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Department of Biology, University of Copenhagen, Copenhagen, Denmark
The ability to control cell volume is pivotal for cell function. Cell volume perturbation elicits a wide array of signaling events, leading to protective (e.g., cytoskeletal rearrangement) and adaptive (e.g., altered expression of osmolyte transporters and heat shock proteins) measures and, in most cases, activation of volume regulatory osmolyte transport. After acute swelling, cell volume is regulated by the process of regulatory volume decrease (RVD), which involves the activation of KCl cotransport and of channels mediating K+, Cl–, and taurine efflux. Conversely, after acute shrinkage, cell volume is regulated by the process of regulatory volume increase (RVI), which is mediated primarily by Na+/H+ exchange, Na+-K+-2Cl– cotransport, and Na+ channels. Here, we review in detail the current knowledge regarding the molecular identity of these transport pathways and their regulation by, e.g., membrane deformation, ionic strength, Ca2+, protein kinases and phosphatases, cytoskeletal elements, GTP binding proteins, lipid mediators, and reactive oxygen species, upon changes in cell volume. We also discuss the nature of the upstream elements in volume sensing in vertebrate organisms. Importantly, cell volume impacts on a wide array of physiological processes, including transepithelial transport; cell migration, proliferation, and death; and changes in cell volume function as specific signals regulating these processes. A discussion of this issue concludes the review.
2 Where relevant, the regulation of specific transporters by the cytoskeleton is discussed in more detail in section VI and VII.
3 An additional gene product was assigned the name SLC9A11 (http://www.gene.ucl.ac.uk/cgi-bin/nomenclature/), but is, to our knowledge, not yet proven to be a functional Na+/H+ exchanger.
4 In the choroid plexus epithelium and brain microvascular endothelial cells, NKCC1 has been found to localize to the apical membrane (735, 819).
5 Alternatively spliced forms exist for both isoforms, yet will not be further discussed here (see Ref. 265).
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