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The Extended Phenotype: The Gene
as the Unit of Selection

by Richard Dawkins

New York: W. H. Freeman, 1982

Exactly what, asked W. D. Hamilton, are the things that natural selection is supposed to select? The fittest what? Some biologists have thought in terms of the individual organism working for its inclusive fitness, while others have gone directly to the level of the gene, 'working' for its survival. Ideally there should be no difference between these two views, but this assumes that the phenotypic characteristics of an organism, to the extent that they are controlled by any genes, are controlled by the genes sitting inside that same organism. If genes in one animal could exert phenotypic effects on the body of another animal, we would have to revolutionize our views of adaptation.

Richard Dawkins shows that some such revolution is, indeed, logically necessary, since genes can be said to have extended phenotypes outside the bodies in which they sit. The argument is developed step by step, with full attention being given to possible objections and difficulties. In the course of this development, the issues tackled include the controversial ones of 'genetic determinism' and 'adaptionism,' the ideas of manipulation and 'arms races,' 'evolutionarily stable strategies,' genetic 'outlaws' and 'selfish DNA.' 'fitness.' animal artefacts and parasitic manipulations, the new concept of genetic 'action at a distance,' and the organism as a phenomenon needing explanation in its own right.

The book is designed for professional evolutionary biologists and their students, but it is written in a style that may appeal to many of the non-professional readers who enjoyed the author's previous book, The Selfish Gene.

Richard Dawkins was born in 1941. He was educated at Oxford University, where he did his doctorate under the Nobel Prize-winning ethologist Niko Tinbergen. From 1967 to 1969 he was an Assistant Professor of Zoology in the University of California at Berkeley, after which he returned to Oxford where he is now a Lecturer in Animal Behaviour. His first book, The Selfish Gene, was published in 1976. Since then he has been clarifying, developing, and modifying the position advocated in The Selfish Gene, with the result that The Extended Phenotype is in some respects a logical development of the earlier book; in other respects it represents a radical departure.

 
   
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