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.     

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.     

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).     

Cobalt, manganese, and iron near the Hawaiian Islands: A potential concentrating mechanism for cobalt within a cyclonic eddy and implications for the hybrid-type trace metals

The vertical distributions of cobalt, iron, and manganese in the water column were studied during the E-Flux Program (E-Flux II and III), which focused on the biogeochemistry of cold-core cyclonic eddies that form in the lee of the Hawaiian Islands. During E-Flux II (January 2005) and E-Flux III (March 2005), 17 stations were sampled for cobalt (n=147), all of which demonstrated nutrient-like depletion in surface waters. During E-Flux III, two depth profiles collected from within a mesoscale cold-core eddy, Cyclone Opal, revealed small distinct maxima in cobalt at 100 m depth and a larger inventory of cobalt within the eddy. We hypothesize that this was due to a cobalt concentrating effect within the eddy, where upwelled cobalt was subsequently associated with sinking particulate organic carbon (POC) via biological activity and was released at a depth coincident with nearly complete POC remineralization [Benitez-Nelson, C., Bidigare, R.R., Dickey, T.D., Landry, M.R., Leonard, C.L., Brown, S.L., Nencioli, F., Rii, Y.M., Maiti, K., Becker, J.W., Bibby, T.S., Black, W., Cai, W.J., Carlson, C.A., Chen, F., Kuwahara, V.S., Mahaffey, C., McAndrew, P.M., Quay, P.D., Rappe, M.S., Selph, K.E., Simmons, M.P., Yang, E.J., 2007. Mesoscale eddies drive increased silica export in the subtropical Pacific Ocean. Science 316, 1017–1020]. There is also evidence for the formation of a correlation between cobalt and soluble reactive phosphorus during E-Flux III relative to the E-Flux II cruise that we suggest is due to increased productivity, implying a minimum threshold of primary production below which cobalt–phosphate coupling does not occur. Dissolved iron was measured in E-Flux II and found in somewhat elevated concentrations (0.5 nM) in surface waters relative to the iron depleted waters of the surrounding Pacific [Fitzwater, S.E., Coale, K.H., Gordon, M.R., Johnson, K.S., Ondrusek, M.E., 1996. Iron deficiency and phytoplankton growth in the equatorial Pacific. Deep-Sea Research II 43 (4–6), 995–1015], possibly due to island effects associated with the iron-rich volcanic soil from the Hawaiian Islands and/or anthropogenic inputs. Distinct depth maxima in total dissolved cobalt were observed at 400–600 m depth, suggestive of the release of metals from the shelf area of comparable depth that surrounds these islands.     

Production and respiration control the marine microbial metabolic balance in the eastern North Atlantic subtropical gyre

Two main contrasted hypotheses have arisen during the last decades about the factors controlling the planktonic net metabolic balance in oligotrophic waters: gross primary production controls net community production vs. variability of net community production is also influenced by changes in microbial respiration. This work discusses both hypotheses analyzing the variability of metabolic rates along a gradient from the margin to the centre of the North Atlantic oligotrophic gyre, i.e. from relatively productive to more oligotrophic conditions. Net community production (NCP) was close to zero (between −3.34 and −11.77 mmol O₂ m⁻² d⁻¹) at the margin of the gyre and tended towards net heterotrophy (−44.03 mmol O₂ m⁻² d⁻¹) to the centre of the gyre as both gross primary production (GPP) and community respiration (CR) decreased. The strong relationships found between nutrient availability and both NCP and GPP suggest that factors controlling GPP are prevalent in determining NCP variability in this biogeographic region. However implementation of existing models to predict NCP from the measured GPP indicates that the precise estimation of NCP in different oligotrophic systems requires consideration of the magnitude and variability of microbial respiration rates.     

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.     

Primary production and nutrients in a tropical macrotidal estuary, Darwin Harbour, Australia

Tropical estuaries are under increasing pressure worldwide from human impacts, but are poorly studied compared with temperate systems. This study examined a tropical macrotidal estuary, Darwin Harbour, in northern Australia, using a combination of direct measurements and literature values to determine the main sources of primary production and the sources of nutrients supporting growth. The main source of primary production was calculated to be the extensive area of fringing mangroves and resulted in a net autotrophic system (P_G:R = 2.1). Much of the carbon in the mangrove forests appears to be retained within the forests or respired, as the water column was also net autotrophic despite the carbon inputs. Phytoplankton were the second largest primary producer on a whole-of-harbour basis, with low biomass constrained by light and nutrient availability. The phytoplankton were likely to be nitrogen (N) limited, based on low N:phosphorus (P) ratios, low dissolved bioavailable N concentrations (ammonium (NH₄⁺), nitrate (NO₃⁻), urea), and evidence that phytoplankton growth in bioassays was stimulated by NH₄⁺ addition. The largest new source of N to the system was from the ocean due to higher N concentrations in the incoming tides than the outgoing tides. Atmospheric inputs via N fixation on the intertidal mudflats and subtidal sediments were substantially lower. The rivers feeding into the harbour and sewage were minor N inputs. Nitrogen demand by primary producers was high relative to available N inputs, suggesting that N recycling within the water column and mangrove forests must be important processes. Darwin Harbour is adjacent to the rapidly growing urban area of Darwin city, but overall there is no evidence of anthropogenic nutrient inputs having substantial effects on primary production in Darwin Harbour.     

Winter and spring phytoplankton composition and production in a shallow eutrophic Baltic lagoon

Taxonomic composition and productivity of winter and spring phytoplankton in a eutrophic estuary have been investigated in order to elucidate the carbon flux under conditions of limitation by physical factors – light and temperature. In spite of the important differences in nutrients, solar radiation and water temperature between winter and spring season, mean concentrations of particulate organic carbon were equal to 13.2 and 13.0 mgC l⁻¹, respectively. Chlorophyll a averaged at 79 μgChl l⁻¹ in winter, that is 69% of spring. Although community respiration accounted for only 6–26% of light saturated photosynthesis, integrated net primary production of the 1.2 m deep water column was negative until April. High attenuation of the water body (K_o = 2.9 m⁻¹) lead to a negative carbon balance (net heterotrophy) below 35 cm for all sampling dates. Thus, the high winter POC and phytoplankton values can only originate from summer or autumn primary production. This assumption was supported by a carbon loss rate of just 3% of total organic carbon per day for the whole water column. The composition of phytoplankton was very constant through both seasons: 39% Chlorophyceae, 33% Cyanobacteria and 25% Bacillariophyceae. As expected, phytoplankton was low light acclimated, having high α values (slope of light limited photosynthesis), but moderate maximum photosynthesis rates at saturating irradiances, which were heavily affected by temperature. Calculation of net carbon flux yet showed net heterotrophy of the Bodden waters in winter and early spring were caused by external physical limitation (low surface irradiance and low temperature) in combination with a high light attenuation of the water body.     

Temporal variation in the structure of autotrophic and heterotrophic communities in the subarctic Pacific

Autotrophic and microheterotrophic plankton populations were monitored in the euphotic zone of the eastern subarctic Pacific during 6 one-month cruises in spring and summer, 1984, 1987 and 1988. Transmitted light, epifluorescence, and electron microscopy were used to identify, enumerate and estimate the biomass of size-populations of species. The 2–10μm size class dominated the biomass of both autotrophs and heterotrophs. The autotrophic flagellate, Phaeocystis pouchetii, was frequently observed in its non-colonial phase. Temporal variation in all the stocks was evident and could be explained only partially by the physical, chemical or biological factors investigated here. The general structure of the autotrophic community was similar to that in the North Atlantic, but major, unexplained variations between cruises occurred. Variation in mixed-layer depth and day length (but not variation in daily insolation) explained 25% of the variation in autotrophic doubling rate. Heterotrophic biomass comprised, in decreasing order of importance, non-pigmented flagellates, dinoflagellates, and ciliates. Ciliates rarely contributed more than 40% to the total. Microheterotrophic biomass rarely exceeded 30μg C 1⁻¹ (avg 15μg C 1⁻¹, 0–60m) whereas autotrophic biomass averaged 20μg C 1⁻¹, 0–60m, and reached 74μg C 1⁻¹ on one occasion, yet the grazing capacity of these microheterotrophs averaged 100% of primary production.     

Influence of basin-scale and mesoscale physical processes on biological productivity in the Bay of Bengal during the summer monsoon

Physical forcing plays a major role in determining biological processes in the ocean across the full spectrum of spatial and temporal scales. Variability of biological production in the Bay of Bengal (BoB) based on basin-scale and mesoscale physical processes is presented using hydrographic data collected during the peak summer monsoon in July–August, 2003. Three different and spatially varying physical processes were identified in the upper 300 m: (I) anticyclonic warm gyre offshore in the southern Bay; (II) a cyclonic eddy in the northern Bay; and (III) an upwelling region adjacent to the southern coast. In the warm gyre (>28.8 °C), the low salinity (33.5) surface waters contained low concentrations of nutrients. These warm surface waters extended below the euphotic zone, which resulted in an oligotrophic environment with low surface chlorophyll a (0.12 mg m⁻³), low surface primary production (2.55 mg C m⁻³ day⁻¹) and low zooplankton biovolume (0.14 ml m⁻³). In the cyclonic eddy, the elevated isopycnals raised the nutricline upto the surface (NO₃–N > 8.2 μM, PO₄–P > 0.8 μM, SiO₄–Si > 3.5 μM). Despite the system being highly eutrophic, response in the biological activity was low. In the upwelling zone, although the nutrient concentrations were lower compared to the cyclonic eddy, the surface phytoplankton biomass and production were high (Chl a – 0.25 mg m⁻³, PP – 9.23 mg C m⁻³ day⁻¹), and mesozooplankton biovolume (1.12 ml m⁻³) was rich. Normally in oligotrophic, open ocean ecosystems, primary production is based on ‘regenerated’ nutrients, but during episodic events like eddies the ‘production’ switches over to ‘new production’. The switching over from ‘regenerated production’ to ‘new production’ in the open ocean (cyclonic eddy) and establishment of a new phytoplankton community will take longer than in the coastal system (upwelling). Despite the functioning of a cyclonic eddy and upwelling being divergent (transporting of nutrients from deeper waters to surface), the utilization of nutrients leading to enhanced biological production and its transfer to upper trophic levels in the upwelling region imply that the energy transfer from primary production to secondary production (mesozooplankton) is more efficient than in the cyclonic eddy of the open ocean. The results suggest that basin-scale and mesoscale processes influence the abundance and spatial heterogeneity of plankton populations across a wide spatial scale in the BoB. The multifaceted effects of these physical processes on primary productivity thus play a prominent role in structuring of zooplankton communities and could consecutively affect the recruitment of pelagic fisheries.     

Net biological oxygen production in the ocean—II: Remote in situ measurements of O2 and N2 in subarctic pacific surface waters

Net community biological production in the euphotic zone of the ocean fuels organic matter and oxygen export from the upper ocean, which has a large influence on the atmospheric pressure of carbon dioxide and is the driving force for metabolite distributions in the sea. We determine the net annual biological oxygen production in the mixed layer of the northeast subarctic Pacific Ocean from in situ O₂ and N₂ measurements. Temperature, salinity, total gas pressure and O₂ were measured every 3 h for 9 months in 2007 at about 3 m depth on a surface mooring at Station P (50°N, 145°W). The concentration of nitrogen gas, N₂, determined from separate total gas pressure and pO₂ measurements, was used as an inert tracer of the physical processes that induce gas departure from thermodynamic equilibrium with the atmosphere. We use a simple model of the ocean’s mixed layer along with the nitrogen concentration to constrain the importance of bubbles, gas exchange and horizontal advection, which are then used in the oxygen mass balance to derive net biological oxygen production. The mixed-layer oxygen mass balance is dominated by exchange with the atmosphere, and we determine a mean summertime oxygen production of 24 mmol O₂ m^?2 d^?1. The annual pattern in the difference between the supersaturation of oxygen and nitrogen in the surface waters reveals very little net oxygen production during the winter at this location. The calculated annual net community production (NCP) of carbon from this new method, 2.5 mol m^?2 yr^?1, agrees to within its error of about×40% with previous determinations at this location from oxygen mass balance, NO₃^? draw down and ²³⁴Th measurements. This value is either indistinguishable from or lower than annual NCP measurements in the subtropical North Pacific, indicating that there is no experimental evidence for differences in annual NCP between the subarctic and subtropical North Pacific Ocean.     

Diatoms in the desert: Plankton community response to a mesoscale eddy in the subtropical North Pacific

As part of the E-Flux project, we documented spatial variability and temporal changes in plankton community structure in a cold-core cyclonic eddy in the lee of the Hawaiian Islands. Cyclone Opal spanned 200 km in diameter, with sharply uplifted isopycnals (80–100 m relative to surrounding waters) and a strongly expressed deep chlorophyll a maximum (DCM) in its central core region of 40 km diameter. Microscopic and flow cytometric analyses of samples from across the eddy revealed dramatic transitions in phytoplankton community structure, reflecting Opal's well-developed physical structure. Upper mixed-layer populations in the eddy resembled those outside the eddy and were dominated by picophytoplankton. In contrast, the DCM was composed of large chain-forming diatoms dominated by Chaetoceros and Rhizosolenia spp. Diatoms attained unprecedented levels of biomass (nearly 90 μg C l⁻¹) in the center of the eddy, accounting for 85% of photosynthetic biomass. Protozoan grazers displayed two- to three-fold higher biomass levels in the eddy center as well. We also found a distinct and persistent layer of senescent diatom cells overlying healthy populations, often separated by less than 10 m, indicating that we were sampling a bloom in a state of decline. Time-series sampling over 8 days showed a successional shift in community structure within the central diatom bloom, from the unexpected large chain-forming species to smaller forms more typical of the subtropical North Pacific. The diatom bloom of Cyclone Opal was a unique, and possibly extreme, example of biological response to physical forcing in the North Pacific subtropical gyre, and its detailed study may therefore help to improve our predictive understanding of environmental controls on plankton community structure.     

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.     

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.     

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.     

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.     

Improving estuarine net flux estimates for dissolved cadmium export at the annual timescale: Application to the Gironde Estuary

Dissolved Cd (Cd_D) concentrations along the salinity gradient were measured in surface water of the Gironde Estuary during 15 cruises (2001–2007), covering a wide range of contrasting situations in terms of hydrology, turbidity and season. During all situations dissolved Cd concentrations displayed maximum values in the mid-salinity range, reflecting Cd addition by chloride-induced desorption and complexation. The daily net Cd_D fluxes from the Gironde Estuary to the coastal ocean were estimated using Boyle's method. Extrapolating Cd_D concentrations in the high salinity range to the freshwater end member using a theoretical dilution line produced 15 theoretical Cd concentrations (Cd_D⁰), each representative of one distinct situation. The obtained Cd_D⁰ concentrations were relatively similar (201 ± 28 ng L⁻¹) when freshwater discharge Q was >500 m³ s⁻¹ (508 ≤ Q ≤ 2600 m³ s⁻¹), but were highly variable (340 ± 80 ng L⁻¹; 247–490 ng L⁻¹) for low discharge situations (169 ≤ Q ≤ 368 m³ s⁻¹). The respective daily Cd_D net fluxes were 5–39 kg day⁻¹, mainly depending on freshwater discharge. As this observation invalidates the existing method of estimating annual Cd_D net fluxes, we proposed an empirical model, using representative Cd_D⁰ values and daily freshwater discharges for the 2001–2007 period. Subsequent integration produced reliable Cd_D net flux estimates for the Gironde Estuary at the annual timescale that ranged between 3.8–5.0 t a⁻¹ in 2005 and 6.0–7.2 t a⁻¹ in 2004, depending on freshwater discharge. Comparing Cd_D net fluxes with the incoming Cd_D fluxes suggested that the annual net Cd_D addition in the Gironde Estuary ranged from 3.5 to 6.7 t a⁻¹, without any clear temporal trend during the past seven years. The annual Cd_D net fluxes did not show a clearly decreasing trend in spite of an overall decrease by a factor 6 in Cd gross fluxes during the past decade. Furthermore, in six years out of seven (except 2003), the annual Cd_D net fluxes even exceeded river borne total (dissolved + particulate) gross Cd fluxes into the estuary. These observations were attributed to progressive Cd desorption from both suspended particles and bottom sediment during various sedimentation–resuspension cycles induced by tidal currents and/or continuous dredging (navigation channel) and diverse intra-estuarine sources (wet deposition, urban sources, and agriculture). Provided that gross fluxes remain stable over time, dissolved Cd exportation from the Gironde Estuary to the coastal ocean may remain at the present level for the coming decade and the estuarine sedimentary Cd stock is forecast to decrease slowly.     

Diapycnal mixing by meso-scale eddies

The mean available potential energy released by baroclinic instability into the meso-scale eddy field has to be dissipated in some way and Tandon and Garrett [Tandon, A., Garrett, C., 1996. On a recent parameterization of mesoscale eddies. J. Phys. Oceanogr. 26 (3), 406–416] suggested that this dissipation could ultimately involve irreversible mixing of buoyancy by molecular processes at the small-scale end of the turbulence cascade. We revisit this idea and argue that the presence of dissipation within the thermocline automatically requires that a component of the eddy flux associated with meso-scale eddies must be associated with irreversible mixing of buoyancy within the thermocline. We offer a parameterisation of the implied diapycnal diffusivity based on (i) the dissipation rate for eddy kinetic energy given by the meso-scale eddy closure of Eden and Greatbatch [Eden, C., Greatbatch, R.J., 2008. Towards a meso-scale eddy closure. Ocean Modell. 20, 223–239.] and (ii) a fixed mixing efficiency. The implied eddy-induced diapycnal diffusivity (κ) is implemented in a coarse resolution model of the North Atlantic. In contrast to the vertical diffusivity given by a standard vertical mixing scheme, large lateral inhomogeneities can be found for κ in the interior of the ocean. In general, κ is large, i.e. up to o(10) cm²/s, near the western boundaries and almost vanishing in the interior of the ocean.     

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.     

Organic carbon and iron modulate nitrification rates in mangrove swamps of Goa, south west coast of India

Nitrification, fuelled by ammonium is the pivotal oxidative pathway to nitrogen cycling. In spite of its ecological significance, the factors regulating nitrification rates in the benthic realm remain poorly understood. The present study therefore examines some of the factors like ammonium, nitrite, nitrate, organic carbon, iron and manganese on down-core variability in benthic nitrification rates in two different mangrove ecosystems, one under the influence of ferromanganese ore mining (experiment) and the other relatively undisturbed (control). We hypothesize that besides organic carbon, iron could also influence the rate of nitrification. The study also contrasts the distributive pattern of autotrophic and heterotrophic nitrifiers in the two regions. The concentration of iron at the control site ranged from 1.1% to 15.1% while at the experimental site it ranged from 2.9% to 46%. The levels of organic carbon at control and experimental sites ranged from 0.02% to 6.9% and 0.1% to 6.5%, respectively. The nitrification rates at the control and experimental sites are comparable and ranged from 3.2 ± 1.2 to 18.4 ± 1.9 ng at-N g(sediment)⁻¹ h⁻¹ and 2.7 ± 1.5 to 18.2 ± 0.6 ng at-N g(sediment)⁻¹ h⁻¹, respectively. While the abundance of heterotrophic nitrifiers at both the sites ranged from 10^2–3 cells g⁻¹ sediment, the autotrophic nitrifiers at the experimental site was higher by an order at 10³ cells g⁻¹ sediment reflecting the relatively higher refractile nature of organic carbon at the experimental site (Straus and Lamberti, 2000). Though organic carbon and nitrification rates are similar in both the sites, the underlying mechanisms governing the processes could be different. Our studies suggest that at the control site, heterotrophic nitrifiers govern nitrification rates (r = 0.28, p < 0.05, n = 64) using organic carbon (r = 0.32, p < 0.01, n = 64). At the experimental site, nitrification was governed more by autotrophic nitrifiers (r = 0.43, p < 0.001, n = 64) at the expense of iron (r = 0.47, p < 0.001, n = 64). Therefore at the experimental site with higher load of iron, autotrophic nitrification could be more important. It is therefore inferred that both the quality of organic carbon and quantity of iron govern nitrification rates in these mangrove swamps.