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The Selfish Gene is a controversial book by Richard Dawkins published in 1976. The phrase "selfish gene" in the title of the book was coined by Dawkins as a provocative way of expressing the gene-centered view of evolution, which holds that evolution can be viewed as acting on genes, and that selection on organisms or populations almost never overrides selection on genes. More precisely, an organism is expected to evolve to maximise its inclusive fitness—the number of copies of its genes passed on globally (rather than by a particular individual). In addition, organisms will tend to evolve towards an evolutionarily stable strategy. The book also introduces the concept of memes, a unit of human cultural evolution analogous to the gene, arguing that replication also happens in culture, albeit in a different sense. Memetics have become the subject of many studies since the publication of the book. Describing genes with the term "selfish" is not meant to imply that they have actual motives or will—only that their effects can be described as if they do. This makes the concept somewhat problematic, as is critically summarised by Andrew Brown: "'Selfish,' when applied to genes, doesn't mean 'selfish' at all. It means, instead, an extremely important quality for which there is no good word in the English language: 'the quality of being copied by a Darwinian selection process.' This is a complicated mouthful. There ought to be a better, shorter word -- but 'selfish' isn't it. 1 A crude analogy can be found in the old joke "A chicken is just an egg's way of making more eggs." Likewise, Dawkins describes biological organisms as "vehicles" used by their genes for making more copies of those genes, regardless of the effect they might have on individuals or species. Genes that help the organism they are in to survive and reproduce also improve their own chances of being passed on; so most of the time "successful" genes will also be beneficial to the organism. But there are exceptions: segregation distortion genes, for example, that are detrimental to their host nonetheless propagate themselves at its expense. Likewise, the existence of junk DNA that provides no benefit to its host, once a puzzle, can be more easily explained. These examples might suggest that there is a power-struggle between genes and their host. In fact, the claim is that the interests of the organism are redundant, from an evolutionary point of view. (This is where the related concept of the extended phenotype comes into play.) When looked at from the point of view of gene-selection, many biological phenomena that in prior models were difficult to explain become easier to understand. In particular, phenomena such as kin selection and eusociality, where organisms act altruistically, against their individual interests (in the sense of health, safety or personal reproduction) to help related organisms reproduce, can be explained as genes helping copies of themselves in other bodies to replicate. In other words, genes act "selfishly" to increase the number of copies of themselves and for no other reason. This is in contrast to a version of group selection, which was common in evolutionary genetics prior to the 1960s. Proponents argue that the central point of the idea, that the gene is the unit of selection, is a more complete and accurate explanation of evolution than Darwin's. (Darwin, however, cannot be faulted for this absence because the basic mechanisms of genetics weren't understood at the time). Critics argue that this view oversimplifies the relationship between genes and the organism. The majority of modern evolutionary biologists accept that the idea is consistent with many processes in evolution. However, the view that selection on other levels such as organisms and populations almost never opposes selection on genes is increasingly viewed as extreme. In the last decade, difficulties with the theory of multi-level selection have been overcome and interaction between genes and between organisms as a force in evolution has again become a topic of research. Another criticism of the book, made by philosophers such as Mary Midgley in her book Evolution as a Religion, is that it discusses philosophical and moral questions that go beyond the biological arguments that Dawkins makes. The idea is sometimes mistakenly believed to support genetic determinism. This is incorrect: knowing that an organism carries a particular allele, we might be able to say that the organism is more likely than otherwise to behave in a certain way, but its actual behavior will depend on its environment and its developmental history. In particular, this applies to human organisms; Dawkins is quick to point out that although we may be influenced by our genes, we are not controlled by them. Even further from Dawkins' concept is the misunderstanding of the idea as predicting (or even prescribing or justifying) that human behaviour must inevitably be "selfish" in a moral or ethical sense. Bibliography
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