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Cytoskeleton and Cell Motility

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Encyclopedia of Complexity and Systems Science

Definition of the Subject

We, as human beings, are made of a collection of cells, which are most commonly considered as the elementary building blocks of all livingforms on earth [5]. Whether they belong to each of the three domains of life (archaea, bacteria or eukarya), cells are smallmembrane‐bounded compartments that are capable of homeostasis, metabolism, response to their environment, growth, reproduction, adaptation throughevolution and, at the cellular as well as multicellular level, organization. In addition, spontaneous, self‐generated movement – alsoknown as motility – is one of the properties that we most closely associate with all life forms. Even in thecase of apparently inanimate living forms on macroscopic scales, like most plants and fungi, constitutive cells are constantly remodeling their internalstructure for the entire organism to perform its metabolism, growth and reproduction [29]. In animalslike human beings, cell motility is at the basis of most – if not all –...

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Notes

  1. 1.

    See, for example,http://www.cytoskeletons.com/database.php.

  2. 2.

    Note that some zoologists also use the term “pseudopodia” or “pseudopods” rather generally to refer to a variety of cell‐surface protrusions. These include the different types of protrusions described here as playing a role in amoeboid motility, but also the long extended processes that some cell types use only as feeding apparatus, like axopodia.

  3. 3.

    A short video of a locomoting Amoeba proteus can be seen on the following website: http://www.bms.ed.ac.uk/research/others/smaciver/A.prot.Loc.mov.

  4. 4.

    Fibroblasts are the cells that synthesize and maintain the extracellular matrix in most animal connective tissues. They provide a structural framework (stroma) for many tissues, and play a crucial role in wound healing. Keratocytes are epithelial cells that have been characterized in the epidermis of fish and frogs, and that have been named so because of their abundant keratin filaments. They are specialized in wound healing, and are one of the most spectacular example of fast and persistent locomotion in cells, with velocities up to 30 µm/min [5,158].

  5. 5.

    Growth cones are structures that are found at the tip of axons and dendrites, by means of which neuron cells extend.

  6. 6.

    Cell‐motility videos can be seen at http://cellix.imba.oeaw.ac.at.

  7. 7.

    The persistence length L p is defined as follows: consider a thin flexible rod of fixed length L, submitted to thermal forces. Its shape is completely specified by the tangent angle \( { \theta(s) } \) in three dimensions along the arc length of the rod [154]. The persistence length L p is defined as the characteristic arc length above which thermal fluctuations of the angle \( { \theta(s) } \) become uncorrelated. Specifically, \( { \langle \cos{[\Delta\theta(s)]}\rangle = \exp{(-s/L_p)} } \), where \( { \Delta\theta(s) } \) is the three‐dimensional angle change over the arc length s. L p is related to the rod's material Young modulus E and its geometrical moment of inertia I by \( { L_p=EI/k_\text{B} T } \), where \( { k_\text{B} T } \) represents thermal energy [81].

  8. 8.

    The hydrolysis reaction of ATP (ATP \( { \rightleftharpoons {\text{ADP}} + \text{P}_i } \), where \( { \text{P}_i } \) designates inorganic phosphate) breaks a high‐energy chemical bond – here a phosphoanhydride bond – to drive many chemical reactions in the cell.

  9. 9.

    Animated movies of this process can be seen at http://www.uni-leipzig.de/%7Epwm/kas/actin/actin.html or http://cellix.imba.oeaw.ac.at/actin-polymerisation-drives-protrusion.

  10. 10.

    Illustrations of these structures can be found at http://cellix.imba.oeaw.ac.at. See also [136,257].

  11. 11.

    Similarly to ATP, GTP is a stored source of energy for the cell that is consumed via a hydrolysis reaction, here GTP \( { \rightleftharpoons {\text{GDP}} + \text{P}_i }\).

  12. 12.

    Animated movies of myosin skeletal fibers' detailed motion can be seen at http://www.scripps.edu/cb/milligan/research/movies/myosin.mov, or http://valelab.ucsf.edu.

  13. 13.

    Up-to-date information about myosin motors can be found at http://www.proweb.org.

  14. 14.

    Animated movie of kinesin's detailed motion can be seen at http://www.scripps.edu/cb/milligan/research/movies/kinesin.mov or http://valelab.ucsf.edu.

  15. 15.

    Up-to-date information about kinesin motors can be found at http://www.proweb.org.

  16. 16.

    In addition to Arp2 and Arp3, which are members of the Actin related proteins (Arp) family in that they have sequences and structures that are similar to actin , the Arp2/3 complex contains five other smaller proteins.

  17. 17.

    Up-to-date informations can be found at http://www.bms.ed.ac.uk/research/others/smaciver/Cyto-Topics/actinpage.htm.

  18. 18.

    See the movies associated with ref [215], as well as the one of a neutrophil cell that chases a bacterium at http://www.biochemweb.org/fenteany/research/cell_migration/movement_movies.html.

  19. 19.

    For an animated illustration, see http://www.jhu.edu/cmml/movies/anim/eBRatchet2.swf.

  20. 20.

    For an introduction to the elasticity of continuous media, see, e. g., [154].

  21. 21.

    An illustration of a paramecium can be seen in Fig. 1, center.

  22. 22.

    The next momentum \( { q_{\alpha\beta}=\langle u_{\alpha}u_{\beta} - d^{-1}\,\boldsymbol{p}^2\,\delta_{\alpha\beta}\rangle } \), where d is the dimension of space, is a symmetric traceless tensor of order two that corresponds to nematic order.

  23. 23.

    Note that only one relaxation time is assumed to characterize the system, as some experiments suggest that a power-law distribution of relaxation times is better suited to describe cytoskeleton dynamics, potentially because of some scale‐invariant dynamical properties in the system [18,61].

Abbreviations

Cell:

Structural and functional elementary unit of all life forms. The cell is the smallest unit that can be characterized as living.

Eukaryotic cell:

Cell that possesses a nucleus, a small membrane‐bounded compartment that contains the genetic material of the cell. Cells that lack a nucleus are called prokaryotic cells or prokaryotes.

Domains of life:

archaea, bacteria and eukarya – or in English eukaryotes, and made of eukaryotic cells – which constitute the three fundamental branches in which all life forms are classified. Archaea and bacteria are prokaryotes. All multicellular organisms are eukaryotes, but eukaryotes can also be single‐cell organisms. Eukaryotes are usually classified into four kingdoms: animals, plants, fungi and protists.

Motility:

Spontaneous, self‐generated movement of a biological system.

Cytoskeleton :

System of protein filaments crisscrossing the inner part of the cell and which, with the help of the many proteins that interact with it, enables the cell to insure its structural integrity and morphology, exert forces and produce motion.

Amoeboid motility :

Crawling locomotion of a eukaryotic cell by means of protrusion of its leading edge.

Molecular motor :

Motor of molecular size. In this context, protein or macromolecular complex that converts a specific source of energy into mechanical work.

Filament:

Here, extended unidimensional structure made of an assembly of repeated protein units that hold together via physical interactions (without covalent bonds). A filament will be either a single polymer (or here biopolymer), a linear assembly of such polymers, or a linear assembly of molecular motors .

Active gel :

Cross‐linked network of linear or branched polymers interacting by physical means, and that is dynamically driven out of equilibrium by a source of energy.

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Acknowledgments

To write this article, I grandly benefited from Karsten Kruse's habilitation thesis, which constituteda very good starting point as an extensive review of the existing biophysical literature on the cytoskeleton . I acknowledge Cécile Sykes andJean‐François Joanny for discussions, careful reading of the manuscript and constructive criticisms and suggestions. I also acknowledge AndrewCallan‐Jones for pointing to me important references.

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  1. Institut Curie, Section de Recherche, Laboratoire Physicochimie Curie (CNRS-UMR 168), Paris, France

    Thomas Risler

  2. Université Pierre et Marie Curie Paris VI, Paris, France

    Thomas Risler

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  1. RAMTECH LIMITED, 122 Escalle Lane, Larkspur, CA, 94939, USA

    Robert A. Meyers Ph. D. (Editor-in-Chief) (Editor-in-Chief)

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Risler, T. (2009). Cytoskeleton and Cell Motility. In: Meyers, R. (eds) Encyclopedia of Complexity and Systems Science. Springer, New York, NY. https://doi.org/10.1007/978-0-387-30440-3_112

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