Response of phytoplankton in an alpine lake to inputs of dissolved organic matter through nutrient enrichment and trophic forcing

Inputs of terrestrially derived dissolved organic matter (DOM) are increasing in alpine lakes due to multiple drivers such as climate change, tree line advancement, and insect epidemics. A 21 d microcosm experiment investigated three potential mechanisms by which increased inputs of terrestrial DOM subsidies might affect phytoplankton density, growth, and species assemblage: (1) directly, by providing nutrients enhancing growth of select phytoplankton species (nutrient stimulation hypothesis); (2) indirectly, through trophic forcing of zooplankton uniformly increasing the total biomass of all zooplankton that selectively graze on phytoplankton (trophic intensity hypothesis); and (3) indirectly, through trophic forcing of zooplankton by favoring zooplankton species that selectively graze on phytoplankton (trophic shift hypothesis). We manipulated DOM (terrestrial DOM additions vs. unmanipulated control), zooplankton (presence vs. absence), and incubation depth (epilimnion vs. hypolimnion) in a full 3 x 3 factorial design. Phytoplankton density and growth increased substantially and species assemblage shifted to near dominance by Asterionella formosa in the presence of DOM. Zooplankton biomass and growth increased with the addition of DOM, yet the species assemblage remained stable across treatments, and contributed to selective grazing effects on phytoplankton. Our data support the nutrient stimulation and trophic intensity hypotheses. While DOM effects have been classically attributed to stimulation by addition of fixed carbon, our experiments indicate that nutrient stimulation is also important. Additionally, the indirect DOM effect of trophic forcing can occur in the absence of selective effects of DOM on zooplankton.