Nitrogen uptake and transformation in a midwestern U.S. stream: A stable isotope enrichment study

This study presents a comprehensive analysis ofnitrogen (N) cycling in a second-order forestedstream in southern Michigan that has moderatelyhigh concentrations of ammonium (mean,16 g N/L) and nitrate (17 g N/L). Awhole-stream ¹⁵NH₄ ⁺ addition wasperformed for 6 weeks in June and July, and thetracer ¹⁵N was measured downstream inammonium, nitrate, and detrital and livingbiomass. Ancillary measurements includedbiomass of organic matter, algae, bacteria andfungi, nutrient concentrations, hydrauliccharacteristics, whole-stream metabolism, andnutrient limitation assays. The resultsprovide insights into the heterotrophic natureof woodland streams and reveal the rates atwhich biological processes alter nitrogentransport through stream systems.Ammonium uptake lengths were 766–1349 m anduptake rates were 41–60 g N m^–2min^–1. Nitrate uptake could not bedetected. Nitrification rates were estimatedfrom the downstream increase in¹⁵N-enriched nitrate using a simulationmodel. The ammonium was removed bynitrification (57% of total uptake),heterotrophic bacteria and fungi associatedwith detritus (29%), and epilithic algae(14%). Growth of algae was likely limited bylight rather than nutrients, and dissolvedO₂ revealed that the stream metabolism washeterotrophic overall (P:R = 0.2). Incubationsof detritus in darkened chambers showed thatuptake of ¹⁵N was mostly heterotrophic.Microbial N in detritus and algal N inepilithon appeared to reach isotopic steadystate with the dissolved ammonium, but theisotopic enrichment of the bulk detritus andepilithon did not approach that of ammonium,probably due to a large fraction of organic Nin the bulk samples that was not turning over. The actively cycling fraction of total N inorganic compartments was estimated from theisotopic enrichment, assuming uptake ofammonium but not nitrate, to be 23% forepilithon, 1% for fine benthic organic matter,5% for small woody debris, and 7% for leaves. These percentages agree with independentestimates of epilithic algal biomass, whichwere based on carbon:chlorophyll ratios in bulksamples and in algal fractions separated bydensity-gradient centrifugation in colloidalsilica, and of microbial N in the detritus,which were based on N released by chloroformfumigations.