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Background  The floral homeotic C function gene  AGAMOUS  ( AG ) confers stamen and carpel identity and is involved in the regulation of floral meristem termination in  Arabidopsis .  Arabidopsis ag  mutants show complete homeotic conversions of stamens into petals and carpels into sepals as well as indeterminacy of the floral meristem. Gene function analysis in model core eudicots and the monocots rice and maize suggest a conserved function for  AG  homologs in angiosperms. At the same time gene phylogenies reveal a complex history of gene duplications and repeated subfunctionalization of paralogs.    Results   EScaAG1  and  EScaAG2 , duplicate  AG  homologs in the basal eudicot  Eschscholzia californica  show a high degree of similarity in sequence and expression, although  EScaAG2  expression is lower than  EScaAG1  expression. Functional studies employing virus-induced gene silencing (VIGS) demonstrate that knock down of  EScaAG1  and  2  function leads to homeotic conversion of stamens into petaloid structures and defects in floral meristem termination. However, carpels are transformed into petaloid organs rather than sepaloid structures. We also show that a reduction of  EScaAG1  and  EScaAG2  expression leads to significantly increased expression of a subset of floral homeotic B genes.    Conclusions  This work presents expression and functional analysis of the two basal eudicot  AG  homologs. The reduction of  EScaAG1  and  2  functions results in the change of stamen to petal identity and a transformation of the central whorl organ identity from carpel into petal identity. Petal identity requires the presence of the floral homeotic B function and our results show that the expression of a subset of B function genes extends into the central whorl when the C function is reduced. We propose a model for the evolution of B function regulation by C function suggesting that the mode of B function gene regulation found in  Eschscholzia  is ancestral and the C-independent regulation as found in  Arabidopsis  is evolutionarily derived.

evodevo

Floral homeotic C function genes repress specific B function genes in the carpel whorl of the basal eudicot California poppy (Eschscholzia californica)