Accessing activity and viscoelastic properties of artificial and living systems from passive measurement

Living systems are complex dynamic entities that operate far from thermodynamic equilibrium. Their active, non-equilibrium behaviour requires energy to drive cellular organization and dynamics. Unfortunately, most statistical mechanics approaches are not valid in non-equilibrium situations, forcing researchers to use intricate and often invasive methods to study living processes. Here we experimentally demonstrate that an observable termed mean back relaxation quantifies the active mechanics of living cells from passively observed particle trajectories. The mean back relaxation represents the average trajectory of a particle after a recent motion and is calculated from three-point probabilities. We show that this parameter allows the detection of broken detailed balance in confined systems. We experimentally observe that it provides access to the non-equilibrium generating energy and viscoelastic properties of artificial bulk materials and living cells. These findings suggest that the mean back relaxation can function as a marker of non-equilibrium dynamics and is a non-invasive avenue to determine viscoelastic material properties from passive measurements. An approach based on the average trajectory of moving particles allows for the quantification of the mechanics of living systems, namely, the non-equilibrium energy and viscoelastic properties of cells, in a non-invasive manner.