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In cell biology, microtubule nucleation is the event that initiates de novo formation of microtubules (MTs). These filaments of the cytoskeleton typically form through polymerization of α- and β-tubulin dimers, the basic building blocks of the microtubule, which initially interact to nucleate a seed from which the filament elongates.
Because nucleation from the centrosome is inherently symmetrical, Golgi-associated microtubule nucleation may allow the cell to establish asymmetry in the microtubule network. In recent studies, the Vale group at UCSF identified the protein complex augmin as a critical factor for centrosome-dependent, spindle-based microtubule generation.
More specifically, TPX2 contributes to microtubule branching during spindle assembly by cooperating with augmin in order to amplify microtubule mass and preserve its polarity. [22] Branching nucleation by TPX2 is observed without RanGTP, though more fan-shaped microtubule structures are formed when both RanGTP and TPX2 are present. [22]
The microtubule-organizing center (MTOC) is a structure found in eukaryotic cells from which microtubules emerge. MTOCs have two main functions: the organization of eukaryotic flagella and cilia and the organization of the mitotic and meiotic spindle apparatus , which separate the chromosomes during cell division .
The gradient triggers release of spindle assembly factors (SAFs) from inhibitory interactions via the transport proteins importin β/α. The unbound SAFs then promote microtubule nucleation and stabilization around mitotic chromatin, and spindle bipolarity is organized by microtubule motor proteins. [27]
Genes from the alpha, beta and gamma tubulin families are found in all eukaryotes. The alpha and beta tubulins represent the major components of microtubules, while gamma tubulin plays a critical role in the nucleation of microtubule assembly. There are multiple alpha and beta tubulin genes, which are highly conserved among species.
The growing ends of microtubules are shown in green (labeled with green fluorescent protein fused to the microtubule plus end binding protein EB1 of Arabidopsis thaliana). N = Nucleus, V = Vacuole, PPB = Preprophase band, MTN = Microtubule nucleation starts at the nuclear envelope, NEB = Nuclear envelope breakdown at the onset of prometaphase.
γ-tubulin is the best understood mechanism of microtubule nucleation, but certain studies have indicated that certain cells may be able to adapt to its absence, as indicated by mutation and RNAi studies that have inhibited its correct expression. Besides forming a γ-TuRC to nucleate and organize microtubules, γ-tubulin can polymerize into ...