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25.3:

Polarity of the Cytoskeleton

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Cell Biology
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JoVE Central Cell Biology
Polarity of the Cytoskeleton

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Polarity in cells results from the asymmetric nature and structural orientation of the cytoskeletal filaments.

Of the three cytoskeletal filaments, microfilaments and microtubules are made up of asymmetric protein monomers.

G-Actin, the monomer of microfilaments, has a globular structure with two distinct domains. When actin polymerizes, all the subunits orient in the same direction, resulting in the filament having two ends, the plus, and the minus.

The polymerization rates at the two ends are different, further contributing to the filament's polarity.

The building blocks of microtubules are the alpha-beta-tubulin dimers. Like actin filaments, these dimers are uniformly oriented. The minus end has the free alpha-tubulins, and the plus end has free beta-tubulins.

The polarity of cytoskeletal filaments controls the directional movement of motor proteins like kinesin and dynein, carrying cargos and vesicles within the cell.

During processes like phagocytosis and cell movement, these filaments are rearranged to determine the direction of motion

25.3:

Polarity of the Cytoskeleton

The intrinsic polarity of cells can be primarily attributed to two factors- i) the asymmetric accumulation of mobile components such are regulatory molecules and subcellular components across the cell and ii) the orientation of polar cytoskeletal filaments that make up the cytoskeletal networks, specifically microfilaments, and microtubules arranged along the axis of polarity. Interactions between the cytoskeletal filaments are crucial for the establishment and maintenance of the polar nature of the cell.

Other factors that contribute to cytoskeletal polarity include the structure of monomers, the structure of individual filaments, the filament’s rate of assembly or disassembly, accessory proteins associated with cytoskeletal filaments, and the overall asymmetric distribution of these filaments within the cell. Microtubules and microfilaments are inherently polar and contribute through all these factors, whereas intermediate filaments are non-polar and contribute only through their asymmetric distribution in a cell.

Cellular Functions Based on Cytoskeleton Polarity

The polarity of cytoskeletal filaments regulates cell motility and transport of molecules within the cell. A cell moves by repeatedly extending and forming an attachment at the front, followed by detachment and retraction of the rear end. The polarity of the microfilaments directs the directionality of such protrusions. In migrating cells, F-actins dominate at the migrating front and promote the formation of lamellipodia or filopodia—membrane protrusions essential for cell movement.

Vesicular transport by motor proteins is regulated by microtubule polarity. Microtubules determine the directionality of cargoes carried by kinesin and dynein. While kinesin moves towards the cell’s periphery with the plus-end of microtubules, dynein moves towards the nucleus with the minus end.

Suggested Reading

  1. Raman, R., Pinto, C. S., & Sonawane, M. (2018). Polarized organization of the cytoskeleton: regulation by cell polarity proteins. Journal of molecular biology, 430(19), 3565-3584. https://doi.org/10.1016/j.jmb.2018.06.028
  2. Svitkina, T. (2018). The actin cytoskeleton and actin-based motility. Cold Spring Harbor perspectives in biology, 10(1), a018267. DOI: 10.1101/cshperspect.a018267.
  3. Wang, H., Robinson, R.C. and Burtnick, L.D., 2010. The structure of native G‐actin. Cytoskeleton, 67(7), pp.456-465.