Relative Entropy in Biological Systems

In this paper we review various information-theoretic characterizations of the approach to equilibrium in biological systems. The replicator equation, evolutionary game theory, Markov processes and chemical reaction networks all describe the dynamics of a population or probability distribution. Under suitable assumptions, the distribution will approach an equilibrium with the passage of time. Relative entropy—that is, the Kullback–Leibler divergence, or various generalizations of this—provides a quantitative measure of how far from equilibrium the system is. We explain various theorems that give conditions under which relative entropy is nonincreasing. In biochemical applications these results can be seen as versions of the Second Law of Thermodynamics, stating that free energy can never increase with the passage of time. In ecological applications, they make precise the notion that a population gains information from its environment as it approaches equilibrium.