Evolutionary significance of epigenetic variation

Several chapters in this volume demonstrate how epigenetic work at the molecular level over the last few decades has revolutionized our understanding of genome function and developmental biology. However, epigenetic processes not only further our understanding of variation and regulation at the genomic and cellular levels, they also challenge our understanding of heritable phenotypic variation at the level of whole organisms and even the process of evolution by natural selection (Jablonka and Lamb 1989, 1995; Danchin et al. 2011). Although many of the epigenetic mechanisms involved in differential gene expression are reset each generation, some epigenetic marks are faithfully transmitted across generations (Jablonka and Raz 2009; Verhoeven et al. 2010a). In addition, we now know that natural variation exists not only at the DNA sequence level but also the epigenetic level (e.g., Vaughn et al. 2007; Herrera and Bazaga 2010). This may be particularly common in plants, and several studies suggest that epigenetic variation alone can cause significant heritable variation in phenotypic traits (e.g., Cubas et al. 1999; Johannes et al. 2009; Scoville et al. 2011). Because of these observations, there is currently increasing interest in understanding the role of epigenetic processes in ecology and evolution (e.g., Richards 2006, 2011; Bossdorf et al. 2008; Johannes et al. 2008; Richards et al. 2010a)