System of microscopic filaments or fibres, present in the cytoplasm of eukaryotic cells (see eukaryote), that organizes other cell components, maintains cell shape, and is responsible for cell locomotion and for movement of the organelles within it. Three major types of filaments make up the cytoskeleton: actin filaments, microtubules, and intermediate filaments. Actin filaments occur as constantly changing bundles of parallel fibres; they help determine cell shape, help the cell adhere to surfaces, help the cell move, and assist in cell division during mitosis. Intermediate filaments are very stable structures that form the cell's true skeleton; they anchor the nucleus within the cell and give the cell its elastic properties.
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Eukaryotic cells contain three main kinds of cytoskeletal filaments, which are microfilaments, intermediate filaments, and microtubules. The cytoskeleton provides the cell with structure and shape, and by excluding macromolecules from some of the cytosol it adds to the level of macromolecular crowding in this compartment.
These filaments, 8 to 12 nanometers in diameter, are more stable (strongly bound) than actin filaments, and heterogeneous constituents of the cytoskeleton. Like actin filaments, they function in the maintenance of cell-shape by bearing tension (microtubules, by contrast, resist compression. It may be useful to think of micro- and intermediate filaments as cables, and of microtubules as cellular support beams). Intermediate filaments organize the internal tridimensional structure of the cell, anchoring organelles and serving as structural components of the nuclear lamina and sarcomeres. They also participate in some cell-cell and cell-matrix junctions.
Different intermediate filaments are:
Microtubules are hollow cylinders about 25 nm in diameter (lumen = approximately 15nm in diameter), most commonly comprised of 13 protofilaments which, in turn, are polymers of alpha and beta tubulin. They have a very dynamic behaviour, binding GTP for polymerization. They are commonly organized by the centrosome.
In nine triplet sets (star-shaped), they form the centrioles, and in nine doublets oriented about two additional microtubules (wheel-shaped) they form cilia and flagella. The latter formation is commonly referred to as a "9+2" arrangement, wherein each doublet is connected to another by the protein dynein. As both flagella and cilia are structural components of the cell, and are maintained by microtubules, they can be considered part of the cytoskeleton.
They play key roles in:
|Cytoskeleton type||Diameter (nm)||Structure||Subunit examples|
|Intermediate filaments||8-10||two parallel helices/dimers, forming tetramers|
|Microtubules||25||protofilaments, in turn consisting of tubulin subunits||α- and β-tubulin|
A fourth eukaryotic cytoskeletal element, microtrabeculae, was proposed by Keith Porter based on images obtained from high-voltage electron microscopy of whole cells in the 1970s. The images showed short, filamentous structures of unknown molecular composition associated with known cytoplasmic structures. Porter proposed that this microtrabecular structure represented a novel filamentous network distinct from microtubules, filamentous actin, or intermediate filaments. It is now generally accepted that microtrabeculae are nothing more that an artefact of certain types of fixation treatment though we have yet to fully understand the complexity of the cell's cytoskeleton.
Some plasmids encode a partitioning system that involves an actin-like protein ParM. Filaments of ParM exhibit dynamic instability, and may partition plasmid DNA into the dividing daughter cells by a mechanism analogous to that used by microtubules during eukaryotic mitosis.
Design and evaluation of Actichip, a thematic microarray for the study of the actin cytoskeleton.(Methodology article)
Aug 29, 2007; Authors: Jean Muller [1,2,4]; André Mehlen ; Guillaume Vetter [1,3]; Mikalai Yatskou ; Arnaud Muller ; Frédéric Chalmel...