Nontopological Saddle-Splay and Curvature Instabilities from Anisotropic Membrane Inclusions

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Abstract
Anisotropic inclusions are shown to induce spontaneous deviatoric bendings in lipidic membranes, by orienting at right angles across the bilayer. In the limit of strong membrane curvatures, a nonanalytical bending energy term is generated that favors saddlelike and cylindrical shapes, without penalizing spherical ones. An “egg-carton” instability results in flat membranes as well as a wormlike shape instability for vesicles.