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.     

A time-series study of particulate matter export in the North Pacific Subtropical Gyre based on Th : U disequilibrium

Depth profiles of total ²³⁴Th (dissolved+particulate) were collected at Station ALOHA (22°45N, 158°00W) in the North Pacific Subtropical Gyre during 9 cruises from April 1999 to March 2000. Samples were collected and processed by a new 2 L technique that enables more detailed depth resolution then previous ²³⁴Th studies. Significant zones of particle export (²³⁴Th deficiency) and particle remineralization (²³⁴Th excess) were measured both temporally and with depth. ²³⁴Th derived particulate carbon (PC) and nitrogen (PN) fluxes were determined with steady-state and non-steady-state models and PC/²³⁴Th and PN/²³⁴Th ratios measured with both in situ pumps and free-drifting particle interceptor traps deployed at 150 m. ²³⁴Th based export estimates of 4.0±2.3 mmol C m⁻² d⁻¹ and 0.53±0.19 mmol N m⁻² d⁻¹, were approximately 60% higher than those measured in PIT style sediment traps from the same time period, 2.4±0.2 mmol C m⁻² d⁻¹ and 0.32±0.08 mmol N m⁻² d⁻¹. Most of this difference is attributable to two large export events that occurred during October and December 1999, when traps undercollected for ²³⁴Th by a factor of 2 to 4. ²³⁴Th export (ThE) ratios based on ²³⁴Th derived PC flux/¹⁴C based primary production ranged from 4% to 22% (average=8.8%). Our results confirm the recent estimates of C export by Emerson et al. (Nature 389 (1997) 951) and Sonnerup et al. (Deep-Sea Research I 46 (1999) 777) and indicate that C export from the oligotrophic ocean must be considered when discussing C sequestration in global climate change.     

Particle export within cyclonic Hawaiian lee eddies derived from Pb–Po disequilibrium

Particle export from the upper waters of the oligotrophic ocean may play a crucial role in the global carbon cycle. Mesoscale eddies have been hypothesized to inject new nutrients into oligotrophic surface waters, thereby increasing new production and particle export in otherwise nutrient deficient regimes. The E-Flux Program was a large multidisciplinary project designed to investigate the physical, biological and biogeochemical characteristics of cold-core cyclonic eddies that form in the lee of the Hawaiian Islands. There, we investigated particle dynamics using ²¹⁰Pb–²¹⁰Po disequilibrium. Seawater samples for ²¹⁰Pb and ²¹⁰Po were collected both within (IN) and outside (OUT) of two cyclones, Noah and Opal, at different stages of their evolution as well as from the eddy generation region. Particulate carbon (PC), particulate nitrogen (PN) and biogenic silica (bSiO₂) export fluxes were determined using water-column PC, PN, and bSiO₂ inventories and the residence times of ²¹⁰Po. PC and PN fluxes at 150 m ranged from 1.58±0.10 to 1.71±0.16 mmol C m⁻² d⁻¹ and 0.22±0.02 to 0.30±0.02 mmol N m⁻² d⁻¹ within Cyclones Opal and Noah. PC and PN fluxes at OUT stations sampled during both cruises were of similar magnitudes, 1.69±0.16 to 1.67±0.16 mmol C m⁻² d⁻¹ and 0.30±0.03 to 0.26±0.03 mmol N m⁻² d⁻¹. The bSiO₂ fluxes within Cyclone Opal were 0.157±0.010 mmol Si m⁻² d⁻¹ versus 0.025±0.002 mmol Si m⁻² d⁻¹ at OUT stations. These results of minimal PC and PN export, but significant eddy-induced bSiO₂ fluxes, agree very well with other studies that used a variety of direct and indirect methods. Thus, our results suggest that using elemental inventories and residence times of ²¹⁰Po is another independent and robust method for determining particle export and should be investigated more fully.     

Nitrogen cycling in deep-sea sediments of the Porcupine Abyssal Plain, NE Atlantic

Rates of transformation, recycling and burial of nitrogen and their temporal and spatial variability were investigated in deep-sea sediments of the Porcupine Abyssal Plain (PAP), NE Atlantic during eight cruises from 1996 to 2000. Benthic fluxes of ammonium (NH₄) and nitrate (NO₃) were measured in situ using a benthic lander. Fluxes of dissolved organic nitrogen (DON) and denitrification rates were calculated from pore water profiles of DON and NO₃, respectively. Burial of nitrogen was calculated from down core profiles of nitrogen in the solid phase together with ¹⁴C-based sediment accumulation rates and dry bulk density. Average NH₄ and NO₃-effluxes were 7.4 ± 19 μmol m⁻² d⁻¹ (n = 7) and 52 ± 30 μmol m⁻² d⁻¹ (n = 14), respectively, during the period 1996–2000. During the same period, the DON-flux was 11 ± 5.6 μmol m⁻² d⁻¹ (n = 5) and the denitrification rate was 5.1 ± 3.0 μmol m⁻² d⁻¹ (n = 22). Temporal and spatial variations were only found in the benthic NO₃ fluxes. The average burial rate was 4.6 ± 0.9 μmol m⁻² d⁻¹. On average over the sampling period, the recycling efficiency of the PON input to the sediment was 94% and the burial efficiency hence 6%. The DON flux constituted 14% of the nitrogen recycled, and it was of similar magnitude as the sum of burial and denitrification. By assuming the PAP is representative of all deep-sea areas, rates of denitrification, burial and DON efflux were extrapolated to the total area of the deep-sea floor (>2000 m) and integrated values of denitrification and burial of 8 ± 5 and 7 ± 1 Tg N year⁻¹, respectively, were obtained. This value of total deep-sea sediment denitrification corresponds to 3–12% of the global ocean benthic denitrification. Burial in deep-sea sediments makes up at least 25% of the global ocean nitrogen burial. The integrated DON flux from the deep-sea floor is comparable in magnitude to a reported global riverine input of DON suggesting that deep-sea sediments constitute an important source of DON to the world ocean.     

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.     

Rapid sediment accumulation and microbial mineralization in forests of the mangrove Kandelia candel in the Jiulongjiang Estuary, China

Rates of sediment accumulation and microbial mineralization were examined at three Kandelia candel forests spanning the intertidal zone along the south coastline of the heavily urbanized Jiulongljiang Estuary, Fujian Province, China. Mass sediment accumulation rates were rapid (range: 10–62 kg m⁻² y⁻¹) but decreased from the low- to the high-intertidal zone. High levels of radionuclides suggest that these sediments originate from erosion of agricultural soils within the catchment. Mineralization of sediment carbon and nitrogen was correspondingly rapid, with total rate of mineralization ranging from 135 to 191 mol C m⁻² y⁻¹ and 9 to 11 mol N m⁻² y⁻¹; rates were faster in summer than in autumn/winter. Rates of mineralization efficiency (70–93% for C; 69–92% for N) increased, as burial efficiency (7–30% for C; 8–31% for N) decreased, from the low-to the high-intertidal mangroves. Sulphate reduction was the dominant metabolic pathway to a depth of 1 m, with rates (19–281 mmol S m⁻² d⁻¹) exceeding those measured in other intertidal deposits. There is some evidence that Fe and Mn reduction-oxidation cycles are coupled to the activities of live roots within the 0–40 cm depth horizon. Oxic respiration accounted for 5–12% of total carbon mineralization. Methane flux was slow and highly variable when detectable (range: 5–66 μmol CH₄ m⁻² d⁻¹). Nitrous oxide flux was also highly variable, but within the range (1.6–106.5 μmol N₂O m⁻² d⁻¹) measured in other intertidal sediments. Rates of denitrification were rapid, ranging from 1106 to 3780 μmol N₂ m⁻² d⁻¹, and equating to 11–20% of total sediment nitrogen inputs. Denitrification was supported by rapid NH₄ release within surface deposits (range: 3.6–6.1 mmol m⁻² d⁻¹). Our results support the notion that mangrove forests are net accumulation sites for sediment and associated elements within estuaries, especially Kandelia candel forests receiving significant inputs as a direct result of intense human activity along the south China coast.     

Relationships between macroalgal biomass and nutrient concentrations in a hypertrophic area of the Venice Lagoon

Macroalgae biomass and concentrations of nitrogen, phosphorus and chlorophyll a were determined weekly or biweekly in water and sediments, during the spring-summer of 1985 in a hypertrophic area of the lagoon of Venice. Remarkable biomass production (up to 286 g m⁻² day⁻¹, wet weight), was interrupted during three periods of anoxia, when macroalgal decomposition (rate: up to 1000 g m⁻² day⁻¹) released extraordinary amounts of nutrients. Depending on the macroalgae distribution in the water column, the nutrients released in water varied from 3·3 to 19·1 μg-at litre⁻¹ for total inorganic nitrogen and from 1·8 to 2·7 μg-at litre⁻¹ for reactive phosphorus. Most nutrients, however, accumulated in the surficial sediment (up to 0·640 and to 3·06 mg g⁻¹ for P and N respectively) redoubling the amounts already stored under aerobic conditions, Phytoplankton, systematically below 5 mg m⁻³ as Chl. a, sharply increased up to 100 mg m⁻³ only after the release of nutrients in water by anaerobic macroalgal decomposition. During the algal growth periods, the N:P atomic ratio in water decreased to 0·7, suggesting that nitrogen is a growth-limiting factor. This ratio for surficial sediment was between 6·6 and 13·1, similar to that of macroalgae (8·6–12·0).     

Wintertime nutrients in the North Atlantic—new approaches and implications for new production estimates

Observations of wintertime nutrient concentrations in surface waters are scarce in the temperate and subarctic North Atlantic Ocean. Three new methods of their estimation from spring or early summer observations are described and evaluated. The methods make use of a priori knowledge of the vertical distribution of oxygen saturation and empirical relationships between nutrient concentrations and oxygen saturation. A south–north increase in surface water winter nutrient concentration is observed. Winter nitrate concentrations range from very low levels of about 0.5 μmol dm⁻³ at 33°N to about 13.5 μmol dm⁻³ at 60°N. Previous estimates of winter nitrate concentrations have been overestimates by up to 50%. At the Biotrans Site (47°N, 20°W), a typical station in the temperate Northeast Atlantic, a mean winter nitrate concentration of 8 μmol dm⁻³ is estimated, compared to recently published values between 11 and 12.5 μmol dm⁻³. It is shown that most of the difference is due to a contribution of remineralised nitrate that had not been recognized in previous winter nutrient estimates. Mesoscale variation of wintertime nitrate concentrations at Biotrans are moderate (less than ±15% of the regional mean value of about 8 μmol dm⁻³). Interannual variation of the regional mean is small, too. In the available dataset, there was only 1 year with a significantly lower regional mean winter nitrate concentration (7 μmol dm⁻³), presumably due to restricted deep mixing during an atypically warm winter. The significance of winter nitrate estimates for the assessment of spring-bloom new production and the interpretation of bloom dynamics is evaluated. Applying estimates of wintertime nitrate concentrations of this study, it is found that pre-bloom new production (0.275 mol N m⁻²) at Biotrans almost equals spring-bloom new production (0.3 mol N m⁻²). Using previous estimates of wintertime nitrate yields unrealistically high estimates of pre-bloom new production (1.21–1.79 mol N m⁻²) which are inconsistent with observed levels of primary production and the seasonal development of biomass.     

Elemental mercury concentrations and formation rates in the Scheldt estuary and the North Sea

Concentrations of Hg⁰ in surface waters and atmosphere of the Scheldt estuary and the North Sea are presented and their relationship with biological processes is discussed. Hg⁰ concentrations in the Scheldt estuary range from 0.1 to 0.38 pmol·l⁻¹ in the winter and from 0.24 to 0.65 pmol·l⁻¹ in the summer and show a positive relationship with phytoplankton pigments. In the North Sea Hg⁰ concentrations range from 0.06 to 0.8 pmol·l⁻¹ and are higher in coastal stations. Transfer velocities across the air–sea interface were calculated using a classical shear turbulence model. Volatilization fluxes of Hg⁰ were calculated for the Scheldt estuary and the North Sea. For the Scheldt estuary the fluxes range from 226–284 pmol·m⁻²·d⁻¹ in winter and 500–701 pmol·m⁻²·d⁻¹ in summer and for the North Sea the fluxes range from 59–1110 pmol·m⁻²·d⁻¹ for an average windspeed of 8.1 m·s⁻¹. These fluxes are comparable to the wet and dry depositional fluxes to the North Sea. Hg⁰ formation rates necessary to balance the volatilization fluxes vary from 0.2 to 4% d⁻¹.     

Seasonal phytoplankton composition, productivity and biomass in the Neuse River estuary, North Carolina

Phytoplankton community composition, productivity and biomass characteristics of the mesohaline lower Neuse River estuary were assessed monthly from May 1988 to February 1990. An incubation method which considered water-column mixing and variable light exposure was used to determine phytoplankton primary productivity. The summer productivity peaks in this shallow estuary were stimulated by increases in irradiance and temperature. However, dissolved inorganic nitrogen loading was the major factor controlling ultimate yearly production. Dynamic, unpredictable rainfall events determined magnitudes of seasonal production pulses through nitrogen loading, and helped determine phytoplankton species composition. Dinoflagellates occasionally bloomed but were otherwise present in moderate numbers; rainfall events produced large pulses of cryptomonads, and dry seasons and subsequent higher salinity led to dominance by small centric diatoms. Daily production was strongly correlated (r = 0·82) with nitrate concentration and inversely correlated (r = −0·73) with salinity, while nitrate and salinity were inversely correlated (r = −0·71), emphasizing the importance of freshwater input as a nutrient-loading source to the lower estuary. During 1989 mean daily areal phytoplankton production was 938 mgC m⁻², mean chlorophyll a was 11·8 mg m⁻³, and mean phytoplankton density was 1·56 × 10³ cells ml⁻¹. Estimated 1989 annual areal phytoplankton production for the lower estuary was 343 gC m⁻².     

Primary production and nutrient assimilation in the Iberian upwelling in August 1998

Primary production was measured during two Lagrangian experiments in the Iberian upwelling. The first experiment, in a body of upwelled water, measured day-to-day changes in phytoplankton activity as the water mass moved south along the shelf break. Nutrient concentrations decreased over a five day period, with concomitant increases in phytoplankton biomass. Initially the maximum phytoplankton biomass was in the upper 10m but after four days, a sub-surface chlorophyll maximum was present at 30m. Depth-integrated primary production at the beginning of the experiment was 70mmolC.m⁻².d⁻¹ (838mgC.m⁻².d⁻¹) and reached a maximum of 88mmolC.m⁻².d⁻¹ (1053mgC.m⁻².d⁻¹) on day 3. On day 1, the picoplankton fraction (<2μm) was slightly more productive than larger (>5μm) phytoplankton, but the increase in overall production during the drift experiment was by these larger cells. Nitrate was the dominant nitrogen source. As nutrient concentrations declined, ammonium became increasingly more important as a nitrogen source and the f-ratio decreased from 0.7 to 0.5. Picoplankton cells (<2μm) were responsible for most (65–80%) of the ammonium uptake. The C:N:P uptake ratios were very close to the Redfield ratio for the first four days but as nutrients became depleted high C:N uptake ratios (11 to 43) were measured. Over the period of the experiment, nitrate concentration within the upper 40m decreased by 47.91mmolN.m⁻². In vitro estimates, based on ¹⁵N nitrate uptake, accounted for 56% of the decrease in nitrate concentration observed in the drifting water mass. Ammonium uptake over the same four day period was 16.28mmolN.m⁻², giving a total nitrogen uptake of 43.18mmolN.m⁻².In the second experiment, an offshore filament was the focus and a water mass was sampled as it moved offshore. Nutrient concentrations were very low (nitrate was <10nmol l⁻¹ and ammonium was 20–40nmol l⁻¹). Primary production rate varied between 36mmolC.m⁻².d⁻¹ (436mgC.m⁻².d⁻¹) and 21mmolC.m⁻².d⁻¹ (249mgC.m⁻².d⁻¹). Picophytoplankton was the most productive fraction and was responsible for a constant proportion (ca 0.65) of the total carbon fixation. Uptake rates of both nitrate and ammonium were between 10 and 20% of those measured in the upwelling region. Urea could be a very significant nitrogen source in these waters with much higher uptake rates than nitrate or ammonium; urea turnover times were ca. one day but the source of the urea remains unknown. Urea uptake had a profound effect on calculated f ratios. If only nitrate and ammonium uptake was considered, f ratios were calculated to be 0.42–0.46 but inclusion of urea uptake reduced the f ratio to <0.1. The primary production of this oligotrophic off-shore filament was driven by regenerated nitrogen.     

Summer algal blooms in a coastal ecosystem: the role of atmospheric deposition versus entrainment fluxes

The nitrogen inputs from atmospheric deposition and bottom water entrainment to the surface layer were modelled in the summer period (May–September) over a 11-year period (1989–1999) and compared to investigate the significance of these fluxes for generating blooms in the Kattegat. In the summer periods the average atmospheric deposition was 2.81 mg N m⁻² d⁻¹ compared to average entrainment fluxes of 5.42 mg N m⁻² d⁻¹, 1.21 mg N m⁻² d⁻¹ and 1.15 mg N m⁻² d⁻¹ for the northern, central and southern part of the Kattegat, respectively. Atmospheric nitrogen deposition alone could not sustain biomass increases associated with observed blooms and entrainment fluxes dominated the high nitrogen inputs to the surface layer. The potential for a bloom through growth was typically obtained after several days of high nitrogen inputs from entrainment in the frontal area of the northern Kattegat and to some extent from atmospheric deposition. The modelled nitrogen input in this area could account directly for 30% of the observed blooms in the Northern sub-basin, and through advective transport 24% and 19% of the observed blooms in the central and southern Kattegat. The direct nitrogen inputs through atmospheric deposition and entrainment to the central and southern sub-basins were small and could not be linked to any bloom observation.     

Factors influencing fatty acid and hydrocarbon composition of sedimenting particles in the northeastern Adriatic Sea

Fatty acids and hydrocarbons of sedimenting particles were investigated in the northeastern Adriatic Sea from November 1988 to December 1989. Particles were collected at approximately monthly intervals, using sediment traps deployed at 30 m depth (2 m above bottom). Seasonal changes in sedimentation of particulate matter were very pronounced. Hydrocarbon fluxes and concentrations were found to vary significantly depending on the season. They averaged 2.69 ± 1.44 mg m⁻² day⁻¹ and 232.4 ± 90.93 μg g⁻¹ in winter, respectively. In late spring-early summer the corresponding values amounted to 0.045 ± 0.015 mg m⁻² day⁻¹ and 13.72 ± 5.56 μg g⁻¹, and they increased towards autumn, when mean values of 0.517 ± 0.228 mg m⁻² day⁻¹ and 98.86 ± 48.72 μg g⁻¹ were obtained. In contrast, fatty acid fluxes and concentrations were low during winter (0.26 ± 0.08 mg m⁻² day⁻¹ and 21.95 ± 3.35 μg g⁻¹), increased slightly towards the summer (0.48 ± 0.12 mg m⁻² day⁻¹ and 139.9 ± 44.6 μ g⁻¹) and reached maximum rate and concentration in autumn, when average values were 1.98 ± 1.30 mg m² day⁻¹ and 489.1 ± 186.7 μg g⁻¹, respectively. The differences in composition, concentrations and fluxes of the fatty acids and hydrocarbons were related to the sources of sedimenting material, reflecting the influence of resuspension of bottom sediments during winter and the appearance of mucus aggregates during summer and their subsequent deposition in autumn.     

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

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

Comparison of factors controlling phytoplankton productivity in the NE and NW subarctic Pacific gyres

The subarctic North Pacific is one of the three major high nitrate low chlorophyll (HNLC) regions of the world. The two gyres, the NE and the NW subarctic Pacific gyres dominate this region; the NE subarctic Pacific gyre is also known as the Alaska Gyre. The NE subarctic Pacific has one of the longest time series of any open ocean station, primarily as a result of the biological sampling that began in 1956 on the weathership stationed at Stn P (50°N, 145°W; also known as Ocean Station Papa (OSP)). Sampling along Line P, a transect from the coast (south end of Vancouver Island) out to Stn P has provided valuable information on how various parameters change along this coastal to open ocean gradient. The NW subarctic Pacific gyre has been less well studied than the NE gyre. This review focuses mainly on the NE gyre because of the large and long term data set available, but makes a brief comparison with the NW gyre. The NE gyre has saturating NO₃ concentrations all year (winter = about 16 μM and summer = about 8 μM), constantly very low chlorophyll (chl) (usually <0.5 mg m⁻³) which is dominated by small cells (<5 μm). Primary productivity is low (about 300–600 mg C m⁻² d⁻¹ and varies little (2 times) seasonally. Annual primary productivity is 3 to 4 times higher than earlier estimates ranging from 140 to 215 g C m⁻² y⁻¹. Iron limits the utilization of nitrate and hence the primary productivity of large cells (especially diatoms) except in the winter when iron and light may be co-limiting. There are observations of episodic increases in chl above 1 mg m⁻³, suggesting episodic iron inputs, most likely from Asian dust in the spring/early summer, but possibly from horizontal advection from the Alaskan Gyre in summer/early fall. The small cells normally dominate the phytoplankton biomass and productivity, and utilize the ammonium produced by the micrograzers. They do not appear to be Fe-limited, but are controlled by microzooplankton grazers. The NW Subarctic Gyre has higher nutrient concentrations and a shallower summer mixed depth and photic zone than Stn P in the NE gyre. Chl concentrations tend to be higher (0.5 to 1.5 μg L⁻¹) than Stn P, but primary productivity in the summer is similar to Stn P (600 mg C m⁻² d⁻¹). There are no seasonal data from this gyre. Iron enrichment experiments in October, resulted in an increase in chl (mainly the centric diatom Thalassiosira sp.) and a draw down of nitrate, suggesting that large phytoplankton are Fe-limited, similar to Stn P.     

Nitrogen cycling in the southern North Sea: consequences for total nitrogen transport

Nitrate and ammonium uptake rates were measured during a series of cruises in the well-mixed region of the southern North Sea from February to September. Water column-integrated uptake rates ranged between 0.01 and 8.7 mmol N m⁻² d⁻¹ and 0.01 and 12.2 mmol N m⁻² d⁻¹ for nitrate and ammonium, respectively, with ammonium uptake dominating after the phytoplankton spring bloom in May. A moored buoy continuously measuring nitrate and chlorophyll a and seabed current meters were also deployed in the central southern North Sea in the region of the East Anglian plume—a permanent physical feature which transports nutrients towards continental Europe. This enabled the flux of water and hence of nutrients across the southern North Sea to be determined and an assessment of the contribution of freshwater nutrients to the flux to be made. A simple box model is developed to relate the phytoplankton uptake of nitrate and ammonium to the transport of nitrate, ammonium and particulate organic matter (POM) across the southern North Sea. This showed the importance of the plume region of the North Sea in the processing of nitrogen, with nitrate dominating total nitrogen transport prior to the spring bloom (10 340×10³ kg N inflow to the plume in March) and transport of nitrogen as ammonium, nitrate and POM in approximately equivalent amounts during summer (2560, 2960 and 2151×10³ kg N inflow to the plume, respectively, in July). The box model also demonstrates more generally the need to assess nitrogen transport as nitrate, ammonium and POM if an improved understanding of the impact of nutrient input in shelf seas is to be achieved.     

Carbon flux in the northwest Mediterranean estimated from microbial production

Spring profiles of microbial production derived from the dark incorporation of tritiated leucine and tritiated thymidine in the northwest Mediterranean show an exponential decline with depth. Assuming this to represent a steady-state balance between microbial respiration and the downward flux of carbon, the downward flux is estimated as (1−/)p/b, where p is the microbial production, their gross growth efficiency and b the coefficient of exponential decline with depth. Summer profiles, ranging over about 3° of latitude and 4° of longitude, were well fitted by a two-component exponential decline, suggesting two distinct microbial substrates. Values of b for the more rapidly declining component varied between 0.01 and 0.06 m⁻¹ according to location. In the case of the slower component, b was estimated as 0.002 m⁻¹, and did not vary significantly over the region. Estimated fluxes of carbon at the surface are 123–335 mg m⁻² d⁻¹ for the fast and 95 mg m⁻² d⁻¹ for the slow component. Below about 200 m, carbon flux is dominated by the slow component. Flux estimates are compatible with flux estimates from sediment traps in the same region. The observed changes between the spring and summer profiles, combined with the horizontal homogeneity of the summer profiles below 200 m, are consistent with a downward transport of about 5–10 m d^–1, implying a significant dispersive component to the observed fluxes.     

Seasonal variations in faunal distribution and activity across the continental slope of the goban spur area (NE Atlantic)

Density, biomass and community structure of macrofauna were estimated together with several sediment characteristics at seven stations ranging from 208 m to 4460 m water depth along the OMEX transect in the Goban Spur area (NE Atlantic) during three seasons (October 1993, May 1994, and August 1995). Median grain size decreased with increasing water depth and showed no differences between the seasons. The percentages of organic carbon and total nitrogen were highest at mid-slope depths (1000 to 1500 m), and were significantly higher in August at the upper part of the slope to a depth of 1500 m. The C:N ratio in the surface layer amounted to 7 to 8 in May, 10 to 12 in August and 14 to 17 in October at all stations (except the deepest at 4460 m, where it was 11 in May and August), indicating arrival of fresh phytodetritus in May, and therefore seasonality in food input to the benthos. Densities of macrofauna decreased exponentially with increasing water depth. Significantly higher densities of macrofauna were found in May at the upper part of the slope to a depth of 1500 m. These differences were mainly due to high numbers of postlarvae of echinoids at the shallowest station and ophiuroids at the deeper stations. Biomass values also decreased with increasing water depth, but biomass was relatively high at the 1000 m station and low at 1500 m, due to relatively high and low mean weights of the individual macrofaunal specimens. No significant differences in biomass were found between the seasons. Respiration was high (15 to 20 mgC·m⁻²·d⁻¹) in May at the upper part of the slope to a depth of 1000 m and low (1–3 mg C·m⁻²·d⁻¹) at the deeper part. At the shallowest stations to a depth of 1000 m respiration was highest in May, at the mid-slope stations (1400–2200 m) it was highest in August, whereas the deepest stations (3600 to 4500 m) did not show any differences in respiration rates. In conclusion; seasonal variation in organic input is reflected in denstiy, community structure and activity of the macrofauna along the continental slope in the NE Atlantic.     

Mesozooplankton biomass and grazing responses to Cyclone Opal, a subtropical mesoscale eddy

As part of E-Flux III cruise studies in March 2005, plankton net collections were made to assess the effects of a cyclonic cold-core eddy (Cyclone Opal) on the biomass and grazing of mesozooplankton. Mesozooplankton biomass in the central region of Cyclone Opal, an area of uplifted nutricline and a subsurface diatom bloom, averaged 0.80±0.24 and 1.51±0.59 g DW m⁻², for day and night tows, respectively. These biomass estimates were about 80% higher than control (OUT) stations, with increases more or less proportionately distributed among size classes from 0.2 to >5 mm. Though elevated relative to surrounding waters south of the Hawaiian Islands (Hawai’i lee), total biomass and size distribution in Cyclone Opal were almost exactly the same as contemporary measurements made at Stn. ALOHA, 100 km north of the islands, by the HOT (Hawaii Ocean Time-series) Program. Mesozooplankton biomass and community composition at the OUT stations were also similar to ALOHA values from 1994 to 1996, preceding a recent decadal increase. These comparisons may therefore provide insight into production characteristics or biomass gradients associated with decadal changes at Stn. ALOHA. Gut fluorescence estimates were higher in Opal than in ambient waters, translating to grazing impacts of 0.11±0.02 d⁻¹ (IN) versus 0.03±0.01 d⁻¹ (OUT). Over the depth-integrated euphotic zone, mesozooplankton accounted for 30% of the combined grazing losses of phytoplankton to micro- and meso-herbivores in Opal, as compared to 13% at control stations. Estimates of active export flux by migrating zooplankton averaged 0.81 mmol C m⁻² d⁻¹ in Cyclone Opal and 0.37 mmol C m⁻² d⁻¹ at OUT stations, 53% and 24%, respectively, of the carbon export measured by passive sediment traps. Migrants also exported 0.18 mmol N m⁻² d⁻¹ (117% of trap N flux) in Cyclone Opal compared to 0.08 mmol N m⁻² d⁻¹ (51% of trap flux) at control stations. Overall, the food-web importance of mesozooplankton increased in Cyclone Opal both in absolute and relative terms. Diel migrants provided evidence for enhanced export flux in the eddy that was missed by sediment trap and ²³⁴Th techniques, and migrant-mediated flux was the major export term in the observed bloom-perturbation response and N mass balance of the eddy.     

Coupling of biological and physical regimes across the Antarctic Polar Front as reflected by nitrogen production and recycling

Oceanographic samples were collected across the Antarctic Polar Front (APF) region in the vicinity of 60°S, 170°W during the US JGOFS program from December 1997 to March 1998. This paper reports the uptake rates of new (nitrate) and regenerated (ammonium and urea) nitrogen measured by ¹⁵N tracer techniques together with the levels of ammonium, urea-N and dissolved free amino acids (DFAAs) during December and mid-February–March. The APF was an important biological boundary, and in December rates of new (nitrate) uptake were greatest south of the APF, exceeding 10 mmol m⁻² d⁻¹ near the retreating ice edge. In February, nitrate uptake rates were an order of magnitude lower. Rates of ammonium uptake in both periods were greater in the warmer water north of the front. Nitrogen f-ratios varied from 0.50 to less than 0.05, with larger values associated with the >5 μm size fraction at the ice edge and generally lower values north of the APF. Urea was an important nitrogen source north of the APF, and lowered f-ratios there by 22% on average when included as part of total nitrogen uptake. Urea uptake was less important south of the APF. Ammonium concentrations increased dramatically south of the APF later in the season, suggesting a system dominated by regeneration. Seasonal changes in the concentrations of regenerated organic compounds such as urea and DFAAs were less obvious, although DFAAs exhibited consistent maxima in the high flow regions of the APF. A mass balance based of ammonium fluxes suggests that nitrification was significant at locations south of the APF in February. In these nitrate-replete waters, light/mixing conditions in the surface water (the Sverdrup criterion) accounted for over 50% of the variance in nitrate uptake rates. The stability responsible for higher new production south of the APF is due both to the separation of this region from the maximum zonal wind field to the north as well as to melt-water contribution from the retreating ice field. Estimated new production and exportable carbon production exceeded 500 mmol nitrate m⁻² yr⁻¹ and 40 g C m⁻² yr⁻¹, respectively, south of the APF. Thus, new production in the marginal ice zone of the Southern Ocean rivals that in coastal systems and indicates that this is an important region for export production.