Modularity and reshuffling of Snail and Slug expression during vertebrate evolution

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Abstract
Gene duplication has been a major mechanism for increasing genomic complexity and variation during evolution. The evolutionary history of duplicated genes has been poorly studied along the vertebrate lineage. Here, we attempt to study that history by analyzing the expression of two members of the Snail family, Snail and Slug, in representatives of the major vertebrate groups. We find a surprising degree of variability in a subset of the expression sites for both genes in different species. Although some of the changes can be explained by neofunctionalization or subfunctionalization, others imply reciprocal changes in the expression of the two genes and the reappearance of expression in sites lost earlier in evolution. Because these changes do not fit easily into current models, we need to invoke additional mechanisms acting on enhancer elements to distribute expression domains and functions of duplicated genes unequally during evolution.