Exploring the role of spatial and stoichiometric heterogeneity in the top-down control in eutrophic planktonic ecosystems.

Understanding the impact of eutrophication on the dynamics of aquatic food webs, remains a long-term challenge in ecology. Mathematical models generally predict the destabilisation of such webs, under increasing eutrophication levels, with large oscillations of species densities that can result in their extinction. This is at odds with a number of ecological observations that show stable dynamics even for high nutrient loads. The apparent discrepancy between theory and observations is known as the Rosenzweig's 'paradox of enrichment' and various solutions to the problem have been proposed over the years. In this study, we explore the stabilisation of dynamics of a tri-trophic plankton model in a eutrophic environment which occurs as a result of interplay of space heterogeneity, ecological stoichiometry, and food taxis of predators. We build a variety of models of increasing complexity, to explore various scenarios of phytoplankton growth, zooplankton food-dependent vertical movement, and different stoichiometric limitations of zooplankton. We show that the synergy among the vertical gradient in phytoplankton growth, phytoplankton structuring in terms of their stoichiometric ratio, and food-dependent vertical movement of zooplankton, would result in a postponing of destabilisation of eutrophic systems as compared to a well-mixed system. Our approach reveals a high complexity of the bifurcation structure of the system when key model parameters, such as the degree of eutrophication and light shading, are varied. We find coexistence of limit cycles and stable equilibria and that the possibility of multiple attractors in the system can result in hysteresis phenomena when the nutrient load is manipulated. These results are relevant and should be taken into account in lake restoration programs.