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This is a revised edition with a new afterword by Daniel Dennett. The Extended Phenotype carries on from where The Selfish Gene takes off. It is a fascinating look at the evolution of life and natural selection. Dawkins's theory is that individual organisms are replicators that have extended phenotypic effects on society and the world at large, thus our genes have the ability to manipulate other individuals. A worldwide bestseller, this book has become a classic in popular science writing.
Heavy-duty advanced neo-Darwinian adaptationist defence with the gene as the unit of selection
| Customer Rating:  | The first thing I will say about The Extended Phenotype (EP) is that it is far from the first book on evolution you should read and as a stand-alone Dawkins book it is a poor choice because it is a sequel of sorts to The Selfish Gene (TSG) and there is much more going on besides. I would suggest Climbing Mount Improbable or The Blind Watchmaker first. Both of those books by Dawkins have a much broader, more generalized, look at natural selection and evolution. TSG covers the basics of the gene view as the unit of selection. After you do this then I recommend that you read what has become known as "Darwin Wars" to some (it is not a book but a collection of writings). The two main critics are Stephen Jay Gould and Richard Lewontin. There are two articles you should read from them before EP. They are called Darwinian Fundamentalism by Stephen Jay Gould and The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme: by Stephen Jay Gould and Richard Lewontin. Both articles appear on the unofficial Stephen Jay Gould archive on-line. You can read about the Darwin Wars in general there also. Author Daniel Dennett has much to say on this also.
EP is a two-fold book. In one respect it is an apologetics in response to an anti-selfish gene (calling this version of neo-Darwinism, Darwinian fundamentalist or the adaptationist program) movement that was also associated with the anti-sociobiology criticisms of E.O Wilson's work. In other respects EP is a straightforward work on evolution that is geared towards professional biologists in an attempt to take matters raised in TSG and to develop them further to encompass how the gene is expressed in the phenotype from the view of the gene and not the traditional view of the individual. Like TSG it will be anti-group selectionist. It is also very much a book on adaptation from the neo-Darwinian perspective. This book will enjoy a dog ear phenotype for those who get it.
Chapter 1 - Necker Cubes and Buffaloes
Dawkins suggests that EP is not really saying anything different than what we understand about phenotypes today. He suggests however that the gene view of phenotypes as adaptations in evolution offers much more for the evolutionist.
Chapter 2 - Genetic Determinism and Gene Selectionism
This is largely an apologetic for those who misunderstood The Selfish Gene to mean that people are puppets of our genes (some foreign book covers with puppet-type illustrations may have implied this) and who didn't get that the robot examples were just analogies. Dawkins sets the record straight, genes are selected for, they give rise to phenotypes and the environment has an impact on the phenotype so it is not possible that everything is determined by the gene alone. Dawkins brings up embryology (developmental biology) to put this gene myth to bed. However he says that it is important to understand that without genes, the phenotype could never be expressed at all. More crucially, how the gene is expressed needs to be understood in a statistical sense. Dawkins even gives examples where talking about traits in terms of genes is misguided although in the end a biologist is forced to admit that without genes the phenotype cannot appear. Genes do determine the phenotype but there is a degree of variation there. The rest of the book talks about what that degree is and how to understand it.
3 - Constraints on Perfection
This is the correct version of the adaptationist program as taken from the standard model of neo-Darwinian adaptation. It shows that the Panglossian critic is not the neo-Darwinian view at all. The Panglossian sees all adaptations as being the best for that environment. The neo-Darwinian doesn't think this. Dawkins tells us that to see all adaptations as just roads to the perfect adaptation ignores physiology constraints such as biochemical restrictions. He gets some less likely suggestions out of the way like how neutral theory shouldn't apply because it's neutral on adaptations, how allometry is probably only a minor role in adaptation and how pleiotropy is influenced by developmental biology in separating out the better phenotype. So this leaves the way for what he sees as the more important considerations for adaptationists in the form of time lags (environment changes that the organism has yet to catch up with), historical constraints (how previous adaptations restrain future ones), available genetic variation (one variation that was sufficient may have taken off before a better variation arrived), constraints on costs and materials, imperfection levels (genes in populations can produce adaptations that fly in the face of Panglossian adaptation) and malevolence (the environment can act against the adaptation, statistically).
4 - Arms races and manipulation
Using various examples Dawkins convinces the reader that organisms can be in the business of directly manipulating other organisms to their own advantage and that evolution causes an arms race between manipulators and victims. The parts of slave making insects can get very heady but it is worth the effort and the hymenopteran sex ratios may be the most difficult topic Dawkins has ever covered in any of his books... but there are nearly equally as difficult topics that lie ahead.
5 - The Active Germ-line replicator
This chapter is devoted to undermining the individual or the group as the unit of selection and promoting the gene as the unit of selection. The gene is reduced, Dawkins introduces the concept of replicators and vehicles, describes each in terms of natural selection, make precise definitions of his replicators (active germ-line replicator) and takes on his critics. It is a genetics heavy chapter.
6 - Organisms, Groups and Memes: Replicators or Vehicles?
On the bases of the previous chapter Dawkins applies the concept of the replicator and vehicle to real world scenarios. This chapter gives one of the best explanations of Gould's and Elderage's Punctuated Equilibrium in print anywhere. It appears Dawkins subscribes to it in principle and then makes probably his first concession that the unit of selection now has to be expanded from the gene to include the species, but he asserts that this is not group selection and that species selection cannot be responsible for any sort of serious biological complexity and is therefore just an outcome of a trend. He then defends his position on the meme and tries to clarify more about replicators and vehicles.
7 - Selfish Wasp or Selfish Strategy
This chapter is about ethology research that Dawkins did on wasps to show why the gene view is important because it produces different results. It is a technical chapter where Dawkins also visits Maynard Smith's concept of the evolutionarily stability strategy (ESS) and does an analysis of the experiment from the view of the gene and then the view of the individual. He believes the gene view gives more accurate results.
8 - Outlaws and Modifiers
This is a challenging chapter that takes a look at natural selection in action within an individual between their own genes. Dawkins covers a number of phenomena within cells that cannot be explained without invoking a gene view. He then deals with the problem of relative-recognition and how the selfish genes are actually deciphered by organisms.
9 - Selfish DNA, Jumping Genes, and a Lamarckian Scare
This takes a look at `Junk' DNA and Dawkins gives his own explanation for it. He addresses the genes that depart from DNA strands in the cell and their fate or evolution. He then talks about the problem of evolution of sex and meiosis before presenting a research paper that was almost Lamarckian and helps the reader to understand Weismann's dogma.
10 - An Agony in Five Fits
Dawkins look at five different ways in which scientists have used the term `fitness' in relation to evolution. He goes through the pros and cons of each model.
11 - The Genetical Evolution of Animal Artefacts
200 pages in and Dawkins now addresses what is the central theme of this book, The Extended Phenotype or the long reach of the gene. Dawkins believes that he has established that behaviour is genetic and is subject to natural selection so there is no reason not to see the results of behaviour, such as the construction efforts of organisms, to be treated as phenotypes and therefore subject to natural selection. He uses Caddis houses, spider webs and beaver dams as examples. Then he introduced group construction and though a gene view using the example of termite mounds, explains that these artefacts are constructed by a combination of gene variation, roles in the colony and the use of cues and signalling. There is a very interesting bee swarm observation at the end.
12 - Host phenotypes of parasite genes
Here Dawkins looks at how parasites modify a host organism so that the host because more suitable for them even at the expense of the host. He brings up a number of examples mostly concerning flukes, snail shell size and snail antlers. He then suggests a new type of biologist who should look beyond just the phenotypes that appear on the organism to include other organisms. He proposes that two biologists studying separate types of organisms that interact could be incorporated in one `extended view'. He calls this type of biologist the `extended geneticist'.
13 - Action at a distance
This gives some more examples of the extended phenotype in action with more snails and the Bruce effect with mice. Dawkins states the formula of the extended phenotype, `An animal's behaviour tends to maximize the survival of the genes 'for' that behaviour, whether or not those genes happen to be in the body of the particular animal performing it'. He questions over what distance this can extend to but expects that its effects diminish with longer distances. He sees no reason why it can't be across continents. He talks about Lovelock's `Gaia' hypothesis and disagrees with this saying that there is no mechanism that supports a feedback system of that type. Dawkins believes the selfish gene view is the only way to view biota on this planet. The chapter closes by Dawkins quoting important evolutionists who appear to critic the gene view and answering or commenting on their statements.
13 - Rediscovering the organism
This chapter takes a look at the individual in a new that that examines the roles of growth and reproduction and how development cycles or stages of growth has advanced and sped up evolution. He finishes by recapitulating everything. You get a glossary of evolution terms at the end.
EP does suffer slightly from being apologetic even if Dawkins does try to overcome it. EP does have those parts in its shadows but to be honest I quite like its apologetic nature. However it takes a fair few chapters before it gets into the gene as the unit of selection (around 4 or 5) and the extended phenotype doesn't really emerge until chapter 11. It is worth the wait or battling through some of the most complicated examples for a laymen's book on biology, but Dawkins states that this is also an advanced evolution book for professionals. Essentially if you are brave enough to read it, then go for it but don't be surprised if you can't follow it. I will admit that even though I am a biology undergraduate I didn't understand a lot of it and it demands several readings.
The argument for the extended phenotype strikes the reader as something so obvious that it is surprising that Dawkins thinks he is establishing something new. Biology doesn't rule out what Dawkins is presenting and in fact states as much, but the language of the extended phenotype is rarely emergent in other works on biology. This is probably due to biologists being inclined to use the individual as the unit of selection, rather than the gene and also apprehension that using the principle of the extended phenotype could be wrong. Dawkins sets the record straight. Talk in terms of genes and the extended phenotype is a fact.
One weakness that does emerge is the criticism that even blind natural selection cannot see genes, only the individual, but Dawkins rightly brushes this off as merely a stiffness of thought and that the genes are still being selected but the criticism goes further to say that genes are selected in groups, not as discreet units. Dawkins has to make the concession that it is genes coexisting (or for their capacity to cooperate with other genes) that are the unit of selection. Is this a form of group selection after all? Dawkins implies that this is a different sense completely and that it is really only the germ-line genes that are being selected. It is frustrating because the reader can see that he is right to forward this gene view only to end up with a fact of evolution that genes cooperate and coexist so that they are potential candidates for a selection process that chooses them in groups. Can you imagine them as discreet units? Yes of course, and this is exactly what Dawkins wants us to see, but nature picks them out in aggregate and not discreetly. Whatever way Dawkins tries to cut the cake of phenotypes it looks like he has to eat a slice of group selection to have a filling of gene selection. Look at the problem the gene view has with selecting deleterious genes, like in a heterozygous state. The only way it seems to overcome this problem is to allow grouped genes as the unit of selection otherwise we end up saying that natural selection is selecting for genes that are disadvantageous to the organism. However here comes the genius, Dawkins rightly suspects that this not the case. Natural selection is not selecting disadvantageous genes, but meiosis in terms of reproduction is reproducing advantageous genes but recombines them in a way that makes them disadvantageous. It seems this view saves the gene view except for one small glitch. Meiosis itself cannot escape its own genes. Dawkins however does bring up one more salient point. Embryology or the development of the biology of the genetic influence is subject to change. Since development is so variable how can it be selected? It isn't. It's the genes that are ultimately selected not developmental biology. Dawkins saves the gene view in light of his critics confusing genetics with embryology. That slice of group selection now appears to be composed entirely of selfish genes. Maybe the cake doesn't taste so bad in light of this.
Dawkins asserts that this is his most important book. I would agree, it ranks as his most complex work. Not to mention that his critics haven't been able to answer it since 1983! and just rehash the same arguments Dawkins addresses in this book. I placed The Selfish Gene as the most important works on evolution outside of Darwin's and R.A Fisher's. I am inclined to push The Extended Phenotype into a slot above it. This is more powerful than all of Dawkins' works combined... and a host of other books on evolution to boot. Dawkins works always resonate in my mind in ways that change my worldview but the extended phenotype is the nuclear detonation of conscious expanding scientific experience on a magnitude that intensifies with every new insight it discloses. |
Hard core biology
| Customer Rating: | It's a tough read at times, since it is aimed at Dawkins' colleagues and biology students. But it gives you a sense that the mechanisms of inheritance and natural selection are getting to be very well understood.
Oddly, the alleged subject of the book is to be found tucked away in the last chapters, the first 10 or so being Dawkins explaining stuff and telling how others are mistaken. |
Theory, without the distractions.
| Customer Rating:  | TEP is primarily a technical treatise. It elaborates the implications of holding a Darwinian explanation for the diversity of life forms across and, most especially, through time. In detail, the reader is alerted to what is conceptually required for a Darwinian theory to be internally consistent, and for it then to be applicable to life as it is and as it has been.
Professor Dawkins persists in using the misleadingly emotive terminology from his previous best seller, The Selfish Gene. However, in TEP it is very much tempered and qualified. The picture he wishes to make vivid is from a vantage entirely remote from human concerns - it is at the level of molecules, and the complex systems which require copies of molecules to be made . From this perspective, to `survive' means to produce a copy, and to be `selfish' is likewise to be inclined towards such replication; the `struggle' to survive, by this account, is simply a tallying of the number of reproductions both across time, and through the generations. It is made clear that the `fighting' takes place between lineages, and hence over a much extended time frame, that is, `in evolutionary time'.
As the participants in this most undramatic of dramas are utterly bereft of any human attributes, the terms are spoken `innocently', that is without any suggestion that any extrapolations can be made from what goes on at the level of goo to the level of human interaction. TSG also claimed to speak innocently, but it protested its innocence in the face of a gale of rhetoric to the contrary - in TEP, the wind has, thankfully, settled.
TEP spends time undermining certain tenets previously central to Darwinian thinking. The notion that the individual organism works for its own reproductive ends is challenged in chapter four, `Arms races and manipulation'. Prioritizing the germ-line over the individual organism is a recurrent theme, perhaps made most stark in chapter six, `Organisms, Groups, and Memes: Replicators or Vehicles?'; here the central issues turn on Darwinian theory requiring a means by which variety is generated in the factors to be `selected', and a means by which these factors are replicated in succeeding generations - dealing with organisms and groups in turn, Dawkins explains troubles with their candidature. In regard groups, while he is not utterly inimicable to selection occurring at this level, he gives good reason to be skeptical that it could account for the development of complex organs and specializations. Oddly, with his own theory of memes he gives a paragraph of objections and these seem far more convincing than the theory itself.
The penultimate chapters expand on the concept of the title. By extending the phenotype, Professor Dawkins is again undermining the notion that the individual organism is the `level of resolution' which we should examine. The level at which forces of selection operate may better be conceived of as organisms plus their direct environmental effects; or, in virtue on focusing on the genetic germ-line, in might be better to focus on an extended lineage of related individuals. Where the limit is drawn is discussed, albeit vaguely, in terms of where there is discernable feedback to the reproductive success of the germ-line in question.
Finally, the author `rediscovers the organism'. The last chapter is, however, more of an invitation to question why life is organized at the level of individual organisms than a celebration of that fact. To ask such a question, once again the germ-line, or the level of the genetic replicator, is made basic.
In sum, TEP asks us to view the panoply of life from the perspective of replicating DNA, and it suggests that such a view helps to order and explain the events of myriad complexity and diversity which otherwise appear wonderful but beyond explanation. The story it tells is full of detail and complextiy itself, and it occurs at a scale of magnitude and a time frame and, ultimately, from a perspective which makes no comment on our actual lives. Remove the vestigial emotive words carried over from TSG, and the TEP stands clearly as an exposition of a technical scientific theory, and a good one at that. |
A scholastic argument
| Customer Rating: | | I read this book because Dawkins in other fora has identified it as his best work. He says it's a version of "The Selfish Gene" for biologists, but as Daniel Dennett points out in the afterword, it is actually an extended philosophical argument. More precisely, it's a scholastic disquisition on why it's only the individual gene that may be denominated as the "unit of selection" for purposes of natural selection as opposed to the organism or some other model. Here's an example of why this is more a medieval "angels on the head of a pin" argument than anything else. Say you are born with a gene mutation that gives you better running ability. Let's say, though, that because your judgment doesn't match your speed, your better running leads you not only to better escape predators, but also to run over the side of a cliff. In that case, is the "running" gene being selected for, or against? The answer really is, neither. Your genome as a whole is being selected against. If you survive long enough to have children and have passed on your mutation, the running gene may have another chance, assuming it's not paired up for life with the "judgment" gene. If you haven't survived long enough, then tough luck. Insisting on the gene as the only possible "unit of selection" is just a semantic argument and doesn't really get you anywhere in terms of understanding the real world. This book is a hard slog, in part because so many paragraphs are digressions to address various biologists who disagree with Dawkins. There were a couple of points that are eye-openers (why are cicada cycle years prime numbers?), but I'm not sure those alone make it worth the effort... |
Why the ridiculously small type?
| Customer Rating: | I was impressed with Richard Dawkins' "The Selfish Gene" and this book is the logical followup. It is more complicated and therefore more difficult to comprehend than the former. So that makes the very small type all the more annoying -- it adds to the difficulty. Seriously, the type size is among the smallest I have seen in any modern mass produced book.
Anyone know if any of the earlier editions have larger type? |
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