The influence of a mature cyclonic eddy on particle export in the lee of Hawaii

Mesoscale eddies may enhance primary production (PP) in the open ocean by bringing nutrient-rich deep waters into the euphotic zone, potentially leading to increased transport of particles to depth. This hypothesis remains controversial, however, due to a paucity of direct particle export measurements. In this study, we investigated particle dynamics using ²³⁴Th–²³⁸U disequilibria within a mesoscale cold-core eddy, Cyclone Opal, which formed in the lee of the Hawaiian Islands. ²³⁴Th samples were collected along two transects across Cyclone Opal as well as during a time-series within the eddy core during a decaying diatom bloom. Particulate carbon (PC), particulate nitrogen (PN) and biogenic silica (bSiO₂) fluxes at 150 m varied spatially and temporally within the eddy and strongly depended on the ²³⁴Th model formulation used (e.g., steady state versus non-steady state, inclusion of upwelling, etc.). Particle fluxes estimated from a steady state model assuming an upwelling rate of 2 m day⁻¹ yielded the best fit to sediment-trap data. These ²³⁴Th-derived particle fluxes ranged from 332±14 to 1719±53 μmol C m⁻² day⁻¹, 27±3 to 114±12 μmol N m⁻² day⁻¹, and 33±20 to 309±73 μmol Si m⁻² day⁻¹. Although PP rates within Cyclone Opal were elevated by a factor of 2–3, PC and PN fluxes were the same, within error, inside and outside of Cyclone Opal. The ratio of PC export to PP remained surprisingly low at <0.03 and similar to those measured in surrounding waters. In contrast, bSiO₂ fluxes within the eddy core were three times higher. Detailed analyses of ²³⁴Th depth profiles consistently showed excess ²³⁴Th at 100–175 m, associated with the remineralization and possible accumulation of suspended and dissolved organic matter from the surface. We suggest that strong microzooplankton grazing facilitated particulate organic matter recycling and resulted in the export of empty diatom frustules. Thus, while eddies may increase PP, they do not necessarily increase PC and PN export to deep waters. This may be a general characteristic of wind-driven cyclonic eddies of the North Pacific Subtropical Gyre and suggests that eddies may preferentially act as a silica pump, thereby playing an important role in promoting silicic-acid limitation in the region.     

The influence of island generated eddies on the carbon dioxide system, south of the Canary Islands

Measurements of surface partial pressure of CO₂ and water column alkalinity, pH_T, nutrients, oxygen, fluorescence and hydrography were carried out, south of the Canary Islands during September 1998. Cyclonic and anticyclonic eddies were alternatively observed from the northwestern area to the central area of the Canary Islands. Nutrient pumping and vertical uplifting of the deep chlorophyll maximum by cyclonic eddies were also ascertained by upward displacement of dissolved inorganic carbon. A model was applied to determine the net inorganic carbon balance in the cyclonic eddy. The fluxes were determined considering both the diffusive and convective contributions from the upward pumping and the corresponding horizontal transport of water outside the area. An increase in the total inorganic carbon concentration in the upper layers inside the eddy field of 133 mmol C m^− 2 d^− 1 was determined. The upward flux of inorganic carbon decreased the effect of the increased primary production on the carbon dioxide chemistry. The reduced fCO₂ inside the cyclonic eddy, 15 μatm lower than that observed in non-affected surface water, was explained by thermodynamic aspects, biological activity, eddy upward pumping and diffusion and air–sea water exchange effects.     

Trophic pathways and carbon flux patterns in the Laptev Sea

The Laptev Sea is a high-Arctic epicontinental sea north of Siberia (Russia) that is one of the least understood regions of the world’s ocean. It is characterized by a shallow and broad shelf plateau, high influx of river water, sediments and nutrients during summer, long-lasting sea-ice cover from October to May, and the formation of a narrow flaw-lead polynya off the fast-ice edge during winter.Here, we describe results of a German–Russian research project (1993-present), presenting the distribution patterns and dynamics of its marine flora and fauna, as well as pathways and processes of coupling between sea-ice, water-column and sea-floor biota.Three ecological zones are distinguished along a combined east–west and Lena-impact gradient, differing in the composition of pelagic and benthic communities. In general, high Chl a concentrations in the sediments indicate a tight coupling between sympagic and pelagic primary production and nutrient supply to the benthos throughout the entire Laptev Sea. However, there were pronounced regional differences between the ecological zones in magnitude of primary production and trophic dynamics. Primary production during the ice-free summer was highest in the estuarine zone most strongly influenced by the Lena River (210 mg C m⁻² day⁻¹). The western and northeastern Laptev Sea yielded 55 and 95 mg C m⁻² day⁻¹, respectively. Moreover, the zones differed in the partitioning of carbon flux between zooplankton and benthic food webs. In the Lena zone zooplankton carbon demand was about 31 mg C m⁻² day⁻¹ whereas in the western zone it was 21 mg C m⁻² day⁻¹ and in the eastern zone 4 mg C m⁻² day⁻¹. Total benthic carbon demand was 32 mg C m⁻² day⁻¹ for the Lena zone, 56 mg C m⁻² day⁻¹ in the western zone and 100 mg C m⁻² day⁻¹ in the northeastern zone.A carbon budget constructed for the Laptev Sea indicates that (1) a high proportion of primary production is channelled through the benthic trophic web, bypassing the pelagic trophic web, and (2) autochthonous primary production in the northeastern and western Laptev Sea might not be sufficient to fuel both pelagic and benthic secondary production and, hence, input of allochthonous organic carbon is required to balance the overall carbon demand.     

Physical dynamics and biological implications of Cyclone Noah in the lee of Hawaii during E-Flux I

Quasi-synoptic observations of the horizontal and vertical structure of a cold-core cyclonic mesoscale eddy feature (Cyclone Noah) were conducted in the lee of Hawai’i from November 4–22, 2004 as part of the E-Flux interdisciplinary collaborative research program. Cyclone Noah appears to have spun up to the southwest of the ‘Alenuihaha Channel (between Maui and Hawai’i) as a result of strong and persistent northeasterly trade winds through the channel. Shipboard hydrographic surveys 2.5 months later suggest that Noah weakened and was in a hypothesized spin-down phase of its life cycle. Although the initial surface expression of Noah was limited in scale to 40 km in diameter and, as evidenced by surface temperatures, 2–3 °C cooler than the surrounding waters, depth profiles revealed a fully developed semi-elliptical shallow feature (200 m), 144 km long and 90 km wide (based on sigma-t=23 kg m⁻³) with tangential speeds of 40–80 cm s⁻¹, and substantial isopycnal doming. Potential vorticity distribution of Noah suggests that radial horizontal flow of the core water was inhibited from the surface to depths of 75 m, with high vorticity confined above the sigma-t=23.5 kg m⁻³ isopycnal surface. Upward displacements of isopycnal surfaces in the eddy's center (50 m) were congruent with enhanced pigment concentrations (0.50 mg m⁻³). Comparisons of the results obtained for E-Flux I (Noah) and E-Flux III (Opal) suggest that translation characteristics of cyclonic Hawaiian lee eddies may be important in establishing the biogeochemical and biological responses of the oligotrophic ocean to cyclonic eddies.     

Mesozooplankton dynamics in nearshore waters of the Great Barrier Reef

Zooplankton dynamics (community composition, juvenile somatic growth rate, adult egg production, secondary production) were studied in coastal waters of the Great Barrier Reef. Two sectors were compared, one adjacent to a catchment of near-pristine land use patterns, the other to a more intensively farmed catchment. Sampling was conducted in the austral winter (August) and summer (January–March) of two succeeding years. Gradients in zooplankton community composition were weak, with only moderate effects of season and sector. Overall, 37% of zooplankton biomass was in the 73–150 μm size fraction, 26% in the 150–350 μm fraction, and 38% was >350 μm. There was no biomass difference and only small differences in community composition between samples taken during the day and at night; ostracods and large calanoid copepods were occasionally more common at night. Carbon-specific growth rates averaged 0.29 d⁻¹ for cyclopoid copepods and 0.35 d⁻¹ for calanoid copepods, with no difference between sectors. Calanoid copepod growth showed a significant relationship to chlorophyll concentration, but cyclopoid copepods did not. Copepod egg production was low (7.9 ± 5.9 eggs female⁻¹ d⁻¹) and apparently food-limited. Copepod secondary production was lower in August (mean = 2.6, range 1.4–4.0 mg C m⁻² d⁻¹) than in January–March (mean = 8.5, range 2.4–15.5 mg C m⁻² d⁻¹). Secondary production by mesozooplankton in the 73–100 μm size range averaged 0.9% of total phytoplankton production.     

Primary production and implications for metabolic balance in Hawaiian lee eddies

Recent discrepancies between geochemical and biological approaches for determining whether ocean ecosystems are net heterotrophic or net autotrophic have led to uncertainty in the net metabolic state of open ocean ecosystems. Geochemical approaches indicate that the oceans are net positive autotrophic, but direct observations based on short-term incubation techniques suggest that the ocean is in a state of net heterotrophy. One hypothesis for the apparent discrepancy is that net autotrophic production occurs in aperiodic “bursts,” which are superimposed on a more constant background state of net heterotrophy. Mixing events, which introduce new nutrients to the surface ocean, provide one mechanism for fueling such aperiodic bursts of net production. In conjunction with the Eddy Flux (E-Flux) program in the lee of the Hawaiian Islands during winter 2004–2005, we examined the relationship between photosynthesis and irradiance (P vs. E) in surface waters inside and outside of two cold-core, cyclonic eddies, and conducted five incubation experiments to examine the metabolic response of mixed-layer plankton communities to nutrient-rich deep-sea water additions. Our results showed that in the mixed layer, maximum rates of light-saturated photosynthesis, derived from photosynthesis–irradiance experiments were not significantly different inside vs. outside the eddies (p=0.35 and 0.44 for E-Flux I and E-Flux III, respectively). Addition of nutrients to mixed-layer water showed that (1) gross primary production (GPP) became decoupled from a more constant rate of respiration and (2) net system metabolism shifted from approximate balance, or slight net heterotrophy, to a demonstrably net autotrophic system. From these results, we determined that the threshold GPP for net autotrophic production for the mixed layer of the study region was 1.65 mmol O₂ m⁻³ d⁻¹, which is consistent with previous estimates for the oligotrophic open ocean.     

The carbon dioxide system and net community production within a cyclonic eddy in the lee of Hawaii

The dynamics of dissolved inorganic carbon (DIC) and processes controlling net community production (NCP) were investigated within a mature cyclonic eddy, Cyclone Opal, which formed in the lee of the main Hawaiian Islands in the subtropical North Pacific Gyre. Within the eddy core, physical and biogeochemical properties suggested that nutrient- and DIC-rich deep waters were uplifted by 80 m relative to surrounding waters, enhancing biological production. A salt budget indicates that the eddy core was a mixture of deep water (68%) and surface water (32%). NCP was estimated from mass balances of DIC, nitrate+nitrite, total organic carbon, and dissolved organic nitrogen, making rational inferences about the unobserved initial conditions at the time of eddy formation. Results consistently suggest that NCP in the center of the eddy was substantially enhanced relative to the surrounding waters, ranging from 14.1±10.6 (0–110 m: within the euphotic zone) to 14.2±9.2 (0–50 m: within the mixed layer) to 18.5±10.7 (0–75 m: within the deep chlorophyll-maximum layer) mmol C m⁻² d⁻¹ depending on the depth of integration. NCP in the ambient waters outside the eddy averaged about 2.37±4.24 mmol C m⁻² d⁻¹ in the mixed layer (0–95 m). Most of the enhanced NCP inside the eddy appears to have accumulated as dissolved organic carbon (DOC) rather than exported as particulate organic carbon (POC) to the mesopelagic. Our results also suggest that the upper euphotic zone (0–75 m) above the deep chlorophyll maximum is characterized by positive NCP, while NCP in the lower layer (>75 m) is close to zero or negative.     

Atmospherically-promoted photosynthetic activity in a well-mixed ecosystem: Significance of wet deposition events of nitrogen compounds

Wet atmospheric deposition of dissolved N, P and Si species is studied in well-mixed coastal ecosystem to evaluate its potential to stimulate photosynthetic activities in nutrient-depleted conditions. Our results show that, during spring, seawater is greatly depleted in major nutrients: Dissolved Inorganic Nitrogen (DIN), Dissolved Inorganic Phosphorus (DIP) and Silicic acid (Si), in parallel with an increase of phytoplanktonic biomass. In spring (March–May) and summer (June–September), wet atmospheric deposition is the predominant source (>60%, relative to riverine contribution) for nitrates and ammonium inputs to this N-limited coastal ecosystem. During winter (October–February), riverine inputs of DIN predominate (>80%) and are annually the most important source of DIP (>90%). This situation allows us to calculate the possibility for a significant contribution to primary production in May 2003, from atmospheric deposition (total input for DIN ≈300 kg km⁻² month⁻¹). Based on usual Redfield ratios and assuming that all of the atmospheric-derived N (AD-N) in rainwater is bioavailable for phytoplankton growth, we can estimate new production due to AD-N of 950 mg C m⁻² month⁻¹, during this period of depletion in the water column. During the same episode (May 2003), photosynthetic activity rate, considered as gross primary production, was estimated to approximately 30 300 mg C m⁻² month⁻¹. Calculation indicates that new photosynthetic activity due to wet atmospheric inputs of nitrogen could be up to 3%.     

Dimethylsulfide production in Sargasso Sea eddies

Lagrangian time series of dimethylsulfide (DMS) concentrations from a cyclonic and an anticyclonic eddy in the Sargasso Sea were used in conjunction with measured DMS loss rates and a model of vertical mixing to estimate gross DMS production in the upper 60 m during summer 2004. Loss terms included biological consumption, photolysis, and ventilation to the atmosphere. The time- and depth (0–60 m)-averaged gross DMS production was estimated to be 0.73±0.09 nM d⁻¹ in the cyclonic eddy and 0.90±0.15 nM d⁻¹ in the anticyclonic eddy, with respective DMS replacement times of 5±1 and 6±1 d. The higher estimated rate of gross production and lower measured loss rate constants in the anticyclonic eddy were equally responsible for this eddy's 50% higher DMS inventory (0–60 m). When normalized to chlorophyll and total dimethylsulfoniopropionate (DMSP), estimated gross production in the anticyclonic eddy was about twice that in the cyclonic eddy, consistent with the greater fraction of phytoplankton that were DMSP producers in the anticyclonic eddy. Higher rates of gross production were estimated below the mixed layer, contributing to the subsurface DMS maximum found in both eddies. In both eddies, gas exchange, microbial consumption, and photolysis were roughly equal DMS loss terms in the surface mixed layer (0.2–0.4 nM d⁻¹). Vertical mixing was a substantial source of DMS to the surface mixed layer in both eddies (0.2–0.3 nM d⁻¹) owing to the relatively high DMS concentrations below the mixed layer. Estimated net biological DMS production rates (gross production minus microbial consumption) in the mixed layer were substantially lower (by almost a factor of 3) than those estimated in a previous study of the Sargasso Sea, which may explain the relatively low mixed-layer DMS concentrations found here during July 2004 (3 nM) compared to previous summers (4–6 nM).     

The distributions of, and relationship between, He and nitrate in eddies

We present and discuss the distribution of ³He and its relationship to nutrients in two eddies (cyclone C1 and anticyclone A4) with a view towards examining eddy-related mechanisms whereby nutrients are transported from the upper 200–300 m into the euphotic zone of the Sargasso Sea. The different behavior of these tracers in the euphotic zone results in changes in their distributions and relationships that may provide important clues as to the nature of physical and biological processes involved.The cyclonic eddy (C1) is characterized by substantial ³He excesses within the euphotic zone. The distribution of this excess ³He is strongly suggestive of both past and recent ongoing deep-water injection into the euphotic zone. Crude mass balance calculations suggest that an average of approximately 1.4±0.7 mol m⁻² of nitrate has been introduced into the euphotic zone of eddy C1, consistent with the integrated apparent oxygen utilization anomaly in the aphotic zone below. The ³He–NO₃ relationship within the eddy deviates substantially from the linear thermocline trend, suggestive of incomplete drawdown of nutrients and/or substantial mixing between euphotic and aphotic zone waters.Anticyclone (A4) displays a simpler ³He–NO₃ relationship, but is relatively impoverished in euphotic zone excess ³He. We suggest that because of the relatively strong upwelling and lateral divergence of water the residence time of upwelled ³He is relatively short within the euphotic zone of this eddy. An estimate of the recently upwelled nutrient inventory, based on the excess ³He observed in A4's lower euphotic zone, is stoichiometrically consistent with the oxygen maximum observed in the euphotic zone.     

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.     

Elevated biomass of mesozooplankton and enhanced fecal pellet flux in cyclonic and mode-water eddies in the Sargasso Sea

Accumulating evidence points to the importance of mesoscale eddies in supplying nutrients to surface waters in oligotrophic gyres. However, the nature of the biological response and its evolution over time has yet to be elucidated. Changes in mesozooplankton community composition due to eddy perturbation also could affect biogeochemical cycling. Over the course of two summers we sampled seven eddies in the Sargasso Sea. We focused on and followed a post-phytoplankton bloom cyclonic eddy (C1) in 2004 and a blooming mode-water anticyclonic eddy (A4) in 2005. We collected zooplankton in all eddies using a Multiple Opening and Closing Net Environmental Sampling System (MOCNESS) and quantified biomass (>0.15 mm, in five size fractions) from 0 to 700 m over nine discrete depth intervals. Zooplankton biomass (>0.5 mm) in the upper 150 m was similarly enhanced at night for the periphery of C1 and the center of A4 at 0.514 g m⁻² and 0.533 g m⁻², respectively, compared to outside (0.183 g m⁻² outside C1 and 0.197 g m⁻² outside A4). Despite minimal chlorophyll a enhancement and dominance by picoplankton in C1, zooplankton biomass increased most for the largest size class (>5 mm). Gut fluorescence for euphausiids and large copepods was also elevated on the C1 periphery. In A4, peak biomass occurred at eddy center coincident with peak primary production, but was highly variable (changing by >3-fold) over time, perhaps resulting from the dense, but patchy distribution of diatom chains in this region. Shifts in zooplankton community composition and abundance were reflected in enhancement of fecal pellet production and active transport by diel vertical migration in eddies. Inside C1 the flux of zooplankton fecal pellets at 150 m in June 2004 was 1.5-fold higher than outside the eddy, accounting for 9% of total particulate organic carbon (POC) flux. The flux of fecal pellets (mostly from copepods) increased through the summer in eddy A4, matching concurrent increases in zooplankton <2 mm in length, and accounting for up to 12% of total POC flux. Active carbon transport by vertically migrating zooplankton was 37% higher on the periphery of C1 and 74% higher at the center of A4 compared to the summer mean at the Bermuda Atlantic Time-series Study (BATS) station. Despite contrasting responses by the phytoplankton community to cyclonic and mode-water eddies, mesozooplankton biomass was similarly enhanced, possibly due to differential physical and biological aggregation mechanisms, and resulted in important zooplankton-mediated changes in mesoscale biogeochemistry.     

Surface Waters of the NW Iberian Margin: Upwelling on the Shelf versus Outwelling of Upwelled Waters from the Rías Baixas

A set of hydrographic surveys were carried out in the Ría of Vigo (NW Spain) at 2–4 d intervals during four 2–3 week periods in 1997, covering contrasting seasons. Residual exchange fluxes with the adjacent shelf were estimated with a 2-D, non-steady-state, salinity–temperature weighted box model. Exchange fluxes consist of a steady-state term (dependent on the variability of continental runoff) and a non-steady-state term (dependent on the time changes of density gradients in the embayment). More than 95% of the short-time-scale variability of the exchange fluxes in the middle and outer ría can be explained by the non-steady-state term that, in turns, is correlated (R²>75%) with the offshore Ekman transport. Conversely, 96% of the variability of exchange fluxes in the inner ría rely on the steady-state term. The outer and middle ría are under the direct influence of coastal upwelling, which enhances the positive residual circulation pattern by an order of magnitude: from 10²to 10³ m³s⁻¹. On the contrary, downwelling provokes a reversal of the circulation in the outer ría. The position of the downwelling front along the embayment depends on the relative importance of Ekman transport (Q_x, m³s⁻¹km⁻¹) and continental runoff (R, m³s⁻¹). When Q_x/ R>7±2 the reversal of the circulation affects the middle ría. Our results are representative for the ‘Rías Baixas’, four large coastal indentations in NW Spain. During the upwelling season (spring and summer), 60% of shelf surface waters off the ‘Rías Baixas’ consist of fresh Eastern North Atlantic Central Water (ENACW) upwelled in situ. The remaining 40% consists of upwelled ENACW that previously enters the rías and it is subsequently outwelled after thermohaline modification. During the downwelling season (autumn and winter), 40% of the warm and salty oceanic subtropic surface water, which piled on the shelf by the predominant southerly winds, enters the rías.     

Nutrient irrigation of the North Atlantic

The North Atlantic, as all major oceans, has a remarkable duality in primary production, manifested by the existence of well-defined high and low mean primary production regions. The largest region is the North Atlantic Subtropical Gyre (NASTG), an anticyclone characterized by bowl shaped isopycnals and low production. The NASTG is surrounded at its margins by smaller cyclonic high-production regions, where these isopycnals approach the sea surface. The most extensive cyclonic regions are those at the latitudinal extremes, i.e. the subpolar and tropical oceans, though smaller ones do occur at the zonal boundaries. In this article we review historical data and present new analyses of climatological data and a selected number of hydrographic cruises in the western/northwestern and eastern/southeastern boundaries of the NASTG, with the objective of investigating the importance of upward epipycnal advection of nutrient-rich subsurface layers (irrigation) in maintaining high primary production in the euphotic layers. In the North Atlantic Subpolar Gyre (NASPG) irrigation implies intergyre exchange caused by the outcropping extension of the Gulf Stream (GS), following the formation of the deep winter mixed-layer. In the eastern boundary of the NASTG irrigation is attained through a permanent upwelling cell, which feeds the Canary Upwelling Current (CUC). In the southeastern corner irrigation occurs in fall, when the Guinea Dome (GD) is reinforced, and in winter, when the CUC reaches its southernmost extension. Other characteristics of the north/south extension of the GS/CUC are the seasonal nutrient replenishing of subsurface layers (spring restratification of NASPG and winter relaxation of the GD) and the maintenance of high levels of diapycnal mixing during the last phase of nutrient transfer to the euphotic layers. Off the Mid-Atlantic Bight the GS transports a total of about 700 kmol s⁻¹ of nitrate, with almost 100 kmol s⁻¹ carried in the surface (σ_θ < 26.8) layers and some 350 kmol s⁻¹ in the intermediate (26.8 < σ_θ < 27.5) layers. A box model suggests that north of Cape Hatteras most surface and upper-thermocline nitrates are used to sustain the high levels of primary production in the NASPG. Off Cape Blanc there is winter along-shore convergence of order 10 kmol s⁻¹ of nitrate in the near-surface layers (possibly larger in summer), with only a small fraction used to sustain local primary production in the coastal upwelling band and the remainder carried to the interior ocean. Nutrients and biomass exported from these cyclonic regions may account for the concentration levels observed within the NASTG.     

Nutrient flux into an intense deep chlorophyll layer in a mode-water eddy

An intense deep chlorophyll layer in the Sargasso Sea was reported near the center of an anticyclonic mode-water eddy by McGillicuddy et al. [2007. Eddy–wind interactions stimulate extraordinary mid-ocean plankton blooms, Science, accepted]. The high chlorophyll was associated with anomalously high concentrations of diatoms and with a maximum in the vertical profile of ¹⁴C primary productivity. Here we report tracer measurements of the vertical advection and turbulent diffusion of deep-water nutrients into this chlorophyll layer. Tracer released in the chlorophyll layer revealed upward motion relative to isopycnal surfaces of about 0.4 m/d, due to solar heating and mixing. The density surfaces themselves shoaled by about 0.1 m/d. The upward flux of dissolved inorganic nitrogen, averaged over 36 days, was approximately 0.6 mmol/m²/d due to both upwelling and mixing. This flux is about 40% of the basin wide, annually averaged, nitrogen flux required to drive the annual new production in the Sargasso Sea, estimated from the oxygen cycle in the euphotic zone, the oxygen demand below the euphotic zone, and from the ³He excess in the mixed layer. The observed upwelling of the fluid was consistent with theoretical models [Dewar, W.K., Flierl, G.R., 1987. Some effects of wind on rings. Journal of Physical Oceanography 17, 1653–1667; Martin, A.P., Richards, K.J., 2001. Mechanisms for vertical nutrient transport within a North Atlantic mesoscale eddy. Deep-Sea Research II 48, 757–773] in which eddy surface currents cause spatial variations in surface stress. The diapycnal diffusivity at the base of the euphotic zone was 3.5±0.5×10⁻⁵ m²/s. Diapycnal mixing was probably enhanced over more typical values by the series of storms passing over the eddy during the experiment and may have been enhanced further by the trapping of near-inertial waves generated within the eddy.     

Remotely sensed sea-surface chlorophyll and POC flux at Deep Gulf of Mexico Benthos sampling stations

Biweekly composite averages of the standing stock of sea-surface chlorophyll (SSC) were derived from SeaWiFS satellite ocean-color data at 44 benthic sampling stations occupied along the continental slope and rise by the Deep Gulf of Mexico Benthos (DGoMB) program. At the 22 DGoMB sites north of 26°N and west of 91°W in the NW Gulf of Mexico, annual average SSC was 0.19 mg m⁻³, ranging at most locations from annual highs of about 0.3 mg m⁻³ in November–February to lows of about 0.1 mg m⁻³ in May–August. Comparison of three years of SeaWiFS data (January 1998–December 2000) showed little inter-annual variation at these NW Gulf stations. In contrast, at the 22 NE Gulf sites north of 26°N and east of 91°W, SSC averaged 2.8 times higher than in the NW Gulf, showing also strong inter-annual variation. Maxima in the NE region occurred in November–February and also during summers. The summer maxima were associated with Mississippi River water transported offshore to the east and southward by anticyclonic eddies in the NE Gulf. The apparent increases in SSC in June–August at NE Gulf stations reached average monthly concentrations >50% greater than in November–February. Based on a primary productivity model and a vertical flux model, the calculated export of particulate organic carbon (POC flux reaching the seafloor) was estimated as 18 mg C m⁻² day⁻¹ at the 22 NE Gulf stations, and 9 mg C m⁻² day⁻¹ at the 22 NW Gulf stations. These estimates are comparable to fluxes measured by benthic lander by others in the DGoMB program, which may drive the differences in west versus east bathymetric zonation and community structure of macrobenthos that were sampled with large box corers by others in the DGoMB program.     

Inorganic carbon in the central California upwelling system during the 1997–1999 El Niño–La Niña event

Sea surface pCO₂ was monitored during 49 cruises from February 1997 to December 1999 along a section perpendicular to the central California Coast. Continuous measurements of the ocean–atmosphere difference of pCO₂ were made on a mooring in the same region from July 1997 to December 1999. The El Niño/La Niña cycle of 1997–1999 had a significant influence on local ocean–atmosphere CO₂ transfer. During the warm anomaly associated with El Niño, upwelling was suppressed and average sea surface pCO₂ was below atmospheric level. High rainfall and river runoff in the late winter and early spring of 1998 produced areas where pCO₂ was depressed by as much as 100 μatm. A flux ranging from 0.3 to 0.7 mol C m⁻² y⁻¹ from the atmosphere into the ocean was estimated for the El Niño period from wind and ΔpCO₂ data. Temperatures and upwelling returned to near normal in the summer of 1998, but a cold anomaly developed during autumn of that year. Temperature and pCO₂ data indicate that upwelling continued throughout much of the 1998–1999 winter and intensified significantly in the spring of 1999. During strong upwelling events, the estimate of ocean to atmosphere flux approached rates of 50 mol C m⁻² y⁻¹. The estimate for the average CO₂ flux from July 1998 to July 1999 was 1.5–2.2 mol C m⁻² y⁻¹ from the ocean to the atmosphere. While the flux estimate for the El Niño time period may be applicable to a larger area, the high ocean to atmosphere fluxes during La Niña might be the result of sampling near a zone of intense upwelling.     

Particle fluxes associated with mesoscale eddies in the Sargasso Sea

We examined the impact of a cyclonic eddy and mode-water eddy on particle flux in the Sargasso Sea. The primary method used to quantify flux was based on measurements of the natural radionuclide, ²³⁴Th, and these flux estimates were compared to results from sediment traps in both eddies, and a ²¹⁰Po/²¹⁰Pb flux method in the mode-water eddy. Particulate organic carbon (POC) fluxes at 150 m ranged 1–4 mmol C m⁻² d⁻¹ and were comparable between methods, especially considering differences in integration times scales of each approach. Our main conclusion is that relative to summer mean conditions at the Bermuda Atlantic Time-series Study (BATS) site, eddy-driven changes in biogeochemistry did not enhance local POC fluxes during this later, more mature stage of the eddy life cycle (>6 months old). The absence of an enhancement in POC flux puts a constraint on the timing of higher POC flux events, which are thought to have caused the local O₂ minima below each eddy, and must have taken place >2 months prior to our arrival. The mode-water eddy did enhance preferentially diatom biomass in its center, where we estimated a factor of three times higher biogenic Si flux than the BATS summer average. An unexpected finding in the highly depth-resolved ²³⁴Th data sets is narrow layers of particle export and remineralization within the eddy. In particular, a strong excess ²³⁴Th signal is seen below the deep chlorophyll maxima, which we attribute to remineralization of ²³⁴Th-bearing particles. At this depth below the euphotic zone, de novo particle production in the euphotic zone has stopped, yet particle remineralization continues via consumption of labile sinking material by bacteria and/or zooplankton. These data suggest that further study of processes in ocean layers is warranted not only within, but below the euphotic zone.     

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.     

The herbivorous impact of microzooplankton during two short-term Lagrangian experiments off the NW coast of Galicia in summer 1998

Microzooplankton (heterotrophic microplankton and heterotrophic nanoflagellates) and their herbivorous activity were estimated from dilution experiments in August 1998 during two Lagrangian drift experiments that sampled contrasting conditions—an upwelling/relaxation event along the shelf edge and an oligotrophic offshore filament. During upwelling/relaxation, heterotrophic microplankton were present at mean surface concentrations between 15,000 and 48,000 cells l⁻¹. Heterotrophic nanoflagellate concentrations were between 200 and 700 cells ml⁻¹ and the most abundant component of the heterotrophic microplankton was the aloricate choreotrich ciliates which increased dramatically in concentration from 6,000 to 24,000 cells l⁻¹ during the first 4 days of the study. Total microzooplankton biomass reached a maximum of 39mgC.m⁻³. In the filament, which developed from the upwelling, cell concentrations were lower and averaged 4,500 cells l⁻¹ for heterotrophic microplankton and 250 cells ml⁻¹ for heterotrophic nanoflagellates. Total microzooplankton biomass was about 10–12mgC.m⁻³. Microzooplankton turned over between 40 and 85% of the phytoplankton standing stock, thereby consuming between 5 and 78mg phytoplankton carbon.m⁻³.d⁻¹. The magnitude of this activity was highest during upwelling/relaxation and was positively correlated to heterotrophic nanoflagellate biomass and chlorophyll-a concentration but not heterotrophic microplankton biomass. The proportion of primary production grazed decreased from 160 to 59% d⁻¹ during upwelling/relaxation and ranged between 60 and 90% d⁻¹ in the filament. Microzooplankton herbivory within the euphotic zone increased from 684 to >2000mgC.m⁻².d⁻¹ during upwelling/relaxation and was between 327 and 802mgC.m⁻².d⁻¹ in the filament. Although microzooplankton herbivory was lower and less variable during the filament study, microzooplankton consumed on average 60% of the phytoplankton standing stocks which was higher than found during upwelling/relaxation. Microzooplankton assimilation efficiency ranged between 3 and 33% during upwelling/relaxation and between 0 and 13% in the filament. Our data demonstrate a close coupling between phytoplankton growth and microzooplankton herbivory in surface waters off the Galician Coast and suggest that microzooplankton may have been a significant sink for phytogenic carbon during August 1998.