Lipid domain structures in biological membranes.

Phase separation represents a possibility for segregation of lipidic membrane components into structurally distinct domains. Freeze-fracture electronmicroscopy is a useful method for detection of lipid domains. Indications of a possible domain-nature of structures are a regular pattern within a separated area, a regular outline of such an area and a local modulation of curvature (evagination or invagination). Candidates for domain structures in biological membranes are smooth particle-free areas and arrays of regularly arranged particles. The interpretation of the particle-free areas is more reliable than that of the arrays with regularly arranged particles. Phase separation in biological membranes can be induced experimentally by lowering the temperature, but physiologically the isothermically induced domains are more important. Factors in control of isothermic domain formation are divalent cations, proteins, cholesterol etc. Suggestions on the biological relevance of domain formation concern mainly their role in the mechanism of membrane fusion, but domains in form of transient or stable membrane structures seem to occur also otherwise and disturbances in domain formation or artificially induced domains can be suitable for pathological alterations.