Nontopological Saddle-Splay and Curvature Instabilities from Anisotropic Membrane Inclusions

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.