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.     

Simultaneous occurrence of denitrification and nitrate ammonification in sediments of the French Mediterranean Coast

Dissimilatory nitrate reductions in coastal marine sediment of Carteau Cove (French Mediterranean Coast) were studied between April 1993 and July 1994. Simultaneous determination of denitrification and dissimilatory nitrate reduction to ammonium was achieved by using a combination of acetylene blockage and 15N techniques. After short incubations (maximum 5 h), a part of 15N labelled nitrate added to the sediment was recovered as ammonium without incorporation in organic matter. The result indicate that a fraction of nitrate was reduced to ammonium by a dissimilatory mechanism instead of denitrifying. Denitrifying and nitrate ammonifying activities ranged from 0 to 19.8 μmol l-1 d-1 and from 2.3 to 83.2 μmol l-1 d-1, respectively. Denitrification rates were highest in early spring whereas nitrate ammonification were highest in fall. The recovery of nitrate reduced as N2O-N plus ammonium was between 40 and 100%, the highest nitrogen losses were recorded in July. Depending on the station and time of year denitrification accounted for between 0 and 43% of the total nitrate reduction whereas dissimilatory nitrate reduction to ammonium (DNRA) accounted for between 18 and 100%. The reduction rate data suggest that the pathway of nitrate reduction to ammonium may be important in coastal sediments. This revised version was published online in July 2006 with corrections to the Cover Date.     

Denitrification of a landfill leachate with high nitrate concentration in an anoxic rotating biological contactor

The denitrification performance of a lab-scale anoxic rotating biological contactor (RBC) using landfill leachate with high nitrate concentration was evaluated. Under a carbon to nitrogen ratio (C/N) of 2, the reactor achieved N-NO₃ ⁻ removal efficiencies above 95% for concentrations up to 100 mg N-NO₃ ⁻ l⁻¹. The highest observed denitrification rate was 55 mg N-NO₃ ⁻ l⁻¹ h⁻¹ (15 g N-NO₃ ⁻ m⁻² d⁻¹) at a nitrate concentration of 560 mg N-NO₃ ⁻ l⁻¹. Although the reactor has revealed a very good performance in terms of denitrification, effluent chemical oxygen demand (COD) concentrations were still high for direct discharge. The results obtained in a subsequent experiment at constant nitrate concentration (220 mg N-NO₃ ⁻ l⁻¹) and lower C/N ratios (1.2 and 1.5) evidenced that the organic matter present in the leachate was non-biodegradable. A phosphorus concentration of 10 mg P-PO₄ ³⁻ l⁻¹ promoted autotrophic denitrification, revealing the importance of phosphorus concentration on biological denitrification processes.     

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.     

Identification of nitrogen sources and transformations within karst springs using isotope tracers of nitrogen

Isotope analyses of nitrate and algae were used to gain better understanding of sources of nitrate to Florida’s karst springs and processes affecting nitrate in the Floridan aquifer at multiple scales. In wet years, δ¹⁵N and δ¹⁸O of nitrate ranged from +3 to +9‰ in headwater springs in north Florida, indicating nitrification of soil ammonium as the dominant source. With below normal rainfall, the δ¹⁵N and δ¹⁸O of nitrate were higher in almost all springs (reaching +20.2 and +15.3‰, respectively) and were negatively correlated with dissolved oxygen. In springs with values of δ¹⁵N-NO₃ and δ¹⁸O-NO₃ greater than +10‰, nitrate concentrations declined 40–50% in dry years and variations in the δ¹⁵N and δ¹⁸O of nitrate were consistent with the effects of denitrification. Modeling of the aquifer as a closed system yielded in situ fractionation caused by denitrification of 9 and 18‰ for Δ¹⁸O and Δ¹⁵N, respectively. We observed no strong evidence for local sources of nitrate along spring runs; concentrations declined downstream (0.42–3.3?μmol-NO₃ L^?1 per km) and the isotopic dynamics of algae and nitrate indicated a closed system. Correlation between the δ¹⁵N composition of nitrate and algae was observed at regional and spring-run scales, but the relationship was complicated by varying isotopic fractionation factors associated with nitrate uptake (Δ ranged from 2 to 13‰). Our study demonstrates that nitrate inputs to Florida’s springs are derived predominantly from non-point sources and that denitrification is detectable in aquifer waters with relatively long residence time (i.e., matrix flow).     

Assessing Nitrification and Denitrification in the Seine River and Estuary Using Chemical and Isotopic Techniques

Downstream from metropolitan Paris (France), a large amount of ammonium is discharged into the Seine River by the effluents of the wastewater treatment plant at Achères. To assess the extent of nitrification and denitrification in the water column, concentrations and isotopic compositions of ammonium (δ¹⁵N–NH₄⁺) and nitrate (δ¹⁵N–NO₃⁻, δ¹⁸O–NO₃⁻) were measured during summer low-flow conditions along the lower Seine and its estuary. The results indicated that most of the ammonium released from the wastewater treatment plant is nitrified in the lower Seine River and its upper estuary, but there was no evidence for water-column denitrification. In the lower part of the estuary, however, concentration and isotopic data for nitrate were not consistent with simple mixing between riverine and marine nitrate. A significant departure of the nitrate isotopic composition from what would be expected from simple mixing of freshwater and marine nitrates suggested coupled nitrification and denitrification in the water, in spite of the apparent conservative behavior of nitrate. Denitrification rates of approximately 0.02 mg N/L/h were estimated for this part of the estuary.     

Patterns and drivers of anaerobic nitrogen transformations in sediments of thermokarst lakes

Significant attention has been given to the way in which the soil nitrogen (N) cycle responds to permafrost thaw in recent years, yet little is known about anaerobic N transformations in thermokarst lakes, which account for more than one-third of thermokarst landforms across permafrost regions. Based on the N isotope dilution and tracing technique, combined with qPCR and high-throughput sequencing, we presented large-scale measurements of anaerobic N transformations of sediments across 30 thermokarst lakes over the Tibetan alpine permafrost region. Our results showed that gross N mineralization, ammonium immobilization, and dissimilatory nitrate reduction rates in thermokarst lakes were higher in the eastern part of our study area than in the west. Denitrification dominated in the dissimilatory nitrate reduction processes, being two and one orders of magnitude higher than anaerobic ammonium oxidation (anammox) and dissimilatory nitrate reduction to ammonium (DNRA), respectively. The abundances of the dissimilatory nitrate reduction genes (nirK, nirS, hzsB, and nrfA) exhibited patterns consistent with sediment N transformation rates, while α diversity did not. The inter-lake variability in gross N mineralization and ammonium immobilization was dominantly driven by microbial biomass, while the variability in anammox and DNRA was driven by substrate supply and organic carbon content, respectively. Denitrification was jointly affected by nirS abundance and organic carbon content. Overall, the patterns and drivers of anaerobic N transformation rates detected in this study provide a new perspective on potential N release, retention, and removal upon the formation and development of thermokarst lakes.     

The denitrification capacity of sediment from a hypereutrophic lake

SUMMARY.

• 1 In sediment from Wintergreen Lake, Michigan, denitrification was not detectable by the acetylene inhibition method at in situ nitrate concentrations. When nitrate was added to sediment slurries, denitrification capacities up to 18.8μg N g^-1 h^-1 were measured. The denitrification capacities decreased with increasing sediment depth and distance from shore.
• 2 The high denitrification capacities in these sediments which under natural conditions had no supply of nitrate and oxygen suggested that denitrifies with alternative mechanisms for anaerobic energy conversion were present. Nitrous oxide was a significant portion of the N-gas produced immediately after the nitrate addition. Small amounts (4–5% of the total N-gas production) of nitric oxide accumulated in the early phase of nitrate reduction. Presumably after depletion of nitrate and nitrite both N₂O and NO were further reduced to N₂.
• 3 About 70%r of the added nitrate was denitrified, and the remainder was assumed to have been reduced to ammonium.

     

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.     

Denitrification,dissimilatory nitrate reduction to ammonium,and nitrogen fixation along a nitrate concentration gradient in a created freshwater wetland

Wetlands are often highly effective nitrogen (N) sinks. In the Lake Waco Wetland (LWW), near Waco, Texas, USA, nitrate (NO₃⁻) concentrations are reduced by more than 90% in the first 500 m downstream of the inflow, creating a distinct gradient in NO₃⁻ concentration along the flow path of water. The relative importance of sediment denitrification (DNF), dissimilatory NO₃⁻ reduction to ammonium (DNRA), and N₂ fixation were examined along the NO₃⁻ concentration gradient in the LWW. “Potential DNF” (hereafter potDNF) was observed in all months and ranged from 54 to 278 μmol N m⁻² h⁻¹. “Potential DNRA” (hereafter potDNRA) was observed only in summer months and ranged from 1.3 to 33 μmol N m⁻² h⁻¹. Net N₂ flux ranged from 184 (net denitrification) to −270 (net N₂ fixation) μmol N m⁻² h⁻¹. Nitrogen fixation was variable, ranging from 0 to 426 μmol N m⁻² h⁻¹, but high rates ranked among the highest reported for aquatic sediments. On average, summer potDNRA comprised only 5% (±2% SE) of total NO₃⁻ loss through dissimilatory pathways, but was as high as 36% at one site where potDNF was consistently low. Potential DNRA was higher in sediments with higher sediment oxygen demand (r ² = 0.84), and was related to NO₃⁻ concentration in overlying water in one summer (r ² = 0.81). Sediments were a NO₃⁻ sink and accounted for 50% of wetland NO₃⁻ removal (r ² = 0.90). Sediments were an NH₄⁺ source, but the wetland was often a net NH₄⁺ sink. Although DNRA rates in freshwater wetlands may rival those observed in estuarine systems, the importance of DNRA in freshwater sediments appears to be minor relative to DNF. Furthermore, sediment N₂ fixation can be extremely high when NO₃⁻ in overlying water is consistently low. The data suggest that newly fixed N can support sustained N transformation processes such as DNF and DNRA when surface water inorganic N supply rates are low.     

Nitrate respiration and diel migration patterns of diatoms are linked in sediments underneath a microbial mat

Diatoms are among the few eukaryotes known to store nitrate (NO₃⁻) and to use it as an electron acceptor for respiration in the absence of light and O₂. Using microscopy and ¹⁵N stable isotope incubations, we studied the relationship between dissimilatory nitrate/nitrite reduction to ammonium (DNRA) and diel vertical migration of diatoms in phototrophic microbial mats and the underlying sediment of a sinkhole in Lake Huron (USA). We found that the diatoms rapidly accumulated NO₃⁻ at the mat-water interface in the afternoon and 40% of the population migrated deep into the sediment, where they were exposed to dark and anoxic conditions for ~75% of the day. The vertical distribution of DNRA rates and diatom abundance maxima coincided, suggesting that DNRA was the main energy generating metabolism of the diatom population. We conclude that the illuminated redox-dynamic ecosystem selects for migratory diatoms that can store nitrate for respiration in the absence of light. A major implication of this study is that the dominance of DNRA over denitrification is not explained by kinetics or thermodynamics. Rather, the dynamic conditions select for migratory diatoms that perform DNRA and can outcompete sessile denitrifiers.     

硝态氮异化还原机制及其主导因素研究进展

硝态氮(NO_3~-)异化还原过程通常包含反硝化和异化还原为铵(DNRA)两个方面,是土壤氮素转化的重要途径,其强度大小直接影响着硝态氮的利用和环境效应(如淋溶和氮氧化物气体排放)。反硝化和DNRA过程在反应条件、产物和影响因素等方面常会呈现出协同与竞争的交互作用机制。综述了反硝化和DNRA过程的研究进展及其二者协同竞争的作用机理,并阐述了在NO_3~-、pH、有效C、氧化还原电位(Eh)等环境条件和土壤微生物对其发生强度和产物的影响,提出了今后应在产生机理、土壤环境因素、微生物学过程以及与其他氮素转化过程耦联作用等方面亟需深入研究,以期增进对氮素循环过程的认识以及为加强氮素管理利用提供依据。     

Influence of natural amphipod (<Emphasis Type="Italic">Victoriopisa</Emphasis><Emphasis Type="Italic">australiensis</Emphasis>) (Chilton, 1923) population densities on benthic metabolism,nutrient fluxes,denitrification and DNRA in sub-tropical estuarine sediment

The influence of natural populations of the sub-surface deposit-feeding amphipod Victoriopisa australiensis on sediment biogeochemistry was assessed by randomly collecting 21 sediment cores in a zone of Coombabah Lake, southern Moreton Bay, Australia, where the benthic infauna was dominated by this species. Cores were incubated sequentially to determine sediment–water column fluxes of oxygen, dissolved inorganic carbon and inorganic N species, followed by incubations to determine rates of denitrification and dissimilatory nitrate reduction to ammonium (DNRA) using the isotope pairing technique. Finally, each core was sieved in order to determine the population and biomass of amphipods present. Whilst all measures of overall benthic metabolism (sediment oxygen demand, and effluxes of inorganic carbon and nitrogen) showed increased with amphipod density, with rates being stimulated 70–220% at the highest categorised density range of 2,500–3,500 ind m⁻², only the correlation with dissolved inorganic carbon was statistically significant. In contrast, there were no discernable trends between amphipod densities and any of the N-cycle processes with the slopes of all correlations being very close to zero. These results highlight the differences in mesocosm simulations of fauna effects, which primarily relate to shifts in rates of organic matter turnover, compared to natural sediments where fauna effects relate more to induced changes in rates of organic matter deposition. Therefore, while mesocosms represent a powerful tool to investigate the mechanisms by which fauna influences microbial metabolism in the sediment, only studies of natural sediments can determine to what extent these mechanisms function in situ. Handling editor: Pierluigi Viaroli     

Denitrification in the periphyton associated with plant shoots and in the sediment of a wetland system supplied with sewage treatment plant effluent

Seasonal variation in denitrification and major factors controlling this process were determined in sediment, microbial communities attached to plant shoots (periphyton) and in the water of a Phragmites and an Elodea-dominated stand of a constructed wetland system between May 1997 and February 1998. The wetland was supplied with effluent from a sewage treatment plant. The denitrification rate in periphyton on plants shoots (expressed per shoot area) was always considerably higher than in the sediment and varied with the chlorophyll-a content of the periphyton in the course of the year. The algae in the periphyton provided attachment surfaces and probably also organic compounds to the denitrifying bacteria. Decreases in periphyton biomass and denitrification rate in the Phragmites and Elodea-dominated stands during the growing season were associated with enhanced shading by Phragmites shoots or a floating layer of macro-algae and Lemna spp., respectively. Light availability and the denitrification rate of periphyton increased again after the Phragmites shoots were cut in October. Nitrate appeared to limit the denitrification rate in the sediment. Periphyton denitrification rates were mostly lower on Elodea shoots than on Phragmites shoots, in spite of the higher living algal biomass on Elodea shoots. This difference was associated with lower nitrate concentrations in the Elodea-dominated stand. In the two stands, the daily denitrification rates in periphyton on shoots of Phragmites australis (44.4–121 mg N m^–2 stand area d^–1) and Elodea nuttallii (14.8–33.1 mg N m^–2 d^–1) were clearly more important than rates in the sediment (0.5–25.5 mg N m^–2 d^–1) or the water (0.4–3.9 mg N m^–2 d^–1). The presence of few bacteria attachment sites or low organic carbon availability possibly resulted in low denitrification rates in the water. Denitrification appeared to be a major process in nitrate removal from the through-flowing water in this wetland system.     

Correlations between foliar δ15N and nitrogen concentrations may indicate plant-mycorrhizal interactions

Nitrogen isotope measurements may provide insights into changing interactions among plants, mycorrhizal fungi, and soil processes across environmental gradients. Here, we report changes in δ¹⁵N signatures due to shifts in species composition and nitrogen (N) dynamics. These changes were assessed by measuring fine root biomass, net N mineralization, and N concentrations and δ¹⁵N of foliage, fine roots, soil, and mineral N across six sites representing different post-deglaciation ages at Glacier Bay, Alaska. Foliar δ¹⁵N varied widely, between 0 and –2‰ for nitrogen-fixing species, between 0 and –7‰ for deciduous non-fixing species, and between 0 and –11‰ for coniferous species. Relatively constant δ¹⁵N values for ammonium and generally low levels of soil nitrate suggested that differences in ammonium or nitrate use were not important influences on plant δ¹⁵N differences among species at individual sites. In fact, the largest variation among plant δ¹⁵N values were observed at the youngest and oldest sites, where soil nitrate concentrations were low. Low mineral N concentrations and low N mineralization at these sites indicated low N availability. The most plausible mechanism to explain low δ¹⁵N values in plant foliage was a large isotopic fractionation during transfer of nitrogen from mycorrhizal fungi to plants. Except for N-fixing plants, the foliar δ¹⁵N signatures of individual species were generally lower at sites of low N availability, suggesting either an increased fraction of N obtained from mycorrhizal uptake (f), or a reduced proportion of mycorrhizal N transferred to vegetation (T _r). Foliar and fine root nitrogen concentrations were also lower at these sites. Foliar N concentrations were significantly correlated with δ¹⁵N in foliage of Populus, Salix, Picea, and Tsuga heterophylla, and also in fine roots. The correlation between δ¹⁵N and N concentration may reflect strong underlying relationships among N availability, the relative allocation of carbon to mycorrhizal fungi, and shifts in either f or T _r. Received: 14 December 1998 / Accepted: 16 August 1999     

Interactions between Benthic Copepods,Bacteria and Diatoms Promote Nitrogen Retention in Intertidal Marine Sediments

The present study aims at evaluating the impact of diatoms and copepods on microbial processes mediating nitrate removal in fine-grained intertidal sediments. More specifically, we studied the interactions between copepods, diatoms and bacteria in relation to their effects on nitrate reduction and denitrification. Microcosms containing defaunated marine sediments were subjected to different treatments: an excess of nitrate, copepods, diatoms (Navicula sp.), a combination of copepods and diatoms, and spent medium from copepods. The microcosms were incubated for seven and a half days, after which nutrient concentrations and denitrification potential were measured. Ammonium concentrations were highest in the treatments with copepods or their spent medium, whilst denitrification potential was lowest in these treatments, suggesting that copepods enhance dissimilatory nitrate reduction to ammonium over denitrification. We hypothesize that this is an indirect effect, by providing extra carbon for the bacterial community through the copepods'' excretion products, thus changing the C/N ratio in favour of dissimilatory nitrate reduction. Diatoms alone had no effect on the nitrogen fluxes, but they did enhance the effect of copepods, possibly by influencing the quantity and quality of the copepods'' excretion products. Our results show that small-scale biological interactions between bacteria, copepods and diatoms can have an important impact on denitrification and hence sediment nitrogen fluxes.     

Experimental measurement of sediment nitrification and denitrification in Hamilton Harbour,Canada

This research examines the role of sediment nitrification and denitrification in the nitrogen cycle of Hamilton Harbour. The Harbour is subject to large ammonia and carbon loadings from a waste-water treatment plant and from steel industries. Spring ammonia concentrations rapidly decrease from 4.5 to 0.5 mg 1⁻¹, while spring nitrate concentrations increase from 1 to 2 mg l⁻¹, by mid-summer. A three-layer sediment model was developed. The first layer is aerobic; in it, oxidation of organics and nitrification occurs. The second layer is for denitrification, and the third layer is for anaerobic processes. Ammonia sources for nitrification include diffusion from the water column, sources associated with the oxidation of organics, sources from denitrification and from anaerobic processes. Diffusion of oxygen, ammonia and nitrate across the sediment-water interface occurs. Temperature effects are modelled using the Arrhenius concept. A combination of zero-order kinetics for nitrate or ammonia consumption with diffusion results in a half-order reaction, with respect to the water column loss rate to sediments. From experimental measurement, the rate of nitrification is 200 mg N 1⁻¹ sediment per day, while that of denitrification is 85 mg N 1–1 sediment per day at 20 °C. The Arrhenius activation energy is estimated as 15 000 cal/ mole-K and 17 000 cal/ mole-K for nitrification and denitrification, respectively, between 10 °C and 20 °C. Calculations of the flux of ammonia with the sediments, using the biofilm model, compare favourably with experimental observations. The ammonia flux from the water column is estimated to account for 20% of the observed decrease in water column stocks of ammonia, while the nitrate flux from the water column is estimated to account for 25% of the total nitrogen produced by the sediments.     

Nitrate and nitrite in interstitial waters of eelgrass beds in relation to the rhizosphere

The distribution of nitrate and nitrite in the interstitial water of the sediment of eelgrass (Zostera marina) bed of Izembek Lagoon, Alaska, were investigated. Their concentrations were relatively high (0 to 9.8 μg-at.N·1^?1, average 4.8 for nitrate; 0 to 4.0 μ-at.N·1^?1, average 1.9 for nitrite) although the sediments were anoxic and contained hydrogen sulphide. The rates of bacterial denitrification measured by ¹⁵N tracer technique ranged from 0.49×10^?10 to 1.2 × 10^?9 g-atN·g^?1·h^?1. When a steady state is maintained, the loss of nitrate and nitrite must be balanced by their production by bacterial nitrification. Experimentally determined rate of nitrification in the sediment was of the same order. A model experiment demonstrated that oxygen is transported from leaves to rhizomes and roots of eelgrass and released into the sediment. The oxygen is used for nitrification in the rhizosphere in anoxic sediments.     

Nitrogen transformations in microenvironments of river beds and riparian zones

The soil of flooded riparian zones, the rhizosphere of riparian plants, biofilms at solid surfaces in the river, and the surface layer of sediments all constitute important environments for the oxidative or reductive transformations of inorganic nitrogen compounds. The exact microzonation and coupling of the processes have recently been studied intensively with ¹⁵N enrichment methods and microsensors for NH₄⁺, NO₂⁻, NO₃⁻, and N₂O. Microsensor analyses of gradients in sediments and biofilms have shown that nitrate production takes place in an aerobic surface zone that has a maximum thickness of a few millimeters in most shallow-water sediments and may be as thin as 100 μm in biofilms from very eutrophic environments. In the anoxic zone, denitrification is also concentrated in a zone of maximum a few millimeters, and typically half of the nitrate produced by nitrification is denitrified while the other half escapes to the water. The supply of nitrate from above is primarily controlled by the oxic layer acting as a diffusion barrier, and therefore denitrification is generally a linear function of the nitrate concentration in the water. The overlying water is thus a much more important source of nitrate for denitrification if the concentration is high. The rate and location of denitrification are also affected by bioturbating animals, benthic microphytes, plants, and bacteria performing dissimilatory nitrate reduction to ammonium (DNRA).