The Selfish Gene
Original cover from the painting The Expectant Valley by zoologist Desmond Morris
Author	Richard Dawkins
Subject(s)	Evolutionary biology
Publisher	Oxford University Press
Publication date	1976
Pages	224
ISBN	ISBN 019857519X
Followed by	The Extended Phenotype

The Selfish Gene is a book on evolution by Richard Dawkins, published in 1976. It builds upon the principal theory of George C. Williams's first book Adaptation and Natural Selection. Dawkins coined the term selfish gene as a way of expressing the gene-centered view of evolution, which holds that evolution is best viewed as acting on genes, and that selection at the level of organisms or populations almost never overrides selection based on genes. An organism is expected to evolve to maximize its inclusive fitness – the number of copies of its genes passed on globally (rather than by a particular individual). As a result, populations will tend towards an evolutionarily stable strategy. The book also coins the term meme for a unit of human cultural evolution analogous to the gene, suggesting that such "selfish" replication may also model human culture, in a different sense. Memetics has become the subject of many studies since the publication of the book. Image File history File links The_Selfish_Gene3. ... Dr Desmond Morris (born 24 January 1928 in the village of Purton, UK) is most famous for his work as a zoologist and ethologist. ... Clinton Richard Dawkins, FRS (born March 26, 1941) is a British ethologist, evolutionary biologist and popular science writer who holds the Charles Simonyi Chair for the Public Understanding of Science at the University of Oxford. ... This article or section does not cite any references or sources. ... A publisher is a person or entity which engages in the act of publishing. ... Oxford University Press (OUP) is a highly-respected publishing house and a department of the University of Oxford in England. ... See also: 1975 in literature, other events of 1976, 1977 in literature, list of years in literature. ... ISBN redirects here. ... A cathedral termite mount – a small animal with a very noticeable extended phenotype The Extended Phenotype: The Long Reach of the Gene is a 1982 book by British ethologist Richard Dawkins. ... This article is about evolution in biology. ... Clinton Richard Dawkins, FRS (born March 26, 1941) is a British ethologist, evolutionary biologist and popular science writer who holds the Charles Simonyi Chair for the Public Understanding of Science at the University of Oxford. ... See also: 1975 in literature, other events of 1976, 1977 in literature, list of years in literature. ... George Williams Professor George Christopher Williams (b. ... Cover of the 1996 edition. ... The gene-centered view of evolution, gene selection theory or selfish gene theory holds that natural selection acts through differential survival of competing genes, increasing the frequency of those alleles whose phenotypic effects successfully promote their own propagation. ... For other uses, see Gene (disambiguation). ... Inclusive fitness encompasses conventional Darwinian fitness with the addition of behaviors that contribute to an organism’s individual fitness through altruism. ... In game theory, an evolutionarily stable strategy (or ESS; also evolutionary stable strategy) is a strategy which if adopted by a population cannot be invaded by any competing alternative strategy. ... For other uses, see Meme (disambiguation). ... Cultural evolution is the structural change of a society and its values over time. ... Memetics is an approach to evolutionary models of information transfer based on the concept of the meme. ...

"Selfish" genes

Describing genes with the term "selfish" is not meant to imply that they have actual motives or will – only that their effects can be accurately described as if they do. The contention is that the genes that get passed on are the ones whose consequences serve their own implicit interests, not necessarily those of the organism, much less any larger level. Some people find this metaphor entirely clear, while others find it confusing, misleading or simply redundant to ascribe mental attributes to something that is mindless. For example, Andrew Brown has written:

"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]

Crude analogy

A crude analogy can be found in the old saying about a chicken being just an egg's way of making more eggs. In a similar inversion, Dawkins describes biological organisms as "vehicles" or survival machines, with genes as the "replicators" that create these organisms as a means of acquiring resources and copying themselves. From an organism-centric perspective, genes can be thought of as a blueprint for some feature that might benefit the organism; but from a gene-centric perspective, the sole implicit purpose is to benefit themselves. A related concept here is outlined in Dawkins' later work, The Extended Phenotype, in which the consequences of the genes to the environment outside the organism are considered. Survival machine is a concept, first proposed by Richard Dawkins in The Selfish Gene, whereby all organisms should be considered as the protection genes construct to survive their surroundings in order to propagate. ... DNA replication. ... A cathedral termite mount – a small animal with a very noticeable extended phenotype The Extended Phenotype: The Long Reach of the Gene is a 1982 book by British ethologist Richard Dawkins. ...

Genes and selection

Dawkins proposes that genes that help the organism in which they happen to be to survive and reproduce tend to also improve their own chances of being passed on, so – most of the time – "successful" genes will also be beneficial to the organism. An example of this might be a gene that protects the organism against a disease, which helps the gene spread and also helps the organism.

Genes can reproduce at the expense of the organism

There are other times when the implicit interests of the vehicle and replicator are in conflict, such as the genes behind certain male spiders' instinctive mating behaviour, which increase the organism's inclusive fitness by allowing it to reproduce, but shorten its life by exposing it to the risk of being eaten by the cannibalistic female. Another good example is the existence of segregation distortion genes that are detrimental to their host but 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. A more controversial example is aging, in which an old organism's death makes room for its offspring, benefiting its genes at the cost of the organism. The selfish gene theory postulates that natural selection will increase the frequency of those genes whose phenotypic effects ensure their successful replication. ... In molecular biology, junk DNA is a collective label for the portions of the DNA sequence of a chromosome or a genome for which no function has yet been identified. ...

Power struggles are rare

These examples might suggest that there is a power-struggle between genes and their host. In fact, the claim is that there isn't much of a struggle because the genes usually win without a fight. Only if the organism becomes intelligent enough to understand its own interests, as distinct from those of its genes, can there be true conflict. An example of this would be a person deciding to use contraception, even though their genes lose out due to this decision. Birth control is the practice of preventing or reducing the probability of pregnancy without abstaining from sexual intercourse; the term is also sometimes used to include abortion, the ending of an unwanted pregnancy, or abstinence. ...

Many phenomena explained

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 gene sets "helping" copies of themselves in other bodies to replicate. Interestingly, the "selfish" actions of genes lead to unselfish actions by organisms. In evolutionary biology, kin selection refers to changes in gene frequency across generations that are driven at least in part by interactions between related individuals, and this forms much of the conceptual basis of the theory of social evolution. ... Meat Eater ant colony swarming Fire ants Eusociality is the phenomenon of reproductive specialization found in some animals. ... For the ethical doctrine, see Altruism (ethics). ...

Prior to the 1960s, it was common for such behaviour to be explained in terms of group selection, where the benefits to the organism or even population were supposed to account for the popularity of the genes responsible for the tendency towards that behaviour. This was shown not to be an evolutionarily stable strategy, in that it would only take a single individual with a tendency towards more selfish behaviour to undermine a population otherwise filled only with the gene for altruism towards non-kin. In evolutionary biology, group selection refers to the idea that alleles can become fixed or spread in a population because of the benefits they bestow on groups, regardless of the fitness of individuals within that group. ...

Acclaim and criticism

The book was extremely popular when first published, and continues to be widely read. It has sold over a million copies, and been translated into more than 25 languages.^[2] Proponents argue that the central point, that the gene is the unit of selection, usefully completes and extends the explanation of evolution given by Charles Darwin before the basic mechanisms of genetics were understood. Critics argue that it oversimplifies the relationship between genes and the organism. For other people of the same surname, and places and things named after Charles Darwin, see Darwin. ...

Other types of selection suggested

Most 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, seldom opposes selection on genes is more controversial. While naive versions of group selectionism have been disproved, more sophisticated formulations make accurate predictions in some cases while positing selection at higher levels. Nevertheless, the explanatory gains of using sophisticated formulations of group selectionism as opposed to Dawkins's gene-centred selectionism are still under dispute. This article or section does not cite any references or sources. ...

Unit of selection or of evolution

Some biologists have criticised the idea for describing the gene as the unit of selection, but suggest describing the gene as the unit of evolution, on the grounds that selection is a "here and now" event of reproduction and survival, while evolution is the long-term trend of shifting allele frequencies.^[3] An allele (pronounced , ) (from the Greek αλληλος, meaning each other) is one member of a pair or series of different forms of a gene. ...

Moral arguments

Another criticism of the book, made by the philosopher 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. For instance, humanity finally gaining power over the "selfish replicators" is a major theme at the end of the book. Dawkins has pointed out that he is only describing how things are under evolution, not endorsing them as morally good.^[4] For other uses, see Philosophy (disambiguation). ... Mary Midgley, née Scrutton, (b. ... Morality (from the Latin manner, character, proper behavior) has three principal meanings. ...

Editions

The Selfish Gene was first published in 1976 in eleven chapters with a preface by the author and a foreword by Robert Trivers. A second edition was published in 1989. This edition added two extra chapters, and substantial endnotes to the preceding chapters, reflecting new findings and thoughts. It also added a second preface by the author, but the original foreword by Trivers was dropped. In 2006, a 30th anniversary edition was published which reinstated the Trivers foreword and contained a new introduction by the author (alongside the previous two prefaces), and also some selected extracts from reviews at the back. Robert L. Trivers, (born 19 February 1943) is an American evolutionary biologist and sociobiologist, most noted for proposing the theories of reciprocal altruism (1971), parental investment (1972), and parent-offspring conflict (1974). ...

30th anniversary celebrations (2006)

For the 30th anniversary of the publication of The Selfish Gene, a festschrift was published entitled Richard Dawkins: How a Scientist Changed the Way We Think. (An anniversary edition of The Selfish Gene was also published as mentioned above.) In March 2006, a special event entitled The Selfish Gene: Thirty Years On was held at the London School of Economics. The event was organised by Helena Cronin, and chaired by Melvyn Bragg. The programme was as follows: In academia, a Festschrift (; plural, Festschriften, ) is a book honouring a respected academic. ... Mascot: Beaver Affiliations: University of London Russell Group EUA ACU CEMS APSIA Universities UK U8 Golden Triangle G5 Group Website: http://www. ... Dr Helena Cronin is a noted Darwinian philosopher and rationalist. ... Melvyn Bragg, Baron Bragg, FRSL, FRTS (born 6 October 1939, in Wigton, Cumberland) is a British author and broadcaster. ...

The Selfish Gene: Thirty Years On (1:22:55) 00:00 – Melvyn Bragg – Introduction 02:20 – Daniel Dennett – The view from Dawkins' mountain 18:35 – John Krebs – From intellectual plumbing to arms races 31:55 – Matt Ridley – Selfish DNA and the junk in the genome 46:15 – Ian McEwan – Science writing: Towards a literary tradition? 1:00:00 – Richard Dawkins – Afterword

Melvyn Bragg, Baron Bragg, FRSL, FRTS (born 6 October 1939, in Wigton, Cumberland) is a British author and broadcaster. ... Daniel Clement Dennett (born March 28, 1942 in Boston, Massachusetts) is a prominent American philosopher whose research centers on philosophy of mind, philosophy of science and philosophy of biology, particularly as those fields relate to evolutionary biology and cognitive science. ... Sir John Richard Krebs (born 1945) is a British biologist and a Fellow of the Royal Society. ... Matthew (Matt) Ridley (born February 7, 1958 at Newcastle upon Tyne) (not to be confused with Mark Ridley) is an English science writer. ... Ian McEwan CBE (born June 21, 1948) is a British novelist. ... Clinton Richard Dawkins, FRS (born March 26, 1941) is a British ethologist, evolutionary biologist and popular science writer who holds the Charles Simonyi Chair for the Public Understanding of Science at the University of Oxford. ... Image File history File links Selfish_Gene_3. ... Image File history File links How_a_Scientist_Changed_the_Way_We_Think. ...

References

External links

• Video introduction by Richard Dawkins
• The Selfish Gene: Thirty Years On and mp3
• A BBC World Service interview with Richard Dawkins about "The Selfish Gene". Accessed 31 July 2007.

v • d • e Richard Dawkins
Books	The Selfish Gene (1976) · The Extended Phenotype (1982) · The Blind Watchmaker (1986) · River out of Eden (1995) · Climbing Mount Improbable (1996) · Unweaving the Rainbow (1998) · A Devil's Chaplain (2003) · The Ancestor's Tale (2004) · The God Delusion (2006)
Related works	Growing Up in the Universe (1991) · Break the Science Barrier (1996) · The Atheism Tapes (2004) · Beyond Belief (2006) · Richard Dawkins: How a Scientist Changed the Way We Think (2006) · The Root of All Evil? (2006) · The Enemies of Reason (2007) · The Oxford Book of Modern Science Writing (2008)
See also	Meme · Gerin oil · Foundation for Reason and Science · Ultimate Boeing 747 gambit · Lalla Ward