Extensional flow of erythrocyte membrane from cell body to elastic tether. I. Analysis

This is the first of two papers on an analytical and experimental study of the flow of the erythrocyte membrane. In the experiment to be discussed in detail in the second paper, preswollen human erythrocytes are sphered by aspirating a portion of the cell membrane into a small micropipette; and long, thin, membrane filaments or "tethers" are steadily withdrawn from the cell at a point diametrically opposite to the point of aspiration. The aspirated portion of the membrane furnished a "reservoir" of material that replaces the membrane as it flows as a liquid from the nearly spherical cell body to the cylindrical tether. In this paper we show that an application of the principle of conservation of mass permits the tether radius (approximately 200 A or less) to be measured with the light microscope as the tether is formed and extended at a constant rate. A static analysis of the axisymmetric cell deformation and tether formation process reveals that the tether radius is uniquely determined by the isotropic tension in the membrane and the elastic constitutive (material) behavior of the tether itself. A dynamic analysis of the extensional flow process reveals that the tether radius must decrease as the velocity of the tether is increased and that the decrease depends on both the viscosity of the membrane and the elasticity of the tether. The analysis also shows that these two factors (membrane viscosity and tether elasticity) are readily decomposed and determined separately when flow experiments are performed at different isotropic tensions.