Pia Romare

Effects of Enrichment on Simple Aquatic Food Webs

Simple models, based on Lotka‐Volterra types of interactions between predator and prey, predict that enrichment will have a destabilizing effect on populations and that equilibrium population densities will change at the top trophic level and every second level below. We experimentally tested these predictions in three aquatic food web configurations subjected to either high or low nutrient additions. The results were structured by viewing the systems as either food chains or webs and showed that trophic level biomass increased with enrichment, which contradicts food chain theory. However, within each trophic level, food web configuration affected the extent to which different functional groups responded to enrichment. By dividing trophic levels into functional groups, based on vulnerability to consumption, we were able to identify significant effects that were obscured when systems were viewed as food chains. The results support the prediction that invulnerable prey may stabilize trophic‐level dynamics by replacing other, more vulnerable prey. Furthermore, the vulnerable prey, such as Daphnia and edible algae, responded as predicted by the paradox of enrichment hypothesis; that is, variability in population density increased with enrichment. Hence, by describing ecosystems as a matrix of food web interactions, and by recognizing the interplay between interspecific competition and predation, a more complete description of the ecosystem function was obtained compared to when species were placed into distinct trophic levels.

Consumption patterns, complexity and enrichment in aquatic food chains

The interactions between consumers and prey, and their impact on biomass distribution among trophic levels, are central issues in both empirical and theoretical ecology. In a long–term experiment, where all organisms, including the top predator, were allowed to respond to environmental conditions by reproduction, we tested predictions from ‘prey–dependent’and ‘ratio–dependent’models. Prey–dependent models made correct predictions only in the presence of strong interactors in simple food chains, but failed to predict patterns in more complex situations. Processes such as omnivory, consumer excretion, and unsuitable prey–size windows (invulnerable prey) increased the complexity and created patterns resembling ratio–dependent consumption. However, whereas the prey–dependent patterns were created by the mechanisms predicted by the model, ratio–dependent patterns were not, suggesting that they may be ‘right for the wrong reason’. We show here that despite the enormous complexity of ecosystems, it is possible to identify and disentangle mechanisms responsible for observed patterns in community structure, as well as in biomass development of organisms ranging in size from bacteria to fish.

Synthesis of theoretical and empirical experiences from nutrient and cyprinid reductions in Lake Ringsjön

The reduction in external phosphorus load to Lake Ringsjön during the 1980s, did not result in improved water transparency during the following ten-year period. Furthermore, a fish-kill in the Eastern Basin of the lake, in addition to a cyprinid reduction programme (biomanipulation; 1988–1992), in contrast to theory, did not lead to any increase in zooplankton biomass or size. This absence of response in the pelagic food chain may have been attributed to the increase in abundance of YOY (0+) fish, following the fish reduction programme. Despite the lack of effect on zooplankton, there was a decrease in phytoplankton biomass, a change in species composition and an increase in water transparency following biomanipulation. In 1989, one year after the fish-kill in Eastern Basin, the Secchi depth (summer mean) increased from 60 cm to 110 cm. In the following years, water transparency increased further, despite an increase in phosphorus loading. An unexpected effect of the biomanipulation was an increase in benthic invertebrate and staging waterfowl abundances, which occurred 2–4 years after fish reduction. Hence, the response in the benthic community following biomanipulation was considerably stronger than in the pelagic community. A likely explanation is that reduction in abundance of the benthic feeding fish species bream (Abramis brama), strongly affected the benthic invertebrate fauna. In this paper, we present what we believe happened in Lake Ringsjön, and which processes are likely to have been important at various stages of the restoration process.