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.

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Although the scale-dependence of ecological patterns and processes is recognized by freshwater ecologists, current knowledge of scale effects is rudimentary and non-quantitative. We review issues of spatial and temporal scale in this paper to highlight conceptual problems relating to scale and some potential solutions. We present examples of how the spatial scale of a study influences observed patterns and their interpretation, and discuss how the size of an experimental arena influences the degree to which the dynamics of studied populations are influenced by exchange processes (immigration and emigration). The results of small-scale field experiments in streams will often be strongly influenced by the per capita exchange rates of organisms and differences in exchange rates may explain differences in the perceived effects of stream manipulations across scales. Spatial extent also influences the amount of spatial heterogeneity within a study site or arena, with important consequences for the outcome of predator-prey interactions. We suggest that changes in the availability of prey refuges may help explain why predator manipulations in streams appear to weaken as arena size increases. We also recommend that new techniques for decomposing and quantifying spatial heterogeneity be applied to characterize scaledependent variation in freshwater systems. Lastly, we discuss the pitfalls of mismatching the temporal scale of experiments and models. Models incorporating spatial heterogeneity and the behaviour of organisms are needed to predict the short-term outcome of perturbations in streams, whereas models predicting long-term dynamics will need to integrate the impacts of episodic disturbance and all life history stages of organisms. In general, we recommend that freshwater ecologists undertake more multi-scale sampling and experimentation to examine patterns and processes at multiple scales, and make greater attempts to match the scales of their observations and experiments to the characteristic scales of the phenomena that they investigate.

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