title: Positive selection on the K domain of the AGAMOUS protein in the Zingiberales suggests a mechanism for the evolution of androecial morphology.
abstract: BACKGROUND
  The ABC model of flower development describes the molecular basis for specification of floral organ identity in model eudicots such as Arabidopsis and Antirrhinum. According to this model, expression of C-class genes is linked to stamen and gynoecium organ identity. The Zingiberales is an order of tropical monocots in which the evolution of floral morphology is characterized by a marked increase in petaloidy in the androecium. Petaloidy is a derived characteristic of the ginger families and seems to have arisen in the common ancestor of the ginger clade. We hypothesize that duplication of the C-class AGAMOUS (AG) gene followed by divergence of the duplicated AG copies during the diversification of the ginger clade lineages explains the evolution of petaloidy in the androecium. In order to address this hypothesis, we carried out phylogenetic analyses of the AG gene family across the Zingiberales and investigated patterns of gene expression within the androecium.
  
  
  RESULTS
  Phylogenetic analysis supports a scenario in which Zingiberales-specific AG genes have undergone at least one round of duplication. Gene duplication was immediately followed by divergence of the retained copies. In particular, we detect positive selection in the third alpha-helix of the K domain of Zingiberales AGAMOUS copy 1 (ZinAG-1). A single fixed amino acid change is observed in ZinAG-1 within the ginger clade when compared to the banana grade. Expression analyses of AG and APETALA1/FRUITFULL (AP1/FUL) in Musa basjoo is similar to A- and C-class gene expressions in the Arabidopsis thaliana model, while Costus spicatus exhibits simultaneous expression of AG and AP1/FUL in most floral organs. We propose that this novel expression pattern could be correlated with the evolution of androecial petaloidy within the Zingiberales.
  
  
  CONCLUSIONS
  Our results present an intricate story in which duplication of the AG lineage has lead to the retention of at least two diverged Zingiberales-specific copies, ZinAG-1 and Zingiberales AGAMOUS copy 2 (ZinAG-2). Positive selection on ZinAG-1 residues suggests a mechanism by which AG gene divergence may explain observed morphological changes in Zingiberales flowers. Expression data provides preliminary support for the proposed mechanism, although further studies are required to fully test this hypothesis.
authors.affiliation: Department of Plant and Microbial Biology, University of California, Berkeley, 111 Koshland Hall, Berkeley, CA 94720 USA ; Programa de Pós-Graduação em Genética e Biodiversidade, Universidade Federal da Bahia, Campus de Ondina, Salvador, BA 40170-290 Brazil.
authors.forename: Ana Maria R
authors.initials: AM
authors.lastname: Almeida
authors.affiliation: Department of Integrative Biology and the University and Jepson Herbaria, University of California, Berkeley, Berkeley, CA 94720 USA ; Muséum National d'Histoire Naturelle, Institut de Systématique, Évolution et Biodiversité. UMR 7205 CNRS, CP39, 16 Rue Buffon, 75231 Paris/Cedex 05, France ; Current address: Corporación Colombiana de Investigación (CORPOICA), Km 14 Vía Mosquera Bogotá, Colombia.
authors.forename: Roxana
authors.initials: R
authors.lastname: Yockteng
authors.affiliation: Departamento de Biologia Vegetal/BIOAGRO, Av. Peter Henry Rolfs s/n, Universidade Federal de Viçosa, Campus Viçosa, Viçosa, MG 36570-900 Brazil.
authors.forename: Wagner C
authors.initials: WC
authors.lastname: Otoni
authors.affiliation: Department of Plant and Microbial Biology, University of California, Berkeley, 111 Koshland Hall, Berkeley, CA 94720 USA ; Department of Integrative Biology and the University and Jepson Herbaria, University of California, Berkeley, Berkeley, CA 94720 USA.
authors.forename: Chelsea D
authors.initials: CD
authors.lastname: Specht
keywords: AGAMOUS; Androecial morphogenesis; Gene duplication; K domain; Petaloidy; Positive selection; Protein divergence; Zingiberales