Petals of animal-pollinated angiosperms have adapted to attract pollinators. Factors influencing pollinator attention include colour and overall size of flowers. Colour is determined by the nature of the pigments, their environment and by the morphology of the petal epidermal cells. Most angiosperms have conical epidermal cells, which enhance the colour intensity and brightness of petal surfaces. The MYB-related transcription factor MIXTA controls the development of conical epidermal cells in petals of Antirrhinum majus. Another gene encoding an R2R3 MYB factor very closely related to MIXTA, AmMYBML2, is also expressed in flowers of A. majus. We have analysed the roles of AmMYBML2 and two MIXTA-related genes, PhMYB1 from Petunia hybrida and AtMYB16 from Arabidopsis thaliana, in petal development. The structural similarity between these genes, their comparable expression patterns and the similarity of the phenotypes they induce when ectopically expressed in tobacco, suggest they share homologous functions closely related to, but distinct from, that of MIXTA. Detailed phenotypic analysis of a phmyb1 mutant confirmed the role of PhMYB1 in the control of cell morphogenesis in the petal epidermis. The phmyb1 mutant showed that epidermal cell shape affects petal presentation, a phenotypic trait also observed following re-examination of mixta mutants. This suggests that the activity of MIXTA-like genes also contributes to petal form, another important factor influencing pollinator attraction.KEY WORDS: Petal, Cell shape, Petunia, Antirrhinum, MYB transcription factor Development 134, 1691Development 134, -1701Development 134, (2007
DEVELOPMENT
1692Emerging data from the study of transcription factors belonging to large families of structurally related proteins suggest that very similar members of phylogenetically-clustered subgroups usually share closely related functions, even though functions may have diverged over the entire family. Structural similarity has led to claims of orthology and functional equivalence between MIXTA, PhMYB1 and AtMYB16 (van Houwelingen et al., 1998;Romero et al., 1998). However, to achieve a general understanding of the control of morphogenesis of petal epidermal cells, the function of new genes needs to be assayed and compared with that of the prototype, MIXTA. In addition, the relevance of these genes to cell shaping and, specifically, to their roles in adapting petals for pollinator attraction, needs to be established in different angiosperm species.We have examined the function of three of the genes encoding proteins very closely related to MIXTA; PhMYB1 from Petunia hybrida, AmMYBML2 from A. majus and AtMYB16 from Arabidopsis thaliana. Structurally, these proteins are most closely related to each other and their genes are therefore orthologous. All three proteins promote directional cell expansion in a bioassay in tobacco. The similarities between the phenotypes induced by these proteins in this bioassay (their biochemical functions) and the similarities in their expression patter...