“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) 
There's a valuable discussion of these issues in the comments that follow the article here.