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

_{Salvador Cordova

April 4, 2008

Darwinism, Evolution, Intelligent Design}

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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

[...] Fisher’s analysis of the effect of gambler’s ruin essentially trashes his own theorem, Fisher’s Fundamental Theorem of Natural Selection. Fisher’s Malthusian notions of “fitness” in his fundamental theorem do not [...]Gambler’s ruin is Darwin’s ruin | Uncommon Descent_{May 3, 2008
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scordova: 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. So, according to the logic in the OP, how can Darwinian evolution produce diversity if by definition, natural selection decreases it? Maybe this should be a question on "Are you smarter than a fifth grader?" The most elementary formulation of evolution that I think anyone is familiar with always characterizes mutation and the like as the engine for producing variants, and natural selection as a culling factor. So what brilliant insight has been conveyed in the OP, or what oversight has been unveiled? The word mutation is only mentioned once in the entire piece: "But even granting this miracle set of mutations can happen, what role can selection possibly play?" This is merely a passing comment in a discussion regarding the evolution of a four chambered heart, and it is that discussion I want to focus on now. The central insight of the author is that if a four-chambered heart appeared, and was superior and thus replaced through natural selection the three-chambered heart, it would be a reduction in diversity, as there would only be one type of heart (4-chambered) whereas previously there were two types. But the logic of this utterly eludes me. From my perspective, a four-chambered heart is more diverse than a three chambered heart. It is certainly more complex, and I think 'complex' and 'diverse' or near synonyms. From my perspective as a software engineer, an increase in fitness always entails an increase in complexity. (We'll get to Occam's razor in a minute.) Let's throw a very instructive cliche into the mix here: "The devil is in the details." Your original understanding of a problem is nearly always incomplete, so your original solution may be very neat and tidy and elegant. But as you start testing it in real world situations, you start encountering exception after exception after exception, and you have to go back and modify your original solution continuously to account for all these exceptions. This is not merely a result of bad planning - you can imagine consumer's giving feedback, "but I want this feature or that feature", or maybe modifications are driven by a need to keep up with competitors or new technologies. So despite Gould's claim that evolution does not necessarily entail an increase in complexity, I would strongly disagree with that. (Do society, technology, laws, etc. get more or less complex over time?) In spite of all this it is certainly and obviously true that if you can ever take some big chunk of your software, and say, "Wait a minute, I could replace this 200 lines of code with 10." Then by all means do it. This is apparently the brilliant insight of Occam. So complexity in an artifact is often an indication of a lack of planning and foresight. By the end of this thread, Scordova is arguing that fitness is not even a meaningful concept:
In math, if objects A and B and C have real values, we can say: if A > B, and B > C, then A > C ( to put hard numbers, for example, consider A = 10, B =8, C = 5) This is the Transitive Property of Inequality. In this manner we have a means of even comparing apples and oranges. We can objectively say, one object has more mass than another. We cannot say the same about biological organisms with respect to fitness. Lewontin demonstrated the difficulty of making such objective statements in defining fitness. For example, in sports competition we can't make the following statement with certainty. If team A beat team B, and if team B beat team C, then team A will beat team C. Lewontin shows this difficulty with respect to the competition among varieties, along with some very distressing situations where the definition fitness varies radically with context. Thus Lewontin showed the difficulty in showing that fitness is anything more than a restatement of known statistics, and not a very good restatement at that. If fitness cannot obey the transitive properties of inequality (which is so fundamental to properties in physics and chemistry, or practically any other scientific discipline except Darwinian evolution), trying to use this concept to make statements about the evolution of complexity become essentially incoherent.
So is scordova or Lewontin claiming that the concept of fitness among football teams is meaningless because the transitive property does not hold there (e.g. If team A beat team B, and if team B beat team C, then team A will beat team C. ) Certainly I don't think Lewontin believes this (or other thoughts scordova credits to him) as I think the football analogy is scordova's. But let's examine the analogy for a minute. If team A beat team B, and if team B beat team C, then it most certainly increases the probability that team A will beat team C (if that's all you know about the teams). If through tradition Michigan has Cincinnati on their schedule every year, and beats them nearly every year and Cincinnati plays East Toledo State every year and beats them nearly every year, then certainly it is highly probable that if Michigan played East Toledo State, Michigan would beat them. Maybe some think these observations are too simplistic for this forum or thread, but apparently they elude scordova. But to continue with the sports analogy, Highly successful programs end up with teams from around the league snatching up all their assistant coaches and thus the strategies, schemes and plans of successful teams are disseminated around the league and persist for decades, long after these schemes are effective (e.g. "The West Coast Offense"). However The fact that fitness is certainly a meaningful concept in football does not imply that there is not also random genetic drift. You can have some obscure conference that for no apparent reason gets hot one year send a few teams to bowls, and then immediately descends back into obscurity. (Perhaps there's a better analogy for genetic drift specifically, however.) Here was the transitivity problem in Lewontin's words: In experiments involving competition of several genotypes taken two at a time, Dobhzansky (1948) showed lack of transitivity in fitness. That is, genotype A is more fit than genotype B in an experiment involving only these two genotypes, and B is more fit than C in two-way competition, but in three way competition C beats A. But does this imply that fitness is a vacuous concept (Or does Lewontin think that - I don't think so.) In basketball, height is a huge advantage. In a one-on-one competition, even more so. But with five players on each side, throw an extremely quick but very short guard into the mix, and suddenly his height is not a factor. Does this make meaningless the concept of tallness or fitness? Hardly. (And for the record, in response to a remark made by someone recently in another thread, I am one of those people that still believes the story of Noah's ark is literally true.)JunkyardTornado_{April 20, 2008
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I read the Lewontin article. It is a fascination article and I am glad it was recommended. It celebrates fitness as an important variable in evolution, not discounts it. In the examples on pages 24-26 it does not discount fitness but essentially says we do not know how to measure it. (Fitness may not be a single gene oriented concept by a gene network concept.) The examples used illustrate that varying environments and ecologies affect reproduction rates and measures the reproduction rates by variant. What can not be determined is what variables/networks determine the reproduction rates. It does not seem to be a random effect so it is variable related. So as the density of the population varies by the different species in the ecology, the reproduction rates vary by variant. What they cannot do is isolate the characteristics/networks within the species that are responsible for the varying reproduction rates when the various external variables are manipulated. The article highlights micro evolution and why macro evolution may be impossible because the variation within the gene pool does not have the variation necessary to do anything really complex. It also defines novelty as trivial and only as changes not seen before. It is not what we usually mean by novelty. I highly recommend that everyone read it and see if they come to the same conclusions that I did, which are different from Sal's conclusions.jerry_{April 8, 2008
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Especially problematic is the phrase "random with respect to fitness". If fitness cannot be defined in general (except for special cases like antibiotic resistance), then how can we even say "random with respect to fitness"? Sanford argues the difficulty with this becuase empirically speaking mutations on the whole do not seem random with respect to fitness. For example environmentally induced mutations (like radiation) seem to be statistically correlated against reproductive success, thus it is not random. On the other hand we have adapative mutation which seem positively correlated to fitness. Ok, for another naturalistic theory, consider the work of James Shapiro (who seems privately favorable to Behe). See Who are the Multiple Designers? James Shapiro offers some compelling answers
Bacteria as natural genetic engineers…. This remarkable series of observations requires us to revise basic ideas about biological information processing and recognize that even the smallest cells are sentient beings.
Thus there are observed non-Darwinian modes of evoltution, an I would argue they constitute 99% of the evolutionary modes present today and in the past. I do believe in ID and even the more extreme hypothesis of special creation. The fundamental theorem of calculus was widely accepted as true even 300 years before it was proven. I think the case for special creation (if true) or ID might be in a similar boat. It may seem reasonable to many, but from a theoretical and empirical standpoint the proofs are not quite mature. Seeing these hypotheses put on a more solid foundation is my personal area of interest. Time will tell if ID proponents and their creationist cousins will be successful in these endeavors.scordova_{April 8, 2008
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What causes these frequency changes?
Jerry, I recommend the examples in the article itself. Frequency change can be due to: 1. Random Drift 2. Bad luck 3. Intitial density of population 4. Mix of population Problematic is defining "fit" in the most general sense as we define mass or force or temperature in physics or even moles in chemistry. If one cannot do this, one has to question the difficulty of putting selectionist theories on par with the other sciences. I think the neutralist theories are at least framed in a way that puts them on a more sound footing scientifically (I say that as one who actually rejects mainstream neutral theory). With respect to anti-biotic and pesticide resistance. The way we are able to argue for the presence of selection is repeated trials. If for example, one trait is observed in 100 petri dishes where anti-biotic resistence occurs and no resistance occurs in 100 petri dishes where the trait is absent, we can say a selectionist metaphor is applicable. In the absence of such repeated trials, how is it possible even in principle to argue selection was the cause or not. The only way I know how is to argue "not" through issues with population resources, as Kimura did using Haldane's dilemma. However form Fisher's fundamental theorem alone, it's kind of hard to argue eukaryotic evolution from prokaryotes since, in terms of Matlthusian reproductive fitness, prokaryotes seem more "fit". One might argue the exitence of allopatric (geographic) or sympatric niches to enable selection to work in selected isolated domains, but this seems like a special pleading. In such case, Goodman's cooperative (versus competition) model would seem more accurate. i.e. a species develops ability to digest a different food source, thus it becomes sympatrically isolated without competing with the parent. The populations then live cooperatively together. This would accord well with symbotic and ecosystem evolution. I think Goodman is closer to the truth, but I say that as someone who personally disagrees with Goodman. PS by the way, I keep forgetting to list "self-organization" as a naturalistic theory. I consider that the main competitor to ID theories. I think Pigluicci of late is quite enamored with self-organization. So is Denton....scordova_{April 8, 2008
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scordova, (68) I found out what I was doing wrong. The article you cite is found between pages 16 and 17, if you look at the bottom of the pages. My browser doesn't give me precise PDF pages; instead I have to scroll down using the pages listed on the PDF. That of course threw me off. It is quite a delightful article.Paul Giem_{April 8, 2008
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"It simply affirms that types change in frequency." What causes these frequency changes?jerry_{April 8, 2008
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Dr. Giem, My apologies for a bit of confusion. Lewontin's essay begins on page 19, but his section on fitness begins on page 23. On page 24:
The difficulties of the concept of fitness are, unfortunately, much deeper than the problem of frequency and density dependence. The problem is that it is not entirely clear what fitness is.
In Fisher's formulation fitness is growth rate. Thus bacteria are more fit than humans. Lewontin points out that these sort of notions are in conflict with Darwin's notion of some sort of inherent biological property of what is "good".
Darwin’s sense of fit has been completely bypassed.
For example, in sheer numbers, the mammalian eye would be less fit than the insect type eye(s). This is problematic for explaining the evolution of the mammalian eye in terms of fitness scores. This problem I think is pervasive....How can we explaing eukaryotes if prokaryotes are more "fit"? Lewontin wasn't the easiest read, but each time I have re-read it I gain something more.
If a type increases in a population then it is, by definition, more fit. But this suffers from two difficulties. First, it does not distinguish random changes in frequencies in finite populations from changes that are a consequence of different biological properties. Finally, it destroys any use of differential fitness as an explanation of change. It simply affirms that types change in frequency. But we already knew that.
scordova_{April 8, 2008
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scordova, (65) I did what you suggested and scrolled to what was labeled page 19 of http://tinyurl.com/4sjthv and was unable to see how page 19 or the immediately surrounding pages demonstrated the difficulty of defining fitness. Could you please check the reference here and/or explain how the printed material makes your point. ThanksPaul Giem_{April 8, 2008
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Allen wrote: The lack of a PhD has not prevented me from teaching either introductory biology nor evolution at Cornell, where I have been nominated for and/or awarded several honors for my teaching, including the Clark Award for Outstanding Teaching at Cornell University.
And this relfects well on Cornell letting the best teachers teach independent of the letters after their name. The IDEA club there spoke highly of you and it is easy to see why. I think I would have very much enjoyed taking the classes you taught. And from my perspective, your Evolution and Design class at Cornell where the Bill Dembski and Michael Behe's works were studied critically was the best course of its kind anywhere.scordova_{April 8, 2008
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Gentleman, This is wonderful discussion. I was en-route 16 hours yesterday, and it is good to be on my laptop again for a few minutes. I see that DaveScot has posted a link to the paper by Lewontin which I mentioned in my original posting. I thank him for finding the paper in the first place since I could not longer find it on the net. Lewontin's paper was highly influential to Stanley Salthe and myself. It is technical, but wow, I learned so much from it. Lewontin was Gould's colleague and also (along with Gould) a mentor to creationist Kurt Wise. When I asked Wise about Lewontin last year at Baraminology 2007, Wise spoke glowingly of Lewontin. It is easy to see why, when reading this paper... Salthe considered this paper fatal to Darwinian evolution becuase it showed the difficulty of defining fitness. Scroll to page 19: http://tinyurl.com/4sjthv To simplify: In Physics the notion of force is measured in Newtons or Pounds. We can measure mass by the amount of pounds of force a gravitational field creates on object. The typical measuring device is a scale. I can even compare apples and oranges using a scale. :-) In math, if objects A and B and C have real values, we can say: if A > B, and B > C, then A > C ( to put hard numbers, for example, consider A = 10, B =8, C = 5) This is the Transitive Property of Inequality. In this manner we have a means of even comparing apples and oranges. :-) We can objectively say, one object has more mass than another. We cannot say the same about biological organisms with respect to fitness. Lewontin demonstrated the difficulty of making such objective statements in defining fitness. For example, in sports competition we can't make the following statement with certainty. If team A beat team B, and if team B beat team C, then team A will beat team C. Lewontin shows this difficulty with respect to the competition among varities, along with some very distressing situations where the defintion fitness varies radically with context. Thus Lewontin showed the difficulty in showing that fitness is anything more than a restatement of known statistics, and not a very good restatement at that. If fitness cannot obey the transitive properties of inequality (which is so fundamental to properties in physics and chemistry, or practically any other scientific discipline except Darwinian evolution), trying to use this concept to make statements about the evolution of complexity become essentially incoherent. We can compare apples and oranges in terms of their weight. Can we do so in terms of fitness? That's the problem. At best we can say the statistics may be evolve in a way analogous to a Darwinian model, or it may not. If things can evolve in a manner that is non-Darwinian, then the question is how much evolution is Darwinian, how much is non-Darwinian. I have suggested based on the channel capacity problem and Haldane's dilemma, that even if fitness could be defined (which as Lewontin showed is a hopless quest in general), only about 2 bits ber 300 generation could be attributable to Darwinian evoltuion. That correlates to less than .1% of the human genome. The rest of the evolutionary story, 99.9% is non-Darwinian. That figure is already roughly accepted for molecular evolution (Kimura and friends). The question then is why shouldn't that figure apply to just about everything else (which is Nei's thesis). I have suggested that Fisher's theorem implies that selection actually has to be substantially relaxed to account for diversification, which will permit the evolution of complexity. The euphamism used by some to say this is "time-varying fitness landscapes". So, on many accounts, independent of ID, I don't think the overwhelming majority of evolution can be Darwinian. Lynch's irritation is so great, he practicallly faults Dawkins and Dennent (as well as ID folks) for making evolution a caricature by ascribing so much power to selection. There are other naturalistic models for evolution that are non-Darwinian: 1. Brain Goodmans evolution via cooperation (rather than competition) 2. Saltationism, mutationism 3. neutral theory Of course I have my ideas. My personal challenge is that it is hard to frame the answer in terms of direct empirical science. If God were readily apparent, I'd have an easier time arguing for special creation. ID on the other hand is at the doorstep of empirical science, but it is unfortunately still only and inference, it is not a direct empirical observation in the sense of observing mass or other physical quantities. The task is complicated because information and complexity (including specified complexity) is not a physical quantity.scordova_{April 8, 2008
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Allen_MacNeill, In (15) (I know, this is late) you wrote a couple of paragraphs which I am still trying to decipher. In the interest of not quoting you out of context, as you warned about, (or at least trying), I will quote them in full (I see one of them was quoted by ericB in (63)).
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.
The above apppears to be making two assumptions (among others). First, that we can study, in a given organism, the processes by which variation are produced, and that these can be identified with the processes whereby the organism itself was originally produced. The second is that if traits are somehow produced that are not adaptive at present, but become adaptive in a changed environment, this is somehow less indicative of foresight than the production of traits that are immediately adaptive. Now perhaps I have misunderstood you, and If so I would appreciate correction. If not, the second point seems weak, and the first counterfactual. Preadaptation seems to me to require more foresight than plain old ordinary adaptation. Adaptation can immediately be preserved and enhanced by natural selection, but preadaptation requires the preservation of traits that are not presently advantageous, and may even be presently deleterious, for some future advantage, and this foresight would seem to be a hallmark of intelligence. One could argue that if the adaptations are complex and specified enough, either one could be evidence of intelligence. But the reversal makes no sense to me. Putting a spare tire and a jack in a car takes at least as much intelligence as putting the original four tires on. But that seems a minor quibble compared to the question that can be raised about the first assumption. What this seems to be saying is that the 47 (and counting) sources of phenotypic variation in a given organism necessarily account for all the variation needed to create it, or to modify it from some previous organism. I'm not sure that this is accurate. An example might illustrate the problem. As you probably know, extensive attempts to take natural variations and artificially select them for the effect of a blue rose have uniformly resulted in failure. However, recently because of some careful gene insertions and manipulations, scientists have been able to produce a rose that can reasonably be described as blue, and that has no known counterpart in nature. Now supposing that we are alien scientists exploring the earth after a disastrous epidemic has wiped out humankind, and enough time has passed so that the products of civilization (including records of what happened to make the roses blue) have disappeared, but the varieties of roses live on. Could we apply those 47+ kinds of phenotypic variation to the roses now (then) existing and explain how the rose became blue? Would we not be tempted to call it "lateral gene transfer" (meaning undirected lateral gene transfer)? And would we not be dead wrong? How could we possibly arrive at the correct answer to the origin of blue roses without allowing for the possibility of intelligent design? Furthermore, imagine an island where blue roses were planted and survived because there were few natural enemies. We might observe the roses exhibiting many variants, and identify experimentally confirmable sources of variation until we were blue in the face, and still not be able to identify the correct reason why these roses differed from the vast majority of those on the mainland. Part of the problem would seem to stem (ahem) from the assumption that all causes that have ever operated are now operating roughly equally to how they operated in the past. For intelligent agents, that may not be a valid assumption. Intelligent agents may very well create episodically, and they are not required to create when we want them to so that we can see how it is done. I don't see how your proposed way of determining whether ID is involved in present-day biology is either sensitive or specific. Perhaps you can clarify.Paul Giem_{April 7, 2008
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Allen_MacNeill (55): "So the question really is, where does the information in the genome come from, and how much does it contribute to the actual phenotypes of organisms? How much of the “design” of an organism is provided by its environment? And can any of this be shown to be foresighted?" Your posts are delightfully informative, as always. And these are good questions. However, there is a logical error in the following. (15): "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." This is assuming that all the new information comes from the engines of variation, with complexity only coming in the bottom-up fashion of being always preceded by a simpler ancestor. That might be so, but it also might not be so. You mentioned a friend's thought that "It’s all been downhill since the Mesozoic." What if there is more "downhill" to evolution than even your friend supposed? You acknowledged "Admittedly, we have a long way to go. We are only just beginning to understand how development is regulated, and the part that the environment plays in it. As we learn more about it, our models of how development and evolution are related inevitably change, to accommodate new research results." This leaves open the possibility of instances front-loaded development where it truly is "downhill" with the crest of a hill as a starting point. Notice that in such a scenario, it is not necessary for the engines of variation to demonstrate any "foresight" in the sense of reliably yielded immediately adapted variation. Nevertheless, the conclusion that "ID is an unnecessary hypothesis" in this scenario is false. One valid hypothesis is that the role of ID was at the point of front-loading the organism with the information, control mechanisms, and/or potential regarding development that makes the diverse variations we see possible. In short, even though it may take no special intelligent intervention to roll downhill, employing variations that are not foresighful, that does not allow us to conclude that intelligent agency had no role, as you suggested. Rather, intelligence may be essential to getting to and starting from the top of a hill. I agree this cannot be decided by speculation. Just wanted to point out that your list of possibilities was too short, with an important omission.ericB_{April 7, 2008
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nullasalus wrote (in #59):
"I’ve seen people sternly correct others about how evolution was just mutation and natural selection, period. This, from people who believe in evolution."
As have I. That was one of my motivations in posting my blog on the 47 mechanisms for producing phenotypic variation. I remember my evolution professor at Cornell railing about precisely the same problem, and about the grossly mistaken notion that natural selection is the principle creative process in evolution. Nothing could be further from the truth, of course. Natural selection produces absolutely nothing. It is an outcome, and is most emphatically not a "creative force." The "creation" of new variation is what the "engines of variation" (the 47 different mechanisms) do, not natural selection. The whole point to natural selection is that it preserves those variations that have the effect of allowing those individuals who have them to survive and reproduce more often than other individuals who have different variations. This is why Darwin preferred the term "natural preservation" to "natural selection", as the latter sounded too active, as well as too "intentional." However, he waited too long to try to change the terminology. By the time he expressed this opinion (in a letter to Charles Lyell; see http://www.darwinproject.ac.uk/darwinletters/calendar/entry-2931.html), "natural selection" had become entrenched in the vocabulary of evolutionary theory. On the other hand, natural selection is the explanation for why complex adaptations can evolve in surprisingly brief periods of time (e.g. 10,000 generations). Selection limits the possible range of variations in each generation to those that are closest to the forms that are already most adaptive.Allen_MacNeill_{April 7, 2008
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Haha, yeah, I suppose so (if having a PhD makes one "smarter than you"). --AllenAllen_MacNeill_{April 7, 2008
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A clarification: I appreciate that you always address me as “Dr. MacNeill”, but the fact is that I do not have a PhD nor any other doctoral degree.
Egad! Does that mean that "Dick the Dawk" is smarter than you (per the Machine video)? ;)russ_{April 7, 2008
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Allen_MacNeill, "Once again, this whole view of development and its relationship to evolution is almost completely outside the bounds of the “modern evolutionary synthesis” (aka “neo-darwinism”). It is, however, supported by massive and increasing amounts of empirical data, and represents the cutting edge of evolutionary theory today." I have to ask. Are you aware that most people - and I mean ID-proponents, ID-opponents, from every religious and areligious stripe - are utterly, blissfully unaware of this? That the typical view of evolution amounts to 'stuff kills other stuff and the stuff that lives reproduces and that's how we got apes'? Kind of excessively reducing it there, but it's not far from the truth. I mean, I've seen people sternly correct others about how evolution was just mutation and natural selection, period. This, from people who believe in evolution. Just wondering.nullasalus_{April 7, 2008
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Sal: Have an enjoyable and safe trip! A clarification: I appreciate that you always address me as "Dr. MacNeill", but the fact is that I do not have a PhD nor any other doctoral degree. I earned a bachelor's degree in biology and a masters degree in science education, both from Cornell, but have been on (unofficial) leave from the PhD program here for nigh on thirty years. I am what is known in academic circles as "ABD"; I was hired over thirty years ago at Cornell without having finished a dissertation, and have never taken the time to do so. I would like to at some point; maybe after I retire? The lack of a PhD has not prevented me from teaching either introductory biology nor evolution at Cornell, where I have been nominated for and/or awarded several honors for my teaching, including the Clark Award for Outstanding Teaching at Cornell University. Nor has it prevented me from doing my own research. As a Senior Lecturer at Cornell, I am a full member of the faculty, but unlike my colleagues in tenure tracks, I can research quite literally anything I want, so long as I am willing to pay for it myself ;-) When I reflect on my lack of a doctoral degree, I consider the fact that neither did Charles Darwin nor Thomas Henry Huxley. Indeed, Darwin's only academic degree was a bachelor's degree in Anglican theology, and Huxley never finished his bachelor's, joining the Royal Navy instead. Ergo, I conclude (perhaps a tad self-servingly) that one's contributions to intellectual progress need not necessarily by tied to a string of letters following one's name.Allen_MacNeill_{April 6, 2008
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As to the assertion that "common sense" can immediately answer questions about these topics, "common sense" tells me that the desk I am writing at is made of completely solid material, not mostly empty space, and that the light streaming down on it from my desk lamp should move faster with the rotation of the Earth than against it. That light also seems to have the characteristics of a wave-like process, as it clearly shows interference patterns at edges (I can see them pretty well, especially if I take off my glasses). And, "common sense" tells me that it should be possible to determine simultaneously both the momentum and position of a moving object, including an electron. After all, that's how you catch a baseball, right? It's just "common sense". But we all know what happened to those "common sense" ideas...Allen_MacNeill_{April 6, 2008
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Allen, he is saying that from an informational perspective the principals that can be eventually said to govern species development with replace the chaotic physical ones that you listed once we understand their limits as well. You are talking about physical processes but ID is about the informational principals that complex specified information not only requires but can be reduced to via Intelligent Design explanation. This is obviously well in the future but Jonathan Wells thinks not too far in the near distant future. That is why Dave phrased his comment ...and then...start trimming away... We have a lot of work to do but materialisms utter inadequacy as an explanatory template for origins requires it.Frost122585_{April 6, 2008
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"Canalization" as Waddington frist proposed it, and as Jablonka and Lamb have described it in their new book, doesn't refer to "canals". Rather, it refers to the idea that, as development proceeds, it develops a kind of "momentum" in which traits that were originally set in motion by genetic programs become modified and stabilized as the resjult of interactions with the environment. The metaphor that Waddington was using was "channelization" - the idea that, when water first runs over a surface (say at the top of a hill), it can flow almost anywhere. However, as it flows (and as more and more water follows it), it scours out channels that "canalize" where subsequent water flows can move. If this sounds like natural selection, that's no accident. What Waddington was arguing for was a process like natural selection, but applied to the process of development, in which the genetic program of an organism is modified by interactions with the environment to produce the phenotype of the organism. Waddington was trying to explain why some genetic changes do not produce significant effects on phenotype. He proposed that the genome "gets the ball rolling", but that most of the actual phenotype is the result of the "ball" wobbling down a series of "epigenetic trajectories", in which historical contingency plays an enormously important role. Once again, this whole view of development and its relationship to evolution is almost completely outside the bounds of the "modern evolutionary synthesis" (aka "neo-darwinism"). It is, however, supported by massive and increasing amounts of empirical data, and represents the cutting edge of evolutionary theory today. Admittedly, we have a long way to go. We are only just beginning to understand how development is regulated, and the part that the environment plays in it. As we learn more about it, our models of how development and evolution are related inevitably change, to accommodate new research results. Again, I would urge ID supporters to recognize that there is nothing intrinsic to evolutionary theory that would necessarily rule out design in nature. Indeed, as I have argued in several venues, nature is packed with design; that's what a genome is - a design for an organism. So the question really is, where does the information in the genome come from, and how much does it contribute to the actual phenotypes of organisms? How much of the "design" of an organism is provided by its environment? And can any of this be shown to be foresighted? All good questions, and all answerable by empirical research. However, absent empirical support, none of them can be answered by theoretical speculation alone.Allen_MacNeill_{April 6, 2008
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I will be on travel for a couple days before I return to this wonderful discussion. I wish Dr. Thomas English the best in his partnership with Robert Mark's Evolutionary Informatics Lab. For the record, I stay out of decisions regarding who and who does not participate, excepting for the fact I can on occasion delete posts on threads that I begin if I feel a troll is in evidence. I have not excercised this option in this discussion so far, and I welcome hearing criticism from Dr. Bob OH and Dr. Allen MacNeill. I leave the moderation decisions to those who have been kind enough to volunteer their time for the task, and to them I extend my thanks for a thankless job. Regarding Natural Selection, my main point is that it must be disengaged in some way for diversity to exist. I think it can be mathematically argued natural selection can influence a mere 2 bits per 300 generations according to Haldane's Dilemma. This is far slower than the slowest modem on Earth! Morse code transmissions are more efficient in infusing information. When I say 2 bits per generation, I mean the net gain. Because of interference selection, natural selection will destroy large numbers of good traits in addition to bad with a net gain of 2 bits per 300 generations according Haldane's work and supported by Kimura. THERFORE, on a percentage basis 99.99% of evolution must be driven by mutation and stochastic factors or better yet, imho, front loading. If anything, Natural Selection gets in the way of increasing complexity and diversity rather than assisting it. The kinds mutations that drove evoltution in the past are probably no longer in operation today. That was John Davison's thesis. There are some of us that believe in special creation (the very thing Darwin argued against). I'm only positing that if there were common descent, it had to proceed in the absence of Natural Selection exactly as Mae Wan Ho asserted:
a relative lack of natural selection may be the prerequisite for major evolutionary advance
and Michael Lynch
many genomic features could not have emerged without a near-complete disengagement of the power of natural selection
I think some are reluctant to put hard figures, but I would venture less than 1% is in the ball park for the influence of selection as far as the evolution of complexity. We know when there are selective sweeps where 99% of the population is wiped out (as in a pesticide or sweeps resulting in antibiotic resistance), it's hard to argue there is much of an net gain of information!!! If we are talking about the speed that an evolutionary algorithm can infuse information into a genome, it's pretty dang SLOW! There are finite speed limits to how fast Natural Selection can modify a genome or epigenetic information. I believe, statistically speaking, most of the mutations that would have had to drive evolution, had to do so without the sheparding of natural selection. I think front loading is the best scenario if one is a mutationist. I personally hold out hope for the special creation model, but this model so far is hard pressed to be demonstrated from scientific first principles, thus I'm left only arguing a few negative cases against prevailing views. From a scientific vantage point, I can only argue to the extent that David Berlinski is arguing, namely, in terms of what appears will not work for sure...and so far I think we can rule out Natural Selection as a major force for the evolution of complexity. Natural Selection seems to be more a destroyer than creator of complexity and diversity.scordova_{April 6, 2008
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DaveScot wrote (in #41):
"...trimming away those which are made unnecessary by the mechanism of intelligent agency."
"Trimming them away" how? Virtually none of the mechanisms I've listed are theoretical constructs. On the contrary, almost all of them are empirically verified processes, which have the effect of producing phenotypic variation. Since variation is the core of what we are all trying to understand here, it would seem much more logical to me to examine each of the mechanisms in the list, to determine exactly what kinds of effects they have on phenotypic change over time, and if any of them show any indication of foresight (a very difficult thing to do, BTW). And the number 47 is already out of date. The list grows longer every day, as more research results are published. Also, most of the entries in the list are simply "headings", encompassing dozens (and in some cases hundreds) of different, related mechanisms of phenotypic variation.Allen_MacNeill_{April 6, 2008
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Huh? Any reason why? Just curious.
Yes, Dave. You even agreed with Thomas English at one point:
Thomas Well, I guess I’ll agree with you that Allen’s 47 ways by which heritable phenotypic modification can occur is not an explanation.
You should see the crap that I have to put up with on my own blog because of my no-censorship policy. At one point I had to enable comment moderation because some trolls insisted on posting gossip about my private affairs.Larry Fafarman_{April 6, 2008
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Huh? Any reason why? Just curious.larrynormanfan_{April 6, 2008
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Thom English is once again no longer with us.DaveScot_{April 6, 2008
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scordova said (comment # 26 ) --
The culling process throws out the good with the bad.
Good point. Fruit flies, for example, are considered to be likely survivors of a nuclear war because of a high resistance to radiation. Does that make the fruit flies "fitter" than other organisms -- including humans -- that are less resistant to radiation? Also, natural selection does not necessarily consist of just culling. For example, a bird that is a mutation of a lizard can enter a new ecological niche, and other lizards are unaffected at least for the moment. That is natural selection, too. My favorite criticism of evolution is the Fundamental Theorem of Co-evolution of Total Co-dependence of Two Different Kinds of Organisms: In such co-evolution, unlike in evolutionary adaptation to widespread fixed physical features of the environment, e.g., water, land, and air, there may be nothing to adapt to because the corresponding co-dependent trait in the other organism is likely to be locally absent. First corollary: Where the corresponding co-dependent traits in both organisms are fatal in the absence of the corresponding co-dependent trait in the other organism, co-evolution by means of random mutations is virtually impossible. Second corollary: Even where the corresponding co-dependent traits are not fatal or harmful in the absence of the corresponding co-dependent trait in the other organism, the creation of only one of the pair of co-dependent traits confers no advantage in natural selection.Larry Fafarman_{April 6, 2008
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Allen_MacNeill (29): "Also, most of these mechanisms of variation are not 'random' in the classical genetic sense, but rather are 'canalized'... They are, in other words, 'guided' into particular adaptive zones as the result of a combination of environmental determinism and historical contingency." Asa Gray, Professor of Natural History (Botany), Harvard University, in his review of Darwin's The Origin of Species, 1860:
[W]e should advise Mr. Darwin to assume in the philosophy of his hypothesis that variation has been led along certain beneficial lines. Streams flowing over a sloping plain by gravitation (here the counterpart of natural selection) may have worn their actual channels as they flowed; yet their particular courses may have been assigned; and where we see them forming definite and useful lines of irrigation, after a manner unaccountable on the laws of gravitation and dynamics, we should believe that the distribution was designed. To insist, therefore, that the new hypothesis of the derivative origin of the actual species is incompatible with final causes and design, is to take a position which we must consider philosophically untenable. We must also regard it as highly unwise and dangerous, in the present state and present prospects of physical and physiological science. We should expect the philosophical atheist or skeptic to take this ground; also, until better informed, the unlearned and unphilosophical believer; but we should think that the thoughtful theistic philosopher would take the other side. Not to do so seems to concede that only supernatural events can be shown to be designed, which no theist can admit -- seems also to misconceive the scope and meaning of all ordinary arguments for design in Nature. This misconception is shared both by the reviewers and the reviewed. At least, Mr. Darwin uses expressions which imply that the natural forms which surround us, because they have a history or natural sequence, could have been only generally, but not particularly designed -- a view at once superficial and contradictory; whereas his true line should be, that his hypothesis concerns the order and not the cause, the how and not the why of the phenomena, and so leaves the question of design just where it was before.
j_{April 6, 2008
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Dave, The idea of canaling is discussed over several pages in the Jablonka and Lamb book. Don't ask me yet for good understanding of it. The book is interesting but often convoluted. The more convoluted the topic gets the more iffy the proposition. There are several good things in this book and not one challenges ID. It just expands what comes under what micro evolution can do. The link to Lamarck in the book is a little stretched but interesting. The people who wrote the book are Darwinist and anything they cannot explain they say was selected. It is their answer to everything unknown. One interesting example they use and it is a thought example is a world where all the creatures had identical DNA but completely different phenotypes. That is because cell types are not based on DNA but epigenetic factors such as methylation which determine which genes get expressed in the cell. The topic of genotype and phenotype matching or not matching or one changing without the other changing is discussed in several places.jerry_{April 6, 2008
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“The only thing that interferes with my learning is my education.” -EinsteinFrost122585_{April 6, 2008
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