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    The Extended Phenotype - How Richard Dawkins Got It Wrong Twice
    By Steve Davis | February 16th 2009 01:44 AM | 138 comments | Print | E-mail | Track Comments
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    “The Extended Phenotype – The Long Reach of the Gene” is the book Richard Dawkins wants you to read “if you read nothing else of mine” because “It is probably the finest thing I shall ever write.”
    It purports to be about science, for scientists, yet at the very beginning there is a quite remarkable disclaimer. Dawkins warns the reader that the book contains nothing new, that it is “unabashed advocacy”, (in other words a mere personal opinion) and that it contains no hypotheses that are testable. In short, the book is declared from the outset to be non-scientific.
     Lulled into a false sense of security by this refreshing dose of honesty, the unwary reader is soon swept along by a tsunami of rhetoric to emerge at the book’s conclusion, convinced that white is black and that all that was previously considered normal and proper was not necessarily wrong, “It’s just that you see it wrong.” This begs the questions as to why a scientist would wish us to read it at all, and why a scientist would regard such a book as his most important work. (Dawkins did explain his reasons for writing the book, to get people to look at biology in a new way, but did not explain his motive for this.) It also signals a remarkable departure from his most famous work, The Selfish Gene, in which the reader was informed at the outset “It is science.”

    But was it science? My criticisms of The Selfish Gene are many, but most come under the broad criticism that the book’s principal propositions are, like those of The Extended Phenotype, untestable and therefore non-scientific. But there were numerous false assumptions and skewed definitions in TSG that were carried over into TEP. One skewed definition is that of the phenotype. On page 235 TSG Dawkins defines phenotype as; “the bodily manifestation of a gene, the effect that a gene, in comparison with its alleles, has on the body, via development.” But this is not a phenotype. A phenotype is the combined effects that the environment and the genotype (all the genes of an organism) have on the development of the organism. It is only by subtle and misleading changes in emphasis such as these, (the same for gene, natural selection, etc.) that Dawkins is able to give any coherence or plausibility at all to selfish gene theory.

    But what was it that Dawkins wants us to look at in a new way?
    The extended phenotype according to Dawkins, is the effect that selfish genes have on their environment; their influence on the world outside the organism they inhabit. “We see the wider world as an arena in which these genetic fragments play out their tournaments of manipulative skill.” Pretty scientific, huh? An example given of an extended phenotype is the beaver dam. I’ll have to quote most of this or you just won’t believe me. From the final chapter of The Selfish Gene 2nd ed.; (This chapter titled The Long Reach of The Gene was added to the book after the publication of The Extended Phenotype, and is described by Dawkins as a summarised version of TEP.)
    “It is not entirely clear what its Darwinian purpose is, but it must have one, for the beavers expend so much time and energy to build it. The lake that it creates probably serves to protect the beavers from predators….Whatever its benefits, a beaver lake is a conspicuous and characteristic feature of the landscape. It is a phenotype, no less than a beaver’s teeth and tail, and it has evolved under the influence of Darwinian selection. Darwinian selection has to have genetic variation to work on. Here the choice must have been between good lakes and less good lakes. Selection favoured beaver genes that made good lakes for transporting trees, just as it favoured genes that made good teeth for felling them. Beaver lakes are extended phenotypic effects of beaver genes, and they can extend over several hundred yards. A long reach indeed!” This is science for scientists?

     I’ll start with “Darwinian purpose”. There is no such thing. Natural selection has no purpose.
     Next, “It is a phenotype…” It is not a phenotype; a phenotype has a clear definition as shown above.
    Next, “Are extended phenotypic effects…” A beaver dam might be a phenotypic effect, except no evidence of genetic origin is given for this, and it could be that dam building is a learned behaviour, given that young beavers stay home working until the next litter is about to appear. Sloppy presentation and assumptions such as these are a regular feature of Dawkins’ work.

    The stated purpose of The Extended Phenotype and hence the beaver dam, was to persuade readers that the gene is the unit of selection, an entity possessing individual power and purpose, exactly the same proposition as was put for The Selfish Gene with the added feature of external influence. (The original sub-title of TEP was The Gene as The Unit of Selection.) He employs a tool to convince us of this important idea. Dawkins constantly reminds the reader to ask of adaptations; who or what benefits from the adaptation? It is that very question that destroys his mission. He believes that the gene causing the adaptation is the only entity that should be considered as the beneficiary. This is the heart and soul of selfish gene theory, but he inadvertently gives us in TEP page 92 the evidence that illuminates and destroys the fallacy.
     Dawkins: “Let me show how easy it is to use the gene as a conceptual unit of selection, while admitting that it is only defined by comparison with its alleles. It is now accepted that a particular major gene for dark coloration in the peppered moth has increased in frequency in industrial areas because it produces phenotypes that are superior in industrial areas. At the same time, we have to admit that this gene is only one of thousands that are necessary for the dark coloration to show itself. A moth cannot have dark wings unless it has wings, and it cannot have wings unless it has hundreds of genes and hundreds of equally necessary environmental factors. But this is all irrelevant. The difference between the light and dark phenotype can still be due to a difference at one locus, even though the phenotypes themselves could not exist without the participation of thousands of genes. And it is the same difference that is the basis of natural selection…However complex the genetic basis of features that all members of a species have in common, natural selection is concerned with differences. Evolutionary change is a limited set of substitutions at identifiable loci.”

    There is a devastating problem for Dawkins in that passage.
     He conveniently forgot to ask the question he lectures the reader to always have at the forefront of these discussions; who or what benefits from the adaptation? In this case, as in all cases, it is not only the gene for the variation that benefits. An adaptation can only be successful when it ensures the survival of the organism. All the genes of the organism benefit. No matter what issue is considered in natural selection and evolution, the group in some form is ever present and must be included. The concept of the all-powerful gene as the basis of the extended phenotype is a fable. Genes do not “play out tournaments of manipulative skill”, they merely contribute, with all the other genes in an organism, to the survival or failure of the organism. The level at which selection occurs is just as important as the unit of selection, perhaps more so.

    Now every fable has an element of truth, and so it is for selfish gene theory. There does exist, in every organism, a form of selfishness. It’s that variety of selfishness necessary for individuals to survive, the same selfishness that Dawkins analysed incorrectly in his treatment of selfish genes. This variety of selfishness was addressed by Konrad Lorenz and Robert Ardrey, but they referred to it as aggressiveness. Ardrey saw the shortcomings of the word for its potential to be confused with violence. As he saw it, aggressiveness is common to all creatures but does not in all cases progress to violence. I think a more suitable term is assertiveness. Assertiveness in this sense is the tendency of even a single-celled animal to remove itself from potentially harmful environmental factors, and to move towards comfort zones or food sources. When we head for the fridge we are displaying the assertiveness necessary for survival. Assertiveness appears similar but is subtly different to individual selfishness, and they have vastly different origins. Assertiveness may well have a genetic component, but selfishness is learned, for it is a feature of those organisms that are nurtured by parents. We learn it in the nest or on the breast. For example, a new born infant is helpless. It’s so dependent on its kinfolk and their attention is so focused on its needs that it spends its first few months of life understandably believing that it is the centre of the universe. As the child matures intellectually it slowly casts off this learned behaviour. (Excessive selfishness in an adult is a sure sign of intellectual immaturity.)
     
    But there is such a thing as a selfish gene. It’s called a segregation distorter, and it’s selfish because it doesn’t cooperate with the other genes in the organism, it acts to increase its representation in the gamete. But far from increasing its representation in the population as selfish gene doctrine should predict, it usually results in sterility of the organism. So much for selfishness as the dominant feature in evolution.

    The Extended Phenotype was written to reinforce the view that the gene is the most important element in evolution. Dawkins believes it is important because, of all the factors and influences involved, the gene is the only one that persists, that survives the death of the organism. This view of the gene has an interesting analogy in archaeology. (Dawkins has a fondness for overly simplistic analogies, so I’m vindicated in advance!) We are told that in certain places civilisations have built cities on the ruins of former cities, using the rubble of the fore-runner as building material. This can happen several times with successive civilisations. But although the same bricks are used, exact copies of former buildings do not appear because different forces contribute to their arrangement, just as genes in general do not produce copies of organisms. Individual bricks can have their own characteristics just as genes do, but a certain feature of a brick in one construction can have an entirely different effect when used in the next. A brick with a hole in it could be used to channel air in one building, water in the next, a reinforcing rod in the next. The same for genes. And as with genes, the survival chances of the building do not rest on one particular trait of the building, but on the building when considered as a unit. It’s no good having the smartest looking roof in town if termites are eating the floorboards. Does this make the brick the most important feature in a building? I think not. Important though bricks are, and despite the fact that bricks can survive several reconstructions, they are just one of countless factors that need to be considered. (I warned you it was simplistic, but then, so is selfish gene theory.)

    It’s very difficult to argue that black is white, and although Dawkins tries several tactics to achieve just that, slip-ups are inevitable. One such slip-up can be found in “A Devil’s Chaplain” (2003). On page 260 the following appears. “If a genetic change has no causal influence on bodies, or at least on something that natural selection can ‘see’, natural selection cannot favour or disfavour it. No evolutionary change will result.”
    Why is this a slip-up? Because it contradicts two basic assertions of selfish gene theory. It shows that the organism is the level at which selection occurs, not the gene. This is surely of crucial importance. (So important that selection levels are an infrequent topic for Dawkins.) And it shows that presenting the gene as the unit of selection, as though this is all that is important in evolution is just plain foolish. If a change in something as fundamental as a gene can have no influence on the phenotype then clearly the gene is no more than a cog in a machine. It is the combined characteristics of the phenotype that result in survival and hence evolutionary change. And if the effect of a gene on the phenotype is limited, or heavily dependent on other factors, then its influence on the world at large, the extended phenotype concept that Dawkins hopes will “illuminate whole areas of biology in new ways”, is sure to be diminished as it will be dependent on other factors to a far greater extent. 

     It’s significant that the only convincing examples Dawkins gave of the extended phenotype concept involved parasites, organisms that we’ve always known as master manipulators. To conclude from this that genes in general are master manipulators, or that the extended phenotype is a general rule of nature is delusional. 
    All that we can take from The Extended Phenotype is that genes are a factor in natural selection, and even in ecology. But then, we already knew that.

    I have a confession to make. I wrote the above as I progressed through the book, but prior to reading the final two chapters. You can imagine my surprise when I found in them a de facto scuttling of selfish gene theory. Not that Dawkins would admit that, what he did was capitulate to his academic opponents by conceding that selection takes place at the level of organisms, then he attempted to reconcile the basis of his two principal books with his newly acquired view of evolutionary biology. Rather a difficult task you might think. Dawkins must have thought so too, because in the second edition of TEP a remarkable situation arose. The second edition gives the preface to the first, followed by a preface for the second. The two are worlds apart in difference of tone. In the first, written in 1981, Dawkins is apologetic, with just a hint of despair, as in “She has kept me going by believing in the project even through the times when I lost my own confidence.” That doesn’t sound like the Dawkins we know at all. In the second, written in 1989, the tone is upbeat and confident, as in a reference to “the now widely accepted selfish gene view of evolution.” That’s the Dawkins of old. My guess is that he was not confident prior to publication that selfish gene theory would survive his capitulation. It should not have survived. Dawkins engaged in some fancy footwork and more smoke and mirrors in an attempt to show that “the version of genic selectionism that can be attacked as naively atomistic and reductionist…is not the view that I am advocating.” But he’s loose with the truth, for in the preface to the second edition the gene is referred to as “the centre of a web of radiating power.” That definitely is naively atomistic and reductionist.
     
    Fancy footwork is not quite the right image; he prostrated himself at the feet of Ernst Mayr in an effort to be onside with “the grand old man of evolution”, then had the effrontery to claim that his reformulated interpretation of gene selection would be acceptable to Mayr. Not according to Mayr, who repudiated the gene’s eye view of evolution repeatedly throughout his career. Keep in mind that Dawkins rolled over, accepted organism selection, but still chose The Gene as The Unit of Selection as the sub-title of the first edition. It is true that unit and level are different concepts, but as Mayr said in 1997, “The term (gene as replicator) is, of course, in complete conflict with basic Darwinian thought…Since the gene is not an object of selection (there are no naked genes) any emphasis on precise replication is irrelevant. Evolution is not a change in gene frequencies as is claimed so often, but the maintenance or improvement of adaptedness and the origin of diversity. Changes in gene frequency are a result of evolution, not its cause. The claim of gene selection is a typical case of reduction beyond the level where analysis is useful.” Mayr has, in that single highlighted sentence, exposed the fallacy on which Richard Dawkins built a career.

    Now, Dawkins has a fondness for putting the knife into his opponents then giving it a little twist by using their names as labels for their alleged misunderstandings, as in “Washburn’s Fallacy”. It only seems fair therefore, that “The Dawkins Fallacy” should be added to the list. It goes as follows; “Evolution is the external and visible manifestation of the differential survival of alternative replicators.” (TEP p82) 
    POSTSCRIPT
    There's a valuable discussion of these issues in the comments that follow the article here.

    Comments

    BRAVO!
    I've spent years telling people that the Selfish Gene hypothesis is the worst kind of nonsensical pseudo-science, nothing more than a bunch of Just So Stories, but never had the energy to actually go through one of the wretched man's books again after reading one of them years ago and enumerate the fallacies. This will save me the bother...

    May Dawkins learn more by reading this very simple review of basic genetic concepts, including the concept of phenotype (question number 7): http://www.biology-questions-and-answers.com/gene.html .

    There is no shortage of egos in the world of science.

    Please explain why the dam-building by beavers is not testable? ( I am not a biologist, but I image that I - cold hearted as I am - could remove a beaver pup (or kitten or whatever it is called in English) from the beaver family - and rear it out of reach of any dam-building beavers.)

    regards

    Claus

    Steve Davis
    It was Dawkins, not me, who declared that The Extended Phenotype contained nothing that is testable. I think your idea is a good one, as long as the experiment could be guaranteed to involve no stress to the pup, (kitten? cub?) which might be difficult to determine. The more I think about it, the less likely it seems that it would be stress free, so it seems ethically invalid just to satisfy "idle" curiousity.
    I think we can avoid the ethical dilemma, by doing some bird-watching. I am pretty sure that many birds never see their parents build a nest ( for socio-economic reasons only known to birds, they leave their parents before the parents start a new nest). I conclude that nest-building for birds has a large genetic component (but I do not rule out some 'learning-by-doing' and imitation either, birds are not that stupid).
    I don't see much difference between nest-building by birds and dam-building by beavers (just one bunch of twigs or another).

    Did you notice that the Selfish Gene theory mirrors Adam Smiths 'Invisible Hand' metaphore ?
    RD has the 'selfish gene'-effect producing the (real) organism, AS has the (real) free market agents producing the 'invisible hand'- effect

    How do you propose to explain, say, the social behaviour of ants? A lot of ants (pretty similar genetically) cooperate to allow one or a few individuals reproduce. In a sense you can consider the ant-hill 'The Organism' and not the individual ants. They are, in this view, just subassemblies, like fingers ( with slightly more freedom of movement).

    Steve Davis
    I don't see much difference between nest-building by birds and dam-building by beavers (just one bunch of twigs or another). And not much difference to home buiding by humans or burrows by rabbits. Except that this is one occasion where Dawkins understates a position, his usual practice being overstatement. Its understated on this occasion because a beaver dam is much more than a phenotypic effect. A beaver dam is an ecological marvel that provides habitat for countless forms of plant and animal life. (Nests, burrows and houses do so also, but to a lesser extent.) But Dawkins does not want us to focus on an ecological marvel because that might distract us from the sterility of his gene-centric view of biology. Let's face it, it's natural wonders such as beaver dams that gets us interested in biology in the first place. There will be no Darwin, Margulis or Mayr of the future who is tempted into a career in biology by contemplating the emptiness of selfish gene theory. 
    I (and many others) have noticed the connection between Dawkins and Smith, it's a subject I hope to treat in more detail in the future.
    And if you are from Europe Claus, you might be able to do some beaver watching. I understand some beavers were recently relocated to a site in Nth Europe that was formerly a beaver habitat.  
    Steve Davis
    I'm sorry Claus, I overlooked your question regarding ant behaviour. You're quite right, the anthill can be regarded as "The Organism" as you out it, in fact DS Wilson has (from memory) argued that all social groups can in certain circumstances be referred to as a "super-organism." I think E O Wilson has done the most notable work on the social insects, that would be the area to look for more info.
    I don't recognise the effect of the Selfish Gene Theory on the young minds you suggest.

    I still marvel at the wonders of nature, and the SGT is just one angle. I read the Selfish Gene close to 30 years ago, and I was almost tempted to switch from Electronic Engineering ( I have a M.Sc) to biology by that book. (I don't think a Scientific Theory should be evalutaed by its how it influences young people - or its moral value - just how True (in Karl Poppers sense) it is).

    Did you ever read 'Unweaving the Rainbow' ? I think Dawkins addresses some of your concerns in that book.
    And I think Dawkins get just as much pleasure from watching nature that I do, I haven't met him, but I glean that from reading his books.

    You are correct, beavers has been introduced at a place in Jutland (in Denmark, I am Danish) and seem to thrive, and Sweden have a native population, too.

    We have several butterfly-species close to eradication, but furry, cuddly creatures are much easier to sell to the public, I am afraid. Nice to see, anyway - and they do create some public interest in biology.

    Steve Davis
    I don't recognise the effect of the Selfish Gene Theory on the young minds you suggest.  The reason I brought it up was not only my personal feelings on the matter, but also that Dawkins himself referred to a reader who read the first edition and became quite depressed. I think you are one of the lucky ones! I also think that many people would have been attracted to it because it relieved them of a sense of guilt for their occasional selfish desires. (If selfishness is genetic then there's no need for guilt.) But if Dawkins gave that impression then it was flawed, and their guilt was baseless, for reasons that I gave in the article.
    Thanks for the info on the relocated beavers. Are the Swedish beavers relocated or native survivors?
    Many thanks for the reference to E.O.Wilson. I have 'Nature Revealed' lying on my desk, but have to finish "The Voyage of the Beagle" first.

    My point about the ants is that an evolutionary theory somehow should explain this - and Dawkins (building on W.D.Hamilton, J.Maynard Smith an others) could have a point. As a model organism it almost seems possible to test some of coupling between genes and social behaviour.

    Sometime I sense a chasm between biologists, who do mathematics, and biologists, who don't. The last group seem to be put very much off by Dawkins.

    The Swedish beavers, I believe, are mostly native . Some may have been relocated in connection with the creation of wildlife sanctuaries, though (Sweden is 8 times bigger than Denmark, and with a very different geography and history. Most earth in Denmark has been tilled or grazed for more than 1000 years, not so in Sweden).

    I was wrong: http://findarticles.com/p/articles/mi_m1200/is_n15_v144/ai_14489012

    Hunters killed off Sweden's beavers during the 1800s. Then, during the 1920s and 1930s, conservationists reintroduced a small number of Norwegian animals, of which 46 survived in 11 locations in Sweden. The Swedish beaver population now numbers 100,000, says Ellegren. In Sweden, he and his colleagues obtained DNA from 31 beavers, 25 that lived in the same river system and six from varying distances away. They also examined genetic material from 15 Norwegian beavers and six Russian ones.

    Steve Davis
    Sometime I sense a chasm between biologists, who do mathematics, and biologists, who don't. The last group seem to be put very much off by Dawkins. There's a good reason for that. The predecessors of Dawkins had the strange view that they could explain many biological phenomena by mathematics. You only have to consider this for about 30 seconds to see the flaws. Even Haldane, for whom I have great respect, tended to this view, although I still think his famous remark about saving two brothers or eight cousins (or whatever the numbers were) was given in jest, after all he was in a pub at the time.
    That's interesting about the Swedish beavers, until recently I thought they were only native to Nth America. Now I'll have to tell the missus that I don't know everything.
    I beg to disagree on the role of mathematics. Mathematics is as much a 'thinking tool' as an 'explanation tool'.

    Trying to create even a very simplified mathematical model, e.g. in the ilk of John Maynard Smiths models, will help to clarify thoughts and ideas. For one self and for others.

    I have seen people going off in the other direction and ending up with a biology, which is wordy, imprecise and tedious (like reading Marx or Hegel). But then, I like mathematics.

    We also have a European Bison, the Wisent ( with small reintroduced populations in Poland, Russia and Belarus, none in Denmark, no space) At one time less than 50 was alive in Zoo and parks. Actually I am off to the Zoo myself, it is 10:20AM here in Copenhagen and I will try to take some pictures. Have a nice day - and thanks for the dialogue.

    Steve Davis
    It's time for my evening meal here in Nth Oz, and thank you.
    I've read your post a few times, & I'm really not sure where you're sticking the fork into Dawkins. While you're right that a phenotype is the combination of genotype & environment, doesn't the gene determine the degree to which the environment is to play a role? Some kinds of phenotypes are more influenced by environment than others. And the degree to which the environment is even allowed to play a role, I'd think, would relate back to the survival of the organism, and consequently to the passing on of genes. That is, some phenotypes are highly influenced by the environment specifically b/c that degree of flexibility gives genes an advantage; others are less tied to the environment b/c that degree of flexibility would harm them.

    It's the same with law-enforcement. Some laws are very context-specific, while others are more general. The degree to which a law is context-specific is determined in order to best suit law's purpose (just as the degree to which a phenotype is influenced by environment is meant to benefit the gene). So while the manifestation in which any particular law (or phenotype) is enforced may be context-specific, that doesn't mean that the context (or environment) is given carte-blanche in the law's formation.

    Individual bricks can have their own characteristics just as genes do

    I'm not sure what to make of the analogy between genes & bricks. The analogy that's given in every high school biology class between blueprints & buildings. So if an element of a blueprint has not effect on the building, then what's is its point? The analogy breaks down when you consider that these "blueprints" are passed on from one building to the next.

    On page 260 the following appears. “If a genetic change has no causal influence on bodies, or at least on something that natural selection can ‘see’, natural selection cannot favour or disfavour it. No evolutionary change will result.” Why is this a slip-up? ...It shows that the organism is the level at which selection occurs, not the gene.

    I'm not sure how this is a contradiction. Oversimplified, Dawkins' model is that genes play out in the organism, which play out in natural selection; and conversely, natural selection affects the organism, which affects the genes. The survival of the organism is integral for the survival of the genes.

    And lastly, what of sexual selection? I haven't read The Extended Phenotype, & I've a hazy recollection of The Selfish Gene, but I've always thought that Dawkins provided a sound explanation for why sexual selection is an integral part of evolution.

    Steve Davis
    Thanks for the questions. "I'm really not sure where you're sticking the fork into Dawkins." I think the last two lines satisfy that metaphor. And it really is a serious fallacy that he is promoting. I see on his web-site that the Dawkins acolytes are repeating the mantra, so what he's doing is spreading misunderstanding and ignorance.
    "doesn't the gene determine the degree to which the environment is to play a role?" Not at all. As Dawkins says himself, genes just are. Each gene carries out its own little role. The genes as a group interact with the environment to produce the phenotype, but Dawkins does not like to talk of group activity. "the degree to which a phenotype is influenced by environment is meant to benefit the gene". There is no meaning. You've unconsciously taken in Dawkins' flawed concept of "Darwinian purpose." "natural selection affects the organism, which affects the genes. The survival of the organism is integral for the survival of the genes." Exactly so. You've put the elements of the process in their correct order, which means that you understand natural selection better than Dawkins. Its not that he is not aware of these obvious truths, as the quote you highlighted shows, the problem is that he has an agenda that forces him to conclude from the process you outlined that the genes are organising the process.
    The genes as a group interact with the environment to produce the phenotype, but Dawkins does not like to talk of group activity.

    So the focus should be on groups of genes? If so, what different conclusions would you draw from that?

    Dawkins says himself, genes just are. Each gene carries out its own little role.

    I still don't see the problem here. Imagine a model similar to Adam Smith's - eg, it is not from the benevolence of the gene that we derive life, but from its regard to its own self-interest. Which is to say that a gene doesn't have to comprehend the whole of the organism in order to do its part, it just has to do whatever facilitates its survival. And when its actions or qualities don't facilitate its survival, it doesn't survive.

    From what I remember, he embraced group activity in order to further his point. This was the purpose of his discussion on ants & bees as discussed above, along with his discussion of seemingly selfless behavior in relation to one's immediate family. The main perceived contradiction, as he often repeated, was between our conceptions of selfishness & the sort he was discussing. Adam Smith's ideas encounter the same knee-jerk reaction when people ask how anyone would get along if citizens all just worked in their own self-interest. The response is that often its in our own self-interest to cooperate. Not to get into a long-winded political debate, but you don't have to be a capitalist, or even agree with Adam Smith, to see that self-interest on one level can cause cooperation on another, and to see why the individual unit - be it the gene or the worker - doesn't require knowledge of the whole - eg, the organism, or the economic sector - in order to do its part.

    The real question, which applies to both of these authors, is *when* it's in our own self-interest to cooperate, and when it isn't. If I'm interpreting Dawkins correctly, the answer roughly corresponds to whether an organism's genes are at stake.

    the problem is that he has an agenda that forces him to conclude from the process you outlined that the genes are organising the process.

    What's his agenda? I always thought his Selfish Gene argument was more along the lines of assuming that the gene is the individual "selfish" unit, and then seeing whether the implications of such an assumption match the real world.

    Steve Davis

    So the focus should be on groups of genes? If so, what different conclusions would you draw from that? The conclusion we draw from that is that the obsession Dawkins has for gene selection is just that, a mere obsession. It is the group that is selected.
    Imagine a model similar to Adam Smith's. The problem with selfish gene theory is exactly the same as for Smith's economic view. You can make all the comparisons you like but they are meaningless. Genes do not act alone. The lone gene is an irrelevant entity. Individuals do not act alone economocally. The lone individual is an irrelevant economic entity. While it can be useful to look at the world that way, it is utter foolishness to conclude that this is the way the world works. What's his agenda? I hope to deal with that in more detail in the near future. I'll just have to keep you dangling.

    What's his agenda? I hope to deal with that in more detail in the near future. I'll just have to keep you dangling.

    I'll be looking forward to it.

    As I side-note, I think that a sad side-effect of modern science's zeal for empiricism is that some scientists hide behind the numbers instead of putting their foot out & really saying something. I've been to so many conferences where I'll go up to, say, a poster-presenter and say something like, "Wow, that's a really interesting study. What do you think is going on?" Then they'll point me to a chart and give a little explanation. And I'll say something similar like, "Hmm, that's also really interesting. What do *you* think is *really* going on here?" And they'll point to the same chart, or to another one, & give a little explanation. And I'll be like, "Yes, I see that. It's really interesting. But what do you think is going on here? What's your theory?" & we'll go in lots of little circles. & I'm always kind about it - not like I want to poke holes in their study - but I just want to learn about the world.

    Anyway, one of things I've always admired about Dawkins is that he's not afraid to say what he really thinks is going on. I think he gets a lot of flack for this, b/c many scientists really just would rather hide behind their data. But there's always something going on, which - if the study was well-crafted - *should* be controversial insofar as it's in line w/some theories & not others. & you do your study injustice if you don't at least sketch the underlying theory, or if you aren't willing to speculate here & there about it. Maybe it's just a pet-peeve of mine, but science doesn't speak for itself, & if you're unwilling to risk saying something wrong, you shouldn't say anything at all.

    Ultimately of course the object of science isn't theory or opinion, it's learning about nature. As slanted or opinionated as Dawkins seems, he explicitly recognizes this in this short video clip (http://www.youtube.com/watch?v=eA86N8K4VBI). Maybe I'm just too quick to buy the rhetoric, but it's refreshing to read the work of an active scientist who is not afraid to say something that maybe wrong; that's kind of what's so fun about Scientific Blogging as well. & I think science would be improved & more engaging if more people were willing to take such risks.

    Steve Davis
    I'm glad you mentioned your questioning of alleged authority figures because I was beginning to get the impression that it was only bloggers for whom you have a healthy dose of scepticism! Ultimately there is no such thing as an authority. Each time we pick up a book, no matter who wrote it, we should have two questions in mind. What can I learn from this, and what can I criticise? With regard to the last of those, for me Dawkins has been the gift that keeps on giving.
    Dear Steve,

    could you please clarify what's your educational background? Frankly speaking, it all looks like you had reading comprehension problems and did not understand half of the book at least, sorry to say that. And you clearly demonstrate very poor understanding of mathematics (yes, there's a lot of mathematics between the lines).

    Steve Davis
    Sorry buster, the only way to demolish my arguments is to demolish my arguments. Can you do that? Personal attacks count for nothing.
    Steve Davis is a troll. On his own blog post.

    Steve Davis
    Yes, of course, but can you criticise the article?!
    Hi Steve Davis,
    Thanks for your article that made me think.
    My highest qualification is BEng Electronic Engineering. I am very much interested in evolutionary biology. The old man (Darwin) is my hero. And sorry for my lack of biology knowledge.

    This is the way I understand it. Ok here we go.
    Natural selection selects on what? It selects on adaptation to the environment.. Let me take an example which happens to be a non living thing- a ball of mud. Now if this ball of mud is in a dry cave, nature selects it. If it is near a river bank it will seize to exist. When a flood comes, of course the particles of the mud ball will exist but the essence of a ball will be destroyed. That is natural selection and for natural selection it is not a requirement for anything to be living. If it were a large boulder, it would have better chance of survival next to a river and after a flood, we would still call it a boulder and the essence of a boulder has survived. Now, if you can see it is not the crystals in the rock or the atoms in the rock but the notion of the boulder that has survived. Same goes the mud ball, it is the notion of the mud and ball shape that has survived in the dry cave. So what survives is nothing but only an idea or arrangement and what we label (notion or essence) that survives. However, there is no evolution as no reproduction in non living.
    Now we come to living things. There is a special thing in living things – well that what we call living at least-. Living things replicate. And they replicate or copy with a possible small change. When natural selection is applied to this, we have a recipe for evolution.
    That now established, let us find the unit of evolution, or the building block of evolution. Let us start from top. Let us take as topmost unit as the notion of “life on earth”. Life on earth is a candidate for this unit of evolution. Evolution is applied to it and gets selected. Well so far it has been selected. Life on Earth copies and changes and the notion of life has survived on earth since it started without interruption So there Si it is a candidate for the unit. Let us go a step down in our search for the unit of evolution. Let us take the next step as “a species”, as the unit of evolution and apply natural selection. For example, a certain red ant species. Well there is evolution there as well. Let us go further down and consider a colony of a certain species and apply natural selection, say red ant colony; well there is evolution there as well. Let us take the notion of an individual animal like the queen ant as a unit of evolution and apply natural selection (NS). We find evolution, so it also is a candidate for the unit. We go further down now to a cell on the leg of an ant as a unit of evolution, and we find evolution when we apply NS to it. It also is a candidate. Now I need to mention that going from the ant colony to the queen is a similar step as going from the queen to the cell on the leg because a colony is also like a large organism. From there now let us try the DNA strand as the unit and apply NS to it. We get evolution there as well. The DNA does not get selected directly as NS might not ‘see’ it is just like the NS might not ‘see’ the ant but the colony. We are concerned with the entity that changes and NS being applied to it or whatever it produces that NS ‘sees’. Then we come to the gene. And it also is a candidate. Here is the fun part; we go further down to the A,C,G, and T, and there we do not see them changing. They always stay the same. It is their arrangement that makes the notion of gene which produces something that NS can see. So we have to stop there. Evolution needs natural selection and also change to make evolution.
    So that is how I come to the conclusion that the best unit of evolution is the gene.
    Sorry for my biology. Thanks. Would love to hear what you have to say as I am always looking for moments where my concepts change.

    Thanks.

    Steve Davis
    Thanks for your thoughts Nishan. I'm sorry that this reply will be brief, but I have to get to work.
    Your description of natural selection and evolution is a good one I believe, although I don't think you have justified the gene as "the best" unit of evolution.
    My objection to selfish gene theory is that its promoters did not stop where you stopped. They took it into the realms of fantasy by trying to attribute to genes, qualities that genes do not have. For example, that genes "strive", or that genes "want" to achieve certain things. In short, they tried to give the impression that genes are living entities. They are not.
    Sir. I do not believe that Dawkins implies that genes "strive" or "want" at all. He makes it very clear in the EP that 'fitness for purpose' (expressed by the better survival and fecundity rates of the organism) causes differential frequencies of that gene, which, over time become established as dominant within the population. He then goes on to explain what factors prevent ' perfection'. I think Dawkins goes out of his way to describe this as a differential, not anthropomorphic, process.

    Steve Davis
    LB, you're quite right. But quoting Dawkins is like quoting from the Bible, there's always an opposing quote. His works are full of contradictions.
    Sir - you are clearly a thoughtful person applying himself to this article. This response doesn't do that effort justice.

    To quote you, "they tried to give the impression that genes are living entities. They are not."

    I don't believe at any stage that Dawkins made this representation - in fact, my memory of all his writings is that he went out of his way to repudiate this inference. Time and time again, he states that evolution is the result of differential survival rates at a gene level, caused by the interaction of the gene's phenotype and external environmental factors. Dawkins clearly and repetitively states that the gene has no character, no forward looking planning abilities and no anthropomorphic character. At least in my mind, there was never any confusion surrounding this.

    The usage of the word "Selfish" was more than adequately addressed in the introduction to the 30th Anniversary Edition of the "Selfish Gene" (ISBN 978 0 19 929115 1) It was not a retraction or a contradiction - just an explanation, and I think contextually it is understandable: "The critical question is which level in the hierarchy of life will turn out to be the inevitably 'selfish' level, at which natural selection acts? The Selfish Species? The Selfish Group? The Selfish organism? The Selfish ecosystem?"... Dawkins originally wanted to use the term "immortal" but realized that would be technically incorrect due to meiosis and mutation.

    What I will categorically agree with you on is that Dawkins can sometimes be a hard read. The Extended Phenotype (which he considers his best work) is too often a multithreaded argument, rebuttal of criticism and introduction of new concepts, rolled into one or two pages. It's difficult, but to claim his work full of contradictions as a result is disingenuous. I often find (for instance when showing in the EP that 'gene' can have multiple definitions) that the issue is one of my misunderstanding of semantics - but yes, the books could do with better glossaries!

    The problem with Dawkins (his agenda perhaps that you refer to) is that he has almost single handedly done more for the understanding of genetics within the scientific and lay community than any other person since Darwin. His problem therefore, being on that lofty pedestal, is that ANY inaccuracy in his work will be seized upon by his detractors as a repudiation of ALL his work (and let's not forget these people include fundamentalist creationists, flat earthers and people who think man co-existed with dinosaurs). He is in a terribly precarious position - he dares to offer alternatives to the status quo - he constructs hypotheses to be proved or disproved by future scientists (in doing so increasing our understanding even if he is wrong) yet he has people wishing him to fail.

    Steve Davis
    LB, your quote in defense of Dawkins is useful: "The critical question is which level in the hierarchy of life will turn out to be the inevitably 'selfish' level, at which natural selection acts?" He has exposed here his lack of understanding of the nature of life itself. Life is all about cooperation, so to waffle on about selfish levels, or the inevitabilty of selfishness, shows that the gene-centric view is based on a flawed understanding of the very matter that biologists are concerned with - life.
    Life is about co-operation ?
    Does the antelope co-operate with the cheetah ? Does the wildebeest co-operate with the lion ? Does the fish co-operate with the shark ? Do mammals fight members of their same species ? Do humans co-operate outside of their social group? Life is ANYTHING but co-operation. Life is a constant struggle against intra-species competition, inter-species competition and environmental factors. Without this struggle there would be no evolutionary process.

    HOWEVER --- you are again misinterpreting the intended meaning of the word SELFISH in Dawkins' book and this really doesn't do you any justice. The usage (as per Dawkins foreword) doesn't mean self interested or self absorbed - it means (or was intended to convey) only that the gene is the level at which natural selection acts - period.

    To imply ANYTHING else signals to me that you have either (a) not read the book or (b) not understood it. Dawkins himself says that more than half his book concerns the examination of the genetic rationale for altruism and the book could have been called the "cooperative gene" - but the point being that the emphasis is on GENE - the level at which natural selection acts. There is absolutely no anthropomorphic inference made in that book.

    I am not affiliated with Dawkins.net, nor am I involved in the Cult of Personality mentioned above. I just feel, having made the significant effort to read these books, that you have misrepresented the man and his work unfairly.

    Steve Davis
    I'm sorry LB, but I won't be taking this discussion any further. You have not taken the trouble to read my other articles on the matter, where some of your concerns are addressed. It's not that the matter is overly complex, it's just that the gene-centrics have made it unnecessarily complex. You have taken up the cause of Dawkins because of his position on creationism. That's not a good basis to defend his poor science. Your statement that the gene is the level at which natural selection acts shows the damage that selfish gene theory has done to biology, and to an understanding of life in general.
    Steve - I haven't taken up anyone's cause. I'm simply responding to your article. In front of me I have books by Stephen J Gould, Bryan Sykes, Neil Shubin, Hamilton and Williams. That's just in front of me. Behind me is an entire bookshelf of works that I value as highly as I do Dawkins' writings.

    You've made a frankly scandalous accusation that my (alleged) views of creationism have somehow affected my ability to critique your publications. That's totally unfair. And on the other hand you GLOAT that the Dawkins.net group haven't posted a response ! When someone does, you cut off contact ? It can't work like that.

    Irrespective, you have done nothing to disprove to me that the gene is NOT the level at which natural selection acts. I understand that the 'gene' can be defined at a number of levels. I understand that genes exist within organisms and depend on that organisms survival (at least until successfully reproduced). I understand that a group may suffer differential survival rates compared to other groups, but ultimately, the only traits that get passed on are genetic.

    Please, rather than cut me off because I haven't read your musings that up until now I didn't even know existed, (a) show me where they are so that I can and (b) please show me your peer reviewed papers from an established journal that sets out your case for evolution at a non gene level. I would be extremely and genuinely interested to see these, in much the same was I was genuinely interested to read this article.

    Steve Davis
    "His problem therefore, being on that lofty pedestal, is that ANY inaccuracy in his work will be seized upon by his detractors as a repudiation of ALL his work (and let's not forget these people include fundamentalist creationists, flat earthers and people who think man co-existed with dinosaurs). He is in a terribly precarious position - he dares to offer alternatives to the status quo - he constructs hypotheses to be proved or disproved by future scientists..." That was you talking LB, you have certainly taken up his cause, with a passion I think. "In front of me I have books by Stephen J Gould, Bryan Sykes, Neil Shubin, Hamilton and Williams. That's just in front of me. Behind me is an entire bookshelf of works that I value as highly as I do Dawkins' writings." It's no good having all those books on your shelf unless you read them critically. An example of your failure to analyse can be seen in the first quote I gave here, "he constructs hypotheses to be proved or disproved by future scientists." That's exactly what he does, yet you see no problem with that. One of the principles of the scientific method is that data is to be collected, then an hypothesis is constructed from the data. That principle was ignored in the formation of selfish gene theory. If you had read The Selfish Gene closely you would have noticed that Dawkins admitted that the hypothesis was formed, then the search for data began. Twelve years after the publication of Hamilton's first paper, when Dawkins' first book appeared, they were still searching for data, as Dawkins admitted. And they had the arrogance to refer to it, or allowed it to be referred to, as a theory! It's all a fairy story, without the redeeming feature of a moral. If you go to the top of the page you'll find a link to all my articles. But first, please, take all those books off the shelf and read them again. Critically!
    Your responses are a little ad hominem here. You should assume I can evaluate a book critically in as much as I should assume that you can publish a coherent, peer reviewed article.

    Not only do your articles NOT disprove the focus of evolution being at the level of the gene, they make lofty assertions that not only do you not prove (through collection and analysis of data) but have been disproved or rejected by your peers. You are no better than your accusations against Dawkins.

    Your articles also repeatedly and willfully misrepresent Dawkins at every turn ESPECIALLY with respect to the anthropomorphic qualities of the word Selfish, when time and time again the author has repudiated this meaning. It's very frustrating to see someone as obviously as intelligent as yourself refusing to move on.

    I feel that Dawkins work runs contrary to your personal beliefs of how life "should be" and that explains the issue. To date, I have not seen a crticial rebuttal of Dawkins' work, from anyone, anywhere, via the use of such scientific method as you describe. These are natural sciences - do you have the same criticism of Darwin, coming up with his Theory of Evolution without vast amounts of statistical data? Your objection would see us without Darwin, without Gould, without Dawkins and unable to hypothesize out of synch with field research. The objection has little merit.

    Steve Davis
    "These are natural sciences - do you have the same criticism of Darwin, coming up with his Theory of Evolution without vast amounts of statistical data?" That's my point exactly LB! You have shown everyone that you have not put any effort into this matter at all, because you should know, as any schoolchild knows, that Darwin did collect vast amounts of data, before during and after the voyage on the Beagle. He then spent years putting the data into a coherent form and was still not satisfied at the time of publication. How different to the gene-centrics, who hoodwinked the world with their half-baked ideas, raced to publication, did very nicely thank you, and still are doing nicely. Like I said, you need to read those books again.
    I see no evidence of this behaviour in your musings either. Note I call them musings as don't approach anywhere near the level required for publication or review. Does this mean i should discount your work equally? I really think you're too fixated on a number of misunderstandings on your part. Remember this is a web blog, not an internationally recognized forum subject to peer review. I can accept that without the organism the genes are useless (as Dawkins does) but to claim this invalidates Dawkins work makes a mockery of your readers..

    Thanks, Steve,

    We humans are living in the middle world. (Dawkins). We do not understand things travelling at light speeds, or space time continuum strong quantum forces because we are living in the middle world.

    Of course the genes do not have a mind of their own and they do not think or strive. We all know that. All they do is get replicated. But personification is a great way to explain things. This is our way of understanding things in the middle world. I think there in lies the explaining power of Dawkins. Here are some translations that I made according to the way I understood Dawkins.

    A gene strives to survive means: A gene blindly gets replicated and if its function would help its surviving machine to survive when NS is applied, the gene gets replicated more.

    a gene is selfish means: A gene blindly gets replicated and if its function would help its surviving machine to survive when NS is applied, at the expense of another surviving machine or even another gene, the gene gets replicated more.

    I have heard some people say that one law of thermodynamics states that particles strive to be in a more disorganised state than an organised state. It is just a way of explaining things. Not that particles have a mind and try to do certain things.

    As for the unit of evolution, you can think of it as the gene or the organism or group of organisms. It is only a way of visualising the complexity. Either way will do. We have to start some where.

    Thanks.

    Steve Davis
    Nishan, I have no problem with your take on these matters, although I still feel the use of the word "selfish" is not justified. And I think Dawkins would agree with your explanation. But Dawkins took it further and described genes as "sitting at the centre of a radiating web of influence" or some such nonsense. You said that "A gene blindly gets replicated and if its function would help its surviving machine to survive when NS is applied, at the expense of another surviving machine or even another gene, the gene gets replicated more." That is true, but it's not the whole story. It's not accurate to talk of a gene surviving or a gene increasing in the population, because it's the organism that must survive for those things to happen. And the organism is the product of its genome and its environment. Its not a gene that survives, its the gene-complex that survives. A particular gene in that complex is entirely subservient to the complex. It is turned on and off by the complex. It is replicated by the complex. Its particular function at a particular time is regulated by the complex. To talk of a gene surviving, or doing anything, can be useful in certain areas, but not in evolution. Which is why the term "selfish" is not appropriate.
    Sir- (a) Dawkins has thoroughly addressed the confusion with the word "Selfish" in later publications.

    (b) Gene survival does not depend on any one organism. That gene is present is many organism through meiosis across the ages. It is the DIFFERENTIAL rate of survival within the entire population that matters. To reduce this argument down to the survival of one organism is "reducio ad absurdium." [sic]. Arguing for the differential survival rate of a gene within the population is not illogical to me - it's quite rational.

    I'm not a Dawkins cult member - I just understand what he's saying, and what you're arguing, but you're coming at him from a tangential argument which isn't making much headway when you understand Dawkin's context.

    Beautifully written Nishan! I have come to believe that in Biology there may not be one single fundamental level at which evolution or perhaps any phenomena happen. Evolution happens at the level of ecology too! When temperatures change or atmospheric composition changes, the ecology will evolve and adapt to suit it. Similarly, a gene can be looked at as a unit of evolution.

    What "The Selfish Gene" provides is another way of looking at evolution and I think it's brilliant! As far as the lack of testability of his hypothesis is concerned, it is certainly no criteria to judge whether something is scientific or not. If tools limit scientific imagination, it is really sad. As long as he is stating that this is just his theory which your intellect may or may not choose to accept, though he will advocate it nevertheless, what is wrong with a little bold imagination?

    Anyway...You really seem to have a very logical and objective way of thinking and I would love to discuss science with you. If you haven't already read it, you must read "One, Two, Three, Infinity" by George Gammow.

    Devyani

    Thanks for that Devyani.
    my contact is nishan at usa dot com

    Steve Davis
    "What "The Selfish Gene" provides is another way of looking at evolution..." That is not acceptable. The selfish gene view is just plain wrong for many reasons, and many of those have been aired here and in other articles. To make such a statement here, you first need to disprove the arguments against selfish gene theory.
    Aitch
    I don't think the Selfless Gene Theory by Charles Foster is apparently much better....according to Steve Bishop
    http://stevebishop.blogspot.com/2010/09/selfless-gene-review.html
    ....but then, it's all pretty subjective, all this theorising, isn't it?
    Aitch
    Wow, great article. Dawkins may be the worst case of Emperor's New Clothing in the history of science.

    The preface to the second edition of the Selfish Gene is a brilliant intellectual swindle. He isn't presenting a theory, he says, but a point of view. "Rather than propose a new theory or unearth a new fact, often the most important contribution a scientist can make is to discover a new way of seeing old theories or facts”

    Really? If the new way of seeing doesn't yield a new theory, then what's the point? Scientists are not poets. They don't invent metaphors for their own sake. The metaphor is supposed to illuminate the real world. What Dawkins has done is science fiction, which he implicitly admits here (although he explicitly denies it in the book). He's created a compelling metaphor for the world, but not OUR world, not the one we actually live in.

    The essential problem, which SJ Gould recognized, is that genes are not agents. They are constituent parts of organisms, used for a particular purpose. Dawkins tries to argue for his point (which, by the way, is not SUPPOSED to be provable, so why argue?) by saying natural selection only acts on replicators, Replicators are things that reproduce themselves at high degrees of fidelity. But by this definition, there is no such thing as a replicator. Genes are replicated by a cellular process. Cells reproduce themselves, but not in the high fidelity way that Dawkins requires. This perversion of the notion of replicator--the agent that makes informationally perfect copies of itself--is the fantasy at the root of his science fiction.

    Steve Davis
    "genes are not agents." JRA, that's it in a nutshell!
    Genes are not agents? But what is then ?

    If the organism is an agent, yet the organism is killed (despite it's fitness for purpose) without reproducing, then the replication of that fitness is not passed on. The organism doesn't put tiny organisms in its gamete ---it puts genes/dna/chromosomes. Therefore we must assume that these are the units by which evolutionary change (differential change) occurs?

    If not, where does it occur??

    Steve Davis
    LB, in one sense you are right. I used the word in its independent/management sense, but it can be used to mean something much simpler as in something that produces an effect. But of course that is not the interpretation used by gene-centrics. They assume independence of action, which is ridiculous.
    I would have thought that gene-centrics - especially Dawkins assumes ONLY that one set of genes can experience differential rates of survival to their alleles, thereby favouring the reproduction of one over the other. I am yet to read any part of Dawkins where he implies genes act independently. I've read Selfish Gene, The Extended Phenotype, The Blind Watchmaker, River out of Eden, The Ancestors Tale and Unweaving the Rainbow - and this is a core theme. I'm not saying that Dawkins is right - I just don't think he's saying what you claim. Maybe I misunderstand ?

    Steve Davis
    "I am yet to read any part of Dawkins where he implies genes act independently." Then you did not read my article LB. How about where he refers to genes “playing out tournaments of manipulative skill"?
    Steve - that's an illustrative abstraction in a foreword describing an analogy to a Necker Cube - ie: how one initially sees organisms, and then one sees the replicating dna within - in much the same way as a cube drawn on a piece of paper can 'flip' orientation (a Necker Cube).

    Dawkins spends the remainder of the book (Extended Phenotype) repeatedly insisting that these "genetic fragments play out their tournament" as neutral pieces, not players as you suggest - they are akin to chess pieces, and the chess player could be considered as "the environmental factors influencing the differential survivability between the gene and its alleles". I feel you are simply misrepresenting Dawkins' work as I don't recognize your objections, having fully read the texts.

    I also can't see any problem with the assertion that genes can affect the environment to assist their survival. Beaver dams are one example. Humankind is an other example (albeit appalling and destructive). Some trees shed leaves or sap that prevent undergrowth. Birds build nests. Termites build air conditioned mounds! Where it benefits the organism, you see many clear examples of behavior intended to shape its local environment. Where it doesn't benefit the organism, you don't (pelagic fish for instance don't build houses). This behavior must have some genetic basis (even the ability to learn how to do it is ultimately a physiological manifestation of the genome and learning specialization).

    Steve Davis
    "I am yet to read any part of Dawkins where he implies genes act independently." Then how about the title of his first book?
    Steve - this is simply ridiculous. You tell me to read Dawkins critically AND THEN propose that we judge him by the TITLE of his book ?

    I've read the entire work yet you remain fixated on the title ?

    Sir - this does you a disservice.

    Sir, Let me quote Dawkins for you.

    "However independent and free genes may be in their journey through the generations, they are very much not free and independent agents in their control of embryonic development. They collaborate and interact in inextricably complex ways, both with each other, and with their external environment."

    "There is no gene which single-handedly builds a leg, long or short. But there may well be a single gene which, other things being equal, tends to make legs longer than they would have been under the influence of the gene's allele."

    So you see here, Dawkins is assuming other things constant when considering a particular gene, which most scientists do ( for ex. you physicists assume constant temperature and pressure all the time, just for the convenience of making an argument, though they very well know and assume others do too, that in reality this is not the case).

    Steve Davis
    LB, this is getting tedious. You stated that you are yet to read any part of Dawkins where he implies genes act independently. I've given two examples, I could give more, but you're like the knight in Monty Python; you don't know when you're done!
    And I've directly answered BOTH instances that you raised, refuting them. The tedium is watching you fixate on a title or phrase that you have actually misunderstood. Did you even read my responses? I took the time to read your prior postings.

    Many of your concepts ( evolution at the organism level) aren't controversial. But neither is the gene centric view. It causes us to think carefully about evolution (why for instance animals are rarely perfectly adapted due to genetic drift and external factors, why eusocial insects behave the way they do (Hamilton)) and advances the field exponentially than if only one viewpoint was used. May I remind you that there is no universal law in physics because of similar issues of conceptual scale ( and that's a field that has data available ad infinitum)

    Ripping chunks out of Dawkins on false premises discredits all the work you have done. Claiming he has done no data gathering over his 40 years as an eminent scientist is laughable You'll have to start citing sources discrediting Dawkins directly (peer reviewed please) because until they are fully refuted they remain an incredibly useful hypotheses that may bring us closer to the truth than the alternative.

    In the meantime, keep going. Even Einstein got it wrong.

    Steve Davis
    Dawkins implying that genes act independenty? Try this; "...we have the beguiling image of independent DNA replicators, skipping like chamois, free and untrammelled down the generations, temporarily brought together in throwaway survival machines, immortal coils shuffling off an endless succession of mortal ones as they forge towards their separate eternities." Not only does that clearly imply independence, it also illustrates the delusional basis of selfish gene theory.
    No. It doesn't. It doesn't give them sentience or free will. It merely describes the process. Basic English comprehension, unbiased, I'm afraid.

    So Steve. How does this work? You've given me your three best examples of Dawkins' "error" and I've refuted them as poor comprehension and / or an unabashed desire to insert anthropomorphic characteristics where there are none in order to further your own agenda. I can continue the process if you want but it will get 'tedious' as you say.

    As a 'scientist' however, when are you going acknowledge that all is not that which you portray? Remember that Dawkins bashing may find some supporters, but really you've offered nothing except poorly constructed criticism and fallacy. Not an eminent contribution to the process now, is it?

    LB, some good points raised - I can see why Steve didn't want to find any more quotes to misunderstand! As someone with such an obvious agenda (selfishness, how ironic), I'm suprised he's kept your comments on the posting at all.

    As you said before, he's obviously a man of intelligence and it's rather a shame to see this mental stagnation in so much of a talented wordsmith.

    Steve, I apologise if you've taken my comments harshly, I just don't understand your obvious hatred of Dawkins. Is it that his concepts are too radical for your liking?
    So how about Newton's "theories"? or Darwin's? Yes, they were wrong in part, but who isn't? Where does science come from if not theories and concepts that initially seem far fetched?
    The world IS round Steve...

    Gerhard Adam
    So how about Newton's "theories"? or Darwin's? Yes, they were wrong in part, but who isn't? Where does science come from if not theories and concepts that initially seem far fetched?
    ...and genes aren't selfish.  I get that Dawkins wanted to make a statement against the "good of the species" crowd, so proposing a completely opposing perspective garnered attention.  However, it isn't accurate, and it has created more false impressions than useful perspectives.  Every time someone wants to try and explain altruism, invariably they resort to the inexplicable nature of the phenomenon because of "selfish genes".  There are far too many examples of the concept of selfishness (not gene-centricism) as being fundamental to biology.  That's rubbish.  It isn't, it never was, and when it does occur, it is abnormal and usually destructive to the organism in question.
    Mundus vult decipi
    No. It doesn't. It doesn't give them sentience or free will. It merely describes the process. Basic English comprehension, unbiased, I'm afraid.

    Awesome post! I hate the nonsense Dawkins writes about evolution. Great analysis! I feel like he is a philosopher masquerading as a scientist at times.

    Steve Davis
    Thanks Natalie.
    Huh? What bit do you hate? The bit you haven't read or the bit you didn't understand? Great commentary and insight you've added there. I feel like I'm reading the minutes of a Luddite support network.

    How do you know what I did or did not read or what I did or did not understand. You and Dawkins are similar in that you both aren't very fond of proof. I am a lowly High School student and have hated the intentionality that Dawkins ascribes to genes in his "The Selfish Gene" since I read it. It isn't science. He's a philosopher who doesn't understand the difference between science and philosophy. As for memes and mimetics, please. What has that done for us? As a future biology student. I also think that your being a real jerk and not doing anything for science by behaving in such an immature manner.

    Honestly... If thats your view ( a problem with " intentionality") then you CLEARLY haven't either read or understood his work at all. Thats proof enough for me.

    To you and every other person reading this blog: despite the misquotes and the out of context postings used here Dawkins has NEVER argued for any form of intentionality (sentience or foresight) on behalf of genes. Don't let this nonsense get in the way of your understanding. Most of the objections here are based on dogma, not science.

    Steve Davis
    "Dawkins has NEVER argued for any form of intentionality (sentience or foresight) on behalf of genes." But he did. As I pointed out before, he argues black is black and black is white, depending on the circumstances. The dogma at the heart of selfish gene theory is that genes are living entities. They are not. Although the gene-centrics have tried to leave behind some of Dawkins' more extreme positions, this central misperception is alive and well. They express it as the ludicrous concept of gene-fitness. Fitness in biology is a clearly defined concept. It is a quality of organisms that reproduce themselves. Genes do not reproduce. They are reproduced by the cell. The purpose of this is to present genes as being at the centre of the evolutionary process. They are not. It is organisms and populations that occupy centre stage in evolution.
    " It is a quality of organisms that reproduce themselves. Genes do not reproduce."

    Organisms reproduce, but they do not reproduce themselves. I cannot create an exact replica of me. I can only mate with another female, and produce an offspring having genes from both me and my mate. So my offspring would have replicated copies of my genes and replicated copies of my mate's genes. So genes reproduce.

    Steve Davis
    Sorry Arjun, that's wrong. Since when has precise replication been a requirement for reproduction? The concept was introduced by Dawkins 30 years ago and it shows how illogical his thinking is. Precise replication gives stability, not change, And it's change that's necessary for evolution. But apart from that, in the example you gave, who reproduced the genes? The cells reproduced the genes. All gene functions are produced, controlled, organised, supervised by the cell.The cell is the primary level of life. Genes are not living entities, as they do not, and cannot, demonstrate metabolism, homeostasis, and reproduction.
    Sir, I mean no disrespect, but you have shown that you know very little of evolution and evolutionary complexity. I myself am a molecular biologist, and can talk from the point of view of molecular evolution.

    ". Since when has precise replication been a requirement for reproduction? "

    I never said that precise replication is a requirement for reproduction. At the same time, one would not expect reproduction to occur, if there is a high error rate in the copying mechanism of genes (DNA). That is why our Polymerase enzyme has such a low error rate of close to 1 in a billion nucleotides. Dawkins DID NOT say that precise replication been a requirement for reproduction. He said the HIGH fidelity is a prerequiste for reproduction and thus evolution. In this he is exactly right. High fecundity allows for a small error rate, and it is this window of gap that allows evolution. That is why evolution occurs over millions of years and not just thousands. Another point I would like to make is, if in fact error rate was high, you would see decrease in complexity of living organisms. This is because most mutations are bad, and natural selection would weed out any lethal mutation, and the ones that are good, will change at a more frequent rate, than the change in environment.

    "who reproduced the genes? The cells reproduced the genes"

    If you trace evolution back to whenever life started, or depending on our definition of life, if you go a little further back,
    the most probable event that started things of is a self-replicating molecule. This molecule is capable of Darwinian evolution because it must have had high fidelity, high fecundity, and high longevity. Now, in a recent paper, it was shown that (sorry i cannot cite the source, I read it a couple of weeks back) spontaneous self assembly of lipid molecules around the replicating RNA molecule occurs in the kind of unstable environment that existed 4 billion years ago. This would have been the first event to trigger assembly of a membrane and thus a primitive cell. And once you have room for "little" change in the copying of the RNA , any mutation that causes the RNA molecule to be more prevalent in number, would in fact be the ones that survive, and so on. So, you see here, that your question gives the illusion of a chicken and egg scenario, but it is not. It can easily be shown that genes reproduce cells and not the otherway around.

    "Genes are not living entities, as they do not, and cannot, demonstrate metabolism, homeostasis, and reproduction."

    Genes as we know it are not living. I agree with that. But this statement if flawed. Metabolism, homeostasis and reproduction are all strictly controlled at the level of gene expression.

    Steve Davis
    "If you trace evolution back to whenever life started, or depending on our definition of life, if you go a little further back, the most probable event that started things of is a self-replicating molecule." That is pure supposition Arjun. You cannot legitimately formulate a view of evolution based on a supposition. Freeman Dyson, for one, has an alternative view, but as a scientist with respect for the principles of science, he has presented it as a possibility. He has not raced to publication and become a headline grabber, unlike the gene-centrics. "Metabolism, homeostasis and reproduction are all strictly controlled at the level of gene expression." A gene does not reproduce itself. It is reproduced by the cell.
    A lot of things in science being suppositions have been legitimately followed through. In this case, this is NOT a pure supposition. A lot of surmounting evidence has pointed towards the Replicator-first model. As you might have already known from this article and many others:

    Vera Vasasa, Eörs Szathmáry and Mauro Santosa. Lack of evolvability in self-sustaining autocatalytic networks: A constraint on the metabolism-first path to the origin of life. PNAS, January 4, 2010 DOI: 10.1073/pnas.0912628107

    the metabolism-first model has taken a blow.

    Freeman Dyson's view of double origins is not parsimonious. I could go on, with the amount of evidence stacked up for replicator first model. But the point i am making is, rejecting legitimate pursuit solely because a hypothesis has not been proved is nonsensical.

    "He has not raced to publication and become a headline grabber, unlike the gene-centrics."
    This neither validates his theory, nor does it succeed in your pathetic attempt to defame the gene-centrics.

    And your last statement? Is that even an argument? I expected something better.

    Steve Davis
    Arjun, it matters little if life began with replicators. The fact that we have to deal with now (or rather, that you have to deal with) is that gene functions are controlled by the cell. This follows, as a matter of logic, from the point you conceded; that genes are not living entities. They do not replicate continuously, their replication is controlled by the cell the same as any other gene function. You said "And your last statement? Is that even an argument?" You're right. It's not an argument. It's a fact. The defenders of selfish gene theory have to face up to the fact that what they are defending is this: "I shall argue that a predominant quality to be expected in a successful gene is ruthless selfishness. This gene selfishness will usually give rise to selfishness in individual behaviour." (The Selfish Gene page 2.) That's the fantasy their science is based on.
    Steve - Gene functions are controlled by the cell, which in-turn are controlled by the gene! However you look at it, the phenotype will boil down to whether a gene is expressed or whether it is turned off. And what controls the turning off of these genes? Other genes. Ofcourse, there is an entire field of epigenetics , which studies changes in gene expression caused by things other than changes in gene sequence. Then again, in the end, all that matters is whether a gene is expressed or not.

    Remember, chromosomes replicate before cell division, in both Mitosis and Meiosis. Components of the cell are absolutely necessary for replication of chromosomes, but it is the genes contained in these chromosomes that dictate the cell what to do! This is the fundamental principle in cell biology. You claiming otherwise, without any data, or any peer reviewed papers says that you do not respect the scientific method.

    Steve Davis
    "Gene functions are controlled by the cell, which in-turn are controlled by the gene!" Arjun, you can't have it both ways. But you have exposed the flaw in the gene-centric view. Either gene functions are controlled by the gene itself, which you have stated is not a living entity, or by the cell, which is living. As non-living entities cannot control gene functions, at all times it is the living cell controlling those functions. So when you say " but it is the genes contained in these chromosomes that dictate the cell what to do" you are referring to genes in the plural, in other words, genes as a group, playing their role as a living entity. It is the group of genes that is alive. So to say "which in-turn are controlled by the gene" is an incorrect view, but is also the fallacy on which selfish gene theory is based. Careless language is the hallmark of selfish gene theory, but of course another reason that biologists have been reluctant to deal with this concept of life is that they are terrified of it. Fear of the unknown! Biology - the science of life, but they cannot describe life itself.
    It is not careless language Steve. It is language that you obviously cannot comprehend.

    "you can't have it both ways. "
    It is irrelevant what I can or cannot have. What is relevant is the fact, and it is a fact that genes control cells.
    "As non-living entities cannot control gene functions,"
    I see your problem now. You have become fixated that the selfish gene theory is flawed because genes are not living.
    It doesnt matter. Natural selection cannot distinguish living and non-living things! What it can distinguish is things that reproduce and things that dont.

    Although it is good to encourage scientific discussions on possible theories, one needs to get the basics right. Otherwise, any discussion is going to be fruitless, like an evolutionist trying to teach a creationist that the world is more than 6000 yrs old ( Please dont quote me on this, I have not taken up the cause of Dawkins because of his postion on creationism..besides, it is wrong to call it "position" , you can only take a position when there is doubt surrounding the issue). So I rest my case.

    Steve Davis
    You can rest all you like Arjun, but you cannot walk away from "...it is a fact that genes control cells." You've done it again; careless language leading to misunderstanding. Your statement implies that individual genes are doing their own thing. Now I do not know if you believe that, but the fact remains that it is genes working as a group, as a living entity, that control the cell. If you accept that, then you do biology a disservice by expressing it in a manner that does not make that clear. As for my "fixation" with selfish gene theory, I am certainly concerned that the concept attributes to genes a significance in natural selection and a role in natural selection that does not exist in reality.
    Dear Steve and other bloggers,

    I have the feeling the discussion is getting a bit too hot for the the thin ice we're all standing on. Please all mind your language, and this is all about language. So, a guy writes a book and takes it to his his editor. They talk about the title.
    Somebody, maybe Dawkins, proposes 'The selfish gene'. "Great!", the editor shouts, "that will sell millions!". The author, who just bought a new house, smiles. If he were honest, he would agree that he doesn't know exactly what he means by 'Selfish', and that there never was a biologist who gave a conclusive definition of the concept 'gene', but he doesn't want to disappoint his editor, and anyway, there's the mortgage to be payed.
    Didn't Darwin himself (PBHN) publish a book 'On the origin of species', while having the blandest notion what a species was? The good man thought that the domestic dog consisted of 3 or 4 species!
    And to Steve:
    "A gene does not reproduce itself. It is reproduced by the cell". What about the gene for DNA-polymerase. Or stiil stricter: pol-genes in RNA-viruses?

    Steve Davis
    "If he were honest, he would agree that he doesn't know exactly what he means by 'Selfish', and that there never was a biologist who gave a conclusive definition of the concept 'gene', but he doesn't want to disappoint his editor, and anyway, there's the mortgage to be payed." That's not a reasonable basis for a title for a science book. "Didn't Darwin himself (PBHN) publish a book 'On the origin of species', while having the blandest notion what a species was?" The difference between Darwin and the gene-centrics is that when Darwin was unsure he was honest about it. "What about the gene for DNA-polymerase. Or still stricter: pol-genes in RNA-viruses?" What about them?
    Steve,

    You may have a fundamental misunderstanding of the difference between two questions involved in the 'Unit of Selection' debate that is blinding you to other logical arguments made by LB, Arjun and others.

    You are mixing up the question of what is acting as the replicator and what is acting as the interactor (as Hull calls them), or Dawkins 'vehicle' (a more confusing term i think)

    Genes are the only level of entity that meet the requirements for replication- not species, groups, or individuals (even asexual individuals), nor even cells can replicate. It is irrelevant that cell macinery is used in the replication as it is the result- the replication- that matters.

    All other 'entities' from chromosomes to communities can be thought of as interactors. Natural Selection acts on these interactors, and may act on more than one. By acting on them, NS selects successful interactors. This selection determines the success of the replicators.
    Interactors are produced by the replicators, giving them enhanced survival over replicators who did not spontaneously produce interactors (by chance- not design) or those that produced inferior interactors.

    It is also repeatedly stated that the use of words of intention in these debates are merely to save time and space. We know genes don't "want" anything, don't "control" anyting, or aren't "selfish". Please drop this line argument as it is juvenile at best. You have been repeatedly dogmatic in your refusal to accept this metaphor, making previous arguments futile. Genes are not "doing their own thing" when it is stated they control the cell, it is merely that the genes that produce products that happen to make a cell replicate that gene do better than those that don't.

    Dawkins main point was that a gene that HAPPENS TO HAVE THE EFFECT of contributing to the interactor in such a way that that interactor can will do better than rival replicators in the gene pool. This may mean a phenotype of longer legs for running, or an extended phenotype of a higher frequency of calling for milk as a baby, affecting the behaviour of the mother. You don't question that the design, colour or chirality of a snail's shell is controlled by genes.

    Why then, do you disregard the ability of genes to influence (not absolutly control) behaviour, dam building, etc? And of course, environment has its role, as it is a phenotype. A beaver may genetically "want" to build a tall dam, but the surrounding wood type may influence the actual dam size. Even with dam building as a learned ability, genes which instill superior abilities to learn from conspecifics will thrive and dispay an extended phenotypic effect on the landscape.

    I am not a Dawkins fanatic, but in my research I become dismayed at seeing illogical refutations of his work such as "genes aren't selfish, they're not alive" that totally misses the point of a metaphor repeated again and again. It is most certainly not an argument against his theory, however else it may be flawed.

    And FYI:
    You create a hypothesis before collecting data. Look up wikipedia, something at the most basic level of science is you Observe, Hypothesize then test/collect data. Its really fundamental. Here actually, i'll paste it for you:

    A linearized, pragmatic scheme of the four points above is sometimes offered as a guideline for proceeding:[46]
    Define the question
    Gather information and resources (observe)
    Form hypothesis
    Perform experiment and collect data
    Analyze data
    Interpret data and draw conclusions that serve as a starting point for new hypothesis
    Publish results
    Retest (frequently done by other scientists)

    Dawkins is merely forming the first hypothesis, and as such does not require concrete data once he proclaims (which he does not) that it is merely what he thinks is happening. A comparison with Darwin is not valid as Darwin had a THEORY after gathering data on a HYPOTHESIS. Once again, schoolboy error.

    Steve,

    You may have a fundamental misunderstanding of the difference between two questions involved in the 'Unit of Selection' debate that is blinding you to other logical arguments made by LB, Arjun and others.

    You are mixing up the question of what is acting as the replicator and what is acting as the interactor (as Hull calls them), or Dawkins 'vehicle' (a more confusing term i think)

    Genes are the only level of entity that meet the requirements for replication- not species, groups, or individuals (even asexual individuals), nor even cells can replicate. It is irrelevant that cell macinery is used in the replication as it is the result- the replication- that matters.

    All other 'entities' from chromosomes to communities can be thought of as interactors. Natural Selection acts on these interactors, and may act on more than one. By acting on them, NS selects successful interactors. This selection determines the success of the replicators.
    Interactors are produced by the replicators, giving them enhanced survival over replicators who did not spontaneously produce interactors (by chance- not design) or those that produced inferior interactors.

    It is also repeatedly stated that the use of words of intention in these debates are merely to save time and space. We know genes don't "want" anything, don't "control" anyting, or aren't "selfish". Please drop this line argument as it is juvenile at best. You have been repeatedly dogmatic in your refusal to accept this metaphor, making previous arguments futile. Genes are not "doing their own thing" when it is stated they control the cell, it is merely that the genes that produce products that happen to make a cell replicate that gene do better than those that don't.

    Dawkins main point was that a gene that HAPPENS TO HAVE THE EFFECT of contributing to the interactor in such a way that that interactor can will do better than rival replicators in the gene pool. This may mean a phenotype of longer legs for running, or an extended phenotype of a higher frequency of calling for milk as a baby, affecting the behaviour of the mother. You don't question that the design, colour or chirality of a snail's shell is controlled by genes.

    Why then, do you disregard the ability of genes to influence (not absolutly control) behaviour, dam building, etc? And of course, environment has its role, as it is a phenotype. A beaver may genetically "want" to build a tall dam, but the surrounding wood type may influence the actual dam size. Even with dam building as a learned ability, genes which instill superior abilities to learn from conspecifics will thrive and dispay an extended phenotypic effect on the landscape.

    I am not a Dawkins fanatic, but in my research I become dismayed at seeing illogical refutations of his work such as "genes aren't selfish, they're not alive" that totally misses the point of a metaphor repeated again and again. It is most certainly not an argument against his theory, however else it may be flawed.

    And FYI:
    You create a hypothesis before collecting data. Look up wikipedia, something at the most basic level of science is you Observe, Hypothesize then test/collect data. Its really fundamental. Here actually, i'll paste it for you:

    A linearized, pragmatic scheme of the four points above is sometimes offered as a guideline for proceeding:[46]
    Define the question
    Gather information and resources (observe)
    Form hypothesis
    Perform experiment and collect data
    Analyze data
    Interpret data and draw conclusions that serve as a starting point for new hypothesis
    Publish results
    Retest (frequently done by other scientists)

    Dawkins is merely forming the first hypothesis, and as such does not require concrete data once he proclaims (which he does not) that it is merely what he thinks is happening. A comparison with Darwin is not valid as Darwin had a THEORY after gathering data on a HYPOTHESIS. Once again, schoolboy error.

    Steve Davis
    Anonymous, thanks for your interest. I'll deal with the main points as they are presented. "Genes are the only level of entity that meet the requirements for replication." This is a common theme of Dawkins, but my response is; so what? Natural selection and evolution are about survival and reproduction, not gene replication. " Natural Selection acts on these interactors, and may act on more than one. By acting on them, NS selects successful interactors. This selection determines the success of the replicators." Again, so what? Changes in gene frequencies are a result of natural selection, not a cause. At least you have not tried to argue as gene-centrics do that natural selection selects genes. I would also have to ask why it was necessary for gene-centrics to call an organism an interactor. " You don't question that the design, colour or chirality of a snail's shell is controlled by genes." I most certainly do!!! Please read my article Genes and Behaviours. "Why then, do you disregard the ability of genes to influence (not absolutly control) behaviour..." Because the intent of the gene-centric view is to give the impression that genes are in control, not just mere influences as you correctly point out. Surely you recall Dawkins saying that "genes sit at the centre of a radiating web of power"? Absolute nonsense. And where is the evidence that dam building by beavers is gene based? More nonsense. You have lectured me on the scientific method, but all you've done is prove my point. The gene-centics did not observe before forming their hypothesis. If they had spent some time in the wilderness they would never have come up with this fantasy.
    Steve, from all your arguments, it seems pretty clear that you are out to play on semantics , a trait attributed to a lot of pseudoscience.
    " Because the intent of the gene-centric view is to give the impression that genes are in control"
    Absolute nonsense. Just because YOU have misinterpreted it, don't make this generalization.
    The full context of the Dawkins' quote is
    "Certainly in principle, and also in fact, the gene reaches out through the individual body wall and manipulates objects in the world outside, some of them inanimate, some of them other living beings, some of them a long way away. With only a little imagination we can see the gene as sitting at the centre of a radiating web of extended phenotypic power. And an object in the world is the centre of a converging web of influences from many genes sitting in many organisms. The long reach of the gene knows no obvious boundaries"

    You have grossly misunderstood the intention, and arguing a strawman. Dawkins is taking a bottom up approach, which any determinist/materialist would and should do. By power, he obviously means the inherent ability of genes to encode information, and its ability to replicate.

    "If they had spent some time in the wilderness they would never have come up with this fantasy."
    A false premise , that one has to be involved in field study in order to come up with a legitimate hypothesis. Watson and Crick figured out the structure of DNA just by thinking.

    Gerhard Adam
    By power, he obviously means the inherent ability of genes to encode information, and its ability to replicate.
    Bottom up or not, it's simply not accurate.  Genes don't replicate and they also don't have any means of ensuring that they are expressed.  It's an overly simplistic approach that barely even works as a viewpoint.

    It conveys no particularly useful information, since its primary intent was to offset the idea of selection working for "the good of the species".  In that respect, taking a self-interested view is an important point, but to focus on genes, is to deal with an incomplete picture of reality.  This is especially true when one considers that Dawkin's definition of a gene isn't recognizable in any biological sense, because he's only referring to any segment of code that can be faithfully replicated, so the entire point is strictly about making copies.  However, a copy is insufficient, since it is only the expression of a gene which renders it relevant to future generations.

    Since there are clear examples of "selfish genes" (in the form of segregator-distorter genes) and they are invariably detrimental to the organism.  So it isn't a question of semantics, it is a question of whether such a description is useful for what happens, and in general, it is simply wrong.
    Mundus vult decipi
    Steve Davis
    Thanks Gerhard. Anonymous, you said, "Certainly in principle, and also in fact, the gene reaches out through the individual body wall and manipulates objects in the world outside..." The gene does nothing of the sort. The gene-complex might do this, but that's not how it's written, is it? Because Dawkins' intent is to show the gene in control. You said, "Watson and Crick figured out the structure of DNA just by thinking." There's nothing wrong with that at all, they presented a model that fitted the known facts and explained difficulties. And at that point thinking was the only option. But the gene-centrics had other options that they refused to pursue. They did not want to be confronted by evidence that might not sit well with their pet hypothesis. It's interesting that most of the early advocates for the hypothesis were mathematicians, who would not recognise natural selection if they were bitten on the backside by a crocodile. You might also want to check out the deliberate misrepresentation by Hamilton and Dawkins of works that did not fit their views.
    Steve,

    you must have too much time on your hands!

    Steve Davis
    You're right, it does take a lot of time to overturn 30 years of propaganda!
    But hey, it was worth it!
    Because of my efforts and those of Gerhard Adam, we don't get any references to gene-centred evolution theory here at Science 2.0 any more.
    The gene-centrics have quietly packed up and moved elsewhere, to quiet backwaters where they don't actually have to think.
    It's strange that so many people trained in science still blur the distinction between philosophy and science. It's obvious that Dawkins is interested in philosophy more than science as is evident in his recent NYT article. The questions that fascinate him are unscientific questions that no one can provide an answer for. What Dawkins' success says to me is that people are still fooled by semantic arguments that rest on logical fallacies (as your article states at length). WHy not just call his version of evolution "Dawkins Design." It has about as much validity as Intelligent Design because it makes evolution some kind of directional force. For that matter, what is the difference between Nabokov (another artist of language) and Dawkins with respect to their viewpoints on evolution. Nabokov wanted to refute Darwin by saying that evolution was aesthetically driven. He was a lover of art and art was everything so naturally he saw science as an extension of art. Dawkins sounds similar. Sure, he doesn't try to refute Darwin, but his desire to add to the theory of evolution is an attempt to inject subjectivity and intentionality into science. THis is certainly a strategy of the crazy postmodernists who claim that science is biased. I guess if you buy Dawkins' version of science it is; however, I find that version disturbing. Frankly, if Dawkins is supposed to be an ambassador for science and evolution, he is failing miserably. His desire to declare anyone who disagrees with him anti-science makes him a kind of fundamentalist who condemns Dawkins "non-believers" to scientific hell. I appreciate the article.

    Steve Davis
    "It has about as much validity as Intelligent Design..."
    Thanks for the comment.
    Yes, it has much in common with creationism.
    Everyone might want to take a look at an article in the 9 September 2011 issue of Science magazine --A Gene for an Extended Phenotype authored by Kelli Hoover and a number of other notable scientists.

    Seems Dawkins was right after all --it appears that a viral gene (egt) can induce phenotypic changes in gypsy moth caterpillars, getting them to eat until their stuffed so the baculoviruses can reproduce. The viruses get the caterpillars to climb up to the tree tops so when they die they rain down viruses and leave infecting particles all over the bark and tree leaves. Kind of cunning, right?

    Expect David Hughes to find a gene that that fungi use to turn ants into zombies, grow spore stalks and spread the fungi spores around infecting more ants.

    Gerhard Adam
    I'm not sure why you think this demonstrates an extended phenotype?

    In fact, it seems that the simplest explanation is to recognize that any parasite is simply modifying the environment they live in to enhance their own survival.  If that environment is another living organism, then we would expect nothing different. 
    The viruses get the caterpillars to climb up to the tree tops so when they die they rain down viruses and leave infecting particles all over the bark and tree leaves. Kind of cunning, right?
    ... and how is this different from making you cough, or bleed out?  Has it become different because some particular behavior has changed? 

    So when we humans modify an organism by genetic engineering, are you suggesting that is a gene that is responsible for making us do this to ensure its survival in future generations?  After all, that's what you're suggesting by giving such credit to the virus.

    The problem with Dawkin's extended phenotype, is that it credits the gene with something the gene can't actually do.  It cannot ensure it's own survival, nor can it ensure that it is actually expressed.  The point that a gene is used to modify the behavior of the caterpillar is as irrelevant as arguing that a gene is used to give you a fever.  While it is obviously true at one level, it is true only in a trivial sense (i.e. we are all the product of genes, although that doesn't mean that genes determine the outcomes).

    In one respect, Dawkin's wanted to counter the group selection ideas which suggested that things were done for "the good of the species" and replaced it with an equally inflexible position by suggesting that everything was done for "the good of the genes".  All that does is replace who you consider the "group" to be.
    Mundus vult decipi
    You seem to miss the point --this behavior-alterning gene is encoding self destructive behavior on the host. Coughing and sneezing is a host response to infection (or perceived infection, ie. allergy); two entirely different things.

    Gerhard Adam
    OK, I specifically mentioned bleeding out (such as in the hemorrhagic fevers), which is not a response to infection, but the means by which the virus spreads.  If we consider other parasites, then the effect is even more noticeable, since specific behaviors are elicited to facilitate the spread of the parasite.

    Even coughing, which as you pointed out, is the body's response to foreign substances in the lung, is being exploited by the virus to facilitate it's spread as an airborne agent.  The mere fact that it enhances the spread of disease, strongly suggests that it is actually an exploited side-effect instigated by the virus.  There are many other instances, such as the hijacking of the immune system, etc.  Each represents the parasite's ability to exploit the environment in which it exists.  You are simply presuming that a behavioral change is somehow a more unique exploit than the other methods employed.  This simply suggests that you are biased towards neural functions rather than the other systems that can be utilized in this fashion.

    However, other examples are also easily observable in well-known cases, such as rabies, where the behavioral changes are quite obvious.  We already know that meningitis and herpes can cause dramatic and radical changes in the brain and consequently behavior.

    For other examples, consider toxoplasmosis in humans:
    http://schizophreniabulletin.oxfordjournals.org/content/33/3/757.full

    I realize that one must be careful in distinguishing those changes that are simply the result of damage due to infection, versus those that are specifically being exploited by the virus (or other agent) to promote it's distribution.  All examples of the latter case, must be considered equally as "clever" as manipulating a caterpillar's behavior.

    Ultimately the problem with the "extended phenotype" is that it can be made to mean anything that even indirectly involves the organism, including learned behaviors.  This is gibberish, since the gene can't be responsible for something in which it plays no role.  In short, it's simply a redefinition of an equally useless term; "instinct".
    Mundus vult decipi
    Gerhard Adam
    Human genes constitute only about 1.5 percent of the genome, whereas regulatory elements appear to take up about three times as much space.
    http://www.sciencecodex.com/read/millions_of_new_regulatory_elements_found_in_human_genome-79640
    Perhaps this might also shed light on why focusing on the genes alone (i.e. selfishness) misses how things actually work.
    Mundus vult decipi
    Steve Davis
    And Gerhard, the other thing that stands out from Marcia's comment is that once again we are expected to believe that parasite behaviour is representative of all organisms, an issue I referred to in the article.
     I never cease to be amazed by the mesmerising power of the selfish gene hypothesis.
    No matter how many instances are given of the faulty reasoning behind it, people are clearly stunned into incomprehension by criticism of Richard Dawkins. I think that Dawkins must establish himself in the minds of readers as being such an overwhelmingly decent and honest person, and I have no doubt that he is, that they cannot accept that he sometimes misrepresents, which he clearly does.
    Gerhard Adam
    Unfortunately there's a certain appeal to the "selfish gene" concept, because it closely resembles another related fallacy; "DNA is destiny".  No matter how well we understand it, it is tempting to find an "suicide gene", or an "obesity gene", or whatever.  It simplifies a complex idea by promoting the notion that there is a singular gene-trait relationship.

    Instead, reality indicates that genes aren't nearly as precisely defined as one can hope.  A significant number of genes are shared in varying capacities and controlled by regulatory elements that further erode the singular relationship of gene-to-trait that people like.  As a result, the entire process becomes muddied and unclear.  In a way, it's like the discomfort people get from the Uncertainty Principle in Physics. 

    The "selfish gene" idea seems to provide order into a complex system, just like determinism seemed like a reasonable view of the universe until quantum mechanics arrived on the scene.

    In my view, biology is vastly more complex than physics, but it doesn't lend itself to the mathematical precision of physics because it is a system that consists of millions of variables, that continuously vary.  In addition, biology has the ethical barrier of having to deal with living systems, instead of inanimate systems that can be disassembled at will.
    Mundus vult decipi
    Steve Davis
     "...it's like the discomfort people get from the Uncertainty Principle in Physics."
    Exactly Gerhard. 
    Gene-grasping is an attempt to find a solid foundation in a biological system that has no solid foundation, as it operates on the principles of quantum physics. 
    The system is one of process, variables as you put it; there is nothing solid to grasp.  
    This discussion has been useful to me. I'm fairly new to all this but am currently reading avidly and am fascinated. I have, of course, read Dawkins and I am a fan but I'm not a blind follower and am interested in further exploring some of the criticisms which have been aired here. I have been interested by your arguments Steve although on balance I am still more convinced by Dawkins' position and the objections posted by LB, Arjun and an anonymous poster. So, I wonder if you could point me to any literature that supports your point of view more comprehensively as I would really like to more fully understand this controversy.

    Gerhard Adam
    One good source would be The Philosophy of Biology by Elliot Sober.  Another good book is Evolution in Four Dimensions by Eva Jablonka and Marion Lamb.

    It becomes easier to see that the difficulty is that the gene isn't nearly well defined enough to be credited with what Dawkin's wishes.  In addition, with the problem that a gene may not be expressed, and that it is rare that a single gene is actually responsible for a specific trait, it is easy to see that the premise is flawed.  After all, it makes little sense to talk about a "selfish gene" and then require that the genes "cooperate" in order to express a trait.

    It is also important to remember, that Dawkin's was presented an alternative view to the group selection notion of "for the good of the species" perspective.  So while it may provide an interesting or more striking approach to the problem, it shouldn't be taken too literally as how the process actually works.  In general, Dawkin's simply tends to overplay his hand on this.  There's no question that genes are critically important, but it's like arguing that words compete for expression in books, or that notes compete for expression in songs.  One example of a gene that could be considered "selfish" is the segregator-distorter gene, which is detrimental to the host, so the "model" just doesn't work.

    In the most obvious problem, it makes little sense to have something like a human being with approximately 10 trillion sets of genes (one for each cell), all "voluntarily" giving up control to allow for the possibility of replication into the future, by giving everything over to the germ cells.  This doesn't indicate "selfishness", but represents the epitome of cooperative behaviors, if you wanted to assign such an attribute to the gene.  It indicates that the gene is subsumed to a role within the organism, and not the driver.
    Mundus vult decipi
    Steve Davis
    Gerhard, thanks for your input.
    Matt, other sources include DS Wilson's series of blogs titled Truth and Reconciliation for Group Selection, in which Wilson does not hold back in his criticism of Dawkins, but he does make the mistake of accepting some of the assumptions of the selfish gene hypothesis, in particular the work of Hamilton on inclusive fitness, which is basically nonsense as was all his work. There is nothing of value in the selfish gene idea.
    Then there's Gabriel Dover's Dear Mr Darwin, an excellent book. Here's a quote that will interest you, 'There's no such thing as a free standing self-replicating molecule in biology, and there probably never has been. DNA relies for its replication on tens (maybe hundreds) of protein enzymes." So there you have it, the basis of the hogwash is...hogwash. What's really disturbing however, is that the gene centrics know all this but choose to ignore it. 
    But an easy place to start would be the twenty or so articles that Gerhard and I have posted here on this subject.
    If you are still a fan of Dawkins after digesting those, I'll be surprised and I would hope for, in fact I would expect, a rebuttal.
    Cheers
    Gerhard Adam
    Steve

    Thought you might find this link interesting:
    http://www.theatlantic.com/magazine/archive/2011/11/e-o-wilson-s-theory-of-everything/8686/?single_page=true
    Mundus vult decipi
    Steve Davis
    Thanks for that Gerhard, a fascinating article.
    I think their idea of genes for sociality is wrong however, genes are by nature social, although that's an overly simplistic way of putting it. Probably better to say that it is genes as a whole that create sociality. We don't have to look for something unique or something pertaining to particular life forms, sociality is a feature of life itself.
    "There's no question that genes are critically important, but it's like arguing that words compete for expression in books"
    Obviously words don't 'compete' for expression in books but I think it's a perfect analogy for me to try and convey what I believe Richard Dawkins means, since the use of words is often based on the environment.

    For this to work, I have to ask you graciously that you don't put too much emphasis on the fact that books are created by intelligent design, that words and books don't reproduce (but rather, might inspire a future author - which is what I will be using in the place of reproduction).

    Here goes: An author writes a book and decides to invent a word to describe something, it might be that with the growing awareness of the limited nature of global resources the author decides to coin the word 'recycle'

    This term 'co-operates' with the other words in the sentence to form a coherent idea, and the term itself is widely understood (due to the context it is used in) to mean 'to extract useful materials from garbage or waste'

    Now, another author decides to write a book, and a situation arises in which he feels the word 'recycle' will adequately display his idea - so he uses it. And thus we see how the word 'recycle' begins to be replicated.

    As you well know, the environment plays a massive part in natural selection, and with the amount of carbon in the atmosphere increasing, the word 'recycle' begins to be used more and more and more.

    The word itself has done nothing consiously to get itself replicated, it just so happens that the abilities it instrinsically has are suitable for alot of authors.

    I realise that I am not a very good writer at all, and that the analogy isn't the best either, but I hope I managed to just about convey how non-consious entities can become more numerous due to the fact that their properties happen to suit them to their environments. It was a bad attempt, I know.

    Steve Davis
    Your analogy is actually quite good. A good description of selfish gene theory, that is. Unfortunately, the selfish gene idea does not describe what actually happens in real life. Because what the idea fails to convey, is that genes are not visible to the forces of natural selection. They are not visible to natural selection because "gene" does not equal "trait." All that a gene does is synthesise a protein. Is a protein a trait? Obviously not. The protein has to undergo further processing before finally making a contribution to a trait. It is the chemical processes taking place inside the cell that produce traits that are then visible to natural selection. Traits are the products of living systems. Genes are not living systems. The fact that gene numbers rise and fall due to the operation of the forces of selection has little to do with evolution because the existence of a gene within a genome is almost irrelevant, as the gene is only activated by the cell. The implied message of the gene-centrics that genes have independent power or influence is a fallacy.
    - Thankyou very much for your time and for replying so quickly and fully -

    I thought that was where the gene-complex comes in. I'm not a biologist or a chemist so I could be way off, but is there a series of synthetic processes in which genes turn what we consume into the proteins which make muscle or saliva and many hundreds of other things? If this is so (which it might well not be if I have grasped the idea wrongly) then it seems fairly sensible to assume that, with the multitude of genes throughout biology, their re-arrangement (through darwinian and sexual selection - and also possibly parasites merging with hosts) would produce not only different substances but also different patterns of substances like hair and fingernails.

    Gerhard Adam
    Actually it's even more complex than that.  Genes interact with one another, but they are also dependent on being "turned on" or "off" and also when, how frequently, duration, etc. that will all affect the final results.  That's why each cell in your body can share exactly the same genetic information despite being radically different in how they actually manifest (i.e. heart cell vs brain cell vs liver cell).  This simple example easily illustrates how the DNA and genetic information is insufficient to explain any particular trait by itself.
    Mundus vult decipi
    Hi, Gerhard. Thankyou for replying.
    I think I'm getting a bit woolly minded here. Just so I can adjust myself to the right level, could you give me what you would definitively call a 'trait'.

    Gerhard Adam
    Sure.  A trait is any variant that represents a characteristic of the phenotype that can be either inherited or influenced by the environment.  An obvious example would be something like blue eyes, although it could be more subtle such as the limitations of muscular development.
    Mundus vult decipi
    If a person has blue eyes then it is inherited, as you mentioned, but what is it that's passed on during reproduction - aside from genes - that could explain them?

    Gerhard Adam
    I'm not sure I understand your question.  Genes are certainly responsible regarding the probability of inheritance (although a small but finite probability of a mutation could occur - not likely).

    However, blue eyes are interesting because they are caused by genes, indirectly, since they occur when a normal gene fails to produce the normal amount of melanin in the iris.  So, the problem isn't the gene, but rather that there is a switch which prematurely "turns off" the gene before the normal (i.e. brown) amount of melanin is produced.

    In other words, brown and blue eyes share exactly the same gene.  What is different is the "switch" that causes reduced melanin production by that gene to manifest the trait of blue eyes.  So the explanation is that blue eyes are produced by the failure to produce brown eyes.
    Mundus vult decipi
    Steve Davis
    Further evidence that the gene influence for eye colour is minimal, is the fact that eye colour can change after a person suffers physical trauma.
    This happened to the climber who was left on Everest because his friends thought he was dead.
    It's also been known to happen to people who are injured in fist fights.  
    Steve Davis
    That's why each cell in your body can share exactly the same genetic information despite being radically different in how they actually manifest (i.e. heart cell vs brain cell vs liver cell).
    That's a very useful point, Gerhard.
    It shows that genes are subservient to the needs of the cell, and ultimately to the needs of the organism.
    And it shows why the gene-centrics obssession with gene replication is misleading.
    That particular fallacy flowed from Fisher's "Fundamental Theorem of Natural Selection", which was merely Fisher's description of changes in gene frequencies as a result of natural selection.
    But the gene-centrics twisted his theorem (mathematical explanation) into a theory of natural selection, (something quite different) in one of the most brazen deceptions in the history of science.
    Hi Steve, thankyou for replying.
    Gerhard also mentioned that genes are 'turned on and off' in certain specialised cells. Surely this could outline further the way in which the genes influence an organisms development. I don't see why the 'needs' of the organism can't be the 'needs' of the genes.
    You may argue that genes are not alive when organisms are but the line between alive and not alive it really a very hazy thing. There must have been a point in history when something not alive developed into something 'alive', though I suppose this idea is at the mercy of how one believes life began in the first place.

    Gerhard Adam
    I don't see why the 'needs' of the organism can't be the 'needs' of the genes.
    Try thinking about it like this.  A gene cannot know what the external needs of an organism are, because that would require a direct, quite specific, cause and effect relationship between the gene and the trait.  So, a gene can't "know" that an organism needs to run fast, etc.  Those are traits that will be selected for in the environment and circumstances the organism is in.

    If running fast is useful, then that trait will tend to be selected.  If running fast is unnecessary, then such a gene will tend to be irrelevant and/or disappear.  However, it is entirely possible that such a gene may still persist over a long period of time, simply by chance and suddenly become relevant when some environmental change causes the ability to run fast to become important.

    In that case, the genetic potential of being able to have the muscles properly respond to training and conditions suddenly becomes an immensely important trait and (in the extreme) may be sufficient to ensure that this organism is capable of out-competing all others and may represent the sole ancestor of future generations.  In other words, all offspring are significantly more successful that the trait eventually becomes fixed in the population.

    In short, the gene has no "interface" to the external environment with which to react or with which it can be selected.  It is the manifestation of a particular trait that will determine its usefulness, but the gene is powerless to exert any influence in the matter.  It can only do whatever it does.  The rest is up to the organism, the environment, and circumstance.

    It is also important to remember that only the whole organism is ever "selected", so it is meaningless to consider a singular gene has having any meaningful way of "competing" for selection in the future. 
    Mundus vult decipi
    Gerhard Adam
    BTW, your point about genes being alive or not is also a useful one to think about.  If the gene isn't alive, then it simply becomes an a process that is irrelevant in discussions about selection, just as respiration or digestion would be.  However, if the gene is alive, then an important question is to consider what it's "environment" is.

    I would suggest that it is plausible to argue that the gene can compete for inclusion in the germ line or potentially among other genes, but it would have to be confined to those aspects of its environment in which it can actually be selected for, and for which the results would be tangible.

    While I don't know anything about you personally, but consider that by suggesting that genes can compete for selection through the organism, is a bit like arguing that you were born simply so that your wife would have someone to marry.  It's a bit of a stretch. 

    (In other words, you could not have foreseen the circumstances that would've given rise to the individual you married, so it's something that can't be postulated as a cause).
    Mundus vult decipi
    Steve Davis
    "...the line between alive and not alive is really a very hazy thing."
    Not really Jack. (These are good points you are raising, by the way.)
    The question of life only seems hazy because biologists in general and gene-centrics in particular, have been reluctant to decide what actually constitutes life.
    The gene-centrics are reluctant because they understand that any realistic evaluation of life will conclude that its essence is cooperation, as I explained in What is Life.
    Once it's understood that life is cooperation, then it follows logically that life is about groups, and this they cannot accept because the selfish gene idea is all about promoting the ideology of individualism, as I explained in Twelve Misunderstandings of Evolution.
    Hey, Steve
    I'm a little confused about your view of the gene as being dependent on the cell. Is it not the case that all cellular functions are intimately effected by interactions with the genome an very short timescales, i.e. the gene controls the cell? You may have noticed a trend amongst your critics here; that you are misrepresenting Dawkins. Perhaps you should consider this as a possibility. I grew up with Dawkins and it has never occured to me that he was implying intentionality to genes. The metaphorical use is understandable, perhaps unavoidable. I have been perplexed by Dawkins' increasingly lengthy assurances that he is using metaphorical language, in that I couldn't imagine that someone would think that the "selfish gene" ascribed motive to a molecule. I think I understand now. You must realize that this is a basic necessity in the sciences; I do not believe that plants or electrons have will, but when talking to a fellow scientist I do not hesitate to describe what they "want" to do. If they were to quarrel over this I would think them intentionally ignorant in the matter. Perhaps you should reread Dawkins while assuming this is the case, suspend your disbelief, and re-evaluate. Also, keep in mind that TEP is one of the few works written at the academic level that has seen a popular audience, and in having students attempt it, perhaps one in three can "get" it. Not an easy work, and it would be no shame to be in the 2/3 that are defeated in the attempt.

    Good Luck!

    Steve Davis
    Rick, thanks for your interest. "I grew up with Dawkins and it has never occured to me that he was implying intentionality to genes. The metaphorical use is understandable, perhaps unavoidable..Perhaps you should reread Dawkins while assuming this is the case, suspend your disbelief, and re-evaluate. ."

    Dawkins would have more credibility if he did not constantly contradict himself. He has stated that the selfish gene is both a metaphor and not a metaphor, so he is totally confused.
    Keep in mind that the intent of the selfish gene hypothesis is to present genes as acting to increase their numbers. This is simplistic nonsense. Genes do not act, as this implies independence. They function as directed by the cell.
    Which brings us to your point that "the gene controls the cell."
    I'm not surprised that most readers of Dawkins think this is so.
    It is wrong because the genome is not a collection of genes with their own agenda.
    The genome is the operations centre for want of a better phrase, for the cell, the cell being a living entity. The genes in that cell are not living entities. Just as a compound has different qualities to the elements of which it is comprised, so the genes in the genome take on different qualities in that the genome acts to preserve the cell and the organism.
    Gene-centrism is based on a number of incorrect assumptions, most of which I have covered in my other articles.
    This is interesting.
    I read the article and the thread, and incidentally I read The Selfish Gene about 15 years ago (and wasn't completely impressed, IIRC).
    I don't know *what* is right, based on the arguments – but I can intuit *who* is right, based on their arguments.
    Interesting, huh?
    Sorry Steve-O.

    Steve Davis
    Don't be sorry, and don't be bashful.
    Just tell me where I'm wrong.
    I was wondering, since you disagree with Dawkins, what you thought about the somewhat recent reaction he had to E.O.Wilson's paper that claimed to disprove kin selection theory. Dawkins behaved like a religious zealot and thought that it was a disgrace that Nature published the paper. The paper claimed that genetic relatedness simply couldn't be used to predict altruism at least according to their mathematical models. I think kin selection theory is attractive to Dawkins because it is obviously centered on genes being the important feature that dictates behaviors like altruism. I am curious as to your thoughts on this issue.

    One more question. How do promoters and enhancers fit into his selfish gene theory? Do these genes also "decide" upon their location within the genome so that they are near a certain promoter or enhancer? What about epigenetics? How would his theory reconcile with the idea that outside environmental factors can methylate histones and alter gene expression and that these effects can be passed down to the next generation? Seems like Dawkins is changing the driver of evolution completely. If not, then he is not saying anything new. The thing is, his language is nebulous rather than giving experimental data then a clear interpretation of that data.

    Gerhard Adam
    Actually the problem is much simpler to describe.  An unexpressed gene cannot be selected by any definition.  Since a gene can't control whether it is expressed, then it cannot serve as "the" unit of selection.  There is absolutely no doubt about the important role genes play, it just isn't up the exclusive level that Dawkins wants to portray.
    Mundus vult decipi
    Thank you very much. Great point. Certainly, genes play a role in evolution, but that's nothing we need Dawkins to elucidate. I agree with you, I am just not aware of any of his argument concerning the exact problems you bring up.

    Gerhard Adam
    The problem is that I'm not sure how seriously to take Dawkins, although I know many people have taken him literally on this.  In the first place, his definition of a gene is absurd because it is considered only to be any replicating portion of the DNA, regardless of whether it would actually be considered a gene normally or expressible as a gene. 

    More to the point, Dawkins was attempting to focus attention on the gene as being the driver of evolution against the group selection view.  He was trying to demonstrate that life is a product of the unbridled selfishness of the gene and not altruistic group behaviors.

    Of course, both extreme viewpoints are wrong, but such simplistic explanations rarely have the necessary depth to approach the truth.  In my view, part of the problem here is that there are simply too many attempts to try to find a singular explanation for what is clearly a widely varying set of phenomenon for which it is highly unlikely a single set of rules would consistently apply.  In short, biology is much more nuanced than many would like it to be.
    Mundus vult decipi
    Steve Davis
    Thanks for your questions Natalie, they are most relevant.
    My reaction to his response to Wilson was exactly the same as yours; this is religious zealotry.
    But we should not be surprised by this.
    What sort of scientist would refer with pride to a scientific view, as "dogma."

    As for gene regulation, I've always been prepared to cut him a little slack on the basis that nothing was known of this when he first published.
    But I've just learned in the last few days that gene regulation was an established fact years before, so this lends weight to my claim that he was promoting an ideology. 
    First of all, thank you for replying so quickly considering this is an old thread. But, hey it's an interesting topic. I have one more question that would really clarify things for me. Last week in Science, a paper was published that showed how adaption to cold occurred via RNA editing not at the genomic level. In this paper, this RNA editing allowed cephalopods to live in cold waters and not compromise their nervous systems by changing how their K+ channels worked (put very simply for sake of time.) Can't we use this example of how selection must take place at the phenotypic level of the organism, because in this case, the genes weren't even changed? It was an adaption via RNA editing! No changes in the genetic information. THe phenotype was selected for not the genes. What do you think? Thanks again in advance for the conversation and thanks again for the article.

    Gerhard Adam
    Even if you take the position that RNA editing only exists to "fix" coding errors, it finishes the gene-centric view, because obviously a gene can't code for its own correction.

    Personally I have a different take on the whole process, but suffice it to say that while DNA is critical for retaining information between generations and ensuring a reasonably faithful replication, that's not the whole story.  It suggests that there are numerous other mechanisms that are sensitive to the environment and also capable of intervening to prevent expression of certain genes based on some [as yet, unknown] criteria.

    http://blogs.scientificamerican.com/observations/2011/12/25/green-glow-shows-rna-editing-in-real-time/
    Mundus vult decipi
    Steve Davis
     "...selection must take place at the phenotypic level of the organism,..." Natalie, this has always been obvious to those not seduced by the selfish gene fantasy.
    How gene selection got the traction it did is staggering.
    "It suggests that there are numerous other mechanisms that are sensitive to the environment and also capable of intervening to prevent expression of certain genes based on some [as yet, unknown] criteria." Exactly Gerhard. As you indicated, genes are important, but they are tools used by the cell. The gene-centrics can't get their heads around the fact that a cell is not a bag of individual genes. 

    Hi Steve,
    I've read your opinion with great interest, and also parts of the discussion. I've also see the phenotype more as the entity of selection than single genes, but I suspect that there can also be conflict of interests between genes within an organism, as there can be conflicts between members of an ant society about reproduction.
    I am interested how would you explain greenbeards, if you only consider the "group of genes" interacting with the environment as the source of selection.
    Greetings
    cvb

    for studies on green beards see:
    Andy Gardner and Stuart A. West 2010. "GREENBEARDS". Evolution, doi:10.1111/j.1558-5646.2009.00842.x
    Queller et al. 2003. Single-Gene Greenbeard Effects in the Social Amoeba Dictyostelium discoideum. Science
    Keller and Ross 1998. Selfishgenes:a green beard in the red fire ant. Nature

    Steve Davis
    Thanks for your interest.
    I see the green-beard effect as a tactic dreamed up by Hamilton in 1964 to give support to his ridiculous "altruism and cooperation are really selfishness" idea.
    At that time Hamilton was making statements about the role of genes that were false, bordering on delusional. He depicted genes as independent agents, and that independent agent aspect to his thinking can be seen in the green-beard idea.
    Like all predictions coming from the selfish gene hypothesis, green-beard predictions can be explained by other influences, so it's no more than speculation.
    Steve Davis - let me explain the peppered moth
    Imagine that there are two peppered moths. They both have exactly the same genotype (very unlikely I know), except that one has a gene which produces the dark pigmentation, the single gene effect (SGE) (as you don't seem to like the word phenotype) of the gene is either to produce a dark pigmentation or not to (as the case may be). If the moths live in an industrial area, the moth with the SGE providing dark pigmentation is more likely to survive and reproduce than the other moth. Now imagine a whole city full of moths. As natural selection is always acting, the gene in question will always be paired with a set of relatively successful genes. In nearly all cases the dark pigment gene will produce dark pigmentation, and individuals with a dark pigmentation SGE will do better on average than those without. The gene with the SGE producing dark pigmentation will therefore produce more copies of itself than its allele. It has the more successful SGE in a city setting and therefore out-competes its allele, even though the process by which this occurs is through the death of whole organisms

    Gerhard Adam
    I'm not sure you're understanding Steve's point.  All natural selection works through the death of the organism [or it's offspring].  If it didn't there wouldn't be anything selected.

    The gene simply produces a dark pigmentation or light pigmentation.  It is the environment that selects which of those pigments will be more successful for the organism.  This is precisely why there  have been more lightly pigmented peppered moths in recent years as the industrial pollution has been cleaned up.  So, the point is that the gene cannot behave in any manner except to produce the trait that it is set to produce.  Nothing more nothing less.  Whether such a trait is useful or not will depend on the environment [of which the gene is completely oblivious].

    This is precisely why any assertion that the gene is attempting to maximize its replication in the gene pool is completely foolishness, since the gene has no idea of what's happening "out there".
    Mundus vult decipi
    Of course, the genes are oblivious to everything, and the idea of competition between genes is just a metaphor.
    But don't you think that whole organisms (perhaps with the exception of humans) are also oblivious and merely follow their instincts. In which case, competition between organisms is simply a metaphor as well? After all, organisms don't know that they are merely a product of evolution, and they don't know that they're evolutionary purpose is to pass on their genes.

    So yes, the idea of rivalry between genes is not real, but it is just a way of looking at things in my view

    Gerhard Adam
    But it's a goofy metaphor.  It's like arguing that words compete with each other for their future presence in books.  It makes no sense.

    More importantly, this "metaphor" has given rise to discussions about how altruism and cooperation could arise from such "selfishness".  That's more than a metaphor.

    Dawkins has even gone so far as to suggest that we are simply robots constructed to do the bidding of our gene masters.  Again, this is more than a metaphor.

    The simple reality is that genes can't even control their own expression.  Between the influence of the environment and epigenetics, the gene isn't capable of competing at any level, even metaphorically.

    Also your point about organisms competing is a good point, because I don't see it as competition.  Organisms simply do what they do in their own self-interest.  That's not competition.  They aren't out do to better than others.  They simply are out to do for themselves.  Again, the terminology clouds the issue.

    Organisms don't need to know that they are the "product of evolution" any more than humans needed that knowledge before Darwin.  Even possessing that knowledge, it's not like humans are capable of directing their own evolution.  As before, they simply do what they do.  No metaphor is necessary.

    However, it also misses the larger picture, because organisms do specifically cooperate.  It's not simply some coincidence or accident of their behavior.  They specifically recognize their own "kind", even bacteria have quorum sensing, which allows them to distinguish between their own numbers and that of others.  Cells within an organism specifically cooperate to create the entire creature, even to the point of programmed cell death when they are superfluous.  They have given up their individual capability of reproducing in most cases, for the greater group.

    I don't accept the word "instincts" since it doesn't provide any meaning to the conversation.  It's generally used to describe some behavior that we don't know the causes of and with a wave of the hand, we use it to explain all manner of behaviors.  Whether these behaviors are genetic, behavioral, cultural, etc. is still largely unknown since whatever else we want to postulate, there aren't any organisms that behave as if they are programmed.  This suggests a fair amount of flexibility even within the context of their genetic background.

    In short, focusing on the genes, is simply wrong.  The genes cannot, by themselves, produce a viable multicelled organism.  They are dependent on too many factors, including other microorganisms to complete the work, and those other members of the microbiota do not share any genetic information.  So, the gene-centric view doesn't work to explain what we actually see.
    Mundus vult decipi
    Steve Davis
    Gerhard, you summed the matter up perfectly.
    The problem with gene-centrics is that they cannot stick to the facts as anonymous has, they have to go further and introduce competition between alleles, and rivalry, and other lies.
    It's all lies!