Reproductive ecology and postpollination development in the hydrophilous monocot Ruppia maritima.

PREMISE: Water-pollination (hydrophily) is a rare but important pollination mechanism that has allowed angiosperms to colonize marine and aquatic habitats. Hydrophilous plants face unique reproductive challenges, and many have evolved characteristic pollen traits and pollination strategies that may have downstream consequences for pollen performance. However, little is known about reproductive development in the life history stage between pollination and fertilization (the progamic phase) in hydrophilous plants. The purpose of this study was to characterize reproductive ecology and postpollination development in water-pollinated Ruppia maritima L. METHODS: Naturally pollinated inflorescences of R. maritima were collected from the field. Experimental pollinations using both putatively outcross and self pollen were conducted in the greenhouse and inflorescences were collected at appropriate intervals after pollination. Pollen reception, pollen germination, pollen tube growth, and carpel morphology were characterized. RESULTS: Ruppia maritima exhibits incomplete protogyny, allowing for delayed selfing. Pollen germinated within 15 min after pollination. The average shortest possible pollen tube pathway was 425 µm and pollen tubes first reached the ovule at 45 min after pollination. The mean adjusted pollen tube growth rate was 551 µm/h. CONCLUSIONS: Ruppia pollen is adapted for rapid pollen germination, which is likely advantageous in an aquatic habitat. Small effective pollen loads suggest that pollen competition intensity is low. Selection for traits such as a long period of stigma receptivity, fast pollen germination, and carpel morphology likely played a larger role in shaping postpollination reproductive development in Ruppia than evolution in pollen tube growth rates.

Functional Characterization of Cnidarian HCN Channels Points to an Early Evolution of Ih.

HCN channels play a unique role in bilaterian physiology as the only hyperpolarization-gated cation channels. Their voltage-gating is regulated by cyclic nucleotides and phosphatidylinositol 4,5-bisphosphate (PIP2). Activation of HCN channels provides the depolarizing current in response to hyperpolarization that is critical for intrinsic rhythmicity in neurons and the sinoatrial node. Additionally, HCN channels regulate dendritic excitability in a wide variety of neurons. Little is known about the early functional evolution of HCN channels, but the presence of HCN sequences in basal metazoan phyla and choanoflagellates, a protozoan sister group to the metazoans, indicate that the gene family predates metazoan emergence. We functionally characterized two HCN channel orthologs from Nematostella vectensis (Cnidaria, Anthozoa) to determine which properties of HCN channels were established prior to the emergence of bilaterians. We find Nematostella HCN channels share all the major functional features of bilaterian HCNs, including reversed voltage-dependence, activation by cAMP and PIP2, and block by extracellular Cs+. Thus bilaterian-like HCN channels were already present in the common parahoxozoan ancestor of bilaterians and cnidarians, at a time when the functional diversity of voltage-gated K+ channels was rapidly expanding. NvHCN1 and NvHCN2 are expressed broadly in planulae and in both the endoderm and ectoderm of juvenile polyps.