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Molecular motors and cytoskeletal filaments work collectively most of thetime under opposing forces. This opposing force may be due to cargo carried bymotors or resistance coming from the cell membrane pressing against thecytoskeletal filaments. Some recent studies have shown that the collectivemaximum force (stall force) generated by multiple cytoskeletal filaments ormolecular motors may not always be just a simple sum of the stall forces of theindividual filaments or motors. To understand this excess or deficit in thecollective force, we study a broad class of models of both cytoskeletalfilaments and molecular motors. We argue that the stall force generated by agroup of filaments or motors is additive, that is, the stall force of $N$number of filaments (motors) is $N$ times the stall force of one filament(motor), when the system is in equilibrium at stall. Conversely, we show thatthis additive property typically does not hold true when the system is not atequilibrium at stall. We thus present a novel and unified understanding of theexisting models exhibiting such non-addivity, and generalise our arguments bydeveloping new models that demonstrate this phenomena. We also propose aquantity similar to thermodynamic efficiency to easily predict this deviationfrom stall-force additivity for filament and motor collectives.

Sufficient conditions for the additivity of stall forces generated by multiple filaments or motors