Application of Finite-Time and Control Thermodynamics to Biological Processes at Multiple Scales

Abstract

AbstractAn overall synthesis of biology and non-equilibrium thermodynamics remains a challenge at the interface between the physical and life sciences. Herein, theorems from finite-time and control thermodynamics are applied to biological processes to indicate which biological strategies will succeed over different time scales. In general, living systems maximize power at the expense of efficiency during the early stages of their development while proceeding at slower rates to maximize efficiency over longer time scales. The exact combination of yield and power depends upon the constraints on the system, the degrees of freedom in question, and the time scales of the processes. It is emphasized that biological processes are not driven by entropy production but, rather, by informed exergy flow. The entropy production is the generalized friction that is minimized insofar as the constraints allow. Theorems concerning thermodynamic path length and entropy production show that there is a direct tradeoff between the efficiency of a process and the process rate. To quantify this tradeoff, the concepts of compensated heat and waste heat are introduced. Compensated heat is the exergy dissipated, which is necessary for a process to satisfy constraints. Conversely, waste heat is exergy that is dissipated as heat, but does not provide a compensatory increase in rate or other improvement. We hypothesize that it is waste heat that is minimized through natural selection. This can be seen in the strategies employed at several temporal and spatial scales, including organismal development, ecological succession, and long-term evolution. Better understanding the roles of compensated heat and waste heat in biological processes will provide novel insight into the underlying thermodynamic mechanisms involved in metabolism, ecology, and evolution.