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The SUMO-conjugating enzyme Ubc9 is an essential enzyme in the SUMO (small ubiquitin-related modifier) protein modification system. Although sumoylation, covalent modification of cellular proteins by SUMO, is considered to regulate various cellular processes, and many substrates for sumoylation have been identified recently, the regulation of Ubc9 expression has not been examined in detail. We analyzed the expression of Ubc9 during rat brain development at the mRNA and protein levels. Northern and Western blot analyses revealed that expression of Ubc9 and SUMO-1 was developmentally regulated, while that of the ubiquitin-conjugating enzyme UbcH7 did not change so dramatically. In situ hybridization analysis revealed that the expression of Ubc9 was high in neuronal stem cells and moderate in differentiated neurons at embryonic stages. In the adult brain, moderate expression was observed in subsets of neurons, such as the dentate granular neurons and pyramidal neurons in the hippocampal formation and the large pyramidal neurons in the cerebral cortex. These results suggest that the Ubc9-SUMO system might participate in the proliferation and differentiation of neuronal cells in the developing brain and in neuronal plasticity in the adult brain.
Premaxillary tooth count tends to be stable amongst toothed dinosaurs, and most theropods have four teeth in each premaxilla. Only one case of bilaterally asymmetric variation is known in theropod premaxillary dentition, and there is no record of ontogenetic or individual variation in premaxillary tooth count. Based on these observations, a tyrannosaurid left premaxilla with three teeth (TMP 2007.20.124) is an interesting deviation and represents an unusual individual of Daspletosaurus sp. with a developmental abnormality. The lower number of teeth is coupled with relatively larger alveoli, each of which is capable of hosting a larger than normal tooth. This indicates that tooth size and dental count vary inversely, and instances of reduction in tooth count may arise from selection for increased tooth size. On the other hand, the conservative number of premaxillary teeth in most theropods implies strong developmental constraints and a functional trade−off between the dimensions of the premaxillary alveolar margin and the size of the teeth. In light of recent advances in the study of tooth morphogenesis, tooth count is a function of two parameters: dimensions of an odontogenic field for a tooth series, and dimensions of tooth positions. A probable developmental cause for the low tooth count of TMP 2007.20.124 is that the dimensions of the alveoli expanded by approximately a third during tooth morphogenesis. Numerical traits such as tooth count are difficult to treat in a phylogenetic analysis. When formulating a phylogenetic character, a potential alternative to simply counting is to rely on the morphological signature for developmental parameters that control the number of the element in question.
Steroid hormones in plants and in animals are very important for physiological and developmental regulation. In animals steroid hormones are recognized by nuclear re­ceptors, which transcriptionally regulate specific target genes following binding of the ligand. In addition, numerous rapid effects generated by steroids appear to be me­diated by a mechanism not depending on the activation of nuclear receptors. Although the existence of separate membrane receptors was postulated many years ago and hundreds of reports supporting this hypothesis have been published, no animal mem­brane steroid receptor has been cloned to date. Meanwhile, a plant steroid receptor from Arabidopsis thaliana has been identified and cloned. It is a transmembrane pro­tein which specifically recognizes plant steroids (brassinosteroids) at the cell surface and has a serine/threonine protein kinase activity. It seems that plants have no intracellular steroid receptors, since there are no genes homologous to the family of animal nuclear steroid receptors in the genome of A. thaliana. Since the reason of the rapid responses to steroid hormones in animal cells still re­mains obscure we show in this article two possible explanations of this phenomenon. Using 1,25-dihydroxyvitamin D3 as an example of animal steroid hormone, we review results of our and of other groups concordant with the hypothesis of membrane steroid receptors. We also review the results of experiments performed with ovarian hormones, that led their authors to the hypothesis explaining rapid steroid actions without distinct membrane steroid receptors. Finally, examples of polypeptide growth factor that similarly to steroids exhibit a dual mode of action, activating not only cell surface receptors, but also intracellular targets, are discussed.
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