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

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