Morphology of small aggregates of red blood cells

Blood can be considered a two-phase liquid composed of plasma as well as cells and cell aggregates. The degree of cell aggregation is an important determinant of blood rheology: The size and shape of the aggregates affect blood viscosity. The microscopic mechanisms of red blood cell adhesion involve a complex interplay of electrostatic, van der Waals, and a range of specific biochemical inter-membrane interactions. Here we use an effective model of these interactions combined with the membrane elasticity theory to calculate the equilibrium shape of a red blood cell doublet and compare it with the experimentally observed red blood cell aggregates both in vitro and in vivo. Special attention is devoted to the shape of doublets formed by dissimilar cells. A possible effect of doublet shape on pathways of the formation of multicellular aggregates is discussed. Red blood cell rouleau formation is expected to take place at intermediate adhesion strengths where the outer doublet surfaces are either concave or flat, whereas in the strong-adhesion regime where the outer doublet surfaces are convex the cells should form rounded clump-like aggregates.