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.     

Aerobic ammonium oxidation in the oxycline and oxygen minimum zone of the eastern tropical South Pacific off northern Chile (∼20°S)

Aerobic NH₄⁺ oxidation rates were measured along the strong oxygen gradient associated with the oxygen minimum zone (OMZ) of the eastern tropical South Pacific off northern Chile (∼20°S) during 2000, 2003, and 2004. This process was examined by comparing NH₄⁺ rates of change during dark incubations, with and without the addition of allylthiourea, a classical inhibitor of the ammonia monooxygenase enzyme of ammonium-oxidizing bacteria. The contribution of aerobic NH₄⁺ oxidation in dark carbon fixation and NO₂⁻ rates of change were also explored. Thirteen samples were retrieved from the oxycline (252 to ⩽5 μM O₂; 15 to ∼65 m depth) and three from the oxygen minimum core (⩽5 μM O₂; 100–200 m depth). Aerobic NH₄⁺ oxidation rates were mainly detected in the upper part (15–30 m depth) of the oxycline, with rates ranging from 0.16 to 0.79 μM d⁻¹, but not towards the oxycline base (40–65 m depth). In the oxygen minimum core, aerobic NH₄⁺ oxidation was in the upper range and higher than in the upper part of the oxycline (0.70 and 1.0 μM d⁻¹). Carbon fixation rates through aerobic NH₄⁺ oxidation ranged from 0.18 to 0.43 μg C L⁻¹ d⁻¹ and contributed between 33% and 57% of the total dark carbon fixation, mainly towards the oxycline base and, in a single experiment, in the upper part of the oxycline. NO₂⁻ consumption was high (up to 10 μM d⁻¹) towards the oxycline base and OMZ core, but was significantly reduced in experiments amended with allylthiourea, indicating that aerobic NH₄⁺ oxidation could contribute between 8% and 76% of NO₂⁻ production, which in turn could be available for denitrifiers. Overall, these results support the important role of aerobic NH₄⁺ oxidizers in the nitrogen and carbon cycling in the OMZ and at its upper boundary.     

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.     

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.     

Phytoplankton growth and mortality during the 1995 Northeast Monsoon and Spring Intermonsoon in the Arabian Sea

Phytoplankton growth rates and mortality rates were experimentally examined at eight stations in the Arabian Sea along the U.S. JGOFS cruise track during the 1995 Northeast Monsoon (January) and Spring Intermonsoon (March–April). Instantaneous growth rates averaged over an entire cruise were approximately twice as high during the NE Monsoon than during the Spring Intermonsoon period (overall averages of 0.84±0.29 (s.d.) versus 0.44±0.19 d⁻¹). Average herbivore grazing (mortality) rates, however, were quite similar for the two seasons (overall averages of 0.35±0.18 and 0.30±0.17 d⁻¹ for the NE Monsoon and Spring Intermonsoon, respectively). The absolute amounts of phytoplankton biomass consumed during each season also were similar (29 and 25% of standing stock consumed d⁻¹ for the January and March–April cruises, respectively), as were the geographical trends of this removal. These seasonal trends in growth and removal rates resulted in net phytoplankton growth rates that were considerably higher during the January cruise (0.48 d⁻¹) than during the March–April cruise (0.14 d⁻¹). That is, phytoplankton production was more closely balanced during the Spring Intermonsoon season (87% of daily primary production consumed) relative to the NE Monsoon season (49% of daily primary production consumed). Station-to-station variability was high for rate measurements during either cruise. Nevertheless, there was a clear onshore–offshore trend in the absolute rate of removal of phytoplankton biomass (μg chlorophyll consumed l⁻¹ d⁻¹) during both cruises. Coastal stations had removal rates that were typically 2–4 times higher than removal rates at oceanic stations.     

Dry atmospheric deposition and diazotrophy as sources of new nitrogen to northwestern Mediterranean oligotrophic surface waters

Atmospheric dry deposition of nitrogen (N) and dinitrogen (N₂) fixation rates were assessed in 2004 at the time-series DYFAMED station (northwestern Mediterranean, 43°25′N, 7°52′E). The atmospheric input was monitored over the whole year. Dinitrogen fixation was measured during different seasonal trophic states (from mesotrophy to oligotrophy) sampled during nine cruises. The bioavailability of atmospherically deposited nutrients was estimated by apparent solubility after 96 h. The solubility of dry atmospheric N deposition was highly variable (from ∼18% to more than 96% of total N). New N supplied to surface waters by the dry atmospheric deposition was mainly nitrate (NO₃⁻) (∼57% of total N, compared to ∼6% released as ammonium (NH₄⁺)). The mean bioavailable dry flux of total N was estimated to be ∼112 μmol m⁻² d⁻¹ over the whole year. The NO₃⁻ contribution (70 μmol NO₃⁻ m⁻² d⁻¹) was much higher than the NH₄⁺ contribution (1.2 μmol NH₄⁺ m⁻² d⁻¹). The N:P ratios in the bioavailable fraction of atmospheric inputs (122.5–1340) were always much higher than the Redfield N:P ratio (16). Insoluble N in atmospheric dry deposition (referred to as “organic” and believed to be strongly related to anthropogenic emissions) was ∼40 μmol m⁻² d⁻¹. N₂ fixation rates ranged from 2 to 7.5 nmol L⁻¹ d⁻¹. The highest values were found in August, during the oligotrophic period (7.5 nmol L⁻¹ at 10 m depth), and in April, during the productive period (4 nmol L⁻¹ d⁻¹ at 10 m depth). Daily integrated values of N₂ fixation ranged from 22 to 100 μmol N m⁻² d⁻¹, with a maximum of 245 μmol N m⁻² d⁻¹ in August. No relationship was found between the availability of phosphorus or iron and the observed temporal variability of N₂ fixation rates. The atmospheric dry deposition and N₂ fixation represented 0.5–6% and 1–20% of the total biological nitrogen demand, respectively. Their contribution to new production was more significant: 1–28% and 2–55% for atmospheric dry deposition and N₂ fixation, respectively. The dry atmospheric input was particularly significant in conditions of water column stratification (16–28% of new production), while N₂ fixation reached its highest values in June (46% of new production) and in August (55%).     

A method for the determination of nitrification rates in oligotrophic marine seawater by gas chromatography/mass spectrometry

An isotope dilution method has been developed to determine by gas chromatography/mass spectrometry (GC/MS) the rates of ammonium and nitrite oxidation in severely oligotrophic marine waters. The method is based on the formation of sudan-1 from nitrite, or from nitrate following reduction to nitrite. Samples were collected by solid phase extraction and purified by high performance liquid chromatography (HPLC). A deuterated sudan-1 internal standard was synthesized, purified by HPLC and used for quantitative analysis. Concentrations of NO₂⁻ and NO₃⁻ were generally < 2 nmol/kg and < 5 nmol/kg respectively, typical of oligotrophic surface waters, and turnover times for the inorganic N pools ranged from < 1 day to > 10 days. Significant rates of nitrification were measured in the surface oligotrophic ocean, with rates of ammonium and nitrite oxidation generally within the range of 10–500 pmol/kg/h. Consequently, a significant proportion of daily NO₃⁻ assimilation by marine phytoplankton is regenerated, and not new. In a case study of the oligotrophic gyre of the North Atlantic, the influence of NH₄⁺ regeneration and nitrification on f-ratio values suggests that in the oligotrophic ocean, f-ratio values may be significantly, and sometimes grossly, overestimated.     

Picophytoplankton dynamics and production in the Arabian Sea during the 1995 Southwest Monsoon

Phytoplankton community structure is expected to shift to larger cells (e.g., diatoms) with monsoonal forcing in the Arabian Sea, but recent studies suggest that small primary producers remain active and important, even in areas strongly influenced by coastal upwelling. To better understand the role of smaller phytoplankton in such systems, we investigated growth and grazing rates of picophytoplankton populations and their contributions to phytoplankton community biomass and primary productivity during the 1995 Southwest Monsoon (August–September). Environmental conditions at six study stations varied broadly from open-ocean oligotrophic to coastal eutrophic, with mixed-layer nitrate and chlorophyll concentrations ranging from 0.01 to 11.5 μM NO₃ and 0.16 to 1.5 μg Chl a. Picophytoplankton comprised up to 92% of phytoplankton carbon at the oceanic stations, 35% in the diatom-dominated coastal zone, and 26% in a declining Phaeocystis bloom. Concurrent in situ dilution and ¹⁴C-uptake experiments gave comparable ranges of community growth rates (0.53–1.05 d⁻¹ and 0.44–1.17 d⁻¹, to the 1% light level), but uncertainties in C:Chl a confounded agreement at individual stations. Microzooplankton grazing utilized 81% of community phytoplankton growth at the oligotrophic stations and 54% at high-nutrient coastal stations. Prochlorococcus (PRO) was present at two oligotrophic stations, where its maximum growth approached 1.4 d⁻¹ (two doublings per day) and depth-integrated growth varied from 0.2 to 0.8 d⁻¹. Synechococcus (SYN) growth ranged from 0.5 to 1.1 d⁻¹ at offshore stations and 0.6 to 0.7 d⁻¹ at coastal sites. Except for the most oligotrophic stations, growth rates of picoeukaryotic algae (PEUK) exceeded PRO and SYN, reaching 1.3 d⁻¹ offshore and decreasing to 0.8 d⁻¹ at the most coastal station. Microzooplankton grazing impact averaged 90, 70, and 86% of growth for PRO, SYN, and PEUK, respectively. Picoplankton as a group accounted for 64% of estimated gross carbon production for all stations, and 50% at high-nutrient, upwelling stations. Prokaryotes (PRO and SYN) contributed disproportionately to production relative to biomass at the most oligotrophic station, while PEUK were more important at the coastal stations. Even during intense monsoonal forcing in the Arabian Sea, picoeukaryotic algae appear to account for a large portion of primary production in the coastal upwelling regions, supporting an active community of protistan grazers and a high rate of carbon cycling in these areas.     

Seasonal variations in the distribution of Fe and Al in the surface waters of the Arabian Sea

Concentrations of dissolved Al and Fe in the surface mixed layer were measured during five cruises of the 1995 US JGOFS Arabian Sea Process Study, Concentrations of both Al and Fe were relatively uniform between January and April, the NE Monsoon and the Spring Intermonsoon period, ranging from 2 to 11 nM Al (mean 5.3 nM) and 0.5 to 2.4 nM Fe (mean 1.0 nM). In July/August, after the onset of the SW Monsoon, surface water Al and Fe concentrations increased significantly (Al range 4.5–20.1 nM; mean=10 nM, Fe range 0.57–2.4 nM; mean=1.3 nM), particularly in the NE part of the Arabian Sea, as the result of the input and partial dissolution of eolian dust. Using the enrichment of Al in the surface waters, we estimate this is the equivalent to the deposition of 2.2–7.4 g m⁻² dust, which is comparable to values previously estimated for this region. Approximately one month later (August/September), surface water concentrations of both Al and Fe were found to have decreased significantly (mean Al 7.4 nM, mean Fe 0.90 nM) particularly in the same NE region, as the result of export of particulate material from the euphotic zone. Fe supply to the surface waters is also affected by upwelling of sub-surface waters in the coastal region of the Arabian Sea during the SW Monsoon. Despite the proximity of high concentrations of Fe in the shallow sub-oxic layer, freshly upwelled water is not drawn from this layer and the NO₃/Fe ratio in the initially upwelled water is below the value at which Fe limitation is through to occur. Continued deposition of eolian Fe into the upwelled water as it advects offshore provides the Fe required to raise this ratio above the Fe limitation value.     

Monsoon-controlled export fluxes to the interior of the Arabian Sea

As a part of the US-JGOFS Arabian Sea Process Study (ASPS), we deployed a mooring array consisting of 16 Mark-7G time-series sediment traps on five moorings, each in the mesopelagic and interior depths in the western Arabian Sea set along a transect quasi-perpendicular to the Omani coast. The array was deployed for 410 days to cover all monsoon and inter-monsoon phases at 4.25-, 8.5- or 17-day open-close intervals, all of which were synchronized at 17-day periods. Total mass flux, fluxes of organic, inorganic carbon, biogenic Si and lithogenic Al (mg m⁻² day⁻¹) were obtained from samples representing 667 independent periods. The average total mass fluxes estimated in the interior depth along this sediment trap array at Mooring Stations 1–5 (MS-1–5) during 1994-5 ASPS were 147, 235, 221, 164 and 63 mg m⁻² day⁻¹, respectively. Mass fluxes during the southwest (SW) Monsoon were always larger than during the northeast (NE) Monsoon at all divergent zone stations, but the difference was insignificant at the oligotrophic station, MS-5. Four major pulses of export flux events, two each at NE Monsoon and SW Monsoon, were observed in the divergent zone; these events dominated in quantity production of the annual mass flux, but did not dominate temporally. Export pulses were produced by passing eddies and wind-curl events, but the direct processes to produce individual export blooms at each station were diversified and highly complex. The onset of these pulses was generally synchronous throughout the divergent zone. Export pulses associated with specific biogeochemical signatures such as the ratio of elevated biogenic Si to inorganic carbon indicate a supply of deep water to the euphotic layer in varying degrees. The variability of mass fluxes at the oligotrophic station, MS-5, also represented both monsoon events, but with far less amplitude and without notable export pulses.     

Biogeochemical responses to late-winter storms in the Sargasso Sea,I—Pulses of primary and new production

We have hypothesized that the weekly/biweekly passage of winter storms in the subtropical open ocean destabilizes the water column leading to pulsed NO₃^? inputs, resulting in new production that is not accounted for in most annual estimates. This paper presents data on nitrogen and carbon cycling in the Sargasso Sea at approximately daily resolution, during the period prior to seasonal stratification in 2004 and 2005; these data permit us to assess the importance of winter storms for introducing NO₃^? and the contribution of these inputs to annual new and export production. The two sampling years were in stark contrast to each other with 2004 characterized by periods of relative calm between winter storms, and 2005 characterized by nearly continuous storm activity. As a result, temporal variability in mixed layer depth (MLD) and euphotic zone [NO₃^?] were very different between years. MLDs in 2004 increased to >150 m in response to the passage of storms and then rapidly shoaled to <100 m leading to the pulsed injection of NO₃^? (～100 nmol l^?1) into the lower half of the euphotic zone, while in 2005 MLDs were consistently >300 m and euphotic zone [NO₃^?]>100 nmol l^?1. Despite the very different [NO₃^?], rates of daily NO₃^? uptake were similar from year to year because of significant nocturnal uptake in 2004. Similar rates of new production did not translate into similar rates of particulate nitrogen and carbon export however, as observed export from the upper 200 m was 2–5-fold greater in 2004 than in 2005. Furthermore, the decrease of particulate nitrogen and carbon flux with depth between 200 and 400 m in 2004 was substantially lower than in 2005; this is consistent with the observed biological response in which diatoms and coccolithophores exhibited rapid growth following pulsed NO₃^? inputs in 2004. A combination of data from the Bermuda Testbed Mooring, which provides a longer temporal record than the cruise, and the observations presented in this study show that in the winter of 2004, there were 8–10 storm events that likely resulted in pulsed NO₃^? inputs. Summed over all the events, new production prior to seasonal stratification was estimated to be ～0.12–0.18 mol N m^?2 or ～14–21% of current annual estimates.     

Diatom biomass and productivity in oceanic and plume-influenced waters of the western tropical Atlantic ocean

Whereas diatoms (class Bacillariophyceae) often dominate phytoplankton taxa in the Amazon estuary and shelf, their contribution to phytoplankton dynamics and impacts on regional biogeochemistry are poorly understood further offshore in the western tropical Atlantic Ocean (WTAO). Thus, relative contribution of diatoms to phytoplankton biomass and primary production rates and associated environmental conditions were quantified during three month-long cruises in January–February 2001, July–August 2001, and April–May 2003. The upper water column was sampled at 6 light depths (100%, 50%, 25%, 10%, 1% and 0.1% of surface irradiance) at 64 stations between 3° and 14°N latitude and 41° and 58°W longitude. Each station was categorized as ‘oceanic’ or ‘plumewater’, based on principal component analysis of eight physical, chemical and biological variables. All stations were within the North Brazil Current, and plumewater stations were characterized by shallower mixed layers with lower surface salinities and higher dissolved silicon (dSi) concentrations than oceanic stations. The major finding was a much greater role of diatoms in phytoplankton biomass and productivity at plumewater stations relative to oceanic stations. Mean depth-integrated bSi concentrations at the plumewater and oceanic stations were 14.2 and 3.7 mmol m⁻², respectively. Mean depth-integrated SiP rates at the plumewater and oceanic stations were 0.17 and 0.02 mmol m⁻² h⁻¹, respectively. Based on ratios of SiP and PP rates, and typical Si:C ratios, diatoms contributed on average 29% of primary productivity at plumewater stations and only 3% of primary productivity at oceanic stations. In contrast, phytoplankton biomass (as chlorophyll a concentrations) and primary production (PP) rates (as ¹⁴C uptake rates) integrated over the euphotic zone were not significantly different at plumewater and oceanic stations. Chlorophyll a concentrations ranged from 8.5 to 42.4 mg m⁻² and 4.0 to 38.0 mg m⁻² and PP rates ranged from 2.2 to 11.2 mmol m⁻² h⁻² and 1.8 to 10.8 mmol m⁻² h⁻² at plumewater and oceanic stations, respectively. A conservative estimate of annual integrated SiP in offshore waters of Amazon plume between April and August is 0.59 Tmol Si, based on mean SiP rates in plumewaters and satellite-derived estimates of the area of the Amazon plume. In conclusion, river plumewaters dramatically alter the silicon dynamics of the WTAO, forming extensive diatom-dominated phytoplankton blooms that may contribute significantly to the global Si budget as well as contributing to energy and matter flow off of the continental shelf.     

Isotopic and elemental indicators of nutrient sources and status of coastal habitats in the Caribbean Sea,Yucatan Peninsula,Mexico

Nutrient inputs associated with coastal population growth threaten the integrity of coastal ecosystems around the globe. In order to assess the threat posed by rapid growth in tourism, we analyzed the nutrient concentrations as well as the δ¹⁵N of NO₃⁻ and macrophytes to detect wastewater nitrogen (N) at 6 locations along a groundwater-dominated coastal seagrass bed on the Caribbean coast of Mexico. We predicted that locations with greater coastal development would have higher concentrations of dissolved inorganic nitrogen (DIN) and phosphorus (P), as well as δ¹⁵N of NO₃⁻, reflecting wastewater sources of N. However, concentrations of NO₃⁻ were not significantly different between developed (3.3 ± 5.3 μM NO₃⁻) and undeveloped (1.1 ± 0.7 μM) marine embayments. The most important control on DIN concentration appeared to be mixing of fresh and salt water, with DIN concentrations negatively correlated with salinity. The δ¹⁵N of NO₃⁻ was elevated at an inland pond (7.0 ± 0.42‰) and a hydrologically-connected tide pool (7.6 ± 0.57‰) approximately 1 km downstream of the pond. The elevated δ¹⁵N of NO₃⁻ at the pond was paralleled by high δ¹⁵N values of Cladophora sp., a ubiquitous green alga (10 ± 1‰). We hypothesize that inputs of nitrogen rich (NO₃⁻ > 30 μM) groundwater, characterized by ¹⁵N enriched signatures, flow through localized submarine groundwater discharges (SGD) and contribute to the elevated δ¹⁵N signatures observed in many benthic macrophytes. However, changes in nitrogen concentrations and isotope values over the salinity gradient suggest that other processes (e.g. denitrification) could also be contributing to the ¹⁵N enrichments observed in primary producers. More measurements are needed to determine the relative importance of nitrogen transformation processes as a source of ¹⁵N to groundwaters; however, it is clear that continued inputs of anthropogenic N via SGD have the potential to severely impact ecologically and economically valuable seagrass meadows and coral reefs along the Caribbean coast of Mexico.     

Nitrogen sources for new production in the NE Arabian Sea

New productivity measurements using the ¹⁵N tracer technique were conducted in the north-eastern (NE) Arabian Sea during six expeditions from 2003 to 2007, mostly in winter. Our results indicate that the NE Arabian Sea has a potential for higher new productivity during blooms. Nitrate uptake by plankton is the highest during late winter. New productivity and f-ratios in the NE Arabian Sea are mainly controlled by hydrodynamic and meteorological parameters such as wind strength, sea surface temperature (SST), mixed layer depth (MLD) and mixed layer nitrate. Deepening of the mixed layer supplies nitrate from below, which supports the observed nitrogen uptake. Higher f-ratios during blooms indicate the strong coupling between surface layers and sub-surface layers. Deepening of mixed layer below 100 m (from its inter-monsoon value between 30 and 40 m) transferred often more than 100 mmol N–NO₃ m^? 2 into the surface layers from below. The observed winter blooms in the region are supported by such input and are sustained for more than a month. Higher new productivity has been found in late winter, whereas transport of nitrate is maximum in early winter. In general, new production varies progressively during winter. Diurnal cycling of the mixed layer could be the reason for the under utilization of entrained nitrate during early winter. New productivity values and wind strength show significant differences during Feb–Mar 03 and Feb–Mar 04. These differences indicate that the winter cooling and parameters related the biological productivity also vary inter-annually. However, the difference between the new productivity values between Feb–Mar 03 and Feb–Mar 04 is much lower than the difference between Jan 03 and Feb–Mar 03. The results suggest that amplitude of seasonal variation is higher than the inter-annual variation in the region. During spring, Fickian diffusive fluxes of nitrate into the surface layer range from 0.51 to 1.38 mmol N–NO₃ m^? 2 day^? 1, and can account for 67% and 78% of the observed nitrogen uptake in the coastal and open ocean regions, respectively. We document the intra-seasonal and inter-annual variations in new productivity during winter and identify sources of nitrate which support the observed productivity during spring.     

Downward transport of organic carbon by diel migratory micronekton in the western equatorial Pacific:: its quantitative and qualitative importance

Using simultaneous sampling with a commercial-sized trawl, a zooplankton net, and a sediment trap, we evaluated the contribution of vertically migrating micronekton to vertical material transport (biological pump) at two stations (3°00′N, 146°00′E and 3°30′N, 145°20′E) in the western equatorial North Pacific. The gravitational sinking particulate organic carbon flux out of the euphotic zone was 54.8 mg C m⁻² day⁻¹. The downward active carbon flux by diel migrant mesozooplankton was 23.53 and 9.97 mg C m⁻² day⁻¹, and by micronekton 4.40 and 2.26mg C m⁻² day⁻¹ at the two stations. Assuming that the micronekton sampling efficiency of the trawl was 14%, we corrected the downward carbon flux due to micronekton respiration to 29.9 and 15.2mg C m⁻² day⁻¹, or 54.6 and 27.7% of the sinking particle flux at the two stations. The corrected micronekton gut fluxes were 1.53 and 0.97mg C m⁻² day⁻¹. The role of myctophid fish fecal matter as a possible food resource for deep-sea organisms, based on its fatty acid and amino acid analysis, is discussed.     

Benthic fluxes in a tidal salt marsh creek affected by fish farm activities: Río San Pedro (Bay of Cádiz,SW Spain)

Benthic fluxes of dissolved inorganic carbon, total alkalinity, oxygen, nutrients, nitrous oxide and methane were measured in situ at three sites of Río San Pedro salt marsh tidal creek (Bay of Cádiz, SW Spain) during three seasons. This system is affected by the discharges of organic carbon and nutrients from the surrounding aquaculture installations. Sediment oxygen uptake rates and inorganic carbon fluxes ranged respectively from 16 to 79 mmol O₂ m^? 2 d^? 1 and from 48 to 146 mmol C m^? 2 d^? 1. Benthic alkalinity fluxes were corrected for the influence of NH₄⁺ and NO₃^? + NO₂^? fluxes, and the upper and lower limits for carbon oxidation rates were inferred by considering two possible scenarios: maximum and minimum contribution of CaCO₃ dissolution to corrected alkalinity fluxes. Average C_ox rates were in all cases within ± 25% of the upper and lower limits and ranged from 40 to 122 mmol C m^? 2 d^? 1. Whereas carbon mineralization did not show significant differences among the sites, C_ox rates varied seasonally and were correlated with temperature (r² = 0.72). During winter and spring denitrification was estimated to account for an average loss of 46% and 75%, respectively, of the potentially recyclable N, whereas during the summer no net removal was observed. A possible shift from denitrification to dissimilatory nitrate reduction to ammonium (DNRA) during this period is argued. Dissolved CH₄ and N₂O fluxes ranged from 5.7 to 47 μmol CH₄ m^? 2 d^? 1 and 4.3 to 49 μmol N–N₂O m^? 2 d^? 1, respectively, and represented in all cases a small fraction of total inorganic C and N flux. Overall, about 60% of the total particulate organic matter that is discharged into the creek by the main fish farm facility is estimated to degrade in the sediments, resulting in a significant input of nutrients to the system.     

The supply of nutrients due to vertical turbulent mixing: A study at the Porcupine Abyssal Plain study site in the northeast Atlantic

As part of a multidisciplinary cruise to the Porcupine Abyssal Plain (PAP) study site (49°00′N 16°30′W), in June and July 2006, observations were made of the vertical nitrate flux due to turbulent mixing. Daily profiles of nitrate and turbulent mixing, at the central PAP site, give a mean nitrate flux into the euphotic zone of 0.09 (95% confidence intervals: 0.05–0.16) mmol N m⁻² d⁻¹. This is a factor of 50 lower than the mean observed rate of nitrate uptake within the euphotic zone (5.1±1.3 mmol N m⁻² d⁻¹). By using our direct observations to ‘validate’ a previously published parameterisation for turbulent mixing, we further quantify the variability in the vertical turbulent flux across a roughly 100×100 km region centred on the PAP site, using hydrographic data. The flux is uniformly low (0.08±0.26 mmol N m⁻² d⁻¹, the large standard deviation being due to a strongly non-Gaussian distribution) and is consistent with direct measurements at the central site. It is demonstrated that on an annual basis convective mixing supplies at least 40-fold more nitrate to the euphotic zone than turbulent mixing at this location. Other processes, such as those related with mesoscale phenomena, may also contribute significantly.     

Nitrogen assimilation and the f-ratio in the northwestern Indian Ocean during an intermonsoon period

Rates of nitrogen assimilation by phytoplankton were measured at 13 stations along a transect in the northwestern Indian Ocean, from the Gulf of Oman, southwards to approximately 8°N, during November and December 1994. Nitrate (NO₃), ammonium (NH₄) and urea assimilation were measured using simulated in situ ¹⁵N incubation techniques. These measurements were supported by simultaneous rate measurements of primary production using ¹⁴C incubation techniques and detailed vertical distributions of temperature and chlorophyll concentrations. Euphotic zone integrated nitrogen assimilation rates varied between 1.1 and 23.6 mmol N m^-2 day^-1, with generally higher rates occurring at the northern and southern ends of the transect. At the majority of stations ammonium was the preferred nitrogen substrate assimilated; the average integrated assimilation rate of ammonium being 3.7 mmol N m^-2 day^-1 compared to 1.6 and 1.8 mmol N m^-2 day^-1 for urea and nitrate respectively. This general preference is reflected in the low f-ratios, which were ⩽0.52 for all stations and in the relative preference indices (RPI) values which were consistently >1 for ammonium and <1 for nitrate. A further examination of the data has lead to an apparent partitioning of the northwestern Indian Ocean into 2 regions; a region north of 17°30′N and a region south of this, to about 8°N. This division is based on: (i) the relationship between the f-ratio and ambient nitrate levels; (ii) nitrogen assimilation and primary production and (iii) the biomass distribution. It is suggested that this partitioning should be investigated further with the development of biogeochemical provinces in mind and the estimation of f-ratios on much larger, horizontal scales.     

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.     

Impact of a cyclonic eddy on phytoplankton community structure and photosynthetic competency in the subtropical North Pacific Ocean

A synoptic spatial examination of the eddy Haulani (17–20 November 2000) revealed a structure typical of Hawaiian cyclonic eddies with divergent surface flow forcing the upward displacement of deep waters. Hydrographic surveys revealed that surface water in the eddy center was ca. 3.5°C cooler, 0.5 saltier, and 1.4 kg m⁻³ denser than surface waters outside the eddy. Vertically integrated concentrations of nitrate+nitrite, phosphate and silicate were enhanced over out-eddy values by about 2-fold, and nitrate+nitrite concentrations were ca. 8× greater within the euphotic zone inside the eddy than outside. Si:N ratios were lower within the upper mixed layer of the eddy, indicating an enhanced Si uptake relative to nitrate+nitrite. Chlorophyll a concentrations were higher within the eddy compared to control stations outside, when integrated over the upper 150 m, but were not significantly different when integrated over the depth of the euphotic zone. Photosynthetic competency, assessed using fast repetition-rate fluorometry, varied with the doming of the isopycnals and the supply of macro-nutrients to the euphotic zone. The physical and chemical environment of the eddy selected for the accumulation of larger phytoplankton species. Photosynthetic bacteria (Prochlorococcus and Synechococcus) and small (<3 μm diameter) photosynthetic eukaryotes were 3.6-fold more numerically abundant outside the eddy as compared to inside. Large photosynthetic eukaryotes (>3 μm diameter) were more abundant inside the eddy than outside. Diatoms of the genera Rhizosolenia and Hemiaulus outside the eddy contained diazotrophic endosymbiontic cyanobacteria, but these endosymbionts were absent from the cells of these species inside the eddy. The increase in cell numbers of large photosynthetic eukaryotes with hard silica or calcite cell walls is likely to have a profound impact on the proportion of the organic carbon production that is exported to deep water by sinking of senescent cells and cells grazed by herbivorous zooplankton and repackaged as large fecal pellets.