Although trophic cascades (indirect effects of predators on plants via herbivores) occur in a wide variety of food webs, the magnitudes of their effects are often quite variable. We compared the responses of herbivore and plant communities to predator manipulations in 102 field experiments in six different ecosystems: lentic (lake and pond), marine, and stream benthos, lentic and marine plankton, and terrestrial (grasslands and agricultural fields). Predator effects varied considerably among systems and were strongest in lentic and marine benthos and weakest in marine plankton and terrestrial food webs. Predator effects on herbivores were generally larger and more variable than on plants, suggesting that cascades often become attenuated at the plant–herbivore interface. Top‐down control of plant biomass was stronger in water than on land; however, the differences among the five aquatic food webs were as great as those between wet and dry systems.

Trophic cascades have been documented in a diversity of ecological systems and can be important in determining biomass distribution within a community. To date, the literature on trophic cascades has focused on whether and in which systems cascades occur. Many biological (e.g., productivity : biomass ratios) and methodological (e.g., experiment size or duration) factors vary with the ecosystem in which data were collected, but ecosystem type, per se, does not provide mechanistic insights into factors controlling cascade strength.Here, we tested various hypotheses about why trophic cascades occur and what determines their magnitude using data from 114 studies that measured the indirect trophic effects of predators on plant community biomass in seven aquatic and terrestrial ecosystems. Using meta-analysis, we examined the relationship between the indirect effect of predator manipulation on plants and 18 biological and methodological factors quantified from these studies. We found, in contrast to predictions, that high system productivity and low species diversity do not consistently generate larger trophic cascades. A combination of herbivore and predator metabolic factors and predator taxonomy (vertebrate vs. invertebrate) explained 31% of the variation in cascade strength among all 114 studies. Within systems, 18% of the variation in cascade strength was explained with similar predator and herbivore characteristics. Within and across all systems, the strongest cascades occurred in association with invertebrate herbivores and endothermic vertebrate predators. These associations may result from a combination of true biological differences among species with different physiological requirements and bias among organisms studied in different systems. Thus, although cascade strength can be described by biological characteristics of predators and herbivores, future research on indirect trophic effects must further examine biological and methodological differences among studies and systems.

Some studies suggest that lotic populations of brown trout (Salmo trutta) are regulated through density-dependent mortality and emigration to the extent that mean growth rates of resident survivors are unrelated to trout densities. To test this, we studied the relationship between density and growth, mortality, and emigration of brown trout in two alpine streams and a set of stream channels in eastern California. We sampled trout at the scale of ''segments'' (5-31 m long riffles, runs, and pools) and ''sections'' (340-500 m in length) of Convict Creek over a 3-yr period. Trout were also sampled during 6 yr in seven 90-m sections of Mammoth Creek. For 2 yr, we manipulated trout densities in Convict Creek by removing trout from two sections and adding trout to two other sections. We also manipulated densities in seven 50-m stream channels, using a natural size distribution of trout in one year and underyearlings only in a second year.In both streams, average size (body length or mass) of underyearlings in fall was negatively related to trout density and was furthermore affected by sampling location and year. The strong, negative relationship between individual mass and density of trout could be detected at the spatial scale of whole sections, but not at the scale of individual segments. The Convict Creek and stream channel experiments also revealed strong negative effects of density on average mass of underyearlings in fall, and on proportional mass increase of yearling and older trout from spring to fall. In contrast, mortality and emigration were unrelated to initial stocking densities in the channels. In all our data, the negative effects on growth were most pronounced at densities Ͻ1 trout/m 2 and the growth-density relationships were well described by negative power curves. Large individuals were always less affected by increasing trout density than were small individuals, suggesting a competitive advantage of large over small trout that increased with density.We conclude that individual growth of brown trout in streams can be affected by trout density to an extent that suggests a substantial influence on population regulation. Results from our multiyear, multiscale, and experimental study indicate that density dependence in the growth of stream salmonids will be difficult to detect in purely observational data, especially in systems with relatively high fish densities (where the growth-density relationship has a flat slope), when data are collected and analyzed at small spatial scales, and when insufficient information is collected to assess the contribution of interannual variation in growth.

We present four lines of evidence that the magnitude of prey exchange (=immigration/emigration) among substrate patches has an overwhelming influence on the perceived effects of predators on prey populations. (1) An extensive review of the literature on predation effects in benthic and littoral freshwater revealed a significant relationship between prey exchange rate and observed predator impact. In streams, studies showing significant predator effects used cages with smaller mesh sizes than studies showing nonsignificant effects. Similarly, there was a highly significant correlation between cage mesh size and the magnitude of predator impact on common prey. Large—scale stream studies indicated that prey drift and colonization rate were inversely related to predator impact on benthic prey. (2) These patterns were confirmed by field experiments and observations where mesh size was directly manipulated or where exchange rates varied among taxa. In Colorado streams we saw greater predator impacts on Baetis prey when immigration/emigration was restricted vs. when the mesh size of the cage was relatively large. Similarly, the effects of trout in California stream pools were greater when prey turnover rates were low. (3) A re—analysis of Peckarsky's (1985) data shows an inverse relationship between predator impact and prey mobility within a field experiment. (4) Finally, a model that incorporates both predation and exchange of prey indicates that we ought to expect a lower magnitude of predator effects when exchange rates are high. These results suggest that some discrepancies in past studies may be explained by differences in the exchange rates of prey, and that differences in predator effects across different systems or habitats may be related to variation in the rates of prey dispersal and colonization.

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