Why do phytoplankton evolve large size in response to grazing?

Phytoplankton are among the smallest primary producers on Earth, yet they display a wide range of cell sizes. Typically, small phytoplankton species are stronger nutrient competitors than large phytoplankton species, but they are also more easily grazed. In contrast, evolution of large phytoplankton is often explained as a physical defense against grazing. Conceptually, this explanation is problematic, however, because zooplankton can coevolve larger size to counter this size-dependent escape from grazing. Here, we hypothesize that there is another advantage for the evolution of large phytoplankton size not so readily overcome: larger phytoplankton often provide lower nutritional quality for zooplankton. We investigate this hypothesis by analyzing an eco-evolutionary model that combines the ecological stoichiometry of phytoplankton-zooplankton interactions with coevolution of phytoplankton and zooplankton size. In our model, evolution of cell size modifies the nutrient uptake kinetics of phytoplankton according to known allometric relationships, which in turn affect the nutritional quality of phytoplankton. With this size-based mechanism, the model predicts that low grazing pressure or nonselective grazing by zooplankton favors evolution of small phytoplankton cells of high nutritional quality. In contrast, selective grazing for nutritious food favors evolution of large phytoplankton of low nutritional quality, which are preyed on by medium-to large-sized zooplankton. This size-dependent change in food quality may explain the commonly observed shift from dominance by small picophytoplankton in oligotro-phic waters with low grazing pressure to large phytoplankton species in nutrient-rich waters with high grazing pressure.