First and second laws of information processing by nonequilibrium dynamical states.

The averaged steady-state surprisal links a driven stochastic system's information processing to its nonequilibrium thermodynamic response. By explicitly accounting for the effects of nonequilibrium steady states, a decomposition of the surprisal results in an information processing first law that extends and tightens-to strict equalities-various information processing second laws. Applying stochastic thermodynamics' integral fluctuation theorems then shows that the decomposition reduces to the second laws under appropriate limits. In unifying them, the first law paves the way to identifying the mechanisms by which nonequilibrium steady-state systems leverage information-bearing degrees of freedom to extract heat. To illustrate, we analyze an autonomous Maxwellian information ratchet that tunably violates detailed balance in its effective dynamics. This demonstrates how the presence of nonequilibrium steady states qualitatively alters an information engine's allowed functionality.

Homeostatic and adaptive energetics: Nonequilibrium fluctuations beyond detailed balance in voltage-gated ion channels.

Stochastic thermodynamics has largely succeeded in characterizing both equilibrium and far-from-equilibrium phenomena. Yet many opportunities remain for application to mesoscopic complex systems-especially biological ones-whose effective dynamics often violate detailed balance and whose microscopic degrees of freedom are often unknown or intractable. After reviewing excess and housekeeping energetics-the adaptive and homeostatic components of a system's dissipation-we extend stochastic thermodynamics with a trajectory class fluctuation theorem for nonequilibrium steady-state, nondetailed-balanced complex systems. We then take up the neurobiological examples of voltage-gated sodium and potassium ion channels to apply and illustrate the theory, elucidating their nonequilibrium behavior under a biophysically plausible action potential drive. These results uncover challenges for future experiments and highlight the progress possible understanding the thermodynamics of complex systems-without exhaustive knowledge of every underlying degree of freedom.