Flat and sigmoidally curved contact zones in vesicle-vesicle adhesion

Using the membrane-bending elasticity theory and a simple effective model of adhesion, we study the morphology of lipid vesicle doublets. In the weak adhesion regime, we find flat-contact axisymmetric doublets, whereas at large adhesion strengths, the vesicle aggregates are nonaxisymmetric and characterized by a sigmoidally curved, S-shaped contact zone with a single invagination and a complementary evagination on each vesicle. The sigmoid-contact doublets agree very well with the experimentally observed shapes of erythrocyte aggregates. Our results show that in identical vesicles with large to moderate surface-to-volume ratio, the sigmoid-contact shape is the only bound morphology. We also discuss the role of sigmoid contacts in the formation of multicellular aggregates such as erythrocyte rouleaux.