Temporal and spatial variability in phytoplankton ammonium and nitrate uptake in the Delaware Estuary

Phytoplankton NH₄⁺ and NO₃⁻ uptake was examined along the longitudinal salinity gradient of the Delaware Estuary over several seasonal cycles using ¹⁵N-tracer techniques. Saturated nitrogen uptake rates increased directly with water temperature and reached a maximum of 380 nmol Nl⁻¹h⁻¹ during summer. This temperature dependence was related primarily to changes in the rate of maximum chlorophyll specific uptake, which varied exponentially between 2 and 70 nmol N [μg Chl h]⁻¹ over a temperature range of 2–28°C. Despite these high uptake rates, balanced growth (C:N7:1) could be maintained over the diel light cycle only by highly efficient nitrogen uptake at low light intensities and dark uptake below the photic zone and at night (dark UPTAKE=25% maximum uptake).Ammonium fulfilled 82% of the annual phytoplankton nitrogen demand in the estuary despite dominance of NO₃⁻ in the ambient dissolved inorganic nitrogen pool. The predominance of NH₄⁺ uptake occurred because of the general suppression of NO₃⁻ assimilation at NH₄⁺ concentrations in excess of 2 μ . This suppression, however, was not as universal as has been reported for other systems, and it is suggested that the extremely high NO₃⁻ concentrations found in the estuary contribute to this pattern. Nitrate was a significant source of nitrogen only during periods of high phytoplankton production in summer, and when NH₄⁺ concentrations were low towards the end of the spring bloom.     

Nutrients (organic C, P, N, Si) in the eutrophic River Loire (France) and its estuary

The Loire estuary has been surveyed from 1982 to 1985 by 13 isochronous longitudinal profiles realized at low tide. Nutrient (SiO₂, NO₃⁻, NH₄⁺, PO³⁻₄, particulate organic carbon or POC) patterns are very variable depending on the season, the estuarine section [river, upper-inner estuary, upstream of the fresh-water-saline-water interphase FSI, the lower-inner estuary characterized by the high turbidity zone (HTZ), the outer estuary] and the river discharge. Biological processes are dominant. In the eutrophied River Loire (summer pigment > 100 μg l⁻¹), the high algal productivity (algal POC > 3 mg l⁻¹) results in severe depletion of SiO₂, PO₄³⁻, NO₃⁻. The enormous biomass (55 000 ton algal POC/year) is degraded in the HTZ where bacterial activity is intense. As a result, there is generally a regeneration of dissolved SiO₂ and PO₄³⁻, a marked NH₄⁺ maximum, while NO₃⁻ is conservative or depleted when the HTZ is nearly anoxic. Other processes can be considered including pollution from fertilizer plans (PO₄³⁻, NH₄⁺) and from a hydrothermal power plant (NH₄⁺). In the less turbid outer estuary, nutrients are generally conservative. Major variations of concentrations are observed in the lowest chlorinity section (Cl⁻ < 1 g kg⁻) and also upstream the FSI, defined here as a 100% increase in Cl⁻. Nutrient inputs to the ocean are not significantly modified for SiO₂ and NO₂⁻, but are increased by 70% and 180% for PO₄³⁻ and NH₄⁺ and depleted by 60% for POC. Odd hydrological events, especially some floods, may perturbate or even mask the usual seasonal pattern observed in profiles.     

Biogeochemical transport in the Loxahatchee River estuary, Florida: The role of submarine groundwater discharge

The distributions of dissolved organic carbon (DOC), Ba, U, and a suite of naturally occurring radionuclides in the U/Th decay series (²²²Rn, ^{223,224,226,228}Ra) were studied during high- and low-discharge conditions in the Loxahatchee River estuary, Florida to examine the role of submarine groundwater discharge in estuarine transport. The fresh water endmember of this still relatively pristine estuary may reflect not only river-borne constituents, but also those advected during active groundwater/surface water (hyporheic) exchange. During both discharge conditions, Ba concentrations indicated slight non-conservative mixing. Such Ba excesses could be attributed either to submarine groundwater discharge or particle desorption processes. Estuarine dissolved organic carbon concentrations were highest at salinities closest to zero. Uranium distributions were lowest in the fresh water sites and mixed mostly conservatively with an increase in salinity. Suspended particulate matter (SPM) concentrations were generally lowest (< 5 mg L^− 1) close to zero salinity and increased several-fold ( 18 mg L^− 1; low discharge) toward the seaward endmember, which may be attributed to dynamic resuspension of bottom sediments within Jupiter Inlet.Surface water-column ²²²Rn activities were most elevated (> 28 dpm L^− 1) at the freshwater endmember of the estuary and appear to identify regions of the river most influenced by the discharge of fresh groundwater. Activities of four naturally occurring isotopes of Ra (^{223,224,226,228}Ra) in this estuary and select adjacent shallow groundwater wells yield mean estuarine water-mass transit times of less than 1 day; these values are in close agreement to those calculated by tidal prism and tidal frequency. Submarine groundwater discharge rates to the Loxahatchee River estuary were calculated using a tidal prism approach, an excess ²²⁶Ra mass balance, and an electromagnetic seepage meter. Average SGD rates ranged from 1.0 to 3.8 × 10⁵ m³ d^− 1 (20–74 L m^− 2 d^− 1), depending on river-discharge stage. Such calculated SGD estimates, which must include both a recirculated as well as fresh water component, are in close agreement with results obtained from a first-order watershed mass balance. Average submarine groundwater discharge rates yield NH₄⁺ and PO₄^− 3 flux estimates to the Loxahatchee River estuary that range from 62.7 to 1063.1 and 69.2 to 378.5 μmol m^− 2 d^− 1, respectively, depending on river stage. SGD-derived nutrient flux rates are compared to yearly computed riverine total N and total P load estimates.     

Effects of nitrogen and iron enrichments on phytoplankton communities in the Northwestern Indian Ocean

Nutrient-enrichment bottle experiments in the northwestern Indian Ocean surface waters were conducted to investigate phytoplankton growth following enrichments with either NH₄⁺, NO₃⁻, Fe or Fe + NO₃⁻. Stimulation of phytoplankton growth could be achieved by the addition of either NH₄⁺ or NO₃⁻ under the ambient Fe concentrations, but the most significant increases in Chl a, POC, and cell densities were observed in the Fe + NO₃⁻-amended culture. Iron addition caused more rapid responses of phytoplankton growth in the Fe + NO₃⁻ treatment than those in the NO₃⁻ and NH₄⁻ treatment. However, the Fe-enrichment treatment revealed minimal growth of phytoplankton because of severe major nutrient deficiency and was similar to the control treatment. Increases in the cell density of diatoms and spherical phytoplankton cells (< 10 μm) were significant in the NH₄⁺-enriched samples, whereas NO₃⁻ enrichment alone had little effect on the diatoms. Simultaneous addition of Fe and NO₃⁻ stimulated maximal growth of phytoplankton, in particular in diatoms, coccolithophorids and Phaeocystis type colonies. However, the dominance of coccolithophorids and Phaeocystis type colonies in the Fe + NO₃⁻ treatment may be interpreted as resulting from Si-limitation. The high N/P ratio for phytoplankton nutrient uptake in the N-amended culture indicates the possibility of some P-limited growth. From these results, we conclude that in the northwestern Indian Ocean, Fe and major nutrients are co-limiting phytoplankton production during the northeast monsoon. Iron appeared to affect the ability of phytoplankton to respond quickly to transient nutrient inputs.     

Nitrogen and phosphorus transport between Fourleague Bay, LA, and the Gulf of Mexico: the role of winter cold fronts and Atchafalaya River discharge

Nutrient fluxes were measured between Fourleague Bay, a shallow Louisiana estuary, and the Gulf of Mexico every 3 h between February 1 and April 30, 1994 to determine how high velocity winds associated with cold fronts and peak Atchafalaya River discharge influenced transport. Net water fluxes were ebb-dominated throughout the study because of wind forcing and high volumes of water entering the northern Bay from the Atchafalaya River. Flushing time of the Bay averaged <8 days; however, more rapid flushing occurred in response to northerly winds with approximately 56% of the volume of the Bay exported to the Gulf in 1 day during the strongest flushing event. Higher nitrate+nitrite (NO₂+NO₃), total nitrogen (TN), and total phosphorus (TP) concentrations were indicative of Atchafalaya River input and fluxes were greater when influenced by high velocity northerly winds associated with frontal passage. Net exports of NO₂+NO₃, TN, and TP were 43.5, 98.5, and 13.6 g s⁻¹, respectively, for the 89-day study. An average of 10.6 g s⁻¹ of ammonium (NH₄) was exported to the Gulf over the study; however, concentrations were lower when associated with riverine influence and wind-driven exports suggesting the importance of biological processes. Phosphate (PO₄) fluxes were nearly balanced over the study with fairly stable concentrations indicating a well-buffered system. The results indicate that the high energy subsidy provided by natural pulsing events such as atmospheric cold fronts and seasonal river discharge are efficient mechanisms of nutrient delivery to adjacent wetlands and nearshore coastal ecosystems and are important in maintaining coastal sustainability.     

Tidal time-scale variation in nutrient flux across the sediment–water interface of an estuarine tidal flat

We determined the range of the tidal variations in nutrient flux across the sediment–water interface and elucidated mechanisms of the flux variation in two estuarine intertidal flats (one sand, one mud) in northeastern Japan. Nutrient flux was measured using in situ light and dark chambers, which were incubated for 2 h, 2–6 times per day. Results showed that nutrient concentration in overlying water varied by tide and was also affected by sewage-treated water inflow. The nutrient fluxes responded quickly to the tidal variation in overlying water chemistry and the range of the variation in flux was as large as the seasonal-scale variation reported in previous studies. In the sand flat, salinity increase likely enhanced benthos respiration and led to increases in both O₂ consumption and PO₄³⁻ regeneration under low illumination, while benthic microalgae were likely to actively generate O₂, uptake PO₄³⁻ and suppress PO₄³⁻ release under high illumination (>900 μmol photons m⁻² s⁻¹). Also in the mud flat, PO₄³⁻ flux was related with O₂ flux, although the range of temporal variation in PO₄³⁻ flux was small. In both the flats, NH₄⁺ flux was always governed by NH₄⁺ concentration in the overlying water; either an increase in NH₄⁺ uptake or a decrease in NH₄⁺ release was observed as the NH₄⁺ concentration rose due to inflow of river water or input of sewage-treated water. Although NO₃⁻ tended to be released in both tidal flats when low NO₃⁻ concentration seawater dominated, their relationship was likely to be weakened under conditions of low oxygen consumption and suppressed denitrification. It is likely that tidal variation in nutrient flux is governed more by the nutrient concentration than other factors, such as benthic biological processes, particularly in the case where nutrient concentration in the overlying water is relatively high and with wide amplitude.     

Effects of irradiance on benthic and water column processes in a Gulf of Mexico estuary: Pensacola Bay, Florida, USA

We examined the effect of light on water column and benthic fluxes in the Pensacola Bay estuary, a river-dominated system in the northeastern Gulf of Mexico. Measurements were made during the summers of 2003 and 2004 on 16 dates distributed along depth and salinity gradients. Dissolved oxygen fluxes were measured on replicate sediment and water column samples exposed to a gradient of photosynthetically active radiation. Sediment inorganic nutrient (NH₄⁺, NO₃⁻, PO₄³⁻) fluxes were measured. The response of dissolved oxygen fluxes to variation in light was fit to a photosynthesis–irradiance model and the parameter estimates were used to calculate daily integrated production in the water column and the benthos. The results suggest that shoal environments supported substantial benthic productivity, averaging 13.6 ± 4.7 mmol O₂ m⁻² d⁻¹, whereas channel environments supported low benthic productivity, averaging 0.5 ± 0.3 mmol O₂ m⁻² d⁻¹ (±SE). Estimates of baywide microphytobenthic productivity ranged from 8.1 to 16.5 mmol O₂ m⁻² d⁻¹, comprising about 16–32% of total system productivity. Benthic and water column dark respiration averaged 15.2 ± 3.2 and 33.6 ± 3.7 mmol O₂ m⁻² d⁻¹, respectively Inorganic nutrient fluxes were generally low compared to relevant estuarine literature values, and responded minimally to light exposure. Across all stations, nutrient fluxes from sediments to the water column averaged 1.11 ± 0.98 mmol m⁻² d⁻¹ for NH₄⁺, 0.58 ± 1.08 mmol m⁻² d⁻¹ for NO₃⁻, 0.01 ± 0.09 mmol m⁻² d⁻¹ for PO₄³⁻. The results of this study illustrate how light reaching the sediments is an important modulator of benthic nutrient and oxygen dynamics in shallow estuarine systems.     

Simulation of upper-ocean biogeochemistry with a flexible-composition phytoplankton model: C, N and Si cycling in the western Sargasso Sea

We report the first application of a biogeochemical model in which the major elemental composition of the phytoplankton is flexible, and responds to changing light and nutrient conditions. The model includes two phytoplankton groups: diatoms and non-siliceous picoplankton. Both fix C in accordance with photosynthesis-irradiance relationships used in other models and take up NO₃⁻ and NH₄⁺ (and Si(OH)₄ for diatoms) following Michaelis-Menten kinetics. The model allows for light dependence of photosynthesis and NO₃⁻ uptake, and for the observed near-total light independence of NH₄⁺ uptake and Si(OH)₄ uptake. It tracks the resulting C/N ratios of both phytoplankton groups and Si/N ratio of diatoms, and permits uptake of C, N and Si to proceed independently of one another when those ratios are close to those of nutrient-replete phytoplankton. When the C/N or Si/N ratio of either phytoplankton group indicates that its growth is limited by N, Si or light, uptake of non-limiting elements is controlled by the content of the limiting element in accordance with the cell-quota formulation of Droop (J. Mar. Biol. Ass. U.K 54 (1974) 825).We applied this model to the Bermuda Atlantic Time-series Study (BATS) site in the western Sargasso Sea. The model was tuned to produce vertical profiles and time courses of [NO₃⁻], [NH₄⁺] and [Si(OH)₄] that are consistent with the data, by adjusting the kinetic parameters for N and Si uptake and the rate of nitrification. The model then reproduces the observed time courses of chlorophyll-a, particulate organic carbon and nitrogen, biogenic silica, primary productivity, biogenic silica production and POC export with no further tuning. Simulated C/N and Si/N ratios of the phytoplankton indicate that N is the main growth-limiting nutrient throughout the thermally stratified period and that [Si(OH)₄], although always limiting to the rate of Si uptake by diatoms, seldom limits their growth rate. The model requires significant nitrification in the upper 200 m to yield realistic time courses and vertical profiles of [NH₄⁺] and [NO₃⁻], suggesting that NO₃⁻ is not supplied to the upper water column entirely by physical processes. A nitrification-corrected f-ratio (f_NC), calculated for the upper 200 m as: (NO₃⁻ uptake—nitrification)/(NO₃⁻ uptake+NH₄⁺ uptake) has annual values ranging from only 0.05–0.09, implying that 90–95% of the N taken up annually by phytoplankton is supplied by biological regeneration (including nitrification) in the upper 200 m. Reported discrepancies between estimates of organic C export based on seasonal chemical changes and POC export measured at the BATS site can be almost completely resolved if there is significant regeneration of NO₃⁻ via organic-matter decomposition in the upper 200 m.     

Eutrophication and the rate of net nitrification in a coastal marine ecosystem

Rates of net nitrification were calculated for four large (13 m³) estuarine-based microcosms that had been subjected to inorganic nutrient enrichment. Calculated rates were based on two years of weekly nitrate and nitrite measurements and ranged from a maximum of 0·55 μmol NO₂₊₃⁻ produced l⁻¹ day⁻¹ in the control tank (no enrichment) to over 13 μmol NO₂₊₃⁻ produced l⁻¹ day⁻¹ in the most enriched tank (receiving 18·6 μmol NH₄ l⁻¹ day⁻¹). Almost all NO₂₊₃⁻ production was pelagic, little was benthic. Net NO₃⁻ production or net NO₂⁻ production dominated the net nitrification rates during different seasons. Good correlations were found between various oxidation rates and substrate concentrations. The calculated net nitrite production rates were 10 to 1000 times higher than previously reported rates for open ocean systems, demonstrating the potential importance of nitrification to estuarine systems.     

Nitrogen dynamics within a wind-driven eddy

Wind-driven cyclonic eddies are hypothesized to relieve nutrient stress and enhance primary production by the upward displacement of nutrient-rich deep waters into the euphotic zone. In this study, we measured nitrate (NO₃⁻), particulate carbon (PC), particulate nitrogen (PN), their stable isotope compositions (δ¹⁵N-NO₃⁻, δ¹³C-PC and δ¹⁵N-PN, respectively), and dissolved organic nitrogen (DON) within Cyclone Opal, a mature wind-driven eddy generated in the lee of the Hawaiian Islands. Sampling occurred in March 2005 as part of the multi-disciplinary E-Flux study, approximately 4–6 weeks after eddy formation. Integrated NO₃⁻ concentrations above 110 m were 4.8 times greater inside the eddy (85.8±6.4 mmol N m⁻²) compared to the surrounding water column (17.8±7.8 mmol N m⁻²). Using N-isotope derived estimates of NO₃⁻ assimilation, we estimated that 213±59 mmol m⁻² of NO₃⁻ was initially injected into the upper 110 m Cyclone Opal formation, implying that NO₃⁻ was assimilated at a rate of 3.75±0.5 mmol N m⁻² d⁻¹. This injected NO₃⁻ supported 68±19% and 66±9% of the phytoplankton N demand and export production, respectively. N isotope data suggest that 32±6% of the initial NO₃⁻ remained unassimilated. Self-shading, inefficiency in the transfer of N from dissolved to particulate export, or depletion of a specific nutrient other than N may have led to a lack of complete NO₃⁻ assimilation. Using a salt budget approach, we estimate that dissolved organic nitrogen (DON) concentrations increased from eddy formation (3.8±0.4 mmol N m⁻²) to the time of sampling (4.0±0.09 mmol N m⁻²), implying that DON accumulated at rate of 0.83±1.3 mmol N m⁻² d⁻¹, and accounted for 22±15% of the injected NO₃⁻. Interestingly, no significant increase in suspended PN and PC, or export production was observed inside Cyclone Opal relative to the surrounding water column. A simple N budget shows that if 22±15% of the injected NO₃⁻ was shunted into the DON pool, and 32±6% is unassimilated, then 46±16% of the injected NO₃⁻ remains undocumented. Alternative loss processes within the eddy include lateral exchange of injected NO₃⁻ along isopycnal surfaces, remineralization of PN at depth, as well as microzooplankton grazing. A 9-day time series within Cyclone Opal revealed a temporal depletion in δ¹⁵N-PN, implying a rapid change in the N source. A change in NO₃⁻ assimilation, or a shift from NO₃⁻ fueled growth to assimilation of a ¹⁵N-deplete N source, may be responsible for such observations.     

Seasonal variations in river discharge and nutrient export to a Northeastern Pacific estuary

Seasonal variations in dissolved nitrogen and silica loadings were related to seasonal variability in river discharge. Dissolved nutrient concentrations measured weekly at three stations in the Yaquina River, Oregon from 1999 through 2001, and then monthly in 2002 were used as the basis for developing a nutrient loading regression as part of a larger agency program for evaluating nutrient processes. Because realistic models of nutrient transport require dense data sets to capture both long and short term fluctuations in nutrient concentrations, data at one freshwater station also were collected hourly for the same years using an in-stream monitor.The effects of storm events on dissolved nutrient transport were examined during three storms, including one in a high rainfall-discharge year, and two in average years, one of which followed a drought year. During the drought year (WY2001), total dissolved nitrate input was considerably less than in wetter years. Dissolved nitrate concentrations, however, were unusually high in the first winter storm runoff after the drought. The freshwater dissolved nitrate nitrogen loads varied from 40,380 kg day⁻¹ during a high-flow storm event to 0.11 kg day⁻¹ during late summer, low flow conditions. Dissolved silica dynamics differed from those of nitrate because during storm events, silica concentrations in the Yaquina River decreased to near zero at the storm height, probably due to dilution by near surface or overland flow, and later recovered.During the time interval studied, over 94% of the dissolved nitrate and silica were transported from the watershed during the winter months of greater rainfall, indicating that seasonality and river flow are primary factors when considering nutrient loadings from this watershed system.     

Inorganic and organic nitrogen cycling in the Southern California Bight

On the basis of mass balance calculations performed for nitrogen (N) uptake experiments in the Southern California Bight (SCB), it has been suggested that a significant portion of dissolved inorganic N (DIN) uptake results in the production of dissolved organic N (DON). To investigate this process, the fate of ammonium (NH₄⁺) and nitrate (NO₃⁻) uptake was quantified within the euphotic zone at three coastal stations in the SCB using ¹⁵N tracer techniques. Several trends in the fate of DIN and the production of DON were observed. First, production of particulate N (PN), from both NH₄⁺ and NO₃⁻, was quantitatively more important in near surface waters, while DON release dominated within the nitracline. Second, the percentage of gross N uptake released as DON was generally higher when NO₃⁻, rather than NH₄⁺, was the substrate. Third, the percentage of N released as DON was higher at night, relative to the day. Fourth, rates of DON release were significantly correlated to NH₄⁺ regeneration, suggesting that similar mechanisms are responsible for both processes—presumably grazing. The results of this study indicate that the DON pool is a sink for DIN uptake on the time scale of hours. One implication of this finding is that new production estimates based on ¹⁵NO₃⁻ uptake rates will likely underestimate particle flux out of the surface layer because the rate of NO₃⁻ uptake is underestimated due to loss of DO¹⁵N during the incubation. On time scales of months to years, however, the N that is taken up as NO₃⁻ and released as DON will likely contribute to export flux via incorporation of the dissolved phase during seasonal mixing into sinking particles or transport. The export of DON on these time scales argues for the use of gross uptake rates to calculate f-ratios.     

Effect of changes in water salinity on ammonium, calcium, dissolved inorganic carbon and influence on water/sediment dynamics

The effect of a sudden increase in salinity from 10 to 37 in porewater concentration and the benthic fluxes of ammonium, calcium and dissolved inorganic carbon were studied in sediments of a small coastal lagoon, the Albufera d'Es Grau (Minorca Island, Spain). The temporal effects of the changes in salinity were examined over 17 days using a single diffusion-reaction model and a mass-balance approach. After the salinity change, NH₄⁺-flux to the water and Ca-flux toward sediments increased (NH₄⁺-flux: 5000–3000 μmol m⁻² d⁻¹ in seawater and 600/250 μmol m⁻² d⁻¹ in brackish water; Ca-flux: −40/−76 meq m⁻² d⁻¹ at S=37 and −13/−10 meq m⁻² d⁻¹ at S=10); however, later NH₄⁺-flux decreased in seawater, reaching values lower than in brackish water. In contrast, Ca-flux presented similar values in both conditions. The fluxes of dissolved inorganic carbon, which were constant at S=10 (55/45 mmol m⁻² d⁻¹), increased during the experiment at S=37 (from 30 mmol m⁻² d⁻¹ immediately after salinity increase to 60 mmol m⁻² d⁻¹ after 17 days).In brackish conditions, NH₄⁺ and Ca²⁺ fluxes were consistent with a single diffusion-reaction model that assumes a zero-order reaction for NH₄⁺ production and a first-order reaction for Ca²⁺ production. In seawater, this model explained the Ca-flux observed, but did not account for the high initial flux of NH₄⁺.The mass balance for 17 days indicated a higher retention of NH₄⁺ in porewater in the littoral station in seawater conditions (9.5 mmol m⁻² at S=37 and 1.6 mmol m⁻² at S=10) and a significant reduction in the water consumption at both sites (5 mmol m⁻² at S=37; 35/23 mmol m⁻² at S=10). In contrast, accumulation of dissolved inorganic carbon in porewater was lower in seawater incubations (−10/−1 meq m⁻² at S=37; 50/90 meq m⁻² at S=10) and was linked to a higher efflux of CO₂ to the atmosphere, because of calcium carbonate precipitation in water (675/500 meq m⁻²). These results indicate that increased salinity in shallow coastal waters could play a major role in the global carbon cycle.     

Benthic fluxes of nitrogen in the tidal reaches of a turbid, high-nitrate sub-tropical river

Benthic fluxes of dissolved inorganic nitrogen (NO₃⁻ and NH₄⁺), dissolved organic nitrogen (DON), N₂ (denitrification), O₂ and TCO₂ were measured in the tidal reaches of the Bremer River, south east Queensland, Australia. Measurements were made at three sites during summer and winter. Fluxes of NO₃⁻ were generally directed into the sediments at rates of up to −225 μmol N m⁻² h⁻¹. NH₄⁺ was mostly taken up by the sediments at rates of up to −52 μmol N m⁻² h⁻¹, its ultimate fate probably being denitrification. DON fluxes were not significant during winter. During summer, fluxes of DON were observed both into (−105 μmol m⁻² h⁻¹) and out of (39 μmol m⁻² h⁻¹) the sediments. Average N₂ fluxes at all sampling sites were similar during summer (162 μmol N m⁻² h⁻¹) and winter (153 μmol N m⁻² h⁻¹). Denitrification was fed both by nitrification within the sediment and NO₃⁻ from the water column. Sediment respiration rates played an important role in the dynamics of nitrification and denitrification. NO₃⁻ fluxes were significantly related to TCO₂ fluxes (p<0.01), with a release of NO₃⁻ from the sediment only occurring at respiration rates below 1000 μmol C m⁻² h⁻¹. Rates of denitrification increased with respiration up to TCO₂ fluxes of 1000 μmol C m⁻² h⁻¹. At sediment respiration rates above 1000 μmol C m⁻² h⁻¹, denitrification rates increased less rapidly with respiration in winter and declined during summer. On a monthly basis denitrification removed about 9% of the total nitrogen and 16% of NO₃⁻ entering the Bremer River system from known point sources. This is a similar magnitude to that estimated in other tidal river systems and estuaries receiving similar nitrogen loads. During flood events the amount of NO₃⁻ denitrified dropped to about 6% of the total river NO₃⁻ load.     

The effect of a tidal cycle on the dynamics of nutrients in a tidal estuary in the Seto Inland Sea, Japan

A 24 hour time series survey was carried out during a spring tide (tidal range ca.2 m) of May 1995 on a tidal estuary in the Seto Inland Sea, Japan, in the context of an integrated program planned to quantify the dynamics of biophilic elements (carbon, nitrogen and phosphorus) and the roles played by the macrobenthos on the processes. Three stations were set along a transect line of about 1.4 km, which linked the river to the rear to the innermost part of the subtidal zone. Every hour, at each station, measurements were made of surface water temperature, salinity and dissolved oxygen concentration, and surface water was collected for the determination of nutrients [NH₄ ⁺−N, (NO₃ ⁻+NO₂ ⁻)−N, PO₄ ³⁻−P and Si (OH)₄−Si]. During the ebb flow, riverine input of silicate and nitrate+nitrite significantly increased the concentrations of both the intertidal and the subtidal stations. Conversely, during the high tide, river nutrient concentrations were lowered by the mixing of fresh water with sea water. As a result, best (inverse) correlations were found at the river station for salinity against silicate (y=-2.9 Sal.+110.7,r ²=0.879) and nitrate+nitrite (y=-1.3 Sal.+48.4,r ²=0.796). In contrast, ammonium nitrogen concentrations were higher at intermediate salinities. Indeed, no significant correlation was found between salinity and ammonium. The effect of the macrobenthos, which is abundant on the intertidal flat, is discussed as a biological component that influences the processes of nutrient regeneration within the estuary. The effect of the tidal amplitude is an important one in determining the extent of the variations in nutrient concentrations at all three stations, which were stronger between the lower low tide and the higher high tide.     

Seasonal dynamics of inorganic nutrients and phytoplankton biomass in the NW Alboran Sea

The seasonal dynamics of inorganic nutrients and phytoplankton biomass (chlorophyll a), and its relation with hydrological features, was studied in the NW Alboran Sea during four cruises conducted in February, April, July and October 2002. In the upper layers, the seasonal pattern of nutrient concentrations and their molar ratios (N:Si:P) was greatly influenced by hydrological conditions. The higher nutrient concentrations were observed during the spring cruise (2.54 μM NO₃⁻, 0.21 μM PO₄³⁻ and 1.55 μM Si(OH)₄, on average), coinciding with the increase of salinity due to upwelling induced by westerlies. The lowest nutrient concentrations were observed during summer (<0.54 μM NO₃⁻, 0.13 μM PO₄³⁻ and 0.75 μM Si(OH)₄, on average), when the lower salinities were detected. Nutrient molar ratios (N:Si:P) followed the same seasonal pattern as nutrient distribution. During all the cruises, the ratio N:P in the top 20 m was lower than 16:1, indicating a NO₃⁻ deficiency relative to PO₄³⁻. The N:P ratio increased with depth, reaching values higher than 16:1 in the deeper layers (200–300 m). The N:Si ratio in the top 20 m was lower than 1:1, excepting during spring when N:Si ratios higher than 1:1 were observed in some stations due to the upwelling event. The N:Si ratio increased with depth, showing a maximum at 50–100 m (>1.5:1), which indicates a shift towards Si-deficiency in these layers. The Si:P ratio was much lower than 16:1 throughout the water column during the four cruises. In general, the spatial and seasonal variation of phytoplankton biomass showed a strong coupling with hydrological and chemical fields. The higher chlorophyll a concentrations at the depth of the chlorophyll maximum were found in April (2.57 mg m⁻³ on average), while the lowest phytoplankton biomass corresponded to the winter cruise (0.74 mg m⁻³ on average). The low nitrate concentrations together with the low N:P ratios found in the upper layers (top 20 m) during the winter, summer and autumn cruises suggest that N-limitation could occur in these layers during great part of the year. However, N-limitation during the spring cruise was temporally overcome by nutrient enrichment caused by an intense wind-driven upwelling event.     

The distribution and δC of dissolved organic carbon and its humic fraction in estuaries of southeastern USA

In this study, we examine the distribution and carbon stable isotope signature of dissolved organic carbon (DOC) and humic substances (HS) along a salinity gradient in the Altamaha and Satilla River estuaries. The maximum DOC concentrations in the Altamaha and Satilla were 10 and 29 mg C l⁻¹, respectively, though concentrations were similar at the mouth of both estuaries. There was a decrease in HS content of DOC from 50 to 80% at the head of the estuaries to 10% at salinities higher than 30‰. The δ¹³C DOC varied between −25.5 and −19‰ and between −27 and −21‰ in the Altamaha and Satilla estuaries, respectively. The tendency towards more depleted δ¹³C DOC in the Satilla, especially in the lower salinity portion of this estuary, suggests greater terrestrial inputs in the Satilla than in the Altamaha. Seasonal fluctuations were observed in the form of increased (two to three times) range in DOC concentration, heavier δ¹³C DOC and increased proportion of estuarine–marine-derived DOC (average enrichment of δ¹³C DOC from +1 to +2) during low river flow (July–October). The δ¹³C HS in both rivers showed a similar trend, but was consistently more depleted than DOC, with an average range from −28 to −24.5‰. This suggests that HS have larger proportions of terrestrial components (a maximum of >60% at the mouth of the estuary) than DOC. The less depleted δ¹³C values of DOC in comparison with HS indicate a different source for the non-humic (non-HS) component of DOC (range in δ¹³C non-HS, −22 to −16‰). That source could either be the decomposition of detrital material derived from saltmarsh environments or microalgal-derived DOC of estuarine or marine origins.     

Microscale gradients of planktonic microbial communities above the sediment surface in a mangrove estuary

The microscale (1 and 4 cm sampling resolution) distributions of chemical (O₂, NH₃, NO₃⁻, NO₂⁻, PO₄³⁻) and biological (Chl a, phytoplankton, bacterioplankton, viruses) parameters were measured in the 16 cm of water immediately overlaying the sediment-water interface (SWI) within a temperate mangrove estuary in South Australia during December 2003 and March 2004. Shear velocities (u_*) during the time of sampling were very low (<0.1 cm s⁻¹), and we consequently predict that resuspension of organisms and materials was negligible. In December 2003, profiles were often characterised by strong gradients in nutrients and organisms, with the highest concentrations often observed within 0.5 cm of the SWI. Microscale patterns in O₂, NH₃, NO₃⁻ and NO₂⁻ indicated that a variety of anaerobic and aerobic transformation processes probably occurred at the SWI and within profiles. Strong gradients in PO₄³⁻ were indicative of nutrient flux across the SWI as a consequence of degradation processes in the sediments. Pico- and nanophytoplankton concentrations were strongly correlated (p < 0.01) to PO₄³⁻, and exhibited 12- and 68-fold changes in abundance, respectively, with highest concentrations observed nearest to the SWI. Several bacterial subpopulations were discriminated using flow cytometry and significant shifts in the ‘cytometric structure’ of the bacterial community were observed within microscale profiles. Two populations of viruses were correlated to the phytoplankton and low DNA (LDNA) bacteria, and each exhibited elevated concentrations within 0.5 cm of the SWI. In March 2004, microscale distributions of O₂ and nutrients were more homogenous than in December 2003, and dissimilar microbial community structure and patterns were observed above the SWI. The patterns observed here support the prediction that benthic processes can strongly influence the ecology of planktonic communities in the overlaying water, and provide further evidence for the existence of microscale variability amongst communities of aquatic microorganisms.     

Carbon and nitrogen primary productivity in three North Carolina estuaries

Uptake of inorganic carbon and ammonium by the plankton community of three North Carolina estuaries was measured using ¹⁴C and ¹⁵N isotope methods. At 0% light, C appeared to be lost via respiration, and at increasing light levels uptake of inorganic carbon increased linearly, saturated (mean I_k = 358±30 μEin m⁻² s⁻¹), and frequently showed inhibition at the highest light intensities. At 0% light NH₄⁺ uptake was significantly greater than zero and was frequently equivalent to uptake in the light (light independent); at increasing light levels NH₄⁺ uptake saturated (mean I_k = 172±44 μEin m⁻² s⁻¹) and frequently indicated strong inhibition. Light-saturated uptake rates of inorganic carbon and NH₄⁺ were a function of chlorophyll a (r² = 0·7−0·9); average assimilation numbers were 625 nmol CO₂ (μg chl. a)⁻¹ h⁻¹ and 12·9 nmol NH₄⁺ (μg chl. a)⁻¹ h⁻¹ and were positively correlated with temperature (r² = 0·3−0·7). The ratio of dark to light-saturated NH₄⁺ uptake tended to be near 1·0 for large algal populations at low NH₄⁺ concentrations, indicating near light independence of uptake; whereas the ratio was lower for the opposite conditions. These data are interpreted as indicative of nitrogen stress, and it is suggested that uptake of NH₄⁺ deep in the euphotic zone and at night are mechanisms for balancing the C:N of cellular pools. A 24-h study using summed short-term incubations confirmed this; the cumulative C:N of CO₂ and NH₄⁺ uptake during the daylight period was 10–20, whereas over the 24-h period the ratio was 6 due to dark NH₄⁺ uptake. Annual carbon and nitrogen primary productivity were respectively estimated as 24 and 4·0 mol m⁻² year⁻¹ for the South River estuary, 42 and 7·3 mol m⁻² year⁻¹ for the Neuse River estuary, and 9·6 and 1·6 mol m⁻² year⁻¹ for the Newport River estuary.     

Temporal variability and characterization of physicochemical properties in the neritic area of Sagami Bay,Japan

Seasonal and interannual variations in physicochemical properties were investigated in the neritic area of Sagami Bay, Kanagawa, Japan, from December 2000 to December 2005. Physicochemical properties (i.e. temperature, salinity, density, dissolved oxygen and dissolved inorganic nutrient concentration) revealed clear seasonal variations, which were similar to each other during all 5 years. Temperature, salinity and dissolved inorganic nutrients showed rapid, drastic variations within a few days and/or weeks. These variations are related to sea levels, principally due to the shifting effects of the Kuroshio Current axis: they were strongly affected by the Kuroshio Water and other waters, when sea level difference was greater than ca. 35 cm and lower than ca. 15 cm, respectively. Temperature difference (DF _T ) increased with sea level difference, and the difference of salinity and dissolved inorganic nutrients (NH₄ ⁺-N, NO₃ ⁻+NO₂ ⁻-N, NH₄ ⁺+NO₃ ⁻+NO₂ ⁻-N, PO₄ ³⁻-P and SiO₂-Si) increased and decreased with DF _T , respectively. All these correlations are significant. Total dissolved inorganic nitrogen (N), phosphate (P) and silicate (Si) revealed seasonal variations in the ranges of 0.57–16.08, 0.0070–0.91 and 0.22–46.38 μM, respectively. From the regression equations between these elements allowed the following relation to be obtained; Si:N:P = 14.8:13.4:1. Dissolved inorganic nutrients were characterized by Si and/or P deficiency, especially in the upper layer (0–20 m depth) during summer. Single and/or combined elements are discussed on the basis of potential and stoichiometric nutrient limitations, which could restrict phytoplankton (diatom) growth as a limiting factor.