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The removal of nitrogen (N) in aquatic ecosystems is of great interest because excessive nitrate in groundwater and surface water is a growing problem. High nitrate loading degrades water quality and is linked to eutrophication and harmful algal blooms, especially in coastal marine waters. Past research on nitrate removal processes has emphasized plant or microbial uptake (assimilation) or respiratory denitrification by bacteria. The increasing application of stable isotopes and other tracer techniques to the study of nitrate removal has yielded a growing body of evidence for alternative, microbially mediated processes of nitrate transformation. These include dissimilatory (the reduction of nitrogen into other inorganic compounds, coupled to energy producing processes) reduction of nitrate to ammonium (DNRA), chemoautotrophic denitrification via sulfur or iron oxidation, and anaerobic ammonium oxidation (anammox), as well as abiotic nitrate removal processes. Here, we review evidence for the importance of alternative nitrate removal pathways in aquatic ecosystems and discuss how the possible prevalence of these pathways may alter views of N cycling and its controls. These alternative pathways are of particular importance for the management of excess N in the environment, especially in cases where nitrate is transformed to ammonium, a biologically available and less mobile N form, rather than to dinitrogen gas.

Nitrous oxide (N 2 O) is a potent greenhouse gas that contributes to climate change and stratospheric ozone destruction. Anthropogenic nitrogen (N) loading to river networks is a potentially important source of N 2 O via microbial denitrification that converts N to N 2 O and dinitrogen (N 2 ). The fraction of denitrified N that escapes as N 2 O rather than N 2 (i.e., the N 2 O yield) is an important determinant of how much N 2 O is produced by river networks, but little is known about the N 2 O yield in flowing waters. Here, we present the results of whole-stream 15 N-tracer additions conducted in 72 headwater streams draining multiple land-use types across the United States. We found that stream denitrification produces N 2 O at rates that increase with stream water nitrate (NO 3 − ) concentrations, but that <1% of denitrified N is converted to N 2 O. Unlike some previous studies, we found no relationship between the N 2 O yield and stream water NO 3 − . We suggest that increased stream NO 3 − loading stimulates denitrification and concomitant N 2 O production, but does not increase the N 2 O yield. In our study, most streams were sources of N 2 O to the atmosphere and the highest emission rates were observed in streams draining urban basins. Using a global river network model, we estimate that microbial N transformations (e.g., denitrification and nitrification) convert at least 0.68 Tg·y −1 of anthropogenic N inputs to N 2 O in river networks, equivalent to 10% of the global anthropogenic N 2 O emission rate. This estimate of stream and river N 2 O emissions is three times greater than estimated by the Intergovernmental Panel on Climate Change.H umans have more than doubled the availability of fixed nitrogen (N) in the biosphere, particularly through the production of N fertilizers and the cultivation of N-fixing crops (1). Increasing N availability is producing unintended environmental consequences including enhanced emissions of nitrous oxide (N 2 O), a potent greenhouse gas (2) and an important cause of stratospheric ozone destruction (3). The Intergovernmental Panel on Climate Change (IPCC) estimates that the microbial conversion of agriculturally derived N to N 2 O in soils and aquatic ecosystems is the largest source of anthropogenic N 2 O to the atmosphere (2). The production of N 2 O in agricultural soils has been the focus of intense investigation (i.e., >1,000 published studies) and is a relatively well constrained component of the N 2 O budget (4). However, emissions of anthropogenic N 2 O from streams, rivers, and estuaries have received much less attention and remain a major source of uncertainty in the global anthropogenic N 2 O budget.Microbial denitrification is a large source of N 2 O emissions in terrestrial and aquatic ecosystems. Most microbial denitrification is a form of anaerobic respiration in which nitrate (NO 3 − , the dominant form of inorganic N) is converted to dinitrogen (N 2 ) and N 2 O gases (5). The proportion of denitrified NO 3 − that is converted to N 2 O rather than N 2 (h...

Abstract. Salinization, a widespread threat to the structure and ecological functioning of inland and coastal wetlands, is currently occurring at an unprecedented rate and geographic scale. The causes of salinization are diverse and include alterations to freshwater flows, land-clearance, irrigation, disposal of wastewater effluent, sea level rise, storm surges, and applications of de-icing salts. Climate change and anthropogenic modifications to the hydrologic cycle are expected to further increase the extent and severity of wetland salinization. Salinization alters the fundamental physicochemical nature of the soil-water environment, increasing ionic concentrations and altering chemical equilibria and mineral solubility. Increased concentrations of solutes, especially sulfate, alter the biogeochemical cycling of major elements including carbon, nitrogen, phosphorus, sulfur, iron, and silica. The effects of salinization on wetland biogeochemistry typically include decreased inorganic nitrogen removal (with implications for water quality and climate regulation), decreased carbon storage (with implications for climate regulation and wetland accretion), and increased generation of toxic sulfides (with implications for nutrient cycling and the health/functioning of wetland biota). Indeed, increased salt and sulfide concentrations induce physiological stress in wetland biota and ultimately can result in large shifts in wetland communities and their associated ecosystem functions. The productivity and composition of freshwater species assemblages will be highly altered, and there is a high potential for the disruption of existing interspecific interactions. Although there is a wealth of information on how salinization impacts individual ecosystem components, relatively few studies have addressed the complex and often non-linear feedbacks that determine ecosystem-scale responses or considered how wetland salinization will affect landscape-level processes. Although the salinization of wetlands may be unavoidable in many cases, these systems may also prove to be a fertile testing ground for broader ecological theories including (but not limited to): investigations into alternative stable states and tipping points, trophic cascades, disturbance-recovery processes, and the role of historical events and landscape context in driving community response to disturbance.

1. Rates of whole-system metabolism (production and respiration) are fundamental indicators of ecosystem structure and function. Although first-order, proximal controls are well understood, assessments of the interactions between proximal controls and distal controls, such as land use and geographic region, are lacking. Thus, the influence of land use on stream metabolism across geographic regions is unknown. Further, there is limited understanding of how land use may alter variability in ecosystem metabolism across regions. 2. Stream metabolism was measured in nine streams in each of eight regions (n = 72) across the United States and Puerto Rico. In each region, three streams were selected from a range of three land uses: agriculturally influenced, urban-influenced, and reference streams. Stream metabolism was estimated from diel changes in dissolved oxygen concentrations in each stream reach with correction for reaeration and groundwater input. . In contrast, ecosystem respiration (ER) varied both within and among regions. Reference streams had significantly lower rates of GPP than urban or agriculturally influenced streams. 4. GPP was positively correlated with photosynthetically active radiation and autotrophic biomass. Multiple regression models compared using Akaike's information criterion (AIC) indicated GPP increased with water column ammonium and the fraction of the catchment in urban and reference land-use categories. Multiple regression models also identified velocity, temperature, nitrate, ammonium, dissolved organic carbon, GPP, coarse benthic organic matter, fine benthic organic matter and the fraction of all land-use categories in the catchment as regulators of ER. 5. Structural equation modelling indicated significant distal as well as proximal control pathways including a direct effect of land-use on GPP as well as SRP, DIN, and PAR effects on GPP; GPP effects on autotrophic biomass, organic matter, and ER; and organic matter effects on ER. 6. Overall, consideration of the data separated by land-use categories showed reduced inter-regional variability in rates of metabolism, indicating that the influence of agricultural and urban land use can obscure regional differences in stream metabolism.

Compared to the well-studied open water of the ''growing'' season, under-ice conditions in lakes are characterized by low and rather constant temperature, slow water movements, limited light availability, and reduced exchange with the surrounding landscape. These conditions interact with ice-cover duration to shape microbial processes in temperate lakes and ultimately influence the phenology of community and ecosystem processes. We review the current knowledge on microorganisms in seasonally frozen lakes. Specifically, we highlight how under-ice conditions alter lake physics and the ways that this can affect the distribution and metabolism of auto-and heterotrophic microorganisms. We identify functional traits that we hypothesize are important for understanding under-ice dynamics and discuss how these traits influence species interactions. As ice coverage duration has already been seen to reduce as air temperatures have warmed, the dynamics of the underice microbiome are important for understanding and predicting the dynamics and functioning of seasonally frozen lakes in the near future.The quality of freshwater, which is tightly tied to many essential ecosystem services, is influenced in large part by the activities of microbial communities. In inland waters, as in other ecosystems, microorganisms are at the hub of most biogeochemical processes and largely control ecosystem functioning via their metabolic activities. However, attempts to describe the taxonomy and ecology of the freshwater microbiome mainly involve samples collected during the icefree season and, hence, largely neglect microbial communities and their activities during the ice-covered period. Knowledge about the year-round ecological and biogeochemical traits of typical freshwater microorganisms is required to understand when and where certain microorganisms will appear and how they will influence other organisms or biogeochemical processes and, hence, water quality. Together, this information will improve our ability to predict and model freshwater ecosystem and biogeochemical dynamics and to determine their role in the landscapes in the face of environmental change.A large number of lakes, particularly those situated at high altitude and the numerous high-latitude lakes in the temperate and boreal climate zones, are seasonally covered by ice for more than 40% of the year (Walsh et al. 1998). Despite this, surprisingly little is known about the ecology, diversity, and metabolism of microorganisms that reside under the ice cover in such lakes (Salonen et al. 2009). The traditional view is that ecosystems subjected to low temperatures are ''on hold'' and that cellular adaptations for survival at low temperatures control the composition of the winter microbial community, which awaits environmental conditions more conducive for growth. This traditional concept fails to recognize that the winter season affects the ecology and metabolic features of freshwater microorganisms, as well as their involvement in food webs and global biogeochemical c...

We measured uptake length of 15 NO { 3 in 72 streams in eight regions across the United States and Puerto Rico to develop quantitative predictive models on controls of NO { 3 uptake length. As part of the Lotic Intersite Nitrogen eXperiment II project, we chose nine streams in each region corresponding to natural (reference), suburban-urban, and agricultural land uses. Study streams spanned a range of human land use to maximize variation in NO { 3 concentration, geomorphology, and metabolism. We tested a causal model predicting controls on NO { 3 uptake length using structural equation modeling. The model included concomitant measurements of ecosystem metabolism, hydraulic parameters, and nitrogen concentration. We compared this structural equation model to multiple regression models which included additional biotic, catchment, and riparian variables. The structural equation model explained 79% of the variation in log uptake length (S Wtot ). Uptake length increased with specific discharge (Q/w) and increasing NO

We measured denitrification rates using a field 15 N-NO { 3 tracer-addition approach in a large, cross-site study of nitrate uptake in reference, agricultural, and suburban-urban streams. We measured denitrification rates in 49 of 72 streams studied. Uptake length due to denitrification (S Wden ) ranged from 89 m to 184 km (median of 9050 m) and there were no significant differences among regions or land-use categories, likely because of the wide range of conditions within each region and land use. N 2 production rates far exceeded N 2 O production rates in all streams. The fraction of total NO { 3 removal from water due to denitrification ranged from 0.5% to 100% among streams (median of 16%), and was related to NH z 4 concentration and ecosystem respiration rate (ER). Multivariate approaches showed that the most important factors controlling S Wden were specific discharge (discharge / width) and NO