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Talk:Haldane's Dilemma

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Early remarks

Despite this article referring to Walter ReMine in the third person, it was actually written and inserted by Walter ReMine himself. Much of the text consists of ReMine referring to his own claims, with no mention that these claims have been addressed, and in some cases refuted (there are, for instance, many references to Haldane's work in mainstream sources - but ReMine claims it is unacknowledged). It is nothing more than self-aggrandisement plus a plug for his website and his book.

The most glaring problem is that of the limit of 1667 beneficial mutations in the human lineage from the last common ancestor with chimps. ReMine states that this is a limit, and says that this is a problem, but explicitly avoids explaining why it is a problem. The universally accepted solution to this 'dilemma' (which ReMine claims doesn't exist) is that there is no problem unless it can be shown that 1667 mutations is insufficient - and it never has been shown. Thus the dilemma may be illusionary.

I recommend this article either be rewritten by an independent reviewer, i.e. someone other than ReMine, or deleted.

Roy 05:26, 11 June 2006 (CDT)

I did my best to describe the dilemma itself and broaden the article out -- any ideas from either Walter or Roy? Ungtss 09:50, 11 June 2006 (CDT)
Roy has this completely backwards. Roy said that, "there is no problem unless it can be shown that 1667 mutations is insufficient". But when the very point in dispute is whether human evolution actually happened, there really is a problem unless it can be shown that 1667 mutations is sufficient. Not the other way around. The burden of proof is on science to explain how the 1667 mutations made it happen. For Roy to say that the inability of science to explain the facts isn't a problem shows that he is assuming an answer to the very point in dispute. This makes evolution unfalsifiable; true by default.
Roy may counter that he is allowed to assume evolution as an answer here because other fields of science substantiate it better. But if he does this, that is a catastrophic problem for science: What if EVERY field of science defers to every other field of science to assume evolution as the answer? Evolution ends up with ice at its foundation on a hot day. Roy's strongest argument here turns out to be an ad hominem against "Walter ReMine." But everyone should know, ad hominem arguments don't prove anything. David Pesta 15:29, 2 September 2011 (PDT)

The anonymous evolutionist, "Roy", seems to think his anonymity gives him credentials for obscuring Haldane's Dilemma -- which is what evolutionists have done for decades.

  • Evolutionists never communicated Haldane's Dilemma to the general public in any coherent fashion, Roy is mistaken to suggest otherwise.
  • Haldane's Dilemma is contradictory and unresolved in the evolutionary genetics literature, and that fact is generally "unacknowledged" by evolutionary geneticists -- again Roy is mistaken to claim otherwise.
  • Roy claims, "The universally accepted solution to this 'dilemma' is that there is no problem unless it can be shown that 1667 mutations is insufficient". Unfortunately, evolutionary geneticists publish no such statement, especially not "universally" -- and emphatically not to the general public. Roy is mistaken to claim otherwise.

Roy's essay attempts to obscure Haldane's Dilemma, delay it, put it off, tire out readers with needless diversions -- rather than tell readers the problem straight up. His essay is not remotely NPOV. Wremine 19:05, 11 June 2006 (CDT)Wremine

Which essay is not npov -- the one Roy wrote on the talkpage, or the one I wrote into the article? Ungtss 19:09, 11 June 2006 (CDT)

The essay as it currently exists is unacceptable -- for its various factual errors and confusions -- also it makes the problem obscure and compels readers to slog through extraneous minutiae that are unnecessary to seeing the problem. That is not NPOV. Wremine 19:24, 11 June 2006 (CDT)Wremine

Assuming you mean the main article -- that was not Roy -- it was me -- and I'd like to work with you in making it a mutually agreeable article. I made the changes I did for the following reasons: The article as it stood did not explain what the dilemma was -- it simply stated your conclusions about its implications for evolution. I tried to give some background before we did our analysis. My intent was not to "bog the reader down" -- but to make sure a new reader understands what it means. But i'm not an expert my any means -- please feel free to make any changes you find to be appropriate -- but keep in mind that the average reader doesn't have the background you have in this topic -- keep it simple for us simple folk:). Ungtss 19:37, 11 June 2006 (CDT)

The current essay has the following difficulties (listed here in the order of their appearance):

  • Haldane’s Dilemma is not a “paradox”, it is a problem. A real, unsolved problem. It is not something easily cast aside as a novelty, such as Zeno’s paradox. Haldane’s Dilemma is described as a mere “paradox” by evolutionists, in an attempt to minimize the issue.
  • Haldane did not actually, explicitly, “introduce” Haldane’s Dilemma, rather it later became vocalized, and named, by others.
  • Haldane’s Dilemma is not about the “efficacy of natural selection” See here. If we had to identify the nexus or source of the problem, it is more accurate to say, “Haldane’s Dilemma challenges the efficacy of evolutionary genetics.”
  • The material about “The dilemma” misrepresents and obscures the problem. The problem is not, “In higher animals, it is much more difficult for natural selection to select for more than one trait at a time, because advantageous genes may not be found in the same individuals.” Where’s the problem? The problem disappeared. No reader will see a problem there. Whoever wrote that sentence into the essay appears to have bought into evolutionist misinformation and obfuscation. This essay should not be based on the evolutionists’ attempts to obfuscate Haldane’s Dilemma. Understand that evolutionists NEVER revealed Haldane’s Dilemma in plain English – they could easily have done so, but they didn’t. We cannot use the evolutionists’ (mis-)definitions of the problem, such as the above example.
  • All the material about “cost of substitution” should appear later, after readers are told what the core problem is. Minutiae about the “cost of substitution” are entirely unnecessary to understanding the core problem. Are 1,667 mutations enough?
  • The “simplified example” should appear later, after readers are told what the problem is. Minutiae about the “cost of substitution” are entirely unnecessary to understanding the core problem. Also, the current “simplified example” itself is needlessly confused, and a better example is available – see the next point.
  • The focus on elimination and “dying” and “killing off” and “kill rate” of the previous genes is needlessly confused. The issue is not the elimination of the previous genes, but rather the growth of the substituting genes – and the extra reproduction rate necessary to make that happen. That’s what the Cost of Substitution is.
  • The essay emphasizes (for an entire paragraph!) that, “This (kill) rate is extremely difficult to measure in practice …” However, the species reproduction rate is quite easy to measure – and that’s what matters. Let me put it this way. A “cost” (such as the cost of substitution) is calculated (not measured), the “payment” is the species actual reproduction rate (which is measured). The essay confuses those.
  • The essay says “Haldane stated the 1,667 limit.” That is not precisely true. Haldane did not, himself, explicitly state the limit of 1,667 substitutions. Indeed, such a clear statement of the problem is exceedingly rare (non-existent?) in evolutionary literature. Rather, Haldane, and most evolutionists to this day, stated things so as to obscure the problem from view. That is, they give a substitution rate, such as “one per 300 generations.” When stated that way, no ordinary person can see any problem. Instead, it was I who explicitly stated the “1,667 limit,” though that figure comes in every meaningful sense from Haldane (plus evolutionary leaders’ ordinary claims about human evolution, plus simple arithmetic: (10 million / 20 / 300=1,667). Does simple arithmetic count as “original research”? That limit is not “my conclusion” (as someone suggested), because I have virtually nothing to do with it, other than state it clearly. Instead it is evolutionists’ conclusion, or better yet, Haldane’s conclusion – only I wrote the words, because evolutionists were afraid to. None of that “conclusion” belongs to me – it all comes from evolutionists. New readers can understand the problem very quickly, without first being dragged through minutiae about the “cost of substitution”, etc.
  • The essay says, “Thus, the mechanisms of evolution could only provide for 1667 beneficial nucleotide differences between humans and the last common ancestor with the chimps.” That is not accurate.
  1. The starting point is an alleged ape-human-like ancestor ten million years ago – not the “last common ancestor with chimps.” (Again, ten million years is being very generous to evolutionists, and this point should not be lost.)
  2. The issue is not about net “differences” between starting point and ending point – because that does not account for the many substitutions that likely need to occur at the same nucleotide site, and for the many substitutions that likely need to occur in order to evolve AROUND barriers in the fitness terrain. Evolution is not a linear straight line from start to finish. The issue is not the NET difference; rather, the issue is whether 1,667 mutations are sufficient to CREATE all the uniquely human adaptations. The business of creating is tougher than “net difference.”
  3. The 1,667 substitutions are not all required to be “single nucleotides” – though that is close enough perhaps for a simplified teaching of the problem – though evolutionists will object, so we must find a suitable middle ground to teach. Rather, according to evolutionary geneticists, almost all of the substitutions will be single nucleotides. That point is needed to fend off one of the classic mistakes about Haldane’s Dilemma, namely, that each “substitution” is an entirely new gene (thousands of new nucleotides!). See here.
  • The essay removed a key point: “Some creationists view Haldane's Dilemma as compelling evidence that evolutionary genetics cannot cope with its most central issues.” Indeed, that is the more likely outcome in the near term, because evolutionists will sooner trash-out evolutionary genetics as a field (and embrace every unfalsifiable, untestable avenue of escape) rather than allow Haldane’s Dilemma to falsify evolution outright.
  • The essay claims evolutionists “typically respond that the 1667 limit is not problematic.” However, evolutionary geneticists claim no such thing: not universally; not even typically; and especially not to the general public. Indeed, they have yet to address the issue in any direct way for the general public. The bald claim that the 1,667 limit “is not a problem” is made instead by Internet evolutionists, almost always anonymously, on transitory, untraceable, shouting matches called Internet “discussion groups.” Haldane's Dilemma cannot be considered solved until the solution exists in ink, by capable scientists willing to put their names and reputations to it. Evolutionary operatives posting their opinions anonymously at unreadable, internet shouting-matches, do not count as solutions. See here

Given this list of difficulties with the current essay, I trust I may be forgiven for reverting back to something more like the previous version. Tell me where I have not kept things sufficiently simple.

The essay may well need sections on responses to evolutionist arguments. I can go into these matters as deeply as you like, though I suspect, the goal here is give reasonably short, illuminating, accurate, summaries, with links to further information. We’ll see what people here want. Wremine 02:45, 12 June 2006 (CDT)Wremine


For what it's worth, I like Walter's article. I'm certainly not an expert, but I now find the issue quite fascinating. I don't agree that it should be rewritten by someone else. I think someone who has done so much research into it and understands the subtleties is the ideal person to write it. -- Tim --Klang 05:12, 12 June 2006 (CDT)
Good work, Mr. Remine -- thanks for working with me -- you're definitely the expert here and i'll defer to your greater knowledge on the topic. As we move forward, I would really like it if we found a way to:
  • Explain what the cost is -- why is the cost 30? That's what I tried to add in the last one, with the illustration about cost as a result of selecting for multiple genes;
  • Address the assumptions in the model -- as I read it, I'm struck by all the assumptions he makes to come to this number -- assumptions about unobservable conditions. For instance, as I understand it, he assumes a stable population. Where in nature do we see stable populations? The human population has never been stable -- 1/3 of Europe died during the black plague -- we had 1.6 billion people at the beginning of the 20th century, and 6.1 billion at the end. How can we assume a stable population? And doesn't an unstable population render his calculation pretty meaningless? Further, he assumes an average generation of 20 -- why? chimps have been known to have babies at 7.5 -- humans at 13-14 -- why a reproductive age of 20? even worse, how can he assume a certain reproductive rate? Only 100 years ago, humans in the West were having 10-12 kids per generation. humans in Africa are still having that many. What reproductive rate did he assume, and why?
  • Explain evolutionary counterarguments (on their own terms), and then give full refutations of them;
  • Try to maintain something of an npov tone in the intro, etc.
'll take a stab at these ... feel free to edit them as you see fit ... i'm not here to prove this true or false -- just to write a good article:). that's the joy of a wiki, right:)? Ungtss 08:08, 12 June 2006 (CDT)

Thanks for the encouragement. Yes, I think Haldane’s Dilemma is an utterly fascinating topic (from several angles), no matter which side of the fence you are on. I’m delighted you are interested too. However, you are now asking for self-contradictory things. That is, you want the essay to explain ‘this, that, and the other thing,’ and to explain these simply (so ordinary folks can understand) – yet you also want it to be a Wiki essay, not a book! Well, it would take a book (or many, many webpages) to explain everything in the simplicity that you want. We have to find the right balance between brevity and completeness, (with emphasis here on brevity). And there are many other sub-topics about Haldane’s Dilemma that are more important for this essay. Keep the ultimate goal in perspective here. Now to your questions:

  • Why is the total cost of substitution 30? I explained that in the essay. If you sum the cost each generation, over the entire substitution, then on average, the total is 30. The more difficult question is why that holds also during multiple concurrent substitutions. The answer is complicated (likely way too complicated for this essay). But I can come up with a simplified example, if readers wanted it. Though I strongly suspect readers want other things handled more.
  • Haldane assumed a constant population size, because:
  1. It is a reasonable assumption for many (most?) species, most of the time. And it is quite suitable for achieving general-purpose results.
  2. There are an infinite number of ways for the population size to fluctuate. And that merely complicates the problem, without solving it! For example, Haldane’s Dilemma says the substitution rate is too slow. Is the problem ‘solved’ by noting that “1/3 of Europe died during the plague”? I’m sorry, that only makes the problem worse. (Indeed, most of the so-called “solutions” to Haldane’s Dilemma actually make the problem worse.)
  3. The fact that human populations quadrupled in the twentieth century is not typical of history. Indeed, it can be proven that such a long-term trend is impossible. In the twentieth century, the high growth was due to technology, not biology, and that does not apply to human pre-history. According to evolutionists, human population sizes were reasonably stable during the period of human evolution. (Note: Why do they claim that? Answer: Lack of pre-human ancestors during the period in question – so evolutionists claim “there must have been consistently small population sizes,” and it is awkward to claim “consistently small population sizes that are unstable.”)
  4. Most importantly, Haldane assumed a constant population size, because his particular cost concept required it, a limitation that has been removed by ReMine’s paper.
  • The key parameter is not the age of reproductive maturity (13 years for humans). Rather, it is the effective generation time, which, according to evolutionary geneticists was at least 20 years during the period in question. (To document this point, various evolutionary geneticists are cited in my book. Again, ALL the key data, models, theory, derivations, and calculations, come from evolutionists.
  • Explain evolutionary counter-arguments on their terms, and give full refutations? Again, you’re asking for a self-contradiction. You’re asking for a book, when you know a book is unsuitably long here. You’ll need to decide what you really want for this essay. You’ll need to decide how much brevity you want. Some answers are more important here than others. Some can be handled implicitly (to save space), some explicitly, and some answers can be referred elsewhere (with links). And some will simply have to be briefly summarized.

Ungtss, you are obviously immersed in the standard evolutionist attempts to obfuscate Haldane’s Dilemma. Indeed, I currently think of you, in effect, as a closet-evolutionist. That’s fine. Though you need to get a grip on your self-contradictory demands for this essay. Keeping the required brevity in mind, think of the most important things this essay must provide.

For example, do we need to specifically itemize and belabor the simple arithmetic (1,667 = 10 million / 20 / 300), as in the current essay? Is that really necessary? It's not a point that anyone disputes. Isn't that needlessly bludgeoning a point that is undisputed anyway? The essay already makes the correct, blanket, covering statement: "It is undisputed that Haldane’s calculations, if correct, would indicate such a limit." That is the essential point, without itemizing every last detail. Isn't this simple, undisputed, arithmetic better handled implicitly, with a link to such tedious details?

I recommend keeping the essay focused on the major issues, not needless tedium that no one disputes.

Wremine 14:20, 12 June 2006 (CDT)Wremine


Thanks for the encouragement.
Gracious -- we're very glad to have you around:).
We have to find the right balance between brevity and completeness,
That's a good point and we should keep that in mind. I'd like to be a little less "brief" and a little more "complete" than we are now, tho, if that's okay with you:).
Why is the total cost of substitution 30? I explained that in the essay. If you sum the cost each generation, over the entire substitution, then on average, the total is 30.
I guess what I'm after is an understanding of what causes each of those costs, how they were calculated, and what assumptions were used to make the calculations.
The more difficult question is why that holds also during multiple concurrent substitutions. The answer is complicated (likely way too complicated for this essay). But I can come up with a simplified example, if readers wanted it. Though I strongly suspect readers want other things handled more.
This particular reader wants it a great deal:).
There are an infinite number of ways for the population size to fluctuate. And that merely complicates the problem, without solving it! For example, Haldane’s Dilemma says the substitution rate is too slow. Is the problem ‘solved’ by noting that “1/3 of Europe died during the plague”? I’m sorry, that only makes the problem worse. (Indeed, most of the so-called “solutions” to Haldane’s Dilemma actually make the problem worse.)
Bingo -- that's what I was thinking, too. I suggest we emphasize less the "precise calculation" (which is so steeped in assumptions that it's not all that meaningful in and of itself) and instead focus on the limits -- the fact that many of the assumptions underlying that number actually make it artificially large, and it is probably even smaller.
The fact that human populations quadrupled in the twentieth century is not typical of history. Indeed, it can be proven that such a long-term trend is impossible. In the twentieth century, the high growth was due to technology, not biology, and that does not apply to human pre-history. According to evolutionists, human population sizes were reasonably stable during the period of human evolution. (Note: Why do they claim that? Answer: Lack of pre-human ancestors during the period in question – so evolutionists claim “there must have been consistently small population sizes,” and it is awkward to claim “consistently small population sizes that are unstable.”)
This goes along with my response to your last comment -- evolutionists assume stable population, but it's an assumption not grounded in evidence or experience. In reality, populations undergo rapid reductins and expansions -- from asteroids to ice ages to dessication -- but those facts help the case against evolution, and we might do well to draw them out.
The key parameter is not the age of reproductive maturity (13 years for humans). Rather, it is the effective generation time, which, according to evolutionary geneticists was at least 20 years during the period in question. (To document this point, various evolutionary geneticists are cited in my book. Again, ALL the key data, models, theory, derivations, and calculations, come from evolutionists.
Indeed ... it is certainly useful to use their own words against them -- I just think we'd do well to point out that the final calculation is based on those assumptions, and so may very well not be accurate in and of itself ... but it does provide us with a general picture of the maximum substitution rate, because the average generation couldn't be too much less than 20.
Ungtss, you are obviously immersed in the standard evolutionist attempts to obfuscate Haldane’s Dilemma. Indeed, I currently think of you, in effect, as a closet-evolutionist. That’s fine. Though you need to get a grip on your self-contradictory demands for this essay. Keeping the required brevity in mind, think of the most important things this essay must provide.
Sir, we're on the same side:). My hopes (not demands) for the essay are not self-contradictory -- I am in favor of providing a brief summary up front, and then a detailed description and analysis to follow. I'm also not a closet evolutionist by any stretch of the imagination, in effect or otherwise -- but I do question the merits of claims made by both sides. Otherwise I'd lose my mind:).
For example, do we need to specifically itemize and belabor the simple arithmetic (1,667 = 10 million / 20 / 300), as in the current essay? Is that really necessary? It's not a point that anyone disputes.
It may not be a point of dispute, but it's a point of clarity. As an ordinary reader (and I'm certainly ordinary), it's very helpful for me to know where all these numbers are coming from. I don't trust Haldane any more than Darwin or Pee Wee Herman -- I'm unwilling to accept ideas on the authority of people, particularly when people disagree. I can only accept an argument when the merits of the argument have been laid out for me. Ungtss 14:48, 12 June 2006 (CDT)

Big Questions:

  • How many simultaneous traits is he modeling for his 1667 number? One? Ten?
  • In the scenario, are we saying that it takes 300 generations for a single mutation to go from single individual to homozygosity in the population? If so, wouldn't our actual number be lower, given the fact that beneficial mutations don't come around very often? Ungtss 15:23, 12 June 2006 (CDT)
 Ungtss 15:11, 12 June 2006 (CDT)

“As a result, Haldane's calculations indicate that there can be a maximum of one beneficial nucleide change in 300 generations.”

That over-states the case. It’s a “typical,” not an absolute “maximum”. [Note of clarification, added later, to eliminate ambiguity about that statement. The time between two specific substitutions may be short or long, and is not the issue. Rather, the issue is the average over the long term. Like cars on a highway, we are interested in the average rate at which they cross the county line, not the interval between two specific crossings (which might be short or long). This average long-term rate, is also a limit, as explained in the article, because it uses wildly unrealistic assumptions in favor of evolution.] Also, the point was already covered sufficiently in the essay, and needs no further emphasis – indeed the over-emphasis of trivial points should be avoided. For sake of brevity, we cannot be adding clarifiers needlessly for every little thing. Stick to main points of interest. Even without that clarifier, the essay is going to get terribly long.

Think of it this way: For everything you add, something else must be left out. Develop a perspective on what should be in, and what should be out. The more important points to put in are the points that evolutionists misrepresent, confuse, attack, or avoid – in other words, live issues. Points that have no serious objections (such as the arithmetic example) are, in effect, a diversion from the real issues.

“… because the cost would have exceeded the reproduction rate, causing the extinction of the line.”

That wording is entirely unnecessary, confused, and ultimately mistaken. The cost of substitution does not, by itself, cause extinction of the species. Rather, it causes extinction of the given evolutionary scenario. No reference to extinction of the species is necessary for a cost argument. Rather, a certain reproduction rate is required, and if the species cannot actually deliver that reproduction rate, then the scenario is implausible. PERIOD! END OF ARGUMENT! This finality comes from the word “required.” For example, if a tank of gas is required for driving to Omaha, and you don’t have a tank of gas, then you’re not driving to Omaha – not going extinct.

The logic of the cost argument is undeniable, and is not helped by introducing needless confusion about “species extinction.” Indeed, this is one of the classic confusions that ReMine’s paper eliminated. Evolutionists argued this way: If a species receives beneficial mutations too rapidly, then the cost of substitution will be too high, and the species will go extinct. Such nonsense was promoted as a way to confuse, and ultimately denounce cost arguments.

Big Question: how many simultaneous traits is he modeling for his 1667 number? One? Ten?

Haldane’s calculations allowed for hundreds even thousands of simultaneous, concurrent substitutions. That is, he assumed each INDIVIDUAL substitution is exceedingly slow (infinitely slow), which gives the ABSOLUTE LOWEST total cost of substitution in each case. Then he allowed many of these concurrently, to get the OPTIMALLY FASTEST substitution rate overall, at the lowest possible cost.

Here’s an analogy. You want to truck products across country at the fastest RATE, but you have limited gas. One approach is to drive a truck fast across country. But that inefficiently uses the limited gas. The better approach would be to use thousands of trucks simultaneously, with each one driving super slow. That will optimize the overall RATE of delivery. In effect, that is what Haldane did. Haldane calculated an optimally fast delivery rate.

… wouldn't our actual number be lower, given the fact that beneficial mutations don't come around very often?

There are two separate limitations to the beneficial substitution rate. It is mutation-rate limited (due to the rarity of beneficial mutations). A good example of this argument was given by evolutionary geneticist, Motoo Kimura, and is covered in my book. His argument tends to be more effective for smaller populations. The other limit is a cost-limit (such as Haldane’s argument), and tends to be more effective in larger populations. Both arguments apply – a mutation-rate limit, and a cost-limit – whichever is slower.

Wremine 16:18, 12 June 2006 (CDT)Wremine


“As a result, Haldane's calculations indicate that there can be a maximum of one beneficial nucleide change in 300 generations.”
That over-states the case. It’s a “typical,” not an absolute “maximum”. Also, the point was already covered sufficiently in the essay, and needs no further emphasis – indeed the over-emphasis of trivial points should be avoided.
I have read the article several times, and obviously still do not understand the heart of the argument. It seems clear to me, then, that the point has not been covered adequately in the essay. If it had been, I would understand it by now:). If what you say above is what you actually mean, then why does the article call this a "limit," and why do you write, "Haldane calculated that organisms with low reproduction rates, such as cows, could substitute new beneficial mutations no more often than one per 300 generations."?
For sake of brevity, we cannot be adding clarifiers needlessly for every little thing. Stick to main points of interest.
I am sticking to the main points of interest:). The merits of the argument:).
Think of it this way: For everything you add, something else must be left out.
That assumes it's impossible to expand the article:). If we expand the article, we can add things without taking anything out:).
Develop a perspective on what should be in, and what should be out. The more important points to put in are the points that evolutionists misrepresent, confuse, attack, or avoid – in other words, live issues.
That is our key point of disagreement. I don't think those are the key points. I think the key issue is whether there is validity in the general theory. Your main points are simply ad hominem arguments, and are totally irrelevant to what I consider to be the "real issue." I think the real issue is, "Does this argument have merit?" And I think the key facts to be considered related to "What this argument actually says."
“… because the cost would have exceeded the reproduction rate, causing the extinction of the line.”
That wording is entirely unnecessary, confused, and ultimately mistaken. The cost of substitution does not, by itself, cause extinction of the species. Rather, it causes extinction of the given evolutionary scenario. No reference to extinction of the species is necessary for a cost argument. Rather, a certain reproduction rate is required, and if the species cannot actually deliver that reproduction rate, then the scenario is implausible. PERIOD! END OF ARGUMENT! This finality comes from the word “required.” For example, if a tank of gas is required for driving to Omaha, and you don’t have a tank of gas, then you’re not driving to Omaha – not going extinct.
Then, good friend, I will need you to explain why a certain reproduction rate is required for a scenario to be plausible. Your conclusion is made without stating the facts that justify it. Help a brutha out:).
Here’s an analogy. You want to truck products across country at the fastest RATE, but you have limited gas. One approach is to drive a truck fast across country. But that inefficiently uses the limited gas. The better approach would be to use thousands of trucks simultaneously, with each one driving super slow. That will optimize the overall RATE of delivery. In effect, that is what Haldane did. Haldane calculated an optimally fast delivery rate.
Understood. So is he saying that this a change/300 generations is the optimal rate, the historical rate, the maximum rate, or a typical rate?
There are two separate limitations to the beneficial substitution rate. It is mutation-rate limited (due to the rarity of beneficial mutations). A good example of this argument was given by evolutionary geneticist, Motoo Kimura, and is covered in my book. His argument tends to be more effective for smaller populations. The other limit is a cost-limit (such as Haldane’s argument), and tends to be more effective in larger populations. Both arguments apply – a mutation-rate limit, and a cost-limit – whichever is slower.
Understood. I think the article could benefit from clarification on that point. Ungtss 16:37, 12 June 2006 (CDT)

I read the original Haldane article, and I see no indication of any limit stated or implied in his paper. He seems to recognize the limitations of his own assumptions, and state 300 as a mean -- not a limit -- and the real rate could change drastically, based on environmental conditions. Just as my intuition had led me to believe To be brutally honest, I'm not impressed at all. The article states this "300" as a limit explicitly, but that 300 is a meaningless, arbitrary number based on extremely tenuous assumptions -- and Haldane himself recognized it. While this provides a very useful method for evaluating questions of this kind, i think it's absurd to claim that this "300" or "1667" is some sort of magical limit. There's no basis for it, my friend. Ungtss 20:47, 12 June 2006 (CDT)


Haldane’s argument uses average values (mean values), of course it does. And that makes it even more compelling, because over the long run there is no getting around the average values. Under Haldane’s argument, there is an average cost (30), and an average payment (0.1), and over the long run there is no getting around the limit they establish (one per 300 generation). Yes, it is a limit, and a rather strong one. Yes, one particular substitution might be faster, and another slower, but over the long run there is no getting around the limit. You cannot claim that over the long run the rate is one substitution per 250 generations – the averages forbid any such claim. The long-term average is a limit, and a rather powerful one.

No, it is not an “average rate” of substitution, because nothing in Haldane’s argument assures that the beneficial mutations will occur, or that they won’t be eliminated by drift (as they would be the vast majority of the time). And nothing in Haldane’s argument assures that the species isn’t stuck on top of a local fitness peak, pinned there by survival of the fittest, and prevented from evolving. Nothing in Haldane’s argument assures there won’t be obstacles in the fitness terrain that prevent evolution, or turn evolution into a convoluted maze, with dead ends, blind alleys, and backtracking. Nothing in Haldane’s argument assures there won’t be extinction. Rather, Haldane assumed-away any and all such obstacles. In short, Haldane’s argument is not sufficient to assure any evolution, but it is sufficient to prevent evolution faster than the Haldane rate. This asymmetry is why it is called a “Limit,” not an average rate.

and the real rate could change drastically, based on environmental conditions.

Yes, the real rate could change drastically – slower. There is no cost problem whatever with a slower substitution rate.

You are correct that Haldane made some “extremely tenuous assumptions” – he even made some highly unrealistic assumptions – and all of them are IN FAVOR of evolution. It is not enough to brush aside Haldane’s assumptions, because that only makes Haldane’s Dilemma worse.

For example, when the environment changes (as you suggest), then the environmental change (like mutational change) is random with respect to the species, and therefore (like mutational change) is generally harmful to the species. In this way, environmental change tends to reduce the species reproduction rate, and increase other costs, such as the cost of environmental deterioration, and the cost of random loss. And all of that makes Haldane’s Dilemma worse, not better. Haldane completely ignored all the ill effects of environmental change, and incorporated highly unrealistic environmental effects – and all of it unrealistically favors evolution.

It’s unfortunate that you’re “not impressed at all” by Haldane’s argument. Because leading evolutionary geneticists were definitely impressed. Motoo Kimura cited Haldane’s Dilemma as his MAIN reason for promoting his theory of neutral evolution. John Maynard-Smith likewise cited Haldane’s Dilemma as his main reason for promoting his revolutionary new view of evolutionary process. Likewise with Sved. Likewise with Wallace’s notion of “soft selection.” Likewise with Warren Ewens advancing his unique view of evolutionary process. These evolutionary geneticists saw Haldane’s argument as deeply impressive enough to require a radical new response. No minor, small, nitpicking would do. Rather, they proposed: (1) some radically new type of evolutionary process, or (2) the identification of some serious deep mistake within Haldane’s reasoning. Whether their radical “solutions” are plausible, or not, is the issue. But one cannot simply brush Haldane’s assumptions, his reasoning, or his conclusions – because that doesn’t solve Haldane’s Dilemma.

I read the original Haldane article, and I see no indication of any limit stated or implied in his paper. …. The article states this "300" as a limit explicitly,

You’re contradicting yourself.

There is nothing “meaningless” or “arbitrary” about Haldane’s derivations, calculations, or conclusion. Your charges are unspecific and baseless.

Wremine 01:33, 13 June 2006 (CDT)Wremine


And that makes it even more compelling, because over the long run there is no getting around the average values.
Oh, but there is:). his average values are based on assumptions that are unverifiable in some cases and untrue in others:(.
The long-term average is a limit, and a rather powerful one.
In short, Haldane’s argument is not sufficient to assure any evolution, but it is sufficient to prevent evolution faster than the Haldane rate. This asymmetry is why it is called a “Limit,” not an average rate.
Only if his assumptions are true. In a scenario in which population is not constant, his limit would be shattered:(. You addressed population decrease, but you avoided population increase. How would his limit apply in a scenario where humans regularly had 8-10 children and population is rapidly increasing, as they did in the days of Isaac? Simple: It would be broken:(.
It’s unfortunate that you’re “not impressed at all” by Haldane’s argument. Because leading evolutionary geneticists were definitely impressed.
I'm afraid that doesn't mean much to me. Those geneticists were also impressed by evolution. They accepted one tenuous idea ... why not another:(?
The problem with this argument, as I see it, is that because it makes reference only to evolutionary assumptions, it will act (in their minds) only as a refinement and correction of their assumptions. I'm sure you observed that with the evolutionists you've discussed this with. This doesn't disprove evolution -- they simply come up with more speculations to get around it. In order to falsify evolution (which is my primary interest), an argument must be based on observable, falsifiable facts, so that nothing can be challenged except the theory itself. And this argument doesn't provide that.
I read the original Haldane article, and I see no indication of any limit stated or implied in his paper. …. The article states this "300" as a limit explicitly,
You’re contradicting yourself.
I apologize for my ambiguous language. This article states "300" as a limit. That article does not.
In any event, thank you for the article -- it contributes a lot to the wiki here -- I do hope you come back around and add more:). Ungtss 07:41, 13 June 2006 (CDT)

The essay doesn’t claim to “disprove evolution.” If falsifying evolution is your only concern, then you should broaden your concerns. There are other ways of invalidating evolution as a science, (such as by showing evolution is unfalsifiable – and that too is in play here, in Haldane’s Dilemma).

You have it backwards. Those geneticists did NOT take Haldane’s Dilemma as a “tenuous” idea, quite the contrary. It is you, not those geneticists, who is brushing aside Haldane’s idea as tenuous. The fact that “it makes reference only to evolutionary assumptions” in FAVOR of evolution, is what makes Haldane’s Dilemma an intriguing anti-evolutionary argument.

his average values are based on assumptions that are unverifiable in some cases and untrue in others:(.

Your argument is invalid. Haldane’s assumptions are wildly unrealistic IN FAVOR of evolution. When you claim the assumptions are “unverifiable or untrue” you have not changed that fact in the slightest, and you have not s-o-l-v-e-d Haldane’s Dilemma.

In a scenario in which population is not constant, his limit would be shattered:(.

That is untrue. Changing the population size does not “shatter” the cost of substitution. There is always a cost of substitution. And you have not shown how the cost of substitution can even be reduced. And you have not s-o-l-v-e-d Haldane’s Dilemma.

How would his limit apply in a scenario where humans regularly had 8-10 children and population is rapidly increasing,

You just gave a different scenario. But your scenario has a HIGHER cost than before. Why? Because it REQUIRES more reproductive excess. And the cost is a LOT higher. Why? Because your scenario claims a “RAPID INCREASE,” and that requires a LOT more reproductive excess. The more rapidly you claim the population increases, the more rapidly you increase the cost. You’ve made the problem worse than before.

Suppose you want to drive a truck from A to B. You can drive with intermittent speed-ups and slow-downs, or, with a steady constant speed. Which one uses less gas? The steady constant speed, because it makes more efficient use of the limited gas. Well, in the cost of substitution, the ‘thing’ in limited supply is not gas, but excess reproduction rate. And it is used most efficiently if the same amount is used each generation – in other words, constant cost all the time. That proof is given in my published paper. The same proof shows that changes in population size do not reduce the cost of substitution, rather the lowest possible total cost of substitution is determined by the number of copies of the new substituting mutation at the beginning and end of the substitution.

Also, you have not accounted for the other costs of evolution (the cost of mutation, the cost of random loss, the cost of segregation, and so forth). You seem to assume the entire species reproduction rate goes to paying the cost of substitution, which is false.

If you don’t see the “300” limit in Haldane’s article, then perhaps you don’t know enough evolutionary genetics. To evolutionary geneticists (and most anyone else), it’s an undisputed fact that it’s in the article.

Wremine 01:22, 14 June 2006 (CDT)Wremine


Better, but still some problems. The opening sentence suggests a hard limit, whereas Haldane explicitly says the number 300 is an estimate (and suggests a value of 43 in specific cicumstances detailed in the paper). The comment that Haldane assumed a species didn't go extinct is probably unnecessary, since gene substitution doesn't take place in extinct species. Although the section on nucleotide numbers correctly states that many beneficial mutations involve a change of multiple nucleotides, it doesn't mention that there will also be neutral and mildly detrimental mutations which also contribute to the total number of nucleotides changed. There's no mention of the effect of sexual vs asexual reproduction on the fixation rates. And maybe there should be more emphasis on the dilemma being dependent on whether a few thousand beneficial mutations is enough for human/chimp divergence. Since Haldane's paper is online at http://www.blackwellpublishing.com/ridley/classictexts/haldane2.pdf, why not link to it? Oh, and Walter, desist from characterising me as anonymous. I am no more anonymous than you are. Roy 07:36, 14 June 2006 (CDT)


The essay doesn’t claim to “disprove evolution.”

The article states that "Creationists and Intelligent Design theorists argue that the limit is sufficiently slow as to call into question the “fact” evolution." Now if by "call into question" you mean "give meaningful reason to believe it is false," I think your argument fails. If, on the other hand, you mean, "Cause one to question today's version of the theory, then I think your argument succeeds, but to no purpose. evolutionists are always calling into question today's version of the theory -- that's how they get themselves famous. Nevertheless, they remain true to the theory itself. As long as we were pointing out mere "areas for refinement," we're not doing anything meaningful.

You have it backwards. Those geneticists did NOT take Haldane’s Dilemma as a “tenuous” idea, quite the contrary. It is you, not those geneticists, who is brushing aside Haldane’s idea as tenuous.

Again, I'm sorry I spoke in such vague terms. Let me make it clearer: These geneticists have accepted evolution (one tenuous idea), as fact. They therefore have little to no credibility with me on issues of origins science (although I'm sure they're quite good when it comes to operational science). Therefore, since they have no credibility with me in that area, the fact that they believe haldane's dilemma is significant, or that the Earth is 4 Billion years old, or that they can fly like superman, means nothing to me. They have no credibility in the area of origins science, so their response to Haldane's dilemma is irrelevent to me.

The fact that “it makes reference only to evolutionary assumptions” in FAVOR of evolution, is what makes Haldane’s Dilemma an intriguing anti-evolutionary argument.

Not to me, I'm afraid. Evolutionary assumptions are so malleable that they can come up with a "new and improved theory" in a second. The more we can replace "evolutionary assumptions" with "facts," the more effectively we are attacking the general theory. Since Haldane's dilemma is virtually all evolutionary assumptions, all it means is that some of their assumptions must be adjusted -- it doesn't nothing to challenge the theory itself:(.
Haldane’s assumptions are wildly unrealistic IN FAVOR of evolution. When you claim the assumptions are “unverifiable or untrue” you have not changed that fact in the slightest, and you have not s-o-l-v-e-d Haldane’s Dilemma.
I'm not trying to solve the dilemma -- I think it's an interesting unsolved issue -- but I think that it is so steeped in those assumptions (whether in favor of evolution or against it) that all one has to do is adjust the assumptions and they can still proclaim that evolution is true.
How would his limit apply in a scenario where humans regularly had 8-10 children and population is rapidly increasing,

You just gave a different scenario. But your scenario has a HIGHER cost than before. Why? Because it REQUIRES more reproductive excess. And the cost is a LOT higher. Why? Because your scenario claims a “RAPID INCREASE,” and that requires a LOT more reproductive excess. The more rapidly you claim the population increases, the more rapidly you increase the cost. You’ve made the problem worse than before.

I'd appreciate it if you could explain that to me further. Haldane is assuming a constant population, and determining what "cost" the population could "afford" in a constant population. But if the population is increasing because much higher reproductive rates, then can't the population "afford" a much higher cost?

If you don’t see the “300” limit in Haldane’s article, then perhaps you don’t know enough evolutionary genetics. To evolutionary geneticists (and most anyone else), it’s an undisputed fact that it’s in the article.

Well in this discussion, Roy (above) and I are disputing that "fact." I see the article as stating it as an estimate and an average, while you are stating it as a "hard limit," which, I'm afraid, is not what he meant. I may not be an expert at evolutionary genetics (I'm not much for fiction), but I am fairly adept at reading comprehension. Ungtss 08:14, 14 June 2006 (CDT)

Analysis from Ungtss

The failure of this argument, in my mind, lies in its failure to grasp the nature of population equilibrium.

Here are my premises:

  • Population stability is simply an artifact of a population's death rate equalling its birth rate. If birth rate exceeds death rate, equilibrium population will increase. If death rate exceeds birth rate, equilibrium population will decrease.
  • Beneficial mutations will by definition either increase the birth rate or decrease the death rate. That's what makes them beneficial.
  • Thus, beneficial mutations will by definition alter the equilibrium population. If mutation lowers a population's death rate or increases its birth rate, then the equilibrium population will increase.
  • Thus, the assumption of a stable population is a fundamental flaw in the model -- by definition, beneficial mutations will increase the equilibrium population, while disadvantageous mutations will decrease the equilibrium population. When you assume constant population, you assume something that is not only never true, but which adjusts to account for the mutations a population is undergoing.

Here are some scenarios to illustrate:

Scenario 1:

Population of 100 individuals with old trait A.

1 gets advantageous mutation B.

Old population A was stable, because reproduction rate was in equilibrium with death rate;

New individual B has children

Population B has a higher equilibrium population than population A, because its new advantage causes its death rate to be lower than population A's, and/or its birth rate to be higher than population A's.

Consequently, Population B grows more quickly than A, because it's reproduction rate exceeds its death rate.

Because of the increase in population B, Total Population (A+B) increases above the equilibrium of population A, toward the equilibrium of population B.

As population B continues to grow at a faster rate than population A, population B will increase relative to population A. As a result of population B's lower death rate/higher birth rate, the total population will continually move to a higher equilibrium population.

Where's the cost?

Scenario 2:

Population of 100 individuals with old trait A.

1 gets advantageous mutation B, and has a few children, creating a small subpopulation.

Rapid environmental change kills off all of population A, leaving only a small population B.

Because of the reduced population, B's death rate is decreased; B's reproduction rate now drastically exceeds its death rate, and it begins to repopulate.

Where's the cost?

Analysis:

I think the above scenarios illustrate the problem with this "cost analysis:" its assumption of a stable population. Populations are only stable if the reproductive rate equals the death rate. If a beneficial mutation reduces a subpopulation's death rate (or increases its birth rate), that subpopulation will move out of equilibrium, and the population will increase. Even more severely, if an environmental change wipes out the pre-mutation population, the decreased population will decrease the post-mutational death rate, so that the post-mutational population will increase (potentially very rapidly).

In the end, this "cost analysis" is meaningless, because the realities of population dynamics belie any claims to a stable population.

Ungtss 10:18, 14 June 2006 (CDT)


This section is from Ungtss, responding (in interleaved fashion) to a post by Wremine (which is now gone, due to the interleaving).

Your scenario slows down evolution, and in ways you have not remotely accounted:

You have evaded addressing the heart of my criticism, in favor of arguing irrelevancies. The point of my argument is this: population is not constant. advantageous mutations increase reproduction rate and equilibrium population, paying the "cost of substitution." therefore this 1667 of yours, based on an assumed stable population, means absolutely nothing.
Let me count the irrelevencies:
  • Your scenario assumes a single substitution – which slows evolution down.
Add however many substitutions you like. Beneficial mutations will still increase the reproduction rate and equilibrium population, paying the cost of substitution.
  • Your scenario assumes the new beneficial mutation has a selection coefficient of nearly 1, which is implausible. Moreover, high selection coefficients increase the cost of substitution. Haldane got the lowest conceivable cost of substitution by assuming tiny selection coefficients, (approaching 0+).
Substitute whatever selection coefficient you like -- Beneficial mutations will still increase the reproduction rate and equilibrium population, paying the cost of substitution.
  • The dramatic reduction of population size also dramatically reduces the rate at which the population receives beneficial mutations – and this dramatically slows evolution.
But it dramatically increases the rate at which old mutations spread throughout the population.
  • The severe inbreeding rapidly fixates numerous harmful mutations, which takes evolution in the wrong direction.
You are engaging in straw man. I'm not talking about a population bottleneck of such an extreme degree -- I'm talking about an environmental change that wipes out the pre-mutation population. If you like, the example can have 1/3 post-mutation population, to avoid your silly population bottleneck side-issue.
  • The drastic population size reduction (down to a few individuals) is likely to cause extinction – especially if this is claimed to occur repeatedly – and that makes your long-term scenario implausible.
Again, this is mere straw man. I'm not talking about extreme population bottleneck. I'm talking about a population decrease that wipes out the old and gives the new room to grow and expand.
  • The cost is still there, and especially incurred when the new population is growing back to its previous size. Pay now, or pay later, but you cannot get around the cost of substitution. You have not solved, nor even reduced, Haldane’s Dilemma.
As you so often do, you have stated a conclusion without justifying it with facts. I am attempting to explain to you that your need for increased reproduction to spread the "new-type" is paid for by the increased growth rate caused by the new-type's survival advantage. Yes, the new-type must have a higher reproduction rate to pay the cost of substitution. And that higher reproduction rate is paid for my its increased birth rate and decreased death rate, as a result of its new genetic advantages. If you've addressed this issue somewhere, please let me know. Otherwise, stop simply dismissing what I have to say with premature cries of victory over a person who is not your opponent.
As I said, you are confusing cost and payment. The scenario you gave ABSOLUTELY REQUIRES higher reproduction rates, therefore it has a higher cost. You falsely assumed – by virtue of simply telling your scenario – that the species can actually supply that reproduction rate. The payment is what the species actually produces – it is observable. To see whether the payment is sufficient, you must make an argument ultimately based upon OBSERVED reproduction rate (not imaginings in your head).
As I have said, the reproduction rate of a subpopulation with a survival advantage will increase by definition, thus paying this "cost of substitution" of yours.
I already explained why the Haldane rate is not an “average rate,” because Haldane’s argument does not assure – and made no attempt to assure – that evolution will happen at his rate. It is a limit, not a guaranteed “average rate.”
It appears to me that you are contradicting yourself. You say above that "over the long run there is no getting around the average values." Now you say it not a "guaranteed 'average rate'" Now you tell me, how do those two statements fit together? An average is the sum divided by the period. If there is no getting around average values in the long run, then why is it not a "guaranteed average rate?"
If, on the other hand, you mean, "Cause one to question today's version of the theory, then I think your argument succeeds, but to no purpose.
You are contradicting yourself.
Perhaps we should review the definition of "contradiction." A contradiction is making two incompatible statements. I have done so such thing. I have said that if your goal is to question today's version of the theory, you've succeeded, but your victory is hollow, because it will only inspire them to come up with a "new and improved version" of the same tripe.
[Concerning their knowledge of genetics, which is their field of expertise] These geneticists … have little to no credibility with me.
You are now impugning your own credibility. Your attempt to brush aside evolutionary geneticists wholesale is wrong.
Perhaps we should review the dangers of accepting ideas purely on the basis of a person's credibility. I have said that I do not accept the opinions of evolutionary geneticists on their credibility, but only on the facts they present. I have not "brushed aside evolutionary geneticists." I have demanded that they make meritorious arguments, rather than sit on high horses like self-proclaimed prophets and tell me what I ought to believe. If they begin to make meritorious arguments, I will accept their arguments on the merits. But the fact that you can name evolutionary biologists who "agree with you" is totally irrelevent, because at heart they don't agree with you -- they're still evolutionists.

Your assumption – that beneficial substitutions must increase the population size – is false. No population geneticist would endorse it. No population geneticist objects to evolution because of a constant population size. You misunderstand population genetics.

You are stating conclusions without justifying them, other than the fallacy of proof by authority. I don't care what population geneticists object to. I want facts, and you continually fail to provide them. If have argued with facts that beneficial mutations will increase reproductive rates and population equilibriums, paying off any "increased costs" of substitution, and rendering any model based on static population size totally meaningless. You have responded that "Nobody believes that." If all you have is proof by authority, proof by assertion, and ad hominem argument, please let me know now so I can terminate our discussion.

You have not yet explained why the increased population growth either does not occur as a result of survival advantage, or does not pay the increased cost of substitution.

Ugtss – You are inserting things into the essay that are mistaken, and for which there exist no publications to support you. If your statements do not come from a reliable source, then they don’t belong in the essay. I understand that you have concerns, but the essay is not your place for them.

Listen, Remine. You have hurled bizarre accusations (like my being some sort of in effect closet evolutionist), you have emphasized the importance of making ad hominem arguments against evolutionists, and you have rested the credibility of your argument against evolution on the opinion of evolutionists. Your performance in these areas has been far less than admirable. Further, you have deleted criticisms at the end of the article, without addressing them directly either in the article or on the talkpage. I haven't run into behavior like this since my old days fighting the evolution mafia on Wikipedia. Let me be perfectly clear. It is not appropriate to delete criticisms, no matter what you think of them. CreationWiki has no policy requiring that arguments be cited or published. If you want to play that game, try and get your arguments to last 3 minutes on wikipedia. The appropriate behavior is as follows: address criticisms in the body of the article. You have thusfar failed to do that. You have simply deleted a criticism because you thought it didn't have merit. Do not delete stuff simply because you disagree with it. Address it in the body of the article, so that readers can gain a deeper understanding of the topic. Ungtss 22:52, 14 June 2006 (CDT)

Comments by Roy

Re: "This ignores the more than ninety-nine percent of the time that Stephen Gould claims species (including pre-human species) are in stasis, where genetic change is not occurring."

This is false. Gould did not say that no genetic change is occurring. I doubt that anyone else would say that either.

Re: "# He assumed the species is not in error catastrophe (also known as mutational meltdown), where genetic deterioration occurs from generation to generation."

Assuming that something that has never been demonstrated to happen is not happening seems reasonable to me. This criticism is groundless.

Re: "In this way, Haldane eliminates the largest source of beneficial mutations (beneficial recessive mutations),"

This is an unsupported claim and makes no sense anyway.

Re: "# He assumed a sufficiently high rate of beneficial mutation."

Of course. When calculating the maximum rate at which beneficial mutations can become fixed, it's futile to use a rate of appearance of mutations so slow that the fixation rate is restricted by having insufficient beneficial mutations to work on. This assumption is necessary.

Re: "# He assumed the same model of genetics and selection still predominant today in evolutionary genetics textbooks – there was nothing special about Haldane’s model."

Naturally. There was no reason to use anything else. Had he used an obscure or specially invented model that differed radically from the predominant one, then you'd have grounds for complaint. But criticising Haldane for using the standard model is just silly.

Re: "# He assumed tiny selection coefficients (approaching 0+),..."

Absolutely false. Actual selection coefficients listed in Haldane's paper include 0.03, 0.69 and 0.1. The 300 generation estimate for fixation assumes a selection coefficient of 0.1, which is definitely not "approacting 0+".

Incidentally, the comment on Crow and Ewens opinion of ReMine's paper being correct omits one very important detail - that they also thought it contributed nothing new.

Roy 08:57, 15 June 2006 (CDT)

Walter: If you would be so kind as to provide cites or qualifications to address Roy's concerns ... for instance ... where does Haldane assume 0+ selection coefficients? on what basis do you argue that beneficial recessive mutations are the most common type? and what, exactly, is "mutational meltdown," and has it ever been observed? Ungtss 10:23, 15 June 2006 (CDT)
Pending Walter's reincarnation, I'll describe "mutational meltdown".
Error catastrophe, or mutational meltdown, is used to describe a situation in which mutation rates are sufficiently high that almost all offspring have detrimental mutations, with the result that each generation is less fit than the preceding one. Although this is easy to demonstrate in simulations, this is typically done by setting (i) an absurdly low population (i.e. less than 20), (ii) an absurdly high mutation rate, and (iii) ensuring that every member of the next generation has exactly one mutation, rather than allowing mutations to occur with an equivalent frequency (see e.g. http://www.answersingenesis.org/home/area/magazines/tj/docs/tjv16n2_weasel.asp ). If any of these conditions are relaxed, error catastrophe does not occur. Note that removing the third criteria - forcing every offspring to have exactly one mutation - will prevent error catastrophe even if the absurdly low populations and absurdly high mutation rates are retained, simply by allowing the mutations to be distributed randomly and hence allowing some of the next generation to have no mutations.
Since in nature mutation rates are low, populations are usually large and mutations are distributed randomly, rather than being parcelled out so that everyone gets one, error catastrophe/mutational meltdown will not occur.
Roy 05:32, 21 June 2006 (CDT)
Following the absence of a response from Walter, is anyone going to remove the falsehoods? Roy 10:33, 6 July 2006 (CDT)
Don't hold your breath. Remine has exclusive rights to the page. PrometheusX303 11:34, 6 July 2006 (CDT)

More changes, but few if any of the above problems have been fixed. For instance, 1 beneficial mutation substituted per 300 generations is still being treated as a hard limit rather than as an estimated average; the false claim that Haldane used negligible selection coefficients (0+) remains; the reference to nucleotide numbers of beneficial mutations only is still there. Instead, additional falsehoods have been added. Specifically:

Re: "That can be compared with the power of beneficial mutations observed today, such as the alleged examples in the beaks of Galapagos finches."

We've only been examining these finches for ~150 years. If Walter's treatment of Haldane's dilemma correct, there hasn't been enough time for any beneficial mutations to fix in these finches; which means Walter's characterisation of the changes in beak size as being due to beneficial mutations, rather than variation in frequency of existing alleles, must be incorrect.

Re: "Haldane’s Dilemma is easy to communicate – a limit of 1,667 beneficial mutations for human evolution."

That isn't Haldane's dilemma.

Re: "Even their technical journals strongly avoided such direct figures, and instead spoke obliquely in terms of substitution rate – such as “one gene substitution per 300 generations,” a phrase that obscures the problem from view, and may as well be in code."

Walter provides a link for his quote - but the link is not to a technical journal, but to his own web-pages. Walter is complaining about others allegedly using a phrase that he uses himself.

Re: "Haldane measured that they have a reproduction rate that leaves only 0.1 leftover to pay for substitutions. In other words, the total cost is 30, and paid-off in installments of 0.1 per generation, which means, over the long term, substitutions can occur no more often than one per 300 generations (=30/0.1)"

False. This is not the calculation Haldane used. Specifically, Haldane's value of 0.1 is a selection coefficient, not a reproduction rate, and is calculated from the number of generations, not vice versa.

Re: "He assumed the population does not get stuck on top of a local fitness peak, held there by survival of the fittest, and forever prevented from evolving."

Fitness landscapes change over time. Thus not only is such an assumption justified, but objecting to it shows a lack of understanding of evolution. In fact, Haldane explicitly refers to changes in the environment which affect fitness.

Re: "He assumed the fitness terrain offers no significant obstacles to evolution, such as impassable ‘mountains’ or deep ‘crevasses’ in the fitness terrain."

False. Haldane was not dealing with any specific fitness terrain. His only assumption was that a series of beneficial mutations were possible. (The reference to "impassable 'mountains'" is another indication that Walter does not understand what fitness landscapes are).

Re: "He also assumed the vast majority of beneficial recessive mutations are eliminated ..."

This is not an assumption, but an easily demonstrated mathematical probability.

Re: "... and completely without cost"

The cost of this elimination them would be subsumed by the cost of selection for other (non-recessive) mutations, or the cost of random 'accidents', since that is what is eliminating them.

Re: "Moreover, since the large majority of mutations are known to be recessive, Haldane theoretically eliminated the largest source of beneficial mutations"

Just because the majority of mutations are recessive does not mean that the majority of beneficial mutations are recessive. The 'logic' here is akin to saying "Most humans have dark hair; Norwegians are humans; therefore most Norwegians have dark hair."

Re: "Haldane’s model implicitly assumes no epistasis (or complex, favorable, interactions between the substituting mutations). The vast majority of biological traits (such as brains, hearts, blood clotting, vision, motility, hair, replication, metabolism, etcetera) are products of numerous genes, whose specific combination (not just any random combination of genes) is vastly more beneficial than any of the same genes taken alone or in smaller combinations."

If this is true, it means Haldane made an assumption which is unfavorable to evolution, since epistatic linkage would assist multiple genes to fix simultaneously. Yet Walter repeatedly says Haldane's assumptions were favourable to evolution.

Re: "And it does not change the fundamental problem – Is 1,667 beneficial mutations enough?"

That isn't a problem, it's a question. If the answer is yes - and Walter has done absolutely nothing to suggest that it isn't - then there is no problem.

Re: "Environmental-change (like mutation) is random with respect to a species, and therefore it tends to harm the species. ... Haldane’s calculations ignored those unfavorable factors.

Utter nonsense. Haldane wrote:

"A sudden change occurs in the environment, for example pollution by smoke, a change of climate, the introduction of a new food source, predator or pathogen, and above all migration to a new habitat. ... The species is less adapted to the new environment, and its reproductive capacity is lowered."

Haldane explicitly mentions the very thing that Walter accuses him of ignoring.

Re: ":That is because truncation selection is not realistic. In effect, it ranks individuals, say, by their number of beneficial mutations, and then eliminates individuals below a certain threshold."

False. Truncation is indeed about eliminating individuals who do not reach a certain threshold, but the thresholds involved are specific physical ones. A good example is the size of a young bird or reptile's egg-tooth; those with egg-teeth below a certain size will be unable to break through the eggshell, and will never hatch. Other examples include size, weight, bone strength etc. Truncation selection does not rank individuals by their 'number of beneficial mutations'.

Roy 11:51, 3 August 2006 (CDT)


Walter ReMine’s Response

The CreationWiki article reports Haldane’s Dilemma as reflected in creationist and evolutionist literature. If you have a new angle, then I encourage you to publish in a science journal, out in the open, where it can be reliably cited – and embraced or denied by evolutionary geneticists. Here is not the place to propose new evolutionary solutions.

Haldane’s Dilemma remains a scandal because evolutionary geneticists, by and large, tolerate and allow widespread contradiction, error, and confusion from their fellows (so long as it favors evolution). Rather than correct those faults, evolutionary geneticists abandoned the topic to lesser evolutionists, non-specialists mostly, who advance those faults on various Internet sites (often anonymously or effectively so). This disconnect between evolutionists and their literature enables the scandal to continue.

Roy, an evolutionist, posts his views here without identifying any support from the evolutionary genetic literature. His views fall into three categories:

  1. Misrepresentations of his opponents.
  2. Many of Roy’s views are common on the Internet and have ‘support’ in the evolutionary literature – but are false, and the falsehood is known to evolutionary leaders. These cases show that evolutionary leaders negligently allow confusion and error to thrive, and demonstrate why Haldane’s Dilemma remains a scandal.
  3. Some of Roy’s views have virtually no serious support in evolutionary genetic literature. They are specious and will likely never be published.

Misrepresentations –

  • “Walter [ReMine] is complaining about others allegedly using a phrase that he uses himself.”
ReMine legitimately uses the phrase – “one gene substitution per 300 generations” – in order to critique it. And Roy misrepresents the objection. The mere phrase is not the objection. The objection is that evolutionists never revealed Haldane’s Dilemma to the public – a limit of 1,667 beneficial mutations for human evolution. Even in their technical journals, instead of revealing the problem, evolutionists use the phrase “one gene substitution per 300 generations” – which obscures the problem from view, and may as well be in code.
Also, that phrase promotes the widespread falsehood that each substitution is a new “gene,” with hundreds or thousands of new differences in nucleotides. Because evolutionary leaders allow that falsehood to thrive, the CreationWiki article takes active steps to correct it. That is, according to evolutionary geneticists, the typical beneficial substitution is a nucleotide, not thousands of new nucleotide differences. (Of course, evolutionary geneticists are welcome to change their view, and may likely do so, as the result of the recent genome projects, which discovered vast numbers of “chromosomal rearrangements” in need of evolutionary explanation. The world awaits their discussion, and consensus, on this new data.)
  • “Haldane explicitly mentions the very thing [i.e., the unfavorable effects of environmental-change] that Walter accuses him of ignoring.”
Haldane mentioned a lot of things. But the CreationWiki article correctly states, “Haldane’s calculations ignored those unfavorable factors.”
  • “If Walter's treatment of Haldane's dilemma [is] correct, there hasn't been enough time for any beneficial mutations to fix in these finches; which means Walter's characterization … must be incorrect.”
In Galapagos finches, Roy says 150 years of observation is not enough time for any beneficial mutations to go from start to fixation. He suggests the observed changes in Galapagos finch beaks are instead due to “variation in frequency of existing alleles” (which, by the way, happens to be a widely advertised creationist position). Roy therefore claims my argument must be incorrect. In reality, he misrepresented my argument. My argument does not require any position on when the finch beak mutations occurred or how far they substituted into the population.
My argument merely requires the identification of ‘beneficial-ness’ within a population. And for that, my argument uses the evolutionists’ own claim that some Galapagos finches have favored beaks due to the possession of (at least one) beneficial mutation. The comparison of various finch beaks is taken to show what beneficial mutation can do. Evolutionists cannot legitimately object to the use of their own widely advertised claims. My argument is straightforward: Use examples of beneficial mutations today as a measure of their power to transform organisms. Is 1,667 powerful enough to explain human evolution?
  • “Just because the majority of mutations are recessive does not mean that the majority of beneficial mutations are recessive. The ‘logic’ here is akin to saying ‘Most humans have dark hair; Norwegians are humans; therefore most Norwegians have dark hair.’”
Roy offers neither a theoretical nor empirical solution. Instead Roy misrepresents the problem through a false analogy, involving Norwegians, whose hair color is well known. His analogy does not apply to beneficial mutations – whose rate, characteristics, and indeed very existence, are all controversial matters, almost always discussed theoretically and rarely empirically. These matters cannot be settled by false analogies, and not by assuming them away (as Haldane and other evolutionists do). These matters ought be widely discussed in evolutionary genetics literature – but are not. The CreationWiki article correctly flags this issue as another theoretical problem that evolutionists assumed-away.

Evidence of evolutionist negligence –

  • “[Haldane’s calculation] assumes a selection coefficient of 0.1, which is definitely not ‘approaching 0+’.” …. “the false claim that Haldane used negligible selection coefficients (0+) remains;”
In calculating the limit on substitution rate, all Haldane’s cost equations assumed tiny selection coefficients (approaching 0+), and this favors evolution by giving the absolute smallest total cost of substitution in each case. Moreover, if you plug-in larger selection coefficients, those equations give a cost of substitution lower than the true value – and that further favors evolution. And his equations become increasingly erroneous in favor of evolution as higher selection coefficients are plugged into his equations. Leading evolutionary geneticists know this is true, but did not bother to clarify it in their literature – thereby confounding generations of researchers, such as Roy. The continued confusion on such fundamentals is further evidence of evolutionist negligence. There is no excuse for it.
  • “Haldane’s value of 0.1 is a selection coefficient, not a reproduction rate”
Haldane’s value of 0.1 is a “selection intensity” (Haldane’s words), not a selection coefficient. And “selection intensity” is one of the needless confusion factors that remain predominant to this day. Haldane’s value of 0.1 amounts to the reproduction rate available specifically to pay the cost of substitution. This fact is known to leading evolutionary geneticists, though they allowed the error to thrive in the literature – thereby confounding generations of researchers, such as Roy. Roy’s error is more evidence of evolutionist negligence. Evolutionists have no excuse for allowing such fundamental errors to prevail since 1957.
  • “The cost of this elimination them [in other words, the cost of eliminating beneficial recessive mutations] would be subsumed by the cost of selection for other (non-recessive) mutations, or the cost of random ‘accidents’, since that is what is eliminating them.”
Haldane assumes the vast majority of beneficial recessive mutations are “eliminated, and completely without cost” (those words belong together to correctly state my point). However, even though they are eventually eliminated, their substitution partway into the population requires extra reproduction rate, just as other substitutions do. Therefore, they incur a portion of the total cost of substitution.
Also, these mutations are beneficial – they use resources, and cause the elimination of less favored individuals – thereby raising the cost of substitution for other beneficial mutations (including those others that are not eliminated).
Roy’s analysis is erroneous. His error arises because he focuses on ‘elimination,’ which is a traditional confusion factor that evolutionary leaders allowed to thrive. Instead, the real issue is the ‘growth’ (or increase in number of copies) of the beneficial mutations (regardless of whether they are later eliminated), and the extra reproduction rate necessary to make it happen. Once again, evolutionary leaders allowed confusion to prevail – thereby confounding generations of researchers, such as Roy.
  • “Fitness landscapes change over time. Thus not only is such an assumption justified, but objecting to it shows a lack of understanding of evolution.”
Roy assumes many problems away. For example, a sharp fitness peak can prevent evolution, even if riding upon a large-region of fitness terrain that “changes over time.” And vice-versa, a sharp large-scale fitness mountain can prevent further evolution, even if smaller regions of the same fitness terrain “change over time.’ On the other hand, too much warping and deforming of the fitness terrain can turn evolution into a random process, where beneficial evolution is a moot point. These and many other problems are silently assumed away by evolutionists, especially in their presentations aimed at the defenseless public. Therefore, the CreationWiki article explicitly states Haldane’s many wildly unrealistic, pro-evolutionary assumptions.
  • “Haldane was not dealing with any specific fitness terrain. His only assumption was that a series of beneficial mutations were possible.”
False. Haldane was inevitably dealing with nature’s fitness terrain – he dealt with it by assuming-away any and all obstacles to evolution. Haldane assumed a series of beneficial substitutions is possible unimpeded by the fitness terrain. But fitness ‘mountains’ can pose impassable obstacles, just as a local fitness peak can stop evolution. (See above.) The CreationWiki article explicitly identifies assumptions that Haldane (and Roy) tried to brush aside.
  • “1 beneficial mutation substituted per 300 generations is still being treated as a hard limit rather than as an estimated average;”
The cost of substitution puts a limit, or threshold, on plausible substitution rates – lower rates are plausible, but higher rates are not plausible. Moreover, Haldane’s calculation made many wildly unrealistic assumptions in favor of evolution – and he made no attempt to guarantee those many favorable assumptions actually hold true. In short, the Haldane rate is an upper limit, an unrealistic maximum – not remotely a guaranteed ‘average rate’ of evolution.

Views unsupported by evolutionary genetics literature –

  • “Haldane made an assumption which is unfavorable to evolution, since epistatic linkage would assist multiple genes to fix simultaneously.”
Roy gives a naïve and mistaken view. He silently assumes away the role of nature’s epistasis when combined with: (1) sexual reproduction (which drastically slows beneficial evolution), and (2) cost theory (which further slows beneficial evolution under these circumstances). The CreationWiki article correctly states that Haldane’s assumption of no epistasis is wildly unrealistic in favor of evolution. If evolutionists want to dispute the matter, then their journals are a good place to start. Evolutionary geneticists ought put their reputations to these matters openly, in their literature, where they can be held accountable. These complex matters cannot be resolved here by “Roy.”
  • “Truncation selection does not rank individuals by their 'number of beneficial mutations'.”
The CreationWiki article is not a treatise on truncation selection and its varieties. Instead the article gives the simplest version of truncation selection as it appears in the literature on Haldane’s Dilemma. Such evolutionary proposals assume, for example, that beneficial mutations have roughly the same selection coefficient, and then individuals are ranked by their fitness – which amounts to ranking individuals by their number of beneficial mutations – then individuals below a certain threshold are eliminated. Roy is mistaken about the literature on this subject. His version of truncation selection is not the version used in cost literature.
Roy makes a common evolutionist error. That is, to solve a given problem at hand, he selects an (atypical) example from nature, and then presumes it represents a general solution. His particular chosen example – involving an egg-shell as a ‘threshold’ – does not make truncation selection a general phenomenon.
Moreover, truncation selection does not even reduce Haldane’s Dilemma, if the truncation selection concerns only one substituting mutation. Contrary to Roy’s claim, ‘the number of beneficial mutations ranked within the truncation selection’ is essential to the discussion.
  • “Since in nature mutation rates are low, populations are usually large and mutations are distributed randomly, rather than being parceled out so that everyone gets one, error catastrophe/mutational meltdown will not occur.”
Error catastrophe shows up on any remotely realistic evolutionary simulation, so long as the mutation rate is sufficiently high, and/or the reproduction rate is sufficiently low. (By the way, both those factors are especially prominent concerning the origin-of-life.) Evolutionists scarcely reveal the problem of error catastrophe to the general public – it is another evolutionist scandal. And Roy misrepresents the problem:
  1. Mutation rates are not “low.” Depending on which evolutionary geneticist you cite, (even after all our highly specialized error correction processes) the measured rate is between 70 and 300 new mutations per progeny – sufficient to make it a serious problem. This prompted evolutionary geneticist, Alexi Kondrashov, to publish his paper, “Why have we not died 100 times over?” where he acknowledges the problem, but does not solve it. There remains no consensus on any evolutionary solution.
  2. Roy fails to mention the species’ limited reproduction rate, which is central to error catastrophe.
  3. Though genetic deterioration is faster in smaller populations, large population size does not solve the problem. Moreover, (due to systematic anti-Darwinian patterns in the fossil record), evolutionists claim most beneficial evolution occurs in small populations (said to leave no fossil record) – precisely the circumstances where genetic deterioration is most rapid.
  4. Roy is mistaken. For example, Dawkins’s famous simulation – “METHINKS IT IS LIKE A WEASEL” – has each progeny receive exactly the same number of mutations, which is unrealistic. When altered to distribute mutations realistically (so mutations are distributed randomly at a specified average rate), then the simulation slows to nearly twice the time needed to reach the target phrase. Then, if the reproduction rate is reduced to a level comparable to the higher vertebrates, the simulation goes into error catastrophe. Contrary to Roy’s claim, a realistic distribution of mutations is not sufficient to prevent error catastrophe. (ReMine, The Biotic Message, p 235-236, also, 229 & 245-253).
  • [The 1,667 limit on human evolution] “isn’t a problem” and “isn’t Haldane’s Dilemma.”
Evolutionary geneticists publish no such statements or consensus in their journals. Evolutionary geneticists must locate their backbones, and their vocal cords, and address Haldane’s Dilemma publicly. They must take responsibility for advancing their solutions, and openly place their reputations behind it – not hide behind unsupported opinions posted on an untraceable, transitory, un-readably-tangled, discussion webpage by “Roy.” Once again we see the scandal of Haldane’s Dilemma – a disconnect between evolutionists and their own literature.

Wremine 15:44, 13 August 2006 (CDT)


Making text more simple

Hi.

I am a secondary shool student and I am making some reaserch in creadion vs. evolution discussion. I found out, that there is the Haldene's dilemma. However, I have to say, that I just can't get the point, of the dillema. Like:

In the introduction to The Cost of Natural Selection Haldane writes that it is difficult for breeders to simultaneously select all the desired qualities, partly because the required genes may not be found together in the stock; but, writes Haldane (p. 511), especially in slowly breeding animals such as cattle, one cannot cull even half the females, even though only one in a hundred of them combines the various qualities desired. That is, the problem is for the cattle breeder is that keeping only those specimens with the desired qualities will lower the reproductive capability too much to keep a useful breeding stock. (text is taken from en.wikipedia.com)

I cannot comprehend the last sentence. Why it will lower the reproductive capability? And from where is the number 300? (those mathematical formulas is too complicated to understand)


Answer:

  • If you cull all the stock that does not carry the desired combination of qualities, you will be left with one cow out of a hundred. That one cow will have to provide all the offspring needed to bring the herd up to its previous numbers. The reproductive capacity of the herd will be one hundredth of what it was before the cull. For a breeder, this is not an economic proposition.
  • The number 300 is Haldane's optimistic estimate of the average number of generations taken to spread a mutation through an entire population of a species of large animals:
    • In conclusion, the total cost of substitution is 30, and paid in installments of 0.1 per generation, which means, over the long term, substitutions can occur no more frequently than one per 300 generations (=30/0.1). --Oelphick 05:09, 24 January 2007 (EST)

My point is, wouldn't it be possible to make an article easy to understand for people, who do not understand biology well (i was taught evolution in school a bit, however there was not a word about the Haldene's dilemma).

Sexual reproduction

Although the article says that Haldane's calculation "allowed for sexual reproduction and for numerous concurrent substitutions overlapping in time in arbitrary ways.", I don't think he did. His comment on multiple selection is as follows: "Can this slowness be avoided by selecting several genes at a time? I doubt it, for the following reason. Consider clonally reproducing bacteria, in which a number of disadvantageous genes are present, kept in being by mutation, each with frequencies of the order of 10^-4. They become slightly advantageous through a change of environment or residual genotype. Among 10^12 bacteria there might be one which possessed three such mutants. But since the cost of selection is proportional to the negative logarithm of the initial frequency the mean cost of selecting its descendants would be the same as that of selection for the three mutants in series, though the process might be quicker. The same argument applies to mutants linked by an inversion. Once several favorable mutants are so linked the inversion may be quickly selected. But the rarity of inversions containing several rare and favorable mutants will leave the cost unaltered." (Haldane, 1957).

Haldane was there referring to a group of mutations 'linking together' and substituting together as one substitution. Haldane points out that the infrequency of this "will leave the cost unaltered". See the appendix in The Biotic Message, where Haldane's logic on this point is explained in greater detail. [Hint: Log(10^-4 * 10^-4 * 10^-4) = Log(10^-4) + Log(10^-4) + Log(10^-4) = the cost when linked together equals the sum of the costs when substituted separately.] His argument applies also to a sexually reproducing species. Wremine 21:42, 26 September 2008 (UTC)

But in a species such as humans, with multiple chromosomes, this is no longer the case. Each mutation can substitute simultaneously, and there will only be significant interference when they are close to substitution. For example, in a population of a million, a gene spends half its substitution time at numbers less than a thousand, and the reproductive excess required to increase each one at the expected rate would therefore only be 1/1000th of the species overall capacity.

Your logic there makes no sense. Wremine 21:42, 26 September 2008 (UTC)

In a clonal species this makes no difference because to complete its substitution a mutation must wipe the others from the board. But with multiple chromosomes a mutation may substitute without reducing the frequency of other beneficial mutations, and indeed with less than full effect on their own rate of increase. Thus, the species will come much closer to using its full capacity. For example, a species with reproductive excess of 0.1 could in theory substitute one mutation every ten generations, if the excess were entirely devoted to improved individuals.

Your logic there is mistaken. For example, say the population size is one million, with one of the individuals possessing a beneficial mutation. A reproductive excess of 0.1 can, at most, increase it by 10 percent per generation (=0.1). After ten generations it will be increased to 2.59 individuals [=(1.1)^10]. Obviously that is far short of one million individuals. Wremine 21:42, 26 September 2008 (UTC)
That same population, over ten generations, will have a reproductive excess of one million, of which 999,998.41 will be available for other mutations to increase. A single mutation may take 300 generations to substitute, but if each of 23 chromosomes substitutes a mutation every 300 generations, that is one every 13 generations. Because chromosomes are readily remixed, this is quite possible.Ozcanute 15:07, 27 September 2008 (UTC)
Reproductive excess is an extra reproduction rate. That is its unit of measure. An extra reproduction rate of "one million" is physical nonsense, especially in the higher vertebrates (such as humans) where Haldane's Dilemma is most problematic. Also, Haldane already allowed for sexual reproduction, and for independent segregation of the substituting mutations -- in your terms, this is like assuming an infinite number of chromosomes -- an unrealistic assumption that works in favor of evolution. In other words, everything you just wrote is false. Please inform yourself on Haldane's Dilemma before posting here. This webpage is not a place to debate Haldane's Dilemma, nor for tutorials. This webpage is for discussing how to make this article better. Wremine 04:19, 28 September 2008 (UTC)

Such effects are easy to simulate now that computers have so much memory.

A second sexual reproduction effect not considered is that of mate selection. Consider a species with no reproductive excess (ie all females have exactly two children survive to adulthood). If a gene renders carrier males more attractive to females, such that carriers have twice as many children as non-carriers, then it will increase at a rate of 50% per generation (0% increase for female carriers, 100% for males). This is an effective reproductive excess of 0.5, without any actual excess for the species.

Females are quite capable of selection at such high intensities even when the actual benefit is low - a male who wins one mating contest can win a great many of them, as we see in many primates.

Combining the two effects a species might well substitute mutations at one every two generations; this is also easy to simulate on a computer, where we find that such perfection is approached but not reached. Still, a species might easily achieve one mutation every 10 generations, and this would allow humans to differ from chimps by 50,000 selected mutations - two for every gene we have, according to the results of the Human Genome Project (see http://www.ensembl.org/Homo_sapiens/index.html). --Ozcanute 13:22, 26 September 2008 (UTC)

Haldane already allowed for sexual reproduction and for numerous concurrent substitutions overlapping in time in arbitrary ways. Numerous evolutionary geneticists (such as Kimura, Crow, Ewens, Maynard-Smith, and many others) already accept that point. I am aware of no evolutionary geneticist who disputes that point in the literature.
That's not the way I read the paper, and certainly not what I saw when I tested it. I would encourage you to find a friendly software engineer and simulate this setup; you might be surprised. I'll be happy to send you a copy of my simulation program if you're interested.Ozcanute 15:07, 27 September 2008 (UTC)
If you have a simulation you think is helpful, then you ought try to publish it in a scientific journal. My guess is you won't even try. I strongly suspect your simulation does not simulate Haldane's paper, but instead simulates something even more unrealistically in favor of evolution. Wremine 04:19, 28 September 2008 (UTC)
I do appreciate your interest in this great unsolved problem, Haldane's Dilemma. However, keep in mind, this webpage is not a debating society. If you think you have a solution to Haldane's Dilemma, you ought try to publish it in a scientific journal, or try to cite where the literature contains your solution. (Neither one of those will happen. You are promoting "solutions" that have no serious support in the literature.) This is not really the place to raw-debate your solution. This webpage is for discussing our article on Haldane's Dilemma, and how to make it better. Wremine 21:42, 26 September 2008 (UTC)
Well, then I would suggest that the article would be better if it had a more realistic figure for the maximum number of beneficial mutations.Ozcanute 15:07, 27 September 2008 (UTC)
Again, I point out that you don't cite any sources. You don't have any support. That is because (1) you engage in empty posturing, and (2) evolutionary geneticists extremely rarely offer any figure whatever on the maximum number of beneficial substitutions available for evolution of the problematic species (i.e., humans, apes, elephants, whales, the higher vertebrates). Evolutionists are profoundly negligent on the matter. They strongly avoid publishing such a figure, and they do not reveal Haldane's Dilemma to the public. Your homework assignment is to find a published evolutionary source for your claims. Wremine 04:19, 28 September 2008 (UTC)
I don't think it's fair to describe my comments as "empty posturing" when I'm offering you the program source on which I'm basing them. However, if you're not interested in anything that hasn't been published, I'll take your assignment... but since you suggest I will need to publish it myself, don't expect a quick response. If you change your mind, let me know.Ozcanute 12:58, 28 September 2008 (UTC)
There is a shortcut, since Nunney's computer simulation already has a paper published. Nunney's paper claims his simulation is faster than the Haldane limit. However, (1) Nunney's paper lacks sufficient description of his simulation's operation, thereby preventing others from replicating his simulation, and (2) Nunney refuses to release his software for inspection of its details, and (3) Evolutionists refuse to call for the release of Nunney's software. [That includes the journal where Nunney published his paper, and the NCSE (National Center for Science Education, which promotes Nunney's simulation as a solution to Haldane's Dilemma), and evolutionists in general (for example, at sci.bio.evolution where this matter was discussed at length).] Nunney's simulation was produced with taxpayer support, and is important evidence in public classrooms -- where evolutionists have monopoly control. So, you could press forward: (A) by reproducing Nunney's simulation exactly as described in his paper and then releasing the source code for inspection -- or publicly acknowledge it cannot be done due to ambiguous/incomplete description in Nunney's paper. Or, (B) By pressing evolutionists to release Nunney's software. I predict you will not succeed at these things, and likely will not even try. Why? Because Haldane's Dilemma is a scandal, and evolutionists are not eager to clear it up. Wremine 08:36, 30 September 2008 (UTC)
OK, I've completed the shortcut. See http://sites.google.com/site/genesubstitution/nunney, and see also http://groups.google.com.au/group/sci.bio.evolution/browse_thread/thread/beba079154185a2b# for a short summary; you can comment further on sci.bio.evolution if you like. From that I can say that 1) Nunney's paper contains sufficient detail to reproduce his results, 2) He therefore doesn't need to release the software, although I think it would be a good thing, and 3) I've done a bit better than call for its release. On the downside, he doesn't include either of the effects that I am talking about, although the program I've released allows you to trial them if you wish. But he does beat your 300 generation limit quite handily; you should at least stop talking about that number as if it were a hard limit rather than an estimate.Ozcanute 11:15, 18 March 2009 (UTC)

my two cents on Ozcanute's reply to Remine

"you should at least stop talking about that number as if it were a hard limit rather than an estimate."

you mean like half-lives? ;)--Nlawrence 13:35, 18 March 2009 (UTC)