Summary model for the origin, evolution, and neofunctionalization of the Pooideae ERF014s. The AP2 family and the common ancestor of the RAV and ERF families have evolved independently in green plants. The split of the RAV and ERF families occurred in the common ancestor of Zygnemophyceae and Embryophyta. The DREB subfamily originated from the ERF subfamily in the common ancestor of Zygnemophyceae after the divergence of Charophyceae and Zygnemophyceae. The ancestral archetypal DREB genes expanded into three archetypal groups of land plants in Zygnemophyceae. One of the three groups, group archetype-II/III, broadly expanded during land plant radiation, resulting in nine subgroups in angiosperms and one moss-specific group in mosses (left panel). Subgroup IIb-1 genes have evolved independently in the radiation of dicots and monocots. After three paleopolyploidy events in Brassicaceae and two in Poaceae, five parologs of subgroup IIb-1 in Brassicaceae, and three in Poaceae were detected in this study, respectively (middle panel). We infer that one duplicate of ERF014s was lost in each of the taxonomic families after the WGDs (dotted lines in middle panel). The orthologs of monocot ERF014s have undergone chromosomal rearrangements to form sHSP-ERF014 locus in Poaceae and neofunctionalized to response to heat stress in Pooideae (right panel). The time shown in the time scales is from Morris et al. (2018) and Wu et al. (2020).
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