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This review discusses the physical mechanisms by which lipids and proteins generate the curvature of biological membranes. It surveys the main proteins of intracellular trafficking pathways that are known to participate in membrane bending and classifies them according to their probable mechanism of action. We introduce membrane curvature and basic geometrical shapes, and show that all of the basic membrane shapes are involved in the generation of intracellular carriers. We distill the applicable theories of membrane deformation, elasticity and bending energy and introduce the effective shape and spontaneous curvature of lipids, which are factors that have an essential role in the generation of membrane curvature. Based on the physics of membrane bending, we quantitatively analyse the conditions under which membrane lipids could create the curvatures that characterize intracellular carriers, and conclude that the requirements for membrane lipid composition are stringent and unusual and that lipid-based mechanisms for the generation of curvature require that proteins provide the necessary energy. Reviewing the mechanism of tubular membrane formation by molecular motors that pull on flat membranes, we quantitatively confirm that it would take just a few molecular motors to induce the required curvature. We formulate the main mechanisms by which membrane-associated proteins can induce curvature: the scaffold mechanism and the local spontaneous curvature mechanism. We formulate the criteria that proteins or their complexes have to satisfy in order to bend membranes according to these mechanisms. Sufficiently rigid scaffold proteins can bend membranes using a membrane-adherent surface that causes the membrane to follow the contour of the protein. Proteins that insert into the lipid bilayer can bend membranes by changing the spontaneous curvature of the local monolayer. In all cases, the most probable way to generate membrane curvature is to use a combination of the scaffold and local spontaneous curvature mechanisms.
Nature Reviews Molecular Cell Biology – Springer Journals
Published: Nov 15, 2005
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