Cycles in Zero-Sum Differential Games and Biological Diversity

Negative frequency-dependent selection (i.e., declining fitness with increased frequency in the population) is thought to be one of the factors that maintains biological diversity. In this paper, we give a concrete mathematical argument supporting this. Our model is as follows: A collection of species derive their fitnesses via a rock-paper-scissors-type game whose precise payoffs are a function of the environment. The new aspect of our model lies in adding a feedback loop: the environment changes according to the relative fitnesses of the species (hence, payoffs change as a function of fitness, which in turn changes as a function of payoffs). The changes in the payoffs are in keeping with the principle of negative frequency-dependent selection, which is widespread in nature. In order to model our game as a continuous time dynamical system, we cast it in the setting of a differential game. We show that for certain parameters, this dynamics cycles, i.e., no species goes extinct and diversity is maintained. We believe that our techniques can be applied to optimization and machine learning to show that first order methods (e.g., gradient descent/ascent) do cycle even in online settings in which the loss function changes with time.