Stratification, multivalency and turnover of the active cortical machinery are required for steady active contractile flows at the cell surface

Many cell membrane proteins that bind to actin form dynamic clusters driven by contractile flows generated by the actomyosin machinery at the cell cortex. Recent evidence suggests that a necessary condition for the generation of these protein clusters on the membrane is the stratified organization of the active agents -formin-nucleated actin, myosin-II minifilaments, and ARP2/3-nucleated actin mesh -within the cortex. Further, the observation that these clusters dynamically remodel, requires that the components of this active machinery undergo turnover. Here we develop a coarse-grained agent-based Brownian dynamics simulation that incorporates the effects of stratification, binding of myosin minifilaments to multiple actin filaments and their turnover. We show that these three features of the active cortical machinery -stratification, multivalency and turnover -are critical for the realisation of a nonequilibrium steady state characterised by contractile flows and dynamic orientational patterning. We show that this nonequilibrium steady state enabled by the above features of the cortex, can facilitate multi-particle encounters of membrane proteins that profoundly influence the kinetics of bimolecular reactions at the cell surface.