Hyphae of the Ascomycota are tubular cells compartmentalized by perforated septa, whose central pore allows the flow of organelles and cytoplasm. While in plants and yeast septation leads to cell separation, in filamentous fungi the formation of crosswalls appears to have an architectural role, limits the extent of mechanical damage thus maintaining hyphal integrity, and also is of fundamental importance as part of cell differentiation. The increasing number of available fungal genome sequences, knockout mutants, versatile tools for protein tagging, and the continuous improvement of fluorescence microscopes have allowed scientists to analyze living cells and reveal the molecular and cellular basis of septation with unprecedented detail. This review summarizes the recent advances in septum ontogenesis in Neurospora crassa. A "septal actomyosin tangle" is the first indication of impending septation. It assembles prior to any visible evidence of plasma membrane inward growth, which occurs concomitantly with the formation and constriction of a contractile actomyosin ring and synthesis of the septum wall. One of the key questions in septum biogenesis is how the septation machinery is assembled to construct a centripetally growing crosswall. Most of the machinery utilized in apical cell wall growth can be expected at septation sites to ensure an organized arrival and supply of vesicles leading to the formation of a septum. Yet, the intrinsically different architecture of the septum may require a different organization and regulation of the wall-synthesizing machinery.
Keywords: Actin; Cell wall; Morphogenesis; Neurospora crassa; Septal pore; Septation.
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