Abstract

The APETALA2 (AP2) genes represent the AP2 group within a large group of DNA-binding proteins called AP2/EREBP. The AP2 gene is functional and necessary for flower development, stem cell maintenance, and seed development, whereas the other members of AP2 group redundantly affect flowering time. Here we study the phylogeny of AP2 group genes in spermatophytes. Spermatophyte AP2 group genes can be classified into AP2 and TOE types, six clades, and we found that the AP2 group homologs in gymnosperms belong to the AP2 type, whereas TOE types are absent, which indicates the AP2 type gene are more ancient and TOE type was split out of AP2 type and losing the major function. In Brassicaceae, the expansion of AP2 and TOE type lead to the gene number of AP2 group were up to six. Purifying selection appears to have been the primary driving force of spermatophyte AP2 group evolution, although positive selection occurred in the AP2 clade. The transition from exon to intron of AtAP2 in Arabidopsis mutant leads to the loss of gene function and the same situation was found in AtTOE2. Combining this evolutionary analysis and published research, the results suggest that typical AP2 group genes may first appear in gymnosperms and diverged in angiosperms, following expansion of group members and functional differentiation. In angiosperms, AP2 genes (AP2 clade) inherited key functions from ancestors and other genes of AP2 group lost most function but just remained flowering time controlling in gene formation. In this study, the phylogenies of AP2 group genes in spermatophytes was analyzed, which supported the evidence for the research of gene functional evolution of AP2 group.

Highlights

  • The genes in AP2/ERF family can be divided into subfamilies according to the number of AP2/ERF domains

  • Blast analysis of spermatophyte AP2 group gene sequences revealed that only some species in Brassicaceae contain five or six AP2 group members

  • We found no orthologs of AP2 group genes in our database searches of alga, moss and fern

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Summary

Introduction

The genes in AP2/ERF family can be divided into subfamilies according to the number of AP2/ERF domains. There are seven conservative domains in a typical AP2 group gene, including one ethylene-responsive element binding factors (ERF) -associated amphiphilic repression motif (EAR) (Kagale et al, 2010) or EAR-like domain, a nuclear localization signal (NLS) domain, two AP2 (AP2-R1 and AP2-R2) domain (Kim et al, 2006), a linkage domain (connecting the AP2R1 with R2), another EAR domain, and a MIR172 target site (Image 1). There are six members in the AP2 gene group in Arabidopsis, TARGET OF EARLY ACTIVATION TAGGED 1-3 (TOE1-3), AP2, SCHLAFMUTZE (SMZ), and SCHNARCHZAPFEN (SNZ). Among these six Arabidopsis genes, AP2, TOE3, and TOE1 contain both complete AP2 domains (AP2-R1 and R2 domains) but there is only one typical AP2 domain (AP2-R1 domain) in TOE2, SMZ, and SNZ (Image 1). The AP2-R2 domain in these three genes are not the same as AP2, TOE3, and TOE1

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