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WIREs Nanomed Nanobiotechnol
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Microtubule‐based force generation

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Microtubules are vital to many important cell processes, such as cell division, transport of cellular cargo, organelle positioning, and cell migration. Owing to their diverse functions, understanding microtubule function is an important part of cell biological research that can help in combating various diseases. For example, microtubules are an important target of chemotherapeutic drugs such as paclitaxel because of their pivotal role in cell division. Many functions of microtubules relate to the generation of mechanical forces. These forces are generally either a direct result of microtubule polymerization/depolymerization or generated by motor proteins that move processively along microtubules. In this review, we summarize recent efforts to quantify and model force generation by microtubules in the context of microtubule function. WIREs Nanomed Nanobiotechnol 2017, 9:e1428. doi: 10.1002/wnan.1428 This article is categorized under: Nanotechnology Approaches to Biology > Cells at the Nanoscale
(a) Force production by polymerization of a microtubule against a boundary. Insertion of new αβ‐tubulin subunits at the plus end exerts a pushing force on the boundary (brown). If the boundary resists movement, this force can cause flux of the microtubule toward the minus end. (b) One possible mechanism for the conformational wave model of microtubule depolymerization exerting a pulling force to separate chromosomes (purple) during mitosis. The kinetochore (yellow oval) is attached to a ring‐shaped complex (yellow annulus) via protein tethers (black lines). As the microtubule depolymerizes, the curling of tubulin protofilaments performs a ‘power stroke’ on the ring‐shaped complex, and a pulling force is exerted on the kinetochore. (c) An alternative mechanism to (b) known as the biased diffusion model. In this model, kinetochore proteins (black lines) interact transiently with a microtubule. As the microtubule depolymerizes, motion of the kinetochore toward the microtubule tip is energetically favored, because when the kinetochore is closer to the microtubule tip, more kinetochore proteins are able to bind to the microtubule lattice. (d) Forces generated by microtubule motors. Cytoplasmic dynein and kinesin‐1 are shown. These are, respectively, the primary minus end‐directed and plus end‐directed motors that transport cargo within the cell. When a motor is anchored at its cargo end, for example to other elements of the cytoskeleton (red lines), the motor exerts a force on the microtubule in the opposite direction of its processivity.
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