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Physiological Reviews, Vol. 81, No. 2, April 2001, pp. 685-740
Copyright ©2001 by the American Physiological Society
Physiology and Biophysics, University Of Texas Medical Branch, Galveston, Texas; and Department of Pharmacology, University Of Western Australia, Nedlands, West Australia
Hamill, Owen P. and
Boris Martinac.
Molecular Basis of Mechanotransduction in Living Cells. Physiol. Rev. 81: 685-740, 2001.
The simplest cell-like structure, the lipid
bilayer vesicle, can respond to mechanical deformation by elastic
membrane dilation/thinning and curvature changes. When a protein is
inserted in the lipid bilayer, an energetic cost may arise because of
hydrophobic mismatch between the protein and bilayer. Localized changes
in bilayer thickness and curvature may compensate for this mismatch.
The peptides alamethicin and gramicidin and the bacterial membrane protein MscL form mechanically gated (MG) channels when inserted in
lipid bilayers. Their mechanosensitivity may arise because channel
opening is associated with a change in the protein's
membrane-occupied area, its hydrophobic mismatch with the bilayer,
excluded water volume, or a combination of these effects. As a
consequence, bilayer dilation/thinning or changes in local membrane
curvature may shift the equilibrium between channel conformations.
Recent evidence indicates that MG channels in specific animal cell
types (e.g., Xenopus oocytes) are also gated directly by
bilayer tension. However, animal cells lack the rigid cell wall that
protects bacteria and plants cells from excessive expansion of their
bilayer. Instead, a cortical cytoskeleton (CSK) provides a structural
framework that allows the animal cell to maintain a stable excess
membrane area (i.e., for its volume occupied by a sphere) in the form
of membrane folds, ruffles, and microvilli. This excess membrane provides an immediate membrane reserve that may protect the bilayer from sudden changes in bilayer tension. Contractile elements within the
CSK may locally slacken or tighten bilayer tension to regulate mechanosensitivity, whereas membrane blebbing and tight seal patch formation, by using up membrane reserves, may increase membrane mechanosensitivity. In specific cases, extracellular and/or CSK proteins (i.e., tethers) may transmit mechanical forces to the process
(e.g., hair cell MG channels, MS intracellular Ca2+
release, and transmitter release) without increasing tension in the
lipid bilayer.
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