Abstract
Stream ecosystems exhibit high degrees of spatial heterogeneity in
environmental conditions, communities of organisms, and ecosystem
processes at nested scales from landscapes to microhabitats. This
heterogeneity may facilitate the co-occurrence of biogeochemical
processes that are favored under incompatible environmental conditions,
like dinitrogen (N2 gas) fixation and denitrification.
We hypothesized that environmental variation at the patch scale (1-10’s
m) would facilitate the co-occurrence of N2 fixation and
denitrification through the formation of ecosystem control points or
patches that show high reaction rates relative to the surrounding area.
We measured rates of N2 fixation and denitrification and
relative abundances of the genes nifH and nirS (genes that
encode for the enzymes nitrogenase and nitrite reductase respectively)
in patches determined by channel geomorphic units and substrate type in
seven streams encompassing a gradient of N and P concentrations. We
found ecosystem control points, where rates of N2 fixation and
denitrification were 1 to 4 times higher than reach-average rates (0.2
– 1400 μg m-2 h-1 and 350 – 60000
μg m-2 h-1, respectively), occurred
in all study streams. Most N2 fixation control points
were in patches with rock substrates, while denitrification rates and
relative abundances of nifH and nirS were higher in fine
sediment patches. Yet, in two of the streams, rates in the top 25% of
all patches for both denitrification and N2 fixation
occurred in the same patches, suggesting that variation in conditions at
the sub-patch scale can also facilitate co-occurrence of these
processes. Across all streams and patches, organic matter and dissolved
oxygen concentrations were important predictors of rates of
N2 fixation, denitrification, and nifH relative
abundance, while P concentration was important to N2
fixation and denitrification. Our results demonstrate that understanding
the spatial ecology of microbially-driven nutrient cycling is required
to characterize nutrient fluxes more completely in stream ecosystems.