Denitrification in nitrate-rich streams: application of N2:Ar and 15N-tracer methods in intact cores.

Rates of benthic denitrification were measured using two techniques, membrane inlet mass spectrometry (MIMS) and isotope ratio mass spectrometry (IRMS), applied to sediment cores from two NO3(-)-rich streams draining agricultural land in the upper Mississippi River Basin. Denitrification was estimated simultaneously from measurements of N2:Ar (MIMS) and 15N[N2] (IRMS) after the addition of low-level 15NO3- tracer (15N:N = 0.03-0.08) in stream water overlying intact sediment cores. Denitrification rates ranged from about 0 to 4400 micromol N x m(-2) x h(-1) in Sugar Creek and from 0 to 1300 micromol N x m(-2) x h(-1) in Iroquois River, the latter of which possesses greater streamflow discharge and a more homogeneous streambed and water column. Within the uncertainties of the two techniques, there is good agreement between the MIMS and IRMS results, which indicates that the production of N2 by the coupled process of nitrification/denitrification was relatively unimportant and surface-water NO3- was the dominant source of NO3- for benthic denitrification in these streams. Variation in stream NO3- concentration (from about 20 micromol/L during low discharge to 1000 micromol/L during high discharge) was a significant control of benthic denitrification rates, judging from the more abundant MIMS data. The interpretation that NO3- concentration directly affects denitrification rate was corroborated by increased rates of denitrification in cores amended with NO3-. Denitrification in Sugar Creek removed < or = 11% per day of the instream NO3- in late spring and removed roughly 15-20% in late summer. The fraction of NO3- removed in Iroquois River was less than that of Sugar Creek. Although benthic denitrification rates were relatively high during periods of high stream flow, when NO3 concentrations were also high, the increase in benthic denitrification could not compensate for the much larger increase in stream NO3- fluxes during high flow. Consequently, fractional NO3- losses were relatively low during high flow.     

Nitrate metabolism in sediments from seagrass (Zostera capricorni) beds of Moreton Bay,Australia

The fate of nitrate in sediments from seagrass (Zostera capricorni Aschers.) beds of Moreton Bay on the subtropical eastern coast of Queensland, Australia, was investigated. Added nitrate was metabolised at rates of 0.4 to 3.4 g N cm^-3 d^-1 when sediments were incubated under anaerobic conditions with a large excess of nitrate. The potential rate of nitrate utilization was as rapid in sediments from subtidal bare areas as from adjacent seagrass beds. Ammonium was produced rapidly from¹⁵N-nitrate by microbial action in all the subtidal sediments examined. After 12 h of incubation, 13 to 28% of the¹⁵N initially added as labelled nitrate was detected as labelled ammonium in the sediments. Denitrification, although not measured directly, appeared to be a relatively minor fate of nitrate. Benthic microbes took up large amounts of¹⁵N but only after a delay of 6 h; this pattern could have been due to induction and synthesis of the enzymes necessary for nitrate uptake, and the assimilation of labelled ammonium. Under field conditions, assimilation by seagrasses and denitrification by bacteria were probably not significant sinks for nitrate in comparison with uptake by benthic microbes and dissimilatory reduction to ammonium.     

Denitrification and the nitrogen budget of a reservoir in an agricultural landscape.

Denitrification is an important process in aquatic sediments, but its role has not been assessed in the N mass balance of upper-Midwestern (USA) reservoirs that receive large agricultural riverine N inputs. We used a 4400-ha reservoir to determine the role of denitrification in the N mass balance and effectiveness in reducing downstream transport of NO(3-)N. Sediment denitrification was (1) measured monthly (March 2002-March 2003) at eight sites in the Lake Shelbyville reservoir in central Illinois using the acetylene inhibition, chloramphenicol technique, (2) scaled to the overall reservoir and compared to N not accounted for in a mass balance, and (3) estimated indirectly using long-term (1981-2003) mass balances of N in the reservoir. Denitrification rates in the reservoir were high during spring and early summer of 2002, when maximum NO(3-)N concentrations were measured (10-14 mg NO(3-)N/L). We estimated that denitrification for the year was between 2580 and 5150 Mg N. Missing N from the mass balance was 3004 Mg N, suggesting that sediment denitrification was the sink. Areal rates of sediment denitrification in the reservoir ranged from 62 to 225 g N x m(-2) x yr(-1), with rates a function of both denitrification intensity (microg N x g dry mass x h(-1)) and the overall mass of sediment present. From 1981 to 2003 the average NO(3-)N inlet flux was 8900 Mg N/yr. About 58% of the total NO(3-)N input was removed, and annual NO(3-)N removed as a percentage of inputs was significantly related to reservoir retention time (average = 0.36 yr for the 23 years, range = 0.21-0.84 yr). By scaling denitrification in Lake Shelbyville to other reservoirs in Illinois, we estimated a sink of 48900 Mg N/yr. When combined with estimated in-stream denitrification, 60900 Mg N/yr was estimated to be removed by sediment denitrification. This reduces riverine export from Illinois to the Gulf of Mexico, where the flux during the 1990s was about 244000 Mg N/yr, and illustrates the importance of reservoir denitrification as an N sink in Midwestern agricultural landscapes.     

Floodplain restoration enhances denitrification and reach-scale nitrogen removal in an agricultural stream

Streams of the agricultural Midwest, USA, export large quantities of nitrogen, which impairs downstream water quality, most notably in the Gulf of Mexico. The two-stage ditch is a novel restoration practice, in which floodplains are constructed alongside channelized ditches. During high flows, water flows across the floodplains, increasing benthic surface area and stream water residence time, as well as the potential for nitrogen removal via denitrification. To determine two-stage ditch nitrogen removal efficacy, we measured denitrification rates in the channel and on the floodplains of a two-stage ditch in north-central Indiana for one year before and two years after restoration. We found that instream rates were similar before and after the restoration, and they were influenced by surface water NO3- concentration and sediment organic matter content. Denitrification rates were lower on the constructed floodplains and were predicted by soil exchangeable NO3- concentration. Using storm flow simulations, we found that two-stage ditch restoration contributed significantly to NO3- removal during storm events, but because of the high NO3- loads at our study site, < 10% of the NO3- load was removed under all storm flow scenarios. The highest percentage of NO3- removal occurred at the lowest loads; therefore, the two-stage ditch's effectiveness at reducing downstream N loading will be maximized when the practice is coupled with efforts to reduce N inputs from adjacent fields.     

Macroinfauna of a Southern New England salt marsh: seasonal dynamics and production

The animal-habitat relationships and seasonal dynamics of the benthic macroinfauna were investigated from November 1986 to October 1988 in the Great Sippe-wissett salt marsh (Massachusetts, USA). Total macrofaunal abundance varied seasonally, displaying a peak in late spring and early summer, then declining sharply during late summer and recovering briefly in fall before collapsing in winter. Three macroinfaunal assemblages were found in the marsh, distributed along gradients of environmental factors. These included a sandy non-organic sediment assemblage, a sandy organic sediment assemblage and a muddy sediment assemblage. The species groups characteristic of unstable sandy non-organic sediments included the polychaetes Leitoscoloplos fragilis, Aricidea jefreyssi, Magelona rosea and Streptosyllis verrilli, the oligochaete Paranais litoralis, and the crustacean Acanthohaustorius millsi. Sandy organic sediments were characterized by the polychaetes Marenzelleria viridis, Capitella capitata, Neanthes succinea, N. arenaceodonta, Polydora ligni and Heteromastus filiformis, the oligochaete Lumbricillus sp., and the mollusc Gemma gemma. In muddy sites, the polychaete Streblospio benedicti and the oligochaetes Paranais litoralis and Monopylephorus evertus were the dominant species. Secondary production of benthic macroinfauna in each of these habitats was estimated. The highest values of biomass and production were recorded in the sandy organic sediments. Secondary production was estimated to be 1850 kJ m^-2 yr^-1 in sandy organic areas, but only 281 kJ m^-2 yr^-1 in sandy non-organic areas and 113 kJ m^-2 yr^-1 in muddy areas. This results in an area-weighted average production of 505 kJ m^-2 yr^-1 for the unvegetated areas of the marsh. The Great Sippewissett salt marsh has an area of 483800 m², the total secondary production of the macroinfauna for the whole unvegetated area of the marsh was estimated as 4651 kg dry wt yr^-1, expressed as somatic growth. This production value seems consistent with production data obtained for other intertidal North Atlantic environments.     

Denitrification across landscapes and waterscapes: a synthesis.

Denitrification is a critical process regulating the removal of bioavailable nitrogen (N) from natural and human-altered systems. While it has been extensively studied in terrestrial, freshwater, and marine systems, there has been limited communication among denitrification scientists working in these individual systems. Here, we compare rates of denitrification and controlling factors across a range of ecosystem types. We suggest that terrestrial, freshwater, and marine systems in which denitrification occurs can be organized along a continuum ranging from (1) those in which nitrification and denitrification are tightly coupled in space and time to (2) those in which nitrate production and denitrification are relatively decoupled. In aquatic ecosystems, N inputs influence denitrification rates whereas hydrology and geomorphology influence the proportion of N inputs that are denitrified. Relationships between denitrification and water residence time and N load are remarkably similar across lakes, river reaches, estuaries, and continental shelves. Spatially distributed global models of denitrification suggest that continental shelf sediments account for the largest portion (44%) of total global denitrification, followed by terrestrial soils (22%) and oceanic oxygen minimum zones (OMZs; 14%). Freshwater systems (groundwater, lakes, rivers) account for about 20% and estuaries 1% of total global denitrification. Denitrification of land-based N sources is distributed somewhat differently. Within watersheds, the amount of land-based N denitrified is generally highest in terrestrial soils, with progressively smaller amounts denitrified in groundwater, rivers, lakes and reservoirs, and estuaries. A number of regional exceptions to this general trend of decreasing denitrification in a downstream direction exist, including significant denitrification in continental shelves of N from terrestrial sources. Though terrestrial soils and groundwater are responsible for much denitrification at the watershed scale, per-area denitrification rates in soils and groundwater (kg N x km(-2) x yr(-1)) are, on average, approximately one-tenth the per-area rates of denitrification in lakes, rivers, estuaries, continental shelves, or OMZs. A number of potential approaches to increase denitrification on the landscape, and thus decrease N export to sensitive coastal systems exist. However, these have not generally been widely tested for their effectiveness at scales required to significantly reduce N export at the whole watershed scale.     

Effects of restoration and reflooding on soil denitrification in a leveed Midwestern floodplain.

River floodplains have the potential to remove nitrate from water through denitrification, the anaerobic microbial conversion of nitrate to nitrogen gas. An important factor in this process is the interaction of river water with floodplain soil; however, many rivers have been disconnected from their historic floodplains by levees. To test the effect of reflooding a degraded floodplain on nitrate removal, we studied changes in soil denitrification rates on the Baraboo River floodplain in Wisconsin, USA, as it underwent restoration. Prior to this study, the site had been leveed, drained, and farmed for more than 50 years. In late fall 2002, the field drainage system was removed, and a gate structure was installed to allow controlled flooding of this site with river water. Soil moisture was extremely variable among zones and months and reflected local weather. Soil organic matter was stable over the study period with differences occurring along the elevation gradient. High soil nitrate concentrations occurred in dry, relatively organic-poor soil samples and, conversely, all samples with high moisture soils characterized by low nitrate. We measured denitrification in static cores and potential denitrification in bulk samples amended with carbon and nitrogen, one year before and two years following the manipulation. Denitrification rates showed high temporal and spatial variability. Static core rates of individual sites ranged widely (from 0.00 to 16.7 microg N2O-N x [kg soil](-1) x h(-1), mean +/- SD = 1.10 +/- 3.02), and denitrification enzyme activity (DEA) rates were similar with a slightly higher mean (from 0.00 to 15.0 microg N2O-N x [kg soil](-1) x h(-1), 1.41 +/- 1.98). Denitrification was not well-correlated with soil nitrate, organic matter content, or moisture levels, the three parameters typically thought to control denitrification. Static core denitrification rates were not significantly different across years, and DEA rates decreased slightly the second year after restoration. These results demonstrate that restored agricultural soil has the potential for denitrification, but that floodplain restoration did not immediately improve this potential. Future floodplain restorations should be designed to test alternative methods of increasing denitrification.     

Nitrogen exchange between a southeastern USA salt marsh ecosystem and the coastal ocean

The salt marsh ecosystem at North Inlet, South Carolina, USA consistently exported dissolved inorganic nitrogen via tidal exchange with the coastal Atlantic Ocean. Concentrations centrations of NH ₄ ⁺ and NO ₃ ^- +NO ₂ ^- displayed distinct tidal patterns with rising values during ebb flow. These patterns suggest the importance of biogeochemical processes in the flux of material from the salt marsh. NH ₄ ⁺ export peaked during the summer (15 to 20 mg m^-2 tide^-1) during a net balance of tidal water exchange. Remineralization of NH ₄ ⁺ within the salt marsh system appears to be contributing to the estimated annual net export of bout 4.7 g NH ₄ ⁺ -N m^-2 yr^-1. NO ₃ ^- +NO ₂ ^- exports were higher in the fall and winter of 1979 (2 to 4 mg N m^-2 tide^-1). The winter export coincided with a considerable net export of water with no distinctive concentration patterns, suggesting a simple advective export. However, the fall peak of NO ₃ ^- +NO ₂ ^- export occurred during a period of net water balance in tidal exchange and an insignificant freshwater input from the western, forested boundary. During the summer and fall, tidal concentration patterns were particularly apparent, suggesting that nitrification within the salt marsh system was contributing to the estimated annual net export of ca 0.6 g NO ₃ ^- +NO ₂ ^- -N m^-2 yr^-1.Contribution No. 637 from the Belle W. Baruch Institute of Marine Biology and Coastal Research     

Effects of mussel aquaculture on the nitrogen cycle and benthic communities in Kenepuru Sound,Marlborough Sounds,New Zealand

Nitrogen pools and transformations and benthic communities at a Perna canaliculus farm and a nearby reference site without direct influence of marine farming in Kenepuru Sound, New Zealand, were compared on four dates between September 1982 and May 1983. The organic nitrogen pool in the top 12 cm sediment was 7.4 to 10.8 mol m^-2 at the mussel farm and 6.1 to 8.9 mol m^-2 at the reference site. The nitrate and nitrite pools were similar in both sediments, but the ammonium pool in the mussel farm sediment was about twice as high as in the reference sediment. In January, the sediment ammonium concentrations ranged from 418 nmol cm^-3 (surface) to 149 nmol cm^-3 (12 cm depth) at the mussel farm and from 86 to 112 nmol cm^-3 at the reference site. The molar C:N ratio of the sediment organic matter was 6.2 to 7.2 at the mussel farm and 7.9 to 10.0 at the reference site. The molar N:P ratio of the sediment organic matter was 4.3 to 7.2 and 3.3 to 6.1 at mussel farm and reference site, respectively. The total nitrogen mineralisation rate in the top 12 cm sediment ranged from 21.7 to 37.1 mmol m^-2 d^-1 at the mussel farm and from 8.5 to 25.0 mmol m^-2 d^-1 at the reference site. Ammonium excretion by mussels was about 4.7% (January) and 7.4% (May) of the combined nitrogen mineralisation by mussels and sediment. The sediment-denitrification rate was 0.7 to 6.1 mmol m^-2 d^-1 at the mussel farm and 0.1 to 0.9 mmol m^-2 d^-1 at the reference site. In January, 76 and 93% of the nitrate reduced in the sediments were denitrified at the mussel farm and reference site, respectively. The denitrification rate on the mussel lines (determined on detritus-covered mussels) was twice the mussel farm sediment-denitrification rate and 10 times the reference sediment-denitrification rate. Total denitrification at the mussel farm was 21% higher than at the reference site. The loss of nitrogen through mussel harvest and denitrification was 68% higher at the mussel farm. The surface layers of both sediments contained about 75 mg m^-2 chlorophyll a. Sediment phaeophytin levels were 52 mg m^-2 at the reference site and 137 mg m^-2 at the mussel farm. While the benthic infauna of the mussel-farm sediment consisted only of polychaete worms, the reference sediment contained also bivalve molluscs, brittle stars and crustaceans.     

Acetylene reduction (nitrogen fixation) in a Delaware,USA salt marsh

Acetylene reduction (nitrogen fixation) was measured in several vegetational areas in a Delaware, USA salt marsh. Samples were collected for 1 yr and the results showed a seasonally variable pattern of acetylene reduction at all stations. Peak rates were generally recorded during the later summer and early fall (September–October). The seasonality was influenced mainly, although not exclusively, by the soil temperature. In addition, samples collected in short Spartina alterniflora stands exhibited rates which were up to 20-fold higher than those found in samples from tall S. alterniflora stands. Over 50% of the total yearly ethylene production occurred from mid-August until the beginning of December at the tall and short S. alterniflora stations. Maximum activity occurred at 5 cm depth for all stations. Surface activity accounted for only 3–4% of the total measured in the top 20 cm. Addition of glucose or mannitol resulted in considerable increases in activity, thus suggesting that heterotrophic acetylene reduction is carbon and/or energy limited. The results obtained in this study indicate that the measured rates are only potential rates and that considerable caution must be used in extrapolating from acetylene reduction rates to nitrogen fixation rates in situ.     

Physiology and chemical composition of nitrogen-fixing phytoplankton in the central North Pacific Ocean

The magnitude and physiological characteristics of biological nitrogen fixation have been studied in the oligotrophic waters of the North pacific gyre. The filamentous blue-green algae Trichodesmium spp. and Richelia intracellularis were the important nitrogen-fixing phytoplankton. Most of the nitrogen fixation occurs in the upper 40 m of the water column, with detectable fixation as deep as 90 m, which corresponds to about the 1 % light depth. There was no evidence of photoinhibition of nitrogen fixation, although CO₂ reduction was depressed slightly at the highest light levels. The rate of nitrogen fixation in the water column varied throughout the day, being highest in mid-morning and in late afternoon. Relatively high fixation rates were also found during periods of darkness. Elevated oxygen concentrations had a marked inhibitory effect on rates of nitrogen fixation, a pO₂ of 0.4 atm causing a 75% inhibition. Data from studies of nitrogen fixation and assimilation rates of ¹⁵N-labelled nitrate, ammonium, and urea indicate that nitrogen fixation furnished about 3% of the total daily fixed nitrogen requirement for phytoplankton growth. Studies with isolated colonies of Trichodesmium spp. indicated that 100% of their nitrogen requirement was met by nitrogen fixation. Chemical composition of the Trichodesmium colonies showed that the C:N ratio was 4.1 and that their phosphorus content relative to carbon or nitrogen was much lower than that of the total particulate material in the water column. Elevated ratios of carbon: adenosine triphosphate (ATP) also suggest that phosphorus deficiency may be limiting the growth of Trichodesmium. The magnitude of nitrogen fixation in the gyre is seasonally dependent, with high rates in late summer and autumn. At these times the water column is stratified, with phosphate and nitrate barely detectable in the upper 100 m. Our data suggest that during these months of stratification, biological fixation of nitrogen amounts to about 33 g-at N/m²/day.     

Effect of monsoonal climate on sulfate reduction in coastal sediments of the central Great Barrier Reef lagoon

Rates of sulfate reduction were measured during the spring dry and summer wet seasons of 1992 in shallow coastal sediments of the central Great Barrier Reef lagoon. In the dry season, sulfate reduction rates, integrated to 18–20 cm depth, ranged from 23.8 to 30.8 mmol S m^-2 d^-1. In the wet season, heavy monsoonal rains and wind-induced waves caused severe disturbance of sediments leading to less anoxic conditions, and to a 2- to 3-fold decline in rates of sulfate reduction (10.2 to 12.8 mmol S m^-2 d^-1). The percentage of reduced ³⁵S recovered as acid-volatile sulfide (% AVS_red) ranged from 15.8 to 44.9% in spring, and was significantly reduced at each station in summer (range 14.5 to 31.4%). An analysis of variance indicated that seasonality accounted for 31.5% of the total variance in total rates of sulfate reduction, with only 7.5% of the variance accounted for by depth into the sediment; there were no station differences. In both seasons, there were no clear patterns in the proportion of radiolabel incorporated into the AVS and chromium-reducible sulfur (CRS) pools with depth into the sediment. These results are contrary to the seasonal patterns and pathways of sulfate reduction in temperate coastal sediments.     

Below-ground nitrogen cycling in relation to net canopy production in mangrove forests of southern Thailand

Rates of accumulation, transformation and availability of sediment nitrogen in four mangrove forests of different age and type in southern Thailand were examined in relation to forest net canopy production. Net ammonification (range: 0.3-2.3 mmol N m^-2 day^-1), nitrification (range: 0-0.7 mmol N m^-2 day^-1) and nitrogen fixation (range: 0-0.6 mmol N m^-2 day^-1) in surface sediments equated to <10% of canopy nitrogen demand (range: 7.5-32 mmol N m^-2 day^-1). By mass balance, we estimated that most of the nitrogen required for tree growth must be derived from root-associated nitrogen fixation and/or mineralisation processes occurring possibly to the maximum depth of live root penetration (75-100 cm). Denitrification, nitrification, rainfall and tidal exchange were comparatively small components of sediment nitrogen flow. Denitrification (range: 0-3.8 mmol N m^-2 day^-1) removed 3-6% of total nitrogen input at three Rhizophora forests, but removed 23% of total nitrogen input in a high-intertidal Ceriops forest. Nitrogen burial ranged from 4% to 12% of total nitrogen input, with the greatest burial rates in two forests receiving the least tidal inundation. Inputs of nitrogen to the forests were rapid (range: 11-37 mmol N m^-2 day^-1), likely originating from upstream sources such as agricultural and industrial lands, sewage and shrimp ponds. Our results indicate that ~70% to 90% of the nitrogen supplied to the forest floor is shunted via the ammonium pool to trees to sustain the rapid rates of net canopy production measured in these forests. Differences in plant-sediment nitrogen relations between the forests appeared to be a function of the interaction between intertidal position and stand age.     

Nitrogen fixation (acetylene reduction) in the phyllosphere of Thalassia testudinum

N₂ fixation (C₂H₂ reduction) associated with the leaves of the sea grass Thalassia testudinum was investigated at 5 sites in South Florida (Biscayne Bay) and one site in the Bahamas (Bimini Harbor). Significant activities were correlated with the occurrence of a heterocystous blue-green alga (Calothrix sp.) on the leaves. C₂H₂ reduction was not stimulated by organic compounds, either aerobically or anaerobically in the light or dark. Therefore, other physiological types of microbes were not important in N₂ fixation. Diurnal and seasonal variations in N₂ fixation occurred, with maximal rates during the daytime and in the late spring and early summer. N₂ fixation was negligible at four stations in Biscayne Bay. At the fifth station, near Fowey Rock, about 5 kg N ha^-1 year^-1 was fixed. In the summer, the N₂ fixed per day (4–5 mg N m^-2) could provide 4 to 23% of the foliar productivity demands of T. testudinum at this site and the station in Bimini Harbor. N₂ fixation at the periphery of a sea-grass patch, near Fowey Rock, could provide 8 to 38% of the daily nitrogen requirement for leaf production, and thereby might compensate for a less effective trapping and recycling of nitrogen from dead leaves in such regions.     

Denitrification mitigates N flux through the stream-floodplain complex of a desert city

The Indian Bend Wash (IBW) flood-control project relies on a greenbelt to carry floods through Scottsdale, Arizona, USA. The greenbelt is characterized by a chain of shallow artificial lakes in a larger floodplain of irrigated turf, which has been protected from encroaching urban development. As such, this urban stream-floodplain complex can be divided into three subsystems: artificial lakes, channelized stream segments, and floodplain. We conducted experiments to evaluate which, if any, of these subsystems were important sites of denitrification, and to explore factors controlling denitrification rates. Denitrification enzyme activity (DEA) bioassays were conducted on sediments from eight lake and six stream segments as well as soil samples from eight floodplain transects. Mass-specific potential denitrification rates were significantly higher in lakes than in streams or floodplains. Nutrient limitation bioassays revealed that nitrate (NO3-) limited denitrification in lake sediments, a surprising finding given that NO3(-)-rich groundwater additions frequently raised lake NO3(-) concentration above 1 mg N/L. Experiments on intact lake cores suggested that denitrification was limited by the rate NO3(-) diffused into sediments, rather than its availability in overlying water. Floodplain denitrification was limited by water content, not NO3(-) or C, and irrigation of soils stimulated denitrification. We constructed a N budget for the IBW stream-floodplain complex based on our experimental results. We found that both lakes and floodplains removed large quantities of N, with denitrification removing 261 and 133 kg N ha(-1) yr(-1) from lake sediments and floodplain soils, respectively, indicating that lakes are hotspots for denitrification. Nevertheless, because floodplain area was >4.5 times that of lakes, floodplain soils removed nearly 2.5 times as much N as lake sediments. Given the desert's low annual precipitation, a finding that floodplain soils are active sites of denitrification might seem implausible; however, irrigation is common in urban landscapes, and it elevated annual denitrification in IBW. Based on our results, we conclude that construction of artificial lakes created hotspots while application of irrigation water created hot moments for denitrification in the stream-floodplain complex, demonstrating that management decisions can improve the ability of urban streams to provide critical ecosystem services like N retention.     

Nutrient regeneration in shallow-water sediments of the estuarine plume region of the nearshore Georgia Bight,USA

Benthic community respiration and the cycling of N and P were seasonally investigated in the unprotected, sandy sediments (Z5m) of the nearshore zone of the Georgia Bight, USA in 1981 and 1982. Nutrient exchange across the sediment-water interface was calculated from a diffusive model, measured by in-situ enclosure experiments and estimated from whole core incubations. Seasonally changing pore water profiles indicated that the sediments were not in steady-state with respect to N and P and showed the characteristics of enhanced interstitial water movement by benthic animals. Over an annual period the total flux of nitrogen measured in situ averaged 1812 mol m^-2 d^-1 from the sediments. NH ₄ ⁺ flux accounted for the vast majority of the total directly measured N flux (77%), followed by nitrate + nitrite (14%), and dissolved organic nitrogen (9%). Phosphorus flux averaged 537 mol m^-2 d^-1. A large ratio of in-situ fluxes to calculated diffusive fluxes (5.2:1) indicated flux enhancement due to benthic animal activity. ammonium fluxes measured in situ did not agree well with the rate of NH ₄ ⁺ produced in incubated whole cores (11.7 mmol m^-2 d^-1). Relative rates of C, N and P release throughout the year fluctuated considerably. Generally, nutrient fluxes were not simply related to respiration or temperature. As respiration was highly correlated with temperature, however, this suggested that respiration-regeneration was temporarily decoupled from exchange across the sediment-water interface. The annual C-N-P flux stoichiometry was 130:3.1:1. Using the rate at which NH ₄ ⁺ was produced in incubated cores the stoichiometry was 120:21:1. The anomalously low N flux measured in situ was attributed to a combination of denitrification and wave-and current-induced sediment nutrient flushing. The potential for sediment flushing is high as experiments showed that sediments were fluidized or resuspended down to 25 cm during large storms. Benthic nutrient flux contributed 40% to the annual P but only 11% to the annual N requirements of the pelagic primary producers.This is Contribution No. 558 from the University of Georgia Marine Institute. This work was supported by the Georgia Sea Grant College Program maintained by the National Oceanic and Atmospheric Administration, US Department of Commerce, under Grant No. NA80AA-D-00091     

Seasonal variations in nitrogen-fixation (acetylene reduction) and sulphate-reduction rates in the rhizosphere of Zostera noltii: nitrogen fixation by sulphate-reducing bacteria

Nitrogen-fixation (acetylene reduction) rates were measured over an annual cycle in meadows of the seagrass Zostera noltii Hornem in the Bassin d'Arcachon, south-west France, between March 1994 and February 1995, using both slurry and whole-core techniques. Measured rates using the slurry technique consistently overestimated those determined on whole cores, probably due to the release of labile organic carbon sources as a result of root damage during preparation of the slurries. Thus, the whole-core technique may provide a more accurate estimate of in situ activity, since disturbance of physicochemical gradients of oxygen, sulphide, nutrients and the relationship between the plant roots and the rhizosphere microflora is minimised. Rates measured by the whole-core method were 1.8- to 4-fold greater (dependent upon season) in the light than those measured during dark incubations, indicating that organic carbon diffusing from the plant roots during photosynthesis was an important factor in regulating nitrogen fixation in the rhizosphere. Additions of sodium molybdate, a specific inhibitor of sulphate-reducing bacteria (SRB) inhibited acetylene-reduction activity by >80% as measured by both the slurry and whole-core techniques throughout the year, inferring that SRB were the dominant component of the nitrogen-fixing microflora. A mutualistic relationship between Z. noltii and nitrogen-fixing SRB in the rhizosphere, based on the exchange of organic carbon and fixed nitrogen is proposed. Acetylene- and sulphate-reduction rates showed distinct summer peaks which correlated with a reduced availability of ammonium in the sediment and the annual growth cycle of Z. noltii in the basin. Overall, these data indicate that acetylene reduction (nitrogen fixation) activity in the rhizosphere of Z. noltii was regulated both by the availability of organic carbon from the plant roots and maintenance of a low NH ₄ ⁺ concentration in the vicinity of the plant roots due to efficient assimilation of NH ₄ ⁺ by Z. noltii during the growth season. Nitrogenfixation rates determined from acetylene-reduction rates measured using the whole-core technique ranged from 0.1 to 7.3 mg N m^-2d^-1, depending on season, and were calculated to contribute between 0.4 and 1.1 g N m^-2yr^-1, or 6.3 to 12% of the annual fixed nitrogen requirement of Z. noltii.     

Denitrification in bottom sediment near oil production facilities off the Louisiana Gulf Coast

There is little information on denitrification in Gulf of Mexico bottom sediment. Potential denitrification rates in surface sediment were measured along transects legs extending 0–800 m from two offshore oil production platforms. The average potential denitrification ranged from approximately 50 mg N m^?2 d^?1 in surface sediment near the platforms to 15 mg N m^?2 d^?1 in sediment 800 m from the platforms. Measured denitrification rates were correlated to a higher organic matter content in sediment nearer the platforms. This research examined only a small component of nitrogen processing in Gulf of Mexico sediment. Additional research should examine the effect of nitrogen loading and temporal and spatial variability on denitrification rate.     

Allocation of nitrogen and carbon in an estuarine salt marsh in Portugal

Above and below-ground biomass and nitrogen and carbon composition ofSpartina maritima, Halimione portulacoides andArthrocnemum perenne, dominating species in plant communities of the lower, middle and higher salt marsh, respectively, were compared in an estuarine salt marsh in Portugal. Plant and soil nitrogen and carbon pools were estimated. For all three species root biomass was significantly higher (70–92% of total biomass) than above-ground biomass. The percentage of root biomass was related to the location of the plants in the marsh: higher values were found in plants growing in the lower salt marsh where the sediment was more unstable and subject to tidal action, which stresses the role of the roots as an anchor. For all three species nitrogen concentrations were highest in leaves, reflecting the photosynthetic role of the tissue. For carbon higher concentrations were found in the stems, with the exception ofS. maritima. In general, lower nitrogen concentrations were found in summer, which can be explained by dilution processes due to plant growth. For both nitrogen and carbon, higher concentrations were found in the soil surface layers. Higher soil nitrogen and carbon levels were associated with higher organic matter contents. Most of the nitrogen in the salt marsh occurred in the sediments (0–40 cm) and only ca. 5.7–13.3% of the total was found in the plants. The greater portion (76.5%–86%) of carbon was found in the sediment.     

Intertidal distribution,population dynamics and production of the amphipod Uhlorchestia spartinophila in a Georgia,USA, salt marsh

The talitrid amphipod Uhlorchestia spartinophila Bousfield and Heard occurs in close association with the smooth cordgrass Spartina alterniflora Loisel in salt marshes along the U.S. Atlantic coast. In order to assess its potential as a prey resource for secondary consumers, we followed the population and production dynamics of this amphipod from 3 November 1990 to 2 October 1991 in a salt marsh on Sapelo Island, Georgia. Semimonthly samples were taken along an intertidal transect, which extended from a vegetated creekbank levee landward 140 m to a site near the upland edge of the marsh. Amphipod densities ranged from 9 to 826 indm^-2 and were greatest in the levee and high marsh habitats at opposite ends of the transect. The highest densities occurred in March to May and the lowest in November and December. Specific growth rates, which ranged from 0.001 to 0.024 mm mm^-1 d^-1 length and 0.003 to 0.068 mg mg^-1 d^-1 AFDW (ash-free dry weight), were least in winter and decreased with increasing amphipod size. Although some reproduction occurred year-round, most of the population's reproductive output was from January to May, when adults had the greatest size-specific mass. Sex ratio usually favored females, which were larger than males. Minimum adult female size was 5.4 mm total length (TL). Broods included from 1 to 28 eggs or young and increased with increasing body size, averaging 7.4 young for an average-size female of 7.6 mm TL. Annual production, which ranged from 0.769 to 1.444 g AFDW m^-2, was least in the low marsh and greatest in the levee habitat. Production:biomass ratios were 15.4 to 17.3 in different habitats. Such high turnover rates, together with the population's broad intertidal distribution, suggest a greater contribution to trophic dynamics than is implied by the relatively low standing stock biomass of U. spartinophila in this marsh system.