Continuous model for the rock–scissors–paper game between bacteriocin producing bacteria

In this work, important aspects of bacteriocin producing bacteria and their interplay are elucidated. Various attempts to model the resistant, producer and sensitive Escherichia coli strains in the so-called rock-scissors-paper (RSP) game had been made in the literature. The question arose whether there is a continuous model with a cyclic structure and admitting an oscillatory dynamics as observed in various experiments. The May-Leonard system admits a Hopf bifurcation, which is, however, degenerate and hence inadequate. The traditional differential equation model of the RSP-game cannot be applied either to the bacteriocin system because it involves positive interaction terms. In this paper, a plausible competitive Lotka-Volterra system model of the RSP game is presented and the dynamics generated by that model is analyzed. For the first time, a continuous, spatially homogeneous model that describes the competitive interaction between bacteriocin-producing, resistant and sensitive bacteria is established. The interaction terms have negative coefficients. In some experiments, for example, in mice cultures, migration seemed to be essential for the reinfection in the RSP cycle. Often statistical and spatial effects such as migration and mutation are regarded to be essential for periodicity. Our model gives rise to oscillatory dynamics in the RSP game without such effects. Here, a normal form description of the limit cycle and conditions for its stability are derived. The toxicity of the bacteriocin is used as a bifurcation parameter. Exact parameter ranges are obtained for which a stable (robust) limit cycle and a stable heteroclinic cycle exist in the three-species game. These parameters are in good accordance with the observed relations for the E. coli strains. The roles of growth rate and growth yield of the three strains are discussed. Numerical calculations show that the sensitive, which might be regarded as the weakest, can have the longest sojourn times.