Grazing behavior and winter phytoplankton accumulation

Recent observations have shown that phytoplankton biomass increases in the North Atlantic during winter, even when the mixed layer is deepening and light is limited. Current theories suggest that this is due to a release from grazing pressure. Here we demonstrate that the often-used grazing models that are linear at low phytoplankton concentration do not allow for a wintertime increase in phytoplankton biomass. However, mathematical formulations of grazing as a function of phytoplankton concentration that are quadratic at low concentrations (or more generally decrease faster than linearly as phytoplankton concentration decreases) can reproduce the fall to spring transition in phytoplankton, including wintertime biomass accumulation. We illustrate this point with a minimal model for the annual cycle of North Atlantic phytoplankton designed to simulate phytoplankton concentration as observed by BioGeoChemical-Argo (BGC-Argo) floats in the North Atlantic. This analysis provides a mathematical framework for assessing hypotheses of phytoplankton bloom formation.