The shapes of red blood cells with fluid interior depend on the properties of their membranes. The red blood cell membrane is composed of a bilayer, containing integral proteins, and the underlying membrane skeleton, which are attached to each other through different specific and nonspecific linkages. A short account is given of theories of shape formation based on the description of the membrane as a single- or multi-layered elastic continuum. Existing experimental evidence is then outlined which indicates that some relevant shape determining factors reside in the specific properties of different red blood cell integral and skeletal proteins. In this context a theoretical approach is introduced, based on the chemical equilibria established by the interacting skeletal and integral proteins. A simple prototype model is analyzed, taking into consideration that the binding between integral membrane proteins and the skeleton depends on membrane curvature. The essential parameters of the model are the gross value of the corresponding interaction energy and the coefficient measuring the dependence of this energy on membrane curvature. The effects of these parameters on RBC shapes and on the lateral mobility of integral membrane proteins are demonstrated.
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