Microtubule nucleation for the assembly of acentrosomal microtubule arrays in plant cells.

Contents Summary I. Introduction II. MT arrays in plant cells III. γ-Tubulin and MT nucleation IV. MT nucleation sites or flexible MTOCs in plant cells V. MT-dependent MT nucleation VI. Generating new MTs for spindle assembly VII. Generation of MTs for phragmoplast expansion during cytokinesis VIII. MT generation for the cortical MT array IX. MT nucleation: looking forward Acknowledgements References SUMMARY: Cytoskeletal microtubules (MTs) have a multitude of functions including intracellular distribution of molecules and organelles, cell morphogenesis, as well as segregation of the genetic material and separation of the cytoplasm during cell division among eukaryotic organisms. In response to internal and external cues, eukaryotic cells remodel their MT network in a regulated manner in order to assemble physiologically important arrays for cell growth, cell proliferation, or for cells to cope with biotic or abiotic stresses. Nucleation of new MTs is a critical step for MT remodeling. Although many key factors contributing to MT nucleation and organization are well conserved in different kingdoms, the centrosome, representing the most prominent microtubule organizing centers (MTOCs), disappeared during plant evolution as angiosperms lack the structure. Instead, flexible MTOCs may emerge on the plasma membrane, the nuclear envelope, and even organelles depending on types of cells and organisms and/or physiological conditions. MT-dependent MT nucleation is particularly noticeable in plant cells because it accounts for the primary source of MT generation for assembling spindle, phragmoplast, and cortical arrays when the γ-tubulin ring complex is anchored and activated by the augmin complex. It is intriguing what proteins are associated with plant-specific MTOCs and how plant cells activate or inactivate MT nucleation activities in spatiotemporally regulated manners.