A comparative (15)N-tracer study of nitrogen dynamics in headwater streams from biomes throughout North America demonstrates that streams exert control over nutrient exports to rivers, lakes, and estuaries. The most rapid uptake and transformation of inorganic nitrogen occurred in the smallest streams. Ammonium entering these streams was removed from the water within a few tens to hundreds of meters. Nitrate was also removed from stream water but traveled a distance 5 to 10 times as long, on average, as ammonium. Despite low ammonium concentration in stream water, nitrification rates were high, indicating that small streams are potentially important sources of atmospheric nitrous oxide. During seasons of high biological activity, the reaches of headwater streams typically export downstream less than half of the input of dissolved inorganic nitrogen from their watersheds.

Region 2 comprises arctic and subarctic North America and is underlain by continuous or discontinuous permafrost. Its freshwater systems are dominated by a low energy environment and cold region processes. Central northern areas are almost totally in¯uenced by arctic air masses while Paci®c air becomes more prominent in the west, Atlantic air in the east and southern air masses at the lower latitudes. Air mass changes will play an important role in precipitation changes associated with climate warming. The snow season in the region is prolonged resulting in long-term storage of water so that the spring¯ood is often the major hydrological event of the year, even though, annual rainfall usually exceeds annual snowfall. The unique character of ponds and lakes is a result of the long frozen period, which aects nutrient status and gas exchange during the cold season and during thaw. GCM models are in close agreement for this region and predict temperature increases as large as 48C in summer and 98C in winter for a 2 Â CO 2 scenario. Palaeoclimate indicators support the probability that substantial temperature increases have occurred previously during the Holocene. The historical record indicates a temperature increase of 418C in parts of the region during the last century. GCM predictions of precipitation change indicate an increase, but there is little agreement amongst the various models on regional disposition or magnitude. Precipitation change is as important as temperature change in determining the water balance. The water balance is critical to every aspect of hydrology and limnology in the far north. Permafrost close to the surface plays a major role in freshwater systems because it often maintains lakes and wetlands above an impermeable frost table, which limits the water storage capabilities of the subsurface. Thawing associated with climate change would, particularly in areas of massive ice, stimulate landscape changes, which can aect every aspect of the environment. The normal spring¯ooding of ice-jammed north-¯owing rivers, such as the Mackenzie, is a major event, which renews the water supply of lakes in delta regions and which determines the availability of habitat for aquatic organisms. Climate warming or river damming and diversion would probably lead to the complete drying of many delta lakes. Climate warming would also change the characteristics of ponds that presently freeze to the bottom and result in fundamental changes in their limnological characteristics. At present, the food chain is rather simple usually culminating in lake trout or arctic char. A lengthening of the growing season and warmer water temperature would aect the chemical, mineral and nutrient status of lakes and most likely have deleterious eects on the food chain. Peatlands are extensive in region 2. They would move northwards at their southern boundaries, and, with sustained drying, many would change form or become inactive. Extensive wetlands and peatlands are an important component of the global carbon budget, and warm...

Phosphorus fertilization of a pristine tundra river for four consecutive summers dramatically changed biological processes and populations at all trophic levels. At the primary producer level, both algal biomass and productivity increased and chlorophyll accumulated on the river bottom in the first two summers. Diatom community composition changed little in spite oflarge chlorophyll changes. However, an increase in grazing insects prevented chlorophyll buildup in the third and fourth summers.Some microbial processes were also stimulated by the increased photosynthesis caused by fertilization. Total respiration of the epilithon, acetate uptake, and decomposition of lignin monomers were all stimulated but only in light-grown epilithon. When epilithon was grown in the dark in the fertilized region of the river, there was no increased respiration. Also, phosphorus did not stimulate the decomposition of Carex litter.Although insects grew more rapidly in the fertilized section of the river, there were community interactions that kept total insect production from appreciable change. The four most abundant large insects did increase their growth rates in response to phosphorus addition and there were increases in populations of Baetis lapponicus and Brachycentrus americanus. These increases were offset by the decline in abundance of the dominant species, the black fly Prosimulium martini, perhaps caused by competition for space from Brachycentrus.Growth of both young-of-the-year and adult grayling (Thymallus arcticus) was strongly stimulated by phosphorus addition in years 3 and 4 (not tested in years 1 and 2). Carbon and nitrogen stable isotope tracers indicated that the measured increases in insect and fish growth were largely attributable to increases in the production of epilithic algae. Overall, the results indicate a strong "bottom-up" response of the riverine food web to additions of the limiting nutrient, phosphorus. The response was modified in later years, however, by a strong "top-down" feedback of insects grazing on epilithic algae and by competitive exclusion of black flies by caddisflies.

A long-term stream fertilization experiment was performed to evaluate the potential eutrophication of an arctic stream ecosystem. During 16 years of summer phosphorus (H 3 PO 4 ) fertilization, we observed a dramatic change in the community structure of the Kuparuk River on the North Slope of Alaska. A positive response to fertilization was observed at all trophic levels with increases in epilithic algal stocks, some insect densities, and fish growth rates. After approximately eight years of P fertilization, bryophytes (mosses) replaced epilithic diatoms as the dominant primary producers in the Kuparuk River. The moss impacted NH 4 ϩ uptake rates, benthic gross primary production, habitat structure, and insect abundance and species composition. This study documents the long-term changes in an arctic tundra stream in response to nutrient enrichment. Predicting stream ecosystem responses to chronic perturbation requires long-term observation and experiments.

Detailed studies of stream N uptake were conducted in a prairie reach and gallery forest reach of Kings Creek on the Konza Prairie Biological Station. Nutrient uptake rates were measured with multiple short-term enrichments of NO 3 Ϫ and NH 4 ϩ at constant addition rates in the spring and summer of 1998. NH 4 ϩ uptake was also measured with 15 N-NH 4 ϩ tracer additions and short-term unlabeled NH 4 ϩ additions at 12 stream sites across North America. Concurrent addition of a conservative tracer was used to account for dilution in all experiments. NH 4 ϩ uptake rate per unit area (U t) was positively correlated to nutrient concentration across all sites (r 2 ϭ 0.41, log-log relationship). Relationships between concentration and U t were used to determine whether the uptake was nonlinear (i.e., kinetic uptake primarily limited by the biotic capacity of microorganisms to accumulate nutrients) or linear (e.g., limited by mass transport into stream biofilms). In all systems, U t was lower at ambient concentrations than at elevated concentrations. Extrapolation from uptake measured from a series of increasing enrichments could be used to estimate ambient U t. Linear extrapolation of U t assuming the relationship passes through the origin and rates measured at 1 elevated nutrient concentration underestimated ambient U t by ϳ3-fold. Uptake rates were saturated under some but not all conditions of enrichment; in some cases there was no saturation up to 50 mol/L. The absolute concentration at which U t was saturated in Kings Creek varied among reaches and nutrients. Uptake rates of NH 4 ϩ at ambient concentrations in all streams were higher than would be expected, assuming U t does not saturate with increasing concentrations. At ambient nutrient concentrations in unpolluted streams, U t is probably limited to some degree by the kinetic uptake capacity of stream biota. Mass transfer velocity from the water column is generally greater than would be expected given typical diffusion rates, underscoring the importance of advective transport. Given the short-term spikes in nutrient concentrations that can occur in streams (e.g., in response to storm events), U t may not saturate, even at high concentrations.