Improved bounds on the entropy production rate in living systems

Living systems maintain or increase local order, working against the second law. Thermodynamic consistency is restored as they dissipate heat, increasing the net entropy of their environment. Recently introduced estimators for the entropy production rate have provided major insights into the thermal efficiency of important cellular processes. In biological experiments, however, many degrees of freedom typically remain hidden to the observer, and in these cases, existing methods are not optimal. Here, by reformulating the problem within an optimization framework, we are able to infer improved bounds on the rate of entropy production from partial measurements of biological systems. Our approach yields provably optimal estimates given certain measurable transition statistics. In particular, it can reveal non-zero heat production rates even when non-equilibrium processes appear time symmetric and so may pretend to obey detailed balance. We demonstrate the broad applicability of this framework by providing improved bounds on the entropy production rate in a diverse range of biological systems including bacterial flagella motors, growing microtubules, and calcium oscillations within human embryonic kidney cells.