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.     

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.     

A nitrate and silicate budget in the equatorial Pacific Ocean: a coupled physical–biological model study

A coupled physical–biological model was developed to simulate the low-silicate, high-nitrate, and low-chlorophyll (LSHNLC) conditions in the equatorial Pacific Ocean and used to compute a detailed budget in the Wyrtki box (5°N–5°S, 180–90°W) for the major sources and cycling of nitrogen and silicon in the equatorial Pacific. With the incorporation of biogenic silicon dissolution, NH₄ regeneration from organic nitrogen and nitrification of ammonia in the model, we show that silicon recycling in the upper ocean is less efficient than nitrogen. As the major source of nutrients to the equatorial Pacific, the Equatorial Undercurrent provides slightly less Si(OH)₄ than NO₃ to the upwelling zone, which is defined as 2.5°N–2.5°S. As a result, the equatorial upwelling supplies less Si(OH)₄ than NO₃ into the euphotic zone in the Wyrtki box, having a Si/N supply ratio of about 0.85 (2.5 vs. 2.96 mmolm⁻² day⁻¹). More Si(OH)₄ than NO₃ is taken up with a Si/N ratio of 1.17 (2.72 vs. 2.33 mmolm⁻² day⁻¹) within the euphotic zone. The difference between upwelling supply and biological uptake is balanced by nutrient regeneration and horizontal advection. Excluding regeneration, the net silicate and nitrate uptakes are nearly equal (1.76 vs. 1.84 mmolm⁻² day⁻¹). However, biogenic silica export production is slightly higher than organic nitrogen (1.74 vs. 1.59 mmolm⁻² day⁻¹) following a 1.1 Si/N ratio. In the central equatorial Pacific, low silicate concentrations limit diatom growth; therefore non-diatom new production accounts for most of the new production. Higher silicate supply in the east maintains elevated diatom growth rates and new production associated with diatoms dominate upwelling zone. In contrast, the new production associated with small phytoplankton is nearly constant or decreases eastward along the equator. The total new production has a higher rate in the east than in the west, following the pattern of surface silicate. This suggests that silicate regulates the diatom production, total new production, and thereby carbon cycle in this area. The modeled mean primary production is 48.4 mmolCm⁻² day⁻¹, representing the lower end of direct field measurements, while new production is 15.0 mmolCm⁻² day⁻¹, which compares well with previous estimates.     

Nitrogen uptake kinetics in the Ross Sea, Antarctica

The first estimates of uptake kinetic parameters for NH₄⁺, NO₃⁻, and urea in the Ross Sea, Antarctica were measured on three cruises during austral late winter–early spring 1996 (pre-bloom), late spring 1997 (bloom development), and summer 1997 (bloom decline). Nitrogen (N) uptake experiments were conducted with water collected at the 50% light penetration depth using trace-metal clean protocols and ¹⁵N tracer techniques. At all sites, ambient NO₃⁻ concentrations ranged from 5.8 to 30.5 μg-at N l⁻¹ and silicic acid concentrations were greater than 62.0 μg-at Si l⁻¹. The following trends were observed. First, based on maximum uptake rates (V_max), apparent N utilization followed the order NO₃⁻>NH₄⁺>urea during the pre-bloom and bloom development cruises. During the summer cruise, as the bloom was declining, the apparent order of utilization was NH₄⁺>NO₃⁻>urea. Second, evidence for possible repression of NO₃⁻ uptake by elevated NH₄⁺ concentrations was only observed at one site. Third, the kinetic parameters of NH₄⁺ uptake rates corrected for isotope dilution were compared with the kinetic parameters determined from uncorrected rates. In this comparison, the measure of substrate affinity, α (α=V_max/K_s) increased by an average of 4.6-fold when rates were corrected for isotope dilution, but values of V_max remained unchanged. Fourth, using bacterial production data, the magnitude of bacterial N uptake was estimated. Assuming that all bacterial N demands were met with NH₄⁺, the estimated bacterial portion of NH₄⁺ uptake ranged from <1%, when the ratio of bacteria to autotrophic biomass was low, to 35%, when bacterial abundance and biomass were highest. Finally, dramatic changes in NH₄⁺ uptake capacity were observed at one station (Stn. O), where kinetic parameters were measured during all three cruises. We hypothesize that a mutualistic relationship exists between phytoplankton and heterotrophic bacteria, and that the creation of microzones of high NH₄⁺ concentrations contributed to the changes seen at this station.     

The effects of light and nutrients on rates of ammonium transformation in a eutrophic river

Rates for nitrification, phytoplankton uptake of ammonium, and regeneration of ammonium were measured in the Delaware River as functions of irradiance and nutrient concentrations, using ¹⁵N labeling methods. Phytoplankton uptake increased and nitrification rates declined with increased light intensity. The irradiance level required for maximum uptake by phytoplankton was similar to that for maximal inhibition of nitrification (about 300μEm⁻² s⁻¹). Daily, water-column averaged rates, calculated by integration of the observed rate-intensity relationships, indicate that light plays a key role in regulating the balance between oxidation of NH₄⁺ by bacteria and assimilation by phytoplankton in the Delaware. The results show that uptake of ammonium by phytoplankton in the dark may exceed uptake in the light in optically thick systems.     

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.     

Seasonal and interannual variability in phytoplankton and nutrient dynamics along Line P in the NE subarctic Pacific

We analysed mixed-layer seasonal and interannual variability in phytoplankton biomass and macronutrient (NO₃ and Si(OH)₄) concentrations from three decades of observations, and nitrogen uptake rates from the 1990s along Line P in the NE subarctic Pacific. Chlorophyll a concentrations near 0.35 mg m⁻³ were observed year-round along Line P except at the nearshore station (P4) where chlorophyll a concentrations in spring were on average 2.4 times the winter values. In contrast, the temporal variability in carbon-to-chlorophyll ratios at the two main end members of Line P (P4 and OSP) was high. Large seasonal and interannual variability in NO₃ and Si(OH)₄ concentration were observed along Line P. Highest upper mixed-layer (top 15 m) nutrient concentrations occurred on the continental shelf in late summer and early fall due to seasonal coastal upwelling. Beyond the shelf, maximum nutrient concentrations increased gradually offshore, and were highest in late winter and early spring due to mixing by winter storms. Interannual variations in upper mixed-layer nutrient concentrations beyond the shelf (>128°W) were correlated with E-W winds and the PDO since 1988 but were not correlated with either climate index between 1973 and 1981. Despite differences in nutrient concentration, nutrient utilization (ΔNO₃ and ΔSi(OH)₄) during the growing season were about 7.5 μM at all offshore stations. Variations in ΔNO₃ were correlated with those of ΔSi(OH)₄. The annual cycle of absolute NO₃ uptake (ρNO₃) and NH₄ uptake (ρNH₄) rates by phytoplankton in the upper mixed-layer showed a weak increasing trend from winter to spring/summer for the period 1992-1997. Rates were more variable at the nearshore station (P4). Rates of ρNO₃ were low along the entire line despite abundant NO₃ and low iron (Fe), at the offshore portion of Line P and sufficient Fe at the nearshore station (P4). As a result, new production contributed on average to only 32 ± 15% of the total nitrogen (N) uptake along Line P. NO₃ utilization in the NE subarctic Pacific is probably controlled by a combination of environmental variables, including Fe, light and ambient NH₄ levels. Elevated ambient NH₄ concentrations seem to decrease the rates of new production (and f-ratios) in surface waters of the oceanic subarctic NE Pacific. Contrary to expectation, phytoplankton biomass, nutrient utilization (ΔNO₃ and ΔSi(OH)₄), and nitrogen uptake (ρNO₃ + ρNH₄) varied relatively little along Line P, despite significant differences in the factors controlling phytoplankton composition assemblages and production. Future studies would benefit from including other variables, especially light limitation, to improve our understanding of the seasonal and interannual variability in phytoplankton biomass and nutrients in this region.     

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.     

Benthic nitrate biogeochemistry affected by tidal modulation of Submarine Groundwater Discharge (SGD) through a sandy beach face, Ria Formosa, Southwestern Iberia

To investigate both the role of tides on the timing and magnitude of Submarine Groundwater Discharge (SGD), and the effect on benthic nitrogen biogeochemistry of nitrate-enriched brackish water percolating upwards at the seepage face, we conducted a study of SGD rates measured simultaneously with seepage meters and mini-piezometers, combined with sets (n = 39) of high resolution in-situ porewater profiles describing NH₄⁺, NO₃⁻, Si(OH)₄ and salinity distribution with depth (0–20 cm). Sampling took place during two consecutive spring tidal cycles in four different months (November 2005, March, April and August 2006) at a backbarrier beach face in the Ria Formosa lagoon, southern Portugal. Our results show that the tide is one of the major agents controlling the timing and magnitude of SGD into the Ria Formosa. Intermittent pumping of brackish, nitrate-bearing water at the beach face through surface sediments changed both the magnitudes and depth distributions of porewater NH₄⁺ and NO₃⁻ concentrations. The most significant changes in nitrate and ammonium concentrations were observed in near-surface sediment horizons coinciding with increased fraction of N in benthic organic matter, as shown by the organic C:N ratio. On the basis of mass balance calculations executed on available benthic profiles, providing ratios of net Ammonium Production Rate (APR) to Nitrate Reduction Rate (NRR), coupled to stoichiometric calculations based on the composition of organic matter, potential pathways of nitrogen transformation were speculated upon. Although the seepage face occasionally contributes to reduce the groundwater-borne DIN loading of the lagoon, mass balance analysis suggests that a relatively high proportion of the SGD-borne nitrogen flowing into the lagoon may be enhanced by nitrification at the shallow (1–3 cm) subsurface and modulated by dissimilatory nitrate reduction to ammonium (DNRA).     

Influence of light intensity on diatom physiology and nutrient dynamics in the Oyashio region

The spring diatom bloom characterizes the plankton and nutrient dynamics in the Oyashio region, the westernmost part of the subarctic Pacific. Previous studies have shown that NO₃ was not depleted during the spring bloom, and an increase in the consumption ratio of Si(OH)₄ to NO₃ (ΔSi(OH)₄:ΔNO₃) was observed as the spring bloom progressed. The increase in ΔSi(OH)₄:ΔNO₃ has been suggested to be caused by growth stresses of diatoms, e.g. light limitation by self-shading. In the present study, incubation experiments of sea-surface water from the Oyashio region under saturated irradiance showed that NO₃ was depleted first and ΔSi(OH)₄:ΔNO₃ was more or less constant until the NO₃ depletion occurred. The increase in ΔSi(OH)₄:ΔNO₃ was observed after the NO₃ depletion had occurred in contrast with the field observation. This result of the increase in ΔSi(OH)₄:ΔNO₃ under saturated irradiance after NO₃ depletion suggests that the in situ increase in ΔSi(OH)₄:ΔNO₃ before the NO₃ depletion might be caused by light limitation for diatoms. Responses to a reduction in irradiance were examined using diatom species isolated from the Oyashio region. Variable responses to a reduced irradiance were observed for cell specific C, N, Si and chlorophyll a (Chl) contents. However, the examined diatom species showed similar tendencies for increases in Si:C and Si:N and decreases in C:Chl ratios with the reduction in irradiance. We conclude that light limitation changes the uptake ratio of nutrients and the elemental composition of diatoms and that light limitation is one of the factors influencing the physiology of diatoms and nutrient dynamics in the Oyashio region during the spring bloom.     

Seasonal variability in nitrogenous nutrition of phytoplankton assemblages in the northeastern subarctic Pacific Ocean

Nitrogen uptake rates, and physical, chemical and biological characteristics of the euphotic zone were studied in winter, spring and late summer during the period 1992–1994 along a transect (Line P) extending from the continental slope off the southwest corner of Vancouver Island (British Columbia, Canada; station P4; 49°N, 127°W) to open waters in the NE Pacific (OSP; 50°N, 145°W). Nitrate (NO₃⁻) and silicic acid (Si(OH)₄) concentrations increased offshore during every season. Lowest NO₃⁻ and Si(OH)₄ values were observed during late summer and spring, and highest during winter throughout the euphotic zone. For spring and late summer, surface depletion of NO₃⁻ was observed at the inshore end of the transect, while offshore concentrations were never limiting for phytoplankton growth. Silicic acid was never depleted at any depth or station during the period covered by this study. Ammonium (NH₄⁺) and urea concentrations exhibited a patchy distribution along the transect, with no seasonal variations. Chlorophyll a and particulate nitrogen did not show a consistent longitudinal pattern from year to year. In general, the highest concentrations of chlorophyll a and particulate nitrogen were measured during the late summer cruises, with lower values in spring and lowest in winter. Phytoplankton assemblages were numerically dominated by flagellates <5 μm throughout the water column on each cruise transect. Ammonium, urea and NO₃⁻ uptake rates represented on average 55, 24 and 21% of the depth-integrated total nitrogen uptake, both longitudinally and seasonally; hence, phytoplankton utilized nitrogen in the following order: NH₄⁺>urea>NO₃⁻ along Line P. Ammonium may have inhibited the uptake rates of NO₃⁻ and urea. Urea uptake rates were lower than those of NH₄⁺, but higher values were occasionally observed at a few depths along the transect, particularly during the spring of 1993. Depth-integrated NH₄⁺ uptake rates were generally higher inshore, while NO₃⁻ uptake rates showed higher values offshore during most seasons. In contrast, urea uptake rates did not exhibit a consistent longitudinal trend. The depth-integrated f-ratio ranged from 0.05 to 0.37 with an average of 0.21 for all stations and cruises, and was overestimated on average by 36% when urea was excluded from the calculation. On a yearly basis, primary productivity in the NE subarctic Pacific was based on regenerated nitrogen.     

Equilibrium and structural studies of silicon(IV) and aluminium(III) in aqueous solution. II. Formation constants for the monosilicate ions SiO(OH)3 and SiO2(OH)2. A precision study at 25°C in a simplified seawater medium

The hydrolysis of silicic acid, Si(OH)₄, was studied in a simplified seawater medium (0.6 M Na(Cl)) at 25°C. The measurements were performed as potentiometric titrations (hydrogen electrode) in which OH⁻ was generated coulometrically. The total concentration of Si(OH)₄, B, and log[H⁺] were varied within the limits 0.00075 B 0.008 M and 2.5 -log[H⁺] 11.7, respectively. Within these ranges the formation of SiO(OH)₃⁻ and SiO₂(OH)₂²⁻ with formation constants log β₋₁₁(Si(OH)₄ SiO(OH)₃⁻ + H⁺) = −9.472 ±0.002 and log β₋₂₁(Si(OH)₄ SiO₂(OH)₂²⁻ + 2H⁺) = −22.07 ± 0.01 was established. With B > 0.003 M polysilicate complexes are formed, however, with -log[H⁺] 10.7 their formation does not significantly affect the evaluated formation constants. Data were analyzed with the least squares computer program LETAGROPVRID.     

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.     

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.     

Diatoms grow faster using ammonium in rapidly flushed eutrophic Dokai Bay,Japan

The importance of the nitrogen source for phytoplankton growth in a highly eutrophic embayment, Dokai Bay, was investigated. The DIN concentration often exceeded 100 μM of which 40–70% was NH₄ ⁺. During two incubation experiments, the natural assemblage of mainly diatoms took up NH₄ ⁺ instead of NO₃ ⁻. The growth of two Skeletonema species isolated in Dokai Bay were significantly faster on NH₄ ⁺ (1.86 and 1.27 div. d⁻¹ respectively) than on NO₃ ⁻ (1.55 and 1.04 div. d⁻¹ respectively). Our results indicated that these diatoms could grow faster by using NH₄ ⁺ compared to NO₃ ⁻ in this eutrophic bay.     

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.     

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.     

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.     

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.     

Close coupling between ammonium uptake by phytoplankton and excretion by Antarctic krill, Euphausia superba

In this study we examined the hypothesis that, under conditions of replete macronutrients and iron in the Southern Ocean, phytoplankton abundance and specific N uptake rates are influenced strongly by the processes of grazing and NH₄ regeneration. NH₄ and NO₃ uptake rates by marine phytoplankton were measured to the northeast and northwest of the island of South Georgia during January-February 1998. Mean specific uptake rate for NO₃ (vNO₃) was 0.0026 h⁻¹ (range 0.0013-0.0065 h⁻¹) and for NH₄ (vNH₄) was 0.0097 h⁻¹ (0.0014-0.0376 h⁻¹). vNH₄ was related positively with NH₄ availability, which ranged from 0.1 to 1.5 mmol m⁻³ within the upper mixed layer. Ambient NH₄ concentrations and vNH₄ were both positively related to local krill biomass values, computed from mean values along acoustic transect segments within 2 km of the uptake measurement stations. These biomass values ranged from ∼1 g krill fresh mass m⁻² in the northwest to >4 kg krill wet mass m⁻² in the northeast. In contrast to the variability found with NH₄ concentrations and uptake rates, vNO₃ was more uniform across the sampling sites. Under these conditions, increasing NH₄ concentration appeared to represent an additional N resource. However, high vNH₄ tended to be found for stations with lower phytoplankton standing stocks, across a total range of 0.24-20 mg chlorophyll a m⁻³. These patterns suggest a coupling between phytoplankton biomass, vNH₄ and krill in this region of variable but high krill biomass. Locally high concentrations of krill in parts of the study area appeared to have two opposing effects. On the one hand they could graze down phytoplankton stocks, but on the other hand, their NH₄ excretion supported enhanced uptake rates by the remaining, ungrazed cells.