Regulatory out-of-equilibrium processes in cells. Part I: Temporal heterogeneities and oscillations

Making decisions is a core task for living matter, and a task that can only be carried out through processes out of equilibrium. While important elements of nonequilibrium processes occur in the spatial distribution of resources and within genetic heterogeneity, temporal dynamics of protein concentrations play a critical yet underappreciated role in cellular decision-making. This review explores our current understanding of how cells use dynamic signaling patterns, such as oscillations, to regulate processes and cell-fate decisions across multiple biological systems. We focus on examples including the tumor suppressor p53, immune signaling via NF- κ B, and oscillatory networks in development. We further highlight theoretical frameworks and the current horizon in our understanding of this fundamental interplay. • Temporal heterogeneity increases adaptability in cells • Non-equilibrium as a key information pathway in biology • Interplay between theoretical and experimental methods • Oscillations provide functional advantages across scale and fields