The role of terrestrial and phytoplankton-derived organic matter in planktonic food webs

Lakes are important global ecosystems and many of them are nutrient-poor (unproductive). Especially in northern boreal latitudes, lakes may be heavily subsidized by terrestrial organic material (t-OM) from peat layers in the catchment. Thus, in addition to heterotrophic bacteria and phytoplankton, zooplankton may also use the particulate fraction of peat layer t-OM (t-POM) as a potential food source in those systems. Inputs of t-OM in northern latitudes are anticipated to increase in the future due to increasing precipitation and temperature. As t-OM is a good substrate for bacterial growth and as bacteria can often outcompete phytoplankton for inorganic nutrients, the proportions of heterotrophic bacteria and phytoplankton are expected to change in unproductive lakes. This may have pronounced impacts on zooplankton population dynamics. The aim of my thesis was to investigate how changes in food quality and quantity will affect metazoan zooplankton performance in unproductive lakes. Three laboratory studies assessed the quality of specific food components (phytoplankton, bacteria and peat layer t-POM) and their effects on Daphnia survival, growth and reproduction. Further, a mesocosm study with a full natural plankton community tested the predictions of the Light:Nutrient-Hypothesis in an unproductive clear water lake in situ by adding carbon and inorganic nutrients and changing light availability. I found that pure bacterial (Pseudomonas sp.) or t-POM diets could not sustain Daphnia populations, even though both were readily ingested. Daphnids needed at least 10-20% phytoplankton (Rhodomonas) in the diet to survive and even higher proportions (≥ 50%) were necessary for the production of viable offspring. Further, I showed that the dilution of non-limiting concentrations of Rhodomonas with increasing proportions of Pseudomonas or t-POM led to decreased Daphnia performance. Both Pseudomonas and t-POM lack essential biochemicals (fatty acids and sterols). In contrast, mineral nutrient limitation only occurred on t-POM-dominated diets as evidenced by a labeling experiment that showed Daphnia can incorporate carbon and phosphorus from Rhodomonas and Pseudomonas with similar efficiencies. Thus, peat layer t-POM was a lower quality food than Pseudomonas. This was corroborated by the finding that intermediate additions of Pseudomonas to limiting amounts of Rhodomonas supported increased Daphnia survival, growth and reproduction while t-POM additions had no beneficial effect. My results suggest that high terrestrial stable isotope signals in metazoan zooplankton are most likely derived from t-OM that is channeled tohigher trophic levels via the microbial loop (i.e. heterotrophic bacteria and/or bacterivorous protozoa) but not from direct metazoan feeding on t-POM. Furthermore, bacteria may serve as an important supplement to zooplankton diets when phytoplankton abundance is low. However, a sufficient proportion of high quality phytoplankton is always necessary to fulfil mineral and especially biochemical requirements of zooplankton in unproductive aquatic systems. The results of the mesocosm study showed that the Light:Nutrient-Hypothesis is not applicable to unproductive clear water systems in which the phytoplankton community is dominated by mixotrophs. In the face of the theoretical predictions, low light levels led to decreased zooplankton biomass. This was most likely caused by a shift in the algal community composition towards less edible taxa. Another reason may have been a weakening of the microbial loop. This is in line with the results of the laboratory studies that point out the importance of the microbial food web for zooplankton nutrition in unproductive lakes