Cytoskeleton Force Exertion in Bulk Cytoplasm

The microtubule and actin cytoskeletons generate forces essential to position centrosomes, nuclei, and spindles for division plane specification. While the largest body of work has documented force exertion at, or close to the cell surface, mounting evidence suggests that cytoskeletal polymers can also produce significant forces directly from within the cytoplasm. Molecular motors such as kinesin or dynein may for instance displace cargos and endomembranes in the viscous cytoplasm yielding friction forces that pull or push microtubules. Similarly, the dynamics of bulk actin assembly/disassembly or myosin-dependent contractions produce cytoplasmic forces which influence the spatial organization of cells in a variety of processes. We here review the molecular and physical mechanisms supporting bulk cytoplasmic force generation by the cytoskeleton, their limits and relevance to organelle positioning, with a particular focus on cell division.