Selection does operate primarily on Genes : In defense of the Gene as the unit of selection

In Francisco José Ayala & Robert Arp (eds.), Contemporary Debates in Philosophy of Biology. Wiley-Blackwell Pub. 127--140 (2010)
Natural selection is an important force that shapes the evolution of all living things by determining which individuals contribute the most descendents to future generations. The biological unit upon which selection acts has been the subject of serious debate, with reasonable arguments made on behalf of populations, individuals, individual phenotypic characters and, finally, individual genes themselves. In this essay, I argue that the usual unit of selection is the gene. There are powerful logical arguments in favor of this conclusion, as well as many real-world examples. I also explore the possibility that epigenetic differences between individuals may be heritable between generations. Although few such examples exist, epigenetic differences provide an exciting source of potentially heritable variation that may allow rapid evolutionary change to occur, perhaps in response to environmental influences.
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William C. Wimsatt (1980). The Units of Selection and the Structure of the Multi-Level Genome. PSA: Proceedings of the Biennial Meeting of the Philosophy of Science Association 1980:122 - 183.
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