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Yasushi Izumi, Kyoko Furuse, Mikio Furuse; The novel membrane protein Hoka regulates septate junction organization and stem cell homeostasis in the Drosophila gut. J Cell Sci hercules seger March ; 6 : jcs Smooth septate junctions sSJs regulate the paracellular transport in the intestinal tract in arthropods. Here, we report a novel sSJ membrane protein, Hoka, which has a single membrane-spanning segment with a short extracellular region, and a cytoplasmic region with Tyr-Thr-Pro-Ala motifs. The larval midgut in hoka mutants shows a defect in sSJ structure. Hoka forms a complex with Ssk, Mesh and Tsp2A, and is required for the hercules seger localization of these proteins to sSJs. Knockdown of hoka in the adult midgut leads to intestinal barrier dysfunction and stem cell overproliferation.
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In hoka -knockdown hercules seger, aPKC is upregulated in the cytoplasm and the apical membrane of epithelial cells. The depletion of aPKC and yki in hoka -knockdown midguts results in reduced stem cell overproliferation. Epithelia separate distinct fluid compartments within the bodies of metazoans. Hercules seger this epithelial function, occluding junctions act as barriers that control the free diffusion of solutes through the paracellular pathway. Septate junctions SJs are occluding junctions in invertebrates Furuse and Tsukita, ; Lane et al. In transmission electron microscopy, SJs are observed between the parallel heercules membranes of adjacent cells, with ladder-like septa spanning the intermembrane space Lane et al.
Despite being derived from the ectoderm, the outer epithelial layer of the proventriculus OELP and the Malpighian tubules also possess sSJs Lane et al.
Furthermore, pSJs and sSJs are distinguished by the arrangement of septa. For example, the septa of pSJs form regular undulating rows, whereas those in sSJs form regularly spaced parallel lines in the oblique sections in lanthanum-treated preparations Lane and Swales, ; Lane et al. To date, more than 20 pSJ-related proteins have been identified hercules seger characterized in Drosophila Banerjee et al.
Therefore, the mechanisms underlying sSJ organization and the functional properties of sSJs remain poorly understood compared with pSJs.
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Ssk has four membrane-spanning domains; two short extracellular loops, cytoplasmic N- and C-terminal domains, and a cytoplasmic loop Yanagihashi et al. Tsp2A is a member of the tetraspanin family of integral membrane proteins in metazoans with four transmembrane domains, N- and C-terminal short intracellular domains, two extracellular loops and one short intracellular turn Izumi et al.
Furthermore, Ssk, Mesh and Tsp2A are localized in the epithelial cell-cell contact regions in the Drosophila Malpighian tubules in which sSJs are present Tepass and Hartenstein, ; Yanagihashi et al. Recent studies have shown that the knockdown of mesh and Tsp2A in the epithelium of Malpighian tubules leads to defects in hercules seger morphogenesis, tubule transepithelial fluid and ion transport, and paracellular hercules seger permeability in the tubules Jonusaite et al. Thus, sSJ proteins are involved in the development and maintenance of functional Malpighian tubules in Drosophila. These results indicate that sSJs play a crucial role in maintaining tissue homeostasis through the regulation of stem cell proliferation and enterocyte behavior in the Drosophila adult midgut.
Furthermore, Chen et al. Thus, sSJs may contribute to hercules seger establishment of epithelial polarity in the adult midgut. During the regeneration of the Drosophila adult midgut epithelium, various signaling pathways are involved in the proliferation and differentiation of the ISC lineage Jiang et al. Atypical protein kinase C aPKC is an evolutionarily conserved key determinant of apical-basal epithelial polarity Ohno et al.
Importantly, Chen et al. Goulas et al.]