Towards developing a general framework for modelling vertical migration in zooplankton

Abstract

Diel vertical migration (DVM) of zooplankton is a widespread phenomenon in both oceans and lakes, and is generally considered to be the largest synchronized movement of biomass on Earth. Most existing mathematical models of DVM are based on the assumption that animals maximize a certain criterion such as the expected reproductive value, the venturous revenue, the ratio of energy gain/mortality or some predator avoidance function when choosing their instantaneous depth. The major shortcoming of this general point of view is that the predicted DVM may be strongly affected by a subjective choice of a particular optimization criterion. Here we argue that the optimal strategy of DVM can be unambiguously obtained as an outcome of selection in the underlying equations of genotype/traits frequency dynamics. Using this general paradigm, we explore the optimal strategy for the migration across different depths by zooplankton grazers throughout the day. To illustrate our ideas we consider four generic DVM models, each making different assumptions on the population dynamics of zooplankton, and demonstrate that in each model we need to maximize a particular functional to find the optimal strategy. Surprisingly, patterns of DVM obtained for different models greatly differ in terms of their parameters dependence. We then show that the infinite dimensional trait space of different zooplankton trajectories can be projected onto a low dimensional space of generalized parameters and the genotype evolution dynamics can be easily followed using this low-dimensional space. Using this space of generalized parameters we explore the influence of mutagenesis on evolution of DVM, and we show that strong mutagenesis allows the coexistence of an infinitely large number of strategies whereas for weak mutagenesis the selection results in the extinction of most strategies, with the surviving strategies all staying close to the optimal strategy in the corresponding mutagenesis-free system.

Keywords: Infinite-dimensional traits; Pontryagin׳s maximum principle; Population fitness; Predator–prey model; Selection.