Talk:Haldane's dilemma/Archive 1

Archive 1

Archive 2

Haldane's dilemma

This edit is very rough yes but at least it talks about the basic ideas behind haldane's dilemma. The preceeding article said that the dilemma comes invalidly assuming that only one allele can be substituted at a time. How can someone with such a poor understanding of the dilemma write this article? Even the cited rebuttals to the dilemma do not take this approach because they at least understand the dilemma. Can the person responsible for reverting the article please talk to me in the discussion section. Cheers

Background

Haldane's mathematical model of the cost of natural selection printed in 1957 in the journal of genetics [ref] is simply an attempt to model how quickly a new beneficial allele would replace the old in a given population. This "cost" can be interpretated as the number of generations required to replace this allele. For example a cost of one means that (in this model) a parent with the beneficial allele reproduced the entire population and the rest of the population died out within the timeframe of one generation. Alternatively a cost of say 40 is interpreted as it taking 40 generations before the old allele is replaced. Haldane's arguments were based on the multiplicative fitness model (link?).

the dillema

The dillema arose when it was found that this cost had a relationship with the initial frequency of the beneficial allele.

that relationship is defined by

• D=-ln(q_0) + other terms

(which is intuitively obvious since it will take longer to reproduce larger populations than smaller). Consider trying to substitute 2 alleles simultaneously. Say in generation T_0

• q_0a = 1/N
• q_0b = 1/N

then for the next generation T_1

• q_0ab ~ 1/N^2 (for large populations)

I find this confusing. It seems that a variable q_0x represents the frequency of a gene x, and q_0xy means the frequency of finding both x and y in one organism (please correct me if I'm wrong). If so, then why does q_0xy matter, particularly when both x and y are extremely rare? Therefore, why does 1/N^2 enter into anything, or 1/N^n (below)?

Now consider trying to substitute n alleles simultaneously.

• q_0a = 1/N
• q_0b = 1/N

.........

• q_0n = 1/N

then for generation T_1

• q_0ab.......n = 1/N^n for large populations.

So the dillema is this. For large populations it takes just as long to substitute n alleles simultaneously as it does to substitute them in series. So the number of substitutions that can be made in say 10^7 years is

• 10^7/(D*|T|) where |T| is the length of 1 generation (note these results are generalized for dominant and recessive genes).

criticisms

Some criticisms of Haldane's dillema are:

• it assumes a constant population size
• soft selection/hard selection....
• it assumes all mutations that are beneficial were initially beneficial (e.g. a neutral or harmful substitution becomes a beneficial substitution after a change in the population's environment)
• it assumes a large population
• who is paying the cost? if I read Haldane's original paper correctly, it comes from the allele that is decreasing. But they are going extinct, so it doesn't matter what costs are for them-except as it modifies the speed the other allele increases.
• others?

recent developments

Walter reMine (link?) recently redefined the substitution cost as "reproductive excess" which is applicable to non-constant population sizes. He also makes the argument for the last two critiques 1. that neutral and harmful substitutions must a) have an upper limit to avoid error catastrophe (link ?) and b) still need to be substituted in and thus incur a higher cost than beneficial substitions. Thus, he argues, there is no reason that neutral or harmful substitutions can increase the total number of possible substitutions in any given time. 2. harmful mutations also spread quicker through a small population thus evolution occurs best at some ideal population P.

Applying Haldane's dillema to human evolution gives the following results: (Assume that all substitutions are point mutations)

• cost:D = 30
• generation length |T| = 20years
• time frame ~ 10^7 years
• maximum number of neuclitides substituded (from some ape-like human-like common ancestor) in this timeframe ~1667

I hope this page gets things started :) cheers

Suggestions for article improvement

First, and most important, there should be a citation and reference for the article in which Haldane introduced the "dilemma".

Second, where does the phrase "Haldane's Dilemma" come from? Who first used it for Haldane's model? And did Haldane himself truly consider it a "dilemma"? If so, did he suggest any way to resolve it?

Third, where did Haldane get his numbers from? Far less was known about genetics back then, and as far as I know, he pulled his numbers out of the air. MrDarwin 17:12, 19 April 2006 (UTC)

Summarize the material in clear laymans's language. Put the eyeglazing math derivation down at the bottom. I understand Wiki to be a general encyclopedia, not a mathematics one. While it is helpful to provide the technical details. Put them down at the bottom in recognition of the frequency of viewers looking for that detail. ie probably 1/1000 viewers.DLH 02:26, 28 June 2006 (UTC)

It's hard to see why anyone thinks that there is a problem

It seems to me that the following statements should be non-controversial. Certainly none of them say anything new, yet they clearly say that Haldane's model was wrong --

1) Most species are pressing against limits of population size imposed by the limited resources of their ecological niche, rather than being on the brink of extinction because of gross maladaptation. That is, Malthus had it right for most species, and not just for human beings: absent resource limitations, populations grow exponentially.

2) Therefore, selection is best thought of as a competition among members of the same species. This competition leads to differing contributions of genes to the next generation: different genes cause differences in mortality, differences in the ability to gather food and rear young, etc. All differences affect the frequency of a gene only through their effect on differences in the contribution of the organism's genes to the next generation's gene pool: call this effect of a gene its "differential contribution".

3) In the critical early stages of progress toward fixation (when a gene goes from one copy in a million to a hundred thousand in a million), favorable differential contributions caused by a gene lead to exponential growth of the frequency of that gene. The rate of the exponential growth depends on the magnitude of differential contribution.

4) Changes in other genes don't have much effect on this process. The rate of change of gene frequencies in a population over generations obviously can't affect how well a particular organism does with the genes it has. Regarding genetic diversity within a single generation, the effect of differentials caused by other genes is merely another aspect of the unpredictability of life. This effect is nothing special, and so it makes no special difference.ǂ

5) Therefore, under ordinary, expected conditions, any number of genes can simultaneously move toward fixation without significant mutual interference.

To simplify, but not too much: If a gene helps an organism, then the gene will spread exponentially through the gene pool of the species. Other genes can do the same without interfering.

Is there a problem here? This all seems obvious and unoriginal to me. It would seem that the only interesting question (though rather academic) is where Haldane went wrong, and why so many people thought that his mathematical model fit the real world.

---

ǂ A numerical example relevant to (4): if a gene were under positive selection with a differential contribution of only 0.03%, then it would approach fixation (99.8% frequency) in a population of 1,000,000 organisms in 100,000 generations. At 20 years per generation (for an unusually slow-breeding species), this would be 2,000,000 years. If 10,000 genes with differential contributions of 0.03% were all at 50% frequency (a high, worst-case number), then the variation in differential contribution from all of them taken together would average about 0.03% times the square root of 10,000 = 3%. In the lottery of life, this genetic variation in success rates would hardly be noticeable. This directly addresses Haldane's argument: small selection pressures are entirely adequate to change an enormous number of genes in a reasonable evolutionary timespan.

Here is what Haldane's Paradox really says!

The previous examples get it wrong. Basically, Haldane's paradox is a bottleneck in probability theory. Haldane, who apparently was not troubled by his model, was simply pointing out that, especially in large vertabrates with long generations and lifespans, evolution could only happen so fast, at least if evolution requires a lot of succcessive fixations of genes.

As Haldane states the problem, there is indeed an issue for evolution. In Haldane's model, multiple mutations are indeed moving towards fixation in the population at the same time. However, each mutation is independent of the others (both functionally and physically, in terms of linkages), and therefore each gene must be paid for in mortality. But the amount of mortality that is available has a practical upper limit (the population itself), and in reality the vast majority of mortality in any population is not related to selection, as Haldane rightly points out.

To understand Haldane's model, imagine that there is a budget constraint in the amount of mortality that is available to pay for favorable selection. Every creature can only die once! And if mutations are functionally and physically independent, then you can only die for one mutation at a time. (Sorry to use economics terminology, but this same model more or less is actually used in economics for other purposes. It is a constrained optimization problem.)

So what Haldane's paradox really says is that only a limited number of genes can move to fixation in a given span of time. Haldane illustrated a scenario (a set of assumptions about mortality, probability, etc.) that would in practice set this rate of fixation at about 1 every 300 generations. In other words, after 5000 generations or so (about 100K years in the case of humans) roughly 16 genes could change to fixation. In fact, this conclusion is independent of the size of the population in Haldane's model. (Of course, in a very small population, factors like genetic drift would be more important.)

Others who have written on Haldane's paradox have relaxed the assumptions Haldane made. In particular, they have shown that the assumption that all mutations moving towards fixation affect mortality independently is certainly wrong. Also, mortality is not the only driver of evolution. Fertility may be far more important. Relaxing those assumptions (and several others) allows more genes to be fixed in 100,000 years, but we are still talking a relatively small number of fixations over time.

Perhaps each mortality event eliminates 5 genes, not 1. That merely increases the number of fixations that are possible several fold. There is still a bottleneck, if we are following this model. What Haldane shows is that, in any kind of classical fixation model, the number of fixations that can occur is constrained over time.

What has really happened in microbiology and evolutionary theory in the last 20 years is that other possible evolutionary mechanisms now seem to be the real drivers of evolution. Recall that at the outset of the Human Genome Project, it was generally assumed that humans have around 100K genes. Today, as the result of the Human Genome Project (and many other genome projects), we know that most vertabrates and complex organisms probably have somewhere in the range of 15K to 35K genes. The complexity of organisms and their evolution turns out to be as much in the expression of the genes as in the genes themsselves.

Haldane wrote at a time when virtually nothing was knows about any genome. Haldane showed that a maximum of perhaps 1000 genes could have been fixed in the time between the human-chimpanzee-bonobo missing link and the present. But Haldane had no idea how many genes there were, or how many might need to be fixed to turn the Missing Link into Homo Sapiens. From the perspective of a half century ago, he simply pointed out one of the hard questions that his own students and successive generations would have to answer.

Today, in 2006, we still don't know how many fixations separate us from those chimps, but it appears that it could easily be a lot less than 1000. Indeed, it may be just a fraction of that number!

Recently, a gene that has been fixed in the last few hundred thousand years was identified. It was a gene that regulated fine motor control and thus speech.

Of course, the Haldane model really assumes that evolution takes place by the independent selection of and replacement of individual homologous (ancestrally and functionally equivalent) genes through natural selection. But it now appears that some of the giant leaps in evolution did not occur by this process at all. Although a major chromosomal mutation (such as the creation of a new copy of a gene or even an entire range of genes) is generally a disaster for the individual organism, a few such mutations appear to take hold. Such changes in the overall structure of chromosomes, when they occur, have probably opened up opportunities for huge changes in genomes, for the emergence of new species. Such large scale mutations allow for rapid change outside the constraints imposed by Haldane's model.

It is best to see Haldane's model as what it is. It's a model, and a really great one too, because we are still talking about it after 50 years. Haldane's paper was one of the greatest theoretical papers of the 20th century, and people will still talk about it in the 22nd and 23rd centuries. What makes it great is not that it was right or wrong. Indeed, it is right, if that's how evolution happens. The model shows what the constraints are. Haldane's model shows how large-scale evolution probably did not happen, and tells us to look in other directions.

Now, would somebody care to rewrite the article, because it is incorrect.

--Metzenberg 03:49, 5 May 2006 (UTC)

I won't be the one to rewrite, as I really don't understand the entire issue. My main concern is that Haldane was almost certainly making his calculations using faulty assumptions.

But one thought regarding population size, mortality, and selection: does selection at the gamete level factor in at all? For example, in a "population" of millions of sperm cells, some (most?) will have mutations that affect their fitness at the biochemical or cellular level. In other words, some will be better than others at reaching their goal (which is fertilization rather than reproduction) and I wonder what impact this has on evolution. MrDarwin 14:13, 5 May 2006 (UTC)

Ok, I can give it a try :-) But don't flame me, if you don't like it. --FreezBee 10:05, 23 May 2006 (UTC)

Hi FreezBee. Actually, Haldane did not say that only one gene could be fixed at a time. Rather, what he said was that if N genes are being fixed, the number of total generations required would be the same whether only one is fixed at a time, or all N are simultaneously being fixed. Also, the 300 generations conclusion was Haldane's rough estimate. I think we are making too much of that rough estimate here. Haldane used that number, and showed what assumptions might produce it, only to demonstrate that there could be a significant bottleneck. It still think the most important thing to understand is that modern genetics is rapidly demonstrating that Haldane's Dilemna is irrelevant, because the complexity of organisms and the differences between species do not depend on there being an large number of mutations over time, a number that might not be feasible by Haldane's Dilemna. This doesn't demonstrate that Haldane was wrong. Rather, Haldane raised a fundamental question that he could not answer in his own time. In this way, he created a framework for others to answer the question later on. --Metzenberg 09:32, 24 May 2006 (UTC)

Hi Metzenberg. Ok, I haven't read Haldane so thoroughly yet to really know, what exactly the problem is, except, as you mention earlier, it's a probabilistic bottleneck. I have taken some inspiration from ReMine's interpretation; but I think he's wrong. I'll try to read some more and see, if I can find a better formulation.

A minor point: my last paragraph (the second to last in the article) was picked up from EvoWiki, and can hardly be based on something Haldane has originally written. And the size given for the human genome is in base pairs, whereas Haldane speaks about genes (the mapping between genes and DNA is somewhat unclear), so the arithmetic is meaningless, I would suppose. Am I right here?

--FreezBee 14:18, 24 May 2006 (UTC)

I have tried to make the arguments clearer - from what I think it all means; I'm far from any expert on these matters :-)

Also I have deleted the last paragraph of the original article:

Haldane was concerned that it would take 300 generations to fix a single gene from initial mutation to ubiquity in a population. While one can debate this figure, it is not the main source of the dilemma. This comes from the invalid assumptions that only one gene can be fixed at a time and that no other changes can accumulate until the ongoing one is fixed. That is why multiplying the number of generations per fixation by the number of total DNA differences makes no sense.

This paragraph would appear to be superfluous now, right?

--FreezBee 16:04, 24 May 2006 (UTC)

Hey I was the one that wrote the original rewrite and was planning to write something more professional. But looks like you guys already did that well done guys! The bit at the end seemed original to me (which i understand to be a problem. Although I'll have to check the EvoWiki article). Besides its hard to see how, based on haldane's method of calculation, changing the number of loci would make a difference. Certainly this is not a well known suggestion for solving the dilemma. Is this based on a new paper? Do you have a reference. Further I suggest a style change. More math is needed. However, I don't want it to become like the error catastrophe article which is impossible to follow for someone outside of that specialist field (e.g. terms like L aren't defined and a couple of rabits are pulled out of the hat). A simple following of Haldane's article won't do either coz thats difficult to follow. The math behind this is actually quite simple so it shouldn't be too hard to make a clearer version than haldane's original (and of course one can stop at the result C~-ln(q) ). Oh and freezbee the dilemma is exactly as someone above mentioned. It takes the same amount of time for N substitutions to occur simultaneously as they do one at a time. Regards - Graham

Hi Graham; sorry, if I have changed the article in a way contrary to your intentions. My own edits are only preliminary, so you are most welcome to change the article to your liking. You appear to know more about it than I do anyway :-) I have no reference for my critique of ReMine's conclusions; but ReMine clearly does change the problem by focusing on single base substitutions than on alleles, and I thought it ok to note that without giving any reference. ReMine gets far more possible differences than Haldane and therefore it's not the same problem. But, as said, you are welcome to rewrite the article, as you see fit. --FreezBee 10:05, 29 May 2006 (UTC)

ReMine does NOT "change the problem" by focusing on single base substititutions rather than alleles. ReMine merely clarifies Haldane's ambiguous wording, in a manner that Haldane would certainly have agreed with. That is, Haldane spoke of "gene substitutions" (wording that was common among evolutionary geneticists at the time), when in fact the "new gene" differs from the "old gene" by merely ONE MUTATION. And according to evolutionary geneticists, that mutation is almost always a single nucleotide. That is a straightforward and uncontroversial clarification of Haldane's Dilemma, not a "changed" problem. The current Wiki essay misrepresents ReMine on this point.

Also, ReMine has been emphatically clear that his argument is NOT about the differences between modern chimps and modern humans. (See http://SaintPaulScience.com/Haldane.htm#chimps-humans), so the current Wiki essay misrepresents ReMine on this point.

The figures used by ReMine come entirely from leading evolutionists (such as Haldane), and ReMine then explains Haldane's Dilemma in a manner the public can understand. That is, using Haldane's figures, all the uniquely human adaptations would have to be explained within a limit of no more than 1,667 beneficial substitutions, (where each "substitution" is typically one nucleotide). The current Wiki essay obscures and buries this simple, central point.

The current Wiki essay amplifies traditional misunderstandings, and overwhelms readers with confusion, especially concerning the cost of substitution. ReMine has emphatically pointed out that Haldane's "selective death" concept (which focuses on the elimination of the previous genotypes) is a major source of confusion. Rather, the cost of substitution quantifies the extra reproduction rate required to make substitutions. ReMine has achieved agreement on that point from leading evolutionary geneticists James Crow and Warren Ewens.

The current Wiki essay is overwhelmed with confusion, error, and misplaced emphasis.

Well, I'll try to look into it - but for future reference, could you please sign your discussion postings, otherwise they might not be considered of relevance. And you are most welcome to rewrite the section dealing with ReMine ;-) Oh, and thanks for the link. --FreezBee 12:16, 30 May 2006 (UTC)

-> Nah that's cool Freezbee you did say "i'll rewrite it but don't flame me if its wrong". But yeah I'm certainly willing to help -Graham

I have actually now read "The Cost of Natural Selection", and can to some changes needed. I'll make those changes now, and you can then see, if there's something you want changed. The paper is rather technical and dense, so it's difficult to have all the little details in the article without making it too technical. ReMine might have gone too far in the opposite direction by ignoring all the technical discussions and focusing only on a few select key numbers. --FreezBee 12:16, 30 May 2006 (UTC)

cool nice changes, although some of the terms need to be made more explicit. eg D=number of deaths but is also the cost, also lambda isn't defined....but it isn't in the paper either.

Hi Graham! I was unsure about how muvh of the technical details to include - the article should be about "Haldane's Dilemma", not an exegesis of "The Cost of Natural Selection". I dis think I made it clear that D = number of deaths = cost of sustitution; but I'll try to make it clearer, if I can :-) As for lambda, I was unsure about how much I should go into details with it. The general assumption is fixation of a gene (whereby, as you write, Haldane in more contemporary language means an "allele"), that is, a move to being homozygous in the entire population. The dufferent values fir lambda indictae how dominant/recessive the allele in question is, and I thought it might make the article too difficult to read, if it became cluttered with discussions about such issues. But I agree that some notes about the meaning of lambda should be included - somehoe. --FreezBee 10:46, 31 May 2006 (UTC)

Rather than say "Robert Williams says the term Haldane's dilemma was first used by ..." might be better to just cite the article (actually he called it the "cost-of-selection dilemma" I'm not entirely sure who first called it the Haldane's dilemma? In anycase Williams is wrong because cost-dillema is used in previous papers eg: Proc. Nat. Acad. Sci. USA Vol. 71, No. 5, pp. 1670-1671, May 1974). Its available http://www.pubmedcentral.gov/pagerender.fcgi?artid=434284&pageindex=1 (since you linked the reference to Haldane's original paper, I'm not sure how to do links).

Thanks for the link. I'll read it and see, if it changes anything - though it's a paper from 1974, and Van Valen's was from 1963. It doesn't appear as if Van Valen's paper is available online, so I haven't read it. --FreezBee 10:46, 31 May 2006 (UTC)

Oh and while the papers said often said gene substitution they actually mean a substitution of a beneficial allele, which at the time was assumed to be a gene. See the cited paper for a direct statement of that in the section "basic assumptions". An allele is more general and can be 1 nucleotide, 10 nucleotides or 50 genes. Remines argument was that a typical new allele (caused by a mutation or duplication) is a point mutation. Actually, even though this paper is old it might provide a good overview of the dilemma.

While the typical new mutant allele will be a point mutation, there is a difference between that and expecting all allele fixations to be based on point mutations. The phenotypic difference between two alleles that differ only in one nucleotide will in most cases be too small to induce any selective pressure. Note that a neutral allele can stay around for a long time and have several point mutations, before it starts climbing towards fixation. --FreezBee 10:46, 31 May 2006 (UTC)

The problem with going to the original paper is its usually explained in difficult terms and even experts have trouble figuring out what they're talking about until later papers explain it in simpler terms (tell me about it I just read a recent paper on counterfactual quantum computation and didn't understand a word of it until a later paper clarified).

Yes, I can relate to that problem myself :-) --FreezBee 10:46, 31 May 2006 (UTC)

But it goes through an interesting discussion of the proposed solutions starting with nuetral substitution theory, removing the assumption of a constant pop, variable selection coefficients, the size of a beneficial allele (I don't see how this one is novel) and a couple of others. The "empirical evidence" that contradicts Haldane's dilemma appears to be in reference to the fossil record: see http://www.pnas.org/cgi/reprint/71/5/1670 . Also the derivation you included was for that of a diploid, which is very relevant but the simpler case might be better? -Graham

I decided to only include the diploid, autosomal case, because it's the one most relevant for ReMine's case of human development. Including the other cases might make it too difficult for readers to figure out, what's relevant and what's not. Haldane's Dilemma as the focus is on it today is mostly ReMine's attempt to use it to deny the possibility of human evolution from apes, and I thought it best to concentrate the article on, what's relevant for that case. --FreezBee 10:46, 31 May 2006 (UTC)

yeah I don't know because the math isn't terribly clear for that particular model in haldanes paper (its harder to follow than the 1st simpler case), I included the step that dq = ... (actually approximately but i couldn't get ~ to work) but Its not clear to me why dq = ....

As for point mutations, is there anything out there that makes an estimate on the average size of a mutation? is it 1.1 nukes? 50? etc. Obviously the typical is 1 so I wouldn't expect the average to be much higher than 1. In anycase I think the last part of the ReMine section is incorrect. Also I noticed he doesn't like comparing modern apes to modern humans because he says "while its interesting to do so it adds too many confusion factors". You mentioned nuetral substitutions, there is an argument that nuetral alleles still have to be substituted in and incur a higher cost due to how it drifts back and forth before the key environment change. You mentioned the Van Valen paper; wasn't that 1st link a link to it? It must not have worked ill try again, http://links.jstor.org/sici?sici=0003-0147(196305%2F06)97%3A894%3C185%3AHDERAH%3E2.0.CO%3B2-0, But yeah your absolutely right the paper I cited was from later sorry. Let me know if you have trouble accessing it. It may only be available to me because Im on a uni server. As for not wanting an exegesis of haldane's paper, I agree and I think it already looks too much like that. Perhaps a greater variety of sources are needed such as the one I mentioned before? Also I like the direct quotes in the first section but the quote in "the cost" section is a bit long winded and perhaps unnesecary. It might be better just to paraphrase that one? Also It would be nice if there was a curt definition of haldane's dilemma in the title. If one wants to focus on the creation-evolution controversy side of things then Haldane's dilemma is that "mathematical models predict that substitution occurs too slowly to be feasable". If one is talking outside that controversy then the dilemma is that "organisms simply cannot cope with environmental changes that require rapid fixation of more than one allele" (so i guess it depends on what should be focused upon). Either way the argument can be summed up that "It takes, on average, the same amount of time to fix N alleles simultaneously as it does to fix them one at a time". I think thats a nice way of saying it. Also the introduction contains "consists of a poor model of natural selection that makes invalid assumptions". I assume thats an artifact of previous edits? Whether or not Haldane's dilemma is solved or not, I don't think that it every rose out of a poor model since the models are still used today. Its genuinely an unsolved problem just like the small vacuum energy is an unsolved problem in cosmology. Oh yeah you do mention the change of environment definition of Haldane's dilemma in another section but I think it belongs in the introduction as well? Anyway here's my proposal for the intro

Haldane's Dilemma refers to a paradox introduced by J. B. S. Haldane concerning the efficacy of natural selection. The dilemma is based upon the substitution cost introduced by Haldane in his 1957 paper The Cost of Natural Selection and refers to the fact that it takes the same amount of time to fix N new alleles simultaneously as it does to fix them one at a time. This result has been used to argue that environmental changes that require a rapid substitution of a number of alleles would cause extinction. It has also been used as a creationist argument to question the feasability of slowly reproducing species.

Any other applications should be included here as well. I wonder if anyone has talked about its application to artificial selection?

No one objected to this intro, and the current one is FAR from NPOV, so I am inserting it. Mdotley 16:28, 17 August 2006 (UTC)

The intro is a remnant from earlier, and you're welcome to change it. As for size of mutations, that's less relevant, because it's the difference between the "ruling" allele and the one moving towards fixation. If the fixation isn't really a fixation, perhaps only 99%, then it's possible that a few alleles that deviate by more than one nucleotide linger around. By I agree that the section about ReMine needs some reworking - it's true that ReMine's point isn't the difference between modern humans and chimps. I'm working on a change here.

As for the various papers dealing with Haldane's Dilemma, I'm afraid I don't have time to write anything about them, so if anybody else has some spare time, there's a place to invest it :-)

I changed the name of the section "The Dilemma" to "Origin of the term 'Haldane's Dilemma'", because it was really my intension with that section.

I can't access the link you gave, because it's on JSTOR, to which I don't have access. But if possible, could you mail it to me at pwemail@hotmail.com?

--FreezBee 10:51, 1 June 2006 (UTC)

remine was comparing humans to the apelike human like common ancestor. Its the comparison between modern apes and modern humans that he "avoids because while its interesting it complicates things" (I think thats the reason). So I changed the part of remines section where you said that he wasnt comparing humans to their ancestor

I edited the section bearing my name. I believe it reads easier now, and more accurately represents my position. -- Walter R.

My omission from the text of the comparison between humans and chimpanzees was to fit with ReMine's currently stated position, and since I haven't read The Biotic Message, I don't know, what his position was back then - only what he writes in the "Table of Contents". If it can be documented that ReMine in The Biotic Message compared humans to chimpanzees, the text should of course note that, and also that ReMine has changed his position. But it would need to be documented.

Walter, I'll try to make your claim as stated at EvoWiki more correctly reflect your position, and, if possible, also make Mark Isaak change the formulation of his CB121 claim. There is no reason to refute non-existing claims.

--FreezBee 12:23, 2 June 2006 (UTC)

ReMine's argument was NEVER focused on comparisons between living humans and living chimps. Rather, that comparison is the way evolutionists insist on misrepresenting ReMine's argument. Marks Isaak's CB121 essay misrepresents ReMine that way (and in many additional ways too). ReMine has not "changed his position". ReMine's argument was ALWAYS focused on the limited number of substitutions available to create human adaptations -- 1,667, according to Haldane's calculations. -- Walter R. P.S. I edited the section bearing my name, but that apparently occurred coincidently with someone else's update, so mine got over-written. I'll post mine again.

DNA differences vs. expression differences

On his webpage Haldane's Dilemma - the trade secret of evolutionary genetics, Walter ReMine in the section "Regulatory genes?" writes

Mutations to regulatory genes can sometimes have a large biological effect. So, are mutations to "regulatory genes" a solution to Haldane's Dilemma?1 Brief answer: It scarcely affects Haldane's Dilemma.

Evolutionists do not get to choose the substitutions, say, as "mutations to regulatory genes" or as "mutations with a large biological effect" or as "gene clusters". Rather, evolutionists must accept what nature doles out – and we can observe what nature doles out. The issue is fundamentally empirical and observable, not one of telling stories about regulatory genes.

However, ReMine might be wrong here. At AiG i happened to find this article: The differences make the difference — differences in gene expression distinguish humans from other primates.

According to the article, which references an article in Nature, the main differences among primates actually is not in the DNA as such, but in the expression of genes. The article explains it like this:

While the DNA sequence of a gene specifies the amino acid sequence of the protein, the expression level is the amount of the protein that is made. In other words, the DNA sequence spells out the code for producing a specific protein whereas the expression level is the number of copies that will be produced. The amount of protein that is produced can make a profound difference, and in some cases a more important difference than a change in the DNA sequence.

There is a table shown in the article of comparisons between humans, chimpanzees, orangutans, and rhesus macaques. The table does show differences in expression levels that are somewhat compatible with the supposed phylogenetic relationships between these primates, though the data are not conclusive. But they are certainly worth a look - and maybe ReMine needs to uodate his position :-)

cheers

--FreezBee 12:59, 2 June 2006 (UTC)

Let me say it again: ReMine's argument is NOT about "comparisons between humans, chimpanzees, orangutans, and rhesus macaques". You are (implicitly) trying to alter ReMine's argument. ReMine's argument is about the limited number of substitutions available to explain human evolution -- 1,667 according to Haldane's calculations.

Beneficial, remember, substitutions - substitutions that increase juvenile survival. And not according to Haldane's calculations, but according to Haldane's estimates.

--FreezBee 11:12, 3 June 2006 (UTC)

Also, your argument about "supposed phylogenetic relationships" is entirely pattern-based (in the classical style of evolutionists). Whereas ReMine's argument focuses on experimental demonstrations -- biological changes we can actually observe taking place. He notes that "evolutionists must accept what nature doles out – and we can observe what nature doles out. The issue is fundamentally empirical and observable, not one of telling stories".

ReMine points out that the substitutions cost the same -- whether they are a point-mutation, a deletion, an insertion, a gene-duplication, the relative ordering of genes on a chromosome, the placement of genes onto different chromosomes, or a "regulatory gene" as opposed to a proteinous gene. (The main exception is recessive substitutions, which cost a lot more.) ANY TIME an evolutionary scenario claims a substitution, there is a cost requirement. Each beneficial substitution has a cost (all approximately cost the same). Your references to "gene expression" are largely irrelevant, because it does not alter the limit -- 1,667 substitutions. Evolutionists do not get to CHOOSE what the substitutions are, rather they must accept what nature doles out, and we can observe what nature does out, for example, in the beaks of Galapagos finches. -- Walter R.

Ok, my argument was based on that it's not sufficient to compare differences in DNA. Two species may produce the same proteins and yet be different due to producing them in different quanta. Try the following experiments: mix one liter of water with one milliliter of alcohol and drink the mixture; then mix one liter of alcohol with one milliliter of water and drink that mixture. You should observe some difference in the effect - or maybe someone observing you should.

Ans, as far as I am informed, we can observe humans, chimpanzees, orangutans and rhesus macaques today, and it appears that they differ more in expression of genes than in genes.

--FreezBee 11:12, 3 June 2006 (UTC)

You're missing the point. A change in "gene expression" is still a change. It still requires a substitution. It still tallies toward the limited number of substitutions. (Which is 1,667 according to Haldane's calculations.) You cannot get around the reasoning: ANY TYPE of substitution is STILL a substitution. It still has a cost of substitution. There is still a limit, and your argument has not changed that limit one iota.

Moreover, evolutionists do not get to CHOOSE the type of substitutions that nature provides. For example, let us assume that living primates differ mostly by gene expression. Fine. That merely aggravates the evolutionists' problems if nature provides few of those substitutions. If MOST beneficial substitutions are not for gene expression, then your argument has made the evolutionists' problem worse than before. If not, then the problem remains the same as before -- Is 1,667 enough?

You are obscuring the central issue, which is quite simple: Is 1,667 beneficial mutations -- of the types observed in nature -- enough to transform a simian into a human? -- Walter R.

Remember that beneficial mutations only relate to increased juvenile survival, so we would have to check human juvenile survival against ape juvenile survival. However, the main difference between apes and humans is that humans can change social organization, invent new technology, and so on - humans are less tied to their genes. So rather we would have to compare human genome to, say, chimpanzee genome and see at how many loci these differ. Assume there to be differences at N loci. The difference between humans/chimps and their supposed last common ancestor would be somewhere between N/2 and N.

--FreezBee 09:13, 6 June 2006 (UTC)

Re. Walter ReMine and Haldane's Dilemma

Now, now, Walter, that is not kosher. You know that WikiPedia is commotted to an NPOV, and while I appreciate that you contribute to this article, I doubt that copying your own material into the article would count as an example of an NPOV.

On your compuserve webpage Haldane's Dilemma you write:

Along the relevant primate line, our supposed pre-human ancestors had an effective generation time of 20 years. (I quote sources and details in my book, so I'll spare you here.) Imagine ten million years ago -- (that is two to three times the age of the alleged chimp-human split) -- that's enough time for 500,000 generations of our presumed ancestors.

Imagine a population of 100,000 of those organisms quietly evolving their way to humanity. For easy visualization, I'll have you imagine a scenario that favors rapid evolution. Imagine evolution happens like this. Every generation, one male and one female receive a beneficial mutation so advantageous that the 999,998 others die off immediately, and the population is then replenished in one generation by the surviving couple. Imagine evolution happens like this, generation after generation, for ten million years. How many beneficial mutations could be substituted at this crashing pace? One per generation -- or 500,000 nucleotides. That's 0.014 percent of the genome. (That is a minuscule fraction of the 2 to 3 percent that separates us from chimpanzees).

That's not a difficult calculation, yet it immediately reveals a problem. Is 500,000 beneficial nucleotides enough to explain the origin of humanity from some chimp-like ancestor?

In the second paragrapf yuo mention a scenario where a substitution per generation occurs - 500,000 nucleotides. If you have read the text you changed, you know that Haldane did not calculate with nucleotides, but with whole genes, so your scenario is of no relevance for Haldane's Dilemma.

And please note, what you write in the parentheses at the end of that paragraph. Yes, it's a paranthetical remark, so I won't conclude that you consider a comparison between humans and chimpanzees vital - but some people might misunderstand that remark.

--FreezBee 08:40, 3 June 2006 (UTC)

The website you refer to as mine, is not mine. You refer to its writings as mine -- on what basis do you do that? You don't have any scholarly basis for it. Why are you so concerned with obscure matters -- such as "parenthetical remarks" on obscure, untraceable websites? Is that the best you can do? Have you lost any sense of scholarship? Why not refer to my book, or my papers, or my website.

Interesting. I admit that I was in doubt about who was the author of the webpage. It begins with "

Walter Remine, author of "The Biotic Message", notes: ...

So, it would seem that it was someone else writing about you. But in these "notes" we have this paragraph:

Evolutionary genetics has trade secrets too. The major one is Haldane's Dilemma, a problem discovered in the 1950s by the famous evolutionary geneticist, J.B.S. Haldane. Journals discussed it through the 60s, and ignored it thereafter. Evolutionists never publicly solved it, rather they brushed it aside. Here are my claims: ...

I have highlighted the word my. This word is the FIRST PERSON singular possessive pronoun. So it would appear as if we are supposed to think that the "notes" are a quote. Can you confirm the authenticity of the quote? If not, I would suggest that you contact the owner of the page; this is apparently possible at rossuk12@hotmail.com.

--FreezBee 09:38, 6 June 2006 (UTC)

If you were seriously concerned about what people "might misunderstand", then you would be deeply concerned about the current Wiki essay, whose purpose is apparently to bury Haldane's Dilemma as much as possible from public view, rather than reveal it.

In what way does "the current Wiki essay" bury Haldane's Dilemma? --FreezBee 09:38, 6 June 2006 (UTC)

Haldane did not "CALCULATE with 'whole' genes", rather he CALCULATED a beneficial SUBSTITUTION RATE (and used the traditional jargon of his profession in speaking of "gene" substitutions). The 'new gene' is identical to the 'old gene' except for one new beneficial mutation -- NOT an entire block of new DNA, not thousands of nucleotides. This point is utterly uncontroversial, and one that Haldane would certainly have agreed with. -- Walter R.

The new gene (allele) is different by the amount at which it is different. This may be one nucleotide, and it may be more than one. Since there are four different nucleotides, and three nucleotides make up a codon, there are 4^3 = 64 different codons; but there are only 20 different amino acids, so far from all single nucleotide mutations will make any difference. --FreezBee 09:38, 6 June 2006 (UTC)

The talk.origins website (essay CB121) misrepresents me thoroughly. For an itemized listing of the misrepresentations see here -- Walter R.

I agree with you here; but right now I see more serious problems, so I'll have to leave the solution of that problem to you.

--FreezBee 09:38, 6 June 2006 (UTC)

I removed text that someone added, because the text is not NPOV, and also misrepresents ReMine:

• Haldane's Dilemma is appropriately called a "dilemma": (1) because that's what evolutionists named it, and (2) because it remains an unsolved problem of central importance in evolutionary genetics. Evolutionists have not shown that 1,667 beneficial mutations is sufficient. The text misrepresents that argument as "illogical".

==

WHY is is appropriately called a 'dilemma'? Because the early geneticists dismissed or did not know about small mutations with large effects? Why is it an 'unsolved problem' for evolutionary genetics? Is it a problem for evolution as such? Or just the modeling? Creationists have not shown that 1667 mutations is insufficient, they merely assert it so. That argument is illogical, because it is premised on a personal belief, not evidence, that 1667 is too few. Walter ReMine knows this, he just edits these pages every time someone points out his errors.

=======

• The text misrepresents ReMine as believing "that this ape-like creature lacked all traits that humans possess".

• The text misrepresents that ReMine "does not indicate what the starting point was". In reality, ReMine gives the starting point as ten million years ago, and lists various traits that evolutionists claim arose during that time. 4.158.234.23 18:07, 5 June 2006 (UTC) Walter R.

—====================

The text misrepresents ReMine as believing "that this ape-like creature lacked all traits that humans possess".

If Walter ReMine does NOT believe this, then why is he so fixated on Haldane's model not being able to account for all of the 'human' traits that ReMine insists had to be derived in 10 million years using no more than 1667 fixed beneficial mutations?

What human traits had to be accoutned for, Walter ReMine? And WHY is 1667 mutations too few to account for them? You never say - in your book, in your many internet forays, etc. You just ASSERT that this is so. Why no substance? ReMine makes some fairly silly allusions to things like appreciation of music, yet never explains why that is a uniquely human trait, how many mutatins he thinks such a trait would need, etc. ReMine's "arguments" here are uninformed personal opinion presented as supported facts.

========

Your post is a mix of confusions. For example, in the cost literature, "q" is not the frequency of the substituting allele, and q does not have "exponential growth". Rather "q" is the frequency of the previous allele being eliminated, and it is decreasing. Also, you use two phrases that are undefined by you and by the literature, namely: "a limited capacity for change" and "the cost to every death" -- both of those are undefined notions. Also, you say "the real problem starts with the assumption that a second trait can only emerge by slowing the emergence of an already evolving trait". No doubt such an assumption would further slow the substitution rate, but Haldane did not make that assumption. Under Haldane's model, given the same amount of reproductive excess (namely ten-percent), the substitution rate turns out to be the same whether the substitutions occur end-to-end, or concurrently (overlapping in time in arbitrary ways) -- the substitution rate doesn't slow (as you suggest), rather it remains the same. Lastly, Bruce Wallace's book ("Fifty Years of Genetic Load") and his arguments about Haldane's Dilemma are based on the concept of "genetic load", which ReMine claims is a needless source of confusion. 4.158.234.215 03:23, 6 June 2006 (UTC) Walter R.

Text moved

I have moved the text of this section to its own page. The reason for this is that I find it best to separate Haldane's Dilemma in general from Walter ReMine's treatment of that dilemma.

--FreezBee 10:11, 6 June 2006 (UTC)

Wow a lot of activity took place while i was gone. I think a summary of ReMine's section is in order with a link to the extra page (ie i don't think remines section should be removed entirely). There will probably need to be quite a few extra pages to explain all the issues, even still i think a summary is in order. I still think an explanation for lambda and dq is in order because the math is too hard to follow otherwise. Also I noticed my intro got changed? It annoys me that someone not contributing to the discussion would change the intro, if he had a problem with what i wrote, i wrote it in here first so that people could discuss its factual merits, NPOV and writing style. Never mind, the intro needs to be changed, any suggestions as to what to? - Graham

FreezBee, you deleted the entire section on my material. But previously you were content enough with that section, when it thoroughly misrepresented me. Indeed, you wrote much of it. 4.158.231.180 06:35, 7 June 2006 (UTC) Walter R.

I didn't delete it, just moved it. Your entire text is there. I simply found that to be the best solution for the moment. You are free to disagree; but the problem is that your text is your POV and therefore not NPOV. How likely is it that you will come up with any critique of your own opinions?

As for misrepresenting you, when you made us aware that you hadn't compared humans to chimpanzees in The Biotic Message, I changed my text accordingly and even supplied a link to your webpage. Ok, someone changed the text to again contain the claim that you had compared Humans to chimpanzees, and you reverted that change. After that, I made a few fixups, including correcting "gene substitutions" to "substitutions" to have the text better reflect, what appears to be your position. How is that misrepresenting you? If you felt misrepresented, what hindered you from writing that on this Talk page? Instead you replaced the entire text with your own!

Graham, as mentioned above, I have moved ReMine's text to its own page, a solution that I under the current circumstances find to be the best. You can read it there, and you can, if you want to, write a condensed version of it in the intro section of that page.

Walter, the current intro section of this article is better than what you wrote earlier. It's not quite NPOV (Haldane didn't mention human evolution); but I'll leave it as is for the moment.

--FreezBee 08:49, 7 June 2006 (UTC)

Re. Intro section

The intro section states:

He [Haldane]calculated that organisms with low reproduction rates, such as cows, could substitute new beneficial mutations no faster than about 1 per 300 generations.

The word "calculated" is suggested replaced with "estimated". The paper ("The Cost ...") contains no calculation of the number, only an estimate of 30 as the average cost multiplier and a suggestion of 300 as the minimum number of generations.

Later scientists saw the meaning of this limitation.

That's a claim - it's still disputed exactly, what "the meaning of this limitation" really is.

For example, starting at a time about ten million years ago, the human lineage could substitute no more than about 1700 beneficial mutations.

Simce only one person is known to have advanced this example, it would be more appropriate to mention that elsewhere.

It is undisputed that Haldane’s calculations, if correct, would indicate such a limit.

Undisputed?

The problem is, can all the uniquely human adaptations be explained within that limit? Such adaptations would include: the tripling of brain size, upright posture, hand dexterity, vocal speech, language, distribution of hair, and appreciation of poetry, to name a few.

Haldane's beneficial substitutions were those who increased juvenile survival, please keep to that.

While most evolutionary geneticists conclude the problem is solved, they disagree about the solution.

I would consider this to be an irrelevant claim. In general this paragraph would be more appropriate in an intro on the page Responses to Haldane's Dilemma.

Much of the controversy arises from the central concept, known as “the cost of substitution,” derived in terms of “genetic death”, and frequently reinterpreted as “substitutional load.”

Too early introduction of technical terms. The averahe reader has no chance of knowing, what this is all about.

A recent clarification claims that the cost of substitution is most clearly defined as a required reproduction rate. In this way, even the fundamentals of Haldane’s Dilemma are in dispute.

Someone might interpret this as thinly-veiled propaganda for a particular author's POV. On sci.bio.evolution (Haldane's Dilemma) that same author received the following moderator notice:

moderator's response: The point is, [name of author], since your book is self-published and available only from you, you are the SOLE SOURCE of non-misrepresentations (sorry for the strangled syntax) of your book. It is simply pointless to suggest that various arguments are addressed in your book, when we are forced to pay you to read said tome. It is incumbent on YOU to make your argument. Thus far you have not, though because I am almost supernaturally patient, I am allowing this thread to stagger on. For now.

As this moderator notice - from october 27, 2000, indicates, the author in question is unwilling to seriously engange in debates over his claims, and I would therefore suggest that references to these claims NOT be made in an intro section, where the average reader might think they refer to something which is generally agreed upon.

best regards

--FreezBee 15:07, 7 June 2006 (UTC)

==

Wiki readers have a right to know what Haldane’s Dilemma is. It is wrong (and not NPOV) to remove any clear statement of the problem from the intro, as some people seem intent on doing. Rather, they seem intent on burying the problem.

It also mistaken (and not NPOV) to present the issue as harmoniously resolved, as some people seem intent on doing. Haldane’s Dilemma is controversial, with evolutionary geneticists contradicting each other even on the basics of the problem, as well as on their “solutions.”

For example, starting at a time about ten million years ago, the human lineage could substitute no more than about 1700 beneficial mutations.

You are mistaken that “only one person is known to have advanced this example” (a limit of 1700 beneficial mutations). Many people now see its importance and advance it. It follows easily from Haldane’s calculations. No evolutionary geneticist has disputed that Haldane’s speed limit, if correct, would indicate such a limit. You are mistaken to suggest otherwise. You are mistaken to suggest a dispute exists where it doesn’t.

Haldane did not merely “estimate” – he derived formulas and calculated in great detail. You are mistaken to pass off his material wholesale as mere “estimate.”

You are mistaken that Haldane's beneficial substitutions were “limited to those that increased juvenile survival.” Though Haldane used juvenile death as an apt tutorial example, he repeatedly stated that the selective deaths were eliminations or their equivalents in reduced fertility – which clearly is not limited to juveniles.

You object to “Too early introduction of technical terms,” which is funny, since the current Wiki essay immediately overwhelms readers with lengthy minutiae that are needlessly tedious, obscure, and confused. That should be avoided:

1. Haldane’s math is very technical, and not essential to understanding Haldane’s Dilemma. Rather, the speed limit of one-per-300-generations is the essential conclusion from Haldane’s paper. And this result should be put in terms readers can readily understand. It is not NPOV to bury this result.
2. Haldane’s version is one version of the problem known as Haldane’s Dilemma. Indeed, his explanation is not the clearest, and is opaque in places – and progress did not end with Haldane. Evolutionary geneticists have given various illuminations of Haldane’s analysis – some of them clearer than Haldane’s. It is mistaken to pretend that Haldane’s text is the clearest available, as that would just be another attempt to bury Haldane’s Dilemma in confusion. In other words, use the clearest available explanation of Haldane’s analysis, and that is not Haldane’s. If you want to focus intensely on Haldane’s detailed text and math, then it belongs on a different page.

For those reasons, I recommend moving all the following technical material together to a separate page on Haldane’s analysis:

• The Substitution Cost
• Selection Intensity
• A Mathematical Model for the Cost of Diploids
• The Magic Number 300.

A recent clarification claims that the cost of substitution is most clearly defined as a required reproduction rate.

You are mistaken to call that a “thinly-veiled propaganda for a particular author’s POV.” The paper was peer-reviewed by evolutionary geneticists, James Crow and Warren Ewens, who acknowledge it is correct. If you want to debate the matter, then do so, or perhaps you could take it up with them, rather than indulge in a personal smear campaign against me. 4.158.231.251 05:23, 8 June 2006 (UTC) Walter R.

==

I do agree with you on some of your points here; e.g. that Haldane's paper (The Cost of Natural Selection) is a very technical paper, and that a somewhat "softer" introduction is warranted. I don'r have the time or qualifications for this and would be happy, if you - or someone else - could do this. But I know and you know that your TJ article is in dispute (ask Joe Felsenstein for his opinion), and therefore it should not be mentioned as a clarification, as if everybody considers it to be that.

It is true that Haldane only used "juvenile deaths" as one example of "selective death". Translating Haldane's calculations to physical terms is not easy from Haldane's own words. And a clarification would certainly be in place; but one translation is not necessarily the only possible translation. The term "selective death" is vague, which also means that any argumentation applying a more precise term would need its own calculations.

Haldane did indeed make calculations, several of them. But the number 300 is not a calculated number; it's a suggested number. It quite simply is.

As for indulging "in a personal smear campaign against" you, please note that I did not in the above refer to the "particular author" as Walter ReMine, so it wasn't against you; it was simply a notice that the "particular author" has a record of bad behavior. Nothing forced you to admit that you were both the author of the new intro and the "particular author".

Walter, to boil what I am saying down to, what I mean: I acknowledge that you know more about Haldane's Dilemma than I do; but I also know that you have an agenda with Haldane's Dilemma and the number 300, and that agenda is not NPOV.

Anyway, to see that not all evolutionists have forgotten good ol' Haldane, I suggest you have a peek at Leonard Nunney's 2003 article The cost of natural selection revisited. It's quite interesting; I promise.

--FreezBee 09:41, 8 June 2006 (UTC)

FreezBee – You engaged in a personal smear campaign against Walter ReMine. That fact is not reduced by your stated belief that he was not here to defend himself. If anything, that makes it worse than before. And no one forced you to do it anonymously.

Your latest attempt to smear him of unspecified “bad behavior” gets worse still, considering that ReMine published a clarification of Haldane’s Dilemma, which evolutionary geneticists, James Crow and Warren Ewens, acknowledge is correct. Again, if you want to debate ReMine’s paper, then do so, or take it up with Crow & Ewens – but your continued smear campaign is out of place.

The fact that Joe Felsenstein contradicts Crow & Ewens is one of countless unresolved fundamental contradictions concerning Haldane’s Dilemma. It is wrong (and not NPOV) of the current Wiki essay to portray Haldane’s Dilemma as harmoniously resolved, as some people here seem intent on doing. Haldane’s Dilemma is controversial, and unresolved, even at its fundamentals.

moved to I think, I suggested

Ethics

Yeah theres been some unethical activity here. Duncharris is just being rude with his reverts. But a lot of edits have been made without discussing them first. I agree that Remine shouldn't have replaced the Remine section without talking about his changes here first (although in his defence I recall Freezbee telling him he was welcome to change it but as always changes should be discussed here first) also Freezbee you should have discussed in here before making the stunt of moving the ReMine section to a new page, i can certainly see why Remine was cut. It looked to him like you were happy to have a page here that had errors that were clearly POV (and erroneous) yet when a page that was factually correct, but perhaps too long and POV, was introduced it didnt just get edited, it got moved completely. Thats double standard.

My reason for the move was that the "Walter ReMine and Haldane's Dilemma" section had become so large that it would be more appropriate to have its own page, and that a discussion of that section would be a separate issue from the discussion of Haldane's Dilemma in general. I feared that we would run into an "edit war" that would didturb work on the main article. I might have judged wrongly, and I apologize, if I did; but it appeared to be the most proper solution at the time, with a possible reintegration later.

--FreezBee 09:51, 9 June 2006 (UTC)

Pending Changes

Ok so walter suggests a new page for the technical treatment, what do people think of that? It seems like a nice idea to me. Freezbee seemed to suggest she wanted revision ( ;-) ) of Walter's section. It seems reasonable to have a small summary of Remines claims with a link explaining it in detail. I wanted a intro and a couple of clarifications in the technical details. Freezbee, the link you cited had a nice summary which can be paraphrased The direct quote is: "Haldane (1957) recognized that his cost translated directly to a maximum rate of evolutionary change. The cost of natural selection can be defined as the number of deaths (in units of N) required for an allele substitution (Grahams note I prefer the second definition, its easier to understand but both are equivalent), or as the minimum time (in generations) between allelic substitutions that is compatible with population persistence. Using Haldane's (1957) approximation that the cost of substituting one allele for another was about 300N deaths, the the maximum rate of evolution was one substitution per 300 generations" (The talk about defining cost as number of deaths is a mathematical artifact, the second definition is a more physical definition). As for Remines section perhaps. "In (year) Walter Remine wrote a paper on the cost of natural selection with the intention of clarifying cost theory. His paper discusses traditional solutions to Haldane's dilemma including nuetral substitution theory, genetic load and genetic death. His equations reduce down to Haldane's original equations. ReMine's book, The Biotic Message, applies Haldane's speed limit to human evolution. Remine argues the speed limit of one new allele per 300 generations to the case of human evolution to show that in the last 10 million years no more than 1667 new alleles could have been fixed. Remine then writes that in standard evolutionary genetics, the new allele differs from the old by a single nucleotide. Thus, according to Remine, all new traits from "speach, walking upright" to the "appreciation of music and poetry" need to be explained within this limit of 1667 nucleotides. Haldane's model of course is still a hot topic for current research and there have been many proposals to lower the cost. For a more detailed explanation a defence of Remines assumptions and his views on the current proposed solutions to Haldane's dillema see (links). Criticism of Remine can be found (links). A summary of some of the proposed solutions to Haldanes dilemma can be found (links, perhaps future articles on Wikipedia)." Hows that? Are they Factually accurate? (it generally speaking will be but i gaurentee you i missed something) Is it NPOV? Is it a good length? - Graham

I noticed the link to walters page has now been voted for deletion? I hope naming it "Walters Soapbox" was not an underhanded method to get rid of it entirely (as soapboxes are specifically talked about in the wiki policy)

Graham, I second your suggestions. People can always later discuss on the talk page, whether it's factual or not, and whether something is missed or not.

As for whether naming Walter's text "Walter's Soapbox" was "an underhanded method to get rid of it entirely", the answer is: no! I wasn't aware that soapboxes are mentioned in the wiki policy, and Walter - and anyone else - was free to rename the section.

--FreezBee 09:58, 9 June 2006 (UTC)

FreezBee – Your naming the section, "Walter's Soapbox", was clearly pejorative, and not remotely neutral POV. You were "free to rename the section" – but you didn't. Don't blame your actions (and your smear campaign) onto me. 4.159.23.70 05:36, 10 June 2006 (UTC) Walter R.

"My choice of name "Walter's Soapbox" was to inform ReMine about my - of course not unbiased - view on his text. If he wanted to, he was free to call it something else. " yeah thats just a bit provocative-Graham

Remines Section

Ill save edits of the introduction for later, but heres my current revision of remines section, If no one objects to it I'll cut and paste it in the next couple of days

"In (2004) Walter Remine wrote a paper on the cost of natural selection with the intention of clarifying cost theory. His paper discusses traditional solutions to Haldane's dilemma including nuetral substitution theory, genetic load and genetic death. His equations reduce down to Haldane's original equations. ReMine's book, The Biotic Message, applies Haldane's speed limit to human evolution. Remine argues the speed limit of one new allele per 300 generations to the case of human evolution to show that in the last 10 million years no more than 1667 new alleles could have been fixed. Remine then writes that in standard evolutionary genetics, the new allele differs from the old by a single nucleotide. Thus, according to Remine, all new traits from "speach, walking upright" to the "appreciation of music and poetry" need to be explained within this limit of 1667 nucleotides. Haldane's model of course is still a hot topic for current research and there have been many proposals to lower the cost. For a more detailed explanation and a defence of Remines assumptions and his views on the current proposed solutions to Haldane's dillema see (links). Criticism of Remine can be found (links)."

I can't see anything about that which isn't sucinct factually accurate NPOV and misrepresenting of Remine. Its short enough to stop it from dominating the main article yet it correctly explains everything he argues without adding bias to whether he's right or not. What do you think

It's certainly ok with me as a start. A few notes though, ReMine's paper is, I believe from 2005. He might of course have written it in 2004; but it was published in TJ 2005. And it's the neutral substitution theory. But apart from those notes, I would say it's a fair compromise.

--FreezBee 09:46, 12 June 2006 (UTC)

Well, what do you say, Walter? Is it ok with you? --FreezBee 10:41, 13 June 2006 (UTC)

The proposed section on ReMine has various faults, errors, and insufficiencies that can be corrected. Its biggest problem, by far, is that it is written with apparent disregard for the remaining dreadful essay. The current essay:

• Conceals Haldane’s Dilemma from view, rather than reveals it.
• Falsely conveys that the problem is harmoniously resolved.
• Misleadingly claims the problem is based on “invalid assumptions,” when Haldane’s assumptions were wildly unrealistic IN FAVOR of evolution, and consistent with today’s practice of evolutionary genetics.
• Conceals the numerous unresolved contradictions between evolutionary geneticists, even on the fundamentals of this issue.
• Obscures and confuses the mechanics of the speed-limit (and the cost of substitution), when clearer understandings are available.
• Mis-states the creationists’ point of view.
• Wearies readers into oblivion with needless emphasis on tedious derivations.
• Over-emphasizes issues that are virtually irrelevant, such as “The origins of the term Haldane’s Dilemma”.

The current essay should be condemned. It is in that light that the proposed section on ReMine is mis-written. That is, given the systematic failings of the current essay, the section on ReMine is inadequate, both in emphasis and coverage. It is not a sufficient remedy or provide overall NPOV. 4.158.231.145 08:43, 14 June 2006 (UTC)Walter R.

• Oh, well, Walter - in what way does the essay conceal Haldane's Dilemma from view? What is there to reveal? Doesn't the essay make it clear that Haldane's Dilemma is about a limit to have fast evolution can occur, if a population is to survive as well? In the essay I mention the case of the pepered moth that Haldane mentioned in The cost of Natural Selection. As Haldane wrote about this case, then during around half a century a rare allele (for dark moths) replaced the most common allele (for light moths - a move from less than 5% to 95% in the period 1848 to 1898 according to Kettlewell. But Haldane wrote that if more than one locus had been under the same severe selection simultaneously, then the peppered moth made have become extinct. I mention this in the essay, and it is the same line of argumentation Van Valen used, so I have to admit that I completely fail to see your problem. Please explain further.
• In what way does the essay falsely convey that the problem is harmoniously resolved? There is a limit to the speed of evolution, nobody denies that - the disagreements are rather about, how hard and how low the speed limit is.
• Haldane’s assumptions were hardly "wildly unrealistic IN FAVOR of evolution", for instance the assumption that there is rarely a synergistic (positive epistasis) between alleles at different loci.
• The essay does not conceal "the numerous unresolved contradictions between evolutionary geneticists, even on the fundamentals of this issue"; it simply doesn't mention them. My idea with this article was to present, what Haldane actually wrote, which ought to make it easier to figure out to what extent later commentators actually address Haldane's Dilemma or some other dilemma, or in what ways parameters and terminology are changed. I had considered addressing the responses to Haldane's Dilemma to be more appropriate for a separate article, so the reader wouldn't get things mixed up and not really know, what was Haldane's Dilemma, what was Kimura's Dilemma, what was ReMine's Dilemma, and so on.
• Regarding the mechanics of the speed-limit, you must remember that we are dealing with Haldane's Dilemma, and therefore the mechanics should be Haldane's - the assumption of an evironmental change, a rare allele moving towards fixation due to differential survival.
• Where is "the creationists' point of view" mentioned? Are you, Walter ReMine, all creationists?
• The "emphasis on tedious derivations" may be wrongly placed; but it should be there. One of Haldane's point was a focus on quantitative analysis in population genetics. And, anyway, how are readers to relate to the number of 300 generations, if it simply pops out of thin air?
• What is wrong in informing people about the “The origins of the term Haldane’s Dilemma”. In a comment it was requested that the essay mentioned something about this. And it's quite common for encyclopedias to mention the origin and history of a term in the encyclopedia. Remember that Wikipedia is an omline encyclopedia, not a website for presentation of personal oppinions.

Walter, remember that a Wiki-project is based on collaboration - I have read CreationWikis article about Haldane's Dilemma and its Talk Page. Don't you see a pattern here? One thing is to write a book or one's own webpage, another thing is participating in a collaboration project, where one must accept to suppress personal opinions.

--FreezBee 11:37, 19 June 2006 (UTC)

Seeing that ReMine hasn't come up with any answers to the above, I have copied the suggested text to the section "Walter ReMine and Haldane's Dilemma". If ReMine is dissatisfied with the text, I suggest that he uses this talk page to address specific points in the text and, if possible, give specific suggestions to changes.

--FreezBee 13:09, 20 June 2006 (UTC)

Suggested change to intro section

The current intro section ends with

though it is not a problem for evolutionary biology since it consists of a poor model of natural selection that makes invalid assumptions.

This is a much too loose sentence. It is true that it was discovered in the 1960s that far from all genes were fixed. Such concepts as heterosis and linkage disequilibrium invalidate some of Haldane's assumptions; but I do think the formulation should be changed.

--FreezBee 10:09, 12 June 2006 (UTC)

I think, I suggested ...

moved from Re. Intro section

Haldane’s calculated beneficial substitution rate – one per 300 generations – is not “vague” and is not limited to some special type of substitution (such as those that increase juvenile survival). Your difficulties with the term, “selective death,” do not change that fact. While Haldane’s math and arguments may not be entirely clear to you, his conclusion is quite direct, and not “vague”– one beneficial substitution per 300 generations. Wiki readers are not served by attempts to hide, obscure, or make “vague” that number.

You claim Haldane’s number 300 is merely a “suggested number” – which is even more misleading than your previous word, “estimate”. Haldane’s number is not “suggested” and not merely an “estimate” – Haldane’s conclusion was based on lengthy, detailed calculations, using the same evolutionary model still predominant in evolutionary genetics textbooks today.

but I also know that you have an agenda with Haldane's Dilemma and the number 300, and that agenda is not NPOV.

You claim the number 300 is an “agenda”. It isn’t. On the contrary, it comes straight from Haldane, and many people here have the agenda of trying to bury that number, obscure it, and hide what Haldane’s Dilemma is.

Haldane’s Dilemma did not come from me. Wiki readers deserve to know the problem, or “dilemma” as evolutionary geneticists named it. That understanding comes, in a clear, direct, easy to understand manner, by taking the case of human evolution – with a limit of 1,667 beneficial substitutions. None of that number comes from me – it comes from Haldane, plus the standard claims from evolutionary leaders (for example, that the last common ancestor with chimps was somewhere between 3 and 5 million years ago), plus simple arithmetic. That number comes from identifiable facts. Your attempt to hide those facts is not NPOV.

4.158.234.105 04:57, 9 June 2006 (UTC) Walter R.

==

Walter, the number 300 is introduced in the "Discussion" section of The Cost ..."; more specifically at p. 521. Haldanes has discussed selection intensities and writes near the top of the page:

I doubt if such high intensities of selection have been common in the course of evolution. I thinl n = 300, which would give I = 0.1, is a more probable figure. Whereas, for example, n = 7.5 would reduce the fitness to e^-4, or 0.02, which would hardly be compatible with survival.

That is, according to Haldane himself, the number of 300 generations for fixation of one gene, is not a hard fact, but what he thinks is compatible with survival.

At the bottom of p. 521-top of p. 522, Haldane writes:

If two species differ at 1000 loci, and the mean rate of gene substitution, as has been suggested, is one per 300 generations, it will take at least 300,000 generations to generate an interspecific difference.

That is, Haldane explicity refers to the number 300 as being "suggested", not as a result of a calculation.

best regards

--FreezBee 09:43, 9 June 2006 (UTC)

==

FreezBee – You are taking Haldane’s words out-of-context. Haldane 1957 derives and calculates at great length. Indeed, his derivations and calculations are painfully obvious – with emphasis on the word pain! Haldane is not required to use the words “derive and calculate” in every sentence, just to keep you from misconstruing his meaning. The words “think” and “suggest” are commonly used by scientists, as pleasant alternative wordings, especially when advancing provocative new ideas, as Haldane was doing. Taken in context, it is clear that Haldane was not merely giving a “suggested number” or offering what he “thinks.” That would misrepresent Haldane, and do a dis-service to Wiki readers.

Moreover, no one has claimed Haldane made any error within his derivations or calculations. His figure for the total cost of substitution (on average, = 30) is widely accepted as correct – even by such evolutionary geneticists as Crow, Kimura, and Ewens (who all confidently use that figure in their papers). Likewise, Haldane’s figure of 0.1, for the “intensity of selection,” has not been a source of serious objection by evolutionary geneticists.

Rather, attempts to solve Haldane’s Dilemma have focused on various (contradictory) interpretations of the cost of substitution, concerning when it does, or does not, apply – matters that are still unresolved in the literature. Or, the proposed solutions advance revolutionary new types of evolutionary process – which, (plausible or not), are still unresolved in the literature. Your claim that Haldane’s figure is merely a “suggested number” misplaces, (and obscures, and conceals), the source of controversy.

4.159.23.70 04:23, 10 June 2006 (UTC) Walter R.

The numbers 30, 300 and 0.1 ar indeed frequently used; but that doesn't change that it doesn't appear as if Haldane meant that 1 substitution per 300 generations was supposed to be any final truth, though maybe the best estimate available. The number 300 still wasn't a calculated value, and remember that it's tied in with the number 30 as the average selective deaths factor. Haldane mentioned 10-20 as typical. Keeping the selction intensity of 0.1 constant, the 300 generations would be the average number of generations required for fixation; but allowing for much faster typical fixations.

--FreezBee 09:58, 12 June 2006 (UTC)

Scientists never claim to have “final truth,” but that does not make all their work a mere “estimate.” It’s not an either-or proposition, either “final truth” or “estimate.” There are plenty of options in between, and in Haldane’s case, his figures are not a mere estimate, thought, or suggestion. Evolutionary geneticists do not claim Haldane’s cost (=30) is too high. Haldane’s figure is the result of lengthy derivation and calculation.

Also, 300 generations is not the “average number of generations required for fixation,” they are, in fact, exceedingly long and slow in Haldane’s case. Rather 300 generations is the average rate at which substitutions (or fixations) occur. Many substitutions are occurring concurrently, and one per 300 generation is, on average, how often a substitution completes. 4.158.234.180 09:15, 13 June 2006 (UTC)Walter R.

• So, "[s]cientists never claim to have 'final truth'". Excuse me for asking, but have you been reformed, Walter? As for, whether "Haldane's figure is the result of lengthy derivation and calculation", please tell me at which page of The Cost of Natural Selection the derivation and calculation occurs.
• As for your second paragraph, you are right that Haldane writes that 300 generations is the (average) number of generations between fixations; it's a poisson process. Thanks for the correction. Still, is this value observed or estimated? One thing is to determine the half-life of radioactive isotopes; but I would say that determining the average time between gene fixations is a very different matter. Do we follow one population for thousands of generations (remember, we are dealing with slow reproducing mammals here!), or do we say that one fixation in one out of 300 populations in one generation is the same as one fixation in one out of 300 generations in one population?

--FreezBee 10:40, 13 June 2006 (UTC)

Regarding your question, please tell me which pages of Haldane’s paper you are missing. Regarding your other material, you missed the point, which is not about poisson processes. Each individual substitution is extremely slow (much longer than 300 generations), but with multiple concurrent substitutions (used by Haldane) a fixation is occurring no more often than one per 300 generations, on average. Your concern with poisson processes is irrelevant here, because it does not change that substitution rate. It would not matter, for example, whether the substitutions occur at exactly 300 generation intervals – because it’s still the same substitution rate, which is our concern here. Also, you confused cost versus payment. The cost is calculated (based on evolutionary genetics and many assumptions that favor evolution). The payment is actually produced by the species (and is observable). 4.158.231.92 07:01, 14 June 2006 (UTC)Walter R.

I am not missing any pages of The Cost of Natural Selection, so maybe the derivation and calculation is in another paper? If they are in The Cost ..., please do tell me at which page, since I am apparently too ignorant to figure that out myself. According to Haldane's suggestion, a fixation would occur on the average each 300 generations; but as Haldane's analysis shows, it's irrelevant if theses are the peaks of sequential substitutions or parallel substitutions. If several loci are under selection simultaneously, then the fixations will take longer time, because still on average only one fixation can occur per 300 generations. The reference to poisson processes is relevant, because it deals with how exactly is the substitution rate determined.

--FreezBee 11:50, 19 June 2006 (UTC)

WalterR, it appears to me that you are a creation scientist whose method in reading the Haldane paper is a form of data mining to try to undermine the theory of evolotion. Rather than understanding Haldane's paper, it appears to me that you understand what you want to understand.

FreezBee understands what Haldane was trying to do, and his/her interpretation of how we should regard that number 300 is correct. Walter R, you do not understand the concept of a theoretical model. Haldane wrote his paper with limited information. At the time he was writing, the genetic code had just been discovered a few years earlier, and geneticists had only a vague idea of how many genes there were in a genome. They had a rough idea of the size of the human genome in base pairs. At the time, no meaningful sequencing of DNA had been done, and the techniques had not even been discovered. The alternative to DNA sequencing at the time was the sequencing of proteins, which is to say, the study of the products of genes themselves. Haldane was not trying to prove once and for all that it takes 300 generations to fix a gene. Rather, he was demonstrating that if evolution consists of the complete replacement of one version of the gene in the entire population with another, then there would be a stochastic bottleneck in how fast evolution can occur, particularly in complex species with long gestation periods, like homo sapiens. User 4.158.231.92 (Walter R), would you please get yourself a Wikipedia login so that we can verify your contributions here in the future. --Metzenberg 06:28, 29 June 2006 (UTC)

Sorry Freezbee and anonymous but I'm having trouble following your argument. I don't see how it being a poisson process makes a difference (perhaps the word you are looking for is stochastic not poisson?). I'm a cricket fan and you can say that Australia are gonna score about 5.4 runs an over. Sure that doesn't mean that every over there is gonna be 5.4 runs. But the longer the game is played the closer the average score is gonna be to 5.4. Same with this. No one denies that it is stochastic. There is a vanishing chance that 10 000 substitutions could be made (vanishing the operative word) but a similar chance of 0 substitutions. But who cares? I don't see how it makes a difference. Further, perhaps the word "estimate" is technically correct but perhaps WR objects to its emotive connotations? A lay reader may be misled as to how reliable it is. For example you could call a fact sheet on a political party "propaganda" and be technically correct. But in the minds of readers the thought of the nazi propaganda machine will pop up and the readers will be left thinking that the document is unreliable even if its not. Perhaps the same applies here?

References

Added template so reference citations show up. Added ReMines paper as the required citation at the end of the sentence per Wiki formalism.71.115.90.62 02:22, 28 June 2006 (UTC)

This is an unreadable pile of nonsense

I wrote a critique of what had been written back in early May. I see that a lot has been added, and it is really not helpful to a non-mathematician who is trying to understand Haldane's dilemma.

Why the mathematical equations? Wikipedia is not the place to parrot the mathematical exposition that Haldane himself gave. The dilemma can be explained in plain English, without the use of mathematics. If you need mathemetics, go read the original article.

Since I wrote a long critique of the article back in May, maybe I will jump in and start editing. I would start by re-writing the introduction. Haldane's model was not a "bad" model of evolution. It was a model based on information that was available in the 1950s, and it posed an important question. It was in fact an excellent model, and it will still be a part of the theoretical study of genetics in the 22nd and 23rd centuries.

A discussion of how the "creation science" movement has misinterpreted the Haldane article as support for some kind of directed evolution would help explain the dilemma, and why it is no longer a dilemma for most biologists. Several excellent pieces have been written in recent years by evolutionary biologists.

Would anybody have a problem with me eliminating the mathematical equations and starting over, writing an article that a high school or college student can read. --Metzenberg 06:10, 29 June 2006 (UTC)

I agree: The current essay "is an unreadable pile of nonsense." Unfortunately, many people here seem to have that as an intentional goal -- of obscuring Haldane's Dilemma, rather than revealing it. So I cleaned up the section on ReMine, to properly portray his position relative to the remaining essay. 4.158.234.225 10:09, 29 June 2006 (UTC)Walter R.

Is it just me, or is it sort of creepy how Walter ReMine always refers to himslef in the third person? ReMine is obsessed with his conspiracy theories, and his self-aggrandizing tales of having somehow disproved evolution (of humans, at least). But until he can demonstrate that 1667 mutations is actually too few, all of his bloviating is just unmitigated hubris.

It is creepy how you anonymously smear ReMine by misrepresenting him as a conspiracy theorist, and so forth. Until evolutionists can demonstrate that 1667 mutations (or any other figure) is enough, they have not solved Haldane's Dilemma. They are wrong to keep Haldane's Dilemma from the public. 4.158.231.23 18:49, 3 July 2006 (UTC)Walter R.

It is no smear - it is obvious from your claims and implications. Your obsession with anonymity is itself creepy, considering how you have posted anonymously at several newsgroups under aliases such as 'IThinkso' and 'laserthing', each time referring to yourself in the third person, each time lauding "ReMine" for his great work.

You have never actually explained how Haldane's dilemma IS a dilemma, you just assert it is because you are one of those anthropocwentric creationists that wants to think that there is some gigantic chasm between humans and all other life. IN order for 1667 to be too few, as is your position, you would have to tell us all what traits the ape-like ancestor lacked such that 1667 fbe's have to accoutn for them. Which you cannot do. Your claims are baseless bombast.

Further, you have never provided any evidence at all that 1,667 (plus several thousand expressed neutrals) can NOT account for it, which is YOUR claim. Your laughable attempts at claiming things like "appreciation of music" requiring fixed beneficial mutations just shows how desparate and pathetic your position is.

Who invited him? 1667 not a problem? please intelligent comments only. C'mon people, this is an encylopedia not a comic strip. If you are going to argue against haldane's dilemma AT LEAST stick to the arguments evolutionists use (I mean the ones that appear in the technical literature). Ps its a bit hypocritical calling walter R anonymous when he signs his name walter R yet you fail to sign yours. - Graham

Vandalism

Slandering another user in the talk page is frowned upon, slandering another user on the actual edit is wrong and consitutes as wiki_vandalism. I removed his comment "don't you think its creepy that walter refers to himself in third person....." from the main page. This is a warning. Walter R's email is easily obtainable from google if you want to insult him but Wiki is not the place to do it - Graham (Look at the history to see what I'm talking about)

Looks to me like the article has been vandalised. The following does not seem appropriate for an encyclopaedia article:

"It is a shame then that Walter ReMine does not ever provide any evidence that 1,667 mutations actually is too few - no, he merely relies on his clumsy prose, silly rhetorical questions (as if they are evidence), and ignorance of gene action and development. For example, he does not seem to realize that 'hand dexterity, vocal speech, language, and the appreciation of music' can all most likely be accounted for by the tripling of brain size. Of course, why ReMine thinks 'appreciation of music' would require a series of fixed, beneficial mutations is anyone's guess. Mine is that he is just desperate and spewed out something that he figured his ignorant target audience would find impressive."

It should be removed. Some individual's opinion about this issue and an accompanying spew is not relevant here. Since the article is clearly in dispute, I won't make the edit. I'll leave it up to the regulars here. The article should provide information about Haldane's dilemma, not the opinons of individuals about why he was "wrong". People appear to be just trying to defend their pet presuppositions and worldviews, not write an informative article. The dilemma is a piece of mathematics, not a criticism of someone's belief structure. There should be no need for this article to be in dispute. -wbhart

As seen on this thread, ReMine's opponents give vigorous lip-service to "neutral point of view" -- but in the article they allow vandalism and blatant non-neutral attempts to smear him. And they do what they can to AVOID revealing Haldane's Dilemma in any coherent, clear, readable fashion. They seem more than content to leave the article vandalized and non-neutral -- so long as it favors evolution. Such are the scandals of Haldane's Dilemma. 4.158.234.185 03:59, 16 August 2006 (UTC) WalterR

Maybe I'm just not as reticent as wbhart, but I went ahead and removed the "offending" (and offensive) paragraph (although it was formatted as part of the paragraph above it). That's the only edit I made in that section. Mdotley 16:47, 17 August 2006 (UTC)

The article has been vandalized again (this time in the "See Also" section), by the same anonymous evolutionist who vandalized it previously. The URL is from Norwich University, where Scott L. Page teaches, and the text bears his infamous style. 4.158.231.39 06:07, 22 August 2006 (UTC)WalterR

I have reverted it. Mdotley 16:54, 22 August 2006 (UTC)

The Real Issue

And the dispute is not about mathematics, nor is the dilemma. It's about the interpretation of it. Does Haldane's model describe the real world? At what level? This is not the place to be answering those questions -- that's what the journals are for. But it is appropriate to report that there is a dispute over those interpretations, and I think that's what most of this article does. Maybe no one cares about my opinion, but that's OK. Mdotley 16:47, 17 August 2006 (UTC)

This WIKI article is a monument to evolutionist attempts to evade, divert, and obscure Haldane's Dilemma from public view. Haldane's Dilemma is not a "concept" -- it is fascinating unsolved problem, which the article strives to bury as deeply as possible in confusion. Haldane's Dilemma is not about the "efficacy of natural selection" -- it is about the inability of evolutionary genetics to cope with its own central issues. Haldane's Dilemma is not about the "extinction of a species" -- it is about the implausibility of high substitution rates. 4.158.231.177 08:52, 19 August 2006 (UTC)WalterR

Yes, yes. You are probably right. However, it is important for an encyclopedia to remain as objective as possible. If you don't take a chill pill, even those who agree with you will soon find you tiresome. Mdotley 16:52, 22 August 2006 (UTC)

On the merits of what you say, perhaps the technical part of the article needs to be moved to a subpage. Mdotley 16:52, 22 August 2006 (UTC)