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    The Cult of Gene-centrism
    By Steve Davis | October 24th 2012 01:39 PM | 31 comments | Print | E-mail | Track Comments
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    An advocate for gene-centrism recently wrote that the concept is a reductive mechanism for the understanding of evolution.
    The first part of that statement is correct; it is reductive, and is therefore lacking those qualities that are necessary for properly understanding a “big-picture” process such as evolution.

    The second part of the statement is not correct. Gene-centrism did not develop as an explanation of evolution; it began as, and has remained a political movement within evolutionary biology, the goal of which is to destroy group selection as a credible evolutionary process.

    I refer to it as a political movement because it presented not an evolutionary view, but a world view, as John Maynard Smith admitted in his review of The Selfish Gene. 

    And of course, the publication of The Selfish Gene was the opening salvo in the war against group selection that swept all before it with extraordinary success. But, in a beautifully ironic twist, a movement based on selfish genes could not stand up to scrutiny once the initial excitement died down, and so we have witnessed the evolution of the movement itself, from the fantasies of selfish genery to the supposedly more rational gene-centrism.

    So is it more rational?
    Barely. The movement is still having to contend with group selection after all these years, so they are attempting to cage  group selection, to tether it within the limits of a gene-centric framework.
    But let’s have a brief look at it’s history.

    The Selfish Gene, published in 1976, was inspired by  Adaptation and Natural Selection, written by GC Williams in 1966, and the nonsensical propositions found in The Selfish Gene can be found in Williams. For example, Richard Dawkins’ description of organisms as “survival machines” constructed by genes is a re-wording of Williams’ belief that “the succession of somatic machinery and selected niches are tools and tactics for the strategy of genes.”

    But more importantly, it was Williams who put purpose back into evolution after Charles Darwin had removed all traces of such mysticism.
    Here’s some more examples of Williams’ misunderstandings.

    He stated that a function or adaptation is something which “is produced by design and not by happenstance.” Intelligent Design, perhaps?
    He stated “Every adaptation is calculated to maximise the reproductive success of the individual, relative to other organisms.” Calculated by whom, we might ask?
    He stated “Each part of the animal is organised for some function tributary to the ultimate goal of the survival of its own genes.” A goal requires purpose. Whose goal, and whose purpose we might ask?

    And as if to declare publicly that gene-centrism is a cult based on ideology, he stated that non-kin altruism is a “biological error.” There can be no such thing in nature. This is ideology at its worst. He might as well have said “Non-kin altruism does not fit with my view so it must be a biological error, because my view cannot be wrong.” And Richard Dawkins gave the ideology traction by repeating this arrogant nonsense in The Selfish Gene with the rhetorical flair that Williams lacked.

    Despite stating more than once that genes are not conscious and do not show purpose, everything else Dawkins wrote pushed the opposite line, the line of GC Williams; that genes have purpose. For example “For our purposes, the word allele is synonymous with rival.” The only way that a gene could have a rival, and hence a purpose, is if the gene is alive, which is clearly not the case. And as genes cannot have purpose as Dawkins already concedes, then they cannot have rivals. So what are “our purposes” exactly, I wonder? (Do not be fooled by those apologists who claim that such uses of language are legitimate “short-cuts” to the truth as they call them. These short-cuts are an abuse of language, designed to deceive.)

    Has the mystical “purpose” of selfish genes been left behind as the idea evolved into gene-centrism? Not at all.

    A student of Dawkins, Alan Grafen, now prominent in the field, in lecturing group selectionists on their lack of careful thought, rather condescendingly made this remarkable and helpful comment to get group selectionists back on track; “Two excellent ‘self-help’ guides are books by G C Williams (1996) and R Dawkins (1976).” Yes, we’ve seen how helpful they are. And we also see from this that gene-centrism will never abandon its roots.

    And as late as 2010 Grafen co-authored a paper titled Capturing the Superorganism : A Formal Theory of Group Adaptation, which, you might assume from the title, is a welcome move towards a more holistic approach by gene-centrics to evolutionary theory. Unfortunately, it was nothing of the sort. As if to placate cult members who might be alarmed by the title, the paper began with a re-assuring quote from Dawkins :
    I have characterized inclusive fitness as ‘that property of an
    individual organism which will appear to be maximized
    when what is really being maximized is gene survival’...
    One might generalize this principle to other ‘vehicles’. A
    group selectionist might define his own version of inclusive
    fitness as ‘that property of a group which will appear to be
    maximized when what is really being maximized is gene
    survival’! Dawkins (1982, p. 187)

    The implied message to the true believers, a message intended as a defence against a charge of heresy, was “Never fear, no matter how much evidence emerges for group selection, it’s still all about genes.”
     Then came this revealing titbit in the very first sentence of the introduction :
    “Darwinism is a theory of the process and purpose of adaptation.”
    Purpose? So, as you can see, in 2010 Williamsian mysticism was alive and well in the minds of the faithful. All that has evolved is the name.

    So why this regression? Why was the mystery injected back into evolution after Darwin went to so much trouble to eject it?
    I see a couple of reasons, both tied to human psychology.

    It’s something of a paradox that despite our bodies being comprised primarily of fluid, despite relationships being fluid, despite life itself being fluid, despite fluidity being the prime quality of existence itself, there are many among us who cannot come to grips with that reality. They crave solidity, a crutch, even if that crutch is no more than mere prejudice. So for the timid and insecure, the discovery of genes was a blessing. Here was something they could hang their hats on, something that provided substance in a world of anarchic chaos. The random nature of natural selection and evolution was no longer quite so unpredictable. Suddenly, with a little imagination and self-deception, they could see purpose. Purpose and mystery combined in one delightful package, what more could the timid and fearful ask for?
    Just a little more, actually. They turned the package into a religion. They portrayed genes as powerful and immortal. Powerful, immortal AND invisible? Sounds like Someone we all know!

    If you doubt the religious aspect to gene-centrism, consider the glowing references to William Paley (of “Natural Theology” fame) by both Williams and Dawkins. Why would they do that? Paley’s adapation-by-Designer argument for the existence of God, ruled supreme until The Origin of Species appeared. Its downfall was swift, its reputation reduced to tatters. But Williams and Dawkins resurrected the concept by simply changing the name of the designer, hence their regard for Paley. Instead of God as designer we were given genes as designer. Yes, you guessed it; designer genes. (Remember the “tools and tactics for the strategies of genes”? Just what a designer needs!) The philosopher David Stove in his discussion of this, wryly observed that just as the many gods of earlier times were replaced by God, so we might eventually see many genes replaced by Gene. Amazingly, this tongue-in-cheek prediction actually came to pass. (http://www.science20.com/darwin_20/selfish_gene_confusion_and_its_interdisciplinary_roots)

     The other psychological aspect to this is equally disturbing.
    Ever since humans first saw that mathematics could explain natural phenomena, numbers have been assumed to have magical qualities. So it’s no coincidence that because the two pioneers of gene-centrism, R A Fisher and W D Hamilton, were mathematicians before they were biologists, both developed aspects of gene-centrism based on mathematics, and consequently numbers have assumed the highest rank in the gene-centric view of evolution. (When Dawkins talks of maximising gene survival, all he’s talking about is numbers.)
    The problem with this is that gene numbers are not evolution, just as a map is not a landscape, and the price of sugar is not sugar. The spread of genes in a population is just one outcome of natural selection. It is not an explanation of natural selection, just as ash is one outcome of fire but is not an explanation of fire. Here’s a definition, the significance of which will be immediately recognised;  “Numerology is any study of the purported divine, mystical or other special relationship between a number and some coinciding observed (or perceived) events....The term numerologist is also used derogatorily for those perceived to place excess faith in numerical patterns and draw scientifically unsound inferences from them, even if those people do not practice traditional numerology.”
     So, scratch a gene-centric and watch a numerologist bleed.

    It’s important to note that gene-centrism was a fringe position with little credibility until Hamilton’s inclusive fitness equation was dusted off and thrown into the mix. It is still regarded with reverential awe by numerologists today, despite being an equation about almost nothing. (http://www.science20.com/gadfly/hamiltons_rule_or_hamiltons_folly)
       But inclusive fitness did conjure up the appearance of another deity to add to the pantheon – the fabled but elusive altruism gene. Well, not so much deity as poltergeist, really. Just as some of the gods of ancient cultures were troublesome busy-bodies, so the altruism gene has driven gene-centrics to distraction as they try to fathom its mysterious machinations, its spiteful interference in their otherwise well-ordered universe.

    So are genes important in natural selection and evolution?
    Of course! Just as every other link in the chain is important. (Please excuse that most unfluid-like metaphor.)

    The main problem with gene-centrism is that genes are not central to anything. All a gene does is synthesize a protein, which it does not do independently. A protein is not a trait, it merely contributes to a trait. And that protein can have different outcomes depending on other inputs. We can really put the alleged “centrality” of genes in perspective by considering the medical condition known as PKU which afflicts a small minority of humans who happen to have a genetic predisposition to it. They are only afflicted if they have a particular diet. So what causes the problem, genes or diet? We cannot say, but what we do know is that genes are not central to it.

    Evolution can be seen as linear, we go from A to B to C after all, but it is a process derived from a network of influences. If one influence in the network changes, the outcome changes, so the only thing central to evolution is life itself.  How often in discussions of evolution do we see life treated in depth? Never. Life is feared because they do not understand it.

    Postscript.
    In the interval between writing this and publishing it, the advocate for gene-centrism referred to in the opening paragraph has become a serial offender.  (http://www.science20.com/alpha_meme/intelligent_design_science_curriculum-95465)  He wrote;
    Design, according to some, needs a designer. However, famous biologists and neo-Darwinists such as D. Dennett say that evolution “designs” by natural selection. If we accept that usage of the term, “design” does not by definition imply an intentional act.
    Nowhere in the piece that followed was there a rejection of that proposition. It was Richard Dawkins all over again, denying intent while implying intent. If a suitable term can found, why use an unsuitable one? So I’ll do the hard work for them, I’ll find the suitable terms. Here’s the process, for those who might be understandably confused by his statement. Environments eliminate those organisms that become less fit as environments change. If that elimination is on a large enough scale across a population, evolution could occur. That’s it. Evolution is merely change over time. And in the process as I’ve explained it there is clearly no designer, so the use of the word is not an aid to clarification, it is the first step towards misunderstanding.

    The misunderstandings continued later in his article with this; However, natural selection needs some background, some statistical ensemble on which selection can happen.
     This is a great example of the influence of numerology on gene-centrism. I can make that assumption because the statement is false. It’s false because selection does not need the magic of numbers in order to occur, it needs variation. Variation, as we’ve seen in the case of PKU, comes from environments as much as genes. (Genes are numbers to a numerologist.) Gene centrism, like selfish gene theory before it, aims to give to genes a significance in evolution beyond that which exists in reality.

    Comments

    Alright, Steve, enough pussy-footing around the subject; tell us what you really think about this stuff. ;)

    Sadly, it seems the religion is growing. Just yesterday I saw a new article on the 'altruism gene'.

    Thanks for a well thought out article. Now, how to get this into the heads of those who aren't already converted to GENE?

    Thor Russell
    "it is reductive, and is therefore lacking those qualities that are necessary for properly understanding a “big-picture” process such as evolution."

    The atomic theory of matter is reductive, therefore it is lacking those qualities necessary for explaining a "big-picture" process such as how our sun generates energy, or how a large lake freezes. It definitely can't explain what I am thinking right now, so I am not made up of atoms... 



    To those rational readers out there:
    It is not a choice between one or the other, there are many levels that selection can act on:
    http://en.wikipedia.org/wiki/Unit_of_selection

    Some levels are better at explaining phenomena than others. The fact that I AM made of atoms doesn't make me "atom centric" and therefore deny the existence of anything to do with higher levels of organization such as cells or that neurons not just atoms are helpful in understanding the brain. To put things another way, imagine there were camps in biology, the "atom centrists" the "molecule centrists", the "cell centrists" the "organ centrists". Each tried to explain everything in biology in terms of their view, e.g "cell centrists" deny that molecular study of the DNA molecule leads to insights. This is how ridiculous "gene centrism" vs "group selection" can seem at times.
     

    Higher levels can sometimes do quite interesting things:
    http://mbio.asm.org/content/3/2/e00036-12
    http://www.sciencedaily.com/releases/2012/03/120327094056.htm
    In these cases, very successful and in fact essential genes could be rejected by the majority of organisms because the "service" is still provided by the group.
    Thor Russell
    Thank you for your article :)

    vongehr
    An advocate for gene-centrism ...
    who wrote in the very article that you link to
    As one who rejects bio-centric evolution, I am automatically immune to extreme “gene centrism”!
    I, a supporter of algorithmic evolution and group selection mechanisms and all of that, am "an advocate for gene centrism"?!? Apparently, the issue is simply beyond your grasp.
    Steve Davis
    "I, a supporter of algorithmic evolution and group selection mechanisms and all of that, am "an advocate for gene centrism"?!?"
    Of course!
    You made that quite clear in your articles, and also in this, "I am automatically immune to extreme gene centrism."
    The mistake you make is to try to distance yourself from naive gene-centrism.
    That's a mistake because all gene-centrism is naive, as I demonstrated in this article.
    I provided links to your articles so that readers could compare what you wrote to what you would subsequently say you wrote. I anticipated that the comparison would be interesting, and that's how it turned out. 
    vongehr
    Design, according to some, needs a designer. However, famous biologists and neo-Darwinists such as D. Dennett say that evolution “designs” by natural selection. If we accept that usage of the term, “design” does not by definition imply an intentional act.
    Nowhere in the piece that followed was there a rejection of that proposition.
    This is where reading comprehension comes in! What else but a rejection is it if the whole text, based on that definition of "design", ends up in demanding ID in the curriculum?
    Look - if advanced mathematical modeling is so mysterious to you as to count as "numerology", you are perhaps wrong on a science site?
    Steve Davis
    "What else but a rejection is it"?
    I actually considered that possibly in your waffle there might be something that could be used as a defence, that perhaps I even got it completely wrong, but I was persuaded against that by the fact that you are a serial offender, as in this from the first article; "That a gene “wants” something or is “selfish” is always a shorthand, and those who use it have always explained that it is such! It is short for: Genes are vitally involved in ("central to") determining behavior,..."
    That is consistent with the same abuse of language you used when discussing design, so I'm happy to conclude that you see no problem with denying intent while implying intent. And I must add that this was not the only error. That genes are vitally involved in determining behaviour is perhaps excusable over-enthusiasm, but no amount of enthusiasm will make them central to anything despite the fervent wishes of the faithful.
     "...if advanced mathematical modeling is so mysterious to you..." The only thing that's mysterious here is why you assume that selection needs numbers to work. You said it, you can retract it any time you like.
    "...you are perhaps wrong on a science site?"
    Well, I'm in good company there, am I not?
    Everyone who takes you to task gets the same suggestion!
    I think I see a pattern of behaviour here, I wonder if it's genetic?
    vongehr
    the same abuse of language
    This, and really only this, is your problem, the point where you get everything wrong. But you are not alone in being unable to grasp that language is the core of the construction of scientific knowledge. You are perhaps simply too old to get to the next level where we realize that science is not about "X" truly really out there, but about how we define "X". You do it again and again, whether it is "life", "centrism", or "design", you want to force your private, usually ill-defined terminology onto others, and instead of waking up to the fact that this results only in useless fights, you rest bitter in the trenches feeling cheated by those who go on and abandon you. As I said many times before, it is sad, but it is perhaps never too late. Perhaps THC can help.
    Steve Davis
    Do you not read your comments at all?
    Have you not noticed that I presented a critique of gene-centrism as a whole, I refuted your arguments in support of gene-centrism, yet all you have done is attack me personally!
    Address the issues or be seen as a fraud.
    A summary of the article:

    Take an idea or quote by a gene-centric writer. Take it out of context and misunderstand it so that it becomes silly. Point out that the idea is now silly. Conclude that gene-centrism is silly. Repeat.

    vongehr
    Yep - thats it, the old straw thingy, except, he advanced this method and now can even do it quoting non-gene-centric writers.
    Steve Davis
    "...and now can even do it quoting non-gene-centric writers."
    Oh, so you're backing away from gene-centrism now.
    Funny that, because you were pretty gung-ho when you posted a comment to Gerhard's article "Inclusive fitness is Group selection."
    Dawkins could have written these; "Inclusive fitness is backed up by numerical simulation that focuses on genes."
    Followed by "...genes that try to invade the gene pool."
    Followed by "Gene centrism avoids mysticism about mechanisms." 
    You are a gene-centric from tip to toe!

     
    John Hasenkam
    The problem isn't that gene centrism is reductive it is the propensity to think of biological objects as possessing intrinsic properties. So when you hear people talking about genes "containing information" think very carefully about that idea. Where is that information stored? If you wish to entertain such an idea consider that if "information" is "transmitted" or "visible" it is only so at the point of interaction with other objects. That is, genes don't contain "information", they are better understood as "response sets" and focus on their relational properties rather than purported intrinsic properties. Unfortunately biology in general tends to think about the things in themselves rather than their relational dynamics.
    Consider inflammatory mediators. I had that appellation. If you read enough about inflammatory mediators you soon realise that these molecules do a host of other things apart from inducing inflammation. So I say to people that they must remember the names given to biological objects often reflect the initial understanding at first discovery but later research reveals a great deal more about the particular object. Hence Wittgenstein: words are posts on which we hang meanings. ... But people stop hanging on the meanings and the word takes on a life of its own. So forget the names, or rather when you think of the name remember all the possible associations, all the possible interactions, all the contextualised effects these objects can bring forth. 

    I'm not even sure we should talk about "brains storing information". Sure that is trivially obvious
    but it presents some enormous problems when you ask "how?" So perhaps even brains can be better perceived as massive response sets and that involves more than neurons; hence, again, if you keep reading the literature you will realise that behavior is not just mediated above the neck but by endocrine factors, the enteric nervous system via the vagus nerve, a host of humoral factors. 
    Enough of my arm waving. Have a good night. 
    Steve Davis
     "...the word takes on a life of its own."
    Thanks John, interesting comments
    I respectfully disagree, John.

    Wittgenstein made a good point, which is that humans interpret words differently depending on their own worldviews. Communication is difficult at times, and perhaps we can agree on that.

    Genes do contain information. The recent release of the human genome for scientific investigation has led to confirmations between the genetic abstractions and archeological evidence of the different paths humans took. Archeology informing biology and vice-versa.

    Genes contain a series of nucleotides (TAGC) the precise sequence of which are the genetic code and are the information the gene contains. If the sequence is changed, the information is changed.

    I suppose one could get into a lengthy discussion of what, exactly, information is. I’m not equipped for that discussion.

    Steve Davis
    Frank, thanks for your input.
    It's a matter of economy. The language of intention occurs in discussions of genes because it is simpler to say "genes try to survive" than "a gene which has a function which tends to promote the successful transmission and propagation of the gene will on average increase in frequency in a natural population over generations". All the popular science books I've read on evolution have a disclaimer somewhere to clarify this. Scientific papers are more tedious to read precisely because they tend more to avoid such useful simplifications.

    Also, simulation and modelling papers almost invariably begin and end with a summary of experiments which establish the assumptions of the model, and there is also a solid body of literature dedicated to testing the predictions of models.

    A also think that there is a misunderstanding of what is meant by gene in the literature and I will agree that the authors could make it more clear, with the exception of Dawkins who makes it quite explicit in a footnote in chapter one of the selfish gene. Genes are sometimes defined as a length of chromosome which codes for a single protein. I prefer the old term 'cistron' for this case, if only because that definition is now a bit shakey given findings in the last decade. At other times, a gene just means whatever underlies a heritable trait - so a 'gene for alturism' may in fact be the result of many cistrons, control elements, epistatic interaction, epigenetics and so on, which has the effect of producing heritable altruistic behaviour.

    Gerhard Adam
    At other times, a gene just means whatever underlies a heritable trait - so a 'gene for alturism' may in fact be the result of many cistrons, control elements, epistatic interaction, epigenetics and so on, which has the effect of producing heritable altruistic behaviour.
    "When I use a word," Humpty Dumpty said in a rather a scornful tone,"it means just what I choose it to mean --- neither more nor less."
    Apologies to Derek for stealing his post.
    Mundus vult decipi
    Steve Davis
    It's not a matter of economy at all, Tim, it's a matter of misunderstanding as you demonstrated.
    For a start, "Genes try to survive" is nonsense because they do nothing of the sort. How can a portion of a molecule try to survive? So how can that be relevant to anything that they actually do? How can that be a shortcut to reality?
    I'm all for economy of language, and if that sentence was an aid to understanding I would support it.
    Now let's consider what the shorthand represented.
    "a gene which has a function which tends to promote the successful transmission and propagation of the gene will on average increase in frequency in a natural population over generations."
    You cannot see the deficiencies in that statement because you've been mesmerised by Dawkinsian rhetoric.
    Your statement completely ignores the primacy of the cell in the transmission of traits.
    It is not the function of a gene to propagate itself.
    The function of a gene is to produce a protein when directed to by the cell.
    If, purely by chance, that function happens to increase the chances of the gene being propagated, it will only be because the functions of the genome as a whole have played a role in the survival of the organism.
    You have simply given evidence for my repeated assertion that gene-centrism is all about giving to genes a greater role in evolution than exists in reality.
    John Hasenkam
    Frank,
    Genes do not always code for the same protein, their position on the chromosome, siliencing RNAs, interfering RNAs, epigenetics, also play an important role in what protein is created from specific genes. The Central Dogma, or at least parts of it, are wrong. There is two way interaction here, it is not just "bottom up". 
    Sorry John. I was objecting to your statement that genes did not contain information. The fact that the information can be modulated, sliced, diced, split, truncated, or otherwise changed in the process of defining a further structure doesn't alter the initial condition. There is information present.

    I note that I should not have responded at all. My intention in reading these topics to gain further understanding from those who have studied these concepts far longer, and deeper, than I ever will. We would likely be better served if I come here in read only mode.

    Gerhard Adam
    Let me toss out a very simplified set of definitions, so that perhaps we can avoid a philosophical quagmire.  I realize that many people may disagree with them, but perhaps it may serve as a point for further refinement.

    I tend to consider this at several levels of definition so that we have:

    (1)  Data - this represents some combination of elements whether they be molecules, or bits [series of 1's and 0's in a computer].  In and of themselves, they are simply raw components.

    (2)  Information - is defined as some interaction that results in the ability to interpret "data" to some degree where decisions or choices can be expressed.

    (3)  Knowledge - is where information can be integrated to create new "information" or benefit from "experience" [i.e. historical precedents].

    I realize that these are necessarily vague descriptions and could warrant considerable discussion as to how applicable they are to the general scheme of things, but perhaps it may provide a way to begin interpreting what we might by the expression of these constituent parts.
    Mundus vult decipi
    So then, if I follow correctly, the DNA nucleotide sequences are merely the data and don't represent information until (or unless) there is a receiver of some sort. Is that how you view this knotty discussion?

    Gerhard Adam
    Yes, because until it is transcribed and interpreted, resulting in the expression of a protein, it doesn't mean much.  This is also why transcription errors can produce different results or problems;  same data, different information.

    That's the underlying problem with gene-centricism, in that it presumes that each gene is selectable based on specific traits and therefore representation in future generations is indicative of such genetic competition. 
    Mundus vult decipi
    Alright. I can accept that. Although, it's getting downright quantum mechanistic seeming. I'll have to wrap my head around this concept and see if I can square this idea with the knowledge that biologists are extracting DNA from 10k year old bodies and using those data to identify it. Thanks.

    We perhaps agree but are saying it differently. If you take my second definition of a gene("that which underlies a heritable trait"), rather than what I rather call a 'cistron' ("all the DNA lying between a stop and a start codon"), then does this disagreement resolve?

    That's the definition I'm advocating, it's also what Dawkins means when he uses the term, though I'd rather argue from the theories and models themselves rather than have semantic discussions about Dawkins wording.

    Gerhard Adam
    Except that's not how Dawkins means it, and in that respect he is simply wrong.
    More handwaving,
    I am using the word gene to mean a genetic unit that is small enough to last for a number of generations and to be distributed around in many copies. ... The more likely a length of chromosome is to be split by crossing-over, or altered by mutations of various kinds, the less it qualifies to be called a gene in the sense I am using the term.

                    ....The Selfish Gene (1989) p.32
    I said that I preferred to think of the gene as the fundamental unit of natural selection, and therefore the fundamental unit of self-interest. What I have now done is to *define* [Dawkins' emphasis] the gene in such a way that I cannot really help being right!

                    ....The Selfish Gene (1989) p.32
    The fact that Dawkins uses the word "gene" in such a non-standard way is not an issue as long as one recognizes that the Dawkins "gene" has nothing to do with the genes that molecular biologists and geneticists talk about. It's not an issue as long as one doesn't try and pretend that Dawkins has avoided handwaving and "clearly" refuted the problems raised by his critics.

    http://sandwalk.blogspot.com/2007/01/richard-dawkins-definition-of-gene-is.html
    Even your phrase "that which underlies a heritable trait" isn't correct, because it "underlies" all traits, and is involved whether it expresses a protein or not.  Such a phrase denies genes for traits that have gone to fixation since their heritability is zero.

    The problem is that it's much more complicated than such simplistic representations allow.  Genes code for certain proteins, which may or may not be expressed [or fully expressed] and will product traits that may or may not be complete and will be further refined by other factors whether it be epigenetics, or involve the microbiota, etc. 

    Even when everything is "done", the organism determines how these traits manifest from a survival perspective.  We can readily see examples of such changes when individuals are born without arms and yet learn to use their feet to write or draw.  One cannot argue that the genes coded feet to be used in this manner, yet the final organism was able to exploit the abilities that did exist and use them.

    If we were to take such an extreme example and consider what would happen if such traits were selectable and produced an advantage, then you would also see the secondary difficulty of establishing whether a trait was adaptive or an adaptation.  This is precisely the situation that gets confused when these simplistic reductionist views of genes take place.

    Perhaps one of the most striking examples of this comes from germ-free mice which are raised free of their microbiota.  This offers a clear view of what genes produce as well as their limitations in producing a viable organism.

    Another problem is DNA repair, since that clearly suggests a higher order regulatory mechanism for ensuring the integrity of the individual genes.  Similarly we have a problem in the production of cells, which the genes are incapable of controlling.  There's no question that genes play an extremely important part in evolution/selection because they are the purveyors of "information" necessary to produce the next generation, but they are hardly the whole story.  So gene-centricism fails to account for or predict the actual results of heredity, since many of the major aspects that produce a new cell or regulate it, are unaccounted for by looking at genes alone.

    I've used the example of letters/words or music notes before and in the same fashion, we can recognize how important these elements are in producing books or music, but one could never conclude that it was these elements alone that determined whether books or music were actually produced.  Books are not simply the means by which words ensure their "survival" into the future.  Although if one examined the evolution of language, a similar argument could be made [to that of gene-centricism].... except that it would be wrong.
    Mundus vult decipi
    vongehr
    I said that I preferred to think of the gene as the fundamental unit of natural selection, and therefore the fundamental unit of self-interest. What I have now done is to *define* [Dawkins' emphasis] the gene in such a way that I cannot really help being right!
    This is why gene-centrism (the defined "gene", being applicable to memes) is not the same as its biocentrist misinterpretation (DNA molecules). He defines it tautologically so that it is fundamentally true. That is the way I do it for "totality" and "fundamental theory" for example.
    Genes code for certain proteins
    Some do.
    Steve Davis
    Dawkins; "What I have now done is to define the gene in such a way that I cannot really help being right!"
    Vongehr; "He defines it tautologically so that it is fundamentally true. That is the way I do it for "totality" and "fundamental theory" for example."
    That would sound so profound except it's rather too close to this; "It’s possible that I shall make an ass of myself. But in that case one can always get out of it with a little dialectic. I have, of course, so worded my proposition as to be right either way." Karl Marx.
    Well, there's plenty of asses commenting on evolutionary biology, but I prefer to think of them as shifty unprincipled con-men.
    vongehr
    Ha ha, yes Steve, such of course blows your mind, so don't worry about it. Gerhard understands what I mean.
    Steve Davis
    Aren't you forgetting something?
    You walked away from this when the going got too hot, then you think you can come back in when it suits.
    Address the issues.
    You still need to explain why natural selection needs the magic of numbers to operate. Sorry, it's not the magic of numbers, it's a "statistical ensemble".
    That sounds so cool!