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Overall, our results highlight the fact that it is impossible to fully understand the evolution of a cellular module in isolation from the genome where it is encoded and the population-level processes that govern evolution. The ability of natural selection to improve any one module depends on the population size, the rate of recombination, the supply, and the fitness effects of all beneficial mutations in the genome and on how these quantities change as populations adapt. Further theoretical work and empirical measurements integrated across multiple levels of biological organization are required for us to understand adaptive evolution of modular biological systems. [Emphasis added.]
Comment: See also: Choosy eggs may pick sperm for their genes, defying Mendel's law