Integration and regulation of hyphal tip growth

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Abstract
Hyphal tip growth is an exquisitely controlled process that forms developmentally regulated, species-specific, even-diameter tubes at rates of up to about 50 μm/min. The traditional view is that this process results from the balance between the expansive force of turgor pressure and the controlled extensibility of the apical cell wall. While these elements are involved, the model places regulation into either the global domain (turgor pressure) or the extracellular environment (the cell wall), neither of which seem well suited to the level of control evinced. Recent evidence suggests that F-actin-rich elements of the cytoskeleton are important in tip morphogenesis. Our current models propose that tip expansion is regulated (restrained under normal turgor pressure and protruded under low turgor) by a peripheral network of F-actin that is attached to the plasmalemma and the cell wall by integrin-containing linkages, thus placing control in the cytoplasm where it is accessible to normal intracellular regulatory systems. The F-actin system also functions in cytoplasmic and organelle motility; control of plasmalemma-located, stretch-activated, Ca2+-transporting, ion channel distribution; vectoral vesicle transport; and exocytosis. Regulation of the system may involve Ca2+, the concentration of which is influenced by the tip-high gradient of the stretch-activated channels, thus suggesting a possible feedback regulation mechanism. Key words: tip growth, fungi, stretch-activated channels, F-actin, Ca2+, hyphae.