Gene Duplication and Differential Expression of Flower Symmetry Genes in Rhododendron (Ericaceae)

Bilaterally symmetric flowers have evolved over a hundred times in angiosperms, yet orthologs of the transcription factors CYCLOIDEA (CYC), RADIALIS (RAD), and DIVARICATA (DIV) are repeatedly implicated in floral symmetry changes. We examined these candidate genes to elucidate the genetic underpinnings of floral symmetry changes in florally diverse Rhododendron, reconstructing gene trees and comparing gene expression across floral organs in representative species with radial and bilateral flower symmetries. Radially symmetric R. taxifolium Merr. and bilaterally symmetric R. beyerinckianum Koord. had four and five CYC orthologs, respectively, from shared tandem duplications. CYC orthologs were expressed in the longer dorsal petals and stamens and highly expressed in R. beyerinckianum pistils, whereas they were either ubiquitously expressed, lost from the genome, or weakly expressed in R. taxifolium. Both species had two RAD and DIV orthologs uniformly expressed across all floral organs. Differences in gene structure and expression of Rhododendron RAD compared to other asterids suggest that these genes may not be regulated by CYC orthologs. Our evidence supports CYC orthologs as the primary regulators of differential organ growth in Rhododendron flowers, while also suggesting certain deviations from the typical asterid gene regulatory network for flower symmetry.

Mechanism of Leaf Vein Coloration and Inheritance of Leaf Vein Color, Flower Form, and Floral Symmetry in Gloxinia

Double-flowered gloxinia (Sinningia speciosa) cultivars with foliar variegation might have a greater market appeal as flowering foliage plants. Crosses were made among 16 gloxinia cultivars and their progenies were analyzed to determine the inheritance of leaf vein color, flower form, and floral symmetry. All plants from self-pollinating white-veined cultivars or crosses between white-veined and green-veined cultivars produced white veins. Progeny derived from self-pollinating plants of white-veined cultivars × green-veined cultivars segregated into a ratio of 3 white-veined:1 green-veined. All plants from self-pollinating or cross-pollinating single-flowered cultivars produced single flowers. Progeny of self-pollination or crosses between double-flowered cultivars segregated into a ratio of 3 double flowers:1 single flower. Contingency chi-square tests revealed that leaf vein color and flower form were inherited independently. New gloxinia progenies with homozygous white veins and double flowers were successfully developed from the F2 segregating population. Plants from self-pollinating or cross-pollinating cultivars with actinomorphic flowers produced actinomorphic flowers. A single dominant gene expressed in the homozygous or heterozygous state resulted in the zygomorphic flowers. Independent inheritance was observed between vein color and floral symmetry. Air spaces between the epidermis and the mesophyll cells were observed in the white, but not in the green, leaf vein portions. Net photosynthesis did not differ significantly between the white vein and adjacent green portion of the same leaf.

Evolution of pollination syndromes and corolla symmetry in Balsaminaceae reconstructed using phylogenetic comparative analyses

Background and Aims Floral diversity as a result of plant–pollinator interactions can evolve by two distinct processes: shifts between pollination systems or divergent use of the same pollinator. Although both are pollinator driven, the mode, relative importance and interdependence of these different processes are rarely studied simultaneously. Here we apply a phylogenetic approach using the Balsaminaceae (including the species-rich genus Impatiens) to simultaneously quantify shifts in pollination syndromes (as inferred from the shape and colour of the perianth), as well as divergent use of the same pollinator (inferred from corolla symmetry). Methods For 282 species we coded pollination syndromes based on associations between floral traits and known pollination systems, and assessed corolla symmetry. The evolution of these traits was reconstructed using parsimony- and model-based approaches, using phylogenetic trees derived from phylogenetic analyses of nuclear ribosomal and plastid DNA sequence data. Key Results A total of 71 % of studied species have a bee pollination syndrome, 22 % a bimodal syndrome (Lepidoptera and bees), 3 % a bird pollination syndrome and 5 % a syndrome of autogamy, while 19 % of species have an asymmetrical corolla. Although floral symmetry and pollination syndromes are both evolutionarily labile, the latter shifts more frequently. Shifts in floral symmetry occurred mainly in the direction towards asymmetry, but there was considerable uncertainty in the pattern of shift direction for pollination syndrome. Shifts towards asymmetrical flowers were associated with a bee pollination syndrome. Conclusion Floral evolution in Impatiens has occurred through both pollination syndrome shifts and divergent use of the same pollinator. Although the former appears more frequent, the latter is likely to be underestimated. Shifts in floral symmetry and pollination syndromes depend on each other but also partly on the region in which these shifts take place, suggesting that the occurrence of pollinator-driven evolution may be determined by the availability of pollinator species at large geographical scales.

Zygomorphic flowers have fewer potential pollinator species

Botanists have long identified bilaterally symmetrical (zygomorphic) flowers with more specialized pollination interactions than radially symmetrical (actinomorphic) flowers. Zygomorphic flowers facilitate more precise contact with pollinators, guide pollinator behaviour and exclude less effective pollinators. However, whether zygomorphic flowers are actually visited by a smaller subset of available pollinator species has not been broadly evaluated. We compiled 53 609 floral visitation records in 159 communities and classified the plants' floral symmetry. Globally and within individual communities, plants with zygomorphic flowers are indeed visited by fewer species. At the same time, zygomorphic flowers share a somewhat larger proportion of their visitor species with other co-occurring plants and have particularly high sharing with co-occurring plants that also have zygomorphic flowers. Visitation sub-networks for zygomorphic species also show differences that may arise from reduced visitor diversity, including greater connectance, greater web asymmetry and lower coextinction robustness of both plants and visitor species—but these changes do not necessarily translate to whole plant-visitor communities. These results provide context for widely documented associations between zygomorphy and diversification and imply that species with zygomorphic flowers may face a greater risk of extinction due to pollinator loss.

Radial or Bilateral? The Molecular Basis of Floral Symmetry

In the plant kingdom, the flower is one of the most relevant evolutionary novelties. Floral symmetry has evolved multiple times from the ancestral condition of radial to bilateral symmetry. During evolution, several transcription factors have been recruited by the different developmental pathways in relation to the increase of plant complexity. The MYB proteins are among the most ancient plant transcription factor families and are implicated in different metabolic and developmental processes. In the model plant Antirrhinum majus, three MYB transcription factors (DIVARICATA, DRIF, and RADIALIS) have a pivotal function in the establishment of floral dorsoventral asymmetry. Here, we present an updated report of the role of the DIV, DRIF, and RAD transcription factors in both eudicots and monocots, pointing out their functional changes during plant evolution. In addition, we discuss the molecular models of the establishment of flower symmetry in different flowering plants.

Ovule and seed production patterns in relation to flower size variations in actinomorphic and zygomorphic flower species

Zygomorphic flower species tend to show lower flower size variation than actinomorphic flower species. Have these differences also brought an association in ovule and seed production that has arisen due to natural selection in these species? Flowers were collected from 29 actinomorphic and 20 zygomorphic flower species, and fruits were collected from 21 actinomorphic and 14 zygomorphic flower species in Miyagi and Aomori prefectures, in Japan. The coefficient of variations (CVs) of flower sizes, mean ovule sizes of flowers, ovule numbers of flowers and mean seed sizes of fruits were calculated. The CV of flower sizes was marginally different between the floral symmetry types; tending to be lower in the zygomorphic flower species than in the actinomorphic flower species. The CVs of mean ovule sizes and ovule numbers of flowers increased with increase in the CV of flower sizes in the actinomorphic flower species but not in the zygomorphic flower species. Mean ovule number of flowers tends to increase with increase in mean flower size in the actinomorphic flower species but not in the zygomorphic flower species. The degrees in variations in ovule size and number of flowers were influenced by the interaction of floral symmetry type and flower size variation, suggesting that floral symmetry also has brought an evolutionary association in ovule production by flowers.

Zygomorphic flowers have fewer potential pollinator species

ABSTRACTBotanists have long identified bilaterally symmetrical (zygomorphic) flowers with more specialized pollination interactions than radially symmetrical (actinomorphic) flowers. Zygomorphic flowers facilitate more precise contact with pollinators, guide pollinator behaviour, and exclude less effective pollinators. However, whether zygomorphic flowers are actually visited by a smaller subset of available pollinator species has not been broadly evaluated. We compiled 53,609 floral visitation records in 159 communities and classified the plants’ floral symmetry. Globally and within individual communities, plants with zygomorphic flowers are indeed visited by fewer species. At the same time, zygomorphic flowers share a somewhat larger proportion of their visitor species with other co-occurring plants, and have particularly high sharing with co-occurring plants that also have zygomorphic flowers. Visitation sub-networks for zygomorphic species also show differences that may arise from reduced visitor diversity, including greater connectance, greater web asymmetry, and lower coextinction robustness of both plants and visitor species — but these changes do not necessarily translate to whole plant-visitor communities. These results provide context for widely documented associations between zygomorphy and diversification and imply that species with zygomorphic flowers may face greater risk of extinction due to pollinator loss.