Microtubule-organizing centers and nucleating sites in land plants.

Microtubule-organizing centers (MTOCs) are morphologically diverse cellular sites involved in the nucleation and organization of microtubules (MTs). These structures are synonymous with the centrosome in mammalian cells. In most land plant cells, however, no such structures are observed and some have argued that plant cells may not have MTOCs. This review summarizes a number of experimental approaches toward the elucidation of those subcellular sites involved in microtubule nucleation and organization. In lower land plants, structurally well-defined MTOCs are present, such as the blepharoplast, multilayered structure, and polar organizer. In higher plants, much of the nucleation and organization of MTs occurs on the nuclear envelope or other endomembranes, such as the plasmalemma and smooth (tubular) endoplasmic reticulum. In some instances, one endomembrane may serve as a site of nucleation whereas others serve as the site of organization. Structural and motor microtubule-associated proteins also appear to be involved in MT nucleation and organization. Immunochemical evidence indicates that at least several of the proteins found in mammalian centrosomes, gamma-tubulin, centrin, pericentrin, and polypeptides recognized by the monoclonal antibodies MPM-2, 6C6, and C9 also recognize putative lower land plant MTOCs, indicating shared mechanisms of nucleation/organization in plants and animals. The most recent data from tubulin incorporation in vivo, mutants with altered MT organization, and molecular studies indicate the potential of these research tools in investigation of MTOCs in plants.