Denitrification potential of a river floodplain during flooding with nitrate-rich water: grasslands versus reedbeds

Denitrification is a major mechanism for nitrogen removal from nitrogen-rich waters, but it requires oxygen-poor conditions. We assessed denitrification rates in nitrate-rich but also oxygen-rich river water during its stay in a floodplain. We measured diurnal oxygen fluctuations in floodwater along the river Rhine, and carried out an experiment to assess denitrification rates during day, evening and night. Denitrification in floodwater and flooded sediment were measured, comparing activity of periphyton and sediment from agricultural grasslands and reedbeds. Floodwater along the river Rhine was oxygen-saturated (> 10 mg O₂/L) during the day, but oxygen largely disappeared during the night (0.4–0.8 mg O₂/L). Independent of oxygen concentrations, denitrification in surface water alone hardly occurred. In flooded sediments, however, denitrification rates were much higher (1.1–1.5 mg N m^–2 h^–1), particularly at dark and oxygen-poor conditions (nighttime). In the experimental jars, reedbed-periphyton bacteria achieved similar denitrification rates as bacteria in sediment, but overall periphyton denitrification was of minor importance when calculated per square meter. Apart from oxygen levels, maximum denitrification appeared to be regulated by nitrate diffusion from water into the sediment, as the maximum quantity of N denitrified in the sediment equalled the quantity of N lossed from the surface water. Assessed 24-hr denitrification rates in the flooded floodplains (c. 15 mg N m^–2 d^–1) were similar in grasslands and reedbeds, and were rather low compared to rates in other floodplains.     

Denitrification efficiency for defining critical loads of carbon in shallow coastal ecosystems

Denitrification efficiency [DE; (N₂ − N/(DIN + N₂ − N) × 100%)] as an indicator of change associated with nutrient over-enrichment was evaluated for 22 shallow coastal ecosystems in Australia. The rate of carbon decomposition (which can be considered a proxy for carbon loading) is an important control on the efficiency with which coastal sediments in depositional mud basins with low water column nitrate concentrations recycle nitrogen as N₂. The relationship between DE and carbon loading is due to changes in carbon and nitrate (NO₃) supply associated with sediment biocomplexity. At the DE optimum (500–1,000 μmol m⁻² h⁻¹), there is an overlap of aerobic and anaerobic respiration zones (caused primarily by the existence of anaerobic micro-niches within the oxic zone, and oxidized burrow structures penetrating into the anaerobic zone), which enhances denitrification by improving both the organic carbon and nitrate supply to denitrifiers. On either side of the DE optimum zone, there is a reduction in denitrification sites as the sediment loses its three-dimensional complexity. At low organic carbon loadings, a thick oxic zone with low macrofauna biomass exists, resulting in limited anoxic sites for denitrification, and at high carbon loadings, there is a thick anoxic zone and a resultant lack of oxygen for nitrification and associated NO₃ production. We propose a trophic scheme for defining critical (sustainable) carbon loading rates and possible thresholds for shallow coastal ecosystems based on the relationship between denitrification efficiency and carbon loading for 17 of the 22 Australian coastal ecosystems. The denitrification efficiency “optimum” occurs between carbon loadings of about 50 and 100 g C m⁻² year⁻¹. Coastal managers can use this simple trophic scheme to classify the current state of their shallow coastal ecosystems and for determining what carbon loading rate is necessary to achieve any future state. Guest editors: J. H. Andersen & D. J. Conley Eutrophication in Coastal Ecosystems: Selected papers from the Second International Symposium on Research and Management of Eutrophication in Coastal Ecosystems, 20–23 June 2006, Nyborg, Denmark     

Seasonal oxygen,nitrogen and phosphorus benthic cycling along an impacted Baltic Sea estuary: regulation and spatial patterns

The regulatory roles of temperature, eutrophication and oxygen availability on benthic nitrogen (N) cycling and the stoichiometry of regenerated nitrogen and phosphorus (P) were explored along a Baltic Sea estuary affected by treated sewage discharge. Rates of sediment denitrification, anammox, dissimilatory nitrate reduction to ammonium (DNRA), nutrient exchange, oxygen (O₂) uptake and penetration were measured seasonally. Sediments not affected by the nutrient plume released by the sewage treatment plant (STP) showed a strong seasonality in rates of O₂ uptake and coupled nitrification–denitrification, with anammox never accounting for more than 20 % of the total dinitrogen (N₂) production. N cycling in sediments close to the STP was highly dependent on oxygen availability, which masked temperature-related effects. These sediments switched from low N loss and high ammonium (NH₄ ⁺) efflux under hypoxic conditions in the fall, to a major N loss system in the winter when the sediment surface was oxidized. In the fall DNRA outcompeted denitrification as the main nitrate (NO₃ ^?) reduction pathway, resulting in N recycling and potential spreading of eutrophication. A comparison with historical records of nutrient discharge and denitrification indicated that the total N loss in the estuary has been tightly coupled to the total amount of nutrient discharge from the STP. Changes in dissolved inorganic nitrogen (DIN) released from the STP agreed well with variations in sedimentary N₂ removal. This indicates that denitrification and anammox efficiently counterbalance N loading in the estuary across the range of historical and present-day anthropogenic nutrient discharge. Overall low N/P ratios of the regenerated nutrient fluxes impose strong N limitation for the pelagic system and generate a high potential for nuisance cyanobacterial blooms.     

Modeling nitrogen cycling in a coastal fresh water sediment

Increased nitrogen (N) loading to coastal marine and freshwater systems is occurring worldwide as a result of human activities. Diagenetic processes in sediments can change the N availability in these systems, by supporting removal through denitrification and burial of organic N (N_org) or by enhancing N recycling. In this study, we use a reactive transport model (RTM) to examine N transformations in a coastal fresh water sediment and quantify N removal rates. We also assess the response of the sediment N cycle to environmental changes that may result from increased salinity which is planned to occur at the site as a result of an estuarine restoration project. Field results show that much of the N_org deposited on the sediment is currently remineralized to ammonium. A rapid removal of nitrate is observed in the sediment pore water, with the resulting nitrate reduction rate estimated to be 130 μmol N cm⁻² yr⁻¹. A model sensitivity study was conducted altering the distribution of nitrate reduction between dissimilatory nitrate reduction to ammonium (DNRA) and denitrification. These results show a 40% decline in sediment N removal as NO ₃ ⁻ reduction shifts from denitrification to DNRA. This decreased N removal leads to a shift in sediment-water exchange flux of dissolved inorganic nitrogen (DIN) from near zero with denitrification to 133 μmol N cm⁻² yr⁻¹ if DNRA is the dominant pathway. The response to salinization includes a short-term release of adsorbed ammonium. Additional changes expected to result from the estuarine restoration include: lower NO ₃ ⁻ concentrations and greater SO ₄ ²⁻ concentrations in the bottom water, decreased nitrification rates, and increased sediment mixing. The effect of these changes on net DIN flux and N removal vary based on the distribution of DNRA versus denitrification, illustrating the need for a better understanding of factors controlling this competition.     

Linkages between organic matter mineralization and denitrification in eight riparian wetlands

Denitrification (N₂ production) and oxygen consumption rates were measured at ambient field nitrate concentrations during summer in sediments from eight wetlands (mixed hardwood swamps, cedar swamps, heath dominated shrub wetland, herbaceous peatland, and a wetland lacking live vegetation) and two streams. The study sites included wetlands in undisturbed watersheds and in watersheds with considerable agricultural and/or sewage treatment effluent input. Denitrification rates measured in intact cores of water-saturated sediment ranged from 20 to 260 mol N m^-2 h^-1 among the three undisturbed wetlands and were less variable (180 to 260 mol N M^-2 h^-1) among the four disturbed wetlands. Denitrification rates increased when nitrate concentrations in the overlying water were increased experimentally (1 up to 770 M), indicating that nitrate was an important factor controlling denitrification rates. However, rates of nitrate uptake from the overlying water were not a good predictor of denitrification rates because nitrification in the sediments also supplied nitrate for denitrification. Regardless of the dominant vegetation, pH, or degree of disturbance, denitrification rates were best correlated with sediment oxygen consumption rates (r ² = 0.912) indicating a relationship between denitrification and organic matter mineralization and/or sediment nitrification rates. Rates of denitrification in the wetland sediments were similar to those in adjacent stream sediments. Rates of denitrification in these wetlands were within the range of rates previously reported for water-saturated wetland sediments and flooded soils using whole core¹⁵N techniques that quantify coupled nitrification/denitrification, and were higher than rates reported from aerobic (non-saturated) wetland sediments using acetylene block methods.     

Nitrogen transformations at the sediment–water interface across redox gradients in the Laurentian Great Lakes

The capacity of a lake to remove reactive nitrogen (N) through denitrification has important implications both for the lake and for downstream ecosystems. In large oligotropic lakes such as Lake Superior, where nitrate (NO₃ ^?) concentrations have increased steadily over the past century, deep oxygen penetration into sediments may limit the denitrification rates. We tested the hypothesis that the position of the redox gradient in lake sediments affects denitrification by measuring net N-fluxes across the sediment–water interface for intact sediment cores collected across a range of sediment oxycline values from nearshore and offshore sites in Lake Superior, as well as sites in Lake Huron and Lake Erie. Across this redox gradient, as the thickness of the oxygenated sediment layer increased from Lake Erie to Lake Superior, fluxes of NH₄ ⁺ and N₂ out of the sediment decreased, and sediments shifted from a net sink to a net source of NO₃ ^?. Denitrification of NO₃ ^? from overlying water decreased with thickness of the oxygenated sediment layer. Our results indicate that, unlike sediments from Lake Erie and Lake Huron, Lake Superior sediments do not remove significant amounts of water column NO₃ ^? through denitrification, likely as a result of the thick oxygenated sediment layer.     

Denitrification in Aquatic Environments: A Cross-system Analysis

A meta-analysis was conducted on 136 data sets of denitrification rates (DR) recorded both during the period of highest water temperature and monthly in five types of aquatic ecosystems: oceans, coastal environments, estuaries, lakes and rivers. There was a gradual increase of DR from the ocean to rivers and lakes at both scales, with the rivers showing the highest DR variability. Denitrification peaked during summertime and showed highest seasonal variability in lakes and rivers. High concentrations of nitrate and interstitially-dissolved organic carbon as well as low oxygen concentration in the overlying water enhanced DR both during summer and at a seasonal scale whereas total phosphorus did at the seasonal scale only. There was a positive linear relationship between overlying nitrate and DR over the range of 1–970 μmol NO₃ (r ² = 0.86, P = 0.001). DR in lakes and rivers might reach values doubling those in the more denitrifying terrestrial ecosystems (e.g. agrosystems). Discrepancies in DR and its controlling factors between site-specific studies and this meta-analysis may arise from environmental variability at two, often confounded, scales of observation: the habitat and the ecosystem level. Future studies on denitrification in aquatic environments should address the topic of spatial heterogeneity more thoroughly.     

The Fate of 15N-Nitrate in Healthy and Declining Phragmites australis Stands

Abstract The dissimilatory nitrate-reducing processes, denitrification, and dissimilatory nitrate-reduction to ammonium were studied in freshwater lake sediments within healthy and degrading Phragmites australis (reed) stands. The samples from the healthy vegetation site contained roots and rhizomes. Cores were supplied with 1.9–5.2 μg ¹⁵N-NO₃ ⁻ g⁻¹ dry sediment in the laboratory and subsequently incubated for 8 h at 20°C, in the dark. The ¹⁵N compounds were determined before (natural percentage of ¹⁵N) and after 1 and 8 h of incubation. The uptake of ¹⁵N by the roots and rhizomes in the healthy vegetation was 61%. Nitrogen losses, interpreted as denitrification, accounted for 25 and 84% of the added ¹⁵N-NO₃ ⁻ in sediment from the healthy and degrading vegetation sites, respectively. The percentages of nitrate reduced to ammonium were 4 and 9% in sediment from the healthy vegetation and degrading vegetation sites, respectively. The percentage of ¹⁵N–total N in the sediment of the healthy vegetation site was 10%, whereas for the degrading vegetation site this percentage was 7%. The percentage of nitrate reduced to ammonium could be potentially underestimated by the percentage of ¹⁵N measured in the sediment. In this case, in healthy and degenerating P. australis stands, the percentage of produced ammonium accounted for 14–16%. The nitrate reduction rates were calculated based on an incubation period of one hour. The denitrification rate in sediment from the degrading vegetation site was higher than from the healthy vegetation site. The rate of dissimilatory nitrate reduction to ammonium was almost tenfold higher in sediment from the degrading vegetation site compared to sediment from the healthy vegetation site. The significantly lower percentages of dissimilatory nitrate reduction to ammonium and denitrification in the healthy stand compared to the degrading stand was probably due to the presence of roots and rhizomes. In the sediments of healthy and degrading P. australis stands, denitrification was the main nitrate-reducing process. Received: 24 July 1996; Accepted: 5 December 1996     

Nitrogen dynamics at the sediment–water interface in shallow, sub-tropical Florida Bay: why denitrification efficiency may decrease with increased eutrophication

Nitrogen (N) dynamics at the sediment–water interface were examined in four regions of Florida Bay to provide mechanistic information on the fate and effects of increased N inputs to shallow, subtropical, coastal environments. Dissimilatory nitrate (NO₃ ^?) reduction to ammonium (DNRA) was hypothesized to be a significant mechanism retaining bioreactive N in this warm, saline coastal ecosystem. Nitrogen dynamics, phosphorus (P) fluxes, and sediment oxygen demand (SOD) were measured in north-central (Rankin Key; eutrophic), north-eastern (Duck Key; high N to P seston ratios), north-western (Murray Key; low N to P ratios), and central (Rabbit Key; typical central site) Florida Bay in August 2004, January 2005, and November 2006. Site water was passed over intact sediment cores, and changes in oxygen (O₂), phosphate (o-PO₄ ^3?), ammonium (NH₄ ⁺), NO₃ ^?, nitrite (NO₂ ^?), and N₂ concentrations were measured, without and with addition of excess ¹⁵NO₃ ^? or ¹⁵NH₄ ⁺ to inflow water. These incubations provided estimates of SOD, nutrient fluxes, N₂ production, and potential DNRA rates. Denitrification rates were lowest in summer, when SOD was highest. DNRA rates and NH₄ ⁺ fluxes were high in summer at the eutrophic Rankin site, when denitrification rates were low and almost no N₂ came from added ¹⁵NO₃ ^?. Highest ¹⁵NH₄ ⁺ accumulation, resulting from DNRA, occurred at Rabbit Key during a picocyanobacteria bloom in November. ¹⁵NH₄ ⁺ accumulation rates among the stations correlated with SOD in August and January, but not in November during the algal bloom. These mechanistic results help explain why bioreactive N supply rates are sometimes high in Florida Bay and why denitrification efficiency may decrease with increased NO₃ ^? inputs in sub-tropical coastal environments.     

Co-occurrence of in-stream nitrogen fixation and denitrification across a nitrogen gradient in a western U.S. watershed

It is frequently assumed that nitrogen (N₂) fixation and denitrification do not co-occur in streams because each process should be favored under different concentrations of dissolved inorganic nitrogen (DIN), and therefore these processes are rarely quantified together. We asked if these processes could co-exist by conducting a spatial survey of N₂ fixation using acetylene reduction and denitrification using acetylene block [with and without amendments of carbon (C) as glucose and nitrogen (N) as nitrate]. Rates were measured on rocks and sediment in 8 southeastern Idaho streams encompassing a DIN gradient of 26–615 µg L^?1. Sampling at each site was repeated in summer 2015 and 2016. We found that both denitrification and N₂ fixation occurred across the gradient of DIN concentrations, with N₂ fixation occurring primarily on rocks and denitrification occurring in sediment. N₂ fixation rates on rocks significantly decreased 100× across the DIN gradient in 1 year of the study, and amended (with N and C) denitrification rates increased 10× across the DIN gradient in both years. Multiple linear regression and partial least squares models with environmental characteristics measured at the scale of entire stream reaches showed that C and phosphorus were positive predictors of amended and unamended denitrification rates, but no significant model could explain N₂ fixation rates across all streams and years. This, coupled with the observation that detectable rates of N₂ fixation occurred primarily on rocks and denitrification occurred primarily on sediment, suggests that microhabitat scale factors may better predict the co-occurrence of these processes within stream reaches. Overlooking the potential co-occurrence of N₂ fixation and denitrification in stream ecosystems will impede understanding by oversimplifying the contribution of each process to the N cycle.     

A meta‐analysis of soil salinization effects on nitrogen pools,cycles and fluxes in coastal ecosystems

Salinity intrusion caused by land subsidence resulting from increasing groundwater abstraction, decreasing river sediment loads and increasing sea level because of climate change has caused widespread soil salinization in coastal ecosystems. Soil salinization may greatly alter nitrogen (N) cycling in coastal ecosystems. However, a comprehensive understanding of the effects of soil salinization on ecosystem N pools, cycling processes and fluxes is not available for coastal ecosystems. Therefore, we compiled data from 551 observations from 21 peer‐reviewed papers and conducted a meta‐analysis of experimental soil salinization effects on 19 variables related to N pools, cycling processes and fluxes in coastal ecosystems. Our results showed that the effects of soil salinization varied across different ecosystem types and salinity levels. Soil salinization increased plant N content (18%), soil NH₄⁺ (12%) and soil total N (210%), although it decreased soil NO₃^? (2%) and soil microbial biomass N (74%). Increasing soil salinity stimulated soil N₂O fluxes as well as hydrological NH₄⁺ and NO₂^? fluxes more than threefold, although it decreased the hydrological dissolved organic nitrogen (DON) flux (59%). Soil salinization also increased the net N mineralization by 70%, although salinization effects were not observed on the net nitrification, denitrification and dissimilatory nitrate reduction to ammonium in this meta‐analysis. Overall, this meta‐analysis improves our understanding of the responses of ecosystem N cycling to soil salinization, identifies knowledge gaps and highlights the urgent need for studies on the effects of soil salinization on coastal agro‐ecosystem and microbial N immobilization. Additional increases in knowledge are critical for designing sustainable adaptation measures to the predicted intrusion of salinity intrusion so that the productivity of coastal agro‐ecosystems can be maintained or improved and the N losses and pollution of the natural environment can be minimized.     

Microbial Nitrogen Cycling Processes in a Sulfidic Coastal Marsh

Sulfide distribution is a key controller of vegetation zonation in coastal ecosystems, but data are limited regarding its impact on the spatial distribution of important N cycling processes. We assessed vegetation distribution and density and, mineral N pool sizes, composition and transformations in a sulfidic coastal marsh in relation to distance from sulfur springs. We observed strong relationships between vegetation attributes (species and density) and mineral N status with greater total inorganic N, NO₃⁻ and denitrification enzyme activity (DEA) in sediment samples from areas populated by Crithmum maritimum (mid-way between S springs and sea shore) than in sediments from areas colonized by either Agropyron repens (closest to the S springs) or mangrove (Rhizophora mangleL., farthest from the springs). Our data also suggest close links between N cycling and SO₄⁻² reduction. The latter resulted in net release of NH₄⁺ ranging from 0.9 mg N kg⁻¹ in the low density C. maritimum to 3.2 mg N kg⁻¹ in the high-density A. repens, during a 3-day incubation. We also tested for microbial adaptation to long-term high sulfide exposure by measuring DEA using the C₂H₂ block method (which has been found to be strongly affected by the presence of sulfide) and amendment of marsh sediment samples with NaMoO₄ to suppress reduced S production. In sediments extracted from sites near the sulfur springs (A. repens and C. maritimum), the C₂H₂ blockage assay yielded similar results without and with NaMoO₄ addition. However, in samples from a mangrove located further downstream from the springs, DEA was substantially lower (2.3 vs. 6.8 mg N₂O-N kg⁻¹ sediment d⁻¹) when production of reduced S was not inhibited by NaMoO₄. These results suggest that denitrifying microbes in the high sulfide areas may have adapted to the presence of sulfide, allowing for high rates of N and S cycling to occur simultaneously in these marshes.     

Denitrification Rates in a Marine Sediment as Measured by the Acetylene Inhibition Technique

A method has been developed for measurement of denitrification activity in sediments by application of the acetylene inhibition technique. Acetylene-saturated water was injected, at close intervals, into sediment cores which were then incubated for a few hours at the in situ temperature. Frozen segments of the cores were assayed for accumulation of N₂O by a combined gas extraction and detection system. The segments were thawed under a stream of helium from which N₂O (and other gases) was collected in a liquid N₂ trap, and the quantity of N₂O was measured by gas chromatography. The maximum rate of denitrification in a coastal marine sediment was 35 nmol of N per cm³ of sediment per day at 2.5°C, and the rate of denitrification for the total sediment was 0.99 nmol of N per m² per day.     

Tracing sources and pathways of dissolved nitrate in forest and river ecosystems using high-resolution isotopic techniques: a review

Nitrogen inputs into stream and river ecosystems, and the factors influencing those inputs, are important for various ecological and environmental concerns. Reliable information on where and how nitrogen compounds flow into aquatic ecosystems is indispensable to understanding the nutrient status of these ecosystems. Such information should include the biogeochemical mechanisms and hydrological controls of nutrient leaching into rivers from terrestrial systems such as forests, agricultural fields, and urbanized areas. Advancements in stable isotopomer measurements over the past two decades have expanded the variety of target substances and the precision with which they can be investigated. The high-throughput microbial denitrifier method allows for simultaneous measurement of nitrogen and oxygen isotope ratios and can provide high-resolution spatiotemporal information on both nitrate sources and biogeochemical processes. Although advanced techniques of stable isotope analysis have been used extensively to detect sources and estimate the relative contributions of multi-source systems in various rivers, there are still new horizons in investigating nitrogen transformations. For example, stable isotopes of oxygen (¹⁸O and ¹⁷O) occurring in nitrate due to atmospheric deposition can be used as natural tracers for evaluating internal nitrogen cycling; these isotopes are distinct from the oxygen within microbially generated nitrate in soils and water bodies. Another future challenge is improved use of nitrous oxide isotopomers in evaluating the relative contributions of nitrification and denitrification. Such analysis could provide insight into the nitrogen transformation that occurs under redox conditions at the boundary between terrestrial and aquatic systems, where nitrification and denitrification often occur simultaneously in soil and aquatic environments.     

Influence of hydrological connectivity of riverine wetlands on nitrogen removal via denitrification

Wetland ecosystems in agricultural areas often become progressively more isolated from main water bodies. Stagnation favors the accumulation of organic matter as the supply of electron acceptors with water renewal is limited. In this context it is expected that nitrogen recycling prevails over nitrogen dissipation. To test this hypothesis, denitrification rates, fluxes of dissolved oxygen (SOD), inorganic carbon (DIC) and nitrogen and sediment features were measured in winter and summer 2007 on 22 shallow riverine wetlands in the Po River Plain (Northern Italy). Fluxes were determined from incubations of intact cores by measurement of concentration changes or isotope pairing in the case of denitrification. Sampled sites were eutrophic to hypertrophic; 10 were connected and 12 were isolated from the adjacent rivers, resulting in large differences in nitrate concentrations in the water column (from <5 to 1,133 μM). Benthic metabolism and denitrification rates were investigated by two overarching factors: season and hydrological connectivity. SOD and DIC fluxes resulted in respiratory quotients greater than one at most sampling sites. Sediment respiration was coupled to both ammonium efflux, which increased from winter to summer, and nitrate consumption, with higher rates in river-connected wetlands. Denitrification rates measured in river-connected wetlands (35–1,888 μmol N m^?2 h^?1) were up to two orders of magnitude higher than rates measured in isolated wetlands (2–231 μmol N m^?2 h^?1), suggesting a strong regulation of the process by nitrate availability. These rates were also significantly higher in summer (9–1,888 μmol N m^?2 h^?1) than in winter (2–365 μmol N m^?2 h^?1). Denitrification supported by water column nitrate (D_W) accounted for 60–100% of total denitrification (Dtot); denitrification coupled to nitrification (D_N) was probably controlled by limited oxygen availability within sediments. Denitrification efficiency, calculated as the ratio between N removal via denitrification and N regeneration, and the relative role of denitrification for organic matter oxidation, were high in connected wetlands but not in isolated sites. This study confirms the importance of restoring hydraulic connectivity of riverine wetlands for the maintenance of important biogeochemical functions such as nitrogen removal via denitrification.     

Woody Vegetation Removal Stimulates Riparian and Benthic Denitrification in Tallgrass Prairie

Expansion of woody vegetation into areas that were historically grass-dominated is a significant contemporary threat to grasslands, including native tallgrass prairie ecosystems of the Midwestern United States. In tallgrass prairie, much of this woody expansion is concentrated in riparian zones with potential impacts on biogeochemical processes there. Although the effects of woody riparian vegetation on denitrification in both riparian soils and streams have been well studied in naturally wooded ecosystems, less is known about the impacts of woody vegetation encroachment in ecosystems that were historically dominated by herbaceous vegetation. Here, we analyze the effect of afforestation and subsequent woody plant removal on riparian and benthic denitrification. Denitrification rates in riparian soil and selected benthic compartments were measured seasonally in naturally grass-dominated riparian zones, woody encroached riparian zones, and riparian zones with woody vegetation removed in two separate watersheds. Riparian soil denitrification was highly seasonal, with the greatest rates in early spring. Benthic denitrification also exhibited high temporal variability, but no seasonality. Soil denitrification rates were greatest in riparian zones where woody vegetation was removed. Additionally, concentrations of nitrate, carbon, and soil moisture (indicative of potential anoxia) were greatest in wood removal soils. Differences in the presence and abundance of benthic compartments reflected riparian vegetation, and may have indirectly affected denitrification in streams. Riparian soil denitrification increased with soil water content and NO₃ ^?. Management of tallgrass prairies that includes removal of woody vegetation encroaching on riparian areas may alter biogeochemical cycling by increasing nitrogen removed via denitrification while the restored riparian zones return to a natural grass-dominated state.     

Sediment Nitrification, Denitrification, and Nitrous Oxide Production in a Deep Arctic Lake

We used a combination of ¹⁵N tracer methods and a C₂H₂ blockage technique to determine the role of sediment nitrification and denitrification in a deep oligotrophic arctic lake. Inorganic nitrogen concentrations ranged between 40 and 600 nmol · cm⁻³, increasing with depth below the sediment-water interface. Nitrate concentrations were at least 10 times lower, and nitrate was only detectable within the top 0 to 6 cm of sediment. Eh and pH profiles showed an oxidized surface zone underlain by more reduced conditions. The lake water never became anoxic. Sediment Eh values ranged from −7 to 484 mV, decreasing with depth, whereas pH ranged from 6.0 to 7.3, usually increasing with depth. The average nitrification rate (49 ng of N · cm⁻³ · day⁻¹) was similar to the average denitrification rate (44 ng of N · cm⁻³ · day⁻¹). In situ N₂O production from nitrification and denitrification ranged from 0 to 25 ng of N · cm⁻³ · day⁻¹. Denitrification appears to depend on the supply of nitrate by nitrification, such that the two processes are coupled functionally in this sediment system. However, the low rates result in only a small nitrogen loss.     

Potential rates of nitrification and denitrification in an oligotrophic freshwater sediment system

Potential rates of nitrification and denitrification were measured in an oligotrophic sediment system. Nitrification potential was estimated using the CO oxidation technique, and potential denitrification was measured by the acetylene blockage technique. The sediments demonstrated both nitrifying and denitrifying activity. E_h, O₂, and organic C profiles showed two distinct types of sediment. One type was low in organic C, had high O₂ and E_h, and had rates of denitrification 1,000 times lower than the other which had high organic C, low O₂, and low E_h. Potential nitrification and denitrification rates were negatively correlated with E_h. This suggests that environmental heterogeneity in denitrifier and nitrifier populations in oligotrophic sediment systems may be assessed using E_h before sampling protocols for nitrification or denitrification rates are established. There was no correlation between denitrification and nitrification rates or between either of these processes and NH₄ ⁺ or NO₃ ^– concentrations. The maximum rate of denitrification was 0.969 nmole N cm^–3 hour^–1, and the maximum rate of nitrification was 23.6 nmole cm^–3 hour^–1, suggesting nitrification does not limit denitrification in these oligotrophic sediments. Some sediment cores had mean concentrations of 6.0 mg O₂/liter and still showed both nitrification and denitrification activity.     

Denitrification kinetics and denitrifier abundances in sediments of lakes receiving atmospheric nitrogen deposition (Colorado, USA)

The transport and deposition of anthropogenic nitrogen (N) to downwind ecosystems is significant and can be a dominant source of new N to many watersheds. Bacterially mediated denitrification in lake sediments may ameliorate the effects of N loading by permanently removing such inputs. We measured denitrification in sediments collected from lakes in the Colorado Rocky Mountains (USA) receiving elevated (5–8?kg?N?ha^?1?y^?1) or low (<2?kg?N?ha^?1?y^?1) inputs of atmospheric N deposition. The nitrate (NO₃ ^?) concentration was significantly greater in high-deposition lakes (11.3?μmol?l^?1) compared to low-deposition lakes (3.3?μmol?l^?1). Background denitrification was positively related to NO₃ ^? concentrations and we estimate that the sampled lakes are capable of removing a significant portion of N inputs via sediment denitrification. We also conducted a dose–response experiment to determine whether chronic N loading has altered sediment denitrification capacity. Under Michaelis–Menten kinetics, the maximum denitrification rate and half-saturation NO₃ ^? concentration did not differ between deposition regions and were 765?μmol?N?m^?2?h^?1 and 293?μmol?l^?1?NO₃ ^?, respectively, for all lakes. We enumerated the abundances of nitrate- and nitrite-reducing bacteria and found no difference between high- and low-deposition lakes. The abundance of these bacteria was related to available light and bulk sediment resources. Our findings support a growing body of evidence that lakes play an important role in N removal and, furthermore, suggest that current levels of N deposition have not altered the abundance of denitrifying bacteria or saturated the capacity for sediment denitrification.     

Seasonal effects of zebra mussels on littoral nitrogen transformation rates in Gull Lake, Michigan, U.S.A.

• 1 Zebra mussels (Dreissena polymorpha) are successful colonisers of lake littoral habitats and they interact strongly with littoral benthos. Previous research suggests that localised areas colonised by zebra mussels may be hotspots of nitrogen (N) cycling.
• 2 The effects of zebra mussels on nitrification and denitrification rates were examined approximately every other month for 1 year in Gull Lake, Michigan, U.S.A. Littoral sediment was collected from an area free of zebra mussels and distributed into shallow trays; rocks colonised with zebra mussels were placed in half of the trays, while uncolonised rocks were placed in the remaining trays. After an incubation period of 6–8 weeks in the lake, sediment and zebra mussels were collected from the trays, replaced with new sediment and zebra mussels, and placed in the lake for the next interval. In the laboratory, sediment nitrification and denitrification rates were measured for each tray.
• 3 Sediment nitrification rates did not increase in the presence of zebra mussels; instead nitrification rates were sensitive to changes in water temperature and increased with increasing exchangeable sediment ammonium. In contrast, denitrification rates increased in sediment trays with zebra mussels in the winter when nitrate (NO₃⁻) availability was high and when Chara did not grow in the trays.
• 4 Sediment denitrification was NO₃⁻‐limited in all seasons, regardless of zebra mussel treatment. However, sediment in the presence of zebra mussels responded less to NO₃⁻ addition, suggesting that NO₃⁻ limitation of denitrification can be reduced by zebra mussel activity. Zebra mussels have a seasonally variable impact on sediment denitrification rates, and this may translate into altered seasonal patterns of N cycling in localised areas of lakes where they are particularly abundant.