Particulate organic carbon export fluxes and size-fractionated POC/234Th ratios in the Ligurian,Tyrrhenian and Aegean Seas

Measurements of ²³⁴Th/²³⁸U disequilibria and particle size-fractionated (1, 10, 20, 53, 70, 100 μm) organic C and ²³⁴Th were made to constrain estimates of the export flux of particulate organic C (POC) from the surface waters of the Ligurian, Tyrrhenian and Aegean Seas in March–June 2004. POC exported from the surface waters (75–100 m depth) averaged 9.2 mmol m⁻² d⁻¹ in the Ligurian and Tyrrhenian Seas (2.3±0.5–14.9±3.0 mmol m⁻² d⁻¹) and 0.9 mmol m⁻² d⁻¹ in the Aegean Sea. These results are comparable to previous measurements of ²³⁴Th-derived and sediment-trap POC fluxes from the upper 200 m in the Mediterranean Sea. Depth variations in the POC/²³⁴Th ratio suggest two possible controls. First, decreasing POC/²³⁴Th ratios with depth were attributed to preferential remineralization of organic C. Second, the occurrence of maxima or minima in the POC/²³⁴Th ratio near the DCM suggests influence by phytoplankton dynamics. To assess the accuracy of these data, the empirical ²³⁴Th-method was evaluated by quantifying the extent to which the ²³⁴Th-based estimate of POC flux, P_POC, deviates from the true flux, F_POC, defined as the p-ratio (p-ratio=P_POC/F_POC=S_Th/S_POC, where S=particle sinking rate). Estimates of the p-ratio made using Stokes’ Law and the particle size distributions of organic C and ²³⁴Th yield values ranging from 0.93–1.45. The proximity of the p-ratio to unity implies that differences in the sinking rates of POC- and ²³⁴Th-carrying particles did not bias ²³⁴Th-normalized POC fluxes by more than a factor of two.     

A time-series study of particulate matter export in the North Pacific Subtropical Gyre based on 234Th : 238U disequilibrium

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

Strong seasonality in particle dynamics of north-western Mediterranean surface waters as revealed by 234Th/238U

²³⁴Th was used to quantify sinking fluxes and residence times of particles in surface waters of the north-western Mediterranean Sea. Measurements of dissolved and particulate ²³⁴Th were made at the DYFAMED station (43°25′N–7°51′E, JGOFS-France program). Sampling covered 1 year on four cruises in 1994 (February 9, April 29, June 3, October 1) and focused on a transition period in mid-spring with six repeated profiles collected during May 1995. ²³⁴Th was nearly in equilibrium with its parent ²³⁸U most of the year, except in spring. The intensive sampling in May shows a rapid evolution throughout the month from a moderate ²³⁴Th deficit to near-equilibrium values. The time-series of ²³⁴Th were treated with steady-state and non-steady-state models. ²³⁴Th particulate fluxes clearly indicate large variability in export, with the highest values observed in spring. Particle residence times in the upper 40 m range from <10 to >250 days, and could increase by a factor of 10 within 2 weeks. POC fluxes from the upper 40 m and export ratios (ThE: ratio of ²³⁴Th-derived POC export to primary production), derived from the ²³⁴Th/²³⁸U disequilibrium in the water column and POC/²³⁴Th ratio on trapped material, decrease from about 9.5 mmol C m⁻² d⁻¹ and >22% in early May to less than 5 mmol C m⁻² d⁻¹ and 15% after mid-May. The ²³⁴Th-derived information is in agreement with the annual variations in Mediterranean Sea productivity.     

Time-series observation of POC fluxes estimated from 234Th in the northwestern North Pacific

Time-series measurements of ²³⁴Th activities and particulate organic carbon (POC) concentrations were made at time-series stations (K1, K2, K3, and KNOT) in the northwestern North Pacific from October 2002 to August 2004. Seasonal changes in POC export fluxes from the surface layer (∼100 m) were estimated using ²³⁴Th as a tracer. POC fluxes varied seasonally from approximately 0 to 180 mg C m⁻² d⁻¹ and were higher in spring–summer than in autumn–winter. The export ratio (e-ratio) ranged from 6% to 55% and was also higher in spring–summer. Annual POC fluxes were estimated to be 31 g C m⁻² y⁻¹ in the subarctic region (station K2) and 23 g C m⁻² y⁻¹ in the region between the subarctic and subtropical gyres (station K3). POC fluxes and e-ratios in the northwestern North Pacific were much higher than those in most other oceans. The annual POC flux corresponded to 69% of annual new production estimated from the seasonal difference of the nutrient in the Western Subarctic Gyre (45 g C m⁻² y⁻¹). These results indicate that much of the organic carbon assimilated in the surface layer of the northwestern North Pacific is transferred to the deep ocean in particulate form. Our conclusions support previous reports that diatoms play an important role in the biological pump.     

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

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

Role of colloidal material in the removal of 234Th in the Canada basin of the Arctic Ocean

The phase partitioning of ²³⁴Th between dissolved (<10-kiloDalton, kD), colloidal (10 kD—0.4 μm), and particulate (⩾0.5 μm) matter across a horizontal transect, from a coastal station to the deep Canada Basin, and a vertical profile in the deep Canada Basin of the western Arctic Ocean was investigated. Concentrations of suspended particulate matter (SPM), dissolved, colloidal and particulate organic carbon, particulate organic nitrogen and nutrients (silicate, phosphate and nitrate) were also measured to assess transport and scavenging processes.Total ²³⁴Th (colloidal+particulate+dissolved) indicated deficiencies relative to secular equilibrium with its parent, ²³⁸U in the upper 100 m, which suggests active scavenging of ²³⁴Th onto particle surfaces. In contrast, at depths >200 m, general equilibrium existed between total ²³⁴Th and ²³⁸U. The inventory of SPM and the specific activity of particulate ²³⁴Th in the Canada Basin was about an order of magnitude higher than the profile reported for the Alpha Ridge ice camp station. This higher concentration of SPM in the southwestern Canada Basin is likely derived from ice-rafted sedimentary particles. Inventories of nutrients, and dissolved organic carbon and nitrogen in the upper 100 m of the Canada Basin are comparable to the other estimates for the central Arctic Ocean. Comparison of the mass concentrations of colloidal and filter-retained particulate matter as well as the activity of ²³⁴Th in these phases indicates that only a very small component of the colloidal material is actively involved in Th scavenging. Lower values of the conditional partition coefficient between the colloidal and dissolved phase indicate that the Arctic colloids are less reactive than colloidal material from other regions. The conditional partition coefficient between the filter-retained and dissolved phases (K_f) is generally higher than that for other regions, which is attributed to the higher complexation capacity of glacio-marine sedimentary particles in these waters. The ²³⁴Th-derived export of POC for the shelf and deep Canada Basin ranges between 5.6 and 6.5 mmol m⁻² d⁻¹, and is in agreement with other estimates reported for the central Arctic Ocean and Beaufort Sea.     

Seasonal changes in thorium scavenging and particle aggregation in the western Arctic Ocean

Seasonal changes in Th scavenging and particle aggregation were determined along two shelf-basin transects in the western Arctic Ocean during the spring (May–June) and summer (July–August) of 2002 and 2004. Measurements of dissolved and particulate ²³⁴Th and ²²⁸Th activities were used to quantify Th residence times and reversible rates of Th sorption and particle aggregation. Prior to the spring bloom in 2002, ²³⁴Th and ²²⁸Th residence times were equal and Th scavenging was concordant, indicating predominately steady-state conditions. In contrast, scavenging of ²³⁴Th and ²²⁸Th in the summer of 2002 and the spring and summer of 2004 was discordant, indicating a departure in scavenging rates from steady-state conditions during periods of seasonally high biological activity and particle export. Rates of particle aggregation and disaggregation were calculated using a one-dimensional reversible exchange model and ²³⁴Th and ²²⁸Th activities in small (1–53 μm) and large (>53 μm) particles. Maximum rates were determined coincident with the chlorophyll maximum (25–100 m) and increased by an order of magnitude between periods of low and high productivity. These Th measurements provide evidence that seasonally enhanced rates of particle aggregation might increase the magnitude of the particulate organic carbon (POC) flux in this Arctic regime.     

Fate of biogenic carbon in the upper 200 m of the central Greenland Sea

Sedimentation of particulate carbon from the upper 200–300 m in the central Greenland Sea from August 1993 to June 1995 was less than 2 g C m⁻² yr⁻¹. Daily rates of sedimentation of particulate organic carbon reached highest values of about 18 mg m⁻² d⁻¹ in fall 1994. For total particulate material, maximum rates of sedimentation of about 250 mg m⁻² d⁻¹ were recorded in spring and fall 1994. For chlorophyll equivalent, highest rates of sedimentation of about 140 μg m⁻² d⁻¹ were recorded in spring 1994. As reported in related investigations, the transient accumulation of DOC in surface waters during summer, as well as respiration and mortality of deep overwintering zooplankton stocks, appeared to dominate the fate of photosynthetically fixed organic carbon. The above processes may account for roughly 43 g C m⁻² in the upper 200 m of the central Greenland Sea. For comparison, the seasonal deficit in dissolved inorganic carbon was reported to be about 23 g C m⁻² in the upper 20 m of surface water, and estimates for new annual production were reported to be about 57 g C m⁻². In our investigation, the biological carbon pump was not unusually effective in transporting carbon out of the productive surface layer.     

Controls of 234Th removal from the oligotrophic ocean by polyuronic acids and modification by microbial activity

To gain new insights into the variability of particulate organic carbon (POC) fluxes and to better understand the factors controlling the POC/²³⁴Th ratios in suspended and sinking particulate matter, we investigated the relationships between POC/²³⁴Th ratios and biochemical composition (uronic acids, URA; total carbohydrates, TCHO; acid polysaccharides, APS; and POC) of suspended and sinking matter from the Gulf of Mexico in 2005 and 2006. Our data show that URA/POC in sediment traps (STs), APS/POC in the suspended particles, and turnover times of particulate ²³⁴Th in the water column and those of bacteria in STs inside eddies usually increased with depth, whereas particulate POC/²³⁴Th (10–50 μm) and the sediment-trap parameters (POC flux, POC/²³⁴Th ratio, bacterial biomass, and bacterial production) decreased with depth. However, this trend was not the case for most biological parameters (e.g., phytoplankton and bacterial biomass) or for the other parameters at the edges of eddies or at coastal-upwelling sites.In general, the following relationships were observed: 1) ²³⁴Th/POC ratios in STs were correlated with APS flux, and these ratios in the 10–50 μm suspended particles also correlated with URA/POC ratios; 2) neither URA fluxes nor URA/POC ratios were significantly related to bacterial biomass; 3) the sum of two uronic acids (G2, glucuronic, and galacturonic acid, which composed most of the URA pool) was positively related to bacterial biomass; and 4) the POC/²³⁴Th ratios in intermediate-sized particles (10–50 μm) were close to those in sinking particles but much lower than those in > 50 μm particles. The results indicate that acid polysaccharides, though a minor fraction (~ 1%) of the organic carbon, act more likely as proxy compound classes that might contain the more refractory ²³⁴Th-binding biopolymer, rather than acting as the original ²³⁴Th “scavenger” compound. Moreover, these acid polysaccharides, which might first be produced by phytoplankton and then modified by bacteria, also influence the on-and-off “piggy-back” processes of organic matter and ²³⁴Th, thus causing additional variability of the POC/²³⁴Th in particles of different sizes.     

Major contribution of autotrophy to microbial carbon cycling in the deep North Atlantic’s interior

Current estimates point to a mismatch of particulate organic carbon supply derived from the surface ocean and the microbial organic carbon demand in the meso- and bathypelagic realm. Based on recent findings that chemoautotrophic Crenarchaeota are abundant in the mesopelagic zone, we quantified dissolved inorganic carbon (DIC) fixation in the meso- and bathypelagic North Atlantic and compared it with heterotrophic microbial activity. Measuring ¹⁴C-bicarbonate fixation and ³H-leucine incorporation revealed that microbial DIC fixation is substantial in the mesopelagic water masses, ranging from 0.1 to 56.7 μmol C m⁻³ d⁻¹, and is within the same order of magnitude as heterotrophic microbial activity. Integrated over the dark ocean’s water column, DIC fixation ranged from 1–2.5 mmol C m⁻² d⁻¹, indicating that chemoautotrophy in the dark ocean represents a significant source of autochthonously produced ‘new organic carbon’ in the ocean’s interior amounting to about 15–53% of the phytoplankton export production. Hence, chemoautotrophic DIC fixation in the oxygenated meso- and bathypelagic water column of the North Atlantic might substantially contribute to the organic carbon demand of the deep-water microbial food web.     

Sediment trap and in-situ pump size-fractionated POC/234Th ratios in the Mediterranean Sea and Northwest Atlantic: Implications for POC export

Measurements of particle size-fractionated POC/²³⁴Th ratios and ²³⁴Th and POC fluxes were conducted using surface-tethered, free-floating, sediment traps and large-volume in-situ pumps during four cruises in 2004 and 2005 to the oligotrophic eastern Mediterranean Sea and the seasonally productive western Mediterranean and northwest Atlantic. Analysis of POC/²³⁴Th ratios in sediment trap material and 10, 20, 53, 70, and 100 μm size-fractionated particles indicate, for most stations, decreasing ratios with depth, a weak dependence on particle size, and ratios that converge to ～1–5 μmol dpm^?1 below the euphotic zone (～100–150 m) throughout the contrasting biogeochemical regimes. In the oligotrophic waters of the Aegean Sea, ²³⁴Th and POC fluxes estimated using sediment traps were consistently higher than respective fluxes estimated from water-column ²³⁴Th–²³⁸U disequilibrium, observations that are attributed to terrigenous particle scavenging of ²³⁴Th. In the more productive western Mediterranean and northwest Atlantic, ²³⁴Th and POC fluxes measured by sediment trap and ²³⁴Th–²³⁸U disequilibrium agreed within a factor of 2–4 throughout the water column. An implication of these results is that estimates of POC export by sediment traps and ²³⁴Th–²³⁸U disequilibrium can be biased differently because of differential settling speeds of POC and ²³⁴Th-carrying particles.     

Seasonal and interannual variability in particle fluxes of carbon,nitrogen and silicon from time series of sediment traps at Ocean Station P, 1982–1993: relationship to changes in subarctic primary productivity

An extended time series of particle fluxes at 3800 m was recorded using automated sediment traps moored at Ocean Station Papa (OSP, 50°N, 145°W) in the northeast Pacific Ocean for more than a decade (1982–1993). Time-series observations at 200 and 1000 m, and short-term measurements using surface-tethered free-drifting sediment traps also were made intermittently. We present data for fluxes of total mass (dry weight), particulate organic carbon (POC), particulate organic nitrogen (PON), biogenic Si (BSi), and particulate inorganic carbon (PIC) in calcium carbonate. Mean monthly fluxes at 3800 m showed distinct seasonality with an annual minimum during winter months (December–March), and maximum during summer and fall (April–November). Fluxes of total mass, POC, PIC and BSi showed 4-, 10-, 7- and 5-fold increases between extreme months, respectively. Mean monthly fluxes of PIC often showed two plateaus, one in May–August dominated by <63 μm particles and one in October–November, which was mainly >63 μm particles. Dominant components of the mass flux throughout the year were CaCO₃ and opal in equal amounts. The mean annual fluxes at 3800 m were 32±9 g dry weight g m⁻² yr⁻¹, 1.1±0.5 g POC m⁻² yr⁻¹, 0.15±0.07 g PON m⁻² yr⁻¹, 5.9±2.0 g BSi m⁻² yr⁻¹ and 1.7±0.6 g PIC m⁻² yr⁻¹. These biogenic fluxes clearly decreased with depth, and increased during “warm” years (1983 and 1987) of the El Niño, Southern Oscillation cycle (ENSO). Enhancement of annual mass flux rates to 3800 m was 49% in 1983 and 36% in 1987 above the decadal average, and was especially rich in biogenic Si. Biological events allowed estimates of sinking rates of detritus that range from 175 to 300 m d⁻¹, and demonstrate that, during periods of high productivity, particles sink quickly to deep ocean with less loss of organic components. Average POC flux into the deep ocean approximated the “canonical” 1% of the surface primary production.     

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.     

Do bottom mixed layers influence 234Th dynamics in the abyssal near-bottom water column?

Dynamics of the natural radioactive particle tracer ²³⁴Th (half-life: 24.1 days) within the abyssal water column up to 1000 m above bottom and within surface sediments of the northeast Atlantic (Porcupine Abyssal Plain; depth: ≈4845 m) were investigated. Distributions of transmissometer voltages and potential temperature indicated a subdivision of the near-bottom water column into a benthic mixed layer (BML; thickness: ≈10–65 m) and the layer above the BML up to the upper boundary of the bottom nepheloid layer (BNL; thickness: ≈1000 m). Comparison of ²³⁴Th fluxes (dpm m⁻²d⁻¹) in sediment traps, vertical fluxes derived from ²³⁴Th/²³⁸U-disequilibrium in the near-bottom water column and excess ²³⁴Th inventories in the surface sediment provided evidence for lateral advection of ²³⁴Th-depleted water and a ‘missing sink’ for ²³⁴Th. A simple one-dimensional steady-state box-model approach was applied to investigate ²³⁴Th dynamics. Estimated residence times suggest the BML and the resuspension zone of the surface sediment to be highly dynamic systems with respect to particle cycling and sorptive reactions on a time scale of weeks. Model results indicate that, through the chemical forcing of changing particle concentration, a thickening BML results in (1) increasing residence times of particulate ²³⁴Th in the BML with respect to the net fluxes across the upper boundary of the BML and into the surface sediment; (2) declining adsorption rate constants in the BML; (3) increasing desorptive fluxes in the BML resulting in (4) enhanced ²³⁴Th decay in the BML; (5) decreasing net fluxes of particulate ²³⁴Th from the BML to the upper BNL above the BML and to the sediment. Potential consequences for carbon cycling in the water column of the deep ocean are discussed.     

Biogeochemical responses to late-winter storms in the Sargasso Sea,III—Estimates of export production using 234Th:238U disequilibria and sediment traps

Direct measurements of new production and carbon export in the subtropical North Atlantic Ocean appear to be too low when compared to geochemical-based estimates. It has been hypothesized that episodic inputs of new nutrients into surface water via the passage of mesoscale eddies or winter storms may resolve at least some of this discrepancy. Here, we investigated particulate organic carbon (POC), particulate organic nitrogen (PON), and biogenic silica (BSiO₂) export using a combination of water column ²³⁴Th:²³⁸U disequilibria and free-floating sediment traps during and immediately following two weather systems encountered in February and March 2004. While these storms resulted in a 2–4-fold increase in mixed layer NO₃ inventories, total chlorophyll a and an increase in diatom biomass, the systems were dominated by generally low ²³⁴Th:²³⁸U disequilibria, suggesting limited particle export. Several ²³⁴Th models were tested, with only those including non-steady state and vertical upwelling processes able to describe the observed ²³⁴Th activities. Although upwelling velocities were not measured directly in this study, the ²³⁴Th model suggests reasonable rates of 2.2–3.7 m d^?1.Given the uncertainties associated with ²³⁴Th derived particle export rates and sediment traps, both were used to provide a range in sinking particle fluxes from the upper ocean during the study. ²³⁴Th particle fluxes were determined applying the more commonly used steady state, one-dimensional model with element/²³⁴Th ratios measured in sediment traps. Export fluxes at 200 m ranged from 1.91±0.20 to 4.92±1.22 mmol C m^?2 d^?1, 0.25±0.08 to 0.54±0.09 mmol N m^?2 d^?1, and 0.22±0.04 to 0.50±0.06 mmol Si m^?2 d^?1. POC export efficiencies (Primary Production/Export) were not significantly different from the annual average or from time periods without storms, although absolute POC fluxes were elevated by 1–11%. This increase was not sufficient, however, to resolve the discrepancy between our observations and geochemical-based estimates of particle export. Comparison of PON export rates with simultaneous measurements of NO₃^? uptake derived new production rates suggest that only a fraction, <35%, of new production was exported as particles to deep waters during these events. Measured bSiO₂ export rates were more than a factor of two higher (p<0.01) than the annual average, with storm events contributing as much as 50% of annual bSiO₂ export in the Sargasso Sea. Furthermore it appears that 65–95% (average 86±14%) of the total POC export measured in this study was due to diatoms.Combined these results suggest that winter storms do not significantly increase POC and PON export to depth. Rather, these storms may play a role in the export of bSiO₂ to deep waters. Given the slower remineralization rates of bSiO₂ relative to POC and PON, this transport may, over time, slowly decrease water column silicate inventories, and further drive the Sargasso Sea towards increasing silica limitation. These storm events may further affect the quality of the POC and PON exported, given the large association of this material with diatoms during these periods.     

Dissolved inorganic carbon,alkalinity, nutrient and oxygen seasonal and interannual variations at the Antarctic Ocean JGOFS-KERFIX site

JGOFS-KERFIX (KERguelen point FIXe) time-series station, located south of the polar front in the Indian sector of the Antarctic Ocean, was occupied monthly between January 1990 and March 1995. Annual cycles of dissolved inorganic carbon (DIC), total alkalinity (TALK), oxygen (O₂) and nutrients (nitrate, silicate, phosphate and ammonia) in the upper ocean are presented for this site. From seasonal drawdown of nutrients and DIC, we estimate a spring–summer net community production of 3.2±0.5 mol m⁻² and C/N/P ratios of 100/16/1. The Si/N ratio varies between 1.8 and 3, suggesting low iron concentrations. The spring–summer biogenic silicon export derived from silicate drawdown is 1.18 mol m⁻², consistent with model estimates of silicate export at this site. Seasonal and interannual variations of oxygen, nitrate and DIC due to physical and biological processes are quantified using a simple month-to-month budget formulation. From these budgets, an annual net community production of 5.7±3.3 mol m⁻² yr⁻¹ is estimated, about twice the averaged spring–summer production, indicating that, at KERFIX, there is a positive net community production throughout the year. Air–sea CO₂ fluxes show that KERFIX is a strong CO₂ sink for the atmosphere of 2.4–5.1 mol m⁻² yr⁻¹ in 1993, depending on the gas exchange formulation used. A 2.1–3.3 mol m⁻² yr⁻¹ outgassing of O₂ is observed at KERFIX except in 1993 and 1994 where a decreasing trend of temperature induces an increase of O₂ solubility.     

Vertical distribution of zooplankton and active flux across an anticyclonic eddy in the Canary Island waters

The vertical distribution (0–900 m) of zooplankton biomass and indices of feeding (gut fluorescence, GF) and metabolism (electron transfer system, ETS) were studied across an anticyclonic eddy south of Gran Canaria Island (Canary Islands). Two dense layers of organisms were clearly observed during the day, one above 200 m and the other at about 500 m, coincident with the deep scattering layer (DSL). The biomass displacement due to interzonal migrants in the euphotic zone was more than 2-fold higher than that previously reported for the southern area of this archipelago. The gut flux estimated (0.14–0.44 mgC m⁻² d⁻¹) was similar to the values previously found in the Canaries. The respiratory flux outside the eddy (1.85 mgC m⁻² d⁻¹) was in the lower range of values reported for this area. Inside the eddy, migrant biomass and respiration rates were 2- and 4- fold higher than in the surrounding waters. Active flux mediated by diel vertical migrants inside the eddy (8.28 mgC m⁻² d⁻¹) was up to 53% of the passive carbon flux to the mesopelagic zone (15.8 mgC m⁻² d⁻¹). It is, therefore, suggested that the anticyclonic eddy enhanced both migration from deep waters and active flux.     

Vertical distribution of zooplankton in Canary Island waters: implications for export flux

Zooplankton biomass, gut fluorescence and electron transfer system (ETS) activity were measured in vertical profiles (0–900 m) in two different size classes (<1 and >1 mm) in Canary Island waters. Both size fractions displayed a typical pattern of distribution with higher biomass, gut fluorescence and ETS in the shallower layers at night. By day, however, the vertical distribution varied between the size fractions, with higher biomass of the small fraction in the 0–200 m and a layer of large organisms at depth (∼500 m). For both size fractions, average ETS activity was higher by day than at night at depths between 200 and 600 m. Similarly, gut fluorescence was slightly higher by day below 200 m. The downward export of respiratory carbon was 1.92 and 4.29 mg C m⁻² d⁻¹ for samples obtained southwest of Gran Canaria Island and west of Tenerife Island respectively, being 2.68 mg C m⁻² d⁻¹ for the whole area. These values represented 16–45% (22–28% for the area) of the calculated passive particulate export production resulting from primary production. The estimated “gut flux” accounted for 0.35 (western zone) and 2.37 mg C m⁻² d⁻¹ (southwest of Gran Canaria), being 1.28 mg C m⁻² d⁻¹ for the whole area and represented between 3 and 25% (11–14% for the whole area) of the estimated passive particle export flux. These results agree with previous estimates and suggest that diel-migrant zooplankton can play an important role in the downward flux of carbon.     

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.     

Microbial respiration in the Levantine Sea: evolution of the oxidative processes in relation to the main Mediterranean water masses

First data on microbial respiration in the Levantine Sea are reported with the aim of assessing the distribution of oxidative processes in association with the main Mediterranean water masses and the changing physical structure determined by the Eastern Mediterranean Transient. Respiratory rates, in terms of metabolic carbon dioxide production, were estimated from measured electron transport system activities in the polygonal area of the Levantine Sea (32.5–36.5 N Latitude, 26.0–30.25 E Longitude) and at Station Geo’95, in the Ionian Sea (35°34.88 N; 17°14.99 E). At the Levantine Sea, the mean carbon dioxide production rate decreased from the upper to the deeper layers and varied from 22.0±12.4 μg C h⁻¹ m⁻³ in the euphotic layer to 1.30±0.5 μg C h⁻¹ m⁻³ in the depth range between 1600 and 3000 m. Significant differences were found among upper, intermediate and bottom layers. The euphotic zone supported a daily carbon dioxide production of 96.6 mg C d⁻¹ m⁻² while the aphotic zone (between 200 and 3000 m) sustained a 177.1 mg C d⁻¹ m⁻² carbon dioxide production. In Station Geo’95, the carbon dioxide production rates amounted to 170.4 and 102.2 mg C d⁻¹ m⁻² in the euphotic and aphotic zones, respectively. The rates determined in the identified water masses showed a tight coupling of respiratory processes and Mediterranean circulation patterns. The increasing respiratory rates in the deep layers of the Levantine Sea are explained by the introduction of younger waters recently formed in the Aegean Sea.