Talk:Haldane's dilemma

From Wikipedia, the free encyclopedia

WikiProject Genetics This article is part of WikiProject Genetics, an attempt to build a comprehensive and detailed guide to genetics on Wikipedia. If you would like to participate, you can edit this page, or visit the project page to join the project and/or contribute to the discussion.
??? This article has not yet received a rating on the quality scale.
??? This article has not yet received an importance rating.
Creationism Portal
This article is supported by the Intelligent design WikiProject.

This project provides a central approach to Intelligent design-related subjects on Wikipedia.
Please participate by editing the article, and help us assess and improve articles to good and 1.0 standards, or visit the wikiproject page for more details.
B This article has been rated as B-Class on the Project's quality scale.
Low This article has been rated as Low-importance on the importance scale.
This article has been marked as needing immediate attention.

The article has been rated for quality and/or importance but has no comments yet. If appropriate, please review the article and then leave comments here to identify the strengths and weaknesses of the article and what work it will need.
Archive
Archives
*Talk:Haldane's_dilemma/Archive 1: April 2006 – Agust 2006


Contents

[edit] Entire article

This entire article has massive WP:NPOV (especially undue weight) problems. Remine's arguments should get a paragraph at most with explanations as to why they are considered to be worthless by mainstream scientists. JoshuaZ 16:59, 22 August 2006 (UTC)

Leading evolutionary geneticists, James Crow and Warren Ewens, acknowledge ReMine’s paper is correct – which is a bit inconvenient for you. Also, the errors and confusions identified by ReMine’s paper are still actively advocated in the Wiki essay – thereby validating ReMine’s claim that evolutionary leaders allowed confusion and error to thrive in this field. 4.159.23.36 04:10, 24 August 2006 (UTC) WalterR
Without ReMine's arguments, there wouldn't be any controversy -- they deserve to be fully described. There are scientists who think the dilemma has been solved, but if you want to claim they think ReMine's arguments are worthless, you'll have to come up with a reputable citation, not just evolutionist 'Net-rants. Mdotley 16:16, 25 August 2006 (UTC)
Excuse me? If no scientists have bothered to comment on Remine other than "Net-rants" then it would go a long way to showing that Remine's work is arguably original research and is almost certainly not notable.JoshuaZ 16:18, 25 August 2006 (UTC)


Excuse me? ReMine's paper was peer-reviewed by evolutionary geneticists (including James Crow and Warren Ewens) who acknowledge it is correct. You have no reason to brush that aside. 4.158.231.1 07:00, 26 August 2006 (UTC) WalterR
JoshuaZ, Remine's arguments have been appropriately published. Responses have also been published; some in 'Net-rants, some in peer-reviewed journals. If you want to claim that mainstream scientists think the arguments are worthless, check the journals, (not the 'Net rants) and get appropriate citations. Mdotley 21:41, 8 September 2006 (UTC)
Mdotley wrote:
"Remine's arguments have been appropriately published."
Not on my reading of the guideline for evaluating the reliability of sources, they haven't. None of Mr Remine's arguments have been published in peer-reviewed scientific literature or any other sources whose credibility on the matter is verifiable. The only sources for most of his arguments are a self-published book and on-line sources (or "Net-rants", as you have so aptly described them). The one known exception, published in TJ-in-depth Journal of Creation, doesn't contain any of the more controversial of Mr Remine's arguments at all. All it contains are:
* An exposition of Mr Remine's revised definition of the cost of selection (which he calls the "cost of substitution");
* Claims that the existing literature is full of confusion and contradictions which have been cleared up by his new definitions;
* Demonstrations that his definitions give the same value as the traditional ones in certain special cases;
* Derivation of conditions under which his cost of substitution will be at a minimum and expressions for the value of the minimum.
I entirely agree with JoshuaZ that the current article gives too much prominence to Mr Remine's views. There are also several places in the current exposition of his views where the wording implies that disputed POVs of his are in fact correct, and no independently verifiable source is cited for the assertion that Warren Ewens and James Crow (among others) have judged his TJ paper to be correct.
Nevertheless, it seems to me that Mr Remine has some justifiable grounds for complaint about the way this article has developed. In the early sixties genuine scientific controversies most certainly did break out over what some people perceived to be implications of Haldane's paper. Some of the participants (E.O.Dodson in particular) made arguments quite similar to some of Mr Remine's. Others, such as Feller and Moran, argued that Haldane's arguments were fallacious; but then their own arguments were in turn rebutted by Kimura and Crow.
A proper encyclopedia article on Haldane's dilemma should give an account of all the significant points of view put forward in this scientific literature, including rebuttals (if any) of the views favoured by the article's editors. It would appear from the history of this article, however, that too many of its editors have merely set out with the principal aim of showing why Haldane---or more particularly Remine---was wrong. In my opinion, the result has been a completely unbalanced article. It seems to me to give far too much prominence to the technical minutiae of Haldane's paper, and not nearly enough (essentially none, in fact) to describing the various points of view of the participants in the above-mentioned scientific controversies. David Wilson 20:45, 20 September 2006 (UTC)

David Wilson is correct that the current article is "completely unbalanced." The article wantonly obscures the problem from view, lulls readers into oblivion with needlessly tedious derivations, and gives a false sense of agreement and resolution among evolutionary geneticists. By comparison, the section on ReMine offers a modest remedy toward sobriety. If you think the current article gives "too much prominence" to ReMine's views, then the solution is to sober-up the rest of the article. That is:

  1. Give a clear, direct statement of the problem (such as the limit on the number of substitutions to explain human evolution).
  2. Give a clear explanation of the cause of the problem (because Haldane's description is not the clearest available).
  3. Identify the most prominent resolutions claimed for the problem.
  4. Acknowledge the lingering confusions and disagreements surrounding Haldane's Dilemma (such as George C. Williams's assertion that Haldane's Dilemma "was never solved").

In other words, give a clear, revealing, up-to-date account. The idea of giving "an account of all the significant points of view put forward in this scientific literature" is not realistic, since that would essentially require a book, and moreover, many of those views are contradictory and "were fallacious". You could forego those views that were explicitly refuted in the literature, unfortunately, rather few of them were. Many fallacious arguments (including Feller, Moran, Hoyle, Felsenstein, and many others) thrive today because they were not explicitly refuted in the literature. Confusion and error were allowed to thrive (so long as it favored a solution to Haldane's Dilemma). The literature remains in a state of unresolved confusion and contradiction, and coping with that appropriately will be a key difficulty for this article. Also, James Crow publicly acknowledged (on the ARN.org website) that he and Warren Ewens judged ReMine's paper to be correct. It is independently verifiable. 4.158.231.65 08:50, 22 September 2006 (UTC) WalterR

WalterR wrote:

'The idea of giving "an account of all the significant points of view put forward in this scientific literature" is not realistic, since that would essentially require a book, ...'

Not at all. There's no need to give a detailed account of every single thing ever written on the subject. The various significant opinions could be quite easily grouped into a small number of about 4 or 5 classes, whose adherents' differences in each case were essentially trivial. All that's necessary is a succinct, accurate summary of each position, with citations from one or two of its most prominent adherents. A much more difficult problem, in my opinion, will be achieving a consensus on which particular positions are sufficiently "significant" to include, how to group them, and what constitutes an unbiased account of each of them.

WalterR wrote:

  1. Give a clear, direct statement of the problem (such as the limit on the number of substitutions to explain human evolution).
  2. Give a clear explanation of the cause of the problem (because Haldane's description is not the clearest available).

Like the current introdution to the article, that procedure would not be NPOV, because it implicitly assumes that one of the more significant POVs, which has never been refuted by anyone, including Walter Remine (in my opinion), is incorrect. The POV in question is that no-one has ever shown that the theory Haldane developed in The Cost of Natural Selection poses any sort of dilemma for evolutionary theory.

It is obvious from the discussion section of Haldane's paper that he himself did not think that it posed any such dilemma at the time when he wrote it. While we know from A Defence of Beanbag Genetics that he was aware of the controversies over the cost of selection which arose in the early sixties, he also gives no hint there, or anywhere else as far as I know, that he ever regarded it as posing a dilemma for evolutionary theory. We also know from the writings of Ewens and Crow that neither of them do either (though for different reasons).

A Wikipedia encyclopedia article is not the place to argue the merits or defects of this (or any other) POV. It is however also not appropriate to suppress it by using a description which implies it is incorrect.

WalterR wrote:

Also, James Crow publicly acknowledged (on the ARN.org website) that he and Warren Ewens judged ReMine's paper to be correct. ...

I presume you are referring to this post by Nikolas Voss, in which he posted a response of Crow's to some discussions that had taken place on the bulletin board where the response was posted. In that response Crow did not say that "he and Warren Ewens judged ReMine's paper to be correct". Here is what he actually wrote:

"I did review ReMine's paper for Theoretical Population Biology. I have forgotten most of the paper, but I recommended rejection on the grounds that, although the paper was essentially correct and made some interesting points, there was not enough new theoretical content to merit publication in this highly technical journal. Another reviewer, Warren Ewens, reached essentially the same conclusion. I also said the paper might be published in a less technical journal."

Omitting the word "essentially" from before "correct" in your indirect quotation gives the possibly misleading impression that Crow and Ewens raised no objection whatever to anything written in the paper. I'm afraid I find that difficult to believe. I have read your paper very carefully, and checked all the calculations and proofs. While I think Crow's assessment of "essentially correct" is quite reasonable, I also think that it would be false to say simply that the paper is "correct" without qualification. For what it's worth, I also agree with Crow's assessment that it contained "not enough new theoretical content to merit publication" in a journal devoted to publishing original research.

In my opinion the paper contains several technical errors. While they can all be corrected without doing much damage to what little of substance it does actually contain, they're also sufficiently serious that any conscientious reviewer would have to demand that this be done before it could be considered for publication.

Apart from the misquotation just indicated, the current wikipedia article is also selective in its choice of exactly which parts of Crow's comments it has chosen to cite. As noted above, Crow also said in his comments on your paper that "there was not enough new theoretical content to merit publication in this highly technical journal" (i.e. the journal of Theoretical Population Biology). He also wrote:

"Let me emphasize that the change of interpretation from Haldane's load to reproductive excess is a small improvement and should not be regarded as comparable in importance to Haldane's original very clever idea. Compared to this, the change in emphasis from genetic deaths to reproductive excess isn't such a big deal."

"There is the additional issue of just how important the Haldane cost is as a limitation to evolutionary change by natural selection, but that is another question. I don't think the Haldane principle is useful as a way of putting a brake on evolution rates; it is too simple and there are too many ways around it."

"I don't want to say that what I wrote 35 years ago is a model of clarity and consistency and couldn't be improved on. But I still think it is reasonably clear and essentially correct, and includes the essence of ReMine's required reproduction rate."

It's simply not appropriate to offer support for a position by selectively misquoting just those parts of someone's comments which agree with the position, while at the same time ignoring other parts which disagree with it. An appropriate use of Crow's comments would be something like the following:

Leading evolutionary geneticists, including James Crow and Warren Ewens, reviewed the paper for possible publication in the journal Theoretical Population Biology. Crow and Ewens acknowledged it is essentially correct but recommended against publication because they considered its technical content was not sufficiently novel to warrant publication in that journal[1]. Remine disagrees with this assessment. His responses to the rejection of his paper can be found here and here

David Wilson 03:52, 24 September 2006 (UTC)

David Wilson (reasonably) proposes that the numerous published views be classified into a small number of groups, and the article then give a summarization of each group -- though many relevant details would be lost. But he also suggests the article should include irrelevant details, such as:

Wilson's text:
"... for possible publication in the journal Theoretical Population Biology. Crow and Ewens ... recommended against publication because they considered its technical content was not sufficiently novel to warrant publication in that journal[1]. ReMine disagrees with this assessment. His responses to the rejection of his paper can be found here and here."

Wilson's text may be relevant for other articles, (say, concerning the peer-review process), but it has no serious relevance for this Wiki article. This Wiki article is not about why ReMine's paper was published in one journal rather than another. This Wiki article is not about the journal Theoretical Population Biology and why it should, or shouldn't, have published ReMine's paper. Wilson's text serves no purpose here (except as a misplaced attempt to besmirch ReMine's paper). This is another example of (to use Wilson's phrase) a "completely unbalanced article." Evolutionists here focus at length on misrepresenting, minimizing, or smearing ReMine, while persistently avoiding any decent coverage of Haldane's Dilemma.

The relevant issue was charged previously by Wilson, and is now answered and independently verified by Wilson: ReMine's paper was peer-reviewed by evolutionary geneticists, Warren Ewens and James Crow, who publicly acknowledged it is essentially correct and made some interesting points.

Though it was correctly peer-reviewed, ReMine's paper contradicts the bulk of the Wiki article. ReMine's paper specifically identifies unnecessary confusions and errors that are widely promoted today -- promoted even within the Wiki article. The Wiki article needs to cope with those in some meaningful way: such as by eliminating them from the article, or alternatively, by acknowledging their existence within the article.

David Wilson writes that Haldane, Crow, and Ewens did not regard Haldane's Dilemma as a dilemma for evolutionary theory. That is highly misleading here, because the evolutionary genetics literature (including those three authors), almost totally avoided discussion of the serious issue -- a limited number (~1,667) of beneficial mutations to explain human evolution. You cannot credit Haldane, Crow, or Ewens for something they did not publish. They and many others were free to publish support that such a limit is not a problem, but they did not. Instead the literature almost always spoke of the matter obliquely, in terms of substitution rate (such as "one substitution per 300 generations"), which obscures the problem from view.

David Wilson wrote:
WalterR wrote:
  1. Give a clear, direct statement of the problem (such as the limit on the number of substitutions to explain human evolution).
  2. Give a clear explanation of the cause of the problem (because Haldane's description is not the clearest available).
... that procedure would not be NPOV, because it implicitly assumes that one of the more significant POVs ... is incorrect. The POV in question is that no-one has ever shown that the theory Haldane developed in The Cost of Natural Selection poses any sort of dilemma for evolutionary theory. [bolding added]

David Wilson argues it would be "not neutral" -- and therefore disallowed by Wiki -- to give a clear, direct statement of the problem and its causes. I regard his argument as scientifically perverse on its face. He attempts to justify it based on the existence of what he calls "one of the more significant points-of-view" -- which happens to be effectively non-existent in the literature. Evolutionary genetics literature does not seriously support (and does not remotely have a consensus) that a limit of something-less-than 1,667 beneficial substitutions is not a problem. On the contrary, (and contrary to Wilson's claim), the literature displays numerous evolutionary geneticists who took Haldane's speed limit as a problem in need of a serious solution. It would be perverse to withhold a clear, direct statement of the problem and its causes.

4.158.231.20 09:09, 28 September 2006 (UTC) WalterR


WalterR writes: "...is highly misleading here, because the evolutionary genetics literature (including those three authors), almost totally avoided discussion of the serious issue -- a limited number (~1,667) of beneficial mutations to explain human evolution. You cannot credit Haldane, Crow, or Ewens for something they did not publish. "

This is true, but trivially so - none of them explicitly discussed human evolution at all, so why make seem like an issue that they did not point out that number and how it is supposedly some big problem? Furthermore, there is nothing anywhere in any literature - ReMine's book or paper, many internet posts, etc. - that actually provides EVIDENCE that, even if correct and relevant, ReMine's '1,667 mutations' even is a 'problem' for evolution. The repeated mantra that it is is itself an unsupported claim. --slpage 21:02, 25 January 2007 (UTC)


[edit] Section on ReMine's opinion

I've removed the section on ReMine for the following reasons: 1) ReMine's views are shared by only a very small minority, namely, like-minded ID advocates. 2) It was way too long, comprised nearly half the article's length. 3) Throughout its length it used non-neutral terms such as "evolutionists" etc. If the opinion is going to be noted in the article then it will need to summarized in a paragraph or two and written using neutral terms. FeloniousMonk 03:20, 20 January 2007 (UTC)

I've been watching this page for some time although I haven't actively edited it. I agree that there were massive factual and POV problems with it but disagree that all reference to Walter Remine and his claims about the supposed "dilemma" should be removed. This is because the idea of "Haldane's Dilemma" has been publicized and perpetuated by Remine and frequently comes up in the creation/evolution dispute (although almost entirely from the creationism side). I think it's fair to say that if it were not for Remine, "Haldane's Dilemma" would be no more than a one-sentence footnote in biology, and the vast majority of people who come to Wikipedia looking for information about it will have heard about it through the creationism connection. MrDarwin 16:27, 20 January 2007 (UTC)
Yeah, I largely agree; so let's write something more appropriate up. I've asked an expert, Arthur Rubin, to assist on another part of the article, perhaps he can give us a hand with this, too. FeloniousMonk 17:10, 20 January 2007 (UTC)
I'm a mathematician, not an evolutionary biologist. I can certainly comment on the "1,667" quoted in an earlier version of this article as the maximum number of "significant" mutations that could have been selected for, but I'm not sure about the consensus in the biology field. — Arthur Rubin | (talk) 17:00, 21 January 2007 (UTC)
Your comments on the "1,667" figure would be most welcome. As you may have seen, ReMine considers it a key figure that should be in Wikipedia. Thanks, AvB ÷ talk 09:55, 24 January 2007 (UTC)
ReMine's opinion strikes me as the very definition of a tiny minority view, and WP:NPOV says tiny minority views need not be mentioned at all. What evidence do we have that is something more than a tiny minority view? FeloniousMonk 03:34, 25 January 2007 (UTC)
At this point I'm viewing his remarks as if they were given on this talk page, not as a source for WP. But we can't do much without verifiable, reliable secondary sources. I've read through a number of his web pages and found them interesting. I haven't seen any acceptable sources for his work to be included in the article but he does come close in the case of the rejected paper in terms of reviewer comments and withdrawn criticisms. They didn't see his credentials as an impediment which is remarkable. Perhaps ReMine is notable in some other way so that his supporters can start an article. AvB ÷ talk 18:34, 25 January 2007 (UTC)

[edit] Heads-up.

This page is linked to from Uncommon Descent via a post today titled "Wikipedia Suppresses Info On Haldane’s Dilemma". grendel|khan 19:37, 23 January 2007 (UTC)

Thanks. Both the relevant material and personal attacks against Wikipedia editors in good standing at uncommondescent.com don't spell any good for "DaveScot" or anyone he may have influenced if they decide to have a go at this WP article.
Nevertheless, some of the criticisms expressed here by Walter ReMine seem valid to me when weighed against Wikipedia's rules. I suggest we disregard its personal and anti-Wikipedia slant and discuss the relevant content here. AvB ÷ talk 09:51, 24 January 2007 (UTC)
Well, if we've got the pseudoscience pov pushers frothing, we must be doing something right. DS has been banned from Wikipedia, due to a particularly shabby record of trolling, disruption and legal threats. FeloniousMonk 03:25, 25 January 2007 (UTC)

It is an anti-wikipedia rant but something that expresses truth. Sorry if this is off-topic but it's quite noticeable to any christian that wikipedia is biased even though they claim to be neutral. --Macguysoft 03:17, 25 January 2007 (UTC)

Really, how so? FeloniousMonk 03:25, 25 January 2007 (UTC)
I think the explanation for his comment can be found in his user profile: "My interests, however, are more into the creation vs evolution controversy. I'm a rare mac user who is a creationist but this alone gets me in serious trouble with mods and grabs attention instantly." In other words, Wikipedia is "biased" against his creationist viewpoint. (B) 27 January 2007 —The preceding unsigned comment was added by 71.218.89.128 (talk) 09:19, 27 January 2007 (UTC).
I've noticed a tendency in some Christians to regard anything devoid of Christian content as biased against Christianity. Unless Macguysoft provide a specific example of bias without pushing a particular POV, I'd say the article is reasonably neutral but not perfect. That said, I think it could be improved: Remine does deserve more than passing mention, even prominent mention, because without him this topic would be a mere footnote in an obscure biology text, instead of a rallying point for the faithful. =Amatulic 03:33, 25 January 2007 (UTC)
Therein lies the rub: This is a marginally notable criticism of evolution that the scientific community considers a complete non-starter. Making ReMine's opinion a tiny minority view, and WP:NPOV says tiny minority views need not be mentioned at all. What evidence do we have that this topic is relevant to anything other than and arguments of creationists and that ReMine's notion is something more than a tiny minority view? FeloniousMonk 03:37, 25 January 2007 (UTC)
Well, that's kind of my point: The only thing that makes this topic notable is Remine. The topic is notable in creationist circles, which I'd argue is sufficient to warrant an article about it (maybe under the Creationism category or perhaps merged into an article about creationis arguments). And Remine is what made this topic notable, so the article should at least give appropriate weight to that fact. -Amatulic 18:57, 25 January 2007 (UTC)
That might be an argument to delete the article more than anything else. There simply aren't any reliable sources which talk about the creationist version of the dilemma. JoshuaZ 19:06, 25 January 2007 (UTC)
Delete the article? if there is an article on the Coulomb phase then I think there is equal grounds for an article like this. Besides, the gallup poll would suggest this is hardly a minority view http://www.galluppoll.com/content/?ci=21814&pg=1 There have been some suggestions for a shorter section on reMines work. I wrote one which should be in the discussion archives somewhere, it was inherently flawed as I believe I ended up sacrificing some factual accuracy for the sake of getting something NPOV (which was my primary goal at the time). Perhaps it should be dug up and discussed
Hmmm, a gallup poll of the general populace has nothing to do with whether we have enough sources to write a Wikipedia article on Haldane's Dillema. I don't know why you would think otherwise. JoshuaZ 08:22, 17 February 2007 (UTC)
FWIW, I've reread ReMine and reviewed the lit and no longer think his views are relevant. Basic high school maths. Walter, I'm prepared to discuss this in depth back-channel. AvB ÷ talk 02:11, 18 February 2007 (UTC)
As you were saying: [2]... FeloniousMonk 03:40, 25 January 2007 (UTC)
Wikipedia is bound to seem biased to anyone who firmly believes things that aren't true. -- 71.102.194.130 15:05, 2 July 2007 (UTC)

I'm amused. So apparently ID isn't religious but somehow Dave thinks he has evidence that we are "religion hating." Slight contradiction there. JoshuaZ 06:36, 25 January 2007 (UTC)

??? perhaps I shouldn't waste my time on comments which are no less than sledging since this discussion section is meant to be about haldane's dilemma and not a soapbox for anti creationists but perhaps there is a need to educate here. There is a venn diagram between ID, religion and creationism. There are certainly many religious people in ID circles but one does not necessarily have to be religious to be an IDer (one prominent member is an agnostic) and one does not have to be an IDer to be religious. Also it is true that some IDers are creationists this is a rare occurance. What I believe though was alluded to before is that generally ID bashing is not done for scientific reasons rather it appears to be for anti-religious reasons. There is no contradiction here
OK, first please sign your comments with four tildes- ~~~~ which the software will replace with your time of edit and your IP (if you get a username which I recommend you do since it only takes a few seconds, that will appear instead of your IP address). Now, you are correct that this page is for discussing the Haldane article and I went slightly offtopic with a throwaway comment. I'm therefore not going to let this degenerate further by responding to this comment here, but if you want I can respond on your talk page in detail explaining why you above comment about the nature of ID , its opponents and its proponents is inaccurate. JoshuaZ 08:22, 17 February 2007 (UTC)
As with most zealots, contradictions are not truly contradiction as the knowledge the zealots displace is revealed and hence inerrant. ROFL. •Jim62sch• 13:58, 26 January 2007 (UTC)

[edit] Haldane's assumptions

What's long overdue, I think, is a close examination of Haldane's most basic assumptions now that we know a whole lot more about genetics in general, about the numbers of genes that make up an individual, how many genes and more precisely how those genes differ between related species. As far as I can tell, Haldane pulled his numbers out of thin air, or at best based his calculations on some very shaky assumptions and I suspect that at least some of these assumptions will turn out to be wildly wrong. I also have to wonder if leaving the haploid (gametic) generation out of the equation is a major source of error, when selection can and does act, and very strongly, on gametes at the molecular and cellular level (e.g., out of millions of sperm cells only one will actually fertilize an egg). Unfortunately I haven't been able to track down any references that address these issues. MrDarwin 19:45, 25 January 2007 (UTC)

The factors that determine which sperm fertilizes the egg have virtually nothing to do with the traits of the offspring, so no, selection does not act "very strongly" on gametes. -- 71.102.194.130 15:12, 2 July 2007 (UTC)

In addition to Haldane's assumptions, a very important aspect of the characterization of Haldane's Dilemma by Walter ReMine and sundry ID/creationsim advocates is a discussion on its relevance to evolution at all. As Haldane had the foresight to concede, his numbers would probably need "drastic revision." This little fact seems to escape the discussion altogether. Further, some important context is overlooked, that is, the time in which Haldane wrote his paper (and the follow-up paper in 1961). At the time, it was assumed that the human genome contained at least 125,000 genes, and that most traits were governed by a discreet gene or set of genes. Haldane simply did not have the information available to him that we do now - basic genetic information like the size of the human (mammalian) genome and the actions and activities of the genes in the genome. Common sense should dictate that with the explosion of new data and technology regarding the field of genetics that one should be cautious about employing standards and mathematical models premised on 30-40+ year-old observations and understandings. In terms of applying Haldane's model to the human evolution question, some important questions that ReMine and his supporters did not, have not, and will not address (I know, I have asked them) must be answered before any relevant application of the model, even if applicable, can be accomplished. Among them:

1. What was the ancestral population from which the lineage leading to humanity arose?

2. What traits did its members possess that cannot be accounted for by the 1,667 fixed beneficial mutations (and some 10s of thousands of expressed neutral mutations), if that number has relevance?

3. How many fixed beneficial mutations ARE required to produce the traits possessed by modern humans that cannot be accounted for (question 1 must be answered first or at least at the same time as this one) by the 'ReMine number'?

These important questions are either ignored, dismissed, or labeled as 'posturing', 'misrepresentation', or 'irrelevant' by ReMine and those that accept his claims regarding Haldane's dilemma. Nevertheless, these questions, at the very least, MUST be addressed by advocates of ReMine's claims before any legitimate scientist takes his applications and claims seriously. As it stands, ReMine's claim is analogous to claiming that you cannot get to New York by driving 55 miles per hour in 10 hours despite not establishing the starting point. --slpage 20:50, 25 January 2007 (UTC)

... or claiming that you cannot get from Los Angeles to New York City in less than ten hours because a horse can only run so fast... MrDarwin 21:02, 25 January 2007 (UTC)

[edit] Is this "dilemma" about Haldane, about ReMine, or about Kimura ?

I recommend the following "tutorial" by Mark Ridley:

http://www.blackwellpublishing.com/ridley/tutorials/Molecular_evolution_and_neutral_theory1.asp

I think it illustrates quite well what the real "Haldane's dilemma" or "Kimura's dilemma" is, as perceived by population geneticists such as Motoo Kimura, James Crow, Warren Ewens and Sewall Wright. It is only indirectly about Haldane(1957), although Haldane's cost model certainly is the primary catalyst. It is not at all about Walter ReMine and his arguments on hominid evolution. First and foremost, it is about Kimura's criticism on the explanatory power of natural selection and about Kimura's promotion of random and neutral change as a better explanation for the vast majority of molecular evolution. The "dilemma" challenging selectionism is related to many observations in molecular evolution, including the claim that in some lineages there have been as many as 6 amino acid changes per generation, which is by any interpretation a more intense rate of change than is allowed by the "cost of natural selection". This would suggest that there are many amino acid sequences, i.e. proteins, which are functionally more of less identical.

Ridley brings forth several key concepts which do relax the genetic load caused by e.g. substitutional load and segregational load. While Ridley's explanations (density-dependent selection, frequency-dependent selection and epistatic fitness interaction) do lower the cost of evolution, it is questionable whether they lower it by many orders of magnitude, as required by Kimura's challenge. Thus, Kimura's argument may still stand. But in any case, arguments mentioned by Ridley remarkably weaken ReMine's argument that at most one adaptation per 300 generations may occur, unavoidably, and thus human evolution somehow becomes impossible. Of course, ReMine's argument never was important to population genetics. ReMine's argument simply isn't the same issue as the real controversy, in contrast to what ReMine would like his readers to believe, partly by resorting to quote mining authorities. ReMine's layman argument may be relevant if changes in gene regulation are included in the argument (as I think they should). That's because apparently we don't yet know how many adaptive changes there have been in gene regulation. But if we only considered changes in coded amino acids, as Kimura apparently did, ReMine's argument would currently fall apart. It seems that mere few hundred amino acids have changed in the human lineage since its divergence from the chimp lineage. If ReMine has a calculation, based on solid data, clearly demonstrating the impossibility of a few hundred adapations during human evolution, then he has demonstrated the existence of a dilemma. It would be novel enough so that he could justifiably name it "ReMine's dilemma". ReMine is invited to present this calculation for criticism, and to use what ever "cost types" he sees fit, and whatever unit of cost he sees fit. But he should publish his calculation in a peer-reviewed journal (TJ or the like hardly count here). So far ReMine has published no rigorously quantified argument anywhere, however -- not even in creationist magazines. Thus, he is requiring others to make public a speculative "dilemma", and to solve it even before it has been demonstrated.

As for "Kimura's dilemma", that selection alone can't explain the fastest observed instances of amino acid change nor the amount of polymorphisms in most genomes, some attempts have been made on it by major authorities. The most radical is by Warren Ewens, apparently to some extent backed by Sewall Wright. This suggested solution was published in "Mathematical Population Genetics, Vol I". It is quoted by Robert Williams on the web site that has been linked from this Wikipedia article. I'm quite puzzled by Warren Ewens' idea. He seems to be claiming that substitutional load is not caused by the difference between the fitness of the theoretic optimal genotype and the average fitness of the population. Instead, he claims that substitutional load is caused by the difference between the fitness of the best actually existing genotype and the average fitness of the population. Ewens is probably right in estimating that beneficial alleles spread in the population in an even fashion, so that the fitness difference between any two living individuals tends to be relatively small. But I find Ewens' core argument questionable. Haldane's idea clearly is that we should compare the fitness of each individual to the optimal genotype, regardless of how likely such a genotype is to manifest itself in a living individual. Even if no individual ever is anywhere near the (current) optimal genotype, selective pressures apply just as mercilessly as they would if some lucky individual happened to hit the optimal genotype. Haldane's key idea seems to be that whenever we postulate the existence of a new beneficial allele, we simultaneously postulate a selection pressure which hurts the fitness of anyone lacking this allele. Ewens seems to ignore the concept of absolute fitness altogether, and only concentrate on relative fitness differences between living individuals. This seems to alter the core idea of Haldane(1957), that there is no beneficial allele without a corresponding selective pressure applied universally to the population. So far as I can tell, Ewens changes Haldane's (apparently quite reasonable) core paradigm without providing any data, or even rational persuasion, to support this assumption. While many of Haldane's assumptions, such as multiplicative fitness interaction, hard selection and omitting intraspecific competition, can reasonably be challenged, I don't think Ewens' change in core assumptions is very persuasive. Obviously, this comes from a layman to challenge an established authority. --Esko Heimonen 11:51, 11 September 2007 (UTC)

Far from being swept under the rug, "Haldane's Dilemma" stimulated quite a bit of discussion in the 1960's and 1970's (and to a lesser extent more recently) and a Google Scholar search will turn up a great number of papers directly addressing it, including examinations of Haldane's assumptions and offering numerous possible solutions to the "dilemma". This is too far outside my area of expertise so I can't contribute much, but some of these papers would certainly be relevant to this article if anybody cares to look through them. MrDarwin 13:05, 11 September 2007 (UTC)
I would like to point out something about "segregational load", which is described in the tutorial linked above. This argument of Kimura against selectionism doesn't really concern the question how polymorphisms evolved at so many loci. It is about how natural selection could maintain these polymorphims. And this does not only concern geologic timescale, it concerns here and now. If heterozygotes had a notable selective edge over homozygotes at hundreds or thousands of loci in many animal populations, these populations should collapse in the very next generation, even if they had been designed last Thursday. Now, Walter ReMine seems to describe Kimura's idea of neutral polymorphisms as an "avenue of escape". I'm interested to learn how the intelligent design theory explains this phenomenon, then. I.e. how does the Designer maintain hundreds or thousands of polymorphisms in many animal populations without resorting to neutralism? If it so happens that the Designer, too, uses the same emergency exit, could Walter ReMine reconsider his choice of words? It sounds implausible that amino acid polymorphisms favoring heterozygotes would be the only neutral mutations. It sounds a lot more plausible that amino acid changes in general are often neutral.

--Esko Heimonen 22:37, 11 September 2007 (UTC)

[edit] More UD discussion

This page is mentioned again today at Uncommon Descent, which has a letter from ReMine up complaining that Nunney (author of a paper mentioned in the references section of the page) has refused to give him access to the simulation software used in the published paper. Someone added a link to the UD article to the external references section, and modified the reference to include the claim that Nunney was refusing access. In the hope of avoiding a lengthy edit battle and much useless screaming, I did not delete either of those edits. I did, however, in the hope of preserving something along the lines of NPOV, edit the claims to make it clear that they are claims, and not statements of independently confirmed fact. Mdunford 19:32, 19 April 2007 (UTC)

[edit] Intraspecific competition

I wonder why the concept of intraspecific competition is not mentioned more often in this whole "Haldane's dilemma" affair. (Nunney *does* mention it, however.) Haldane(1957) is exclusively focused on a population adapting to environmental selection pressures. But individuals in a population do not only strive to overcome external challenges imposed by the environment. They also compete with each other in hogging the vital resources of their ecological niche: food, shelter, good territory, good mate (not to mention harem species!), social status, etc. Even advertising one's fitness to predators is in a sense intraspecific competition: "do yourself a favor and go bother the guy next to me instead". There is no obvious reason why such competition should affect the average fitness of the population and, thus, no reason to expect that allele substitutions always have to be "paid" by increased mortality which then has to be countered by ReMine's cost unit, reproductive excess. In intraspecific competition, it is not about the increased number of deaths, it is simply about the *distribution* of deaths, or, inversely, about the distribution of reproductive success. In summary: intraspecific competition does not require genetic deaths/reproductive excess to occur, and it is capable of driving substitutions just as efficiently as environmental selection pressures. As a layman, I'd suggest that intraspecific competition should be considered as an important factor especially among primates, because they are generally known for their complex social hierarchies and the amount of energy spent on intraspecific interaction.

--Esko Heimonen 12:03, 23 May 2007 (UTC)

Perhaps that hasn't come up in the experts' discussions b/c it is a distinction without a difference. Differential reproduction (or if you prefer, the distribution of deaths) in response to outside pressure is the result of intraspecific competition. An external predator is an environmental factor. I don't understand what you're trying to get at, here. Mdotley 02:12, 25 May 2007 (UTC)
Fair enough. I will try to illustrate the difference more clearly. Consider a population with territorial behavior. In such a population, not being able to conquer a territory at all can be considered a "juvenile death" sentence, and not being able to conquer a *good* territory probably bears fitness penalties. Are fights over territory within the population a matter of pure luck? Of course not. They are genetic viability tests just as overcoming any environmental selection pressure. Only this time we are considering internal competition, and not selection pressures external to the population. When a subpopulation becomes better at winning territorial battles, this does nothing obvious to the average fitness of the population. There are still just as many territories available in the ecological niche, and just as many individuals competing over them. The population as a whole doesn't lose anything, or win anything, in terms of how fit, on average, they are to survive in their environment. Only the *distribution* of the territories between individuals in the population will change, to favor this new "conqueror" genotype. Such a favorable distribution will, then, bring fitness benefits to the optimal genotype and deprive fitness benefits from other genotypes. But that condition is quite sufficient to drive an allele substitution. No need to postulate excess mortality. This was just one example -- I suggest that there are many material and social resources that individuals in a population are likely to compete over.
The difference to Haldane(1957) is that, in intraspecific competition, we do not have to assume any temporary reduction in the average fitness of the population and, thus, no excess mortality and, thus, no reproductive excess. In Haldane's model, which excludes intraspecific competition, changes in environmental pressures would make each territory slightly less hospitable to its host, thus lowering the average fitness of the population and increasing mortality. Haldane assumes that a tiny subpopulation possesses formerly insignificant alleles which now happen to counter the environmental changes, such as the appearance of a new predator. These alleles will obviously form a new optimal genotype, which then becomes fixed. During this substitution process, a large amount of individuals most distant from the optimal genotype will suffer a juvenile genetic death, due to the new environmental challenges, and these deaths are *additional* to whatever the average mortality was before the environmental change. This excess mortality needs to be countered by reproductive excess, or the population faces extinction. These condititions suggest a limit to intensity of selection, which Haldane(1957) estimates to be I=0.1, on average, which in turns allows substitutions to occur no faster than one per 300 generations. It is from this number that ReMine directly draws his "magic number" 1,667.
I maintain that the concept "intraspecific competition" makes a genuine, qualitative difference compared to competition of genotypes exclusively based on differential survival in a changing environment. The difference is that in intraspecific competition, individuals fare better by "oppressing" others in the population, not by directly adapting to the environment surrounding the population. I also maintain that this concept is overlooked by Haldane(1957) while apparently being taken into account by e.g. Nunney(2003). (Apologies for any bad wikiquette in these writings, I am a first timer.) --Esko Heimonen 07:10, 25 May 2007 (UTC)
Firstly, please edit before saving -- stepping through your edits one at a time can be trying.  :-)
Secondly, keep in mind that the "I=0.1" factor is FAR more intense than has ever been observed in nature. That would make the replacement time much longer, and more problematic for molecules-to-man evolution.
Thirdly, I haven't read Nunney, nor am I an expert, but I fail to see how your distinction makes a difference. The "story" one tells to justify why some reproduce more effectively into the next generation is completely irrelevant. It doesn't matter whether it's a change in the environment that makes the majority less fit or a change in the genome making a minority more fit. The effect is that a minority now has an advantage over the majority, and the biomathematical problem is the same: given a degree of "more fit-ness", how long does it take for the minority allele to be fixed in the population? Mdotley 01:11, 31 May 2007 (UTC)
Those repeated saves was one of the things I apologized for. :) In the future, I will try and preview my posts more carefully before saving.
According to Haldane(1957), the I=0.1 factor is certainly not "FAR more intense than has ever been observed in nature". Just look at Haldane's very examples in that paper. In fact, Haldane's estimate of the mean value for intensity of selection (I=0.1) is far less intense than his own examples. But since you apparently claim to have data demonstrating a much smaller mean value, could you present it? What kind of intensities have been measured and reported in primary literature, and what is the mean value for this intensity according to these measurements?
I will try to illustrate my point of intraspecific competition yet once more, although I believe you should re-read my previous explanation. (I think the difference between intraspecic competition and adaptation to environmental changes, in terms of "cost" and, thus, also in terms of various "magic numbers" - both Haldane's and ReMine's - is both explained and emphasized in my previous contribution.)
Consider a population with initial *absolute fitness* 0.75. That is, assume a population in which each generation suffers 25% mortality in terms of "juvenile deaths". This in turn means that this population needs a reproduction rate 1.33 to prevent a gradual collapse in the population size. Let us first apply a change compliant with Haldane's model to this population, and then apply a change compliant with the concept of intraspecific competition, to better see their differences.
Consider, in compliance with Haldane's model, that there is a "hostile" change in environmental selection pressures with selection coefficient s=0.1. Conveniently, this value happens to coincide with intensity I=0.1. In terms of absolute fitness, this means that the average fitness of the population drops to (1.0 - 0.1) * 0.75 = 0.675. Now, there is 32.5% juvenile mortality, and a requirement for 1.48 reproduction rate (0.48 reproductive excess). Unless the population can meet this reproductive requirement, it risks extinction. This is the reason why intensity of selection needs a limitation in Haldane's model. While the required reproduction rate might not be very severe in this case, it would obviously be harder to meet if we had assumed an initial fitness lower than 0.75, or selection more intense than 0.1. Now, what about the *substitution* part? Haldane's model assumes that among the genetic variance in the gene pool, there is a one or more alleles, each with a tiny intial frequency, whose combined phenotypic effects happen to compensate for the fitness penalty imposed by the environmental change. That is, there is an optimal genotype which continues to have absolute fitness 0.75 despite the environmental change. This genotype becomes fixed in the population, as standard population genetics demonstrates. The time required for this substitution, however, is dependent on the intensity of selection: the more intense selection we have, or in other words, the bigger the difference between the fitness of the optimal genotype (0.75) and the fitness of the vast majority of the population (0.675), the less time the substitution takes. So there is a conflict of interests between avoiding extinction and a high rate of allele substitutions.
But consider, instead, that this population has territorial behavior. Individuals compete over the finite territories in their habitat, and they also try to win the *best* territories, at least to some extent, because not all territories support reproductive success equally. In practice, this means that the fitness of an individial is partially dependent on the territory it is inhabiting (and guarding against others in its species). Not having a territory at all might bring fitness all the way down close to zero, but let us assume here that everyone gets a territory. Instead, assume inhabiting a "decent" territory only brings absolute fitness 0.65 and inhabiting a "good" territory brings absolute fitness to 0.85. Also, let half of available territories be "decent" and half be "good", which brings the average fitness of the population to 0.5*0.65 + 0.5*.85 = 0.75, in accordance with our initial assumption. Initially, no individual is better than others in winning a good territory instead of a decent one. But this is a naive assumption, of course. Surely there are genetic differences which affect territorial contests, just as there are genetic differences affecting one's ability to overcome challenges imposed by the environment surronding the population. So, let us change this assumption. Let there be an allele, initially with a tiny allele frequence, which gives an individual a 75% chance of winning a "good" territory, instead of the earlier 50% chance. In practice, this means that this subpopulation of individuals has an average fitness about 0.75*0.85 + 0.25*0.65 = 0.80. (Initially the subpopulation is tiny enough so that they will in practice only fight against the old genotype, and not against each other - yet.) This new genotype being the new optimal genotype, there is a tendency in the population to fix the genotype, just as in Haldane's scenario. (In the previous scenario, the selection coefficient was s=0.10. Here it is initially about s=1.0-0.75/0.8=0.06.) Initally, the presence of the new genotype doesnt really affect the fitness of the less optimal genotype; the new genotype is so rare that individuals of the old type barely ever get beaten by it in territorial contests. So the fitness of the old genotype initially goes just barely below 0.75. But notice, then, that the average fitness of the population remains exactly the same, in *dramatic* contrast to Haldane's model. Whatever the new genotype wins, in terms of fitness, is lost by the old genotype, but these gains and losses are balanced by the number of individuals in each group:
avg fitness of new genotype * frequency of new genotype + avg fitness of old genotype * frequency of old genotype = 0.75 = avg fitness of the whole population
Also notice that when the new genotype increases frequency during the subsitution process, the fitness of both genotypes starts to decrease (but not the average fitness of the population, because the optimal genotype remains above 0.75!). This is because it becomes more likely to meet the optimal genotype in territorial contests, which hurts both the fitness of the old genotype and the fitness of the optimal genotype itself. When the frequency of the old genotype has become tiny, the (average) fitness of the new genotype has "returned" to 0.75; members of the new genotype now practically always fight against each other with just the old 50% winning chance. The fitness of the old genotype has dropped to 0.25*0.85 + 0.75*0.65 = 0.70; they practically always fight against the new genotype. So, at the time the old genotype becomes extinct, the fitness distribution in the population has returned back to the initial state. Such "no-one-wins-in-the-long-run" result is common in all evolution, I believe. Adapting to environmental changes, too, tends to bring only temporary benefits, especially when considering an arms race between predator and prey.
In this example of intraspecific competition, at no point was there any reason to assume (a) a change in the average fitness of the population or (b) a change in the reproductive rate of any subpopulation. Yet an allele substitution took place. Using Haldane's cost unit of excess "genetic deaths", there simply was no cost. Using ReMine's cost unit of reproductive excess, the "cost" was exclusively "paid" by so-called "effective producers". The latter means in practice that while no individual increased its reproductive rate, the allele frequencies still changed (the "reproduction rate" of the new allele increased above 1.0 and the "reproduction rate" of the old allele decreased below 1.0).
I hope these thought plays finally illustrate the difference between the concept "intraspecific competition" and between Haldane's model. And before anyone asks: yes, I have tried to explain this concept to ReMine over and over again. In our discussions, he did not address this concept by a single word and completely ignored even the fact that the concept was mentioned. To my best knowledge, he has not aknowledged the existence of this concept anywhere. I'm looking forward to ReMine finally reacting to it, and I hope Nunney's model forces him to. Although I, too, am intererested to see exactly what are the details of Nunney's model --Esko Heimonen 08:46, 31 May 2007 (UTC)

The effect is that a minority now has an advantage over the majority, and the biomathematical problem is the same: given a degree of "more fit-ness", how long does it take for the minority allele to be fixed in the population?"

A separate comment on the quoted sentence. The time (in generations) required by any individual subsititution depends (a) on the initial frequency of the substituting allele and, more interestingly, (b) on the selection coefficient of the substituting allele. Of these, (a) is a value dependent on the population size and on the mutation rate for the locus in question. The more interesting value (b) is, first of all, limited by reality: as Haldane notes, the selective effect of any single allele is rarely very high. Another limit for (b) is the whole point of Haldande(1957): too intense selection creates a risk of extiction. This latter limit does not concern just a single allele but the whole combined intensity of all allele substitutions taking place in the population at any one time. Haldane's paper shows that in his evolutionary scenario, and asssuming a fairly low intensity, it makes little difference in terms of substitution rate to study a bunch of separate loci and their substitutions (concurrent substitution), compared to a simplified model of just one locus with a selection coefficient which combines the fitness effects of many separate loci (sequential substitution simulating the fitness effects of concurrent substitution). (Using multiplicative or additive fitness interaction, it makes little difference which one when I<=0.1.) In intraspecific competition, concurrency makes a whole lot of difference. Intensity of selection related to intraspecific competition is not limited by the risk of extinction, so the combined effect of fairly intense selection at hundreds of loci could reach a value that would seem ridiculous if simulated by a single locus (as Haldane effectively does in his model). So I claim that Mdotley's focus on the substitution time of a single allele is not warranted. Although perhaps a valid oversimplification in Haldane's model, it applies poorly to intraspecific competition. In the latter, I think one should consider the concurrent substitution taking place in the whole genome, i.e. over several loci. For example, one should be at least prepared for the possibility that the total substitution rate could in principle exceed one per generation, which is impossible at a single locus. --Esko Heimonen 10:52, 31 May 2007 (UTC)

[edit] POV

This is ridiculous. It has no rebuttals to Haldane's dilemma at all, and in the first paragraph sasys that the dilemmma is unsolved and completely ignored by scientists as if that was fact. Adam Cuerden talk 04:51, 30 May 2007 (UTC)

[edit] More on the magic number 1,667

For further discussion see Haldane's Non-Dilemma at The Panda's Thumb. MrDarwin 14:52, 2 July 2007 (UTC)

Excellent discussion there, making it clear that this article is at the very least misnamed. I suggest that it be deleted, and an item about it be added to a section under Intelligent Design listing fallacious and erroneous arguments -- a very long list. And perhaps a section can be added to Walter ReMine's page documenting his blatant falsehoods. -- 71.102.194.130 15:28, 2 July 2007 (UTC)
The PT article in question does not give a reference to any peer-reviewed study on the number of adaptations in our regulatory sequences. According to evo-devo, it is changes in gene expression that has given us our brain size/structure, upright walking, hand dexterity and speech capabilities. I dare say it is anatomical features like these that not only creationists but mostly everyone else is interested in human evolution, as opposed to e.g. the unique features of our immune system. Hence, I find declarations like the one above premature and, frankly, somewhat uncritical.--Esko Heimonen 06:45, 3 July 2007 (UTC)

[edit] This article is a scandal

This article demonstrates only a determination to obscure Haldane's Dilemma from public view.

  • The key figure -- a limit of 1,667 beneficial mutations to explain human evolution -- arises solely from evolutionary theory, evolutionary genetics, and J.B.S. Haldane. This key figure was repeatedly expunged from the Wikipedia article, leaving readers with no idea about the severity of Haldane's Dilemma.
  • The above key figure was never revealed to the general public, and this fifty-year history of suppression was likewise expunged from the Wikipedia article.
  • The renowned evolutionary theorist, G.C. Williams (1992), wrote, "In my opinion the [Haldane's Dilemma] problem was never solved, by Wallace or anyone else." That admission was repeatedly expunged from the Wikipedia article.
  • Evolutionary geneticists, James Crow and Warren Ewens, peer-reviewed Walter ReMine's paper and publicly acknowledge it is correct. Nonetheless, the Wikipedia article repeatedly expunged all the clarifications given in ReMine's paper, and repeatedly promotes the various confusion factors identified in ReMine's paper. In other words, the Wikipedia article promotes known confusions, and withholds known clarifications, as known to James Crow and Warren Ewens. The Wikipedia article presents a garbled basis for the problem.
  • Instead of giving real insight, the Wikipedia article wearies readers into giving up -- through its needlessly tedious mathematical derivations, and its opaque definitions of key terms. Rather than illuminate the real problem of Haldane's Dilemma today, the article wearies readers with a relatively fruitless and misleading journey into the "origin of the term" in 1963. The article seems intentionally designed to wear-out readers, rather than deliver understanding.
  • The Wikipedia article falsely pretends Haldane's 1957 description is the clearest available -- as though no clarifications occurred since then.
  • The Wikipedia article does not reveal the great breadth of confusion and contradiction that remains unresolved among evolutionary geneticists on this topic. The evolutionary literature on Haldane's Dilemma is a quagmire of confusion and contradiction among authorities, and the Wikipedia article conceals it.
  • The Wikipedia article does not reveal the problem of Haldane's Dilemma. Nor does it provide solutions. Instead the Wikipedia article falsely pretends Haldane's Dilemma was solved.

The Wikipedia article demonstrates how evolutionists actively suppress Haldane's Dilemma from public view -- and that is the only reason this Wikipedia article is worth reading.

Compare it with a more revealing article. WalterR 17:47, 31 August 2007 (UTC)

Here are some counter-arguments to both the above post and to the link provided in the post.
  • Haldane's number n=300 (generations per substitution) is not from "Haldane's calculations". (See ReMines's suggested article: I find it "revealing", too, but for entirely different reasons.) I challenge Walter ReMine to demonstrate the existence of such "calculations". Instead, Haldane proposes a model for calculating the costs of natural selection. Based on this model, Haldane presents a vague estimate for the mean intensity of selection (in an average lineage?). In fact, allow me to quote directly the "calculation" for the magic number 300 from Haldane(1957).
To be concrete, if a species had immigrated into an environment where its reproductive capacity was half that obtainable after selection had run its course, so that I=ln2=0.69, n would be 43. This represents, in my opinion, fairly intense selection, of the order of that found in Biston betularia, where it has a rapid effect because it was concentrated on a phenotypic change due mainly to a single gene. I doubt such high intensities of selection have been common in the course of evolution. I think 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.
In other words, Haldane does not provide any rigorous calculation for any exact value for the intensity of selection, and, consequently, for the number of generations per substitution. Instead, Haldane prodives what he finds to be a "probable figure" for the mean intensity of selection. Would Haldane be prepared to increase this mean value to, say, I=0.2 in some lineage if data so suggested? Certainly, as nothing in his "calculations" stands against it. Thus, ReMine effectively tries to mislead his readers into believing that there is some kind of a rigorous mathematical reason forcing Haldane to set the number 0.1 in stone. There is none, and he does not set it in stone. He only makes a vague proposition.
  • Haldane's estimate for the total selection coefficient s=0.1 means that a population needs a reproduction rate 1/(1.0-0.1)=1.11 to not risk going extinct (2 extra offspring per 10 mothers). It is clear that even the slowest breeders like elephants can do much, much better than that. In this light, it is amusing that Walter ReMine accuses Haldane of ignoring a wide variety of causes that might slow down evolution. To the contrary, it is quite clear that the reproductive capacity of any species would meet this requirement with no effort at all. Thus, Haldane clearly applies some other criteria in addition to reproductive capacity when giving his estimate for I. ReMine should think about this when declaring what "Haldane's Assumptions" were.
  • Modern population genetics largely disagrees with the way Haldane(1957) dealt with density-dependent evolution. In his discussion on moths, Haldane assumes that growing population density leads into a flaring parasite infection, which kills larvae randomly. I.e. in his example Haldane divides the "juvenile" phase further into two stages, where first random mortality is applied and only then selection. While this might apply well to some moth population, it is hardly well grounded to apply it universally. It is likely that assuming random density-dependent mortality is largely how Haldane explains the low value I=0.1. Many modern authorities argue that density-dependent mortality is not random in general, but subject to selection. They also argue that density-dependent mortality and other kinds of mortality may well occur in parallel. This effectively means that density is reduced by other kinds of mortality before density-dependent mortality (in the extreme case, sheer starvation) takes place in all severity. This is what soft selection means. It is not an "accounting trick". It's simply saying that if by getting killed by a predator you might prevent someone else in your species from starving or from not finding a free territory to live in. (Note that neither starving nor being unable to find or conquer a territory would be random affairs, though -- they could still be dependent on your genes, even if relaxed population density certainly helps.)
  • Haldane assumes that all selection pressures are imposed on individuals from the outside of the population. Hence the problem of avoiding extinction when selection is intense, and, consequently, when the rate of allele substitutions is high. Thus, Haldane entirely omits intraspecific competition. I have already explained this concept, and its difference to Haldane's selection model, in length above. While we do not know for certain how important intraspecific competition has been for hominid evolution, I would suggest that it is indeed a notable factor for social primates, who spend a lot of time and energy on e.g. maintaining or improving their social status and for fighting with other clans over territory.
  • ReMine seems to assume that stasis plays a role here. In fact, one of the reasons why Gould and Eldredge proposed the concept of punctualism was to explain why, in many lineages, the measured rates of evolution seemed to be much faster than required by the estimate of total evolutionary change in the history of those lineages. Haldane certainly could not have taken punctualist ideas into account in 1957, but Walter ReMine is dishonest by trying inject punctualism on top of Haldane's model. Perhaps the mean intensity of selection is very, very small in most lineages when calculated over several successive geologic periods. But such a mean value need not automatically limit the intensity of selection at some specific lineage and epoch of ReMine's choice. Specifically, ReMine's argument is focused on hominid evolution, and he apparently tries to inject stasis into this evolution independently of data. Would Gould and Eldredge agree with him, that hominid evolution is a classic example of stasis? I have my doubts. Does stasis pose a problem for evolution if it is applied in the more general fashion it was first proposed? I hardly think so because the idea was inspired by lots and lots of data, and any estimates for the relative importance of stasis are derived from data. The punctualists' "magic number" 99% is not set in stone, because it is an estimate based on observations, and it is freely adjustable based on future observations. Just as is Haldane's "magic number".
  • ReMine claims that Haldane somehow favored concurrent evolution at many, many loci, assuming tiny selection coefficient for each locus. As is clear from e.g. my quote above, Haldane did not restrict himself to tiny selection coefficients. In fact, I'm not aware of any example where Haldane would have promoted such a notion. To the contrary, Haldane's examples seem to discuss no more than 10 loci, which happen to have huge selection coefficients for a multi-locus scenario. Specifically, Haldane did not embrace nearly-neutral evolution, as such an idea hadn't even been presented in 1957. Walter ReMine, however tries to make the most of this idea, in order to turn a model on selection into a model on drift in his revisionist history. We have a lot to learn from Walter in avoiding "confusion".
  • Haldane did not promote additive fitness interaction between loci. He only approximated multiplicative fitness with additive fitness in some of his calculations -- he used a lot of approximations in them. This practice is well grounded when the total intensity of selection is no more than 0.1. Beyond that, the approximation starts to fail. For example, if there are 10 selection coefficients each with s=0.1, the resulting additive fitness effect is 1.0 - 10x0.1 = 0, but the corresponding multiplicative fitness is (1.0-0.1)10=0.35 (requires reproductive excess 1.9 to counter, i.e less than six offspring per mother, still within the reach of even elephants).
  • Why does Walter ReMine continue to claim that his paper was acknowledged "correct" by its reviewers, after he was unwilling to discuss the matter with David Sloan Wilson in the past on this very page? According to Wilson, ReMine is likely misleading his readers on purpose.
  • Most importantly, why does Walter ReMine not tell his readers how many adaptations ARE needed for producing Homo sapiens from an ape-like ancestor? For example, development biologist Sean Carroll's estimate is a few thousand adaptations, including changes in exons and changes in gene regulation. Would such a number, with the same order of magnitude as ReMine's "magic number" but still considerably higher, satisfy ReMine? I suspect that this is not the case. ReMine's comment on whether 1,667 adaptations might be sufficient is: "Who are they kidding?" To me, this suggests that ReMine tries to assure his readers, by mere rhetorics, that it is somehow self-evident that there should be an order of magnitude, or perhaps many orders of magnitude, more adaptations. It also seems to me that ReMine does not want to be explicit about it. The latter is because modern experts would likely disagree with him and require him to produce data for his claim (and he has none). On the other hand, if Walter ReMine agreed that the required mutations are likely numbered in thousands at most, his arguments would lose all credibility. For starters, Haldane's number 300 is not set in stone by any "calculation".
  • Equally importantly, where can one find a calculation where Walter ReMine applies his "cost types" to a real population, coming up with a "calculated" upper limit for the intensity of selection compatible with survival? When it comes to the "cost of mutation" for hominids, I advice ReMine not to rely on Nachman(2000) too literally, as it still assumes we have 70,000 genes (U=3) when HGP has shown that we in reality have about one third of that (U=1).

--Esko Heimonen 15:32, 3 September 2007 (UTC)

Futhermore, I think that ReMine grossly mispresents the effect of Haldane(1957) in the history of evolutionary biology.
The cost model presented in Haldane(1957) was important in promoting the neutral theory. This importance is clearly stated in undergraduate university textbooks on evolutionary biology. Why Haldane's cost model is not mentioned in high school textbooks should be self-evident: students never get as far as mathematical population genetics.
While Haldane(1957) apparently did pose a "dilemma" in the past, it is (a) different from what ReMine claims and (b) largely solved. The real impact of Haldane's cost model was that adaptations could not explain the whole of molecular evolution. But genetics discovered that most of the DNA in our genes is not under constraint, i.e. only a small part of our genome codes for proteins (and even those parts experience many silent mutations). In other words, adaptations do not need to explain all of evolution because the actual genetic information forms only a fraction of the DNA in most species. Most of our DNA can change freely with little or no effect on phenotype.
Besides, it seems that molecular evolution is largely caused by gene duplications. ReMine insists that we should concentrate on point mutations, because they are the most common type of mutation. But ReMine ignores the fact that a gene duplication can involve tens of thousands of nucleotides (counting introns and exons). So duplications don't need to be common at all, and they still may well beat point mutations in importance in molecular evolution. For example, I believe that the better part of molecular differences between humans and chimps are attributed to the about 1,500 fixed duplications that these species have experienced since their divergence. So far as ReMine's argument is about molecular evolution, then gene duplications and neutral evolution can easily explain the quick molecular changes, and Haldane's cost model has little to say about it. Presumably this debate is about morphologic adaptations, however, because otherwise Haldane's paper would likely not be under focus. (Of course, gene duplications likely play a relevant role in morphological adaptations as well, because they are a source of new genes. But ReMine's argument deals with the quantity of adaptations rather than the quality. Admittedly, protein dosages seem to be selected, in which case some gene duplications could fit under Haldane's cost scenario.)
If ReMine claims that there exists such a thing as "Haldane's dilemma" which concerns morphologic adaptations, then ReMine should d-e-m-o-n-s-t-r-a-t-e that it exists, before demanding others to s-o-l-v-e it. Has any evolutionary biologist announced that the cost of natural selection prevents human evolution? I doubt it. If no one has demonstrated the dilemma, why should educators make it public?
If ReMine wants to demonstrate the dilemma, he should first produce data which tells us the number of adaptations needed for a specific linege during a specific time period. He should also demonstrate that there indeed is an unavoidable "magic number" telling the maximum number of adaptations, which causes a problem for the evolutionary scenario being studied. Only then is it possible to see the dilemma. So far Walter ReMine has done neither. At best, he has made the weak point that there is possibly a conflict between the cost of natural selection and the rate of hominid evolution. But with little else currently known except perhaps the number of changes in the exons of our ancestors (mere few hundred adptations, it would currently seem), it is impossible to know whether this possibility is also the reality.

--Esko Heimonen 10:16, 8 September 2007 (UTC)

XXXXXX

Nearly the entirety of Esko Heimonen's arguments (above) are irrelevant here, as they are his personal arguments unpublished anywhere, much less in permanent form or peer-reviewed. If he thinks he has a solution to Haldane's Dilemma, he ought seriously publish them. If he thinks he can rebut ReMine, he ought seriously publish them. If he thinks his arguments have any value whatever, he ought seriously publish them. But his arguments will likely never be published, because most of them are false, and he indicates elsewhere that he has no intention of publishing.

Heimonen's arguments have virtually nothing to do with this Wikipedia article, or with this discussion page, which should be about the Wikipedia article. Heimonen did not seriously address any of the objections to this Wikipedia article, and nothing he wrote can fix this Wikipedia article. The Wikipedia article is a scandal, and Heimonen's lengthly essay (above) is an attempt to divert attention away from the scandal. The Wikipedia article documents the evolutionist contempt for revealing Haldane's Dilemma to the public. That is the only reason it is worth reading. Compare it with a more appropriate article.

Like the Wikipedia article, Heimonen makes the following mistakes. He falsely pretends no clarifications have been made (by ReMine or anyone else) since Haldane 1957. He uses known confusion factors, such as elimination of the previous-type individuals (i.e., through genetic death, mortality, and selection intensity) -- Haldane's Dilemma is better understood by focusing on the favored individuals and their required reproduction rate. He ignores all other costs, (such as the cost of mutation, the cost of segregation, the cost of random loss, and the cost of continuity), and thereby gives the false impression that large amounts of the specie's reproduction rate can be applied to paying the cost of substitution. He falsely pretends Haldane's Dilemma is solved. WalterR 03:24, 8 September 2007 (UTC)

XXXXXX

I'm happy to let Walter ReMine have the last word here, because I find his comments and his persistent avoidance quite self-defeating in itself.

--Esko Heimonen 10:16, 8 September 2007 (UTC)

[edit]     

Esko Heimonen wrote:

Why does Walter ReMine continue to claim that his paper was acknowledged "correct" by its reviewers, after he was unwilling to discuss the matter with David Sloan Wilson in the past on this very page? ...

While I'm flattered to have been mistaken for David Sloan Wilson I'm not so sure he would be equally flattered to have been mistaken for me, so I guess I'd better own up to being the author of the David Wilson half of the exchange with ReMine referred to above.

According to Wilson, ReMine is likely misleading his readers on purpose.

I agree that much of what Mr ReMine writes is erroneous and is therefore likely to mislead some of his readers, but I have always presumed that he genuinely believes he is providing accurate information. Nor do I believe there is any implication to the contrary in what I have written above, but just in case, let me make it perfectly clear that none was intended.

—David (J) Wilson (talk · cont)

Concerning Haldane's Dilemma, David (J) Wilson is more careful than most of the Internet commentators, and that is much appreciated. However, he is overly silent concerning this condemnable Wikipedia article which seeks to obscure Haldane's Dilemma from public view. Also, he has not identified any errors in ReMine's work, (more precisely, Wilson's only attempt is rebuffed by ReMine, see ReMine's footnote #4). WalterR 21:10, 10 September 2007 (UTC)
I have responded to WalterR's comments on my talk page here.
—David Wilson (talk · cont) 23:40, 11 September 2007 (UTC)

[edit] 1600 may be more than enough

According to the draft chimpanzee genome the total number of differences that are the result of selection is about 585. even if these are unevenly divided between the chimp and human lines, there is more than enough room for all of them.(I Know professor ReMine was trying to be generous with his number, but until he comes with a more precise estimate, there in NO dilemma). 66.73.48.200 20:12, 10 October 2007 (UTC) fingal

This talk page is not the place to debate Haldane's Dilemma. Rather, it is the place to debate this Wikipedia article, which currently seeks to obscure Haldane's Dilemma from public view. Moreover, no one has published any such solution to Haldane's Dilemma, and until they do, it deserves no place in this Wikipedia article. Lastly, the central question -- Is 1600 substitutions enough? -- is not even mentioned in this Wikipedia article, which makes this article even more condemnable. This article demonstrates evolutionists' contempt for revealing Haldane's Dilemma to the public. WalterR 21:07, 15 October 2007 (UTC)
It seems as if the entire article is WP:SYN, and should trimmed to a raw statement of the dispute without attempt to justify it either being a "dilemma" or not. — Arthur Rubin | (talk) 21:09, 15 October 2007 (UTC)
I think this article should feature more prominenently in the "External links" section: at present, it's a link on the TalkOrigins page which is in turn linked in the article. Like Esko Heimonen's excellent posts above, it explains why Haldane's Dilemma is no longer considered to be a problem. As for WalterR's insistence on a "published" solution: as a solution has been "published" in various places (including this talkpage), I presume what's demanded here is publication in a peer-reviewed journal such as Science or Nature. But therein lies a problem: I think it's unlikely that such a journal would publish a solution that is already common knowledge and in the "public domain". Wouldn't that be rather like publishing an article claiming that the apparent movement of the planets in the night sky can better be explained by a heliocentric model rather than a geocentric one? Scientific understanding has moved on too far for such an article to be considered "newsworthy" nowadays. --Robert Stevens 10:58, 16 October 2007 (UTC)
I do not claim that "Haldane's Dilemma is no longer considered to be a problem". So far as "Haldane's Dilemma" refers to Kimura's various challenges to conventional population genetics, Haldane's Dilemma may well continue to be a problem unsolved by selectionists. In fact, some suggested selectionist solutions to the problem seem to completely overhaul conventional population genetics, or "beanbag genetics" as Ernst Mayr called it. If we emphasize ideas like density-dependent selection, frequency-dependent selection and epistatic fitness interaction between loci, basically nothing is left of Haldane's beanbag genetics. The fitness of an individual can no longer be approximated by multiplying or adding static fitness coefficients over multiple loci. Fitness is now a much more complicated concept depending on, for example: (a) the entire genome of an individual in an epistatic, "holistic" sense, (b) the frequencies of various genotypes in the population, and (c) how densely packed the population is. In effect, Haldane's and Kimura's attempt to apply "beanbag genetics" over entire genomes becomes ridiculous -- even if "beanbag genetics" still managed to sometimes produce successful approximations when applied to individual loci or simple systems consisting of just a few loci. If this is the case, I think it would be valid criticism to ask modern population geneticists whether they still support the old "Haldanian" paradigm in their writings, or whether they have accepted a "Mayrian" approach when studying large amounts of loci. This kind of criticism would be extremely hard, because it essentially says that population genetics is ridiculous in pretending to be able to model the evolution of entire genomes. But this kind of criticism has not been the substance of ReMine's writings, of course. To summarize, I think it is fair to say either that Haldane's Dilemma remains unsolved or that Haldane's oversimplified model has been overthrown altogether.
So far as "Haldane's Dilemma" refers to ReMine's specific claims about human evolution, I fail the see a well-substantiated problem in the first place. Currently there is no solid reason to believe that more than 1667 morphological adaptations have taken place during human evolution -- although I don't think that there currently is a solid reason to believe the opposite, either. As long as exons are concerned, the number seems to been some hundreds. And we don't currently know the number of regulatory adaptations. As for the "magic number" 1667, there is no solid mathematical reason behind it. It is a fairly vague estimate. Even though the very order of magnitude in this estimate seems sufficient to demonstrate that Kimura's challenges can not be addressed by selectionists without changing Haldane's core assumptions, the estimate hardly guarantees that, say, a few thousand adaptations could not have become fixed in the human lineage during the past few million years. (And many more, if we abandon "beanbag genetics".)
In my opinion, one major problem in this wikipedia article has been, in the past, that the article accepted at face value ReMine's hijacking of the term "Haldane's Dilemma". Yes, Haldane(1957) is the catalyst behind both ReMine's arguments and Kimura's arguments, but that fact alone does not make the arguments equivalent. Kimura's arguments are that molecular evolution is at least in some cases many orders of magnitude too fast for being driven by selection, and that selection can't maintain hundreds of polymorphisms in the gene pool. If ReMine's argument truly was that human evolution required hundreds of thousands or even millions of adaptations, then I could understand why he compares his argument to the problem (Haldane's/Kimura's Dilemma) that selectionists have tried to solve. But so long as ReMine does not explicitly state this to be the case, I think he misuses the term Haldane's Dilemma, and quote-mines this term in a potentially dishonest way. Or course, if ReMine did explicitly assume millions of adaptations during human evolution, he would face the task of producing evidence for his assumption. Additionally if ReMine commits himself to the real Haldane's Dilemma, he should himself try and answer Kimura's challenge concerning segregational load. If we grant that a Designer designed a few hundred polymorphisms for a population, how does he maintain those polymorphisms without resorting to either neutralism or frequency-dependent selection? --Esko Heimonen 15:32, 16 October 2007 (UTC)
I want to clarify this earlier claim of mine: While Haldane(1957) apparently did pose a "dilemma" in the past, it is (a) different from what ReMine claims and (b) largely solved. While I still agree with (a), I now disagree with (b). Kimura's argument was specifically that some amino acid sequences change faster than we should expect if the changes became fixed due to selection (and if the assumptions of beanbag genetics were valid!). In the 60's, amino acid sequences were available while DNA sequences were not. Most of our DNA is now known to be non-coding, explaining why the majority of our DNA, as well as our DNA length, can often change fast with little or no constraint. But this observation alone does not weaken Kimura's argument, because the argument specifically considers fast evolution of coding sequences. I apologize for the confusion. --Esko Heimonen 21:54, 16 October 2007 (UTC)
While we're on the subject of clarification: Esko, do you still agree that ReMine's version is either "largely solved" or that he has failed to demonstrate that there's a problem? (which is tantamount to the same thing: he has the burden of proof, so if he can't prove it, the problem is effectively "solved" from the scientific perspective). Because that's what my earlier post should have alluded to: I can see why the "dilemma" would still be an issue in some cases, but I'm not surprised that Science or Nature haven't published a solution to "ReMine's Dilemma". --Robert Stevens 08:16, 17 October 2007 (UTC)
Agreed, basically. I'm nit-picking here. It comes back to the old question on how one should approach creationist arguments. Saying that the "problem has been solved" may create the false impression that there initially was a valid problem to be solved, i.e. "a controversy". But there never was any, as far as I can see. Instead, a different (and valid) problem was falsely associated with the non-existing (or at least undemonstrated) problem, to give the latter credibility and support from confusing quote-mining, without actually taking the trouble of rigorously demonstrating it. --Esko Heimonen 10:38, 17 October 2007 (UTC)