Alternative pathways control actomyosin contractility in epitheliomuscle cells during morphogenesis and body contraction.

In adult Hydra, epitheliomuscle cells form the monolayered ecto- and endodermal epithelia. Their basal myonemes function as a longitudinal and circular muscle, respectively. Based on the observation that a Rho/Rock pathway, controlling the cell shape changes during detachment of Hydra buds, is not involved in body movement, at least two actomyosin compartments must exist in these cells: a basal one for body movement and a cortical one for cell shape changes. We therefore analyzed the regional and subcellular localization of the Ser19-phosphorylated myosin regulatory light chain (pMLC20). Along the body column, pMLC20 was detected strongly in the basal myonemes and weakly in the apical cell compartments of ectodermal epitheliomuscle cells. In cells of the bud base undergoing morphogenesis, pMLC20 was localized to intracellular stress fibers as well as to the apical and additionally to the lateral cortical compartment. Pharmacological inhibition revealed that pMLC20 is induced in these compartments by at least two independent pathways. In myonemes, MLC is phosphorylated mainly by myosin light chain kinase (MLCK). In contrast, the cortical apical and lateral MLC phosphorylation in constricting ectodermal cells of the bud base is stimulated via the Rho/ROCK pathway.

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