Secondary pollen presentation is the developmental relocation of pollen from the anthers onto another floral organ which then functions as the pollen presenting organ for pollination. Nine different types have been identified in sixteen angiosperm families according to which organ is used for presentation, whether the pollen is exposed or concealed within a structure and how pollen is loaded onto the presenting surface: (1) Enveloping bloom presenters (Araceae); (2) Perianth presenters with exposed pollen presentation (Epacridaceae); (3) Androecial presenters (Santalaceae); (4) Terminal stylar presenters with passive pollen placement and concealed stigmas (Rubiaceae and Proteaceae); (5) Terminal stylar presenters with passive pollen placement and sub-terminal stigmas (Marantaceae and Polygalaceae); (6) Terminal stylar presenters with active pollen placement (Asteraceae, Calyceraceae and Lobeliaceae); (7) Sub-terminal stylar presenters (Campanulaceae, Cannaceae, Fabaceae and Myrtaceae); (8) Exposed stigmatic presenters (Rubiaceae); (9) Indusial stigmatic presenters (Goodeniaceae and Brunoniaceae). Secondary pollen presentation occurs in three monocotyledon and thirteen dicotyledon families. The presentation types appear to have been independently derived indicating that secondary pollen presentation is a character with a selective advantage.In all but the enveloping bloom type of secondary pollen presentation, developmental relocation of pollen requires simultaneous, introrse anther dehiscence and a close association of the presenting organ to the anthers prior to anthesis.The various secondary pollen presentation systems may be modified to promote xenogamy or autogamy and this can even change during anthesis. Most plants which have secondary pollen presentation, display reduced herkogamy within the flower to facilitate pollination. Increased risk of self-pollination due to this may be overcome through dichogamy, herkogamy within inflorescences, dry stigmas, self-incompatibility systems and passive or active control over pollinator behaviour. Enhanced male function of the flowers of secondary pollen presenting plants is also evident through extension of the male phase by the protection, controlled release and precise placement and receipt of pollen. Plants displaying secondary pollen presentation are almost always protandrous.
Merkel cell carcinoma (MCC) is an aggressive skin cancer of neuroendocrine origin with a high propensity for recurrence and metastasis. Merkel cell polyomavirus (MCPyV) causes the majority of MCC cases due to the expression of the MCPyV small and large tumor antigens (ST and LT, respectively). Although a number of molecular mechanisms have been attributed to MCPyV tumor antigen-mediated cellular transformation or replication, to date, no studies have investigated any potential link between MCPyV T antigen expression and the highly metastatic nature of MCC. Here we use a quantitative proteomic approach to show that MCPyV ST promotes differential expression of cellular proteins implicated in microtubule-associated cytoskeletal organization and dynamics. Intriguingly, we demonstrate that MCPyV ST expression promotes microtubule destabilization, leading to a motile and migratory phenotype. We further highlight the essential role of the microtubule-associated protein stathmin in MCPyV ST-mediated microtubule destabilization and cell motility and implicate the cellular phosphatase catalytic subunit protein phosphatase 4C (PP4C) in the regulation of this process. These findings suggest a possible molecular mechanism for the highly metastatic phenotype associated with MCC.IMPORTANCE Merkel cell polyomavirus (MCPyV) causes the majority of cases of Merkel cell carcinoma (MCC), an aggressive skin cancer with a high metastatic potential. However, the molecular mechanisms leading to virally induced cancer development have yet to be fully elucidated. In particular, no studies have investigated any potential link between the virus and the highly metastatic nature of MCC. We demonstrate that the MCPyV small tumor antigen (ST) promotes the destabilization of the host cell microtubule network, which leads to a more motile and migratory cell phenotype. We further show that MCPyV ST induces this process by regulating the phosphorylation status of the cellular microtubule-associated protein stathmin by its known association with the cellular phosphatase catalytic subunit PP4C. These findings highlight stathmin as a possible biomarker of MCC and as a target for novel antitumoral therapies.
In Crinum flaccidum the anthers are versatile and tetrasporangiate with a secretory tapetum of binucleate cells. Successive cytokinesis in microspore mother cell results in isobilateral and decussate microspore tetrads. The mature pollen grains are single, spheroidal, disulculate, echinate and 2-celled. In the mature anthers, fibrous thickenings develop not only in the endothecium but also in two or three middle cell layers and the connective tissue before latrorse dehiscence. A lobed tissue in each of the three locules of the ovary serves ovular and placental functions. Each extension of the 5-7 paired lobes represents an ategmic ovule. The development of the female gametophyte conforms to the Polygonum type. Usually only one gametophyte is present in each lobe but occasionally several may occur. Bulb growth is monopodial with normally three umbels produced per plant, each carrying an average of 10 flowers, only two or three of which are open at any one time. Nectar sugar concentration was measured at 14.2% (w/w), of which 44.8% of solids was sucrose and 3.9% either glucose or fructose. The protandrous flowers are phalenophilous, pollinated by sphingid moths. The endosperm formation is of the nuclear type. In the absence of seed coats and the nucellus at maturity, the outer layers of the endosperm become corky following the activity of a phellogen. Embryogeny appears to be of the Asterad type. The mature embryo is straight and chlorophyllous. The large (5.3 g) seeds are 89% water and show no dormancy, germinating without an external supply of water, sometimes while still on the parent plant.
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