State diagram for wall adhesion of red blood cells in shear flow: from crawling to flipping.

Red blood cells in shear flow show a variety of different shapes due to the complex interplay between hydrodynamics and membrane elasticity. Malaria-infected red blood cells become generally adhesive and less deformable. Adhesion to a substrate leads to a reduction in shape variability and to a flipping motion of the non-spherical shapes during the mid-stage of infection. Here, we present a complete state diagram for wall adhesion of red blood cells in shear flow obtained by simulations, using a particle-based mesoscale hydrodynamics approach, multiparticle collision dynamics. We find that cell flipping at a substrate is replaced by crawling beyond a critical shear rate, which increases with both membrane stiffness and viscosity contrast between the cytosol and suspending medium. This change in cell dynamics resembles the transition between tumbling and tank-treading for red blood cells in free shear flow. In the context of malaria infections, the flipping-crawling transition would strongly increase the adhesive interactions with the vascular endothelium, but might be suppressed by the combined effect of increased elasticity and viscosity contrast.