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Fig. 1. Complex traits, such as behavior, are composed of numerous lower-level
(subordinate) traits, themselves interrelated in a strongly hierarchical
fashion. In general, natural and sexual selection will tend to act more
strongly at higher levels of biological organization, as indicated by the
relative thickness of the black arrows. As typically viewed by organismal and
evolutionary biologists, selection acts on phenotypic variation (which
reflects variation in gene expression), but does not generally act directly on
genetic variation (e.g. at the level of DNA sequences). Exceptions to this
point can occur via such phenomena as genomic conflict (e.g.
Stearns and Hoekstra,
2005).