“Selection pressures” are metaphors. So are the “laws of physics.”

September 29, 2013 • 8:10 am

I mentioned offhand the other day that the notion of “selection pressures” in evolutionary biology is a metaphor, not dissimilar to the metaphor of a “selfish gene,” but Matthew convinced me that this idea was more profound than I thought (LOL!), so I’ll write a bit more about it.

A quick refresher: the other day Andrew Brown wrote one of his usual muddled columns in the Guardian, claiming, as he often does, that Richard Dawkins is a malign influence on evolutionary biology.  His argument was the usual blather that gene aren’t really “selfish,” and that this metaphor has led to deep confusion.  In the comments after his piece, Brown repeats this claim in response to a criticism (h/t to moleatthecounter for finding this):

Picture 1If you can’t read the above because the print’s too small, Brown says that “. . . Dawkins got confused by his own title. This confusion was intermittent but it’s absurd to pretend it didn’t happen.” (He also give props to Mary Midgley, who deeply misunderstood the book, attacked Dawkins in a quite vitriolic way, and got a strong response from Richard. Although the original papers don’t seem to be on the Internet, you can read about that controversy here. [Note: reader pacopicopedira has found Dawkins’s reply online; he/she gives the link in the comments but you can read it here.] If you think that Richard’s responses to Midgley were “disgusting”, try reading what she wrote in the first place.)

At any rate, Brown, fulminating about a metaphor that supposedly misled not only Dawkins’s readers, but Dawkins himself, said the following in his Guardian piece:

In particular, the ascription of agency to genes led him and his followers into endless confusion. The point is not merely whether genes can be selfish or generous, but whether they can be said to have any activity at all in the world. This is a point which he freely concedes and then forgets – his manner of dealing with most criticism. If a gene is defined, as he defines it, as a piece of chromosomal material subject to the pressures of selection, it is the pressures of selection which are the active and changing parts of the picture, and the DNA sequence is entirely passive.

Sadly, here Brown is hoist with his own petard. What he doesn’t realize is that there is no such thing as “the pressures of selection”—it is a metaphor, a descriptor of what happens when different genes (i.e. “alleles”, or forms of a single type of gene) leave different number of copies.   That differential reproduction of genes is what constitutes natural selection,, and it is a process of gene sorting.

There are no “pressures” of selection imposed on the organism from the outside. What happens, as everyone knows who learns introductory evolution, is that, in a given environment, some genes leave more copies than others, usually because they increase the reproductive output of their possessor.  Take, for example, a population of brown bears that somehow find themselves in a white-colored environment, like the Arctic.  There are genes affecting coat color, and imagine that a given gene comes in several forms, one of which makes the bear lighter in color than do the alternative forms.  This being a population of bears, there will be variation among their genes due to mutation. Those bears carrying the “light” forms of genes might do better than their browner confrères because they’re more camouflaged in the snow, and thus better at sneaking up on seals and killing them.  “Light-gene” bears will be better fed, and thus have better survival and (crucially) more offspring. (If the color change affects survival but not reproductive output, no natural selection ensues.)  In the next generation, the proportion of color genes having the light form will be higher than before. And the average color of the bear population will be a bit lighter.

If this continues over many generations, and other mutations occur that yield even lighter coats, natural selection will move the bears from brown to white. Presumably this is what happened in polar bears, whose ancestors were probably brown. And it’s happened in many Arctic animals whose ancestors were brown but evolved white color (either pemanently or seasonally) via natural selection. Such animals include the Arctic fox, the Arctic hare, the ptarmigan, the snowy owl, the harp seal, and so on (see a list here).

What “selection” pressures can do: white Arctic mammals.

Note that the environment isn’t exerting any “pressure” here. It is simply providing a milieu in which one gene has an advantage over another. The environment cannot see the genes and their constituent DNA. We speak of “selective pressure to become light-colored” as simple shorthand for the process I’ve described above.

Now I suppose people like Brown could say, “But that’s confusing! Saying that the environment exerts selective pressures could mislead people into thinking that the environment is in someway animate, and can exert a force on animals to mold them one way or another. It could lead to vitalism!”  But it’s not really confusing. In fact, it’s so not confusing that Brown used the metaphor himself, without realizing it.  When I teach my students introductory evolution, I’m always careful to note that “selection pressures” is a shorthand term for something a bit more complicated.

But if Brown can easily recognize this metaphor, so can everybody else, and the notion of “selection pressures” has not been deeply confusing. Likewise, “selfish gene” is not confusing if you have sufficient neurons to see that Richard what Dawkins meant: the differential sorting of genes that is natural selection involves genes behaving as if they were selfish: trying to outcompete their mates and get into the next generation of bodies. Apparently Midgley had such a problem.  But Dawkins certainly did not, despite what Brown claims.

Metaphors are useful if as they enlighten rather than confuse. I claim that both “selection pressures” and “selfish genes” are enlightening, and, judging by the sales of Dawkins’s book, so do most readers.

Let me add, since I’m writing about physics for my book now (kudos to Sean Carroll, my Official Physics Tutor™), that the term “laws of physics” is also a metaphor, or a shorthand descriptor.  There are no “laws of physics,” but just regularities in the universe that appear to be ubiquitously “followed”.  People like Brown might argue that the term “laws” implies “lawgiver” and hence God—a confusing misconception unwittingly promulgated by atheistic physicists.  But again, this metaphor is not confusing—except, perhaps, to theists who want to see those laws having been decreed by God.

As one reader pointed out yesterday, metaphors, often involving the use of anthropomorphizing, are ubiquitous in all of science, not just evolutionary biology. “Selfish gene” is just one of these.  I would argue that anyone who can’t understand, after a minute’s explanation, that selfish genes really aren’t conscious and malevolent entities, is lacking some crucial rationality.  Likewise, it’s not hard to see how “selfish genes” can lead to cooperative behavior.  That, too, is easily explained, and has been done many times by Dawkins and others. Nevertheless, people like Midgley and many theologians argue that the notion of selfish genes simply can’t explain unselfish behavior.  They’re ignorant—often willfully so.


99 thoughts on ““Selection pressures” are metaphors. So are the “laws of physics.”

  1. I’m inclined to agree with your wilfull ignorance hypothesis. The selfish gene metaphor is very easy to understand – if you’ve actually read the book.

    1. Agreed. I read the book and found it to be rather insightful, especially from Chapter 3 onward. The “selfish gene” metaphor is difficult to misunderstand if one has read at least half the book.

    2. Yep, I agree too.

      Certainly there can be good metaphors (good in the sense of helping get a point across more effectively) and bad metaphors. But I tire of creationists and pedants objecting to any use of metaphor in biological language on the basis of biological mechanisms not being conscious agents. We know they aren’t. With the exception of a few woomeisters, everyone knows this; get the frak over it and focus on understanding the gist of what the speaker is trying to say, rather than complaining about how they say it.

    3. Agreed. I’m a software engineer with a love for science. I readily understood the metaphor when I read “The Selfish Gene”. So I can only conclude that people like Andrew Brown are evincing willful ignorance given that they otherwise appear to be intelligent (thus negating the stupidity hypothesis).

      1. hey, greetings, dear colleague. But you should note that our domain of activity has been plagued by crappy metaphors :
        _Building a bridge(shitty consultants, completely different kind of job)
        _Chain production(even shittier consultants, even worse, as when a software is done, you don’t need to fabricate another one, you just copy it or wrap it for external calls)
        _Gardening(Jeff Atwood, not that bad, but still not convincing).
        _Cooperative game(Alistair Cockburn, rather good, but still not perfect).

        In brief : a metaphor is just an entrance door. Interesting things lie behind(and the theory behind the cooperative game is really excellent). Only fools lie attacks on the metaphor.

    1. Thanks for finding that! As usual, the most vocal of the willfully ignorant are those whose meal ticket depends on their willful ignorance.

  2. “There are no “laws of physics,” but just regularities in the universe that appear to be ubiquitously “followed”. People like Brown might argue that the term “laws” implies “lawgiver” and hence God—a confusing misconception unwittingly promulgated by atheistic physicists.”

    An analogy is that Andrew Brown is an irregularity in the universe. We need to come up with a metaphor for that. 🙂

    1. I think that the Laws of Physics imply the existence of a democratically elected parliament of Metaphysics.

      And I demand rigidly defined areas of doubt and uncertainty.

  3. Andrew Brown regularly attacking Dawkins’s book like this has just become pathetic. Give it up Andrew. Dawkins’s book is successful and we get that “selfish” is a metaphor. Just stop it now.

    Good grief! I read about evolution as a child in my children’s books about prehistoric animals, anthropology and more and I didn’t get confused — and I was A CHILD! I can see how people could over think something and get muddled up but honestly, if they’re rational at all, they’d work their way out of the confusion. This whole thing is just silly. I’m glad you brought up the laws of physics because when physicists say things like something is “forbidden” you just know someone is going to be obtuse enough to think someone is forbidding it.

    BTW the cute white animal pictures were squee worthy!

    1. I was just thinking about this recently…I learned a little about evolution as a small child, through books (not very child-oriented, but good enough), museum visits, and parents answering my questions.

      As for the white animals: I’d add the beluga whale and the snow bunting (a small sparrow-like bird).

      1. My thoughts on the white animals were “polar bears are known to interbreed with grizzlies in zoos and (more rarely) in the wild” ; and “Scottish hares are a good case ; white fur in winter, brown in the rest of the year ; and woe betide the hare that changes coat at the wrong time!”

    2. And in particle physics in the vacuum of quantum fields everything is allowed unless expressly “forbidden” by symmetries – but that merely means such events are really, really improbable, not impossible. (Seems fluctuations can do anything, given enough time.)

  4. My graduate degree is from a music conservatory, so I have to dig a bit to retrieve memories of science classes, but one thing that really made an impression was a chemistry professor, at the end of the course, saying that most of what she’d taught us was a metaphorical description devised for ease of understanding (!) and that in reality, things are more complicated.

    Reality is weird.

    1. Not only is reality weird, it is the thing of which our metaphors are made to describe. All thoughts are metaphors representing what we are thinking about. It appears that the Guardian columnist doesn’t appreciate this fact.

  5. There are no “laws of physics,” but just regularities in the universe that appear to be ubiquitously “followed”.

    Of course there are laws of physics.

    There are no laws of nature, so “laws of nature” could be said to be a metaphor. However, for laws of physics, the physicists who came up with those laws are the law givers.

    1. Well, I would say that depends on how you interpret the word “law”. If you take human laws (the laws of the land) as giving the one paradigmatic, literal meaning of the word, then there are no laws of physics in that sense. The “laws” of physics don’t dictate to nature how she should behave! Physicists don’t play a role analogous to that of legislators.

      But I would also say that the distinction between literal and metaphorical meaning is a fuzzy one. Words can have more than one literal meaning, and the scientists’ meaning of the word “law” could be considered a second literal meaning.

    2. My favorite way of making this distinction is that there are “descriptive laws” and “proscriptive laws.” The laws of nature are descriptive and the laws governing human behavior are proscriptive.
      If our knowledge of an aspect of nature were to change, scientists would then be motivated to change the relevant law to reflect the new description.
      Before science (a few centuries ago when the Church was in charge), there was no such thing as a descriptive law. This is why Anselm made some poor assumptions.

  6. I must say that of all the “metaphors” bandied about in science the “selfish gene” metaphor is perhaps the most appropriate and most representative of reality. The mathematical description of evolution – Evolutionary Game Theory – rests on describing the “strategies” of alternative competing entities. These strategies are essentially algorithmic, defining how something behaves in differing competitive conditions (which in turn produces various “fitness payoffs”). Evolutionary Game Theory absolutely abounds in using similar descriptive terms (metaphors) for many various strategies –Hawk, Dove, Prober, Bourgeois, Assessor, Retaliator etc. etc. Rather that using an incredibly longwinded algorithmic descriptor (e.g. Strategy= “for iteration n use an asymmetry of the environment [typically possession] to determine if action is fight [possession=yes] or flight [possession=no]) the whole strategy is summarised by the term BOURGEOIS. Well, after all, this strategy is very …. very.. bourgeois! Now underlying ALL genetic strategy, at a base genetic level, is SELFISH – the genes promotion of it’s own individual reproductive success. This can, as Jerry points out, lead in certain Game circumstances to co-operation and even altruism at some higher level of organisation – e.g. the phenotype. But this does not negate the SELFISH base itself.

    As for Selection Pressure this too has a mathematical equivalence in summarising the overall dynamic effects from the “payoffs” in the payoff matrix.

    Yes..any criticism of using such metaphors is a certain sign of gross or willful ignorance of the subject. I can’t imagine anyone can retain that level of ignorance if they actually read “The Selfish Gene”.

    1. Thank you, hkornstein, for pointing me to the Pinker article arguing that Dawkins might have carried the selfish gene metaphor even further. I won’t attempt to reproduce Pinker’s arguments, but point readers again to the pdf, “Yes, Genes Can be Selfish”. It’s short and extremely tightly reasoned. Basically Pinker is looking at evolution from various vantage points — information, molecules, mind/brain, organism.

  7. It’s rather sad that Andrew Brown, who clearly doesn’t understand evolutionary theory, gets such a loud megaphone with which to erroneously criticize someone who does. Surely in any just world, a reasonable newspaper would get a biologist to critique another biologist’s theories?

    1. The Guardian a “reasonable newspaper”? I think not.

      I read the Guardian’s website daily but it’s the last one I read. Not as bad as Faux Noise, but not too far from being a bad ideological joke where the facts of every newsworthy situation are coerced into fitting a moderately Marxist point of view.

      1. The Guardian has great cryptic crosswords. I particularly like the ones by Araucaria. But since I discovered you can download the crosswords from the web site I’ve stopped buying the paper!

  8. It would be interesting if anyone can point out, in the genre of science fiction,a sentient, artificial intelligence (alien, super-computer, human-made life form, etc.) whose primary concern is not self-preservation.

    1. Well, there’s the smart bomb in the movie Dark Star, whose overriding mission is to blow up.

      Similarly, the Terminator robots are more mission-oriented than survival-oriented.

      Intelligent eusocial aliens, such as the Bugs in Heinlein’s Starship Troopers, would presumably put the welfare of the hive above their own individual survival.

      In a somewhat different vein, writers from John Varley to Greg Egan have depicted futures in which people (natural or artificial) routinely back up their brain states to protect against death of the body. So self-preservation means something completely different to them than it does to us.

    2. Um…hello? Asimov’s Three Laws?

      Self-preservation is at the bottom of the list.

      Of course, most of the dramatic interest in the stories comes from fiddling with the laws, including at least one case where the robot’s self-preservation takes precedence over all else, but that’s just part of a good murder-mystery.



      1. & I think you are referring my favourite Asimov robot short story” Little Lost Robot. I loved how they got the robot to reveal himself.

      2. I wonder how many budding SF writers have put their first manuscript into the envelopes for posting, then retired with a cup of cocoa and “I, Robot” … and thrown the manuscripts into the shredder, un-posted.
        There has probably been better SF written since, but The Good Dr is a terribly imposing standard to compare yourself with. (Which is part of the reason that I subscribe to a short-story SF magazine ; to give the next generations of writers somewhere to fledge.)

    1. Thanks for the link! I think Dawkins was very patient and polite in the way he defended himself, as I’ve always found him. It’s truly laughable to characterize his response as “disgusting”. Shame on you Brown!

      I did like that Dawkins managed to get in some nice (not disgusting) zingers, my favourite being:

      One cannot, after all, be expected to read every single word of a book whose author one wishes to insult.

      Think how irritating it must’ve been to have to take the time to defend your work because someone was being such a loudmouth that it could confuse people. The passages Dawkins quotes are indeed awful both for misunderstanding really obvious things (like the Chicago gangster part) and just being insulting – like actually calling his “typical readers” egoists:

      people with vaguely egoist leanings about individual human psychology – willing to follow him in losing touch with the observed facts of motivation altogether and taking off for the empyrean with the Gene’

      LOL! How can anyone take such a reviewer seriously?

  9. At the risk of fogging Jerry’s neat take-down of Andrew Brown’s nonsense…

    The distinction between literal and metaphorical meaning is a fuzzy one. Some of the word meanings we think of as literal probably began their life as metaphors. But the original literal meaning has gone out of use, leaving only the metaphorical one. If we don’t realise or don’t pay attention to the fact that there was once another meaning, we are likely to think of the newer meaning as a literal one. And there’s nothing wrong with that. Languages change gradually over time, and there’s no particular moment when a once-metaphorical sense becomes a literal sense. Should we forever consider “computer network” to be metaphorical, just because “computer” once referred to a person who computes, and “network” once referred (I believe) to fishing nets?

    We could say that in its literal sense selection requires a selector (some kind of intelligent system with intentions), and that therefore the evolutionary biologist’s usage of the word should be considered metaphorical. But we could also describe what evolutionary biologists have done as extending the sense of the word, making it broader. I think Daniel Dennett would be inclined to accept this latter description, as ISTR that he sometimes talks about such usages being “extended senses”. To be clear, I’m not saying that one description or the other is the right one. They are equally valid ways of describing the same thing.

    1. The problem Richard, is that we are humans, and not computers. Our brains have not evolved to think in pure abstract mathematical constructs or symbology which can EXACTLY represent reality. We often think, and understand in terms of physical or emotional symbologies that we are evolutionarily more familiar with and more adapted to handle, which exist in our human environment. We anthropomorphise. Nothing wrong with that if the analogies to the mathematical constructs are accurate(as with “selfish genes”). It helps our understanding far more than struggling with the more mathematically abstract construct. Why knock something that aids understanding, if it mimics mathematical reality so well?
      Why knock it, indeed.

  10. An interesting example of possible selection for white (or whiter bears) was shown a few years ago in National Geographic. Lovely pictures and a summary can be found at Spirit Bears These are a population of brown bears that are isolated on islands in British Columbia. The population has an elevated frequency of a recessive mutation that makes them whitish. It is suspected that the white color makes them less visible to their fish prey.

    1. Do they hibernate? There might be a link between hibernation/activity in winter period or maybe just availability of food… Is the summer the ‘fallow’ period for polar bears?

  11. Metaphors have heuristic value and are therefore informative (useful). Danger lies where metaphor belies reality. Complex, adaptive systems, like evolutionary biology, often use metaphors for communicative purposes, and are occasionally adapted as the reality. An instance on which I will not elaborate here, is the adaptive landscapes used as metaphors to demonstrate variable fitness in greater than 2 dimensions. I therefore do not have a problem with the “selection pressures” or “evolutionary forces” we assign to genotype-environment interactions.

    It has been about six years since I last read “The Selfish Gene,” and like many older texts, it was out of cultural context and I was already aware of the arguments from my previous training. I do have in my notes, however, a statement similar to one of Brown’s points. In his antepenultimate paragraph he describes assignment of agency in the title, and I agree with that as well as anthropomorphizing any molecule. I obviously didn’t feel as strongly about it as Brown to include that in a public essay, but I agreed nonetheless. I interpreted the title as being what Dawkins has built his popular career on–being provocative. Some people are like that; I am not, but I don’t feel enough conviction to judge.

  12. Some other things to have philosophical debates about:

    0) Genes cannot be selfish.

    1) There are no holes in a black hole.

    2) The sun is not really very “attractive” for a comet that falls into it.

    3) The Kuiper belt can’t be worn.

    and so on and so forth. Or, perhaps, the “genes are not selfish” people could just read George Lackoff.

  13. Brown is a knob. He not only doesn’t understand science but he doesn’t even understand metaphor, something that a writer should really be well-acquainted with.

    When we say a cation is attracted to an anion we know it’s a metaphor and the ions don’t have cognitive functions.

    I vote for Brown being partially willfully ignorant but also a victim of selective understanding based on his emotional animus toward Dawkins (just my opinion).

  14. Jerry:

    the term “laws of physics” is also a metaphor, or a shorthand descriptor. There are no “laws of physics,” but just regularities in the universe that appear to be ubiquitously “followed”. People like Brown might argue that the term “laws” implies “lawgiver” and hence God—a confusing misconception unwittingly promulgated by atheistic physicists.

    Your statement of what physical laws are is entirely correct, however I’m not so sure the term “laws of physics” is a metaphor, since I don’t think physicists have ever intended the term to imply a lawgiver, even a metaphorical one.

    Instead, the physicists use of “law” is a primary meaning of the word. E.g. Oxford Dictionaires: “Law: 3: a statement of fact, deduced from observation, to the effect that a particular natural or scientific phenomenon always occurs if certain conditions are present.”

    There is nothing metaphorical about that. (My own take on the matter at: “What are laws of physics?”)

    1. “Law” in “law of physics” has two meanings – 1) an objective pattern of sufficient scope and level to be considered in the domain of physics, or 2) a partial reconstruction of said pattern propsitionally.

  15. How could someone who claims to have read the bible claim that metaphors are misleading because some people might take them literally?!

  16. The really stupid thing about Brown’s claim that Dawkins literally interpreted his own metaphor is that anyone who knows the molecular basis of genes and biochemical mechanisms of heredity couldn’t possibly be confused about whether “selfishness” is used metaphorically. Well, almost anyone (not Andrew Brown!). Just as you said, Jerry, you only have clarify that “selective pressure” is a metaphor to introductory students, to whom the mechanisms of heredity and natural selection are not old hat – anyone cognizant of the basics knows what you mean when speak of “selective prsssure” or “selfish genes”.

  17. For people like Brown it’s too hard to understand that a word like “pressure” can mean one thing in one context (natural selection) and something else in another (say a pressure cooker).

  18. “There are no “laws of physics,”
    Rather you will be rewarded with say the Nobel Price when you design an experiment that shows that two bodies with mass can repel each other – thus violating several laws of physics.

  19. Both Brown and Midgely have agendas and this drives them to make mistakes which they then cannot acknowledge because the mistake is the bais of their argument and, without it they have none.

  20. Brilliant article!

    I would just point out that I think there is one problem with the otherwise very useful metaphors that good science writers employ, and that is that they encourage teleological thinking. That is, they suggest an ultimate purpose to what happens when really the world’s history could just be summed up by ‘Shit happening’, not ‘Someone or something did some shit’.

    Incidentally, I think I found Midgley’s original, courtesy of the Wayback Machine:


    Dawkins’s reply on his old website, with a link to a pdf version:


    And Midgley’s reply to that:


    1. I had a comment eaten, but I will try again to just make a small point about the use of metaphors. They can be incredibly revelatory; one reads a good one and the truth of the natural processes being described hits one between the eyes.

      But there is one problem, I think; they can encourage teleological thinking (and we are prone to teleological thinking anyway). That is, the history of the world could be summed up as ‘Shit happening’, not ‘Someone or something did some shit’. Many metaphors, like the two discussed above, suggest the second, when the truth is as the article describes, the first.

      Of course, I think many people find it hard to believe that all this could just happen, but I guess that may ultimately be the difference between believers and non-believers.

  21. Theories of science as not predicated on metaphors! Both Darwin & Wallace came up with the same term to express evolutionary theory causation, “natural selection”. “Selective pressure” is simply an alternative way to express “natural selection” NS. As Darwin noted selective breeding mimics natural selection but the “pressures” here are conscious not naturally unconscious.

    While both survival & reproduction are internal organism mechanisms that provide something to select on ie the SELECTEE, outside forces are required and provided by the environment ie the SELECTOR. Organism groups may act as a selector. This process became known as the “Baldwin effect” & was proposed in Darwin’s era.

    Because of the “revised central dogma” no gene is directly selectable ie each gene can only be selected via its polypeptide phenotype. More than just the one coded by are used to form proteins. Thus a minimal MODEL SELECTEE comprises four alleles situated on two loci not Dawkins single gene. This has a profound effect on Dawkins selfish gene model predicated entirely on Hamilton’s Inclusive Fitness since it requires genetic epistasis e, to now be included in Hamilton’s Rule

    rb>c ……(1)

    The new epistatic variable e must be minimally set to e=2 making the rule:

    (r^e)b>c …(2)

    Now Haldane requires 4 brothers related 0.5 IBB via (2) to die for not just the two brothers as calculated by (1). As e increases the rule & selfish geneism becomes inoperable.

    1. No…. you are splitting hairs here.
      What we are dealing with is the lowest level of selection, not the lowest level of subdivision (why not go all the way down to quarks?) This lowest genetic level is strategically “selfish” or none of the maths works. BTW On the subject Hamilton’s rule -the rule as stated is verified by field evidence
      Or perhaps you would suggest that the title of Dawkins book should have been “The Selfish Alleles on Two Loci”

      1. DNA segments were & remain unelectable (as previously outlined). DNA was and remains 100% protected from direct DNA selection via the revised central dogma which only minimally allows polypeptides to be selected. Proteins that form selectable phenotypes are complex polypeptide composites requiring many more than just the one “selfish” allele to code for them. As previously stated this basic fact requires genetic epistasis to be included not excluded from Hamilton’s Rule. This means at least two loci with a total of four alleles, (not just Dawkins single allele) are required as a model selector. Dawkins continues to misuse Hamilton’s tautological model in which cause & effect reverses.

        1. You’re wrong here; single genes with two alleles or more can experience natural selection. And we have many examples, including lab experiments on E. coli.

          And I think you’re misunderstanding epistasis as the interaction between products of different genes and epistasis as a two (or more)-locus model in which two (or more) interacting genes each have two alleles. Selection can operate in both cases, but is not, contrary to your claim, limited to the latter case.

          1. E. coli genes are dependently selected via competing, single, E. coli cells containing one entire genome. A minimal model is therefore, one hypothetical cell containing two loci with four alleles ie not just one fitness “selfish” allele quietly forgetting about the coded polypeptide via which it is selected. Epistasis dependently combines the fitness of at least two loci. Proposing that single alleles can be independently selected is just a misused, simplified/oversimplified model. The contradiction I previously explained stands as proof: while we are related over 95% NON IBD to chimps, employing Hamilton’s gene centric IBD we are not related. Dawkins et al cannot have it both ways.

            1. Sorry but you don’t know what you’re talking about. One can put genes into bacteria (like antibiotic resistance genes) and watch selection operate based on differences at single loci. You have given no reason why organisms differing in a single gene can’t undergo selection, and there’s lots of evidence that two varying genes aren’t required (lactose tolerance in humans, color in B. betularia, etc.) I’m sorry, but your ignorance of evolution is deep here, and I can’t let you continue to make misleading statements without any evidence. Neither theory or evidence supports your “more-than-one-gene” model.

  22. Theories of science as not predicated on metaphors! Both Darwin & Wallace came up with the same term to express evolutionary theory causation, “natural selection”. “Selective pressure” is simply an alternative way to express “natural selection” NS. As Darwin noted selective breeding mimics natural selection but the “pressures” here are conscious not naturally unconscious.

    While both survival & reproduction are internal organism mechanisms that provide something to select on, i.e. the SELECTEE, outside forces are required and provided by the environment i.e. the SELECTOR. Organism groups may act as a selector. This process became known as the “Baldwin effect” & was proposed in Darwin’s era. It must not be confused with the organism group acting as a selectee (group selection).

    Because of the “revised central dogma” no gene is directly selectable i.e. each gene can only be selected via its polypeptide phenotype. More than just the one polypeptide coded by one DNA “gene” are used to form proteins. Thus a minimal MODEL SELECTEE comprises four heritable alleles situated on two loci not Dawkins single heritable gene. This has a profound effect on Dawkins selfish gene model predicated entirely on Hamilton’s Inclusive Fitness since it requires genetic epistasis (e), to now be included in Hamilton’s Rule:

    rb>c ……(1)

    The new epistatic variable (e) must be minimally set to e=2 making the rule:

    (r^e)b>c …(2)

    Now Haldane requires 4 brothers related 0.5 (IBD) via (2) to die for not just the two brothers as calculated by (1). As e increases the rule & “selfish geneism” becomes inoperable.

    Hamilton’s gene-to-gene relatedness probability known as “Identical By Decent” (IBD) cannot validly replace adult organism to adult organism relatedness since when it does so it provides the unacceptable contradiction that IBD we are not at all related to chimps whereas non IBD we are over 95% related. Dawkins et al cannot have it both ways.


    John Edser
    Independent Researcher

    Twitter: @intelligent50
    discussion: sci.bio.evolution

  23. Yes, the problem of being lead on because we use language that relates to our ordinary lives happens often in science. There is always a balance between metaphors that get concepts across, and ones that are taken for more than they justify. You can see another example that the religious use against us in Anthropomrphizing Fine-Tuning.

  24. BS. In the 30th anniversary edition of the book, the sentence “Let us try to teach generosity and altruism, because we are born selfish.” was retracted by the publisher. As Frans de Wall notes in his latest book, this in fact equates the selfishness of genes with psychological selfishness.

    1. The quoted sentence (in the first chapter of the first edition) was altered by the author (not Oxford University Press), who was apparently the first to notice that it did not correctly express the idea he presented more precisely and at length in the final chapter.

      1. Thanks for pointing that out, it was not obvious to me. However, the point being is that Dawkins indeed fell for his own metaphor. Semantics matter.

  25. Those critics who claim the the title ‘Selfish Gene’ is confusing to the typical reader of populist science literature can easily be proven wrong. Has the title resulted in hords of devoted readers grossly misinformed by the metaphor?  Thanks to Dawkins it has become pretty obvious that those of us that have truly read the book have come away with a much better understanding of how genes effect evolution. If anything the reader will become even more aware that genes are not little intellegent designers.

    1. Not to mention the fact that E.O. Wilson, the father of the field of sociobiology has moved away from the idea of kin selection being solely behind altruism. Wilson now favors an emergent multi-level selection. Dawkins is behind the times..

        1. I also listened to an interview with Dawkins a couple of weeks ago and he said the word “emergent” about three times. Seems like Dawkins may indeed be seeing the light.

  26. Victor Stenger in The Fallacy of Fine Tuning notes that laws of physics are written to be “independent” of time and place. Theoretically the laws apply in all possible universes in addition to our own. Yet, we know that these laws don’t apply in black holes. Michio Kaku notes that we know nothing about what happens inside black holes (can anything be said to happen within a black hole). Stenger’s conclusion: the laws of science are created by scientists (to allow us to make verifiable predictions).

    1. Yes, the Universe does what it does, and we model it with what we call “laws.” We can’t know if they represent what is really going on at the deepest level, just that they make predictions that we can test. The validity of such laws in a speculative context of some other “Universe” cannot be tested. That means that we can’t know if we are missing something that does not make a difference here, but would make a difference there.

  27. What are you talking about? Who was talking about atheism in this post? Get down from that pedestal, crazy person, none wants to listen to your ramblings.

  28. Thanks for the ‘brown bear on snow’ thought experiment. But there is something deliciously intellectual hidden behind why exactly that ‘thought example’ of evolution in action is inaccessible to most religious people, especially to Creationists. I have written extensively upon the ‘brain problem’ that prevents religious people from acquiring knowledge (in my looong book ‘Origins of Belief and Behaviour’) It goes like this…

    Religious people are part of a a larger group (I call Drones) who have no option but to seek an hierarchy in which to live and work. Look at the historic development of hierarchical social organisations such as institutions; places where academics, administrators, teachers, professionals all live and work. (In the case of Medicine and The Law, the ‘institution’ is contained in the voluminous case-law books and medical books of those professions) In the case of the religious, that hierarchy is supernatural, and was defined by Drones three thousand years ago in their holy books.

    Religious people, like royalists, show astonishing allegiance to those they think are at the top of the hierarchy. I coined the phrase, Brain Operating System’ to describe the assumptions or precepts that allow a human brain to operate. One of the precepts or assumptions that is foundational to the religious mind is the idea of ‘intention’. They figure that if there is intention in all things, then there must be gods who have that ‘intention’. It is logical. Religious people see that logic, and therefore claim that their religion is based upon logic. But they fail to appreciate that logic raised upon false assumptions, remains false.

    There is a problem in them giving complete allegiance to the authority-figure, because almost everything in the world; all the objects and processes, conflict with the simplistic worldview constructed by the religious hierarchy. And so early on in life, the religious sensibility learns to distrust ‘experiential information’. Finally, they are obliged to deny experiential information.

    Another hidden precept of the Drone is a denial of change over time. All Drones suffer temporal distortion. They already have difficulty with the passing of time. They often find confusion in the idea of the past, present and the future. The medieval intellectuals marvelled at Moorish fountains where the rising water was still, but ever changing. They see that life may change, but is always the same! You can see that problem clearly in all religious thinking… they talk of the living jebus, and talk of biblical stories as if they are contemporary. They cannot see death as an end but merely as a simple transformation by reincarnation. For them, the human soul changes by reincarnation, but remains the same.

    When presented with the idea of evolution (which came to some of us naturally by the age of ten!!) they cannot accept the possibility of change over time without some supernatural ‘agent of change’. That is why the popular parodies of evolution voiced by Creationists generally have the evolving lizard ‘deciding’ to grow wings on its back. As Prof. Ceiling Cat pointed-out, for too long scientists have unfortunately fed the Creationist fantasy by including metaphors of ‘intention’. David Attenborough is terrible at it.
    “…the monotremes had to find a way of feeding their young…” rather than..
    “…the monotremes evolved, by trial and error over time, and the many early deaths along the way, a method of feeding their young by exuding belly-milk…”

    There is, of course, a further handicap shared by the religious mind in its inability to seize and understand metaphor such as ‘The Selfish Gene’. Once again it is a problem stemming from their concept of intention behind all objects and processes. You can see the religious mind of Andrew Brown wrestling with, and losing, any understanding of the metaphor. He is only able to take those words literally. It is beyond the religious mind to see that biology moves as if with intention, when there is only blind chance that changes my prove beneficial to the organism.

    1. A very interesting thesis, but impossible to prove. Still, interesting food for thought.
      I’ve alway been attracted to Dawkin’s model of memes, combined perhaps with Dennett’s little metaphor on liver flukes, which take over the “mind” of an ant. Mathematically, memes can be given numeric power ratings (“fitness”) which essentially encompass their effectiveness on the “hot buttons” of human motivation…survival, safety, love, belonging etc. Nothing beats religion as a meme, as it offers endless life, long term security for oneself and family, explanation, fellowship, parental love equivalents (along with parental discipline). On the flip side it offers maximum threat (eternal damnation) as an “antidote” to its rejection. No wonder the religion meme becomes so entrenched in the human psyche.

  29. The hollow pigment free fibres of the polar bear’s fur scatter light so appear white. There are other traits than whiteness alone which enable the polar bear to survive on the ice, for example its feet.

  30. I read Midgley’s criticisms years ago. And then I read them again because I genuinely thought I’d misunderstood her arguments.

    I couldn’t – and still can’t and don’t – believe that someone could so comprehensively misunderstand such a simple and elegant metaphor. Especially so since Richard never tires in the book of explaining very carefully, clearly and repeatedly that it’s a metaphor.

    Then and since he’s written a great deal about it being a metaphor, about the dangers of teleology and imprecise language. He’s explicitly pointed out and disarmed areas that might conceivably cause confusion.

    In the book he notes many times that genes can’t really ‘want’ anything but that they behave as though they do. And after he has stated this clearly and at length, he continues to remind the reader about it throughout the book. And then again in most of his subsequent books.

    What more could anyone *possibly* do?

    I can only conclude that Midgley indeed did not read the book. And that either she didn’t read Richard’s rebuttal or felt she’d become too invested in her error to pull out.

    I haven’t read Brown’s piece yet, but if it’s anything like every other article of his I’ve ever read, I’ll be spending the rest of the week in a fury.

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