Distribution of active forces in the cell cortex

In this work, we study in detail the distribution of stochastic forces generated by the molecular motors activity, in the actin cortex of pre-muscular cells. By combining active and passive rheology experiments, performed on the same micro-bead bound to the actin network through membrane adhesive receptors, we measure the auto-correlation function C_ff (t) of the average force pulling on the bead. As for any out-of-equilibrium system, the force distribution differs from the thermodynamical equilibrium one, especially at long time scale t ~ 1 s where the bead motion becomes partially directed. Thus the fluctuation-dissipation theorem does not apply and one can measure the distance from equilibrium through its violation. We investigate the influence of different parameters on the force distribution, focusing particularly on the role of ligand density : a detailed study shows how the amplitude of active forces increases when the bead is more tightly attached to the cortex. We introduce and study a model, which takes into account the number of bonds between the bead and the cytoskeleton, as well as the viscoelastic properties of the medium. This model faithfully accounts for the experimental observations. Also, it is shown that the amplitude of active forces increases with temperature. Finally, our data confirm that ATP depletion in the cell, or partial inhibition of the actomyosin activity, leads to a decrease of the amplitude of the force distribution. Altogether, we propose a consistent and quantitative description for the motion of a micrometric probe interacting with the actin network, and for the amplitude of the stochastic forces generated by molecular motors in the cortex surrounding this probe.