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Haldane's view of Haldane's Dilemma

by Walter ReMine
August 2007

Summary:
Introduction
Statements of faith
Haldane's attempt at a solution

Conclusion

Footnotes
In 1957 J.B.S. Haldane discovered the evolutionary speed limit1 that later became known as Haldane's Dilemma and remains unsolved. Though Haldane attempted to solve the problem, his attempt was superficial and is irrelevant today except as a back-page in the history of science, of interest only to historians of science. Nonetheless, Internet evolutionists2 (not the professional literature) are falsely trying to make it an issue today. 

This article:

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Introduction —

My book offers cutting-edge technical issues on origins, in a manner the general public can understand.3 Those two goals often conflict, and achieving both goals was not easy. Most readers had scarcely heard of population genetics, much less had any interest in it, and Haldane's Dilemma can be especially technical. So getting readers to slog through such technical material was a challenge. To do that, I first orient readers to the journey ahead and indicate why it is worth their effort. I do that in the first paragraph. 

[O]ur brains tripled in size within a million years or so (Gould, 1980, p 131)

In the 1950s the evolutionary geneticist, J. B. S. Haldane, calculated the maximum rate of genetic change due to differential survival. He reluctantly concluded there is a serious problem here, now known as Haldane's Dilemma. His calculations show that many species of higher vertebrate could not plausibly evolve in the available time. We begin our study with a simplified account of the calculations. (Walter ReMine, The Biotic Message, p 208, first paragraph of Chapter 8, Haldane's Dilemma.)

That paragraph specifically refers to Haldane's "calculations" — not his conclusions, his beliefs, or his statement of faith — and the chapter details precisely what Haldane's "calculations" refers to. That does not misrepresent Haldane. Rather it is a simple introduction to a chapter, accurately telling my readers what they are about to read. 

Evolutionist Ian Musgrave accuses that paragraph of misrepresenting Haldane.

In [Haldane's] entire paper (nor in his later 1961 paper), there is NO mention of any serious problem. .... One of the main points of my essay was the Walter ReMine seriously misled people over Haldane’s dilemma, attributing to Haldane a position Haldane did not in fact take. (Ian Musgrave, 2007)4

I did not misrepresent Haldane. 

First note the context of my book. My book argues that in each instance, evolutionary theory (large-scale evolution) is either false, or unfalsifiable — and either way it is not scientific, by the same criteria that evolutionists used in all their court cases. There are instances where Classical Darwinism and various evolutionary theories are falsified, and on those instances modern evolutionists save their theory by making it unfalsifiable, which gives evolutionary theory its essential modern character — untestable, unfalsifiable, un-risky, and therefore unscientific. We've all heard the fluff stories that can explain-away anything and its opposite — How the leopard got its spots. In practice, evolutionary theory is a structureless smorgasbord of just-so storytelling, for which the theory is renowned. This large portion of evolutionary thinking is dubious and soft. This is the context of my book, and my readers are well immersed in it by the time they get to Chapter 8, Haldane's Dilemma. 

Compared to those usual evolutionary faults, Haldane correctly saw his cost concept as firm science that poses serious quantitative problems for evolutionary theorists to solve. He saw it as serious enough to publish papers on it. And he concluded, "I am convinced that quantitative arguments of the kind here put forward should play a part in all future discussions of evolution." Yes, Haldane saw it as a serious problem, and I give him credit for that. (Ian Musgrave omitted Haldane's statement, because it contradicts his accusation.) 

Haldane made a superficial attempt to solve the problem, but he acknowledged, "I am quite aware that my conclusions will probably need drastic revision", which shows he did not really consider the problem solved.

Kimura and Crow (1969) explicitly endorsed Haldane's conclusion that cost arguments "should play a part in all future discussions of evolution." And evolutionary geneticists themselves named the problem, "Haldane's Dilemma." Kimura and Maynard-Smith each cited Haldane's Dilemma as the "main reason" for their revolutionary new views of evolutionary process. Many evolutionary geneticists saw it as a serious problem. 

Statements of faith —

Haldane remained a committed evolutionist. His cost papers did not acknowledge any refutation of large-scale evolution, but then again, they did not address large-scale evolution, other than superficially. Instead, they stated his naked belief that his speed limit "accords with the observed slowness of evolution." and "The agreement with the theory here developed is satisfactory." Such statements of faith — statements substantially disconnected from the evidence — are routine in evolutionary papers, especially on highly problematic topics, starting with the origin-of-life and continuing through countless unsolved evolutionary issues.

Here is how the process works. An evolutionary specialist on a particular group of organisms reluctantly acknowledges an inability to find ancestors, and then gives a bald statement of faith, claiming the agreement with evolutionary theory is "satisfactory." This situation, repeated again and again, for all kinds of organisms, gives a rather different picture of the evidence — the ancestors are systematically absent. This occurs again on other anti-evolutionary evidences, such as gradualism, stasis, morphological gaps, the Cambrian Explosion, "convergence," and the rest. The evolutionists' statements of faith are unworthy of scientific respect, yet they are abundantly given, and abundantly quoted, to convince the unwary public that evolution is a "fact." It is the worst kind of quotation mongering. 

This goes far beyond Haldane. Evolutionary leaders operate like a high priesthood, reassuring the congregation with statements of faith, and a wave of the hand. Then evolutionary activists (such as Ian Musgrave) quote those statements of faith as though it had deep scientific merit. Never mind the abundant scientific evidence against macro-evolution, instead evolutionists quote their leader's belief in evolution as though that should be convincing. 

How is a science author to deal with that? My book does not cite the numerous instances where evolutionists give statements of faith. Instead, my book periodically reminds readers that the cited experts are committed evolutionists. This relieves readers from much highly repetitive material of dubious scientific value, while duly acknowledging the evolutionists' beliefs. This process starts in the Preface:

This is a science book about the modern creation/evolution debate, with emphasis on biological issues. I quote frequently. I have excluded quotation from anti-evolutionary literature, such as the books by Denton, Fix Hitching, P. E. Johnson, and G. R. Taylor. I quote only twice from creationists, and then only to disagree. (These two cases are clearly marked.) Except for those two instances, I quote exclusively from committed evolutionists. I have concentrated on mainstream evolutionary literature, with special effort to survey all the anti-creation material. In fact, this book is driven by the issues raised by modern anti-creationists, I quote from these sources to authenticate data or an opinion that I discuss. This is not to imply the quoted author would necessarily agree with the point I make. I quote no authors who would presently have the slightest favor for my views. (Walter ReMine, The Biotic Message, Preface, p 2, underlining added)

My book periodically re-emphasizes the point. For example: 

For documentation, I quote only evolutionists. Each of them firmly believes that large-scale phylogeny exists, and their statements to that effect are ubiquitous, if not unavoidable, Yet there can be (and is) a difference between what they believe and what they observe. For the sake of clarity, I try to separate the two and cite the observations. I also avoid citing their illusory statements [which are discussed in the immediately previous chapter]. That is the context of these quotations. (Walter ReMine, The Biotic Message, p 302) 

In this same way, Haldane's statement of faith (in evolution) was disconnected from serious support. It carries little more scientific weight than facing east and bowing deeply.

Haldane's attempt at a solution —

Haldane 1957 established a compelling theoretical basis for an evolutionary speed limit. Then he attempted to square it with evolution, in twelve sentences. That evolutionists would give it serious weight is another of the scandals surrounding Haldane's Dilemma. 

Haldane's attempt focused on small-scale evolution, such as the difference between two closely related species, and the emergence of a new species from an immediately previous species. Haldane used the species concept to calibrate his discussion, and it was central to his discussion. Yet species and speciation remain today as two of the most ill-defined and controversial concepts among evolutionists themselves, and that provides a large fudge-factor at the center of Haldane's brief discussion. For example, contemplate how many 'species' there were in a given lineage over ten million years. Was it two species? Or was it two thousand species? Haldane's discussion relies on that ambiguity.

Haldane used the average duration (or "mean life") of a species.5 For example, if the average duration of a species is one million years, and we allow ten million years for evolution, then a given lineage would expect a sequence of ten species. This implicitly assumes anagenesis, which is the focus of Classical Darwinism. In anagenesis, a species is transformed into a daughter species, and therefore cannot coexist with a daughter species. Each 'species' replaces the one before it. However, as claimed by Stephen Jay Gould and other paleontologists, fossil species strongly exhibit stasis (or lack of change) for their entire duration, and therefore most evolutionary change must have occurred in sudden rapid bursts at speciation events (or the splitting of a lineage, known as cladogenesis), and this would mean each ancestor species can overlap in time with numerous daughter species. In other words, Haldane underestimated the number of 'species' in each lineage. (Say, by a factor of a thousand?) These matters remain unresolved even today, but they were thoroughly obscure in Haldane 1957. Once again, Haldane's focus on 'species' provided ambiguity that overwhelmed his discussion. 

Haldane assumed the origin of a new 'species' requires merely one thousand substitutions. Besides being wildly speculative, his assumption hinges again on the ambiguous term, species. For example, compare coyote, fox, dog, and wolf, which are classified as separate species. Yet these organisms are linked by the ability to interbreed, which indicates a high degree of genetic identity. There are many examples like this.6 [Such organisms are within the same created kind (or more precisely, the same monobaramin), according to creationists.] Such organisms would have a relatively small number of functional genetic differences. Haldane calibrated his discussion based on "closely related species," yet if they are too closely related, as in the above examples, then it would give a false underestimate of the requirements for large-scale evolution. Yet Haldane did not say what sort of "closely related species" he meant in his above assumption. He did not identify what species he was referring to. This added another layer of ambiguity to his discussion. 

The ambiguity allowed equivocation, where the meaning of a key-term is shifted to have different meanings within the same argument. The term "species" meant one thing (large morphological change) when discussing the fossil record, and a rather different thing (a small number of loci differences) when discussing the genetics of "closely related species" living today. These equivocations were packed into the term species, and Haldane's brief discussion did not clarify it. This all increased the slack and ambiguity of his discussion, which worked to evolution's favor. 

Next focus on the genetics. There was ambiguity over the terms "loci" and "gene." In Haldane's day these were presumed to have a one-to-one relationship with polypeptides (proteins and enzymes) that are actually produced in the organism. In effect, one loci equals one gene, equals one protein. Alternatively, these were identified as phenotypic traits (or genetic linkage blocks) that are inherited intact (without recombining). Those definitions are now known to conceal much biological complexity. Today these terms are more abstract, and harder to define. Haldane had unknowingly hidden much complexity within the terms loci and gene, thereby making the genome seem less complex than it really is.  

Also, gene regulation is seen today as centrally important. But in Haldane's cost papers, it was either ignored, or it was implicitly 'packed' somehow within the concept of loci and gene. Either way, it artificially minimized his depiction of the genome's information content. 

Following tradition in his profession, Haldane wrote of "gene substitutions," which created the false impression that large blocks of entirely new DNA are being substituted into the population. That habit further obscured the severity of Haldane's Dilemma from public view. The substituting 'thing' is not a gene, instead it is a mutation (typically one nucleotide, according to evolutionary geneticists). 

Haldane 1957 said the number of loci in a vertebrate species has been estimated at 40,000. He then implicitly argues that 1000 substitutions represents a substantial portion (one-fortieth) of the genome. His argument was misleading, even at the moment he wrote it. "40,000 loci" is not remotely a sufficient measure of the genome's information content, because, for example, it does not account for the organization of genes and loci. (The ordering of genes within a given chromosome arm. Gene inversions. And so forth.) There is no reason this information could be ignored. Yet Haldane ignored it, and that further minimized his depiction of the genome's information content. He had no grounds for his argument. 

Modern research reveals genomic structure of astonishing complexity, for which there are many examples. Overlapping genes. Genes within genes. Genes overlapping on opposite strands of DNA, coding in both directions. Many examples of a gene that codes for five and six different proteins, depending on how the transcripts are spliced together. Genes that have more than one effect on the organism (i.e., pleiotropy). Nearly all of the genome is transcribed and is not silent. Even the traditional "silent codons" sometimes have a substantial biological effect. The traditional evolutionist concept of "Junk DNA" is evaporating. And different groups of organisms have different genomic organization (which evolutionists are now calling "structural rearrangements"). Haldane profoundly underestimated the information content of the genome, and thereby gave a misleading perspective on the number of substitutions required. 

Now examine Haldane's actual statement, starting with his central reasoning:

Presumably this same kind of process occurs in evolution. The number of loci in a vertebrate species has been estimated at about 40,000. 'Good' species, even when closely related, may differ at several thousand loci, even if the differences at most of them are very slight. But it takes [the same cost of substitution] to replace a gene by one producing a barely distinguishable phenotype as by one producing a very different one. If two species differ at 1000 loci, and the mean rate of gene substitution, as has been suggested, is one per 300 generations, it will take at least 300,000 generations to generate an interspecific difference. It may take a good deal more, [if multiple substitutions are required at the same locus]. (Haldane, 1957, p 521-522, underlining added)

Haldane's statement can also be challenged on its own grounds. Start with our alleged last common ancestor with chimpanzees, which would be a pre-chimpanzee of some type. Haldane's statement would mean the very next species would have to be humans — from pre-chimp to human in one step. Such a claim should have set off alarms among evolutionists.

Continuing: 

"Simpson (1953) finds the mean life of a genus of Carnivora to be about 8 million years. That of a species in horotelic vertebrate evolution may average about a million. 

Zeuner (1945), after a very full discussion of the Pleistocene fossil record, concluded that in mammals about 500,000 years were required for the evolution of a new species, though in the vole genera Mimomys and Arvicola the rate was somewhat greater. Some insects evolved more slowly. He estimated the total duration of the Pleistocene at 600,000 years, but some later authors would about half this figure. On the other hand, environmental changes during the Pleistocene were unusually rapid, and evolution, therefore, probably also unusually rapid." (Haldane, 1957, p 522)

Haldane's twelve sentences (quoted above) are his entire rational for leaping to conclude, "The agreement with theory here developed is satisfactory." No evolutionary genetics literature cares to defend those twelve sentences as a sufficient resolution of Haldane's Dilemma. 

Haldane's speed limit applies with special force to organisms with low reproduction and long generation times — such as the origin of elephants, whales, bears, apes, and humans — but he did not examine these most problematic cases. Instead he mentioned large ambiguous groupings of organisms, most of which have high reproduction and short generation times, such as insects, molluscs, voles, the lower vertebrates, and lower mammals.

Also, Haldane switched horses midstream. He began with units of "generations" and then switched to "years", without translating between the two. Since he did not identify the species (or their generation time) this added yet another layer of ambiguity to his discussion.

Haldane cited Zeuner (1945) concerning speciation rates during the Pleistocene. Haldane then casually mentioned that later authors would about half the total duration of the Pleistocene, but he did not mention this would double the speed requirements. This added further ambiguity to Haldane's discussion. Just who's figures did he use? 

Haldane notes that "environmental changes during the Pleistocene were unusually rapid, and evolution, therefore, probably also unusually rapid." This is just-so storytelling, disconnected from his cost concept. Haldane gave no serious grounds for why rapid environmental changes should (over the long term) lower the amount of genetic death, or lower the cost, or increase the substitution rate. It was simply a just-so story. Environmental change, like mutational change, is random with respect to organisms, and is predominantly harmful to organisms. Environmental-change is one of the traditional confusion factors surrounding Haldane's Dilemma, and addressed in my book and papers. 

My book describes the problem directly, without ambiguity, by avoiding the intermediary concepts of species, gene, and loci, and by focusing on the most problematic case that everyone can understand. That is, according to Haldane's calculations, evolutionists would have to explain the origin of all the human adaptations over ten million years within a maximum limit of 1,667 beneficial substitutions, (where each substitution is a mutation, and a typical substitution is said to alter one nucleotide, but might occasionally be an inversion, a duplication, a deletion, the relocation of a gene on a chromosome, a mutational change to gene regulation, and so forth). This Haldane limit does not depend on how species, loci, and gene are defined. Haldane could have easily stated it this directly, but he did not. Instead his handling obscured the issue with ambiguity, and then brushed it aside with a statement of faith. This was to Haldane's discredit.

Conclusion —

Haldane's discovery of the cost concept was brilliant. Though his attempt to solve the issue was brief, ambiguous, and superficial. In short, it was scientifically juvenile. This is forgivable, because a solution was not the focus of his papers, and he fairly acknowledged his conclusions "will probably need drastic revision". However, this matter is  not forgivable in the following generations of evolutionary geneticists, who obscured Haldane's Dilemma and brushed it aside as solved, when it was never solved. 

Internet evolutionists claim Haldane was unsure about quantitative cost arguments, when in reality he was justifiably confident these "should play a part in all future discussions of evolution." And they claim he was confident about his solution to the problem, when he actually said it will "probably need drastic revision."7 They have it exactly backwards. 

The Haldane speed limit is based on genetic models and assumptions still current today in evolutionary genetics textbooks. In fact, those are wildly unrealistic in favor of evolution, which makes Haldane's Dilemma all the more problematic. (See Haldane's assumptions.) Internet evolutionists claim the Haldane speed limit is based on "out-dated models and assumptions from the 1950s," when it actually remains current — and they cite Haldane's statement of faith (in evolution) as definitive today, when it is actually scientifically worthless. They have it exactly backwards.

Haldane's statement of faith (in evolution) was a discredit for him, and would have been a tedious detour for my book and the general public. The issue may be interesting to academic historians of science, but it has little relevance, one way or the other, to readers of the modern origins debate, who want to know the nature of Haldane's Dilemma and whether it is solved today. On this point, Haldane's statement of faith is irrelevant. If evolutionists wish to make an issue of that, they are welcome to try.

The writer's art requires not abusing reader's patience with side issues, especially when the material is already complex. My book had to judiciously choose which issues my readers would find relevant, and had to explain complex details of population genetics accurately, yet in a manner the general public can cope with. I am immensely pleased my readers found the material readable, understandable, interesting, and compelling. It is fair to say that without my book and its pro-active readership, evolutionists would have allowed Haldane's Dilemma to remain where they had placed it, in oblivion. The reason Haldane's Dilemma is discussed today is because my many readers see its importance and press evolutionists for answers. I am absolutely unapologetic about that.

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Footnotes — 

1    Haldane, J.B.S., "The Cost of Natural Selection", J. Genet. 55:511-524, 1957.

2    With rare exceptions Internet evolutionists are non-specialists who operate anonymously. They abundantly sow confusion, so long as it favors evolution. More significantly, evolutionary leaders negligently remain silent and allow the confusions to thrive, so long as it favors evolution. This ill-health occurs abundantly. This tactic allows all evolutionists to benefit from confusion, without evolutionary leaders themselves having to take responsibility for it. This webpage gives more examples. 

3    The Biotic Message, Walter ReMine, http://SaintPaulScience.com 

4    Ian Musgrave, at PandasThumb.org, July 1, 2007, entry 3202 and a follow-up post, www.pandasthumb.org/archives/2007/07/haldanes_nondil.html  Ian Musgrave repeats an accusation made previously on the Internet by evolutionist David Wilson (groups.google.com/group/talk.origins/msg/c2967947318fe625?hl=en&). Musgrave and Wilson misconstrue the meaning of my words. My words, "serious problem", simply means serious problem, a view that Haldane did in fact take. 

5    Haldane's discussion used the average duration (or "mean life") of a species. Musgrave mistakenly referred to that as the "average rate of speciation". Those are different things. The two are connected (and reciprocal) only if Musgrave was using Classical Darwinism (and anagenesis) as his model, which favors evolutionists here by under-estimating the number of 'species' required. That model is no longer in favor. It has been substantially replaced by punctuated equilibrium and an emphasis on cladogenesis, as explained above. 

6    There are many examples of organisms linked together by an ability to interbreed. Such as: the camel group (including camel and llama); the horse group (including horse, zebra, donkey, quagga, [and mule]); the canine group (including dog, fox, wolf, and coyote); the feline group, (including lion, tiger, and many other cats); the Anatidea family (including swan, duck, goose, and pintail); the various bears (including polar bear, brown bear, and grizzly). 

7   Haldane 1957 wrote, "I am quite aware that my conclusions will probably need drastic revision". He was referring specifically to his "conclusions," recited in his closing paragraph, not to quantitative cost arguments, which he was rightly confident "should play a part in all future discussions of evolution." 

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Updated Sep 10, 2007