Genomic imprinting and methylation: epigenetic canalization and conflict

Imprinted genes have patterns of expression that depend on the parent of origin of their alleles. Establishment of imprinting at a locus requires that the two alleles be differentially marked in oogenesis and spermatogenesis, that these marks escape reprogramming after fertilization, and that they are reliably transmitted through development. Recent work on the mammalian DNA methyltransferases involved in these processes suggests mechanisms of epigenetic canalization, which might contribute to the stability of epigenetic inheritance. At the same time, the interactions that determine whether a particular modification will be transmitted or reprogrammed are destabilized by evolutionary conflicts, as the genes and gene products controlling these processes are subject to divergent selective forces. This review summarizes many of the recent advances in our understanding of mammalian systems of epigenetic gene regulation in the context of the long-running evolutionary conflicts that have created them.