The Rosea1, Rosea2, and Venosa genes encode MYB-related transcription factors active in the flowers of Antirrhinum majus. Analysis of mutant phenotypes shows that these genes control the intensity and pattern of magenta anthocyanin pigmentation in flowers. Despite the structural similarity of these regulatory proteins, they influence the expression of target genes encoding the enzymes of anthocyanin biosynthesis with different specificities. Consequently, they are not equivalent biochemically in their activities. Different species of the genus Antirrhinum, native to Spain and Portugal, show striking differences in their patterns and intensities of floral pigmentation. Differences in anthocyanin pigmentation between at least six species are attributable to variations in the activity of the Rosea and Venosa loci. Set in the context of our understanding of the regulation of anthocyanin production in other genera, the activity of MYB-related genes is probably a primary cause of natural variation in anthocyanin pigmentation in plants.

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...

Summary• Pigment stripes associated with veins (venation) is a common flower colour pattern. The molecular genetics and function of venation were investigated in the genus Antirrhinum, in which venation is determined by Venosa (encoding an R2R3MYB transcription factor).• Pollinator preferences were measured by field tests with Antirrhinum majus. Venosa function was examined using in situ hybridization and transient overexpression. The origin of the venation trait was examined by molecular phylogenetics.• Venation and full-red flower colouration provide a comparable level of advantage for pollinator attraction relative to palely pigmented or white lines. Ectopic expression of Venosa confers pigmentation outside the veins. Venosa transcript is produced only in small areas of the corolla between the veins and the adaxial epidermis. Phylogenetic analyses suggest that venation patterning is an ancestral trait in Antirrhinum. Different accessions of three species with full-red pigmentation with or without venation patterning have been found.• Epidermal-specific venation is defined through overlapping expression domains of the MYB (myoblastoma) and bHLH (basic Helix-Loop-Helix) co-regulators of anthocyanin biosynthesis, with the bHLH providing epidermal specificity and Venosa vein specificity. Venation may be the ancestral trait, with full-red pigmentation a derived, polyphyletic trait. Venation patterning is probably not fixed once species evolve full-red floral pigmentation.

In higher plants several isoforms of starch synthase contribute to the extension of glucan chains in the synthesis of starch. Different isoforms are responsible for the synthesis of essentially linear amylose chains and branched, amylopectin chains. The activity of granule-bound starch synthase I from potato has been compared with that of starch synthase II from potato following expression of both isoforms in Escherichia coli. Significant differences in their activities are apparent which may be important in determining their specificities in vivo. These differences include affinities for ADPglucose and glucan substrates, activation by amylopectin, response to citrate, thermosensitivity and the processivity of glucan chain extension. To define regions of the isoforms determining these characteristic traits, chimeric proteins have been produced by expression in E. coli. These experiments reveal that the C-terminal region of granule-bound starch synthase I confers most of the specific properties of this isoform, except its processive elongation of glucan chains. This region of granule-bound starch synthase I is distinct from the C-terminal region of other starch synthases. The specific properties it confers may be important in defining the specificity of granule-bound starch synthase I in producing amylose in vivo.Keywords: amylopectin; amylose; starch synthase.In plants, starch synthases form starch by catalysing the transfer of the glucosyl moiety from ADPglucose to the nonreducing ends of glucan chains through an a1±4 linkage. Starch is composed of two types of glucan polymer: amylose, a predominantly linear a1±4-linked glucan and amylopectin, in which the a1±4 linked chains are highly branched by a1±6 linkages, which are formed by starch branching enzymes. Several isoforms of starch synthase are involved in starch formation and these can be classified according to their localization within the plastid; one isoform, granule-bound starch synthase I (GBSSI), is bound exclusively to the starch granule and others are localized predominantly in the soluble phase (stroma) of the plastid. In general, GBSSI is a protein of < 60 kDa. Through mutant analysis GBSSI is known to be solely responsible for the synthesis of amylose, because the waxy mutants of cereals, the amf mutant of potato, the lam mutants of pea and GBSSI antisense potato lines all involve the reduction or elimination of GBSSI activity and a specific reduction or elimination of amylose in the starch of the storage organs of these species [1±4].Among the starch synthase isoforms found in the plastid stroma of higher plants, starch synthase II (SSII) is also found bound to starch granules in significant amounts [5±9]. However, SSII cannot compensate for the effects of loss of GBSSI on amylose synthesis in amf mutant potatoes, indicating that SSII is not involved in amylose synthesis in vivo [8]. In the storage organs of different higher plants, SSII may make quite different quantitative contributions to total starch synthase activity. Mutant analysis in pea...

A major innovation in angiosperms is the recruitment of animal pollinators as a means to enhance the efficiency and specificity of pollen transfer. The implementation of this reproductive strategy involved the rapid and presumably coordinated evolution of multiple floral traits. A major question concerns the molecular identity of the genetic polymorphisms that specify the phenotypic differences between distinct pollination syndromes. Here, we report on our work with Petunia, an attractive model system for quantitative plant genetics and genomics. From interspecific crosses, we obtained F2 plants that differed in the length of the floral tube or the size of the limb. We used these plants to study the behaviour of the hawkmoth pollinator, Manduca sexta. Plants with larger limbs were preferentially visited, consistent with the notion that flower size affects visibility under low light conditions. The moths also displayed an innate preference for shorter tubes. However, in those cases that flowers with long tubes were chosen, the animals fed for equal time. Thus, the perception of tube length may help the moths, early on, to avoid those plants that are more difficult to handle.

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HighlightPlants carefully control where and when flowers are made through activators and repressors. We show that spatially the shoot meristem is key in responding to the repressors of flowering TFL1.

Switches between pollination syndromes have happened frequently during angiosperm evolution. Using QTL mapping and reciprocal introgressions, we show that changes in reproductive organ morphology have a simple genetic basis. In animal-pollinated plants, flowers have evolved to optimize pollination efficiency by different pollinator guilds and hence reproductive success. The two Petunia species, P. axillaris and P. exserta, display pollination syndromes adapted to moth or hummingbird pollination. For the floral traits color and scent, genetic loci of large phenotypic effect have been well documented. However, such large-effect loci may be typical for shifts in simple biochemical traits, whereas the evolution of morphological traits may involve multiple mutations of small phenotypic effect. Here, we performed a quantitative trait locus (QTL) analysis of floral morphology, followed by an in-depth study of pistil and stamen morphology and the introgression of individual QTL into reciprocal parental backgrounds. Two QTLs, on chromosomes II and V, are sufficient to explain the interspecific difference in pistil and stamen length. Since most of the difference in organ length is caused by differences in cell number, genes underlying these QTLs are likely to be involved in cell cycle regulation. Interestingly, conservation of the locus on chromosome II in a different P. axillaris subspecies suggests that the evolution of organ elongation was initiated on chromosome II in adaptation to different pollinators. We recently showed that QTLs for pistil and stamen length on chromosome II are tightly linked to QTLs for petal color and volatile emission. Linkage of multiple traits will enable major phenotypic change within a few generations in hybridizing populations. Thus, the genomic architecture of pollination syndromes in Petunia allows for rapid responses to changing pollinator availability.