Floral Development of Rhamnaceae and Origin of Its Unique Floral Features.

Rhamnaceae flowers have a peculiar morphology, including keeled sepals, one stamen whorl closely related to the petals, and a broad perigynous hypanthium that supports a voluminous nectary. In the present investigation, we detailed the flower development of five Rhamnaceae species to understand the origin of such specific floral characteristics. Floral buds and flowers were processed for surface and histological analyses. The sepals emerge in sequential order and the other organs in simultaneous order. The development of the perigynous hypanthium renders the floral apex broad and concave. The sepals undergo abaxial thickening early on, forming a keel and strongly influencing the floral merosity. Petals and stamens appear close to each other on the same radius in a very short plastochron. The carpels unite soon after their emergence, forming a syncarpous ovary and free style branches. Differences in intercalary carpel growth promote the formation of inferior (Gouania virgata) and semi-inferior ovaries (Colubrina glandulosa, Hovenia dulcis, and Sarcomphalus joazeiro). Rhamnidium elaeocarpum does not undergo such growth, and the resulting ovary is superior. The keeled sepals promote the isolation of the petal-stamen pair inside the flower bud. The possibility of a common primordium that the originates petal and stamen is refuted. Comparisons with other Rosales families provide insights into the floral origin and diversification of Rhamnaceae.

Evolution of the Subgroup 6 R2R3-MYB Genes and Their Contribution to Floral Color in the Perianth-Bearing Piperales.

Flavonoids, carotenoids, betalains, and chlorophylls are the plant pigments responsible for floral color. Anthocyanins, a class of flavonoids, are largely responsible for the red, purple, pink, and blue colors. R2R3-MYB genes belonging to subgroup 6 (SG6) are the upstream regulatory factors of the anthocyanin biosynthetic pathway. The canonical members of these genes in Arabidopsis include AtMYB75, AtMYB90, AtMYB113, and AtMYB114. The Aristolochiaceae is an angiosperm lineage with diverse floral groundplans and perianth colors. Saruma henryi exhibits a biseriate perianth with green sepals and yellow petals. All other genera have sepals only, with colors ranging from green (in Lactoris) to a plethora of yellow to red and purple mixtures. Here, we isolated and reconstructed the SG6 R2R3-MYB gene lineage evolution in angiosperms with sampling emphasis in Aristolochiaceae. We found numerous species-specific duplications of this gene lineage in core eudicots and local duplications in Aristolochiaceae for Saruma and Asarum. Expression of SG6 R2R3-MYB genes examined in different developmental stages and plant organs of four Aristolochiaceae species, largely overlaps with red and purple pigments, suggesting a role in anthocyanin and flavonoid synthesis and accumulation. A directed RNA-seq analysis corroborated our RT-PCR analyses, by showing that these structural enzymes activate during perianth development in Aristolochia fimbriata and that the regulatory genes are expressed in correlation with color phenotype. Finally, the reconstruction of the flavonoid and anthocyanin metabolic pathways using predicted peptides from transcriptomic data show that all pivotal enzymes are present in the analyzed species. We conclude that the regulatory genes as well as the biosynthetic pathway are largely conserved across angiosperms. In addition, the Aristolochiaceae emerges as a remarkable group to study the genetic regulatory network for floral color, as their members exhibit an outstanding floral diversity with elaborate color patterns and the genetic complement for SG6 R2R3-MYB genes is simpler than in core eudicot model species.

Gene expression underlying floral epidermal specialization in Aristolochia fimbriata (Aristolochiaceae).

BACKGROUND AND AIMS: The epidermis constitutes the outermost tissue of the plant body. Although it plays major structural, physiological and ecological roles in embryophytes, the molecular mechanisms controlling epidermal cell fate, differentiation and trichome development have been scarcely studied across angiosperms, and remain almost unexplored in floral organs. METHODS: In this study, we assess the spatio-temporal expression patterns of GL2, GL3, TTG1, TRY, MYB5, MYB6, HDG2, MYB106-like, WIN1 and RAV1-like homologues in the magnoliid Aristolochia fimbriata (Aristolochiaceae) by using comparative RNA-sequencing and in situ hybridization assays. KEY RESULTS: Genes involved in Aristolochia fimbriata trichome development vary depending on the organ where they are formed. Stem, leaf and pedicel trichomes recruit most of the transcription factors (TFs) described above. Conversely, floral trichomes only use a small subset of genes including AfimGL2, AfimRAV1-like, AfimWIN1, AfimMYB106-like and AfimHDG2. The remaining TFs, AfimTTG1, AfimGL3, AfimTRY, AfimMYB5 and AfimMYB6, are restricted to the abaxial (outer) and the adaxial (inner) pavement epidermal cells. CONCLUSIONS: We re-evaluate the core genetic network shaping trichome fate in flowers of an early-divergent angiosperm lineage and show a morphologically diverse output with a simpler genetic mechanism in place when compared to the models Arabidopsis thaliana and Cucumis sativus. In turn, our results strongly suggest that the canonical trichome gene expression appears to be more conserved in vegetative than in floral tissues across angiosperms.

Análisis morfológico comparativo de las estructuras reproductivas masculinas de los géneros Orthopterygium y Amphipterygium (Anacardiaceae)

Se presenta una descripción detallada de la inflorescencia y flor masculina del género Orthopterygium (A. Gray) Hemsl. en comparación con su género hermano Amphipterygium Standl., ambos anteriormente ubicados en la familia Julianiaceae, con la finalidad de corroborar sus estatus taxonómicos. Se incluyen todas las especies de ambos géneros, O. huaucui, A. molle, A. adstringens, A. simplicifolium y A. glaucum. Se realizaron medidas de las piezas florales (longitud de la flor, longitud y número de sépalos, longitud de las anteras y longitud de los filamentos) así como la longitud de la inflorescencia, éstas fueron sometidas a un análisis de componentes principales. Las estructuras diferenciales más significativas entre ambos géneros corresponden a la longitud y número de sépalos. Esta observación soporta y corrobora los resultados ya reportados por otros autores y constituye el primer avance en el establecimiento del actual estatus taxonómico de Orthopterygium, se sostiene por tanto la consideración de ambos géneros como entidades independientes.

We present a descriptive study on the male inflorescence and flowers of the genus Orthopterygium (A. Gray) Hemsl. in comparison with their sister genus Amphipterygium Standl., formerly recognized as Julianiaceae, with the aim of corroborate their current taxonomic status. We included all species of both genera O. huaucui, A. adstringens, A. molle, A. simplicifolium and A. glaucum. We measured floral pieces (flower length, sepal length, anther length and filament length) and inflorescence. These data were compared in a principal component analysis. Sepals length and sepals number are the most significant discriminant characters between both genera. This work supports and corroborates the results obtained by other authors and comprises the first initiative on the evaluation of the actual taxonomic status of Orthopterygium and establishes the distinction of both genera as independent entities.

Floral colleters in Pleurothallidinae (Epidendroideae: Orchidaceae).

PREMISE OF THE STUDY: The term colleter is applied to trichomes or emergences positioned close to developing vegetative and floral meristems that secrete a sticky, mucilaginous, and/or lipophilic exudate. Several ecological functions are attributed to these glands, but none are exclusive to colleters. Patterns of morphology and distribution of colleters may be valuable for systematics and phylogeny, especially concerning problematic and large groups such as the subtribe Pleurothallidinae, and are also essential to understand the evolution of these glands in Orchidaceae as a whole. METHODS: We used scanning electron and light microscopy to examine the structure and occurrence of trichomes on bracts and sepals and in the invaginations of the external ovary wall (IEOW) in flowers in several developmental stages from species in seven genera. KEY RESULTS: The exudate was composed of polysaccharides, lipophilic, and phenolic compounds. Colleters were secretory only during the development of floral organs, except for the glands in the IEOW that were also active in flowers at anthesis. After the secretory phase, fungal hyphae were found penetrating senescent trichomes. CONCLUSIONS: Trichome-like colleters seem to be a widespread character in Epidendroideae, and digitiform colleters are possibly the common type in this subfamily. Mucilage from IEOW colleters may aid in the establishment of symbiotic fungi necessary for seed germination. The presence of colleters in the IEOW may be a case of homeoheterotopy, in which extrafloral nectaries that produce simple sugar-based secretions (as in other orchid species) have changed to glands that produce secretions with complex polysaccharides, as in Pleurothallidinae.