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The composite cytoskeleton, comprising interacting networks of semiflexibleactin and rigid microtubules, actively generates forces and restructures usingmotor proteins such as myosins to enable key mechanical processes includingcell motility and mitosis. Yet, how motor-driven activity alters the mechanicsof cytoskeleton composites remains an open challenge. Here, we perform opticaltweezers microrheology on actin-microtubule composites driven by myosin IImotors to show that motor activity increases the linear viscoelasticity andelastic storage of the composite by active restructuring to a network oftightly-packed filament clusters and bundles. Our nonlinear microrheologymeasurements performed hours after cessation of activity show that themotor-contracted structure is stable and robust to nonlinear forcing. Uniquefeatures of the nonlinear response include increased mechanical stiffness,memory and heterogeneity, coupled with suppressed filament bending followingmotor-driven restructuring. Our results shed important new light onto theinterplay between viscoelasticity and non-equilibrium dynamics in activepolymer composites such as the cytoskeleton.

Active restructuring of cytoskeleton composites leads to increased mechanical stiffness, memory, and heterogeneity