Nitrogen dynamics during the Arabian Sea Northeast Monsoon

This investigation focused on the weaker and less well understood of the two Arabian Sea monsoonal wind phases, the NE Monsoon, which persists for 3–4 months in the October to February period. Historically, this period has been characterized as a time of very low nutrient availability and low biological production. As part of the US JGOFS Arabian Sea Process Study, 17 stations were sampled on a cruise in January 1995 (late NE Monsoon) and, 15 stations were sampled on a cruise in November 1995 (early NE Monsoon). Only the southern most stations (10° and 12°N) and one shallow coastal station were as nutrient-depleted as had been expected from the few relevant prior studies in this region. Experiments were conducted to ascertain the relative importance of different nitrogenous nutrients and the sufficiency of local regeneration processes in supplying nitrogenous nutrients utilized in primary production. Except for the southern oligotrophic stations, the euphotic zone concentrations of NO₃⁻ were typically 5–10-fold greater than those of NO₂⁻ and NH₄⁺. There was considerable variation (20–40-fold) in nutrient concentration both within and between the two sections on each cruise. All nitrogenous nutrients were more abundant (2–4-fold) later in the NE Monsoon. Strong vertical gradients in euphotic zone NH₄⁺ concentration, with higher concentrations at depth, were common. This was in contrast to the nearly uniform euphotic zone concentrations for both NO₃⁻ and NO₂⁻. Half-saturation constants for uptake were higher for NO₃⁻ (1.7 μmol kg⁻¹ (s.d.=0.88, n=8)) than for NH₄⁺ (0.47 μmol kg⁻¹ (s.d.=0.33, n=5)). Evidence for the suppressing effect of NH₄⁺ on NO₃⁻ uptake was widespread, although not as severe as has been noted for some other regions. Both the degree of sensitivity of NO₃⁻ uptake to NH₄⁺ concentration and the half-saturation constant for NO₃⁻ uptake were correlated with ambient NO₃⁻ concentration. The combined effect of high affinity for low concentrations of NH₄⁺ and the effect of NH₄⁺ concentration on NO₃⁻ uptake resulted in similarly low f-ratios, 0.15 (s.d.=0.07, n=15) and 0.13 (s.d.=0.08, n=17), for early and late observations in the NE Monsoon, respectively. Stations with high f-ratios had the lowest euphotic zone NH₄⁺ concentrations, and these stations were either very near shore or far from shore in the most oligotrophic waters. At several stations, particularly early in the NE Monsoon, the utilization rates for NO₂⁻ were equal to or greater than 50% the utilization rates for NO₃⁻. When converted with a Redfield C : N value of 6.7, the total N uptake rates measured in this study were commensurate with measurements of C productivity. While nutrient concentrations at some stations approached levels low enough to limit phytoplankton growth, light was shown to be very important in regulating N uptake at all stations in this study. Diel periodicity was observed for uptake of all nitrogenous nutrients at all stations. The amplitude of this periodicity was positively correlated with nutrient concentration. The strongest of these relationships occurred with NO₃⁻. Ammonium concentration strongly influenced the vertical profiles for NO₃⁻ uptake as well as for NH₄⁺ uptake. Both NO₂⁻ and NH₄⁺ were regenerated within the euphotic zone at rates comparable to rates of uptake of these nutrients, and thus maintenance of mixed layer concentrations did not require diffusive or advective fluxes from other sources. Observed turnover times for NH₄⁺ were typically less than one day. Rapid turnover and the strong light regulation of NH₄⁺ uptake allowed the development and maintenance of vertical structure in NH₄⁺ concentration within the euphotic zone. In spite of the strong positive effect of light on NO₂⁻ uptake and its strong negative effect on NO₂⁻ production, the combined effects of much longer turnover times for this nutrient and mixed layer dynamics resulted in nearly uniform NO₂⁻ concentrations within the euphotic zone. Responses of the NE Monsoon planktonic community to light and nutrients, in conjunction with mixed layer dynamics, allowed for efficient recycling of N within the mixed layer. As the NE Monsoon evolved and the mixed layer deepened convectively, NO₂⁻ and NO₃⁻ concentrations increased correspondingly with the entrainment of deeper water. Planktonic N productivity increased 2-fold, but without a significant change the new vs. recycled N proportionality. Consequently, NO₃⁻ turnover time increased from about 1 month to greater than 3 months. This reflected the overriding importance of recycling processes in supplying nitrogenous nutrients for primary production throughout the duration of the NE Monsoon. As a result, NO₃⁻ supplied to the euphotic zone during the NE Monsoon is, for the most part, conserved for utilization during the subsequent intermonsoon period.     

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.     

Modelling phytoplankton deposition to Chesapeake Bay sediments during winter–spring: interannual variability in relation to river flow

The often-rapid deposition of phytoplankton to sediments at the end of the spring phytoplankton bloom is an important component of benthic–pelagic coupling in temperate and high latitude estuaries and other aquatic systems. However, quantifying the flux is difficult, particularly in spatially heterogeneous environments. Surficial sediment chlorophyll-a, which can be measured quickly at many locations, has been used effectively by previous studies as an indicator of phytoplankton deposition to estuarine sediments. In this study, surficial sediment chlorophyll-a was quantified in late spring at 20–50 locations throughout Chesapeake Bay for 8 years (1993–2000). A model was developed to estimate chlorophyll-a deposition to sediments using these measurements, while accounting for chlorophyll-a degradation during the time between deposition and sampling. Carbon flux was derived from these estimates via C:chl-a = 75.Bay-wide, the accumulation of chlorophyll-a on sediments by late spring averaged 171 mg m⁻², from which the chlorophyll-a and carbon sinking fluxes, respectively, were estimated to be 353 mg m⁻² and 26.5 gC m⁻². These deposition estimates were ∼50% of estimates based on a sediment trap study in the mid-Bay. During 1993–2000, the highest average chlorophyll-a flux was in the mid-Bay (248 mg m⁻²), while the lowest was in the lower Bay (191 mg m⁻²). Winter–spring average river flow was positively correlated with phytoplankton biomass in the lower Bay water column, while phytoplankton biomass in that same region of the Bay was correlated with increased chlorophyll-a deposition to sediments. Responses in other regions of the Bay were less clear and suggested that the concept that nutrient enrichment in high flow years leads to greater phytoplankton deposition to sediments may be an oversimplification. A comparison of the carbon flux associated with the deposition of the spring bloom with annual benthic carbon budgets indicated that the spring bloom did not contribute a disproportionately large fraction of annual carbon inputs to Chesapeake Bay sediments. Regional patterns in chlorophyll-a deposition did not correspond with the strong regional patterns that have been found for plankton net community metabolism during spring.     

Benthic metabolism and the fate of dissolved inorganic nitrogen in intertidal sediments

We determined patterns of benthic metabolism and examined the relative importance of denitrification (DNF) and dissimilatory nitrate reduction to ammonium (DNRA) as sinks for nitrate (NO₃⁻) in intertidal sediments in the presence and absence of benthic microalgal (BMA) activity. By influencing the activity of BMA, light regulated the metabolic status of the sediments, and, in turn, exerted strong control on sediment nitrogen dynamics and the fate of inorganic nitrogen. A pulsed addition of ¹⁵N-labeled NO₃⁻ tracked the effect and fate of dissolved inorganic nitrogen (DIN) in the system. Under illuminated conditions, BMA communities influenced benthic fluxes directly, via DIN uptake, and indirectly, by altering the oxygen penetration depth. Under dark hypoxic and anoxic conditions, the fate of water column NO₃⁻ was determined largely by three competing dissimilatory reductive processes; DNF, DNRA, and, on one occasion, anaerobic ammonium oxidation (anammox). Mass balance of the added ¹⁵N tracer illustrated that DNF accounted for a maximum of 48.2% of the ¹⁵NO₃⁻ reduced while DNRA (a minimum of 11.4%) and anammox (a minimum of 2.2%) accounted for much less. A slurry experiment was employed to further examine the partitioning between DNF and DNRA. High sulfide concentrations negatively impacted rates of both processes, while high DOC:NO₃⁻ ratios favored DNRA over DNF.     

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.     

Carbon cycling in a high-arctic marine ecosystem – Young Sound, NE Greenland

Young Sound is a deep-sill fjord in NE Greenland (74°N). Sea ice usually begins to form in late September and gains a thickness of 1.5 m topped with 0–40 cm of snow before breaking up in mid-July the following year. Primary production starts in spring when sea ice algae begin to flourish at the ice–water interface. Most biomass accumulation occurs in the lower parts of the sea ice, but sea ice algae are observed throughout the sea ice matrix. However, sea ice algal primary production in the fjord is low and often contributes only a few percent of the annual phytoplankton production. Following the break-up of ice, the immediate increase in light penetration to the water column causes a steep increase in pelagic primary production. Usually, the bloom lasts until August–September when nutrients begin to limit production in surface waters and sea ice starts to form. The grazer community, dominated by copepods, soon takes advantage of the increased phytoplankton production, and on an annual basis their carbon demand (7–11 g C m⁻²) is similar to phytoplankton production (6–10 g C m⁻²). Furthermore, the carbon demand of pelagic bacteria amounts to 7–12 g C m⁻² yr⁻¹. Thus, the carbon demand of the heterotrophic plankton is approximately twice the estimated pelagic primary production, illustrating the importance of advected carbon from the Greenland Sea and from land in fuelling the ecosystem.In the shallow parts of the fjord (<40 m) benthic primary producers dominate primary production. As a minimum estimate, a total of 41 g C m⁻² yr⁻¹ is fixed by primary production, of which phytoplankton contributes 15%, sea ice algae <1%, benthic macrophytes 62% and benthic microphytes 22%. A high and diverse benthic infauna dominated by polychaetes and bivalves exists in these shallow-water sediments (<40 m), which are colonized by benthic primary producers and in direct contact with the pelagic phytoplankton bloom. The annual benthic mineralization is 32 g C m⁻² yr⁻¹ of which megafauna accounts for 17%. In deeper waters benthic mineralization is 40% lower than in shallow waters and megafauna, primarily brittle stars, accounts for 27% of the benthic mineralization. The carbon that escapes degradation is permanently accumulated in the sediment, and for the locality investigated a rate of 7 g C m⁻² yr⁻¹ was determined.A group of walruses (up to 50 adult males) feed in the area in shallow waters (<40 m) during the short, productive, ice-free period, and they have been shown to be able to consume <3% of the standing stock of bivalves (Hiatella arctica, Mya truncata and Serripes Groenlandicus), or half of the annual bivalve somatic production. Feeding at greater depths is negligible in comparison with their feeding in the bivalve-rich shallow waters.     

Nitrogen isotopic discrimination by water column nitrification in a shallow coastal environment

Temporal changes in nitrogen isotopic composition (δ¹⁵N) of the NO₃ ⁻ pool in the water column below the pycnocline in Ise Bay, Japan were investigated to evaluate the effect of nitrification on the change in the δ¹⁵N in the water column. The δ¹⁵N of NO₃ ⁻ in the lower layers varied from −8.5‰ in May to +8.4‰ in July in response to the development of seasonal hypoxia and conversion from NH₄ ⁺ to NO₃ ⁻. The significantly ¹⁵N-depleted NO₃ ⁻ in May most likely arose from nitrification in the water column. The calculated apparent isotopic discrimination for water column nitrification (ɛ_nit = δ¹⁵N_substrate − δ¹⁵N_product) was 24.5‰, which lies within the range of previous laboratory-based estimates. Though prominent deficits of NO₃ ⁻ from hypoxic bottom waters due to denitrification were revealed in July, the isotopic discrimination of denitrification in the sediments was low (ɛ_denit = ∼1‰). δ¹⁵N_NO3 in the hypoxic lower layer mainly reflects the isotopic effect of water column nitrification, given that water column nitrification is not directly linked with sedimentary denitrification and the effect of sedimentary denitrification on the change in δ¹⁵N_NO3 is relatively small.     

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.     

A comparative study of microalgal production in a marine versus a river-dominated temporarily open/closed estuary, South Africa

Temporarily open/closed estuaries (TOCEs) account for almost 71% of the total number of South African estuarine systems. To date the dynamics of microalgal production and biomass in TOCEs is poorly understood. A survey was therefore conducted in two TOCEs along the KwaZulu-Natal coast: the river-dominated Mdloti and the marine-dominated Mpenjati. The aims of this investigation were: (1) to compare the variability of benthic and pelagic microalgal production and biomass in these two contrasting systems; (2) to determine the key environmental parameters influencing primary production in these TOCEs. Results show a similar pattern of primary production and biomass in both estuaries. High benthic microalgal biomass with low primary production and low phytoplankton biomass with high rates of pelagic primary production were observed in both estuaries. Possible explanations for this include: (a) optimum light conditions in the water column, compared to the sediment surface; (b) high impact of grazing by zooplankton in the water column; and (c) settling of phytoplankton. Significant correlations were observed between primary production and environmental parameters (K_d, PAR, temperature, DIN and DIP), during the open and closed phases of both estuaries. A BIOENV (PRIMER) analysis showed that patterns of primary production in both estuaries were influenced primarily by interactions of multiple independent parameters. Comparisons between measured hourly pelagic primary production rates at the Mpenjati and Mdloti and the composite parameter (P_c) of Cole and Cloern (1987) revealed highly significant relationships (r² = 0.8; p < 0.001) between these differently derived variables.     

Blooms of <Emphasis Type="Italic">Trichodesmium erythraeum</Emphasis> in the South Eastern Arabian Sea during the onset of 2009 summer monsoon

This study presents in situ evidence for the blooms of Trichodesmium erythraeum observed in the shelf waters of the South Eastern Arabian Sea (SEAS) during the onset of the southwest monsoon in June 2009. Evidence showed that water surface discoloration was caused by the accumulation of T. erythraeum, and that the water column contained a colony of T. thiebautii. The surface water color in the bloom region varied from pale brown to pinkish red. Pale brown indicated healthy algae at the peak of its photosynthetic activity, while pinkish red indicated the presence of photosynthetically less active filaments. Zooplankton abundance, especially copepodites, in the bloom area substantiated the theory that Trichodesmium filaments are excellent epiphytes to which the copepodites cling. The bloom area was very fertile with copious quantities of dissolved oxygen (6.85 ml L^?1), PO₄-P (0.108 μmol L^?1) and SiO₄ (1.29 μmol L^?1). Lower NO₃-N (0.028 μmol L^?1) values in the bloom area did not appear to affect Trichodesmium growth from molecular nitrogen fixation. However, lower NO₃-N values altered the normal phytoplankton composition of this area.     

Changes in sediment processes across the western Irish Sea front

Sediment characteristics, sediment respiration (oxygen uptake and sulphate reduction) and sediment–water nutrient exchange, in conjunction with water column structure and phytoplankton biomass were measured at five stations across the western Irish Sea front in August 2000. The transition from thermally stratified (surface to bottom temperature difference of 2.3 °C) to isothermal water (14.3 °C) occurred over a distance of 13 km. The influence of the front on phytoplankton biomass was limited to a small region of elevated near surface chlorophyll (2.23 mg m⁻³; 50% > biomass in mixed waters). The front clearly marked the boundary between depositional sediments (silt/clays) with elevated sediment pigment levels (≈60 mg m⁻²) on the western side, to pigment impoverished (<5 mg m⁻²) sand, through to coarse sand and shell fragments on the eastern side. Maximal rates of sedimentary respiration on the western stratified side of the front e.g. oxygen uptake S2 (852 μmol O₂ m⁻² h⁻¹) and sulphate reduction at S1 (149 μmol SO₄²⁻ m⁻² h⁻¹), coupled to significant efflux of nitrate and silicate at the western stations indicate closer benthic–pelagic coupling in the western Irish Sea. Whether this simply reflects the input of phytodetritus from the overlying water column or entrapment and settlement of pelagic production from other regions of the Irish Sea cannot yet be resolved.     

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.     

Annual Variation of Benthic Nutrient Fluxes in Shallow Coastal Waters (Gulf of Trieste, Northern Adriatic Sea)

Abstract. Benthic fluxes of dissolved N. Si and P nutrients, alkalinity, dissolved inorganic C (DIC), and O₂ from sediments in the Gulf of Trieste (northern Adriatic, Italy) were measured monthly for 16 months, using laboratory incubated flux chambers at in siru temperatures in the dark. The annual average fluxes were: 02 = -19.3 ± 8.2, DIC = 13.7 ± 9.6, NO3 = -0.04 ± 0.16, NH4 = 0.3 ± 0.4. PO₄= 4.001 ± 0.01, Si = 0.9 ± 0.1 mmol m-² d^-1, with strong temporal fluctuations. The highest effluxes of all nutrients and DIC were observed in the summer. Small effluxes of DIC and NH4 and influxes of Si and PO4 were observed in late winter. Only NH₄ (ca. 50%) and Si (ca. 70%) fluxes were significantly correlated with temperature. This correlation suggests that the rate of downward input and the quality of sedimented organic matter (autochthonous and allochthonous) were superimposed on the temperature fluctuations. High DIC, NH₄ and Si effluxes observed in May 1993 during low temperature were due to the degradation of sedimentary organic matter produced by an early spring bloom of benthic microalgae which occurred about 6 weeks earlies while the autumn phytoplankton bloom was simultaneously reflected in enhanced benthic fluxes due to higher temperature. The role of benthic biological advection in this transport across the sediment-water interface, evaluated by comparison between measured benthic and calculated diffusive fluxes from nutrient pore water concentrations, was of minor importance. This is probably due to low infaunal activity throughout the year it was localized mostly in the narrow surficial layer. The annual average diffusive fluxes of NH4 and PO4 were higher than those measured, probably due to the presence of nitrificationdenitrifi-cation processes and redox-dependent chemical reactions at the oxic sediment-water interface, respectively. Only during bottom-water hypoxia in September 1993 did strong PO₄ effluxes prevail. Calculations based on the Redfield stoichiometry of oxic decomposition of organic N to NH4 and NO3, and differences between diffusive and measured NH₄ fluxes showed that denitrifkation averaged 0.8 mmol m^-2 d^-1. Significant correlations between NH₄ and PO₄ DIC and Si, and NH4 and Si fluxes suggested their parallel regeneration and utilization at the sediment-water interface. The nutrient fluxes observed were not significantly linked to O₂ consumption, suggesting also that anaerobic oxidation processes were important at the sediment-water interface in the gulf. The N, P and Si nutriqnts released from sediment pore waters are probably utilized in benthic microalgal and bottorn-hater primary production. This indicates that pelagic and benthic communities in the central part of the Gulf of Trieste function relatively independently of each other.     

A role for nitrite in the production of nitrous oxide in the lower euphotic zone of the oligotrophic North Pacific Ocean

Understanding the role of the oceans in the Earth's changing climate requires comprehension of the relevant metabolic pathways which produce climatically important trace gases. The global ocean represents one of the largest natural sources of nitrous oxide (N₂O) that is produced by selected archaea and/or bacteria during nitrogen (N) metabolism. In this study, the role of nitrite (NO₂⁻) in the production of N₂O in the upper water column of the oligotrophic North Pacific Subtropical Gyre was investigated, focusing primarily on the lower euphotic zone where NO₂⁻ concentrations at the primary NO₂⁻ maximum reached 195 nmol L⁻¹. Free-drifting sediment trap arrays were deployed to measure N cycle processes in sinking particulate material and the addition of selected N substrates to unpreserved sediment traps provided an experimental framework to test hypotheses regarding N₂O production pathways and controls. Sinking particles collected using NO₂⁻-amended, unpreserved sediment traps exhibited significant production of N₂O at depths between 100 and 200 m. Subsequent stable isotope tracer measurements conducted on sediment trap material amended with ¹⁵NO₂⁻ yielded elevated δ¹⁵N values of N₂O, supporting N₂O production via a NO₂⁻ metabolism pathway. Experiments on seawater collected from 150 m showed N₂O production via NO₂⁻ metabolism also occurs in the water-column and indicated that the concentration of NO₂⁻ relative to NH₄⁺ availability may be an important control. These findings provide evidence for the production of N₂O via nitrifer-denitrification in the lower euphotic zone of the open ocean, whereby NO₂⁻ is reduced to N₂O by ammonia-oxidizing microorganisms.     

Carbon to nitrogen (C:N) stoichiometry of the spring–summer phytoplankton bloom in the North Water Polynya (NOW)

The carbon to nitrogen (C:N) stoichiometry of phytoplankton production varied significantly during the spring–summer bloom in the North Water Polynya (NOW), from April through July 1998. The molar ratio of particulate organic carbon (POC) to nitrogen (PON) production by phytoplankton (ΔPOC:ΔPON) increased from 5.8 during April through early June to 8.9 in late June and July. The molar dissolved inorganic carbon (DIC) to nitrate+nitrite (NO₃) drawdown ratio (ΔDIC: ΔNO₃) increased from 6.7 in April and May, to 11.9 in June (no estimate for July because of ice melting). The discrepancy between ΔPOC:ΔPON and ΔDIC:ΔNO₃ was likely due to dissolved organic carbon (DOC) production. Increased ΔPOC:ΔPON of phytoplankton and surface water ΔDIC:ΔNO₃ throughout the phytoplankton blooms resulted from changes in physical properties of the upper water column, such as reduced thickness of the surface mixed layer that exposed phytoplankton to increased photosynthetically available radiation (PAR), accompanied by NO₃ depletion. This is expected to have significant effects on the cycling of carbon (C) and nitrogen (N) in pelagic ecosystems, as the increased C:N ratio of organic matter decreases its quality as substrate for grazers and microbial communities. Based on ΔPOC:ΔPON, the ratio of POC to chlorophyll a (Chl) production (ΔPOC:ΔChl) and the relationship between Chl yields and NO₃ depletion, we estimate that 71±17% and 46±20% of the depleted NO₃ went to PON production in the euphotic zone over the polynya from April to early June, and late June to July, respectively. The remaining NO₃ was likely channelled to dissolved organic nitrogen (DON) and heterotrophic bacteria, which were not returned to the dissolved inorganic nitrogen (DIN) pool through recycling during the course of the study. Hence, the autotrophic production of organic N and its recycling by the microbial food web were not coupled temporally.     

Rates of C,N, P and Si recycling and denitrification at the US Mid-Atlantic continental slope depocenter

Benthic fluxes of O₂, titration alkalinity (TA), total inorganic carbon (TIC), Ca²⁺, NO₃⁻, NH₄⁺, PO₄³⁻, and Si(OH)₄ were measured by in situ benthic flux chamber incubations at 13 locations on the North Carolina continental slope. The majority of measurements were made at water depths of approximately 700–850 m, in the previously identified upper slope depocenter. This region is characterized by extremely high organic matter deposition rates and near saturation bottom water oxygen concentrations. Measured benthic fluxes of TA are reasonably correlated with O₂ benthic fluxes. Because bottom waters are supersaturated with respect to calcite and aragonite at these shallow water depths, these results demonstrate the importance of metabolically driven dissolution in this region. Subtraction of the calcium carbonate dissolution contributions from the TIC benthic fluxes suggests rates of organic matter remineralization ranging from 0.97 to 3.9 mol C m⁻² yr⁻¹ at the depocenter sites, a factor of 3–10 greater than estimated for the adjacent continental rise and upper slope areas. Because biological primary production in the overlying waters does not follow this pattern, these extremely high values are most likely supported by lateral inputs of highly reactive organic matter. Mass balance calculations indicate that despite the oxygenated bottom water conditions, 68% of the organic nitrogen released during organic matter remineralization processes is ultimately denitrified. The release of PO₄³⁻ from the depocenter sediments is equivalent to or larger than that predicted from the remineralization of Redfield organic matter. This implies either that PO₄³⁻ is preferentially released in this setting and that the accumulating sediments must be depleted in PO₄³⁻ relative to organic carbon or that another, non-organic, phase is contributing PO₄³⁻ to the system. The molar ratio of the Si benthic flux and organic carbon remineralization rate ranges from 0.30 to 0.86. This is significantly greater than the ratio reported for most pelagic diatoms. Possible reasons for this high ratio include the deposition of benthic diatoms that may have a larger Si : C ratio than pelagic diatoms, the near-bottom lateral input of partially reworked organic matter that may have an elevated Si : C ratio relative to fresh diatoms, preferential loss of carbon in sinking particulates or the release of Si from non-opaline materials.     

Nitrogen transformations as inferred from the activities of key enzymes in the Arabian Sea oxygen minimum zone

Vertical distributions of the potential activities of some key enzymes mediating nitrification and denitrification were investigated within the oxygen (O₂) minimum zone of the Arabian Sea at a number of locations between latitudes 17°N and 21°N and longitudes 63°E and 68°E so as to get an insight into the predominant biochemical mode(s) of production and consumption of nitrous oxide (N₂O). Results revealed that the dissimilatory nitrate (NO⁻₃) reduction activity was generally very low or absent within the σ_θ range 26.6–26.8, which corresponds to the Persian Gulf Watermass (PGW). Depth profiles of nitrate reductase (NaR), nitrite reductase (NiR) and ammonia monooxygenase (AMO) activities were compared with those of O₂, NO⁻₃, nitrite (NO⁻₂) and N₂O, and it is concluded that nitrifier denitrification rather than heterotrophic denitrification is active within the core of PGW. The presence of multiple peaks of AMO activity coinciding with distinct maxima in the O₂ profile and with a trend opposite to that of NaR activity indicates that the two processes, viz., classical and nitrifier denitrification, occur in discrete layers, probably determined by the variations in the ambient O₂ concentrations at various depths surrounding the PGW core. Further, it appears that at the depths where nitrifier denitrification is active in the absence of heterotrophic denitrification, N₂O builds up as its consumption may be inhibited by O₂. Possible reasons for the occurrence of appreciable nitrate deficit within the core of PGW, where dissimilatory NO⁻₃ reduction is lacking, are discussed.     

Phytoplankton spring bloom of the Gironde plume waters in the Bay of Biscay: early phosphorus limitation and food-web consequences

During the spring 1995 (2–25 May), a cruise was carried on the RV Poseidon (Germany) on the continental shelf of the south Bay of Biscay. The objective was a comprehensive study of the planktonic food web within the Gironde plume waters. In these waters phosphate was present at very low concentrations (undetectable to < 0.1 μmol.L⁻¹), whereas nitrate, silicate and ammonium concentrations were much higher (several μmol·L⁻¹ for nitrate and silicate and 0.5 to 1.0 μmol·L⁻¹ for ammonium). The size distribution of the phytoplankton biomass (estimated from chlorophyll a measurements by high performance liquid chromatography) and primary production (measured by ¹⁴C in situ method) showed a great proportion of small (40 to 70 % < 3 μm) and active autotrophic cells (growth rates estimated from 0.4 to 0.8 d⁻¹ for the entire euphotic layer). Considering the very high values of NO₃-N:PO₄-P ratios and the high C:P and N:P ratios for the particulate organic matter, it is suggested that an early phosphorus depletion limits the spring bloom phytoplankton and particularly the new production (nitrate uptake coming from the Gironde waters).From these results and other simultaneous observations on the heterotrophic processes (such as grazing of microzooplankton), we can conclude that the planktonic food web would be close to a maintenance system as defined by Platt et al. The possible generalisation of these results for each spring is discussed with respect to the scarcity of previous and reliable phosphate data.     

Coupling iodine speciation to primary,regenerated or “new” production: a re-evaluation

In two recent studies (Tian et al., Deep-Sea Res. I 43 (1996) 723–738; Campos et al., Deep-Sea Res. II 43 (1996) 455–466), in order to explain the observed temporal variations in the distributions of the dissolved iodine species at the time-series stations in the Mediterranean Sea, the North Atlantic and the North Pacific, diverse assumptions were invoked on the relationships between changes in the speciation of dissolved iodine in the surface oceans and biological production such that the surface enrichment of I⁻ was linked to both regenerated production and primary production while the surface depletion of [IO₃⁻+I⁻] was linked to “new” production. However, while some of the major conclusions in these studies are critically dependent on these assumptions, the validity of the assumptions has yet to be verified with experimental evidence. On the other hand, while there are still significant unknowns in the understanding of the cycling among dissolved iodine species in the surface oceans, presently available data from laboratory cultures and field observations are consistent with an alternative conceptual model in which IO₃⁻ and NO₃⁻ are taken up at some ratio to each other during NO₃⁻ uptake and almost all of the IO₃⁻ taken up is exuded as I⁻. Thus, the depletion of IO₃⁻ and the enrichment of I⁻ in the surface water are linked to NO₃⁻ uptake. This alternative model is also consistent with the data sets presented by Tian et al. (1996) and Campos et al. (1996). By linking the surface depletion of IO₃⁻ to NO₃⁻ uptake, significantly different biogeochemical behavior of the marine dissolved iodine system may be inferred. The extent to which I⁻ may be oxidized to IO₃⁻ within the euphotic zone during the residence time of the water in the surface ocean is still an open question.     

Nitrous oxide and N-nutrient cycling in the oxygen minimum zone off northern Chile

Measurements of dissolved gases (O₂, N₂O), nutrients (NO₃⁻, NO₂⁻, PO₄³⁻), and oceanographic variables were performed off northern Chile (∼21°S) between March 2000 and July 2004, in order to characterize the existing oxygen minimum zone (OMZ) and identify processes involved in N₂O cycling. Both N₂O and NO₃⁻ displayed sharp, shallow peaks with concentrations of up to 124 nM (1370% saturation) and 26 μM, respectively, in association with a strong oxycline that impinges on the euphotic zone. NO₂⁻ accumulation below the oxycline's base reached up to 9 μM. The vertical distribution of physical and chemical parameters and the existing relationships between apparent oxygen utilization (AOU), apparent N₂O production (ΔN₂O), and NO₃⁻ revealed three main layers within the upper OMZ. The first layer, or the upper part of the oxycline, is located between the base of the mixed layer and the mid-point of the oxycline (around σ_t=25.5 kg m⁻³). There the O₂ declines from ∼250 to ∼50 μM, and strong (but opposing) O₂ and NO₃⁻ gradients and their associated AOU–ΔN₂O and AOU–NO₃⁻ relationships indicate that nitrification produces N₂O and NO₃⁻ in the presence of light. The second layer, or lower part of the oxycline, represents the upper OMZ boundary and is located between the middle and the base of the oxycline (25.9<σ_t<26.1 kg m⁻³). In this layer NO₃⁻ reduction begins at O₂ levels ranging from ∼50 to ∼11 μM and accumulation of 41–68% of the ΔN₂O pool occurs. The accumulation of N₂O (but not of NO₂⁻ or NH₄⁺) and the observed AOU–ΔN₂O and AOU–NO₃⁻ relationships (which are opposite to those of the overlying first layer) suggest that a coupling between nitrification and NO₃⁻ reduction is involved in N₂O cycling in this second layer. The third layer is the OMZ core, where the O₂ concentration remains constant (O₂<11 μM). It coincides with σ_t>26.2 kg m⁻³, which is typical of Equatorial Subsurface Water (ESSW). In this layer, N₂O and NO₃⁻ continue to decrease, but a large NO₂⁻ accumulation is observed. Considering all the data, a biogeochemical model for the upper OMZ off northern of Chile is proposed, in which nitrification and denitrification differentially mediate N₂O cycling in each layer.