Interhemispheric exchanges of mass and heat in the Atlantic Ocean in January–March 1993

Hydrographic, geochemical, and direct velocity measurements along two zonal (7.5°N and 4.5°S) and two meridional (35°W and 4°W) lines occupied in January–March, 1993 in the Atlantic are combined in an inverse model to estimate the circulation. At 4.5°S, the Warm Water (potential temperature θ>4.5°C) originating from the South Atlantic enters the equatorial Atlantic, principally at the western boundary, in the thermocline-intensified North Brazil Undercurrent (33±2.7×10⁶ m³ s⁻¹ northward) and in the surface-intensified South Equatorial Current (8×10⁶ m³ s⁻¹ northward) located to the east of the North Brazil Undercurrent. The Ekman transport at 4.5°S is southward (10.7±1.5×10⁶ m³ s⁻¹). At 7.5°N, the Western Boundary Current (WBC) (17.9±2×10⁶ m³ s⁻¹) is weaker than at 4.5°S, and the northward flow of Warm Water in the WBC is complemented by the basin-wide Ekman flow (12.3±1.0×10⁶ m³ s⁻¹), the net contribution of the geostrophic interior flow of Warm Water being southward. The equatorial Ekman divergence drives a conversion of Thermocline Water (24.58⩽σ₀<26.75) into Surface Water (σ₀<24.58) of 7.5±0.5×10⁶ m³ s⁻¹, mostly occurring west of 35°W. The Deep Water of northern origin flows southward at 7.5°N in an energetic (48±3×10⁶ m³ s⁻¹) Deep Western Boundary Current (DWBC), whose transport is in part compensated by a northward recirculation (21±4.5×10⁶ m³ s⁻¹) in the Guiana Basin. At 4.5°S, the DWBC is much less energetic (27±7×10⁶ m³ s⁻¹ southward) than at 7.5°N. It is in part balanced by a deep northward recirculation east of which alternate circulation patterns suggest the existence of an anticyclonic gyre in the central Brazil Basin and a cyclonic gyre further east. The deep equatorial Atlantic is characterized by a convergence of Lower Deep Water (45.90⩽σ₄<45.83), which creates an upward diapycnal transport of 11.0×10⁶ m³ s⁻¹ across σ₄=45.83. The amplitude of this diapycnal transport is quite sensitive to the a priori hypotheses made in the inverse model. The amplitude of the meridional overturning cell is estimated to be 22×10⁶ m³ s⁻¹ at 7.5°N and 24×10⁶ m³ s⁻¹ at 4.5°S. Northward heat transports are in the range 1.26–1.50 PW at 7.5°N and 0.97–1.29 PW at 4.5°S with best estimates of 1.35 and 1.09 PW.     

Diapycnal nutrient fluxes on the northern boundary of Cape Ghir upwelling region

In this study we estimate diffusive nutrient fluxes in the northern region of Cape Ghir upwelling system (Northwest Africa) during autumn 2010. The contribution of two co-existing vertical mixing processes (turbulence and salt fingers) is estimated through micro- and fine-structure scale observations. The boundary between coastal upwelling and open ocean waters becomes apparent when nitrate is used as a tracer. Below the mixed layer (56.15±15.56 m), the water column is favorable to the occurrence of a salt finger regime. Vertical eddy diffusivity for salt (K_s) at the reference layer (57.86±8.51 m, CI 95%) was 3×10⁻⁵ (±1.89×10⁻⁹, CI 95%) m² s⁻¹. Average diapycnal fluxes indicate that there was a deficit in phosphate supply to the surface layer (6.61×10⁻⁴ mmol m⁻² d⁻¹), while these fluxes were 0.09 and 0.03 mmol m⁻² d⁻¹ for nitrate and silicate, respectively. There is a need to conduct more studies to obtain accurate estimations of vertical eddy diffusivity and nutrient supply in complex transitional zones, like Cape Ghir. This will provide us with information about salt and nutrients exchange in onshore–offshore zones.     

The thermohaline structure and evolution of the deep waters in the Canada Basin,Arctic Ocean

Below the sill depth (at about 2400 m) of the Alpha-Mendeleyev ridge complex, the waters of the Canada Basin (CB) of the Arctic Ocean are isolated, with a ¹⁴C isolation age of about 500 yr. The potential temperature θ decreases with depth to a minimum θ_m≈−0.524°C near 2400 m, increases with depth through an approximately 300 m thick transition layer to θ_h≈−0.514°C, and then remains uniform from about 2700 m to the bottom at 3200–4000 m. The salinity increases monotonically with depth through the deep θ_m and transition layer from about 34.952 to about 34.956 and then remains uniform in the bottom layer. A striking staircase structure, suggestive of double-diffusive convection, is observed within the transition layer. The staircase structure is observed for about 1000 km across the basin and has been persistent for more than a decade. It is characterized by 2–3 mixed layers (10–60 m thick) separated by 2–16 m thick interfaces. Standard formulae, based on temperature and salinity jumps, suggest a double-diffusive heat flux through the staircase of about 40 mW m⁻², consistent with the measured geothermal heat flux of 40–60 mW m⁻². This is to be expected for a scenario with no deep-water renewal at present as we also show that changes in the bottom layer are too small to account for more than a small fraction of the geothermal heat flux. On the other hand, the observed interfaces between mixed layers in the staircase are too thick to support the required double-diffusive heat flux, either by molecular conduction or by turbulent mixing, as there is no evidence of sufficiently vigorous overturns within the interfaces. It therefore seems, that while the staircase structure may be maintained by a very weak heat flux, most of the geothermal heat flux is escaping through regions of the basin near lateral boundaries, where the staircase structure is not observed. The vertical eddy diffusivity required in these near-boundary regions is O(10⁻³) m² s⁻¹. This implies Thorpe scales of order 10 m. We observe what may be Thorpe scales of this magnitude in boundary-region potential temperature profiles, but cannot tell if they are compensated by salinity. The weak stratification of the transition layer means that the large vertical mixing rate implies a local dissipation rate of only O(10⁻¹⁰) W kg⁻¹, which is not ruled out by plausible energy budgets. In addition, we discuss an alternative scenario of slow, continuous renewal of the CB deep water. In this scenario, we find that some of the geothermal heat flux is required to heat the new water and vertical fluxes through the transition layer are reduced.     

Absolutely referenced geostrophic velocity and transport on a section across the North Atlantic Current

A transect of CTD profiles crossing the North Atlantic Current (NAC) along WOCE line ACM6 near 42.5°N during August 1–7, 1993, provides geostrophic shear velocity profiles, which were absolutely referenced using simultaneous POGO transport float measurements and velocity measurements from a ship-mounted acoustic doppler current profiler (ADCP). The NAC absolute transport was 112±23×10⁶ m³ s⁻¹, which includes a portion of the transport of the Mann Eddy, a large permanent anticyclonic eddy commonly adjacent to the NAC. The NAC transport estimated relative to a level of no motion at the bottom would have underestimated the true total absolute transport by 20%. A surprisingly large 58×10⁶ m³ s⁻¹ flowed southward just inshore of the NAC. This flow, centered near 1500 dbars about 200 km offshore of the shelf-break, was fairly barotropic with a peak velocity of greater than 20 cm s⁻¹, and the water mass characteristics were of Labrador Sea Water. These absolute transport observations suggest southward recirculation inshore of the NAC at 42.5°N and a stronger NAC than has previously been observed.     

Xanthophyll cycling in Phaeocystis globosa and Thalassiosira sp.: a possible mechanism for species succession

The ability of phytoplankton species to initiate photo-protective mechanisms and the rates by which they do so have been suggested to be partly responsible for species succession. To examine whether this is also true in the case of diatom spring blooms preceding Phaeocystis globosa, cultures of P. globosa and Thalassiosira sp. were investigated under controlled laboratory conditions for differences in their xanthophyll cycling rates and abilities. It was found that P. globosa exhibited active and rapid xanthophyll cycling when cultures photoacclimated to 10, 50 and 100 μmol quanta m⁻² s⁻¹, were shifted to 150 and 250 μmol quanta m⁻² s⁻¹. The early spring diatom Thalassiosira only exhibited xanthophyll cycling when acclimated to 10 μmol quanta m⁻² s⁻¹. P. globosa always exhibited faster xanthophyll cycling rates than Thalassiosira, giving P. globosa a possible competitive edge over Thalassiosira sp. It was also found that the magnitude of xanthophyll cycling correlates with the intensity of light changes during the one-hour xanthophyll cycling experiments, and thus may be regarded as related to photoacclimation.     

Basin-scale variability of phytoplankton biomass,production and growth in the Atlantic Ocean

The latitudinal distributions of phytoplankton biomass, composition and production in the Atlantic Ocean were determined along a 10,000-km transect from 50°N to 50°S in October 1995, May 1996 and October 1996. Highest levels of euphotic layer-integrated chlorophyll a (Chl a) concentration (75–125 mg Chl m⁻²) were found in North Atlantic temperate waters and in the upwelling region off NW Africa, whereas typical Chl a concentrations in oligotrophic waters ranged from 20 to 40 mg Chl m⁻². The estimated concentration of surface phytoplankton carbon (C) biomass was 5–15 mg C m⁻² in the oligotrophic regions and increased over 40 mg C m⁻² in richer areas. The deep chlorophyll maximum did not seem to constitute a biomass or productivity maximum, but resulted mainly from an increase in the Chl a to C ratio and represented a relatively small contribution to total integrated productivity. Primary production rates varied from 50 mg C m⁻² d⁻¹ at the central gyres to 500–1000 mg C m⁻² d⁻¹ in upwelling and higher latitude regions, where faster growth rates (μ) of phytoplankton (>0.5 d⁻¹) were also measured. In oligotrophic waters, microalgal growth was consistently slow [surface μ averaged 0.21±0.02 d⁻¹ (mean±SE)], representing <20% of maximum expected growth. These results argue against the view that the subtropical gyres are characterized by high phytoplankton turnover rates. The latitudinal variations in μ were inversely correlated to the changes in the depth of the nitracline and positively correlated to those of the integrated nitrate concentration, supporting the case for the role of nutrients in controlling the large-scale distribution of phytoplankton growth rates. We observed a large degree of temporal variability in the phytoplankton dynamics in the oligotrophic regions: productivity and growth rates varied in excess of 8-fold, whereas microalgal biomass remained relatively constant. The observed spatial and temporal variability in the biomass specific rate of photosynthesis is at least three times larger than currently assumed in most satellite-based models of global productivity.     

Oxygen dynamics in the North Atlantic subtropical gyre

Dissolved oxygen (DO) in the ocean is a tracer for most ocean biogeochemical processes including net community production and remineralization of organic matter which in turn constrains the biological carbon pump. Knowledge of oxygen dynamics in the North Atlantic Ocean is mainly derived from observations at the Bermuda Atlantic Time-series Study (BATS) site located in the western subtropical gyre which may skew our view of the biogeochemistry of the subtropical North Atlantic. This study presents and compares a 15 yr record of DO observations from ESTOC (European Station for Time-Series in the Ocean, Canary Islands) in the eastern subtropical North Atlantic with the 20 yr record at BATS. Our estimate for net community production of oxygen was 2.3±0.4 mol O₂ m⁻² yr⁻¹ and of oxygen consumption was −2.3±0.5 mol O₂ m⁻² yr⁻¹ at ESTOC, and 4 mol O₂ m⁻² yr⁻¹ and −4.4±1 mol m⁻² yr⁻¹ at BATS, respectively. These values were determined by analyzing the time-series using the Discrete Wavelet Transform (DWT) method. These flux values agree with similar estimates from in-situ observational studies but are higher than those from modeling studies. The difference in net oxygen production rates supports previous observations of a lower carbon export in the eastern compared to the western subtropical Atlantic. The inter-annual analysis showed clear annual cycles at BATS whereas longer cycles of nearly 4 years were apparent at ESTOC. The DWT analysis showed trends in DO anomalies dominated by long-term perturbations at a basin scale for the consumption zones at both sites, whereas yearly cycles dominated the production zone at BATS. The long-term perturbations found are likely associated with ventilation of the main thermocline, affecting the consumption and production zones at ESTOC.     

Contrasting life-histories,secondary production,and trophic structure of Peracarid assemblages of the bathyal suprabenthos from the Bay of Biscay (NE Atlantic) and the Catalan Sea (NW Mediterranean)

The life-histories and the secondary production of four dominant peracarid crustaceans (the mysids Boreomysis arctica and Parapseudomma calloplura, the amphipod Rhachotropis caeca, and the isopod Ilyarachna longicornis) in bathyal depths of the Bay of Biscay (NE Atlantic; between 383 and 420 m) and the Catalan Sea (Northwestern Mediterranean; between 389 and 1355 m) were established. Both the Atlantic and the Mediterranean populations of the major part of the target-species had two generations/year with mean cohort-production intervals (CPI) ranging from 5.5 mo for Ilyarachna longicornis to 6.3 mo for Parapseudomma calloplura. The Hynes method showed secondary production to vary in the Bay of Biscay between 0.113 mg DW m⁻² yr⁻¹ for I. longirostris and 3.069 mg DW m⁻² yr⁻¹ for P. calloplura, with P/B ratios between 4.57 (I. longirostris) and 7.93 (Boreomysis arctica). In the Catalan Sea, production varied between 0.286 mg DW m⁻² yr⁻¹ for I. longirostris and 1.096 mg DW m⁻² yr⁻¹ for P. calloplura, with P/B between 5.72 (I. longirostris) and 6.66 (P. calloplura). Application of two different empiric models to the whole peracarid assemblage gave similar levels of secondary production in both study areas (between 29.26 and 32.14 mgDWm⁻² yr⁻¹ in the Bay of Biscay; between 26.23 and 26.54 mg DW m⁻² yr⁻¹ in the Catalan Sea). From the analysis of gut contents of 22 species the dominant species in each study area were assigned to two basic trophic levels, detritus feeders and predators. Also, cumulative curves of dominance showed high diversity (low dominance) for peracarid assemblages distributed at mid-bathyal depths (524–693 m) both in the Bay of Biscay off Arcachon and in the Catalan Sea off Barcelona. We also discuss and compare, both within and between areas, how environmental features may explain the observed diversity patterns, the trophic structure, and the production results obtained for the suprabenthos assemblages.     

Latitudinal distribution of microbial plankton abundance,production, and respiration in the Equatorial Atlantic in autumn 2000

Phytoplankton and bacterial abundance, size-fractionated phytoplankton chlorophyll-a (Chl-a) and production together with bacterial production, microbial oxygen production and respiration rates were measured along a transect that crossed the Equatorial Atlantic Ocean (10°N–10°S) in September 2000, as part of the Atlantic Meridional Transect 11 (AMT 11) cruise. From 2°N to 5°S, the equatorial divergence resulted in a shallowing of the pycnocline and the presence of relatively high nitrate (>1 μM) concentrations in surface waters. In contrast, a typical tropical structure (TTS) was found near the ends of the transect. Photic zone integrated ¹⁴C primary production ranged from ∼200 mg C m⁻² d⁻¹ in the TTS region to ∼1300 mg C m⁻² d⁻¹ in the equatorial divergence area. In spite of the relatively high primary production rates measured in the equatorial upwelling region, only a moderate rise in phytoplankton biomass was observed as compared to nearby nutrient-depleted areas (22 vs. 18 mg Chl-a m⁻², respectively). Picophytoplankton were the main contributors (>60%) to both Chl-a biomass and primary production throughout the region. The equatorial upwelling did not alter the phytoplankton size structure typically found in the tropical open ocean, which suggests a strong top-down control of primary producers by zooplankton. However, the impact of nutrient supply on net microbial community metabolism, integrated over the euphotic layer, was evidenced by an average net microbial community production within the equatorial divergence (1130 mg C m⁻² d⁻¹) three-fold larger than net production measured in the TTS region (370 mg C m⁻² d⁻¹). The entire region under study showed net autotrophic community metabolism, since respiration accounted on average for 51% of gross primary production integrated over the euphotic layer.     

Total organic and inorganic carbon exchange through the Strait of Gibraltar in September 1997

The total organic carbon (TOC) and total inorganic carbon (C_T) exchange between the Atlantic Ocean and the Mediterranean Sea was studied in the Strait of Gibraltar in September 1997. Samples were taken at eight stations from western and eastern entrances of the Strait and at the middle of the Strait (Tarifa Narrows). TOC was analyzed by a high-temperature catalytic oxidation method, and C_T was calculated from alkalinity–pH_T pairs and appropriate thermodynamic relationships. The results are used in a two-layer model of water mass exchange through the Strait, which includes the Atlantic inflow, the Mediterranean outflow and the interface layer in between. Our observations show a decrease of TOC and an increase of C_T concentrations from the surface to the bottom: 71–132 μM C and 2068–2150 μmol kg⁻¹ in the Surface Atlantic Water, 74–95 μM C and 2119–2148 μmol kg⁻¹ in the North Atlantic Central Water, 63–116 μM C and 2123–2312 μmol kg⁻¹ in the interface layer, and 61–78 μM C and 2307–2325 μmol kg⁻¹ in the Mediterranean waters. However, within the Mediterranean outflow, we found that the concentrations of carbon were higher at the western side of the Strait (75–78 μM C, 2068–2318 μmol kg⁻¹) than at the eastern side (61–69 μM C, 2082–2324 μmol kg⁻¹). This difference is due to the mixing between the Atlantic inflow and the Mediterranean outflow on the west of the Strait, which results in a flux of organic carbon from the inflow to the outflow and an opposite flux of inorganic carbon. We estimate that the TOC input from the Atlantic Ocean to the Mediterranean Sea through the Strait of Gibraltar varies from (0.97±0.8)10⁴ to (1.81±0.90)10⁴ mol C s⁻¹ (0.3×10¹² to 0.56×10¹² mol C yr⁻¹), while outflow of inorganic carbon ranges from (12.5±0.4)10⁴ to (15.6±0.4)10⁴ mol C s⁻¹ (3.99–4.90×10¹² mol C yr⁻¹). The high variability of carbon exchange within the Strait is due to the variability of vertical mixing between inflow and outflow along the Strait. The prevalence of organic carbon inflow and inorganic carbon outflow shows the Mediterranean Sea to be a basin of active remineralization of organic material.     

The natural diet of a hexactinellid sponge: Benthic–pelagic coupling in a deep-sea microbial food web

Dense communities of shallow-water suspension feeders are known to sidestep the microbial loop by grazing on ultraplankton at its base. We quantified the diet, rates of water processing, and abundance of the deep-sea hexactinellid sponge Sericolophus hawaiicus, and found that, like their demosponge relatives in shallow water, hexactinellids are a significant sink for ultraplankton. S. hawaiicus forms a dense bed of sponges on the Big Island of Hawaii between 360 and 460 m depth, with a mean density of 4.7 sponges m⁻². Grazing of S. hawaiicus on ultraplankton was quantified from in situ samples using flow cytometry, and was found to be unselective. Rates of water processing were determined with dye visualization and ranged from 1.62 to 3.57 cm s⁻¹, resulting in a processing rate of 7.9±2.4 ml sponge⁻¹ s⁻¹. The large amount of water processed by these benthic suspension feeders results in the transfer of approximately 55 mg carbon and 7.3 mg N d⁻¹ m⁻² from the water column to the benthos. The magnitude of this flux places S. hawaiicus squarely within the functional group of organisms that link the pelagic microbial food web to the benthos.     

Export fluxes of particulate organic carbon in the Chukchi Sea: A comparative study using 234Th/238U disequilibria and drifting sediment traps

Large-volume sampling of ²³⁴Th and drifting sediment trap deployments were conducted as part of the 2004 Western Arctic Shelf–Basin Interactions (SBI) spring (May 15–June 23) and summer (July 17–August 26) process cruises in the Chukchi Sea. Measurements of ²³⁴Th and particulate organic carbon (POC) export fluxes were obtained at five stations during the spring cruise and four stations during the summer cruise along Barrow Canyon (BC) and along a parallel shelf-to-basin transect from East Hanna Shoal (EHS) to the Canada Basin. ²³⁴Th and POC fluxes obtained with in situ pumps and drifting sediment traps agreed to within a factor of 2 for 70% of the measurements. POC export fluxes measured with in situ pumps at 50 m along BC were similar in spring and summer (average = 14.0 ± 8.0 mmol C m^− 2 day^− 1 and 16.5 ± 6.5 mmol C m^− 2 day^− 1, respectively), but increased from spring to summer at the EHS transect (average = 1.9 ± 1.1 mmol C m^− 2 day^− 1 and 19.5 ± 3.3 mmol C m^− 2 day^− 1, respectively). POC fluxes measured with sediment traps at 50 m along BC were also similar in both seasons (31.3 ± 9.3 mmol C m^− 2 day^− 1 and 29.1 ± 14.2 mmol C m^− 2 day^− 1, respectively), but were approximately twice as high as POC fluxes measured with in situ pumps. Sediment trap POC fluxes measured along the EHS transect also increased from spring to summer (3.0 ± 1.9 mmol C m^− 2 day^− 1 and 13.0 ± 6.4 mmol C m^− 2 day^− 1, respectively), and these fluxes were similar to the POC fluxes obtained with in situ pumps. Discrepancies in POC export fluxes measured using in situ pumps and sediment traps may be reasonably explained by differences in the estimated POC/²³⁴Th ratios that arise from differences between the techniques, such as time-scale of measurement and size and composition of the collected particles. Despite this variability, in situ pump and sediment trap-derived POC fluxes were only significantly different at a highly productive station in BC during the spring.     

Microbial biomass and viral infections of heterotrophic prokaryotes in the sub-surface layer of the central Arctic Ocean

Seawater samples were collected for microbial analyses between 55 and 235 m depth across the Arctic Ocean during the SCICEX 97 expedition (03 September–02 October 1997) using a nuclear submarine as a research platform. Abundances of prokaryotes (range 0.043–0.47×10⁹ dm⁻³) and viruses (range 0.68–11×10⁹ dm⁻³) were correlated (r=0.66, n=150) with an average virus:prokaryote ratio of 26 (range 5–70). Biomass of prokaryotes integrated from 55 to 235 m ranged from 0.27 to 0.85 g C m⁻² exceeding that of phytoplankton (0.005–0.2 g C m⁻²) or viruses (0.02–0.05 g C m⁻²) over the same depth range by an order of magnitude on average. Using transmission electron microscopy (TEM), we estimated that 0.5% of the prokaryote community on average (range 0–1.4%) was visibly infected with viruses, which suggests that very little of prokaryotic secondary production was lost due to viral lysis. Intracellular viruses ranged from 5 to >200/cell, with an average apparent burst size of 45±38 (mean±s.d.; n=45). TEM also revealed the presence of putative metal-precipitating bacteria in 8 of 13 samples, which averaged 0.3% of the total prokaryote community (range 0–1%). If these prokaryotes are accessible to protistan grazers, the Fe and Mn associated with their capsules might be an important source of trace metals to the planktonic food web. After combining our abundance and mortality data with data from the literature, we conclude that the biomass of prokaryoplankton exceeds that of phytoplankton when averaged over the upper 250 m of the central Arctic Ocean and that the fate of this biomass is poorly understood.     

The western Arctic boundary current at 152°W: Structure,variability, and transport

From August 2002 to September 2004 a high-resolution mooring array was maintained across the western Arctic boundary current in the Beaufort Sea north of Alaska. The array consisted of profiling instrumentation, providing a timeseries of vertical sections of the current. Here we present the first-year velocity measurements, with emphasis on the Pacific water component of the current. The mean flow is characterized as a bottom-intensified jet of O (15 cm s⁻¹) directed to the east, trapped to the shelfbreak near 100 m depth. Its width scale is only 10–15 km. Seasonally the flow has distinct configurations. During summer it becomes surface-intensified as it advects buoyant Alaskan Coastal water. In fall and winter the current often reverses (flows westward) under upwelling-favorable winds. Between the storms, as the eastward flow re-establishes, the current develops a deep extension to depths exceeding 700 m. In spring the bottom-trapped flow advects winter-transformed Pacific water emanating from the Chukchi Sea. The year-long mean volume transport of Pacific water is 0.13±0.08 Sv to the east, which is less than 20% of the long-term mean Bering Strait inflow. This implies that most of the Pacific water entering the Arctic goes elsewhere, contrary to expected dynamics and previous modeling results. Possible reasons for this are discussed. The mean Atlantic water transport (to 800 m depth) is 0.047±0.026 Sv, also smaller than anticipated.     

Factors controlling silicon and nitrogen biogeochemical cycles in high nutrient,low chlorophyll systems (the Southern Ocean and the North Pacific): Comparison with a mesotrophic system (the North Atlantic)

A 1-D coupled physical-biogeochemical model is used to study the seasonal cycles of silicon and nitrogen in two High Nutrient Low Chlorophyll (HNLC) systems, the Antarctic Circumpolar Current (ACC) and the North Pacific Ocean, and a mesotrophic system, the North Atlantic Ocean. The biological model consists of nine compartments (diatoms, nano-flagellates, microzooplankton, mesozooplankton, two types of detritus, nitrate, ammonium and silicic acid) forced by irradiance, temperature, mixing and deep nitrate and silicic acid concentrations. At all sites, nanophytoplankton standing crop variations are low, in spite of variations in primary production, because of a “top–down” control by microzooplankton. Although nanophytoplankton sustain more than 60% of the annual primary production in these areas, their contribution to the export production does not exceed 1% of the total. The differences in the seasonal plankton cycle among these regions come mainly from differences in the dynamics of large phytoplankton (here diatoms). In the ACC, the chlorophyll maximum remains <1.5 mg m⁻³, as an unfavourable light/mixing regime and a likely trace-metal limitation keep diatoms from blooming. In the northeast Pacific, trace-metal limitation seems to keep diatoms from blooming throughout the year. In both these systems, light or iron limitations induce high Si/N uptake ratios. Incidentally these high Si/N uptake ratios lead to a net excess of silicic acid utilization over nitrate, and to a subsequent silicic acid limitation during the summertime. In the North Atlantic, under favourable light/mixing regime and nutrient-replete conditions at the onset of the growing period, diatoms outburst and sustain a bloom >3.5 mg Chl-a m⁻³. Thereafter, mesozooplankton grazing pressure and silicic acid limitation induce the collapse of the chlorophyll maximum and the persistence of lower chlorophyll concentrations in summer. Although the ACC and the North Pacific show HNLC features, they support a high biogenic silica production (1.9 and 1.07 mol Si m⁻² yr⁻¹) and export flux (0.79 and 0.61 mol Si m⁻² yr⁻¹), compared to the North Atlantic (production: 0.23 mol Si m⁻² yr⁻¹, export: 0.12 mol Si m⁻² yr⁻¹). The differences in Si production and export between the HNLC systems and the mesotrophic North Atlantic come from both higher Si concentrations and Si/N uptake ratios in the HNLC areas compared to the North Atlantic. Also, the low dissolution rate of biogenic silica compared to nitrogen degradation rate, and the inhibition of nitrate uptake by ammonium, reinforce the net excess of silicic acid utilization over nitrate. As a result, the model also illustrates the efficiency of the silica pump for the three sites: about 50% of the biogenic silica synthesized in the euphotic layer is exported out of the first 100 m, while only 4–11% of the particulate organic nitrogen escapes recycling in the surface layer.     

Variability of chlorophyll and primary production in the Eastern North Atlantic Subtropical Gyre: potential factors affecting phytoplankton activity

Size-fractionated chlorophyll-a and carbon incorporation rates were determined on a series of 13 cruises carried out from 1992 to 2001with the aim of investigating the patterns and causes of variability in phytoplankton chlorophyll and production in the Eastern North Atlantic Subtropical Gyral Province (NASE). Averaged (±SE) integrated chlorophyll-a concentration and primary production rate were 17±1 mg m⁻² and 253±22 mg C m⁻² d⁻¹. Small-sized cells (<2 μm) formed the bulk of phytoplankton biomass (71%) and accounted for 54% of total primary production. A clear latitudinal gradient in these variables was not detected. By contrast, large seasonal variability was detected in terms of primary production, although integrated phytoplankton biomass, as estimated from chlorophyll-a concentration, remained rather constant and did not display significant changes with time. Variability in primary production (PP) was related mainly to variability in surface temperature and surface chlorophyll-a concentration. The control exerted by surface temperature was related to nutrient availability. By contrary, euphotic-zone depth, depth of maximum concentration of chlorophyll-a and integrated chlorophyll-a did not contribute significantly to the high variability in primary production observed in this oligotrophic region.     

The metabolic balance between autotrophy and heterotrophy in the western Arctic Ocean

The goal of this study was to explore how net community production (NCP) is influenced by the relationship between primary production and community respiration in the western Arctic Ocean. Plankton NCP and respiration were determined by measuring changes in oxygen in light and dark bottle incubations, respectively. Rates of NCP averaged over shelf, slope and basin waters were positive in summer 2002 (57±191 mmol O₂ m⁻² d⁻¹) and spring 2004 (85±86 mmol O₂ m⁻² d⁻¹) and negative in summer 2004 (−25±176 mmol O₂ m⁻² d⁻¹). Determinations of NCP obtained from bottle incubations were similar to rates inferred from in situ changes in dissolved inorganic carbon. An examination of the spatial variability of primary production and community respiration indicated that respiration is distributed more uniformly than primary production. A spatial offset between photosynthesis and respiration from the shelf to the Arctic basin was present in spring 2004, but was not seen at other times. NCP and the potential for export appear to be dependent on an uncoupling of primary production and community respiration. NCP continued into the summer after the stock of NO₃⁻ had been depleted. Our data suggest that the uniform distribution of respiration relative to primary production is an important factor influencing NCP and the potential for export in the western Arctic.     

Coccolithophorids on the continental slope of the Bay of Biscay – production,transport and contribution to mass fluxes

Coccoliths collected by sediment traps deployed on the slope of the Bay of Biscay (northeastern Atlantic), from June 1990 to August 1991, were examined to determine their contribution to the transport of carbonate on a mid-latitude continental margin. They also were used as tracers of particle transfer processes on this slope. Two traps located at 1900 m, respectively at 2300 (Mooring Site 1) and 3000 m (Mooring Site 2) water depths provided high-resolution (4–7 days) time-series samples covering a 14-month period at MS2 and a 3-month period at MS1. Coccoliths from 28 species were identified over the course of the experiment, among which Emiliania huxleyi was always dominant (relative abundance range: 59–93%). Total coccoliths number fluxes were high but variable, ranging from 390×10⁶ to 1610×10⁶ coccoliths m⁻² day⁻¹ at MS1, and from 58×10⁶ to 1500×10⁶ coccoliths m⁻² day⁻¹ at MS2. The time-weighted mean flux, calculated for the whole experiment at MS2, was 499×10⁶ coccoliths m⁻² day⁻¹. Estimate of coccoliths minimal contribution to total carbonate flux at 1900 m depth averaged 12%, which represented a weighted mean flux of 7.3 mg m⁻² day⁻¹ (2.7 g m⁻² yr⁻¹). Lateral transport of coccoliths resuspended from shelf and/or upper slope sediments seems to be the dominant transfer process to depth on this northeastern Atlantic slope. Nevertheless, the clear seasonal succession observed in the species composition implies that the deposition/resuspension/transport sequence is rapid (presumably less than a few months). Several short and unsmoothed signals directly issued from coccoliths bloom events also were recorded in our traps, a result that indicates rapid settling rates. The overall coccolith sedimentation processes appear as being quite diversified, but quantitative and qualitative analyses of aggregates collected by the traps suggest that they are important carriers of coccoliths in this margin environment.     

Diel variations of the bathymetric distribution of zooplankton groups and biomass in Cap-Ferret Canyon,France

The bathymetric distribution, abundance and diel vertical migrations (DVM) of zooplankton were investigated along the axis of the Cap-Ferret Canyon (Bay of Biscay, French Atlantic coast) by a consecutive series of synchronous net hauls that sampled the whole water column (0–2000 m in depth) during a diel cycle. The distribution of appendicularians (maximum 189 individuals m⁻³), cladocerans (maximum 287 individuals m⁻³), copepods (copepods<4 mm, maximum 773 individuals m⁻³, copepods>4 mm, maximum 13 individuals m⁻³), ostracods (maximum 8 individuals m⁻³), siphonophores (maximum >2 individuals m⁻³) and peracarids (maximum >600 individuals 1000 m⁻³) were analysed and represented by isoline diagrams. The biomass of total zooplankton (maximum 18419 μg C m⁻³, 3780 μg N m⁻³) and large copepods (>4 mm maximum 2256 μg C m⁻³, 425 μg N m⁻³) also were determined. Vertical migration was absent or affected only the epipelagic zone for appendicularians, cladocerans, small copepods and siphonophores. Average amplitude of vertical migration was about 400–500 m for ostracods, some hyperiids and mysids, and large copepods, which were often present in the epipelagic, mesopelagic, and bathypelagic zones. Large copepods can constitute more than 80% of the biomass corresponding to total zooplankton. They may play an important role in the active vertical transfer of carbon and nitrogen.     

Community respiration/production and bacterial activity in the upper water column of the central Arctic Ocean

Community metabolism (respiration and production) and bacterial activity were assessed in the upper water column of the central Arctic Ocean during the SHEBA/JOIS ice camp experiment, October 1997–September 1998. In the upper 50 m, decrease in integrated dissolved oxygen (DO) stocks over a period of 124 d in mid-winter suggested a respiration rate of ∼3.3 nM O₂ h⁻¹ and a carbon demand of ∼4.5 gC m⁻². Increase in 0–50 m integrated stocks of DO during summer implied a net community production of ∼20 gC m⁻². Community respiration rates were directly measured via rate of decrease in DO in whole seawater during 72-h dark incubation experiments. Incubation-based respiration rates were on average 3-fold lower during winter (11.0±10.6 nM O₂ h⁻¹) compared to summer (35.3±24.8 nM O₂ h⁻¹). Bacterial heterotrophic activity responded strongly, without noticeable lag, to phytoplankton growth. Rate of leucine incorporation by bacteria (a proxy for protein synthesis and cell growth) increased ∼10-fold, and the cell-specific rate of leucine incorporation ∼5-fold, from winter to summer. Rates of production of bacterial biomass in the upper 50 m were, however, low compared to other oceanic regions, averaging 0.52±0.47 ngC l⁻¹ h⁻¹ during winter and 5.1±3.1 ngC l⁻¹ h⁻¹ during summer. Total carbon demand based on respiration experiments averaged 2.4±2.3 mgC m⁻³ d⁻¹ in winter and 7.8±5.5 mgC m⁻³ d⁻¹ in summer. Estimated bacterial carbon demand based on bacterial productivity and an assumed 10% gross growth efficiency was much lower, averaging about 0.12±0.12 mgC m⁻³ d⁻¹ in winter and 1.3±0.7 mgC m⁻³ d⁻¹ in summer. Our estimates of bacterial activity during summer were an order of magnitude less than rates reported from a summer 1994 study in the central Arctic Ocean, implying significant inter-annual variability of microbial processes in this region.