Natural Selection and Adaptation

Cornelius Hunter seems very confused.

 …This brings us back to the UC Berkeley “Understanding Evolution” website. It abuses science in its utterly unfounded claim that “natural selection can produce amazing adaptations.”

In fact natural selection, even at its best, does not “produce” anything. Natural selection does not and cannot influence the construction of any adaptations, amazing or not. If a mutation occurs which improves differential reproduction, then it propagates into future generations. Natural selection is simply the name given to that process. It selects for survival that which already exists. Natural selection has no role in the mutation event. It does not induce mutations, helpful or otherwise, to occur. According to evolutionary theory every single mutation, leading to every single species, is a random event with respect to need.

He has forgotten what “adaptation” means.  Of course he is correct that “Natural selection is simply the name given to [differential reproduction]”.  And that (as far as we know), “every single mutation …is a random event with respect to need”.

And “adaptation” is the name we give to variants that are preferentially reproduced. So while he would be correct to say that “natural selection” is NOT the name we give to “mutation” (duh); it IS the name we give to the very process that SELECTS those mutations that promote reproduction.  i.e. the process that produces adaptation.

Cornelius should spend more time at the Understanding Evolution website.

ETA: CharlieM points out below that…

When CH says that natural selection does not produce adaptations he is talking about individual organisms. He is discussing mutations in individuals and adaptations in individuals. Natural selection has nothing to do with the first appearance of an adaptation in an individual.

 

And that of course is the confusion – I hadn’t seen just where Cornelius’s confusion lay.  Because, of course, the term “adaptation” is a population-level concept. At the level of the individual, the equivalent would be  “advantageous mutation”.  And that makes the Understanding Evolution website absolutely correct.

843 thoughts on “Natural Selection and Adaptation

  1. stcordova: We can’t use the fact a trait is selectively favored today as evidence it was selectively favored in the deep past.

    Who does that? Niches fill, close and become available. The environment is constantly changing and where long-lived stable niche environments are found, organisms that find and occupy those niches are under less selective pressure.

  2. stcordova,

    At that point, what defines a eugenically ideal phenotype is a functionally defective creature relative to its ancestors. An entire population can continue to be reduced functionally by reductive evolution while all the while getting more eugenically “fit” in the sense of population genetics. Absurdity.

    So you are saying that the only mutations that ever become fixed are those that would not be competitive if the ancestors still lived? That’s not supported by anything. Including Lenski.

    But if it was … what celestial mechanism opposes this assumed degenerative tendency?

  3. So you are saying that the only mutations that ever become fixed are those that would not be competitive if the ancestors still lived?

    No! I was pointing out the absurdity of the notion of “fit” used by population geneticists to the notion of “fit” in the Darwinian blindwatchmaker functional sense. Equivocation abounds and one that Dennett resorts to in his algorithm that population keep getting fitter (true in a population genetic sense) but not necessarily true in a functional sense.

    You’re also making the mistake of defining function in terms of fitness. Sickle cell anemia is highly fit, but it is not functional except in the sense blowing up a bridge is functional to prevent an invasion. An X-box is functional, it’s functionality is not defined by the ability to create reproductive success. Birth control pills are an intelligently designed pharmaceutical — their intelligently designed functionality should not be defined by reproductive success.

    There is a lot of extravagance in biology that is selectively unfavored in the long run, but still evidences interdependent Rube Goldberg complex function.

  4. stcordova,

    Indeed.

    Heck, for the biologist, the reason our cells decay and become old and die, is because that is more fit than an individual that lives forever.

    Why can’t humans cut off their legs and just grow back new ones in a few days? Well, because not being able to grow back new ones is a sign of greater fitness.

  5. phoodoo: Why can’t humans cut off their legs and just grow back new ones in a few days? Well, because not being able to grow back new ones is a sign of greater fitness.

    Wrong, because not being able to grow back new ones is a sign of god’s love for you, little human. Don’t question the almighty god’s plan.

    Oh, can’t help myself from asking:

    Why doesn’t god ever heal amputees?

  6. stcordova,

    No! I was pointing out the absurdity of the notion of “fit” used by population geneticists to the notion of “fit” in the Darwinian blindwatchmaker functional sense. Equivocation abounds […]

    That ‘E’ word again. It must be a US thing. No-one uses the term ‘fit’ in an attempt to be ambiguous. Nor to cavil con dodge elude escape eschew evade falsify fence fib flip-flop fudge hedge jive lie palter parry prevaricate pussyfoot quibble shuck shuffle sidestep stonewall tergiversate waffle beat around the bush beg the question cloud the issue cop out cover up double-talk give run around hem and haw mince words pass the buck sit on the fence tell white lie or weasel They even rigorously define their bloody terms, as we have been doing here! That’s a strange kind of equivocation.

    ‘Fit’ as in ‘fit for purpose’ or ‘physically fit’ is not the sense in which population geneticists use ‘fit’. It was almost an in-joke – that quality that makes ‘survival of the fittest’ true became what population-genetic ‘fitness’ meant. It all reduces to producing more net offspring than your rivals, integrating survival and reproduction.

    Dawkins has a nice chapter on this in The Extended Phenotype – “An Agony In Five Fits”. He prefers to avoid the term altogether. Yes, it can be confusing. But that confusion is easily remedied.

    You’re also making the mistake of defining function in terms of fitness […]

    I never even mentioned function, so I don’t know where you got that from.

    Sickle cell anemia is highly fit

    Not really. The heterozygote has certain advantages in certain environments, but not others. The homozygote is stronly detrimental. Cherry-picking ‘broken’ genes or vestigial traits is not really much of a guide to the entirety of evolution’s capacity, either.

  7. stcordova: Selectionist theory is next to worthless in study of long term evolution. What works well for describing truncation selection in anti-biotic and pesticide resistance isn’t applicable to changes over deep time primarily because what defines “fit” is equivocated, redefined, and auto-renormalized over many generations.

    With respect Sal, I find most of the above gobbledygook. What is “selectionist theory”? Do you mean the theory of Natural Selection? And why do you say that “what defines ‘fit’ is equivocated”? Do you mean that people use different definitions in different contexts? Or that they don’t define what they mean? Or that they don’t know what they mean? Because it seems to me that people use “fit” in very precisely defined ways. Can you give me an example of equivocation?

    stcordova: In species that have insulin receptors and use insulin to regulate their metabolism individuals unable to synthesize insulin will be selected against (since it’s pretty much lethal) and hence we say insulin is selectively favored.

    Evolutionists falsely presume this is evidence insulin came about through natural selection since it is life critical. The problem however is we have no data on the selective pressure on insulin when the machinery for an insulin metabolism isn’t present. As St. Mivart pointed out, selection can’t select for non-existent traits! We can’t use the fact a trait is selectively favored today as evidence it was selectively favored in the deep past.

    What “evidence” are you referring to that “evolutionists falsely presume” is evidence that insulin evolved? Can you cite your source for this claim?

    stcordova: Lewontin points out fitness is an ill-defined concept in his Santa Fe 2003 papers, and that paper pretty much unDarwinized Stanley Salthe.

    Fitness is not an ill-defined concept. It’s an extremely well-defined concept. Ask Joe Felsenstein.

    stcordova: Suppose random drift causes a population of cave fish to be all blind or any given population for that matter to lose some functional trait. At that point, what defines a eugenically ideal phenotype is a functionally defective creature relative to its ancestors. An entire population can continue to be reduced functionally by reductive evolution while all the while getting more eugenically “fit” in the sense of population genetics. Absurdity.

    Any misrepresentation of a hypothesis runs the risk of being an absurdity. Nobody “defines a eugenically ideal phenotype”. It’s not a concept that plays any role in evolutionary theory or biology. Fitness is defined entirely in relation to the current population in the current environment.

  8. phoodoo,

    Indeed.

    Heck, for the biologist, the reason our cells decay and become old and die, is because that is more fit than an individual that lives forever.

    Why can’t humans cut off their legs and just grow back new ones in a few days? Well, because not being able to grow back new ones is a sign of greater fitness.

    You might want to take the absence of these capacities up with your Designer. Built-in obsolescence and irreplaceable parts. Cheers, God!

  9. Joe:

    I am glad to hear that we all knew all along that a population growing without limit could see one allele taking over because of that-bias-that-isn’t-drift-or-mutation.

    That bias-that-isn’t-drift-or-mutation is relative fecundity in my scenario. Selection (by Darwin’s and Mayr’s and Gould’s and Allan’s and my definition) is absent, because the environment doesn’t favor either of the two variants. And yes, even Archimedes could have figured out that if variant B produces twice as many offspring as variant A, and all offspring survive and reproduce, then B will come to dominate the population. We certainly didn’t have to wait until 1903 and the development of the quantitative theory. To call that ‘selection’ is a definitional choice, not an empirical one.

    My point is that you can define “natural selection” as you wish, but that is not binding on evolutionary theorists, who use it my way.

    Not so fast. Some do, some don’t. There is no canonical definition among either scientists in general or evolutionary theorists in particular. See my response to Steve. (I’d say Ernst Mayr and Stephen Jay Gould qualify as ‘evolutionary theorists’, wouldn’t you?)

    I’ll ask again:

    In pop gen, “selection coefficient” is defined as it is as a matter of convenience, and I don’t begrudge you that. What I don’t see is why the pop gen definition should prevail in the current discussion, which is about natural selection’s role in the production of adaptations. As far as I can see, Lizzie has offered no good reason why it should prevail.

  10. Allan,

    Careful circumscription of one’s population of interest is of course very important. One has the thorny issues of how ‘the population’ is bounded in complex bacterial ecosystems … and sex.

    True, though in my scenario the population is quite easily circumscribed: it’s the original population plus all descendants.

  11. With respect Sal, I find most of the above gobbledygook.

    Well thanks anyway for reading and responding. 🙂

    What is “selectionist theory”? Do you mean the theory of Natural Selection?

    Selectionist theory vs. neutralist theory or other evolutionary theory — the primary mode of speculated evolution.

    The phrase is not mine alone, it is used here:
    http://www.nbi.dk/~natphil/salthe/Critique_of_Natural_Select_.pdf

    And why do you say that “what defines ‘fit’ is equivocated”?

    Population geneticists use the notion of “fit” in terms of reproductive schedules. Sickle cell anemia is considered “fit” in a malaria environment. Tay-sac disease is considered fit in the Darwinian sense as well as well as blindness in cavefish, winglessness in beetles and all the trashing created by reductive evolution and the mass destruction of function via extinction via natural selection.

    Clearly the idea of fit as in the medical and engineering sense is equivocated with the definition of “fit” in population genetics which defines it in terms of reproductive success for an existing species line. If you argue that selection is the mechanism that created functional proteins, that’s equivocation on many levels.

    A protein being essential for life today is no evidence it was selected for in the past, especially before it even existed. You can’t have selection pressure toward non-existent traits. That’s like working toward a pre-meditated unseen goal which the blindwatchmaker cannot do as a matter of principle. If anything a life critical trait is evidence against selection being the mechanism of evolution, because without the trait, the organism dies, ergo, no evolution! For evolution to work, the trait has to be not-so-critical, maybe even neutral.

    So trait being critical today is equivocated with the idea it was favorably selected for in the past (usually even before the trait existed!).

    when asked which traits are most likely to be able to evolve, evolutionary biologists, again citing Fisher’s theorem, will reply, “those that have more variability in fitness”. That is to say, traits that have been most important in the lives of organisms up to this moment will be least likely to be able to evolve further! So Fisher’s theorem is “schizoid” when one compares its postures facing the future or the past. And once again one faces the possibility of single traits evolving sequentially, building up by way of ontogenetic agency an overall adapted phenotype subjected to an increasing genetic load directed at maintenance.

    Stanley Salthe

    Furthermore, what about all that elimination of function through extinction via natural selection (like we humans dominating the biosphere and torching inferior species, bwahaha)? How is net reduction of functional diversity a net gain of functional diversity?

    The confused thinking is not mine, it’s in the mind of evolutionary biologists.

    Andreas Wagner said, “fitness is hard to define”.

    However, fitness is hard to define rigorously and even more difficult to measure….An examination of fitness and its robustness alone would thus not yield much insight into the opening questions. Instead, it is necessary to analyze, on all levels of organization, the systems that constitute an organism, and that sustain its life. I define such systems loosely as assemblies of parts that carry out well-defined biological functions.

    Andreas Wagner

    Wager instead focuses on systems and defines them as “assemblies of parts that carry out well-defined biological functions”. Compare that to Behe:

    A single system composed of several well-matched, interacting parts that contribute to the basic function of the system

    Now add to all this Lewontin

    The problem is that it is not entirely clear what fitness is. Darwin took the metaphorical sense of fitness literally. The natural properties of different types resulted in their differential “fit” into the environment in which they lived. The better the fit to the environment the more likely they were to survive and the greater their rate of reproduction. This differential rate of reproduction would then result in a change of abundance of the different types.

    In modern evolutionary theory, however, “fitness” is no longer a characterization of the relation of the organism to the environment that leads to reproductive consequences, but is meant to be a quantitative expression of the differential reproductive schedules themselves. Darwin’s sense of fit has been completely bypassed.

    How, then, are we to assign relative fitnesses of types based solely on their properties of reproduction? But if we cannot do that, what does it mean to say that a type with one set of natural properties is more reproductively fit than another? This problem has led some theorists to equate fitness with outcome. If a type increases in a population then it is, by definition, more fit.

    But this suffers from two difficulties. First, it does not distinguish random changes in frequencies in finite populations from changes that are a consequence of different biological properties. Finally, it destroys any use of differential fitness as an explanation of change. It simply affirms that types change in frequency. But we already knew that.

    Lewontin
    Santa Fe 2003

    I’m just relating to you what leading evolutionary theorist have finally realized, and something creationists have been pointing out for a while. Darwinian theory is incoherent on many levels.

    Cornelius Hunter is not the one confused, evolutionary theorist are.

  12. stcordova: I’m just relating to you what leading evolutionary theorist have finally realized, and something creationists have been pointing out for a while. Darwinian theory is incoherent on many levels.

    So is physics and chemistry, and yet we have transistors. One can choose to quote mine and be a shit, or one can attempt to find coherence amidst popular writing.

  13. stcordova: Clearly the idea of fit as in the medical and engineering sense is equivocated with the definition of “fit” in population genetics which defines it in terms of reproductive success for an existing species line.

    No, it is not “equivocated” Sal. Lots of words have more than one meaning. It is not “equivocation” to use them, as long as you, or the context, makes it clear which sense you are using them in. “Fit” in a medical sense is not the same as “fit” in a population genetics sense.

    If you argue that selection is the mechanism that created functional proteins, that’s equivocation on many levels.

    In what way?

    As for your Lewontin quote, it is taken from an essay that you might want to read again in its entirety. It certainly doesn’t suggest that Darwinian theory is “incoherent”.

  14. keiths,

    (by Darwin’s and Mayr’s and Gould’s and Allan’s and my definition)

    Steady on! I’m not an absolutist in these things! I’m more of a shilly-shallying flip-flopper. Selection is fundamentally about alleles having differential effects on rates-of-increase. It doesn’t have to be the environment that determines these rates. Or rather, the environment of any allele includes all the others.

  15. stcordova,

    Clearly the idea of fit as in the medical and engineering sense is equivocated with the definition of “fit” in population genetics which defines it in terms of reproductive success for an existing species line.

    Clearly it isn’t, as any reference to a relevant glossary will show.

  16. Best to avoid words like fitness and stick with differential reproductive success.

    That covers everything from beneficial and detrimental mutations to asteroids.

  17. stcordova,

    If anything a life critical trait is evidence against selection being the mechanism of evolution, because without the trait, the organism dies, ergo, no evolution! For evolution to work, the trait has to be not-so-critical, maybe even neutral.

    If it is not fixed, then clearly it is not essential in those organisms that don’t have it. This does not mean it must therefore be neutral. This is a ridiculous argument. Once it is fixed, other features can come to rely on the facility provided. You think people imagine a fully-functional eye with all wiring fixing in one go by selection?

  18. Allan Miller: Once it is fixed, other features can come to rely on the facility provided.

    An allele does not need to be fixed in a population before individuals having can participate in additional evolution.

  19. keiths:

    Joe:

    That bias-that-isn’t-drift-or-mutation is relative fecundity in my scenario. Selection (by Darwin’s and Mayr’s and Gould’s and Allan’s and my definition) is absent, because the environment doesn’t favor either of the two variants. And yes, even Archimedes could have figured out that if variant B produces twice as many offspring as variant A, and all offspring survive and reproduce, then B will come to dominate the population. We certainly didn’t have to wait until 1903 and the development of the quantitative theory.To call that ‘selection’ is a definitional choice, not an empirical one.

    I disagree. There were several attempts to build a quantitative theory of evolution before 1900 (notably Fleeming Jenkin, Galton, and Delboeuf) but they all failed because there was no understanding of how genetics worked. There was no clear distinction between genotype and phenotype. Some of the confusions can be seen even in Darwin’s writings. As late as 1908 members of the Genetical Society in Britain seemed to think that an allele would spread in a population simply because it was dominant.

    So no, Archimedes would not have been able to do this, as simple as the mathematics would be.

    [regarding populaion genetic definitions of fitness] Not so fast. Some do, some don’t.There is no canonical definition among either scientists in general or evolutionary theorists in particular. See my response to Steve.(I’d say Ernst Mayr and Stephen Jay Gould qualify as ‘evolutionary theorists’, wouldn’t you?)

    Ernst Mayr was not a deep thinker when it came to population genetics.

    In any case, I do not see in those quotes anything that is incompatible with the population-genetic definitions of fitness.

    No doubt our definitions of fitness are oversimplified. They are designed for use in abstract models. Making the models more realistic (for example, allowing overlapping generations) we need more complicated definitions of fitness. Waving one’s hand and dismissing the mathematical theory is easy. What is hard is to put anything coherent in its place. If you’ve got a replacement, let’s hear about it.

  20. Allan,

    Steady on! I’m not an absolutist in these things!

    I’m not either. I have no problem with pop gen terminology and concepts, and I don’t want to change them. I use them myself when working out pop gen practice problems.

    I’m simply suggesting that the pop gen version of selection isn’t the appropriate one for the current discussion.

    It is Steve and Joe who are being absolutist:

    Steve:

    It’s not binding on everyone, of course — only on those who wish to discuss evolutionary biology scientifically.

    Joe:

    My point is that you can define “natural selection” as you wish, but that is not binding on evolutionary theorists, who use it my way.

    I don’t know Steve very well, but I’m surprised to see Joe being so doctrinaire.

    There is no single, canonical definition of “natural selection” among evolutionary biologists or scientists generally.

  21. keiths:

    It is Steve and Joe who are being absolutist:

    Joe:

    I don’t know Steve very well, but I’m surprised to see Joe being so doctrinaire.

    There is no single, canonical definition of “natural selection” among evolutionary biologists or scientists generally.

    There is when you use population genetics.

  22. BTW Allan, I had a similar discussion over at UD not to far back in which I questioned how you could tell the effects of drift from the effects of selection, which Elizabeth then started a thread about here.

    I was maintaining that people were conflating neutral evolution and genetic drift and arguing that even mutations under selection were not immune to genetic drift. So if I read you correctly you would agree.

    So how does one distinguish between the two?

  23. hotshoe_:Why doesn’t god ever heal amputees?

    I’m with you. Why doesn’t God force everyone to be perfect or just kill them off?

  24. Mung:
    BTW Allan, I had a similar discussion over at UD not to far back in which I questioned how you could tell the effects of drift from the effects of selection, which Elizabeth then started a thread about here.

    I was maintaining that people were conflating neutral evolution and genetic drift and arguing that even mutations under selection were not immune to genetic drift. So if I read you correctly you would agree.

    So how does one distinguish between the two?

    You can only do it statistically.

    If a mutation has increased in prevalence with a rapidity improbable under the null hypothesis of drift, you can infer that is probably conferring a selective advantage.

  25. Mung: I’m with you. Why doesn’t God force everyone to be perfect or just kill them off?

    No, you’re not with me.

    I’ve never in my life been a horrible enough person to wonder why god doesn’t “just kill them off”.

  26. keiths:

    I don’t know Steve very well, but I’m surprised to see Joe being so doctrinaire.

    There is no single, canonical definition of “natural selection” among evolutionary biologists or scientists generally.

    Joe:

    There is when you use population genetics.

    Your claim was far broader:

    My point is that you can define “natural selection” as you wish, but that is not binding on evolutionary theorists, who use it my way.

    That’s simply wrong. Not all evolutionary theorists use your definition, Joe. As I’ve already demonstrated.

  27. Joe:

    So no, Archimedes would not have been able to do this, as simple as the mathematics would be.

    Come on, Joe. Archimedes would have had absolutely no trouble with this:

    There are ten of A and ten of B. Every two minutes, each A gives birth to another A. Every one minute, each B gives birth to another B. No A or B ever dies. After an hour, will there be more A’s or B’s? After one year? After one century?

    You don’t need the quantitative theory of evolution to figure it out.

    Ernst Mayr was not a deep thinker when it came to population genetics.

    If true, so what? He was an evolutionary theorist (and a well-known one) and his definition of natural selection differed from yours, contrary to your claim:

    My point is that you can define “natural selection” as you wish, but that is not binding on evolutionary theorists, who use it my way.

    Joe:

    In any case, I do not see in those quotes anything that is incompatible with the population-genetic definitions of fitness.

    Think about my scenario. There is no death, which means there is no elimination, which means there is no selection according to Mayr’s definition. According to the pop gen definition, there is selection. The definitions are incompatible when applied to my scenario.

    Waving one’s hand and dismissing the mathematical theory is easy.

    I’m not dismissing the mathematical theory at all, nor am I suggesting that it needs to change. I’m simply pointing out that the pop gen definition of selection doesn’t automatically trump all other definitions.

    By all means, continue to use the definition you find most convenient for your work, but don’t pretend that it is the only right definition or the only scientific one.

  28. keiths:

    Joe:

    [Re: Mayr]

    If true, so what? He was an evolutionary theorist (and a well-known one) and his definition of natural selection differed from yours, contrary to your claim:

    Joe:

    Think about my scenario. There is no death, which means there is no elimination, which means there is no selection according to Mayr’s definition.According to the pop gen definition, thereis selection.The definitions are incompatible when applied to my scenario.

    Just goes to show Mayr was prone to make population-genetic howlers.

    He forgot that two genotypes that had the same mortality, in this case none, but differed in fertility/fecundity thereby had different fitness.

    Just goes to show that the particular definition of fitness that he used, and you use, is not useful.

  29. Joe:

    Just goes to show Mayr was prone to make population-genetic howlers.

    He wasn’t talking about population genetics — he was talking about natural selection, which is a concept shared between population genetics and all of the other evolutionary subdisciplines.

    He forgot that two genotypes that had the same mortality, in this case none, but differed in fertility/fecundity thereby had different fitness.

    Huh? He doesn’t even mention fitness in that quote:

    Natural Selection is Really a Process of Elimination

    The conclusion that these favored individuals had been selected to survive requires an answer to the question, Who does the selecting? In the case of artificial selection, it is indeed the animal or plant breeder who selects certain superior individuals to serve as the breeding stock of the next generation. But, strictly speaking, there is no such agent involved in natural selection. What Darwin called natural selection is actually a process of elimination. The progenitors of the next generation are those individuals among their parents’ offspring who survived owing to luck or the possession of characteristics that made them particularly well adapted for the prevailing environmental conditions. All their siblings were eliminated by the process of natural selection.

    Joe:

    Just goes to show that the particular definition of fitness that he used, and you use, is not useful.

    You’ve lost the thread completely. I haven’t offered a definition of fitness. Mayr’s quote doesn’t mention fitness. We’ve been talking about natural selection, Joe.

  30. Elizabeth: Have you actually read On the Origin of Species by Means of Natural Selection Gregory?

    Now that the question has been answered in the affirmative, anyone want to actually face what Darwin himself said about NS, which he would have changed to NP? Funny the equivocation thread in contrast…confident Darwin groupies who promote false Darwin.

  31. jeez Gregory, we are talking about science here!

    Are you going to attack cosmologists who promote false Newton?

    The validity of evolutionary theory is not to be found in a close textual reading of Darwin, but in the development operationalisation and testing of the theory.

  32. “jeez Gregory, we are talking about science here!”

    No, Lizzie, we’re talking about LIFE. Do you actually wish to deny Darwin wrote that letter to Lyell?!

    Schizophrenia studies, music & architecture won’t save you from committing a direct answer.

    Is this thread really mainly about “the validity of evolutionary theory”?

  33. Mung:

    Do you make healing the same thing as “forcing something to be perfect?” If not, that was quite an “evolution” you made to hotshoe’s remark.

  34. Mung,

    BTW Allan, I had a similar discussion over at UD not to far back in which I questioned how you could tell the effects of drift from the effects of selection, which Elizabeth then started a thread about here.

    I was maintaining that people were conflating neutral evolution and genetic drift and arguing that even mutations under selection were not immune to genetic drift. So if I read you correctly you would agree.

    Yes, I would completely agree.

    So how does one distinguish between the two?

    You only get an overall stat, so you can’t strictly distinguish. If you think of the kinds of things that could be selected, they only matter when they matter. So if you escape from a lion, your ‘escape’ genes get a leg-up, and are more likely to persist. But the rest of the time, your escape genes aren’t being tested, and random factors prevail. But also, your ‘escape’ genes get a leg-up every time some slowpoke is eaten, even if you aren’t involved in that transaction. So, the final s value is always an integral of bias and drift.

    This doesn’t mean you can’t spot selection, however. For instance, one can get a ratio of the amount of synonymous to nonsynonymous substitution in codons. The rate of substitution of the latter gives a baseline for drift (with caveats), and the former is the departure from the neutral expectation.

    Neutral alleles form the baseline, and you can calculate the expected time for them – but this is a probability distribution, so data for a ‘real’ neutral allele could legitimately appear selected. From theory, as soon as you depart from neutrality, two things happen: population size becomes a factor, and expected times to fixation shift. The further your measurements depart from the baseline, the more confidence you can have that selection, not visiting the extremes of the distribution (which must happen, for it to be a distribution), is the deciding factor.

  35. keiths:
    Joe:

    He wasn’t talking about population genetics — he was talking about natural selection, which is a concept shared between population genetics and all of the other evolutionary subdisciplines.

    Huh? He doesn’t even mention fitness in that quote:

    Joe:

    You’ve lost the thread completely. I haven’t offered a definition of fitness. Mayr’squote doesn’t mention fitness.We’ve been talking about natural selection, Joe.

    (I am a bit hampered by WordPress, which loses quotes-within-quotes when it generates a reply quoting you).

    Anyway, sorry for the inattentive reply. OK, let’s talk about Mayr’s definition of natural selection (not fitness).

    Apparently it doesn’t include differences in fertility (or fecundity)? Just in viability?

    So if genotype A and genotype B have the same viability, and then genotype A comes to have higher fecundity than genotype B, we expect genotype A to increase, but Uncle Ernie wouldn’t call that natural selection? Hmmm.

  36. Lizzie,

    I’m not confused. I’m suggesting that you are.

    What specifically do you think I’m confused about, and why?

  37. Joe:

    So if genotype A and genotype B have the same viability, and then genotype A comes to have higher fecundity than genotype B, we expect genotype A to increase, but Uncle Ernie wouldn’t call that natural selection?

    If they are perfectly viable (as in my scenario), with no individuals being eliminated, then his definition clearly precludes selection:

    Natural Selection is Really a Process of Elimination

    The conclusion that these favored individuals had been selected to survive requires an answer to the question, Who does the selecting? In the case of artificial selection, it is indeed the animal or plant breeder who selects certain superior individuals to serve as the breeding stock of the next generation. But, strictly speaking, there is no such agent involved in natural selection. What Darwin called natural selection is actually a process of elimination. The progenitors of the next generation are those individuals among their parents’ offspring who survived owing to luck or the possession of characteristics that made them particularly well adapted for the prevailing environmental conditions. All their siblings were eliminated by the process of natural selection.

    So yes, a man who is considered one of the greatest evolutionary biologists of the 20th century disagrees with you on the meaning of “natural selection”.

    Now do you see how silly it was for Steve to write this…

    It’s not binding on everyone, of course — only on those who wish to discuss evolutionary biology scientifically.

    …and for you to write this?

    My point is that you can define “natural selection” as you wish, but that is not binding on evolutionary theorists, who use it my way.

  38. keiths:
    Joe:

    If they are perfectly viable (as in my scenario), with no individuals being eliminated, then his definition clearly precludes selection:

    So yes, a man who is considered one of the greatest evolutionary biologists of the 20th century disagrees with you on the meaning of “natural selection”.

    Now do you see how silly it was for Steve to write this…

    [somehow WordPress deletes this in my reply — JF]

    …and for you to write this?

    [likewise]

    Well I’ll just leave it with these points:

    1. I shook Ernst Mayr’s hand once.

    2. I corresponded with him over his request that I read the chapters on phylogeny methods in his (and Peter Ashlock’s) 1991 revised version of Principles of Systematic Zoology. I gave him lots of corrections, and he thanked me and acknowledged me in the book.

    3, I have many times talked to other theoretical population geneticists about Ernst Mayr’s misunderstandings of theoretical population genetics. We all agreed on that.

    So I have some perspective on Mayr.

    If that’s his definition of “natural selection”, I disagree with it.

    And so should you.

  39. Joe:

    If that’s his definition of “natural selection”, I disagree with it.

    Obviously, although why you think there can only be one scientifically valid definition is beyond me. This is hardly the only example of a technical term that has different definitions depending on the context.

    And so should you.

    Why?

  40. keiths: Why?

    Because science is critically dependent on effective operational definitions.

    The definition that emerged from population genetics does not, in my view, contradict Darwin’s – what it does is show that the scenarios Darwin envisaged were a special case of something that could be formulated more generally.

    In discussing the theory of natural selection, it makes no sense to ring-fence it with boundaries that turned out not, in fact, to be boundaries.

    Darwin’s idea was even more brilliant than he realised, basically, and capable of being expressed in formal mathematical terms.

    That formalisation continues to be refined to take account of new discoveries, including genetics, which Darwin didn’t even know about.

  41. Lizzie,

    Because science is critically dependent on effective operational definitions.

    Sure, but Mayr’s definition is hardly “ineffective”. It just differs from the pop gen definition.

    The definition that emerged from population genetics does not, in my view, contradict Darwin’s – what it does is show that the scenarios Darwin envisaged were a special case of something that could be formulated more generally.

    The pop gen definition is more inclusive, so it only contradicts less-inclusive ones in the areas of non-overlap.

    In discussing the theory of natural selection, it makes no sense to ring-fence it with boundaries that turned out not, in fact, to be boundaries.

    The boundaries are created by the definitions. The pop gen boundaries enclose more but are no more real than the boundaries created by Mayr’s definition.

    Later today I’ll elaborate on why I think Mayr’s definition is the appropriate one to use in the context of this thread.

  42. The Mayr quote refers to NS as “What Darwin called natural selection”. He is not offering his own, but adopting Darwin’s.

    I share a degree of keiths’s reservation that population genetics needn’t have the last word on how to look at evolution. I have argued in the past against evolution being defined as “change in allele frequency”, when change in lineage is clearly important too. The get-out that a new mutation is a change in frequency too isn’t very satisfactory. But … I’m not that bothered about it either!

  43. Allan Miller: The get-out that a new mutation is a change in frequency too isn’t very satisfactory.

    How does mathematics define the change in frequency from zero to non-zero?

    The allele frequency definition reminds me of the definition of a human: a featherless biped with broad flat nails.

    All this strikes me as apologetics. Quibbling over semantics.

    Evolution in its broadest sense is change in populations over time. Most of the heritable change is in the genome, but there are organelles that could, at least in theory, evolve.

    I think it would be more productive to discuss how and why alleles become prevalent or fixed. I would like to see a Larry Moran thread that discusses his assertion that most alleles fix as a result of drift.

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