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• W2048333710 abstract "What is the ‘levels of selection’ question about? It's about the level of the biological hierarchy at which natural selection acts, e.g. individual, group, gene, community, species, etc. Usually when we think about selection, we think of it acting at the level of the individual organism, favouring the fittest individuals over the less fit and thus leading to evolutionary change. But individual-level selection is only one possibility among many. For the key requirements of evolution by natural selection — variation, associated differences in fitness and heritability — can in principle be met by entities at many levels, above and below that of the individual organism. For example, a selective process could quite easily operate on groups of organisms, favouring some types of groups over others. This idea is known as ‘group selection’.What is the origin of the levels of selection debate? Like much in evolutionary biology, it traces back to Darwin. Though Darwin primarily discussed individual-level selection, he was aware of other possibilities. In The Descent of Man (1879), he tackled the problem of how self-sacrificial and other ‘altruistic’ behaviours could have evolved in early hominids. As such behaviours reduce an individual's fitness, it is clear that they cannot have evolved by selection at the individual level. Darwin suggested that group selection may be the answer. Groups in which altruistic behaviour was prevalent may have enjoyed a selective advantage over groups in which it was absent, he argued. Another early evolutionist who discussed the levels question was August Weismann, mainly in relation to selection at the sub-organismic level. However, the modern debate only really took off in the 1960s with the rise of social evolution theory, and the ensuing controversy over group selection.Why did ‘group selection’ become such a chequered concept in the 20th century? Primarily due to George C. Williams' trenchant critique; also influential was the work of John Maynard Smith. They argued that group selection was theoretically possible but unlikely to be a major evolutionary force, and was not needed to explain the known biological phenomena anyway. Also, they stressed the fallacy of assuming that selection on individual organisms would automatically lead to outcomes that benefit the group. This fallacy was surprisingly common in mid-20th century biology and is still encountered today. The rise of ‘kin selection’ also contributed to the demise of group selection. Biologists such as Richard Dawkins and John Maynard Smith argued that kin selection or inclusive fitness theory, first articulated by William D. Hamilton in the 1960s, provided a better explanation of phenomena such as altruism which had traditionally been taken as evidence for group selection. However, the true relation between kin and group selection is a controversial matter. Many modern theorists argue that suitably understood they are in fact equivalent, so do not constitute alternative scientific hypotheses at all.But isn't the gene the real unit of selection? Dawkins originally presented his ‘selfish gene’ idea as an empirical alternative to individual and group-level selection, but later realised the error of his ways. Almost any selection process, at any level, will ultimately lead to a change in gene frequencies, so can be described as a process in which one gene spreads at the expense of its alleles. Thus, it is generally wrong to contrast ‘gene selection’ with either individual or group selection — there is no empirical issue at stake here. This was the point that Dawkins later captured with his ‘replicator–vehicle’ distinction; others have marked it by contrasting units and levels of selection. In recent literature, ‘gene-level selection’ is often used in a restricted sense, to mean selection between genes within a single organism, as occurs in cases of intra-genomic conflict, e.g. meiotic drive. In this sense, most selection processes cannot be described as gene selection.How does the levels of selection question relate to the ‘major evolutionary transitions’? Such transitions, as defined by Eors Szathmary and John Maynard Smith, occur when free-living biological units, capable of surviving and reproducing alone, coalesce into a single larger unit, giving rise to a new, higher-level individual — for example, in the evolution of multi-celled organisms from single-celled ancestors. In transitions of this sort, there is the potential for selection to act on both the smaller and the larger units — the individuals and the groups. For the groups to evolve into ‘real’ individuals, i.e. integrated cohesive units, group-level selection must trump individual-level selection. So multiple levels of selection are intimately involved with evolutionary transitions.What about species selection? This was an idea defended by Stephen J. Gould and other macroevolutionists, which says that selection may operate on whole species over geological time, favouring those species best able to survive or reproduce (i.e. speciate). This could explain why certain types of species, e.g. ecological specialists, become more common than others, e.g. generalists, in a given clade, and thus indirectly help explain long-term evolutionary trends. Species selection is certainly a logical possibility, but it is difficult to assess how important an evolutionary process it has been." @default.
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• W2048333710 title "Levels of selection" @default.
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• W2048333710 doi "https://doi.org/10.1016/j.cub.2010.01.025" @default.
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