“Second extrinsic organizational mechanism” for orienting cellulose: modeling a role for the plasmalemmal reticulum

Summary. Oriented deposition of cellulose fibers by cellulose-synthesizing complexes typically occurs across the plasma membrane from microtubule bundles and is guided by them. However, aligned movement of the complexes can be shown even after applied oryzalin has depolymerized microtubules. Further, there is a claim that when (1) microtubules are depolymerized with oryzalin, (2) a microtubule-orienting stimulus is applied temporarily, and (3) oryzalin is washed out, the newly forming cellulose fibers are oriented with respect to the stimulus. With this in mind, the present paper gathers evidence from a diverse literature to suggest that the plasmalemmal reticulum, a major and structurally important form of cytoskeleton which connects cortical cytoplasm with wall, is a candidate to both independently and cooperatively participate in orienting microtubules and routing movements of cellulose-synthesizing complexes. Critical to this proposed function, the adhesion sites of the plasmalemmal reticulum have some morphological and molecular similarities to animal cell adhesion sites, known to play numerous integrative roles. The reticulum itself may be the morphological manifestation of the so-called lipid raft, previously known only on the basis of biochemical properties. According to the working model, the trusses interconnecting the adhesion sites shape the reticulum into apparently situation-dependent geometries. For example, in nongrowing or nonpolarized cells in which cellulose is deposited in brushy meshes, they form a nonpolar or weakly polar net; however, in elongating cells with oblique or otherwise polarized microtubules and newly forming cellulose fibers, there is suggestive evidence that net formation is dominated by trusses organized with correspondingly biased orientation. Consideration of such geometries and roles of the reticulum suggests several tests that could affirm, deny, or replace key aspects of this proposal to expand the theory of the peripheral cytoskeleton.