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Microtubule organizing center


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 MTOC is a major site of microtubule nucleation and can be visualized in cells by detection of γ-tubulin. The morphological characteristics of MTOCs vary between the different phyla and kingdoms. In animals, the two most important types of MTOCs are the basal bodies associated with cilia and the centrosome associated with spindle formation.

Microtubule-organizing centers function as the site where microtubule formation begins, as well as a location where free-ends of microtubules attract to. Within the cells, microtubule-organizing centers can take on many different forms. An array of microtubules can arrange themselves in a pinwheel structure to form the basal bodies, which can lead to the formation of microtubule arrays in the cytoplasm or the 9+2 axoneme. Other arrangements range from fungi spindle pole bodies to the eukaryotic chromosomal (flat, laminated plaques). MTOCs can be freely dispersed throughout the cytoplasm or centrally localized as foci. The most notable MTOCs are the centrosome at interphase and the mitotic spindle poles.

Centrioles can act as markers for MTOCs in the cell. If they are freely distributed in the cytoplasm, centrioles can gather during differentiation to become MTOCs. They can also be focused around a centrosome as a single MTOC, though centrosomes can work as an MTOC absent of centrioles.

Most animal cells have one MTOC during interphase, usually located near the nucleus, and generally associated closely with the Golgi apparatus. The MTOC is made up of a pair of centrioles at its center, and is surrounded by pericentriolar material (PCM) that is important for microtubule nucleation. Microtubules are anchored at the MTOC by their minus ends, while their plus ends continue to grow into the cell periphery. The polarity of the microtubules is important for cellular transport, as the motor proteins kinesin and dynein typically move preferentially in the "plus" and "minus" directions respectively, along a microtubule, allowing vesicles to be directed to or from the endoplasmic reticulum and Golgi apparatus. Particularly for the Golgi apparatus, structures associated with the apparatus travel towards the minus end of a microtubule and aid in the overall structure and site of the Golgi in the cell.


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