Body Size, Light Intensity, and Nutrient Supply Determine Plankton Stoichiometry in Mixotrophic Plankton Food Webs

Trophic strategy determines stoichiometry of plankton. In general, heterotrophic zooplankton have lower and more stable C∶N and C∶P ratios than photoautotrophic phytoplankton, whereas mixotrophic protists, which consume prey and photosynthesize, have stoichiometry between zooplankton and phytoplankton. As trophic strategies change with cell size, body size may be a key trait influencing eukaryotic plankton stoichiometry. However, the relationship between body size and stoichiometry remains unclear. Here we measured plankton size-fractionated C∶N ratios under different intensities of light and nutrient supply in subtropical freshwater and marine systems. We found a unimodal body size-C∶N ratio pattern, with a maximum C∶N ratio at ∼50 μm diameter in marine and freshwater systems. Moreover, the variation in C∶N ratios is explained mainly by body size, followed by light intensity and nutrient concentration. To investigate the mechanisms behind this unimodal pattern, we constructed a size-based plankton food web model in which the trophic strategy and C∶N ratio are an emerging result. Our model simulations reproduce the unimodal pattern with a C∶N ratio of photoautotrophs ≤50 μm increasing with body size due to increase of photosynthetic carbon, whereas C∶N ratios of organisms >50 μm decrease with size due to decreasing photoautotrophic but increasing heterotrophic uptake. Based on our field observations and simulation, we extend the classic "light-nutrient" theory that determines plankton C∶N ratio to include body size and trophic strategy dependency. We conclude that body size and size-dependent uptake of resources (light, nutrients, and prey) determine plankton stoichiometry at various light and nutrient supplies.