mirrored file at http://SaturnianCosmology.Org/ For complete access to all the files of this collection see http://SaturnianCosmology.org/search.php ========================================================== From a post from Fall, 95: An interesting drama has been playing itself out on talk.origins over the last few months. A serious geneticist, Walter Remine, has demonstrated a problem which utterly destroys the theory of evolution in anything remotely like any of its present forms. This problem was first described by the noted geneticist J.B.S. Haldane in the late fifties, and is called Haldane's dilemma. The logical implications of the problem are that even the gigantic time frames normally used by evolutionists as an enabling mechanism for evolution, the 4 - 7 billion year age of the Earth which you are familiar with, is nowhere even remotely close to being enough. The talk.origins crew has tried to use all of its normal tricks on Remine, with the end result that they, Darwin, Darwinism, and all other modern encarnations of evolution have been made to look very silly indeed. In ten years of watching and participating (as arch villian) in talk.origins, this is the very worst beating I have ever seen these people absorb. There is no way that anybody with any talent or brains could have been reading all of this and still believe in evolution. The talk.origins crew surely would prefer that viewers in other forums did not see any of this material; naturally, I've been collecting as much of it as possible, and hereby provide you with a sampler: The action began with a couple of posts of mine and Robert Bass' on talk.origins, and Remine's on other forums. ............................................................. My post: A number of the big theories in science are on the ropes today, evolution and the big-bang theory being the most notable. There are numerous problems with evolution, any one of which, were the ideological stakes not so high, should have long since laid evolution to rest. Aside from the impossible problem of abiogenesis, the problem of catastrophism destroying the time-scales normally given out, or the programmatic problems (such as with flying birds), you have the problem of trying to get from something resembling an ape to modern man in ten million years. Walter ReMine notes: >Evolutionary genetics has trade secrets too. The major one is Haldane's >Dilemma, a problem discovered in the 1950s by the famous evolutionary >geneticist, J.B.S. Haldane. Journals discussed it through the 60s, and >ignored it thereafter. Evolutionists never publicly solved it, rather they >brushed it aside. Here are my claims: >1) Haldane's Dilemma is invisible in evolutionary genetics textbooks today, >you will be lucky to find information on the problem. The little available >information is cryptic and opaque, even to serious students. >2) The standard model of evolutionary genetics -- prominently displayed in >every evolution textbook -- is massively inadequate to solve the problem. >Yet evolutionists continue to sell that model because it makes evolution >seem easy and inevitable. >3) Even if I arm you with information about the problem, you will find >precious little in evolutionary textbooks that *might* be taken as a >plausible solution. >4) The problem is robust and firm -- the phenomenon can even be >demonstrated in computer simulations, such as the same one Dawkins used in >his book _The Blind Watchmaker_. >In short, Haldane's Dilemma is a thorough trade secret of evolutionary >geneticists. >My book, _The Biotic Message_, has two chapters (and an appendix) detailing >Haldane's Dilemma and rebuffing the many attempts to solve it. Here I'll >draw from that material to describe the problem, and bring you up to speed. >Then I'll answer your questions, and perhaps eventually we'll have our usual >rip-snortin' debate. I'll keep my descriptions short and easy reading. >Along the relevant primate line, our supposed pre-human ancestors had an >effective generation time of 20 years. (I quote sources and details in my >book, so I'll spare you here.) Imagine ten million years ago -- (that is >two to three times the age of the alleged chimp-human split) -- that's >enough time for 500,000 generations of our presumed ancestors. >Imagine a population of 100,000 of those organisms quietly evolving their >way to humanity. For easy visualization, I'll have you imagine a scenario >that favors rapid evolution. Imagine evolution happens like this. Every >generation, one male and one female receive a beneficial mutation so >advantageous that the 999,998 others die off immediately, and the population >is then replenished in one generation by the surviving couple. Imagine >evolution happens like this, generation after generation, for ten million >years. How many beneficial mutations could be substituted at this crashing >pace? One per generation -- or 500,000 nucleotides. That's 0.014 percent >of the genome. (That is a minuscule fraction of the 2 to 3 percent that >separates us from chimpanzees). >That's not a difficult calculation, yet it immediately reveals a problem. >Is 500,000 beneficial nucleotides enough to explain the origin of humanity >from some chimp-like ancestor? >The problem gets worse. The scenario favored evolution in wildly >unrealistic ways. I could name several, but one is simple: There is no >possible way for a female primate to produce 100,000 offspring each >generation!!! Here's the lesson: > Evolution requires the substitution of old prevalent traits > with new rare traits. But the substitution rate is limited by > the species' reproductive capacity. If an evolutionary scenario > requires an implausibly high level of reproductive capacity, > then the scenario is not plausible. >Haldane saw this problem and posed it within the framework of mathematical >population genetics. We will discuss his calculations later, but his >conclusion was easy to understand. He calculated that the higher >vertebrates (such as mammals) have only enough reproductive capacity to >sustain an average rate of 300 generations per substitution. The literature >seldom states the figure, but when it does, that is the only one offered. >Haldane's Dilemma is glaringly plain. Take the population we discussed >above. In ten million years, it could substitute 1,667 beneficial >nucleotides. That is less than 50 millionths of one percent of the genome. >(And that is *before* we make deductions. For example, Gould says species >typically spend *at least* 90% of their time in stasis, where little or no >evolution occurs. There are other deductions we'll discuss later, but >together they reduce the figure far below 1,667.) Is that enough to explain >the origin of upright posture, speech, language, and appreciation of music, >to name just a few of our uniquely human capacities? Is 1,667 beneficial >nucleotides enough to make a sapien out of a simian? >Haldane's Dilemma is fundamentally simple. Anyone can understand it. >Anyone with a pencil can calculate it and see. Computer simulations clearly >demonstrate the problem. So evolutionists cannot claim they were unaware. >Nonetheless they were cryptic, effectively concealing the problem for nearly >forty years. Few people have heard of it, and evolutionary geneticists >offer no unified coherent solution. Haldane's Dilemma is a major scandal. What Haldane is saying, in effect, is that starting from any ape-like or creature ten million years ago and using assumptions which favor evolutionary change in the direction of modern man in more ways than the evolutionists' fairy godmother could imagine, by 1995 you'd be lucky to get to an ape with a slightly shorter tail. The fact that anybody could try to claim, as they apparently do, that Haldane's assumptions do not favor evolution enough, is an indication of the esxtent to which these people (evolutionists) have succeeded in brainwashing themselves. Let's look at one assumption which is implicit in just about everything you see on evolution and which Haldane is also making: the idea that while unfavorable mutations are known to occur far more often than beneficial ones, it is ALWAYS the beneficial mutation which survives and propogates through a population, just as we know from experience that it is always the good which drives out the bad in government work, academia, politics... You only have to look at this a little bit to realize how stupid it really is. You are starting out with apes ten million years ago, in a world of fang and claw with 1000+ lb. carnivores running amok all over the place, and trying to evolve your way towards a more refined creature in modern man. What's wrong with that? Let's start from about ten million years back and assume we have our ape ancestor, and two platonic ideals towards which this ape ancestor (call him "Oop") can evolve: One is a sort of a composite of Mozart, Beethoven, Thomas Jefferson, Shakespeare, i.e. your archetypal dead white man, and the other platonic ideal, or evolutionary target, is going to be a sort of an "apier" ape, fuzzier, smellier, meaner, bigger Johnson, smaller brain, chews tobacco, drinks, gambles, gets into knife fights... Further, let's be generous and assume that for every one chance mutation which is beneficial and leads towards the gentleman, you only have 1000 adverse mutations which lead towards the other guy. None of these mutations are going to be instantly fatal or anything like that at all; Darwinism posits change by insensible degree, hence all of these 1000 guys are fully functional. The assumption which is being made is that these 1000 guys (with the bad mutation) are going to get together and decide something like: "Hey, you know, the more I look at this thing, we're really messed-up, so what we need to do is to all get on our motorcycles and pack all our ole-ladies over to Dr. Jeckyll over there (the guy with the beneficial mutation), and try to arrange for the next generation of our kids to be in better genetic shape than we are..." Now, it would be amazing enough if that were ever to happen once; Darwinism, however, requires that this happen EVERY GENERATION from Oop to us and, as far as I can tell, Haldane is assuming this also. I haven't even mentioned the gambler's problem, which says that in each generation the guy with the good mutation is starting out with one dollar, the biker gang with 1000 dollars, and the house, i.e. the general population, with $100,000 or something like that. And yet, Colby and others claim that Haldane is making unreasonable assumptions. .................................................................... Phil Nichols responded with the usual talk.origins technique, calling ReMine a liar and me an idiot: Path: news3.digex.net!news2.digex.net!howland.reston.ans.net!news.sprintlink.net!uunet!in1.uu.net!library.erc.clarkson.edu!ub!newserve!rebecca!pn8886 From: pn8886 at csc.albany.edu (Phil Nicholls) Newsgroups: talk.origins Subject: Re: Haldane's dilemma: the real story Date: 30 Mar 1995 20:06:30 GMT Organization: University at Albany, SUNY Lines: 168 Message-ID: <3lf306$8gj at rebecca.albany.edu> References: <3l949u$n0j at rebecca.albany.edu> <3ld9ep$dft at ra.lib.ucalgary.ca> NNTP-Posting-Host: thor.albany.edu >> Phil > Ted >> Haldane himself pointed out that the selective deaths would >> be less detrimental to the population if they came in >> embryonic or juvenile stages. > I haven't really had time to try to totally comprehend that > paper yet, but it appears obvious that Haldane realized he had > just wiped the theory of evolution, and was trying in a number > of ways to soften the blow. Please don't project TEDOLUTION onto Haldane's work, especially before you have "totally comprehended" it. Haldane's dilemma is a dilemma because work by Dobzhansky and others in the 50's had provided experimental confirmation of natural selection, race formation and speciation. >Haldane's dilemma applies to very large populations. > Given standard assumptions, the human population would > have always been "very large" during the time under discussion. Ted, population has a specific meaning in biology. The human species (Homo sapiens) or any species, for that matter, is made up of many populations or demes. A population is a group of organisms living belonging to the same species living in the same place at the same time. Some human populations today are large, some are small and relatively isolated. Through much of the pleistocene, "human" (i.e. hominid) populations were small. Large population concentrations did not occur until the neolithic and the invention of argiculture. > Haldane appears to be saying that you'd have to get the population > down near zero before you could have some new trait spread > through the surviving population as it rebuilt itself, but that > a genetic change which would drive the population that low would > destroy it. Actually, that is not what Haldane is saying at all. You are casting Haldane after Velikovsky. >> Under punctuated equilibrium Gould and Eldredge have proposed >> that the rate of evolution is best described as long periods >> of relative stasis punctuated by periods of more rapid >> evolution. The bursts of evolution take place in SMALL >> PERIPHERAL POPULATIONS. > Velikovsky's version of this whole thing assumes a flood or > some other worldwide catastrophy, all animal groups either > being exterminated or coming to some very low population > point, and then old and mutated species reforming populations > as they expand into a world all theirs. The populations would > quickly grow and spread and become diverse. Whether change > could be caused by chance processes in such a scenario is > still an open question (I strongly doubt it), but at least > inbreeding in a small space over a long time does not become a > factor. There is no evidence of a worldwide catastrophy of this magnitude since the K/T event of 65 million years ago. > Gould and Eldridge appear to be claiming that inbreeding is > the source of all genetic progress. There are real reasons > why cousins are not allowed to marry eachother in most > nations. You only have to think about that one a tiny bit to > understand how silly it is. As I pointed out earlier, punctuated equilibrium does not involve "inbreeding." >> Hence the cost of selection is > Again, anything can happen in a fairy-tale. Your standard response when you can't deal with the facts. It is a FACT that smaller populations reduce selection cost. That is a built in assumption in Haldane's models. >> It is not that Haldane's assumptions "do not favor >> evolution." Haldane's conclusions are based on a mathematical >> model and as we all know all models make certain assumptions. > The assumptions which ReMine made favored evolution in evume, as did Haldane, that this is a population of diploid organisms. That means that if we have 100,000 organisms we are talking about 200,000 alleles. Since ReMine says a new allele (singular) appears in two individuals by mutation we must assume that the individuals are heterozygous at this locus. If A is the new allele and a is the old allele. The in two individuals we have a mutation from a ----> A. Let p be the frequency of a and q be the frequency of A, 2 q = --------- = 0.00001 therefore p = 0.99999 200,000 The initial frequency of the new allele is 0.00001. A very very low frequency and if you will recall the lower the frequency of q the higher the cost of selection. Therefore this assumption, contrary to ReMine's claim, is not the best possible case. See Crow and Kimura's book if you want to confirm this. Now, the cost of selection per generation can be calculated as follows: Maximum Fitness - Mean Fitness C = --------------------------------- Mean Fitness This is one of two models that Haldane bases his model on. Now ReMine tells is that the two individuals with the new allele survive, all the rest are killed off. The fitness of the individuals with the new allele is 1.0, the fitness of the individuals with the old allele is 0. Mean Fitness is then 2/100,000 = .00002 Hence in ReMine's model, where Maximum fitness is 1.0: 1 - .00002 C = ------------------- .00002 C = 4999!! Now look at you the article by Haldane you will find that he estimated the Cost of selection should be between 20 and 100 per generation, with an average of 30. Not 4999 In other words, ReMine has used an example of ultra-hard selection and this was chose specifically to maximize C. He chose the smallest possible frequency of q, which also maximized C. You've been HAD, Ted. You've been lied to. He sounds impressive, but when to take apart his argument you find deception. Doesn't that bother you? ----------------------------------------------- Marked as a lo"Why post evidence, when I can simply post lies, and avoid all de facts, unlike dem t.o. guys." So Ted boot up, and he post de posts, and he write dem stories, "how can I b.s. de most?" his mind be fried, he spew dat pollution, an' all he keep talkin' 'bout is REMINEOLUTION... Apologies to David Iain Greig -- "To ask a question you must first know most of the answer." - Robert Sheckely Phil Nicholls (pn8886 at thor.albany.edu) .......................................................................... The arrogance, along with David Iain Greig's obvious lack of talent, might be tolerated if Nicholls knew what he was talking about... I replied: Phil Nicholls writes: >> Velikovsky's version of this whole thing assumes a flood or >> some other worldwide catastrophy, all animal groups either >> being exterminated or coming to some very low population >> point, and then old and mutated species reforming populations >> as they expand into a world all theirs. The populations would >> quickly grow and spread and become diverse. Whether change >> could be caused by chance processes in such a scenario is >> still an open question (I strongly doubt it), but at least >> inbreeding in a small space over a long time does not become a >> factor. >There is no evidence of a worldwide catastrophy of this >magnitude since the K/T event of 65 million years ago. There is a mountain of such evidence, and you can view a tiny fraction of it at: http://access.digex.com/~medved/Catastrophism.html >> Gould and Eldridge appear to be claiming that inbreeding is >> the source of all genetic progress. There are real reasons >> why cousins are not allowed to marry eachother in most >> nations. You only have to think about that one a tiny bit to >> understand how silly it is. >As I pointed out earlier, punctuated equilibrium does not >involve "inbreeding." Sounds like you need to read up on punc/eek... Inbreeding is normally taken to be small isolated groups of individuals breeding only amongst themselves over a period of time which needn't be more than a century or two to cause major problems, at least amongst humans, and we are talking about proto-humans and humans here. If that isn't what Gould and Eldridge are talking about, then what ARE they talking about? The peripheral population undergoing a very substantial change in twenty years and then taking over and replacing the main population??? >diploid organisms. That means that if we have 100,000 >organisms we are talking about 200,000 alleles. Since ReMine >says a new allele (singular) appears in two individuals by >mutation we must assume that the individuals are heterozygous >at this locus. If A is the new allele and a is the old >allele. The in two individuals we have a mutation from >a ----> A. >Let p be the frequency of a and q be the frequency of A, 2 >q = --------- = 0.00001 therefore p = 0.99999 > 200,000 >The initial frequency of the new allele is 0.00001. A very >very low frequency and if you will recall the lower the >frequency of q the higher the cost of selection. >Therefore this assumption, contrary to ReMine's claim, is not >the best possible case. See Crow and Kimura's book if you >want to confirm this. >Now, the cost of selection per generation can be calculated as >follows: > Maximum Fitness - Mean Fitness > C = --------------------------------- > Mean Fitness >This is one of two models that Haldane bases his model on. >Now ReMine tells is that the two individuals with the new >allele survive, all the rest are killed off. The fitness of >the individuals with the new allele is 1.0, the fitness of >the individuals with the old allele is 0. >Mean Fitness is then 2/100,000 = .00002 >Hence in ReMine's model, where Maximum fitness is 1.0: > 1 - .00002 > C = ------------------- > .00002 > C = 4999!! >Now look at you the article by Haldane you will find that he >estimated the Cost of selection should be between 20 and 100 per >generation, with an average of 30. Not 4999 >In other words, ReMine has used an example of ultra-hard >selection and this was chose specifically to maximize C. >He chose the smallest possible frequency of q, which also >maximized C. ....................................................................... Right... Let's look at what ReMine is saying: >Imagine a population of 100,000 of those organisms quietly evolving their >way to humanity. For easy visualization, I'll have you imagine a scenario >that favors rapid evolution. Imagine evolution happens like this. Every >generation, one male and one female receive a beneficial mutation so >advantageous that the 999,998 others die off immediately, and the population >is then replenished in one generation by the surviving couple. Imagine >evolution happens like this, generation after generation, for ten million >years. How many beneficial mutations could be substituted at this crashing >pace? One per generation -- or 500,000 nucleotides. That's 0.014 percent >of the genome. (That is a minuscule fraction of the 2 to 3 percent that >separates us from chimpanzees). So far, he hasn't said a word about genetics, Haldane, or anything like that at all. What he IS talking about is the population-replacement-in-toto fairy doing her thing every twenty years, and he notes that even THAT is not nearly enough to get from anything like an ape to a human in 10 million years. He goes on to note that: > Evolution requires the substitution of old prevalent traits > with new rare traits. But the substitution rate is limited by > the species' reproductive capacity. If an evolutionary scenario > requires an implausibly high level of reproductive capacity, > then the scenario is not plausible. >Haldane saw this problem and posed it within the framework of mathematical >population genetics. We will discuss his calculations later, but his >conclusion was easy to understand. He calculated that the higher >vertebrates (such as mammals) have only enough reproductive capacity to >sustain an average rate of 300 generations per substitution. The literature >seldom states the figure, but when it does, that is the only one offered. I.e. he notes that the instant any sort of a real-world estimate of times for a change due to mutation to spread over a large population, things get even more ridiculous and out of hand. All of that strikes me as pretty simple. I have emailed ReMine a copy of your post and will get back to you if he adds anything beyond what appears obvious to me, which is that you are assuming that gentic principals somehow enter into the scenario involving the population-substitution fairy. Understanding where you are coming from is further made difficult by the observation that your comments on hard selection appear to undermine any case for ape->human in 10 million years being possible rather than support any such case, as one would assume you intended to do. Beyond that, ReMine has recently been quoted as follows: "Note of clarification: That formula is not the definition of cost. The cost of a given situation is defined as its genetic deaths per survivor. (And that cost must be "paid" by the reproduction of the survivors, in births per survivor.) The actual cost depends on the specific situation. The above formula gives the theoretical absolute minimum cost of any single substitution under any and all situations. It gives an unrealistically low estimate for real world situations. "Remember, Haldane-style cost arguments use the term "cost" in a very specific way. Anything going from 'few' to 'many' incurs a reproductive cost. Nothing can go from 'few' to 'many' without paying that cost. The reproduction rate limits the ability to pay, and thereby limits the speed of substitutions. It's quite mechanical and unavoidable. It even shows up clearly in computer simulations. "When you say, "A beneficial trait was substituted into the population" you are simultaneously saying, "All the organisms without the trait died off without heirs" which means "They had to be replenished by the reproduction of the survivors." The fact that the trait happens to be beneficial is virtually irrelevant. *Any* trait that is substituted into a population incurs a cost of substitution. What this means, amongst other things, is that the population-replacement- fairy scenarios, far having a cost of 5000, are free; one second after the fairy does her thing, the max and mean fitness are the same and the cost of the entire transaction is zero. Or, if that isn't obvious, think of it in the terms noted above by ReMine: there is no population loss (cost) to make up. >You've been HAD, Ted. You've been lied to. He sounds >impressive, but when to take apart his argument you find >deception. >Doesn't that bother you? The arrogance here is sort of like icing; the cake is totaly misunderstanding the nature of the non-cost involved in the scenario in which the population- replacement fairy picks up the tab. ....................................................................... And then the heavy cavalry arrived: From: us236454 at dawn.mmm.com (Walter Remine) Newsgroups: talk.origins Subject: Haldane's Dilemma Date: 19 Apr 1995 12:03:17 -0500 Message-ID: <3n3fol$a04 at dawn.mmm.com> *** Haldane's Dilemma -- continued *** Andy Peters offers what he feels are "by far the strongest arguments against" Haldane's Dilemma: >(1) Soft selection and truncation selection radically reduce the >demographic cost (and therefore the rate limitation) of selection. Andy says "soft selection" and "truncation selection" are important solutions to Haldane's Dilemma. Okay, then these topics should be especially prominent and clear in the textbooks. They should be front and center, unavoidable. Right? No student should be able to miss it, right? There should be empirical support for these, right? These should be linked, indelibly, unmistakably to the model of evolutionary genetics, right? ***But you won't find that.*** At least, it will be so exceedingly rare you won't find it on your own. The typical evolutionary genetics textbook doesn't discuss them -- you'll be lucky to find a few pages devoted to it -- the description will be cryptic -- the empirical support will be completely absent -- and they will be disjointed and disconnected, with little linking them to a coherent model of evolutionary genetics. If you find these topics at all, you will wonder: What the heck is it doing in this book? You rarely hear about truncation selection -- unless you challenge evolutionists on Haldane's Dilemma. *Then* they will tell you about it!!! And as always, they will put you down, saying YOU are ignorant, because YOU didn't go to the libraries, because YOU didn't read all those books, (because YOU didn't decipher the mysterious code-words, because YOU didn't slog your way through all the misdirection and the half-stated arguments). They will blame YOU!!! That is standard procedure for evolutionists, (and a recurring theme of my exposes'). Truncation selection is difficult to sell, and evolutionists rarely sell it. Rather, the standard model of evolutionary genetics -- prominent in *all* evolution books -- has truncation selection nowhere in sight. Let me make this point even clearer. John Maynard Smith was into Haldane's Dilemma early, his 1968 paper offered a "solution" -- it was truncation selection. Does it (or "soft selection") show up in his 1989 college textbook, _Evolutionary Genetics_? Nope. Andy's post offers two supposedly "important" solutions -- but they are both effectively absent from the textbooks. His post shows what a scandal Haldane's Dilemma is. Neither the problem, nor its supposed solutions are conveyed to the student. >(2) Sexual reproduction allows several traits to spread through the >population without detrimentally affecting the spread of other traits; >i.e., replacements don't have to - and probably almost never do - occur >sequentially. Both I and Haldane allow that replacements can occur non-sequentially, and that this does not solve Haldane's Dilemma. Sexual reproduction combined with the standard version of evolutionary genetics -- the one model prominently displayed in all the textbooks -- cannot ease Haldane's Dilemma, because traits spread through the population -- on average -- with no effect on the spread of other traits. On average, they neither help nor harm the spread of other traits. Each substitution incurs its own cost. The cost is paid separately, and the payment for one beneficial trait cannot, on average, pay for the substitution of another. >(3) (To me, by far the most damaging argument) Haldane's predictions >depended non-robustly on the assumption that there is a limit to the >amount of variation a population can support at a given time. Low >variation -> low rate of substitution. Less than ten years after Haldane >pointed this out, however, it was shown that more variation than *anyone* >suspected is in fact found in most natural populations. High variation -> >high rate of substitution. .... Andy's argument is mistaken on several levels. First, high genetic variation does *not* inherently speed evolution (contrary to popular evolutionary thought). That was an illusion created by mis-applying Fisher's Fundamental Theorem of differential accrual (commonly mis-named as the "Fundamental Theorem of Natural Selection"). That misapplication was most prominently made by evolutionary geneticist James Crow. He used the theorem to calculate the speed of evolution based on the measured levels of high genetic variation, but he made several errors. One major error was that the theorem does not apply (it is void) to situations of real biological change where mutation, recombination, and migration are injecting new variation into a population. When the theorem applies to a situation it predicts the evolutionary process will grind to a halt -- it predicts precisely the opposite of what Crow was claiming. Correctly understood, the theorem is no help whatever to macro-evolutionists. (Again all details and documentation are in my book.) Second, genetic variation does not solve the problem. Haldane's Dilemma is about substitutions, their reproductive costs, and how they are paid -- and genetic variation cannot pay the costs. No amount of genetic variation can pay the costs or reduce the costs. Genetic variation is not a solution. Third, the observation of high genetic variation in natural populations only deepened the problem. Selectionists tried to explain this as the result of heterozygote advantage, and the neutralists quickly pointed out that this mechanism -- all by itself -- would incur a reproductive cost higher than mammals could plausibly pay. If a mammalian species' entire reproduction couldn't pay the costs of heterozygote advantage, then it certainly couldn't pay the additional costs of substitution -- thus this compounded Haldane's Dilemma. The selectionists responded by de-emphasizing heterozygote advantage, in favor of somewhat less costly balancing selection. None of this eased Haldane's Dilemma. >It's ReMine who's got it wrong. ... the cost of substitution is simply >one form of genetic load (termed, among other things, "lag load"). Cost and load are not the same thing. They are quite different, but their mechanics have similarities. For example, there are costs of: substitution, segregation, and mutation. And those have parallel's in: substitutional load, segregational load, and mutational load -- which are various forms of genetic load. Thus, substitutional load is a form of genetic load. But "substitutional load" and "cost of substitution" are different concepts -- that is Andy's mistake. Such mixups between "load" and "cost" are common. >[Haldane's Dilemma] hasn't been forgotten at all. >It's just not called "Haldane's dilemma" anymore, ... Thank you, Andy, for that admission. Haldane's Dilemma is the trade secret of evolutionary genetics. Rarely can you find it even mentioned, much less clearly described. Students get no vision, no hint, no idea of the problem, and no coherent vision of a solution. >.... Perhaps ReMine should look up "lag load" and "genetic load." >I recommend a recent book by Bruce Wallace, "50 Years of >Genetic Load." Wallace did not solve the problem. His material is an extended demonstration of how the concept of genetic load helped conceal, rather than reveal, Haldane's Dilemma. Walter ReMine Standard disclaimer: These thoughts are mine. P.O. Box 19600 Saint Paul, MN 55119 From: us236454 at dawn.mmm.com (Walter Remine) Newsgroups: talk.origins Subject: Haldane's Dilemma Date: 20 Apr 1995 21:01:42 -0500 Message-ID: <3n73m6$llu at dawn.mmm.com> *** More on Haldane's Dilemma *** First a brief review. I claimed that both Haldane's Dilemma and its supposed solutions (for example, soft selection and truncation selection) are effectively absent from evolutionary genetics textbooks. Chris Colby then claimed he found them in every textbook. The talk.origins crowd then engaged in ad hominem attacks on me, claiming I am a "liar" and ignorant, among other things. But Andy Peters now confirms my point. He claims it is no longer called Haldane's Dilemma (therefore it is absent from textbooks), and he admits that the solutions are typically absent from textbooks. Andy claims the solutions (soft selection and truncation selection) are absent because they are too difficult and "it would be silly to spend a lot of time talking about them." As justification he identifies several specialized topics from evolutionary theory that are also seldom found in textbooks. (He lists topics such as the modeling of evolutionarily stable strategies, ESS.) But that does not justify the situation. Those topics are narrow (and do not adversely impact the standard model of evolutionary genetics), so they may legitimately be discussed separately. But Haldane's Dilemma is not a narrow issue affecting some narrow arcane ecological situation. Rather, it is broad and fundamental, and it severely limits all macroevolution (especially for any species with low reproduction, such as mammals). The standard model of evolutionary genetics (the one model prominent in all the textbooks) is massively inadequate to solve the problem, yet evolutionists teach that model anyway -- without even mentioning the problem or its supposed solutions. This is a scandal, and there is no excuse for it. Andy shows confusion about what the solution is. One moment he claims that soft selection and truncation selection are key solutions, the next he claims the observed high levels of genetic variation "prove that the problem doesn't exist." Though Andy didn't intend it, this is typical of the run-around that evolutionists currently give on this issue. It is difficult to get a straight answer, what with a dozen or so different (erroneous) answers floating around. The confusion is evidence that the problem was never actually solved. The confusion also helped obscure the problem for decades. (More on this below.) ****** I wrote: >> Truncation selection is difficult to sell, and evolutionists >>rarely sell it. Rather, the standard model of evolutionary >>genetics -- prominent in *all* evolution books -- has >>truncation selection nowhere in sight. Andy's latest post confirms my point above. Then he attempts to excuse it. He says "This is primarily because the math of truncation selection ... is a real hassle, even for the best minds in the field." It is unfortunate that the math is complicated, but that is no excuse for leaving truncation selection out of the textbooks -- especially if evolutionists claim it is an essential part of evolution and an essential part of solving Haldane's Dilemma. Truncation selection is difficult to sell, not because of the math, but because of a lack of empirical support, and because of a lack of theoretical justification that it is a plausible way for nature to behave. Truncation selection, if it operated, would also have side effects harmful to the evolutionary process. Moreover, teaching truncation selection might raise student awareness of the problem it was intended to solve -- Haldane's Dilemma. All these issues (not just the math) are entirely avoided in the textbooks. Students get no hint, not the foggiest idea, of the problems. Evolutionary genetics is a theoretical smorgasbord, where evolutionists pick and choose models: The answer they give depends on the question you ask. The problems of one model are ignored by conveniently choosing a different model. Ask them about sexual reproduction, and they will emphasize the critical importance of epistasis. Ask them about Haldane's Dilemma, and they will emphasize truncation selection. But generally they will emphasize their standard model, which is a lot easier to sell. This smorgasbord approach shows up in Andy's post: >If you look in the literature - particularly current literature >in the maintenance of sexual reproduction and mating >systems evolution - you'll see that epistasis is playing an >extremely important part in explaining a number of >phenomena. But, for a very large set of evolutionary >questions, ... it's just a lot easier to use the simpler model. ****** Sexual reproduction, combined with the standard model of evolutionary genetics, is massively inadequate to solve Haldane's Dilemma. Andy disagrees. To support this he starts with an "extreme" situation, one which he is "not claiming would ever happen in the real world" (his words). He has us imagine that one individual has somehow (magically?) acquired 100 new beneficial mutations. He says these are then substituted altogether, all as one -- 100 beneficial substitutions for the cost of one. However, Andy and I both agree that such a situation is "extreme," unrealistic, and cannot be relied on over the long haul to improve the substitution rate. Individuals don't suddenly receive 100 new beneficial mutations. Moreover, sexual reproduction tears apart genomes and mixes them every generation. This will rapidly split apart the 100 mutations so they are no longer substituted as one unit. This way they would each incur the full cost of substitution. Andy insists there is something in sexual reproduction that lowers the cost of substitution. But there is *nothing* in the standard genetic model to allow that. Sex mixes traits all around the population. Sex brings traits together, and breaks them apart with equal facility. The traits are substituted, on average, without helping or hindering the substitution of other traits. Sexual reproduction within the standard model cannot solve Haldane's Dilemma. Evolutionary geneticists like John Maynard Smith recognized that. That is why they proposed radically different genetic models, such as truncation selection, as solutions to Haldane's Dilemma. >I'm saying that the *very same models* that predict a limit >to the rate of evolution (Haldane's Dilemma) also predict that >there should be very little genetic variation in natural populations. That is incorrect. Haldane's Dilemma does not require or predict low genetic variation. The problem remains for both high and low genetic variation. However, the observed high levels of genetic variation deepened the problem by increasing the cost incurred in maintaining the variation -- which takes a bigger bite out of the available reproduction -- which leaves less reproduction to pay the costs of substitution -- which ultimately slows the maximum plausible rate of substitution. In this way the discovery of high genetic variation made Haldane's Dilemma worse. ((NOTE #1: Andy explained his reasoning in only one obscure sentence: >This is because the same demographic cost that limits substitution >of alleles should manifest itself when there are large amounts >of variation across the genome within populations. His explanation makes no sense as it stands, much is left out, and I suspect he is making the classic mistake of using the word "cost" when he means "load".)) ((NOTE #2: Haldane's concept of cost and the concept of genetic load are different, with different meanings, and different formulas, leading to two different approaches to analyzing Haldane's Dilemma and the rate of substitution. Andy says cost and load are the same thing. We are in total disagreement here.)) Creationist Paul Nelson recently brought the following substantial support to my attention. George C. Williams is an expert evolutionist of considerable renown. His latest book, _Natural Selection: Domains, Levels, and Challenges_, has a chapter appropriately titled "Other challenges and anomalies." In it, his first section is "Haldane's dilemma" -- Yes, that is what he calls it. He writes: "In my opinion the problem [of Haldane's dilemma] was never solved, by [Bruce] Wallace or anyone else. It merely faded away, because people got interested in other things. They must have assumed that the true resolution lay somewhere in the welter of suggestions made by one or more of the distinguished population geneticists who had participated in the discussion." He briefly recounts the problem and the failed of attempts to solve it. He concludes: "I think the time has come for renewed discussion and experimental attack on Haldane's dilemma." Haldane's Dilemma is the trade secret of evolutionary genetics, it was never solved. Walter ReMine Standard disclaimer: These thoughts are mine. P.O. Box 19600 Saint Paul, MN 55119 From: us236454 at dawn.mmm.com (Walter Remine) Newsgroups: talk.origins Subject: Haldane's Dilemma Date: 24 Apr 1995 12:57:15 -0500 Message-ID: <3ngopr$lng at dawn.mmm.com> *** Haldane's Dilemma, and the textbooks *** The latest batch of posts concentrates on the extent to which textbooks do or don't deal with Haldane's Dilemma. My opponents claimed the issues are discussed in every book they looked in, and they selected several examples, but there were no surprises. We still have very different perspectives. Let me summarize the literature in several stages. First, there are the books aimed for wide public consumption. These are generally books that sell evolution to the public, as a precise and testable science. To do that, evolutionists inevitably reach for population genetics as the convincer. The simple, easy-to-sell genetic model is the one sold -- truncation selection and soft selection (the alleged solutions to Haldane's Dilemma are nowhere in sight. All non-college (and most college-level) evolution books and articles follow that practice. I can bury you with examples, but will here offer only Kitcher's anti-creation book, _Abusing Science_. He sells evolution via the simple genetic model, and adds, "Mathematical population genetics tells us *precisely* how the forces of immigration, emigration, mutation, and natural selection produce evolutionary change." (Contrast that with my opponents' admissions in this thread recently.) All those books are wrong for doing that. If truncation and/or soft selection are essential to solving Haldane's Dilemma, *then they should be taught* (and defended) and any genetic model lacking them should have appropriate disclaimers. No excuses. This matter gets especially bad when you get to textbooks *specializing* in evolutionary genetics. Any of these -- even mere introductory texts -- should have the solutions to Haldane's dilemma indelibly connected to the evolutionary model (not necessarily with all the math), and it should be painfully clear to any student that these are not optional. On the other hand, if a problem of such broad importance as Haldane's dilemma were unsolved, then it would be inexcusable to leave it out of the books. Either the solution, or the problem, or both, must be clearly displayed in the book. But that is not what we find. John Maynard Smith is a world authority on Haldane's Dilemma and cost issues, and in the 1960s proposed truncation selection as a solution. The preface for his recent textbook, _Evolutionary Genetics_, says the book is intended for advanced undergraduates, is hopefully useful to graduates, and that "proper training in science requires that undergraduates are confronted by the problems of contemporary science." The book discusses the usual standard version of evolutionary genetics. But it doesn't discuss the following: Haldane's problem or any version of it, cost of substitution, substitution load, truncation selection, or soft selection. Everything about the problem and its solutions is absent. This is commonplace in textbooks on evolutionary genetics. Here is an additional brief sampling of modern books, where everything is absent. Molecular Evolutionary Genetics, Nei, 1987 Genetics of Populations, Hedrick, 1983 (***Mentions soft selection, but doesn't tie it in as essential.) Population Genetics and Molecular Evolution, edited by Ohta and Aoki, 1985 (***Mentions truncation selection in one paragraph, but doesn't define it or tie it in as essential.) Rates of Evolution, edited by Campbell and Day, 1987 (About half this book is evolutionary genetics, but the stated topic of the book makes the omission of both the Haldane problem and solution especially notable.) Genetics, Paleontology, and Macroevolution, Levinton, 1988 (***Mentions truncation in one paragraph, but doesn't tie it in as essential.) Theoretical Population Genetics, Gale, 1990 Population and Evolutionary Genetics: A Primer, Ayala, 1982 Now we move to books that at least mention Haldane's problem. Exceedingly few of these refer to it as "Haldane's dilemma". (Most evolutionists will get angry if you call it a "dilemma", as you have already seen in our ongoing discussion here.) The books dramatically downplay it, insisting it's not a problem. So they often give short-shrift to the real mechanics of the issue, with little more than a few sentences, perhaps a paragraph. Even then they generally fail to tie-in truncation selection and soft selection as essential to the solution and essential to the evolutionary model. (Futuyma's text is an example here.) Finally we get to those few texts that actually discuss Haldane's problem. But first, let me briefly restate what that problem was. In 1957, Haldane recognized that it takes excess reproduction to drive (or "pay" for) a substitution, and that each substitution incurs a cost that must be paid. He rigorously defined that cost, and estimated that, on average, it is 30. In effect, 30 entire populations must selectively perish during the course of one substitution. He estimated that, on average, in each generation the typical mammalian population could reproduce itself plus provide enough excess to pay for all sorts of things (random death, and deaths caused by the influx of harmful mutation, and deaths caused by other processes like heterozygote advantage) and still have remaining a reproductive excess of ten percent of its population size. This excess reproduction could pay for substitutions. Thus, on average, the cost is 30, and it is paid in installments of 0.1, so it takes 300 generations to pay for one substitution. People can understand reproduction "payments" and that there are firm limits to it -- female primates cannot possibly average 50 births each! So the key to Haldane's argument was understanding the reproductive costs of substitution. Let me put it this way: Nothing can go from 'few' to 'many', without a reproductive cost being paid, and there are firm theoretical limits to how low that cost can be. Understand that simple concept, and you can overturn some of the commonplace pseudo-solutions to Haldane's Dilemma. Put it all together right, and you have a broad argument that many people can understand. But it requires a clear explanation of cost and its unavoidability, otherwise Haldane's dilemma is incomprehensible to the student. Haldane's argument made the costs and payments explicit, and that increased clarity. For example, tables showed how cost depends on dominance of the mutation. The exceedingly high cost of beneficial recessive mutations became clear, and that prompted some discussion about how to avoid it. In effect, some evolutionists attempted to throw away *beneficial* mutations -- when recessive -- because the cost is so high as to drag down the entire evolutionary scenario into implausibility. That problem was never really solved, but at least the problem was clear in a few presentations. Today, not only is the name "Haldane's dilemma" almost completely absent from the modern textbooks -- so is Haldane's problem, as he formulated it. I mean two things here. His cost/payment problem is either absent altogether, or it is substantially replaced by a recasting of the problem into a total genetic load argument. But either way, Haldane's original cost/payment analysis is underdiscussed, and this leaves students ill-prepared to grasp the genetic load argument. Most textbooks don't adequately explain what cost is, why it is unavoidable, and that there are firm limits it can't go below. They give little feel for cost, and there is even less on details like the high cost of beneficial recessives. Within a few years, evolutionists had recast Haldane's problem in a different direction, whose focus is the total genetic load of all the substitutions together. It is called a genetic load argument, and though much of the underlying mechanics is the same as before, its presentation is dramatically different: with different formulas, and many new unnecessary confusion factors. The presentations argue in the opposite direction from Haldane, and tend to focus almost exclusively on matters that don't show up in the cost/payment analysis. That is why I say the cost and load arguments are different. The load argument goes something like this. You start by assuming evolution is possible via the substitution of distinct mutations, and by assuming some overall substitution rate that would make evolution plausible to you. Typically, those assumptions are only vaguely stated, and can often go unexamined. The numbers are typically given in terms of "gene" or "amino-acid" substitutions, (rather than "nucleotides") and in substitutions "per year" (instead of "per generation"), this makes their interpretation less transparent, less immediate, and less transferable to other situations where the generation time may be different. Some presentations use estimates of the genome size and the number of normal-sized genes. Some presentations then add an estimate of the beneficial mutation rate. These factors take the student ever further away from things that are known, and increase confusion, which is unfortunate because the argument can be stated without them and thus made more compelling. On the other hand, the issue of the high cost of beneficial recessives is totally ignored, and many students are unaware of it. The total genetic load is then calculated. Actually, it is just the total substitutional load, but this detail is often not pointed out, which means all other types of load are implicitly ignored. Those omissions dramatically tilt the presentation in favor of the evolutionary scenario. The analysis results in a number for the load, and the final task it to interpret it to see if it makes sense for the species under consideration. If it does seem plausible, then the underlying substitution scenario seems plausible. But the immediate results are way too implausible, and the load is far too high. This has resulted in a number of approaches intended to explain away the situation. These generally take the form "The load only seems way too high, in reality perhaps we are substantially incapable of seeing the effects of load in natural populations. Perhaps the populations are actually carrying a high load and it's just that we have difficulty seeing it." The issue then becomes, how much can such rationalizations be stretched? Thus, the literature moved from Haldane's original cost argument to a load argument, and the underdiscussion of the former left students ill-prepared to understand the latter. Key relevant issues were left out, and many unnecessary confusion factors were brought in. That was not intentional, but in my view it is the result of the evolutionists' lack of pursuit to truly teach and truly solve the problem. Let me summarize the literature: the material on Haldane's problem is generally entirely absent, or too little and too cryptic for the intended audience. The textbooks taught it poorly. The ultimate proof is two-fold: (1) Evolutionists claim the problem is solved, but they do not remotely agree on what that solution is. (2) The problem was thought solved, when it wasn't, One last point. Haldane laid at our doorstep the ability to calculate the number of substitutions available for human evolution. Using his result, in ten million years an ape-human species with a 20 year generation time could substitute no more than 1,667 beneficial nucleotides. It's an easy calculation, yet in nearly forty years no evolutionist saw fit to publish such a figure. Is it possible they all thought it uninteresting and irrelevant? We can understand that scientists have debates, and we can understand that any given scientist sometimes says things that are later overturned and withdrawn, that is how science progresses. But in this case it is the widespread, persistent, long-term silence that is so inexplicable. Walter ReMine Standard disclaimer: These thoughts are mine. P.O. Box 19600 Saint Paul, MN 55119 From: us236454 at dawn.mmm.com (Walter Remine) Newsgroups: talk.origins Subject: Haldane's Dilemma Date: 27 Apr 1995 12:33:54 -0500 Message-ID: <3noki2$s7f at dawn.mmm.com> *** Haldane's Dilemma, issues on sexual reproduction *** Andy Peters says sexual reproduction (ie. recombination) can reduce the cost of substitution and thereby speed up evolution. Andy feels this can at least help solve Haldane's Dilemma. It can't, because sex ***slows*** evolution. "Populations would have to be in the millions for recombination [ie. sex] to be important in increasing the spread of favorable mutations." (G.C. Williams) Sex cannot make evolution go faster, unless the population sizes are quite large, at which point the cost of substitution is quite large and Haldane's Dilemma especially bad. Andy mistakenly cited John Maynard Smith as supporting his position. Maynard Smith agrees with what I said above, and discusses it, albeit briefly, in his textbook (pages 240-1). Moreover, researchers now agree that if there is epistasis (fitness interactions among the various mutations), then sex dramatically slows down evolution. This is a very robust result. Once again I emphasize that evolutionary theory (and evolutionary genetics) is a smorgasbord -- the answer that evolutionists give you depends on the question you ask. Ask them a question, say, about how populations get rid of harmful mutations fast enough to avoid error catastrophe -- and they will usually tell you the critical importance of epistasis. Ask them about Haldane's Dilemma, and they will emphasize the importance of sex and small population sizes. But in small populations, sex slows evolution. Moreover, sex and epistasis together make evolution vastly slower, under the widest of circumstances. Evolutionists cannot have it both ways, yet they continue the double-speak. Under our current understanding, sex cannot solve Haldane's Dilemma. Therefore, if you have trouble visualizing Haldane's Dilemma in sexual species, then switch to asexual species, where evolution is faster yet the dilemma is clearer and even more compelling. A historical note here. The origin (and maintenance) of sex is a major problem for evolutionary theory (traditionally one of its trade secrets). Continually evolutionists tell their stories about how an advantage of a minuscule fraction of one percent is sufficient to cause the eventual elimination of all who lack that trait. But sex has a whopping disadvantage, well over a 50 percent disadvantage, it's an immovable tough hurdle. Evolutionists claimed it evolved anyway. For decades they said sex was an advantage because it helped species evolve faster. They felt this advantage was self-evident, and sufficient to overcome that whopping 50+ percent hurdle. Then the population geneticists showed that sex actually slows evolution -- and this immediately caused a 180 degree turn-around. Evolutionists now claimed, with straight-faced conviction, that the advantage of sex is that it made species evolve slowly. This 180 degree turn-around was observed and lamented by evolutionist Graham Bell in his fine encyclopedic book, _The Masterpiece of Nature_. The incident is now a memorial to the incredible flexibility of evolutionary storytelling. (For additional details and documentation, see my book, _The Biotic Message_.) Walter ReMine Standard disclaimer: These thoughts are mine. P.O. Box 19600 Saint Paul, MN 55119 From: us236454 at dawn.mmm.com (Walter Remine) Newsgroups: talk.origins Subject: Haldane's Dilemma Date: 28 Apr 1995 12:17:31 -0500 Message-ID: <3nr7vb$c1n at dawn.mmm.com> *** Haldane's Dilemma and genetic variation *** Andy claimed that the observation of high genetic variation is incompatible with, and therefore disqualifies, Haldane's Dilemma. Major evolutionists have published the same claims. It took several posts before Andy revealed his full line of reasoning, and the truth turns out to be dramatically opposite what he was claiming. But before I show that, let me first document some versions of what he was saying, so you can see the difference between what he said and what he actually meant. >I'm saying that the *very same models* that predict a >limit to the rate of evolution (Haldane's Dilemma) also >predict that there should be very little genetic variation >in natural populations. This is because the same >demographic cost that limits substitution of alleles should >manifest itself when there are large amounts of variation >across the genome within populations. Thus, the large >amount of variation we see shows that the assumptions >which predicted less variation (and, therefore, the >assumptions which drive Haldane's Dilemma) must be wrong. ... >the genetic variation doesn't solve [Haldane's] problem >so much as prove that the problem doesn't exist. ... >The existence of the variation was (and, in fact is) >indeed a dilemma. The question still remains, "how is >that variation maintained without incurring a demographic >cost and wiping out populations left and right." >However, as I've said above, far from "deepening the >problem" of Haldane's Dilemma itself, it erased it: >clearly, the assumptions which led to both the prediction >of low variation and the prediction of low rates of >evolution must be wrong. ... >the concept of genetic load exposed the truly interesting >phenomena behind the concept of Haldane's Dilemma, once >the existence of large amounts of genetic variation in >natural populations showed Haldane's Dilemma to be nonexistent. ... >Which of the numerous assumption(s) are invalid is still >unclear. That's the real current problem pertaining to >this stuff: exactly what is it that we don't understand >about selection that allows this variation to exist (and, by >extension, allows relatively rapid evolution by selection)? >... that particular problem hasn't yet been solved. There is no such link between Haldane's argument and a requirement for low variation. Haldane's argument is compatible with both low and high levels of variation. I can refute Andy's claim several ways. First, let me momentarily assume Andy's claim is true, and then show that it leads to a contradiction. Begin with a situation where there is low genetic variation, in full agreement with Andy's requirement for the validity of Haldane's argument. Haldane's argument then places a valid limit on the maximum plausible rate of evolution, and the population must slowly evolve within those limits. Now give the population a large supply of harmful mutations, say with x-rays, chemicals, or some special genetic technique. The population is worse off than before. But it now has a high level of genetic variation, and according to Andy's thinking this sets the population free to evolve faster. That is the contradiction. In effect, Andy claims the harmful mutations allow evolution to go faster. I say the contrary. The harmful mutations are an extra burden for the population. They slow down evolution because the increased harmful mutation causes an increased level of genetic death, and more of the population's reproductive capacity must be devoted to replacing (or "paying" for) those genetic deaths. Therefore less of the reproduction can go to paying for substitutions. On the other hand, take the same example above, and instead of injecting harmful mutations, inject neutral mutations instead. This likewise creates a measurable "high level of genetic variation" but there is no reason whatever that this should impact Haldane's argument. The neutral mutations do no harm, they do not have to be eliminated, so they do not increase the genetic deaths. They take no bite out of the available reproduction, so the reproduction can go on paying for substitutions at the same rate as before. Thus, I have refuted Andy's claim by showing it leads to contradiction, and by giving two counter-examples. Andy's original claim was confusing, and it didn't become clear until he gave the details of his argument. Now it is clear that when he said "low genetic variation" he actually meant low harmful mutation load, which is a different thing. His argument is about harmful mutations and their drastic consequences on populations. Also he inadvertently created confusion by focusing on how it is "maintained." >The question still remains, "how is that variation >maintained without incurring a demographic cost and >wiping out populations left and right." You don't want to "maintain" a high load of harmful mutation -- you want to *get rid of it.* To speak of "maintaining it" just adds confusion, and it took me a while to decode what he meant. (There are times when evolutionists legitimately seek to maintain high levels of genetic variation, such as by heterozygote advantage or balancing selection, but that is a different issue altogether from harmful mutation, which you just want to get rid of.) My next post will examine the innards of Andy's argument. Walter ReMine Standard disclaimer: These thoughts are mine. P.O. Box 19600 Saint Paul, MN 55119 From: us236454 at dawn.mmm.com (Walter Remine) Newsgroups: talk.origins Subject: Haldane's Dilemma Date: 28 Apr 1995 13:26:42 -0500 Message-ID: <3nrc12$h50 at dawn.mmm.com> *** Haldane's Dilemma and genetic variation -- continued *** Andy Peters claimed that the observation of high genetic variation is incompatible with, and therefore disqualifies, Haldane's Dilemma. My previous post refuted his claim by showing it leads to contradiction, and by giving two counter-examples. This post deals with the specifics of his argument, which are even more revealing. Here is Andy's argument again for the sake of documentation. I'll explain his argument down below. >as a result of the exact same assumptions as those for >substitution load - there should be a limit on the amount >of variation that can be maintained in a population. > >Yet the amounts of variation in natural populations are >far beyond what could be expected under such a limit. >For illustration, imagine a single individual with 100 >mutations across its entire genome. Even if each >mutation by itself would only decrease fitness by 1%, >the individual's fitness is .99^100 =~.36 . Estimates >of the numbers of loci at which real-world organisms >might be expected to have deleterious alleles range >from 1000 to 5000, and a 1% decrease in [fitness] is very >likely. That leads to mean fitnesses of 4 e -5 and >1 e -22. If we apply the same criterion to this result >as Haldane applied to get the substitution rate >limitation - asserting a maximum proportion of selective >deaths which should be allowable to the population every >generation (Haldane used 10%) - we can see real-world >populations should all be extinct if the assumptions >behind Haldane's model hold (Proportion of selective >deaths should be 99.996% for 1000 mutated loci, >99.99999999999999999999% for 5000 mutated loci - the >average individual would have to produce 100000 or 10^22 >offspring to expect only one to survive, and that >doesn't even begin to take into account deaths due to >ecological and accidental occurrences). In other words, >under Haldane's model, *the amount of variation that >exists in the real world, *can't* exist*! > >There's clearly something wrong with the assumptions >leading to the predictions of both [Haldane's] rate >limitation and variation limitation. Andy begins with the standard model of evolutionary genetics. He calls this "Haldane's model" but there is nothing about it that is unique to Haldane's model. It's just the standard model of evolutionary genetics, no epistasis. It is the one model prominently displayed in all evolution books. Then Andy says there are very good empirical grounds to believe the typical organism has from 1000 to 5000 harmful mutations, each causing roughly a 1% decrease in selective value. Then, Andy happens to choose a level of selective death that is the same as Haldane used, ten percent. But there is no connection, he could just as well have arrived at a different figure and justified it separately from Haldane. But it's all a red-herring anyway, because Andy never uses the figure. In fact, even a figure near 100% would still not affect his conclusion. Andy then calculates the harmful mutation load based on the standard model. The result dramatically conflicts with reality. The result indicates that such species would rapidly go extinct, but since they aren't extinct there seems to be a contradiction. Here is Andy's mistake. He pinned the discrepancy on Haldane's argument (just as many evolutionists do) and left it at that, as though Haldane's argument was the thing at fault, and the existence of "high genetic variation" (really high mutation load) is just a research problem yet to be solved. The truth is a bit different. Andy's calculations leave three, and only three, options: (1) Perhaps the data is wrong. Andy employed measurements of data (the number of harmful mutations in each individual, and their selective disadvantage) and those measurements might be somewhat in error. But Andy expressed strong confidence in the data. Also Andy is using his argument as a strongly reliable reason to throw out Haldane's Dilemma, this is additional evidence that Andy feels the data is strongly reliable. So Andy would probably consider option #1 as invalid, and so continue to option #2. (2) The many observed species (and presumably all species) are in error-catastrophe, and are deteriorating genetically, as Andy concluded they are. Andy rejected that possibility because he assumes evolution must be true, and the situation he described is evidence against evolution. (Note: That all species are genetically deteriorating is a possibility acceptable to the worldviews of many creationists.) Also, option #2 means that Haldane's Dilemma remains valid because the model on which it is based remains valid. So, option #2 provides an additional independent line of evidence against evolution. The problems don't just add, they magnify each other. All by itself the mutation load seems too high, and the population close to, if not within, error catastrophe. And all by itself the cost of substitution is too high for macroevolution to be plausible. These problems aggravate each other, because evolution must somehow accomplish these tasks simultaneously, but the costs are paid separately out of the limited supply of reproduction. Thus, making macroevolution especially problematic. Andy may find option #2 unacceptable, and so continue to option #3. (3) The standard model of population genetics is wrong. This is precisely what I have been saying in my book and my posts: It is the one model prominently taught in every evolution book, yet it is seriously inadequate to justify macroevolution. And Andy's argument shows that evolutionary geneticists *know it.* So why are they teaching that model universally, almost exclusively, with none of the appropriate disclaimers, for decades after they knew the discrepancy? It is because the standard model is easier to sell, and population genetics is central to selling evolution. That is the scandal I have been talking about. Phil Nicholls writes: >If reality and the model are wrong, toss the model, not reality. Precisely my point. Why haven't evolutionists tossed the standard model? They've known for decades that it can't justify evolution. Moreover, there are several, independent, simple ways of showing that, and the necessary data and arguments were available decades ago. (See my book, _The Biotic Message_, for details.) Andy garbled the truth the same way evolutionists have for decades. Like this: "Haldane's Dilemma is a non-problem -- the observed high level of genetic variation is an arcane puzzle -- sell the public on the standard model of evolutionary genetics -- evolution is a fact." Instead, Andy should be agreeing with me right now, and shouting -- right along side me -- that the standard model of evolutionary genetics is manifestly inadequate to justify macroevolution. If Andy believes his own argument, that is what he must do. The critical pressure point is not genetic variation, but rather the standard model. There is irony in this. As this situation unfolds we may well see creationists defending, and evolutionists attacking, the standard model displayed in all those textbooks. Walter ReMine Standard disclaimer: These thoughts are mine. P.O. Box 19600 Saint Paul, MN 55119 From: us236454 at dawn.mmm.com (Walter Remine) Newsgroups: talk.origins Subject: Haldane's Dilemma Date: 29 Apr 1995 18:49:25 -0500 Message-ID: <3nuja5$bpf at dawn.mmm.com> I made a clear claim: "Haldane laid at our doorstep the ability to calculate the number of substitutions available for human evolution. .... It's an easy calculation, yet in nearly forty years no evolutionist saw fit to publish such a figure." Tim Ikeda claimed to have a specific counter-example. I called his bluff, and he lost. He shrugs it off, saying I am "technically correct." But my claim is completely correct. Lewontin (and all evolutionists) gave no figure, not even to an accuracy of one significant digit. So Tim attempts to change the issue. Evolutionists completely ignored the simple, obvious, readily understandable, highly relevant issue of total substitutions, and instead spoke obscurely in terms of substitution rate. Let me put this in greater focus. Of the few evolution books that mention Haldane's problem, less than a third state Haldane's substitution rate. Of those, nearly all state it cryptically, by speaking of "gene" or "allele" substitutions, as in "one gene substitution per 300 generations" or some variant of that wording. That is how Lewontin did it (page 224). That practice is nearly universal, and it effectively removed the magnitude of Haldane's Dilemma from understanding by students and the public. A figure like "1,667 nucleotides" draws immediate attention and consternation. (It is roughly 50 millionths of one percent of the human genome.) Then even the ordinary person sees the importance. But "one gene substitution per 300 generations" is effectively opaque, even to most evolutionary specialists. The problem is effectively concealed. One renowned evolutionary geneticist/professor chose to argue with me recently. He insisted the substitutions are new 'genes' not 'nucleotides,' and therefore the number of beneficial nucleotides available for human evolution is vastly higher than 1,667. It took a while to convince him of his error, he had been so habitually used to thinking of it as "gene substitutions." In fact, I cannot find even one evolutionary textbook that disabuses students of that fallacy about Haldane's substitution rate. Haldane's dilemma is most severe for species with long generation times and low reproduction, such as humans and simians -- species of key interest. No evolutionary specialist could have overlooked that or found it irrelevant. Yet for nearly forty years they never once took the obvious step of publishing the number of beneficial substitutions available for human evolution. There are no excuses for that, though the talk.origins crowd surely tries. Tim Ikeda says Lewontin "understood the implications" of Haldane's argument. Fine, then why didn't evolutionists publish the implications completely and clearly -- and truly share them with the world? Why did they universally fail to do that? Chris Colby flatly asserts that a limit of 1,667 nucleotides would not be a problem. Fine, then why didn't evolutionists publish that? But no evolutionist published it -- not one. That is now history, and it cannot be changed. We are faced with a puzzle -- the evolutionists' widespread, persistent, long-term silence on Haldane's dilemma. Evolutionary experts did not reveal the magnitude of the problem. Instead, at every turn they downplayed it and discussed it cryptically. Their silence allowed key aspects of the problem to fall into obscurity. Their silence also allowed the unchallenged decades-long promulgation of over a dozen phony "solutions." Their silence also allowed their internal disagreement to go unnoticed -- so that virtually all evolutionists agreed the problem was solved (and few recognized they didn't remotely agree on what the solution was). Lastly, most evolutionary textbooks are silent on both the problem and its alleged solutions. Again I emphasize, it is the evolutionists peculiar pattern of silence that is so damning. I encourage each of you to explain that for the public -- for those many ordinary work-a-day folks who want to understand how science operates. Tell them what Tim, Chris, and others said here. And watch the reaction. Walter ReMine Standard disclaimer: These thoughts are mine. P.O. Box 19600 Saint Paul, MN 55119 From: us236454 at dawn.mmm.com (Walter Remine) Newsgroups: talk.origins Subject: Haldane's Dilemma Date: 2 May 1995 01:18:34 -0500 Message-ID: <3o4irq$8oi at dawn.mmm.com> *** Haldane's Dilemma and sexual reproduction *** Andy Peters writes: > I'm conceding the point that, under most reasonable conditions, > recombination can't erase - or even dramatically reduce - > the cost of substitution. Andy concedes that sexual reproduction (ie. recombination) cannot reduce the cost of substitution or solve Haldane's dilemma. I emphasize that his previous misunderstanding was not his fault. In fact, such misunderstandings about recombination are rather common, especially in discussions of Haldane's dilemma. Yet Andy has had extensive training in this field, and it reinforces my point that the textbooks do not teach Haldane's problem adequately (or the problems of the maintenance of sex adequately). Andy asked for more references documenting the evolutionary difficulties with sex, recombination, and epistasis. I'll offer a few here, (there's more in my book.) We've already accomplished what we need concerning sex in this thread, so I won't be pursuing it further: "[T]he occasional production of extremely fit genotypes in the sexual population will have no permanent significance, as long as fitness depends at all on heterosis and epistasis. Given any plausible level of heterosis and complex interactions among loci [ie. epistasis], almost all of a sexual population will have suboptimal genotypes, no matter how long selection continues. All that this means is what everybody knows, that sexual reproduction generates recombinational load, but I suggest that this mundane fact may be the primary significance of sexuality in evolution. It greatly retards the final stages of multilocus adaptive change and severely limits the attainable precision of adaptation." (G.C. Williams, 1975, p 150) "To save the situation then, we must perform a complete volte-face: just as it was self-evident to Weismann, Fisher and Muller that a faster rate of evolution would benefit a population, so we must now contrive to believe in the self-evident desirability of evolving slowly." (Graham Bell, 1982, p 100) "[I]t has proven difficult to produce explicit models that show that mixis [ie. the genetic mixing caused by sexual reproduction] accelerates adaptive evolution." (Michod and Levin, 1988, p 2) Epistasis not only slows down evolution in sexual species, it also makes the origin and maintenance of sex much harder to explain: "[S]election will always favour a reduction in the rate of recombination, provided that there is any degree of epistatic interaction between loci, .... there is remarkable unanimity among theorists that the ineluctable suppression of recombination is not only a very general but also a very robust result." (Graham Bell, 1982, p 407) Walter ReMine Standard disclaimer: These thoughts are mine. P.O. Box 19600 Saint Paul, MN 55119 From: us236454 at dawn.mmm.com (Walter Remine) Newsgroups: talk.origins Subject: Haldane's Dilemma Date: 2 May 1995 01:38:00 -0500 Message-ID: <3o4k08$91d at dawn.mmm.com> Andy Peters writes: >ReMine keeps implying that the selective use of >assumptions in model building is a bad thing. >It's not. .... >This is, of course, the crux of the biscuit. ReMine seems to advocate an >all-or-nothing approach to this, however: the "standard model of >population genetics" is "manifestly inadequate," so it should be thrown >out altogether. The fact, of course, is that the standard model of >selection does predict very precisely lots of population phenomena. It's >clear that selection works in natural populations, so throwing away the >most general model of selection entirely would be throwing the baby out >with the bath water (I've always wanted to use that expression). Tweaking >various assumptions of the model (and thereby generating ReMine's >"Smorgasbord") is the *appropriate* response to such a situation - though >ReMine insinuates that it's dishonest. Andy says that selective use of assumptions in model building is not a bad thing. I say evolutionary genetics texts fail that on four counts: 1) The textbooks must *say* when they employ the selective use of assumptions, and why. 2) The textbooks must *identify* the assumptions in each model. 3) The textbooks must delineate the various models, their strengths *and weaknesses,* and their role (or not) in evolutionary theory. 4) The textbooks must not give contradictory models and pass them off as one unified solution to problems. I say evolutionary genetics textbooks routinely fail on these four points, (especially #3 and #4). The following examples are macroscopic problems that apply broadly, they are not narrow ecological issues. 1) Error catastrophe (also mutation load) 2) The origin and maintenance of sexual reproduction (recombination) 3) Haldane's Dilemma -- puts limits on the rate of macroevolution 4) Kimura's problem with the limitation on the rate of beneficial mutation, which also puts a limits on the rate of macroevolution. It is a contradiction to embrace epistasis to solve one problem (such as error catastrophe), and then renounce epistasis to solve another problem (such as sexual reproduction). It is a contradiction to embrace "small population sizes" as a solution to Haldane's Dilemma, and also embrace "large population sizes" as a solution to Kimura's problem. It is a contradiction to embrace "sex and small population sizes" as a solution to Haldane's dilemma, when those two together make evolution slower, especially when combined with epistasis. The textbooks routinely employ those contradictions in a smorgasbord fashion, where they fail to identify the contradictions. The answer they give you depends on the question you ask. The problem is not the selective use of models, per se. Rather, the problem is the hidden contradictions, the absence of disclaimers, and the absence of disclosure. The textbooks present evolutionary genetics as a uniform, coherent, testable theory. It is not. Andy claims I am "throwing the baby out with the bath water." I am not. I have consistently said things are *absent* from evolutionary textbooks that ought not be absent. I'm not throwing out anything. I say the textbooks should *bring in* some things -- things they left out for decades. Walter ReMine Standard disclaimer: These thoughts are mine. P.O. Box 19600 Saint Paul, MN 55119 From: us236454 at dawn.mmm.com (Walter Remine) Newsgroups: talk.origins Subject: Haldane's Dilemma Date: 4 May 1995 19:40:57 -0500 Message-ID: <3obs6p$gu at dawn.mmm.com> *** Haldane's Dilemma and genetic variation *** This is my response (part 1 of 3) to Andy Peter's latest post on genetic variation. Andy asserts that the observed high mutation load invalidates Haldane's Dilemma. Andy's post often re-states his assertion (in slightly different words), over and over again, without justifying it or explaining it. It is a repetitious and unilluminating post in that sense. Also, it departs from Andy's normally more patient style and engages in microscopic sniping, taking every opportunity to be contrary and to merely re-state his assertion yet another time, as if that will drum it into us better. That is unfortunate for several reasons. While I can sort Andy's post, I fear many readers here cannot and are left more confused than enlightened. Moreover, evolution textbooks don't remotely teach Haldane's Dilemma adequately -- even evolutionary specialists are confused -- and I've documented that point repeatedly. Even Andy now offers the self-incriminating excuse, "I don't give a flying fajita whether or not Haldane's Dilemma is dealt with to [the student's] satisfaction in textbooks and popular literature." To make matters worse, evolutionists frequently accuse their opponents of ignorance. (We've seen that often in this thread.) But they have no high ground for any such charge. Evolutionists themselves are responsible for the rampant ignorance. For decades they literally sold ignorance in their textbooks. Finally, I cannot find Andy's refutation of Haldane's dilemma detailed anywhere in the literature, and have encouraged him to cite it if he knows examples. Since he has not done that, it appears Andy's refutation is being given here for the first time anywhere, so it is especially important that he teach it clearly and patiently, as it is the only example of its kind. Altogether, Andy has no grounds for sniping, for shortness, or for charging me with ignorance. ****** Let me now eliminate some confusion factors. Andy claims the observed "high level of genetic variation" invalidates Haldane's Dilemma. To refute his claim, I provided two counter-examples. In one counter-example, the high level of genetic variation is due to neutral mutations, which do not affect the beneficial substitution rate or Haldane's argument. Andy accepts my point as "valid," thus we both agree that some kinds of genetic variation (variations that have neutral effect) do not invalidate Haldane's Dilemma. Then Andy appears to contradict himself: >Under Haldane's model, however, genetic variation and >mutation load are in fact exactly the same thing. No, Haldane's model does not require genetic variation and mutation load to be the same thing. I just gave an counter-example and Andy already acknowledged it as "valid." Perhaps we are arguing over the term "genetic variation," but I continue to emphasize that not all genetic variation is the same. Harmful, neutral, and beneficial mutations create different types of "variation" that are all rather different in their effects and in their problems. Many evolutionary arguments achieve their illusory power by using the ambiguous term "variation" and then shifting back and forth between the various meanings. To end such mischief we must identify the type of variation and use that identification consistently. If the "variation" is actually "harmful mutations" or "mutational load", then use these latter terms instead, as they are less ambiguous. I will make that replacement when I quote Andy, as that seems to be his intended meaning throughout. That replacement makes Andy's mistakes more visible. For example: >The [load of harmful mutations] doesn't *allow* the >population to evolve faster (well, actually it does ... That is nonsense. Harmful mutations are a burden to be eliminated. They do not allow a population to evolve faster in any sense whatever. They slow evolution. Another example is when Andy discusses "the problem of maintaining high genetic variation." Unfortunately, his confusing wording is *identical* to legitimate statements about an altogether different problem involving an altogether different type of genetic variation. (eg. Mechanisms such as the neutral theory, heterozygote advantage, or balancing selection are legitimately used to explain the maintenance of special types of genetic variation.) So I thought that's what Andy meant. But I later realized Andy meant something different -- "the problem of maintaining [high levels of harmful mutations]." Even that was confusing, because there is no such maintenance problem. The problem is not "maintaining" harmful mutations, rather the problem is *getting rid of them* "without wiping out populations left and right." Or to put it differently, the problem is "maintaining the population" not the harmful mutations. I recommend clearer wording, such as "the problem of eliminating harmful mutation." Again the nexus of confusion is the term "genetic variation" and its many ambiguous meanings. I'm not picking on Andy here. The terms he used are commonplace in the literature. (continued in part 2) Walter ReMine Standard disclaimer: These thoughts are mine. P.O. Box 19600 Saint Paul, MN 55119 From: us236454 at dawn.mmm.com (Walter Remine) Newsgroups: talk.origins Subject: Haldane's Dilemma Date: 4 May 1995 19:56:34 -0500 Message-ID: <3obt42$n9 at dawn.mmm.com> *** Haldane's Dilemma and genetic variation *** (This is the continuation (part 2 of 3) of my response to Andy Peter's latest post on genetic variation.) Let me call this Andy's assertion: The observed high mutation load invalidates Haldane's Dilemma. An earlier post of mine refuted Andy's assertion. My argument momentarily assumed Andy's assertion is true, and then shows it leads to a contradiction. Andy responded to my argument, and I now show where his response errs. I had Andy imagine a population (any kind he wishes) that meets *his* criteria for the validity of Haldane's limitation on the rate of evolution. According to Andy this would require low genetic variation -- low mutation load. Thus my argument begins by accepting Andy's assertion on Andy's terms. What is there for him to complain about? But Andy bails out of my argument at that point, apparently because he cannot even conceive of any population being subject to Haldane's limitation under any circumstances whatever. He bails out by inventing yet another reason for rejecting Haldane's limitation. Here it is in action: >[J]ust because our hypothetical population has low >genetic variation, ***that doesn't mean that the >selective mechanisms that reduce the cost of >selection don't exist.*** [wjr: my emphasis] In other words, Andy spontaneously invents unspecified mechanisms for removing Haldane's limitation. (He lamely calls them "selective mechanisms that reduce the cost," but that is just an empty name.) He never identifies what those mechanisms are, but they are central to his argument, for it is no longer "high genetic variation" that overturns Haldane's limitation, rather it is now Andy's unknown mechanisms. In that way, Andy dodges my argument an avoids having his position revealed as a self-contradiction. He then merely repeats his assertion several more times. In summary, Andy asserts that high mutation load "shows Haldane's assumptions to be false." I just disproved that. You start by giving Andy everything he could want (including Haldane's assumptions and Andy's requirement for low variation), then Andy bails out and invents another reason to reject Haldane's conclusion. For sake of documentation, here is the entire quote from Andy's post, starting with my original argument, alternating with his responses: > First, let me momentarily assume Andy's claim is true, and then show that > it leads to a contradiction. Begin with a situation where there is low > genetic variation, in full agreement with Andy's requirement for the > validity of Haldane's argument. Haldane's argument then places a valid > limit on the maximum plausible rate of evolution, and the population must > slowly evolve within those limits. Not necessarily. The existence, or lack thereof, of genetic variation in any given population doesn't have any direct effect on the cost of substitution in that population. In other words, just because our hypothetical population has low genetic variation, that doesn't mean that the selective mechanisms that reduce the cost of selection don't exist. All the existence of large amounts of variation in natural populations does is to *demonstrate* that the cost of substitution doesn't exist - it doesn't actively reduce the cost itself. > Now give the population a large supply > of harmful mutations, say with x-rays, chemicals, or some special genetic > technique. The population is worse off than before. But it now has a > high level of genetic variation, and according to Andy's thinking this > sets the population free to evolve faster. Nope. The variation doesn't *allow* the population to evolve faster (well, actually it does, .... > That is the contradiction. In > effect, Andy claims the harmful mutations allow evolution to go faster. .... (continued in part 3) Walter ReMine Standard disclaimer: These thoughts are mine. P.O. Box 19600 Saint Paul, MN 55119 From: us236454 at dawn.mmm.com (Walter Remine) Newsgroups: talk.origins Subject: Haldane's Dilemma Date: 4 May 1995 20:02:31 -0500 Message-ID: <3obtf7$se at dawn.mmm.com> *** Haldane's Dilemma and genetic variation *** This is the continuation (part 3 of 3) of my response to Andy Peter's latest post on genetic variation. Here are three models of evolutionary genetics: 1) The model that supports Haldane's dilemma. 2) The model that supports the concept of mutational load. 3) The standard model, the one in all the books, the one used to sell evolution to the public. I say all three are the same model. In fact, merely use "standard model" whenever Andy uses "Haldane's model," and his post still reads as true. Andy agrees that 1 and 2 are the same model, and most of his post labors to establish that. (I was already convinced a long time ago.) But Andy didn't establish that models 1 and 3 are different, or that 2 and 3 are different. ***He didn't even try.*** Instead he takes the problem of mutational load, and blames it on "Haldane's model", citing its huge failure to avoid error catastrophe and extinction. But he tars the standard model with the same brush, because they're the same model. So I cheer him forward! Andy is making my case for me. If Andy believes his own argument, the standard model of evolutionary genetics massively contradicts macroevolution. And one of them must go. There is much that Andy and I agree on here. The difference is: (1) Andy wants to scapegoat the problems onto "Haldane's model." (2) Andy wants to continue selling the standard model of genetic evolution without revealing the massive contradiction that Andy himself identified. ****** There was an option that Andy arbitrarily rejected, the option that many species (presumably all species) actually are in error-catastrophe and are deteriorating genetically. Andy explainss: >I rejected [that] possibility because it goes against the evidence. >Natural selection happens, whether or not all life shares a common >ancestor. It's been demonstrated countless times. It's not an >"assumption" on my part - it's a conclusion, based on evidence. That's a classic move. When your argument gets in trouble just say, "Natural selection happens!" and we're supposed to all fall over. When it comes to evolutionary illusions, there is none bigger or more intricate than natural selection. Let me make it clear. If every species on the planet were genetically deteriorating, then that would be totally compatible with natural selection. The moths, the finches, and the laboratory flies could take two steps up the hill, and slide three steps back down the hill. The species could "survival of the fittest" all the way down to extinction. Let me broaden that. Take Haldane's argument, mutation load, the standard model of evolutionary genetics, and natural selection -- They are all totally compatible with genetic deterioration, even on a global scale. None of them predicts against it. Yet Andy rejects genetic deterioration. He claims (mistakenly) that Haldane's model makes a "prediction" against genetic deterioration. In truth, he gets that notion from outside -- from somewhere else. He gets it from his worldview. Because global genetic deterioration is incompatible with macroevolution. ****** Andy writes: >Under Haldane's assumptions, however, almost every >population in existence should have been wiped out >centuries ago, given the amount of variation we've >seen in them. Let me clarify Andy's statement: 1) "Haldane's assumptions" = "the standard model of evolutionary genetics" 2) You can't make measurements on organisms **today** and conclude they "should have been wiped out centuries ago." That's hyperbolic (as well as impossible). Change it to "should be extinct sometime in the near future." 3) Change "variation" to "harmful mutation", which is what Andy means. Then the statement is accurate and true to Andy's argument. Like this: >Under [the standard model of evolutionary genetics], >however, almost every population in existence [should >be extinct sometime in the near future], given the amount >of [harmful mutation] we've seen in them. That statement ought to be in the textbooks. It is simple and clear, anyone can interpret its meaning for themselves. So why does Andy garble it beyond the public's understanding? Why did it take so long for Andy to fully reveal his key argument? Why did it take so many posts to strip away layer after layer of misdirection and confusion? Andy totally believes his own argument, so why didn't he just shout it in plain text? Throughout my posts I have shown that evolutionists persistently avoid key issues inconvenient to them. On such matters they speak cryptically, if at all. Andy has just given a demonstration of it. Walter ReMine Standard disclaimer: These thoughts are mine. P.O. Box 19600 Saint Paul, MN 55119 From: us236454 at dawn.mmm.com (Walter Remine) Newsgroups: talk.origins Subject: ReMine statistics Date: 6 May 1995 16:41:21 -0500 Message-ID: <3ogqe1$g00 at dawn.mmm.com> In 1957, the renowned evolutionary geneticist J.B.S. Haldane published a problem that eventually bore his name "Haldane's Dilemma." He saw that reproductive capacity is limited, and this puts a limit on the speed of evolution. He showed that in species with low reproductive capacity (such as higher vertebrates, mammals and man), the long-term average beneficial substitution rate cannot be faster than one substitution per 300 generations. These substitutions are single nucleotides. I wrote (in my April 24 post): "Haldane laid at our doorstep the ability to calculate the number of substitutions available for human evolution. Using his result, in ten million years an ape-human species with a 20 year generation time could substitute no more than 1,667 beneficial nucleotides. It's an easy calculation, yet in nearly forty years no evolutionist saw fit to publish such a figure. Is it possible they all thought it uninteresting and irrelevant? We can understand that scientists have debates, and we can understand that any given scientist sometimes says things that are later overturned and withdrawn, that is how science progresses. But in this case it is the widespread, persistent, long-term silence that is so inexplicable." 1,667 beneficial nucleotide substitutions amounts to about 50 millionths of one percent of the human genome. And that is *before* making several deductions, such as the 90% reduction due to the 90% of the time spent in stasis, according to Gould. Is a maximum of 1,667 beneficial nucleotides enough to make a sapien out of a simian? Chris Colby responds: >What ReMine hasn't done is discuss >his assertion that 1667 adaptive >[nucleotide] substitutions is too few >to explain our divergence from the chimp/ >human ancestor. Even if we accept his number >*****(and I'll admit that it is within the ballpark),***** [wjr: my emphasis above] >there is no indication that it is inadequate. >.... a limit of 1600 nucleotide changes may >not be a problem. Chris should defend that view in a journal. He will be the first evolutionist *ever* to even try. ****** Chris Colby repeatedly misrepresents my posts to the thread "Haldane's Dilemma." For example, he writes: >[ReMine] claims that Haldane's dilemma is absent from evolutionary texts. In my defense, I will merely quote portions of my original post: <> Let me summarize the literature in several stages. First, there are the books aimed for wide public consumption. These are generally books that sell evolution to the public, as a precise and testable science. To do that, evolutionists inevitably reach for population genetics as the convincer. The simple, easy-to-sell genetic model is the one sold -- truncation selection and soft selection (the alleged solutions to Haldane's Dilemma are nowhere in sight. All non-college (and most college-level) evolution books and articles follow that practice. I can bury you with examples, but will here offer only Kitcher's anti-creation book, _Abusing Science_. He sells evolution via the simple genetic model, and adds, "Mathematical population genetics tells us *precisely* how the forces of immigration, emigration, mutation, and natural selection produce evolutionary change." (Contrast that with my opponents' admissions in this thread recently.) All those books are wrong for doing that. If truncation and/or soft selection are essential to solving Haldane's Dilemma, *then they should be taught* (and defended) and any genetic model lacking them should have appropriate disclaimers. No excuses. This matter gets especially bad when you get to textbooks *specializing* in evolutionary genetics. Any of these -- even mere introductory texts -- should have the solutions to Haldane's dilemma indelibly connected to the evolutionary model (not necessarily with all the math), and it should be painfully clear to any student that these are not optional. On the other hand, if a problem of such broad importance as Haldane's dilemma were unsolved, then it would be inexcusable to leave it out of the books. Either the solution, or the problem, or both, must be clearly displayed in the book. But that is not what we find. John Maynard Smith is a world authority on Haldane's Dilemma and cost issues, and in the 1960s proposed truncation selection as a solution. The preface for his recent textbook, _Evolutionary Genetics_, says the book is intended for advanced undergraduates, is hopefully useful to graduates, and that "proper training in science requires that undergraduates are confronted by the problems of contemporary science." The book discusses the usual standard version of evolutionary genetics. But it doesn't discuss the following: Haldane's problem or any version of it, cost of substitution, substitution load, truncation selection, or soft selection. Everything about the problem and its solutions is absent. This is commonplace in textbooks on evolutionary genetics. Here is an additional brief sampling of modern books, where everything is absent. Molecular Evolutionary Genetics, Nei, 1987 Genetics of Populations, Hedrick, 1983 (***Mentions soft selection, but doesn't tie it in as essential.) Population Genetics and Molecular Evolution, edited by Ohta and Aoki, 1985 (***Mentions truncation selection in one paragraph, but doesn't define it or tie it in as essential.) Rates of Evolution, edited by Campbell and Day, 1987 (About half this book is evolutionary genetics, but the stated topic of the book makes the omission of both the Haldane problem and solution especially notable.) Genetics, Paleontology, and Macroevolution, Levinton, 1988 (***Mentions truncation in one paragraph, but doesn't tie it in as essential.) Theoretical Population Genetics, Gale, 1990 Population and Evolutionary Genetics: A Primer, Ayala, 1982 Now we move to books that at least mention Haldane's problem. Exceedingly few of these refer to it as "Haldane's dilemma". (Most evolutionists will get angry if you call it a "dilemma", as you have already seen in our ongoing discussion here.) The books dramatically downplay it, insisting it's not a problem. So they often give short-shrift to the real mechanics of the issue, with little more than a few sentences, perhaps a paragraph. Even then they generally fail to tie-in truncation selection and soft selection as essential to the solution and essential to the evolutionary model. (Futuyma's text is an example here.) <> My original post has some additional discussion. Then I concluded: "Let me summarize the literature: the material on Haldane's problem is generally entirely absent, or too little and too cryptic for the intended audience. The textbooks taught it poorly. The ultimate proof is two-fold: (1) Evolutionists claim the problem is solved, but they do not remotely agree on what that solution is. (2) The problem was thought solved, when it wasn't." Walter ReMine Standard disclaimer: These thoughts are mine. P.O. Box 19600 Saint Paul, MN 55119 From: us236454 at dawn.mmm.com (Walter Remine) Newsgroups: talk.origins Subject: Haldane's Dilemma Date: 6 May 1995 18:24:45 -0500 Message-ID: <3oh0ft$h2f at dawn.mmm.com> Tim Ikeda writes: >Perhaps one way to examine whether it is inevitable that genetic >deterioration eventually leads to extinction would be to look at >organisms with rapid doubling times, relatively high mutation >rates, or ones which seem to have been around for a very long >time. Viruses and bacteria could be useful subjects in the >first two instances, and perhaps, horseshoe crabs in the latter. >To a first approximation, could we suppose that bacteria and >viruses would approach "genetic oblivion" at a higher rate than, >say, birds? The coding and efficiency of bacterial chromosomes >seem tighter than that found in vertebrates. Genetic deterioration depends on the genome size, particularly that portion that is non-inert. (Leaving ploidy out of the discussion for the moment.) Organisms with large genomes will tend to reach error catastrophe sooner. Also organisms such as mammals have many genome duplications that occur in the interval between egg fertilization and the fertilizable eggs of the next generation. (The same with sperm.) Each of these duplications can introduce errors. The process of recombination (ie. sexual reproduction) adds additional chances for error. These tend to increase the per generation mutation rate in multicellular organisms. Also, species with low reproductive capacity (such as mammals) cannot fend off error catastrophe for as long as bacteria, which have very high reproductive capacity. Reproductive capacity is a critical factor. Using the standard model of evolutionary genetics (ie. no epistasis), we can make a simple calculation with data already available to Motoo Kimura and others in the 70s. Assume 97 percent of the human genome is totally inert, and mutation to this portion of the genome does absolutely no harm in this generation or any generation to come. This assumption dramatically favors the standard model, as you will see, because it dramatically reduces the number of harmful mutations experienced by the organism. Then only 3% of the genome is available to suffer harmful mutation. Using the genome size and the mutation rates (data that Kimura and others regularly used in their discussions, and which are still consistent with today's figures) the following can be shown. To avoid error catastrophe, human females would have to conceive more than 16.3 children. That is the cost of harmful mutation. And that is before adding in the additional costs of substitution, segregation, and random death, which place additional demands on reproductive capacity. (See my book for details.) This argument, and Andy Peter's mutation load argument, both address the problem of harmful mutation. And the results point in the same direction. (1) Either the data is wrong (which is getting progressively more unlikely). (2) Or, many organisms are in error catastrophe. (3) Or, the standard model is wrong. The subject of error catastrophe is interesting. But it gets us too far afield from our subject ... which is Haldane's Dilemma. So I won't be addressing it further in this thread. Let me just point out again, that several independent lines of evidence indicate that the standard model of evolutionary genetics contradicts macroevolution. And everything necessary to show this ... was available decades ago. Walter ReMine Standard disclaimer: These thoughts are mine. P.O. Box 19600 Saint Paul, MN 55119 --Kurt Vonnegut from "Galapagos" From: us236454 at dawn.mmm.com (Walter Remine) Newsgroups: talk.origins Subject: Haldane's Dilemma Date: 8 May 1995 14:06:00 -0500 Message-ID: <3olq2o$k0j at dawn.mmm.com> Andy Peter's latest post has little new from his previous efforts, and much confusing thrashing about. Andy continues to flog the standard model of evolutionary genetics (a.k.a. "Haldane's model") -- with even greater vigor now than before. And I continue to cheer him on. The more he flogs, the more ridiculous his position becomes. I could say several things about the details of his mutation load argument ... but will restrain myself. He is self-destructing quite enough already. ****** There is a cost of substitution, it is unavoidable, it must be paid from a species' limited reproductive capacity, and this places a limit on the rate of substitution. The discovery of "high levels of genetic variation" *aggravates* that problem, no matter what evolutionary genetic model you choose. Higher levels of harmful mutation, or heterosis, or balancing selection, or random death, all take higher bites out of the available reproduction, leaving less reproduction to pay the costs of substitution -- thus slowing evolution. Start with *any* of the evolutionary models, and higher levels of harmful mutation (or higher levels of mutation load) make the problem worse. Andy is wrong to even hint otherwise. The observed high levels of harmful mutation doesn't invalidate the problem, as Andy suggested. Rather, it invalidates "solutions." For example, Andy's mutation load argument massively invalidates the standard model of evolutionary genetics. And the problems don't go away. The costs still remain no matter what genetic/selection model you choose. The different models impact the analysis in different ways. But the problems must still be solved, not just brushed aside, as Andy attempts to do. I have before seen evolutionists *claim* that genetic variation invalidates Haldane's conclusions. (eg. Mayr, 1982, p 594) But Andy offered the first example of actually filling in that argument. I recognize Andy's argument as new (and have asked him for references if it is not). I also recognize that Andy's interpretation is the ***opposite*** of G.C. Williams (evolutionist and author of several well received books). Though Williams uses the exact same example that Andy used. "There are perhaps 100,000 loci in a mammalian genome, and we might expect at least 10,000 of them to be polymorphic, ... Perhaps an average individual has a less than ideal genotype at 10% of these loci, or 1000 in all. In a comparable example, Wallace (1989) gives an average individual a suboptimal genotype at 5000 loci. If the average fitness loss per locus is 1% at 1000 loci, the average individual will only be 0.99^1000, or about 10e-5 of the fitness of the best possible genotype. The variance in fitness would be such that almost no individual would be of even 1% of the maximum. Since maximum fitness is no assurance of survival, the population envisioned would have an utterly inadequate gene pool for survival beyond the current generation." (G.C. Williams, 1992, p 145) Williams' argument is identical to Andy's argument, right down to the numbers. But Williams (correctly) presents it the opposite of Andy. Williams' immediately preceding sentence is: "A bit of arithmetic shows the ***severity*** of Haldane's dilemma." (my emphasis) Williams doesn't try to cover it up, or brush it aside, or misinterpret it, as Andy does. ****** Andy cites one of Haldane's papers on variation, dated 1937, a full 20 years before Haldane's paper on the limitations to substitution rate. They are separate papers, dealing with separate issues, in separate parts of Haldane's career. Yet Andy reaches into that (irrelevant?) paper and pulls out an (irrelevant?) quote: "In a species in equilibrium, variation is ***mainly*** due to two causes. ... [heterosis; and mutation- selection balance]." [my emphasis] Andy concludes: >It's clear: under Haldane's assumptions, variation = mutation or segregation load. Wrong. Even in that old paper it is clear that Haldane leaves room for other types of variation, such as neutral mutations. That example constitutes the *entirety* of Andy's attempt to detach the standard model away from Haldane's Dilemma. Andy is wandering around in history, trying to pin the problem on Haldane ... and bury it with him. I say it again -- the cost of substitution and the cost of harmful mutation are each serious problems for the standard model. (Even Andy's mutation load argument shows that.) The existence of neutral variation neither helps, nor harms, nor invalidates, the problems, and Andy has already acknowledged this point as "valid." (his word) ****** >The standard model (which is, indeed, largely Haldane's) does not >"massively contradict" macroevolution. In its explanation of allele >substitution, for instance, it describes the spread of an allele through a >population. ReMine's position is, "No, it doesn't! The assumptions of >the model are wrong - Andy's shown that himself." By Andy's own mutation load argument, the standard model massively contradicts macroevolution. So Andy changes the subject. He wants to talk about "the spread of an allele", never mind that the population "should already be extinct"!!! (His words.) He blames the problems on Haldane, and he wants to keep selling the standard model to the public, without revealing the contradictions. ****** >The fact is, however, that *the very same math* describes >precisely the spread of an allele through the population >under soft selection - it's merely the assumption about the >way selection acts on the population that changes, and that >doesn't change the genetic dynamics. Andy makes an interesting point: soft selection uses "the very same math" as the standard model. This is because soft selection takes the standard model and merely emphasizes the role of frequency-dependent and density-dependent selection. That provides nothing to eliminate harmful mutations more efficiently, and is just as prone to error catastrophe. Thus Andy's mutation load argument is a serious problem for soft selection too. Combine that with the fact that soft selection doesn't even solve Haldane's dilemma, the cost of substitution. (As pointed out by G.C. Williams, 1992, p 143-148) ****** >I'm not sure about how well the standard model describes >the spread of an allele under truncation selection, >but it's certainly at least a fair approximation. I'd challenge that point. Truncation selection is under-discussed and 'selectively' discussed. It's part of the evolutionary smorgasbord. Evolutionists discuss truncation rarely, and only then to "solve" cost issues, like Haldane's dilemma. Then they ignore it thereafter. They pick its effects that they like. And they ignore its effects that they don't like. (Truncation selection does have bad effects for evolutionists, but I've never seen them discuss those.) It's not enough to say "truncation selection!" and expect us to fall over. If evolutionists are serious about it as a solution, then they must hold it high, defend it, and pursue it as a major feature of evolution. ... They haven't done that. ****** Andy injects yet another "model" into our discussion. He calls it the "general model." He praises it glowingly for its precision and science. I call it a structureless smorgasbord of conflicting and contradictory mechanisms, instantly ready to be shaped by the evolutionists' whims of the moment. >The model isn't perfect - such is the burden of science. It remains, >however, the most successful general model to explain the phenomena of >population genetics: because it still offers a good to excellent >approximation of real world events; and because it's still unclear to what >extent soft selection, truncation selection, and other selective >mechanisms act in nature, the general model is the best model. Indeed, if >it turns out (as it very probably will) that these mechanisms are of >varying importance under different conditions, the general model is still >the best model. > >That's the way science works. I disagree. That is not how Science works. Science does not work by obscuring the problems, hiding the contradictions, and misrepresenting the models -- which is what evolutionary geneticists have done for decades. ****** I previously pointed out to Andy that his use of the term "genetic variation" is ambiguous and confusing about key issues -- that it serves to conceal the serious problem of harmful mutation. Nonetheless, Andy's post continues using his ambiguous term -- now more resolutely than ever, without even once clarifying it. So my concern now is for those many ordinary folks who struggle to understand the issues. Though Andy doesn't intend it, his ambiguous language places hurdles in their path. Evolutionists have no right to accuse anyone of ignorance -- until they take seriously their obligation to stop selling it. Perhaps evolutionists are just lazy about communication? Not so! They can be emphatically clear, when they really want to. They are loud and unequivocal ... when attacking any fleck of creationism in the schools. So why are they persistently cryptic about the deeper problems with evolution? Walter ReMine Standard disclaimer: These thoughts are mine. P.O. Box 19600 Saint Paul, MN 55119 From: us236454 at dawn.mmm.com (Walter Remine) Newsgroups: talk.origins Subject: Haldane's Dilemma Date: 8 May 1995 19:18:23 -0500 Message-ID: <3omccf$507 at dawn.mmm.com> Chris Colby writes: >[I]f Williams is an evolutionary biologist and doesn't >cover up [Haldane's dilemma]; how can you claim evolutionary >biologists have covered it up for decades? I claim it, because it's true. For decades, the problem was pronounced solved. The mechanics of the problem were taught cryptically, and rarely, and fell into obscurity. Well over a dozen phony "solutions" lasted in the technical literature for decades, with little effort to remove them, even though many of the solutions were silly. Even the major solutions (soft selection and truncation selection) were mentioned fleetingly in the textbooks and were not merged into a coherent evolutionary model. That is my claim, and the recent quote from Williams confirms it: "In my opinion [Haldane's dilemma] was never solved, by [Bruce] Wallace [and soft selection] or anyone else. It merely faded away, because people got interested in other things. They must have assumed that the true resolution lay somewhere in the welter of suggestions made by one or more of the distinguished population geneticists who had participated in the discussion." Williams is the exception to the rule. He is the *only* leading evolutionist to state that Haldane's dilemma remains unsolved. That is not a ringing vindication of evolutionists. On the contrary, it is an admission of a four-decade *failure* to teach and pursue the truth. Chris uses the words "cover up", which I did not use. I do not think the obfuscation of Haldane's dilemma was intentional, but then again it was not a random accident of nature. Evolutionists did it, and it is for them to explain. Walter ReMine Standard disclaimer: These thoughts are mine. P.O. Box 19600 Saint Paul, MN 55119 From: us236454 at dawn.mmm.com (Walter Remine) Newsgroups: talk.origins Subject: Haldane's Dilemma Date: 9 May 1995 13:43:42 -0500 Message-ID: <3ood4u$krr at dawn.mmm.com> *** Haldane's Dilemma is not invalidated by genetic variation *** Andy Peter's writes: >In his latest post, ReMine offers nothing but flat nay-saying of my >points: he continues to make no effort to address any specific argument >I've made. Wrong. Andy has (somehow) missed the fact that I turned his own argument against him. His mutation load argument is ***his*** argument. He finds it ***totally*** convincing. Therefore, he cannot dodge its conclusion: The standard model of genetic evolution -- the one featured in all evolution books -- is clearly false and massively contradicts macroevolution. Andy keeps trying to dodge his own conclusion. By Andy's own argument -- using the standard model of genetic evolution -- species' are so full of harmful mutations that they could not survive even two generations, and would go extinct. Andy challenges me to overturn his argument. Why should I? Andy dug himself a hole. Why should I dig him out? I repeatedly asked Andy for literature references that make the same argument he does. I know of none, and he has not supplied any. I also supplied a reference (G.C. Williams, 1992) where Andy's identical same mutation load analysis is employed to reach a conclusion opposite from Andy's. It is used to show the "severity of Haldane's dilemma." To that, Andy offers no comment. Andy writes: >ReMine's arguments ... have basically been comprised >of insinuations that: [1] all variation is neutral Wrong. I never said all variation is neutral. I said that neutral variation neither helps, nor harms, nor invalidates Haldane's dilemma (or Andy's mutation load problem for that matter). Moreover, I have *agreed* with Andy that the level of harmful mutation is quite high. This is a serious problem for *any* model of genetic evolution. >... and [ReMine makes] insinuations that selection doesn't >happen in the real world. Wrong. I never said that selection (ie. differential survival) doesn't happen. I said that Andy's "Natural selection happens!" argument hid the central issues. ****** Andy's post is another repetitious re-statement of his previous posts, while being completely non-responsive toward my arguments. It seems the discussion has reached an impasse, so I will state the conclusion again, for those readers who may have arrived late. Andy's "genetic variation" argument does not invalidate Haldane's Dilemma, as Andy once claimed it had. Rather it invalidates the standard model of genetic evolution. The cost of substitution and the cost of harmful mutation are unavoidable, and they remain serious problems no matter what genetic model you choose. These problems did not die with Haldane, or with Andy's refutation of the standard model. The problems remain unsolved. Walter ReMine Standard disclaimer: These thoughts are mine. P.O. Box 19600 Saint Paul, MN 55119 From: us236454 at dawn.mmm.com (Walter Remine) Newsgroups: talk.origins Subject: Haldane's Dilemma Date: 10 May 1995 01:11:53 -0500 Message-ID: <3oplf9$ma8 at dawn.mmm.com> *** Haldane's Dilemma and soft selection *** Chris Colby offers soft selection as "at least part of the solution" to Haldane's Dilemma. (He's already hedging away from previous statements.) Chris's discussion follows that of Bruce Wallace, the originator of the "soft selection" concept. While the following criticisms are directed at Chris's text, they apply to Wallace's material also, as they share the same errors. ****** The cost of substitution is often equated (dubiously) with genetic load, and Chris follows that tradition. Genetic load focuses on selective value, also known as "fitness". Thus, discussions of cost are diverted into discussions of load, which are diverted in discussions of fitness, which are diverted into discussions of: "optimal" versus "average" fitness; "absolute" versus "relative" fitness; scaling of fitness; normalizing fitness values to 1; whether the most fit (or the least fit) organism should be normalized to a fitness of 1; and the normalizing of population sizes. (And that's just for starters. That's before we even try to physically interpret the concept of load.) Because of these confusion factors, discussions of load are frequently opaque, even to evolutionary specialists. Chris follows that tradition also. For sake of documentation, here are examples. Notice the frequent, confusing focus on "fitness." (Notice also the incorrect use of the term "cost.") Don't get lost, I'll explain things down below: In [hard selection], the presence of relatively more fit alleles decreases the absolute fitness of other alleles. .... [In hard selection], the appearance of more relatively fit alleles lowered the _absolute_ fitness of other alleles. Under [soft selection], + alleles raise mean fitness because it is not scaled. Put another way, in [hard selection], the appearance of more relatively fit alleles lowered the _absolute_ fitness of other alleles. In [soft selection], this is not the case. A less fit allele does not induce a cost to more fit alleles. This is a more realistic way of modeling evolution (IMHO). Let's say a type of invertebrate normally lays 500 eggs; would the old type that lays 500 eggs impose a cost that had to be paid by the 550 egg layers? Would the presence of 550 egg layers lower the amount of eggs 500 egg layers laid? It would have to in order to accord to the assumptions of [hard selection]. But, you would not expect it to. Formulated this way, one can see that the details of how fitness differs at a loci is important to seeing if there is a cost imposed by a new allele. Specifically, how much does the new allele lower the absolute fitness of the other allele(s) at that locus. ****** I prefer the term "cost of substitution" because it is clear, and it immediately focuses your attention on the central issue -- substitution. Substitution has a cost. If a species is to substitute a trait, then there is a cost. * * Here is the rule: * * If a trait is be substituted into a population, * then the trait must go from 'few' to 'm