The modern versus extended evolutionary synthesis: Sketch of an intra-genomic gene's eye view for the evolutionary-genetic underpinning of epigenetic and developmental evolution

Studying the phenotypic evolution of organisms in terms of populations of genes and genotypes, the Modern Synthesis (MS) conceptualizes biological evolution in terms of 'inter-organismal' interactions among genes sitting in the di erent individual organisms that constitute a population. It 'black-boxes' the complex 'intra-organismic' molecular and developmental epigenetics mediat- ing between genotypes and phenotypes. To conceptually integrate epigenetics and evo-devo into evolutionary theory, advocates of an Extended Evolutionary Synthesis (EES) argue that the MS's reductive gene-centrism should be abandoned in favor of a more inclusive organism-centered ap- proach. To push the debate to a new level of understanding, we introduce the evolutionary biology of 'intra-genomic con ict' (IGC) to the controversy. This strategy is based on a twofold rationale. First, the eld of IGC is both ‘gene-centered’ and 'intra-organismic' and, as such, could build a bridge between the gene-centered MS and the intra-organismic elds of epigenetics and evo-devo. And second, it is increasingly revealed that IGC plays a signi cant causal role in epigenetic and developmental evolution and even in speciation. Hence, to deal with the ‘discrepancy’ between the ‘gene-centered’ MS and the ‘intra-organismic’ elds of epigenetics and evo-devo, we sketch a conceptual solution in terms of ‘intra-genomic con ict and compromise’ – an ‘intra-genomic gene’s eye view’ that thinks in terms of intra-genomic ‘evolutionarily stable strategies’ (ESSs) among numerous and various DNA regions and elements – to evolutionary-genetically under- write both epigenetic and developmental evolution, as such questioning the ‘gene-de-centered’ stance put forward by EES-advocates.