The role of the cytoskeleton in the morphogenesis and function of stomatal complexes

Microtubules (MTs) and actin filaments (AFs) form highly organized arrays in stomatal cells that play key roles in the morphogenesis of stomatal complexes. The cortical MTs controlling the orientation of the depositing cellulose microfibrils (CMs) and affecting the pattern of local wall thickenings define the mechanical properties of the walls of stomatal cells, thus regulating accurately their shape. Besides, they are involved in determination of the cell division plane. Substomatal cavity and stomatal pore formation are also MT-dependent processes. Among the cortical MT arrays, the radial ones lining the periclinal walls are of particular morphogenetic importance. Putative MT organizing centers (MTOCs) function at their focal regions, at least in guard cells (GCs), or alternatively, these regions either organize or nucleate cortical MTs. AFs are involved in cell polarization preceding asymmetrical divisions, in determination of the cell division plane and final cell plate alignment and probably in transduction of stimuli implicated in stomatal complex morphogenesis. Mature kidney-shaped GCs display radial AF arrays, undergoing definite organization cycles during stomatal movement. They are involved in stomatal movement, probably by controlling plasmalemma ion-channel activities. Radial MT arrays also persist in mature GCs, but a role in stomatal function cannot yet be attributed to them. Contents Summary 613 I. Introduction 614 II. Cytoskeleton and development of the stomatal complexes 614 III. Cytoskeleton and stomatal cell shaping 620 IV. Stomatal pore formation 624 V. Substomatal cavity formation 625 VI. Stomatal complex morphogenesis in mutants 626 VII. Cytoskeleton dynamics in functioning stomata 628 VIII. Mechanisms of microtubule organization in stomatal cells 631 IX. Conclusions-perspectives 634 References 635.