Control across scales: signals, information, and adaptive biological mechanical function

Biological systems perform an astonishing array of dynamical processes -- including development and repair, regulation, behavior and motor control, sensing and signaling, and adaptation, among others. Powered by the transduction of stored energy resources, these behaviors enable biological systems to regulate functions, achieve specific outcomes, and maintain stability far from thermodynamic equilibrium. These behaviors span orders of magnitude in length and time: from nanometer-scale molecular motors driving morphogenesis to kilometer-scale seasonal migrations, and from millisecond reflexes to millennia of evolutionary adaptations. While physical laws govern the dynamics of biological systems, they alone are insufficient to fully explain how living systems sense, decide, adapt, and, ultimately, control their dynamics. In this article, we argue that control theory provides a powerful, unifying framework for understanding how biological systems regulate dynamics to maintain stability across length and time scales far from equilibrium.