.5% asymptotically to 10%) in relation to the genic variance in fitness. In the example above, the "genic" fitness variance is maximal at the initial condition of $20,000 portfolio value. It is clear as the fitness increases, there must be a corresponding drop in variance in genic fitnesses (in this case the weight of the portfolio). As the portfolio becomes more "fit", having a near 10% interest rate, the corresponding variance in genic fitness is lower since the portfolio is now overweighted in the account bearing 10%. This also implies the increase in fitness (because of lower genic fitness variance), will be less and less -- as I said, an asymptotic increase capped at a 10% interest rate. It is not hard to see that a Malthusian formulation of fitness is practically analogous to a diversified interest bearing portfolio. Thus the less fit have a smaller proportion in the composite population as time goes on. This is a reduction in diversity on a percentage basis. I welcome your comments if this if fundamentally incorrect. Yes, I'm aware of variations to my simplistic example, but I think it is materially correct.scordova
April 4, 2008
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Sal - @2: OK, but, as I have pointed out, it is not even necessary. @5: There will be a reduction in the frequency of genotypes after selection, but then there will be mating and recombination, which (unless there is assortative mating) will regenerate these genotypes. Do you actually know what And Fisher's Fundamental Theorem's states? It certainly does not say that diversity is reduced.Bob O'H
April 4, 2008
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Secondly, selection can also act to increase diversity, for example if selection is disruptive.
I thank you for this comment and I encourage the readers to ponder it. However disruptive selection is evidenced by the reduction in diversity of the genotypes in the center of the distribution, so it's still a reduction of diversity in genotypes. I look forward to your futher comments as you are indeed a professional population geneticist.scordova
April 4, 2008
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IDist, Seleciton probably has some role, but when we actually quantify the numbers, we have to conclude it was weak, and if it was weak it cannot be the major explanation for the features of life. The point however, is that I was trying to demonstrate the fundamental incoherency of trying to explain diversity through mechanisms of natural selection. The two are fundamentally contradiction. Let's say selection accounts for 1% of how biology is shaped. What accounts for the other 99%? Lynch and Kimura goes into beautiful math as to where selection is almost indistinguishable from a random walk, and thus casts doubts on the role of natural selection in the majority of cases in the first place. For molecular evolution, there is strong belief that the majority of molecular evolution had to happen in the absence of selection. This is primarily known as neutral theory. However, how does neutral theory account for irreducible complexity. Neutral theory is thus fatally flawed as well...scordova
April 4, 2008
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Thanks Sal for a great and thought provoking post! I have some questions that I am willing to ask, but first I must state that I am no biologist nor do I have much knowledge of biology (I am actually planning to study more biology in depth after finishing college, hopefully I will have time for this), and I believe in ID more-or-less on intuitive bases rather than in-depth scientific consideration of the matter. That said, I was thinking about your post and I am not sure if I got your point. I cannot see why the neccessary lack of natural selection in the begining of speciation invalidates the role played by it later on. I thought of an example (not a really good one!) of a car that is moving through a path full of falling rocks. The car (and therefore its engine) has to turn off for a while to let the rock fall then moves until there is another rock, and so on. Can we say that because the car engine had to stop many times through the walk that it actually played no role in moving the car through?IDist
April 4, 2008
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Firstly, a reduction in diversity can occur due to drift or bottlenecks. Hence, an observation of a reduction in diversity does not equate to an observation of selection.
See my note #3 Bob. You're wrong to claim that I didn't acocunt for that possibility.scordova
April 4, 2008
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Without going into the deep details, we conventionally assert selection exists if there has been a reduction in diversity in a population.
As a population geneticist who works on quantitative genetics, I can confidently assert that this is rubbish. Firstly, a reduction in diversity can occur due to drift or bottlenecks. Hence, an observation of a reduction in diversity does not equate to an observation of selection. Secondly, selection can also act to increase diversity, for example if selection is disruptive. Sal, before declaring that Fisher's Fundamental Theorem destroys Darwinism, I suggest you find out what it actually says. Yes, directional selection will usually have the effect of reducing additive genetic variance, but there are other forces which increase it (e.g. mutation, migration).Bob O'H
April 4, 2008
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