Asymmetry in community regulation: effects of predators and productivity.

Eutrophication and predator additions and extinctions are occurring in ecosystems worldwide. Although theory predicts that both will strongly alter the distribution of biomass in whole communities, empirical evidence has not been consolidated to quantitatively determine whether these theoretical predictions are generally borne out in real ecosystems. Here we analyze data from two types of trophic cascade studies, predator removals in factorial combination with fertilization and observed productivity gradients, to assess the role of top-down and bottom-up forces in structuring multi-trophic communities and compare results from these analyses to those from an extensive database of trophic cascade studies. We find that herbivore biomass declines and plant biomass increases in the presence of predators, regardless of system productivity. In contrast, while plants are increased by fertilization, this effect does not significantly increase herbivores in either the presence or absence of predators. These patterns are consistent among marine, freshwater, and terrestrial ecosystems and are largely independent of study size and duration. Thus, top-down effects of predation are transferred through more trophic levels than are bottom-up effects of eutrophication, showing strong asymmetry in the direction of control of biomass distribution in communities.