Microbial ecoenzyme stoichiometry, nutrient limitation, and organic matter decomposition in wetlands of the conterminous United States.

Microbial respiration (Rm) and ecoenzyme activities (EEA) related to microbial carbon, nitrogen, and phosphorus acquisition were measured in 792 freshwater and estuarine wetlands (representing a cumulative area of 217,480 km2) across the continental United States as part of the US EPA's 2011 National Wetland Condition Assessment. EEA stoichiometry was used to construct models for and assess nutrient limitation, carbon use efficiency (CUE), and organic matter decomposition (- k). The wetlands were classified into ten groups based on aggregated ecoregion and wetland type. The wetlands were also assigned to least, intermediate, and most disturbed classes, based on the extent of human influences. Ecoenzyme activity related to C, N and P acquisition, Rm, CUE, and (- k differed among ecoregion-wetland types and, with the exception of C acquisition and (- k, among disturbance classes. Rm and EEA were positively correlated with soil C, N and P content (r = 0.15-0.64) and stoichiometry (r = 0.15-0.48), and negatively correlated with an index of carbon quality (r = - 0.22 to - 0.39). EEA stoichiometry revealed that wetlands were more often P- than N-limited, and that P-limitation increases with increasing disturbance. Our enzyme-based approach for modeling C, N, and P acquisition, and organic matter decomposition, all rooted in stoichiometric theory, provides a mechanism for modeling resource limitations of microbial metabolism and biogeochemical cycling in wetlands. Given the ease of collecting and analyzing soil EEA and their response to wetland disturbance gradients, enzyme stoichiometry models are a cost-effective tool for monitoring ecosystem responses to resource availability and the environmental drivers of microbial metabolism, including those related to global climate changes.

Urban infrastructure influences dissolved organic matter quality and bacterial metabolism in an urban stream network.

1. Urban streams are degraded by a suite of factors, including burial beneath urban infrastructure, such as roads or parking lots, which eliminates light and reduces direct organic matter inputs to streams from riparian zones. These changes to stream metabolism and terrestrial carbon contribution will likely have consequences for organic matter metabolism by microbes and dissolved organic matter (DOM) use patterns in streams. Respiration by heterotrophic biofilms drives the nitrogen and phosphorus cycles, but we lack a clear understanding of how stream burial and seasonality affect microbial carbon use. 2. We studied seasonal changes (autumn, spring, and summer) in organic matter metabolism by microbial communities in open and buried reaches of three urban streams in Cincinnati, OH. We characterised DOM quality using fluorescence spectroscopy and extracellular enzyme profiles, and we measured the respiration response to carbon supplements in nutrient diffusing substrata (NDS). We hypothesised: (1) that algal production would lead to higher quality DOM in spring compared to other seasons and in open compared to buried reaches, (2) lower reliance of microbial respiration on recalcitrant carbon sources in spring and in open reaches, and (3) that microbial respiration would increase in response to added carbon in autumn and in buried reaches. 3. Several fluorescence metrics showed higher quality DOM in spring than autumn, but only the metric of recalcitrant humic compounds varied by reach, with more humic DOM in open compared to buried reaches. This likely reflected open reaches as an avenue for direct terrestrial inputs from the riparian zone. 4. Extracellular enzyme assays showed that microbes in buried reaches allocated more effort to degrade recalcitrant carbon sources, consistent with a lack of labile carbon compounds due to limited photosynthesis. Nitrogen acquisition enzymes were highest in autumn coincident with riparian leaf inputs to the streams. Buried and open reaches both responded more strongly to added carbon in autumn when terrestrial leaf inputs dominated compared to the spring when vernal algal blooms were pronounced. 5. Our data show that stream burial affects the quality of the DOM pool with consequences for how microbes use those carbon sources, and that heterotrophic respiration increased on carbon-supplemented NDS in buried and open stream reaches in both seasons. Different carbon quality and use patterns suggest that urban stream infrastructure affects spatiotemporal patterns of bacterial respiration, with likely consequences for nitrogen and/or phosphorus cycling given that carbon use drives other biogeochemical cycles. Management actions that increase light to buried streams could shift the balance between allochthonous and autochthonous DOM in urban streams with consequences for spatiotemporal patterns in bacterial metabolism.

A synoptic survey of microbial respiration, organic matter decomposition, and carbon efflux in U.S. streams and rivers.

We analyzed ecoenzyme activities related to organic matter processing in 1879 streams and rivers across the continental U.S. as part of the USEPA's National Rivers and Streams Assessment. Ecoenzymatic stoichiometry was used to construct models for carbon use efficiency (CUE) and decomposition (-k). Microbial respiration (Rm) was estimated from sediment organic carbon stocks, CUE and -k. The streams and rivers were classified by size (headwaters: 1st-order; streams: 2nd-3rd order; small rivers: 4th-5th order; big rivers 6th-7th order; and great rivers ≥ 8th order) and condition class (least, intermediate and most disturbed), and grouped into nine ecoregions. There were ecoregion, stream size, and condition class effects for CUE, -k, and Rm, with Rm increasing from eastern ecoregions through the plains to the western ecoregions. CUE, -k, and Rm decreased with increasing streams size and increased with increasing disturbance. Rm, CUE, and -k were correlated with water and sediment chemistry; CUE and -k were also correlated with stream bed fine sediments; and CUE was further correlated with catchment land cover. Rm was extrapolated to ecoregional and national scales, and the results suggest that microbial assemblages account for 12% of the total CO2 outgassing, and nearly 50% of the aquatic metabolism C losses, from U.S. streams and rivers. Cumulative respiratory C losses increased from headwaters to small streams, then decreased with increasing stream size. This U-shaped respiration curve was not evident when streams were viewed by disturbance classes, suggesting that anthropogenic disturbances mask the expected organic matter processing signature of the river continuum.

Comparisons of soil nitrogen mass balances for an ombrotrophic bog and a minerotrophic fen in northern Minnesota.

We compared nitrogen (N) storage and flux in soils from an ombrotrophic bog with that of a minerotrophic fen to quantify the differences in N cycling between these two peatlands types in northern Minnesota (USA). Precipitation, atmospheric deposition, and bog and fen outflows were analyzed for nitrogen species. Upland and peatland soil samples were analyzed for N content, and for ambient (DN) and potential (DEA) denitrification rates. Annual atmospheric deposition was: 0.88-3.07kg NH4(+)ha(-1)y(-1); 1.37-1.42kg NO3(-)ha(-1)y(-1); 2.79-4.69kg TNha(-1)y(-1). Annual N outflows were: bog-0.01-0.04kg NH4(+)ha(-1)y(-1), NO3(-) 0.01-0.06kgha(-1)y(-1), and TN 0.11-0.69kgha(-1)y(-1); fen-NH4(+) 0.01-0.16kgha(-1)y(-1), NO3(-) 0.29-0.48kgha(-1)y(-1), and TN 1.14-1.61kgha(-1)y(-1). Soil N content depended on location within the bog or fen, and on soil depth. DN and DEA rates were low throughout the uplands and peatlands, and were correlated with atmospheric N deposition, soil N storage, and N outflow. DEA was significantly greater than DN indicating C or N limitation of the denitrification process. We highlight differences between the bog and fen, between the upland mineral soils and peat, and the importance of biogeochemical hotspots within the peatlands. We point out the importance of organic N storage, as a source of N for denitrification, and propose a plausible link between organic N storage, denitrification and N export from peatlands. Finally, we considered the interactions of microbial metabolism with nutrient availability and stoichiometry, and how N dynamics might be affected by climate change in peatland ecosystems.

Quantifying Urban Watershed Stressor Gradients and Evaluating How Different Land Cover Datasets Affect Stream Management.

Watershed management and policies affecting downstream ecosystems benefit from identifying relationships between land cover and water quality. However, different data sources can create dissimilarities in land cover estimates and models that characterize ecosystem responses. We used a spatially balanced stream study (1) to effectively sample development and urban stressor gradients while representing the extent of a large coastal watershed (>4400 km(2)), (2) to document differences between estimates of watershed land cover using 30-m resolution national land cover database (NLCD) and <1-m resolution land cover data, and (3) to determine if predictive models and relationships between water quality and land cover differed when using these two land cover datasets. Increased concentrations of nutrients, anions, and cations had similarly significant correlations with increased watershed percent impervious cover (IC), regardless of data resolution. The NLCD underestimated percent forest for 71/76 sites by a mean of 11 % and overestimated percent wetlands for 71/76 sites by a mean of 8 %. The NLCD almost always underestimated IC at low development intensities and overestimated IC at high development intensities. As a result of underestimated IC, regression models using NLCD data predicted mean background concentrations of NO3 (-) and Cl(-) that were 475 and 177 %, respectively, of those predicted when using finer resolution land cover data. Our sampling design could help states and other agencies seeking to create monitoring programs and indicators responsive to anthropogenic impacts. Differences between land cover datasets could affect resource protection due to misguided management targets, watershed development and conservation practices, or water quality criteria.

Urban Stream Burial Increases Watershed-Scale Nitrate Export.

Nitrogen (N) uptake in streams is an important ecosystem service that reduces nutrient loading to downstream ecosystems. Here we synthesize studies that investigated the effects of urban stream burial on N-uptake in two metropolitan areas and use simulation modeling to scale our measurements to the broader watershed scale. We report that nitrate travels on average 18 times farther downstream in buried than in open streams before being removed from the water column, indicating that burial substantially reduces N uptake in streams. Simulation modeling suggests that as burial expands throughout a river network, N uptake rates increase in the remaining open reaches which somewhat offsets reduced N uptake in buried reaches. This is particularly true at low levels of stream burial. At higher levels of stream burial, however, open reaches become rare and cumulative N uptake across all open reaches in the watershed rapidly declines. As a result, watershed-scale N export increases slowly at low levels of stream burial, after which increases in export become more pronounced. Stream burial in the lower, more urbanized portions of the watershed had a greater effect on N export than an equivalent amount of stream burial in the upper watershed. We suggest that stream daylighting (i.e., uncovering buried streams) can increase watershed-scale N retention.

Ecosystem services altered by human changes in the nitrogen cycle: a new perspective for US decision making.

Human alteration of the nitrogen (N) cycle has produced benefits for health and well-being, but excess N has altered many ecosystems and degraded air and water quality. US regulations mandate protection of the environment in terms that directly connect to ecosystem services. Here, we review the science quantifying effects of N on key ecosystem services, and compare the costs of N-related impacts or mitigation using the metric of cost per unit of N. Damage costs to the provision of clean air, reflected by impaired human respiratory health, are well characterized and fairly high (e.g. costs of ozone and particulate damages of $28 per kg NO(x)-N). Damage to services associated with productivity, biodiversity, recreation and clean water are less certain and although generally lower, these costs are quite variable (<$2.2-56 per kg N). In the current Chesapeake Bay restoration effort, for example, the collection of available damage costs clearly exceeds the projected abatement costs to reduce N loads to the Bay ($8-15 per kg N). Explicit consideration and accounting of effects on multiple ecosystem services provides decision-makers an integrated view of N sources, damages and abatement costs to address the significant challenges associated with reducing N pollution.

Ecoenzymatic stoichiometry of microbial organic nutrient acquisition in soil and sediment.

Biota can be described in terms of elemental composition, expressed as an atomic ratio of carbon:nitrogen:phosphorus (refs 1-3). The elemental stoichiometry of microoorganisms is fundamental for understanding the production dynamics and biogeochemical cycles of ecosystems because microbial biomass is the trophic base of detrital food webs. Here we show that heterotrophic microbial communities of diverse composition from terrestrial soils and freshwater sediments share a common functional stoichiometry in relation to organic nutrient acquisition. The activities of four enzymes that catalyse the hydrolysis of assimilable products from the principal environmental sources of C, N and P show similar scaling relationships over several orders of magnitude, with a mean ratio for C:N:P activities near 1:1:1 in all habitats. We suggest that these ecoenzymatic ratios reflect the equilibria between the elemental composition of microbial biomass and detrital organic matter and the efficiencies of microbial nutrient assimilation and growth. Because ecoenzymatic activities intersect the stoichiometric and metabolic theories of ecology, they provide a functional measure of the threshold at which control of community metabolism shifts from nutrient to energy flow.

Analysis of macroinvertebrate assemblages in relation to environmental gradients among lotic habitats of California's Central Valley.

We assessed relationships between environmental characteristics and macroinvertebrate assemblages in lotic habitats of California's Central Valley with community metric and multivariate statistical approaches. Using canonical ordination analyses, we contrasted results when assemblage structure was assessed with macroinvertebrate metrics, as suggested for use in indices of biotic integrity, or with genera abundances. Our objectives were to identify metrics or taxa diagnostic of lotic environmental stressors and compare the capacity of these approaches to detect stressors in order to suggest how they might be used to diagnose stressors. For macroinvertebrate metrics, redundancy analysis (RDA) extracted three axes correlated with channel morphology and substrates. For genera abundances, canonical correspondence analysis (CCA) extracted two axes correlated with soluble salts and with channel morphology and substrates but did not separate these gradients onto different axes. Cluster analyses identified five RDA and five CCA site groups, which exhibited differences for environmental variables, metrics, or genera abundances, and agreement between the analyses in partitioning of sites was greater than if sites were partitioned randomly. These approaches differ in their ability to detect environmental stressors, because they measure different aspects of assemblages and would be complementary in design of new metrics diagnostic of stressors.

Sensitivity of indices of biotic integrity to simulated fish assemblage changes.

Multimetric indices of biotic integrity (IBIs) are commonly used to assess condition of stream fish assemblages, but their ability to monitor trends within streams over time is largely unknown. We assessed the trend detection ability of two IBI formulations (one with traditional scoring and metrics, and one with nontraditional scoring and region-specific metrics) and of similarity and diversity indices using simulations that progressively altered the fish assemblages of 39 streams in the United States mid-Atlantic Highlands region. We also assessed responses to simulated 50% variability in fish abundances, as a measure of background "noise." Fish assemblage indices responded little to changes that affected all species proportionally despite substantial changes in total fish number. Assemblage indices responded better to scenarios that differentially affected fish species, either according to life history traits or by increasing dominance of already common species, but even these changes took some time to detect relative to background variability levels. Ordinations of stream fish assemblage data suggested that differences among sites were maintained even after substantial alterations of fish composition within sites. IBIs are designed to detect broad assemblage differences among sites while downplaying abundance changes and variability increases that were the first indications of within-site changes, and they appear more suited to detecting large departures from natural fish assemblages than for monitoring gradual changes such as those our simulations produced. Inferences about causes of assemblage changes should be made with caution because of correlations among species traits and interdependence among IBI component metrics. Site trend assessments should be made based on all available data rather than just by summary indices.