Violations of the fluctuation-dissipation theorem reveal distinct nonequilibrium dynamics of brain states.

The brain is a nonequilibrium system whose dynamics change in different brain states, such as wakefulness and deep sleep. Thermodynamics provides the tools for revealing these nonequilibrium dynamics. We used violations of the fluctuation-dissipation theorem to describe the hierarchy of nonequilibrium dynamics associated with different brain states. Together with a whole-brain model fitted to empirical human neuroimaging data, and deriving the appropriate analytical expressions, we were able to capture the deviation from equilibrium in different brain states that arises from asymmetric interactions and hierarchical organization.