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Memes are items that evolve in an evolutionary process covering the social and cultural domain only. They are sociocultural analogues to genes or viruses.

The idea of a cultural item that evolves the same way as biological entities do goes back at least to Herbert Spencer. The notion that culture is passed on like a disease also goes back to the same period. But both metaphors have usually been treated as just that—suggestive similes of no serious import. The first person to take the evolutionary idea seriously was zoologist Richard Semon, a student of Ernst Haeckel, in his 1904 book Die Mneme (published in English as The Mnemein 1924). Semon's mneme idea was not limited to culture; it also included any impression left on an organism by external factors (Engramm), that is, acquired inheritance.

The differentiation of cultural evolution from biological evolution appeared in 1965 in the work of Donald T. Campbell, who influenced Karl Popper to develop an evolutionary epistemology in 1972, and in various other authors, including Julian Huxley in 1957 and Stephen Toulmin in1972. In 1981, C. J. Lumsden and E. O. Wilson and proposed the notion of a “culturgen” (culture–gene) as the target of evolution in culture. A number of other “dual inheritance” models of culture have been proposed. But the modern fascination with the idea of a meme derives mainly from the writings of the theorist who coined the term, Richard Dawkins.

Dawkins developed a generalized theory of evolutionary processes independent of the physical substrate in which they were realized and termed it universal Darwinism. The relevant entities of all evolutionary processes included what Dawkins called a replicator, an entity that copies itself according to three criteria,namely that it is fecund, it is high fidelity, and it is long-lived. “Longevity, fecundity, and fidelity” is the slogan of this new revolution. Dawkins's view of evolution has come to be known as the gene's eye view.

Replicators in biology are DNA based, Dawkins said. But in cultural evolution, they are something else—memes,a term coined to be similar to gene, with which it was grouped as a replicator. Dawkins used two analogies for memes. One was that memes, like genes, are the things that evolve, setting up a problem as to the contents of memes. The other was that memes are mobile hereditary elements like viruses, taking over the resources of a host brain the same way as a virus subverts the cellular machinery of its hosts to replicate.

Advocates of the memes-as-germs approach, as one might call it, tend to focus on the epidemiological aspect of culture. In this regard, this approach is closely allied to the anthropological conception of diffusion. Advocates of the memes-as-genes approach, on the contrary, focus on the selectionist aspects of memes, their fitness, and so forth. There are strong advocates and critics of both approaches.

Because the notion of a meme is tied into the universal Darwinism of Dawkins, and the consequent ontology of phenotype analogues (vehicles in Dawkins's terms, interactors in Hull's terms), two questions that have engendered considerable debate arise. First, what is a meme? Second, what is the phenotype of a meme? Answers have included that memes are neural structures or activated neural constellations and that they are artifacts, behaviors, or the subject of selection pressures in cultural transmission. Examples have included dress practices, snatches of tunes, phrases, pottery styles, and architectural styles. But it remains unclear whether these are the things that replicate or the things that express what is replicated. Some hold that memes are the physical things that are recreated, the practices, or the artifacts. Others hold them to be the ideas or symbolic representations that are passed on and that cause the external behaviors and products of behavior. In short, in the former view, memes are concrete things that get reproduced; in the latter view, memes are abstract things that result in concrete phenotypic behaviors and products. A “multilevel selection” approach, as may be found in modern biological accounts, applied to memes suggests that memes, as selective targets, may be replicators at any level of the hierarchy of the evolutionary domain, and so memes may be both within heads and between them.

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