Periodic coexistence in the chemostat with three species competing for three essential resources.

A chemostat model of three species of microorganisms competing for three essential, growth-limiting nutrients is considered. J. Husiman and F.J. Weissing [Nature 402 (1999) 407] show numerically that this model can generate periodic oscillations. The present contribution is concerned with rigorous analysis regarding the existence of periodic oscillations in this model. Our analysis is based on the following observation made by Huisman and Weissing: there is a cyclic replacement of species, if each species becomes limited by the resource for which it is the intermediate competitor. Using a permanence theory, an index theory, and a Poincaré-Bendixson theory for three-dimensional competitive systems, we analytically succeed to give sufficient conditions for the existence of periodic orbits in the limit sets in this model. The results in this paper suggest that with a wide range of parameter values, sustained periodic oscillations of species abundances for the model are possible, without involving external disturbances. Our results also suggest that competition is not necessarily destructive, i.e., in the case of existence of sustained periodic oscillations, if one of three competitors is absent, one of the other two rivals cannot survive.