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                                                          Membrane Structure






              Anna Seelig
              Joachim Seelig
              University of Basel



                I. Introduction
               II. Membrane Lipids
              III. The Membrane–Water Interface
              IV. Hydrophobic Core Region
               V. Phase Behavior of Lipids and Membrane
                 Domain Formation
              VI. Interaction of Membrane Lipids with Amphiphilic
                 Molecules and Transmembrane Proteins
              VII. Concluding Remarks








              GLOSSARY                                            fast molecular motions such as rotational and flexing
                                                                  movements are thus characterized, on the average, by
              Biological membrane A very thin sheath of biological  a cylindrical symmetry with the bilayer normal as the
                material (thickness ∼10 nm to 15 nm) which consti-  axis of motional averaging.
                                                                                                  2
                tutes the envelope of living cells and also of intracellu-  Deuterium nuclear magnetic resonance ( H-NMR) By
                lar organelles, separating them from the environment.  means of chemical synthesis or biochemical incorpo-
                Membranes are made up from a lipid bilayer into which  ration, protons in lipid molecules can be selectively
                proteins are embedded. They are highly organized but  replaced by deuterium atoms. Since the van der Waals
                are nevertheless fluid enough to allow considerable  radii of the two isotopes are identical, this substitution
                translational, rotational, and flexing movements of the  leaves the membrane virtually unchanged, which is in
                constituent lipid and protein molecules.          contrasttootherbulkierreportergroups.Deuteriumnu-
              Lipid bilayer A double layer of lipid molecules orga-  clear magnetic resonance provides information on the
                nized in a tail-to-tail arrangement. It is an anisotropi-  order and mobility of the molecule. Structural informa-
                cally ordered fluid that has a number of properties in  tion comes from the deuterium quadrupole splitting,
                common with smectic liquid crystals. The normal to the  dynamic information is derived from NMR relaxation
                surface of the lipid bilayer constitutes an axis of mo-  times.
                tional averaging. From an optical point of view, bilayer  Quadrupolar splitting (∆ν Q ) The deuterium nucleus
                membranes thus behave like uniaxial crystals with the  has a spin = 1 and, due to its electric quadrupole mo-
                bilayer normal as the optical axis (director axis  n). All  ment, the anisotropic motion within the membrane will



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