title: Tinkering with the C-function: a molecular frame for the selection of double flowers in cultivated roses.
abstract: BACKGROUND
  Roses have been cultivated for centuries and a number of varieties have been selected based on flower traits such as petal form, color, and number. Wild-type roses have five petals (simple flowers), whereas high numbers of petals (double flowers) are typical attributes of most of the cultivated roses. Here, we investigated the molecular mechanisms that could have been selected to control petal number in roses.
  
  
  RESULTS
  We have analyzed the expression of several candidate genes known to be involved in floral organ identity determination in roses from similar genetic backgrounds but exhibiting contrasting petal numbers per flower. We show that the rose ortholog of AGAMOUS (RhAG) is differentially expressed in double flowers as compared to simple flowers. In situ hybridization experiments confirm the differential expression of RhAG and demonstrate that in the double-flower roses, the expression domain of RhAG is restricted toward the center of the flower. Conversely, in simple-flower roses, RhAG expression domain is wider. We further show that the border of RhAG expression domain is labile, which allows the selection of rose flowers with increased petal number. Double-flower roses were selected independently in the two major regions for domestication, China and the peri-Mediterranean areas. Comparison of RhAG expression in the wild-type ancestors of cultivated roses and their descendants both in the European and Chinese lineages corroborates the correlation between the degree of restriction of RhAG expression domain and the number of petals. Our data suggests that a restriction of RhAG expression domain is the basis for selection of double flowers in both the Chinese and peri-Mediterranean centers of domestication.
  
  
  CONCLUSIONS
  We demonstrate that a shift in RhAG expression domain boundary occurred in rose hybrids, causing double-flower phenotype. This molecular event was selected independently during rose domestication in Europe/Middle East and in China.
authors.affiliation: Reproduction et Développement des Plantes, Université Lyon, Lyon, France.
authors.forename: Annick
authors.initials: A
authors.lastname: Dubois
authors.forename: Olivier
authors.initials: O
authors.lastname: Raymond
authors.forename: Marion
authors.initials: M
authors.lastname: Maene
authors.forename: Sylvie
authors.initials: S
authors.lastname: Baudino
authors.forename: Nicolas B
authors.initials: NB
authors.lastname: Langlade
authors.forename: Véronique
authors.initials: V
authors.lastname: Boltz
authors.forename: Philippe
authors.initials: P
authors.lastname: Vergne
authors.forename: Mohammed
authors.initials: M
authors.lastname: Bendahmane
mesh_terms: D035264:Flowers / Q000033:anatomy & histology / Q000235:genetics* / Q000254:growth & development; D020869:Gene Expression Profiling; D018507:Gene Expression Regulation, Developmental; D018506:Gene Expression Regulation, Plant; D014644:Genetic Variation; D006824:Hybridization, Genetic; D017403:In Situ Hybridization; D026161:MADS Domain Proteins / Q000235:genetics*; D008953:Models, Anatomic; D010641:Phenotype; D010940:Plant Proteins / Q000235:genetics*; D020133:Reverse Transcriptase Polymerase Chain Reaction; D027826:Rosa / Q000033:anatomy & histology / Q000737:chemistry / Q000235:genetics*; D055549:Volatile Organic Compounds / Q000032:analysis
chemical_list: D026161:MADS Domain Proteins; D010940:Plant Proteins; D055549:Volatile Organic Compounds