Relationships between picophytoplankton and optical properties in the Azores Front region in the Atlantic Ocean

The picophytoplankton evolved to become extremely effective harvesters of light and, thus, dominating productivity in the open ocean. In this study, their distribution in relation to the underwater light field was examined in the Azores Front region of the North Atlantic. In this region, attenuation coefficients of downward irradiance varied between 0.038 and 0.065 m^− 1. Maximum absorptions were at the red and green parts of the light spectrum, typical of the oligotrophic ocean. The euphotic zone ranged from 70 to 120 m. The deep chlorophyll maximum (DCM) was found at depths where subsurface light ranged between 0.1 and 1% of its surface values. Prochlorococcus was the dominant phytoplankton group while Synechococcus (cyanobacteria) and picoeukaryotes were much less abundant. The ability of chlorophyll to absorb light, i.e., the absorption coefficient of chlorophyll (a^⁎), was found to be dependent on the photoacclimation of the cells and was lower at low-light intensities. Due to a packaging effect and probable pigment changes, a^⁎ at the DCM was the lowest while the chlorophyll per Prochlorococcus cell was the highest. This is a major adaptation of the picophytoplankton to low light (less than 1% surface light), which allows them to bloom at the DCM. This study indicates that the Atlantic Ocean models for the estimation of phytoplankton concentration and/or primary production should not use a constant carbon biomass-to-chlorophyll ratio for phytoplankton based on phytoplankton size, and should take into account the variation in chlorophyll-absorption ability.     

Particulate organic carbon in the global ocean derived from SeaWiFS ocean color

Satellite remote sensing offers new means of quantifying particulate organic carbon, POC, concentration over large oceanic areas. From SeaWiFS ocean color, we derived 10-year data of POC concentration in the surface waters of the global ocean. The 10-year time series of the global and basin scale average surface POC concentration do not display any significant long-term trends. The annual mean surface POC concentration and its seasonal amplitude are highest in the North Atlantic and lowest in the South Pacific, when compared to other ocean basins. POC anomalies in the North Atlantic, North Pacific, and global concentrations seem to be inversely correlated with El Niño index, but longer time series are needed to confirm this relationship. Quantitative estimates of POC reservoir in the oceanic surface layer depend on the choice of what should represent this layer. Global average POC biomass is 1.34 g m^?2 if integrated over one optical depth, 3.62 g m^?2 if integrated over mixed layer depth, and up to 6.41 g m^?2 if integrated over 200-m layer depth (when assumed POC concentration below MLD is 20 mg m^?3). The global estimate of total POC reservoir in the surface 200-m layer of the ocean is 228.61×10¹³ g. We expect that future estimates of POC reservoir may be even larger, when more precise calculations account for deep-water organic-matter maxima in oligotrophic regions, and POC biomass located just below the seasonal mixed layer in spring and summer in the temperate regions.     

Carbon and biogenic silica export influenced by the Amazon River Plume: Patterns of remineralization in deep-sea sediments

The Amazon River Plume delivers freshwater and nutrients to an otherwise oligotrophic western tropical North Atlantic (WTNA) Ocean. Plume waters create conditions favorable for carbon and nitrogen fixation, and blooms of diatoms and their diazotrophic cyanobacterial symbionts have been credited with significant CO₂ uptake from the atmosphere. The fate of the carbon, however, has been measured previously by just a few moored or drifting sediment traps, allowing only speculation about the full extent of the plume's impact on carbon flux to the deep sea. Here, we used surface (0.5 m) sediment cores collected throughout the Demerara Slope and Abyssal Plain, at depths ranging from 1800 to 5000 m, to document benthic diagenetic processes indicative of carbon flux. Pore waters were extracted from sediments using both mm- and cm-scale extraction techniques. Profiles of nitrate (NO₃) and silicate (Si(OH)₄) were modeled with a diffusion-reaction equation to determine particulate organic carbon (POC) degradation and biogenic silica (bSi) remineralization rates. Model output was used to determine the spatial patterns of POC and bSi arrival at the sea floor. Our estimates of POC and Si remineralization fluxes ranged from 0.16 to 1.92 and 0.14 to 1.35 mmol m⁻² d⁻¹, respectively. A distinct axis of POC and bSi deposition on the deep sea floor aligned with the NW axis of the plume during peak springtime flood. POC flux showed a gradient along this axis with highest fluxes closest to the river mouth. bSi had a more diffuse zone of deposition and remineralization. The impact of the Amazon plume on benthic fluxes can be detected northward to 10°N and eastward to 47°W, indicating a footprint of nearly 1 million km². We estimate that 0.15 Tmol C y⁻¹ is remineralized in abyssal sediments underlying waters influenced by the Amazon River. This constitutes a relatively high fraction (~7%) of the estimated C export from the region.; the plume thus has a demonstrable impact on C_org export in the western Atlantic. Benthic fluxes under the plume were comparable to and in some cases greater than those observed in the eastern equatorial Atlantic, the southeastern Atlantic, and the Southern Ocean.     

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.     

The distribution and speciation of mercury in the South and equatorial Atlantic

Mercury concentration and speciation were measured in surface and deep ocean waters of the South and equatorial Atlantic. In the surface waters, total Hg was 2.9±1.7 pM on average, with a significant fraction present as reactive Hg (1.7±1.2 pM). The reactive Hg fraction consisted of elemental Hg (Hg°) as the dominant species (1.2±0.8 pM). Measurements in surface waters also showed that Hg partitioned to the “colloidal” phase (0.33±0.28 pM) and was associated with particulate matter (0.1±0.05 pM). No dimethylmercury (DMHg;<0.01 pM) or monomethylmercury (MMHg; <0.05 pM) was detected in mixed layer samples. The highest DMHg concentrations were found in recently formed deep waters – Antarctic Intermediate Water and Antarctic Bottom Water and in the equatorial sub-thermocline region. Higher concentrations of DMHg coincided with higher values of apparent oxygen utilization, indicative of a link between microbial activity and methylated Hg production. The lowest-deep water DMHg concentrations were found in the core of the North Atlantic Deep Water. Incubation experiments on-board demonstrated that light enhanced the decomposition of DMHg, with MMHg as the major product. In deep waters, Hg° was still an important constituent and is likely formed as a decomposition product from MMHg. These results suggest that methylated Hg production occurs primarily in regions of high biological activity, and that ionic Hg is strongly complexed to organic matter (colloidal material) in open ocean surface waters.     

The diffusive component of particulate organic carbon export in the North Atlantic estimated from SeaWiFS ocean color

The diffusive component of the particulate organic carbon (POC) export from the ocean's surface layer has been estimated using a combination of the mixed layer model and SeaWiFS ocean color data. The calculations were carried out for several example sites located in the North Atlantic over a 10-year time period (1998–2007). Satellite estimates of surface POC derived from ocean color were applied as an input to the model driven by local surface heat and momentum fluxes. For each year of the examined period, the diffusive POC flux was estimated at a 200 m depth. The highest flux is generally observed in the spring and fall seasons, when surface waters are weakly stratified. In addition, the model results demonstrate significant interannual and geographical variability of the flux. The highest diffusive POC flux occurs in the northern North Atlantic and the lowest in the subtropical region. The interannual variability of the diffusive POC flux is associated with mixed layer dynamics and underscores the importance of atmospheric forcing for POC export from the surface layer to the ocean's interior.     

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.     

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.     

Factors influencing the abundance of deep pelagic bioluminescent zooplankton in the Mediterranean Sea

Measurements of the density of deep pelagic bioluminescent zooplankton (BL) were made with the Intensified Silicon Intensifier Target (ISIT) profiler in the Ligurian, Tyrrhenian, Adriatic, Ionian Seas and the Strait of Sicily from ～300 m to near seafloor. Mean BL densities ranged from 2.61 m^?3 at 500–1000 m depth in the Adriatic Sea to 0.01 m^?3 at 4000–5000 m depth in the E Ionian Sea. We investigated drivers of spatial variation of deep pelagic BL density. Linear regression was applied between surface chlorophyll a (Chl a) concentration and underlying BL density. Chl a values were determined from satellite derived 100 km radius composites (six 10-day means per ISIT deployment, over preceding 60 days). At 500–1000 m depth we found a significant positive relationship between mean BL density and mean Chl a in the period prior to 0–10 days (at 1% level) and prior to 10–40 days (at 5% level). Beyond 40 days no relationship between BL density and Chl a was found at this depth. At depths 1000–1500 m BL density values were low and no significant relationship with Chl a was detected. Generalised additive modelling (GAM) was used to assess the influence of benthic hotspots (seamount; cold water coral mound; mud volcano) on overlying BL density. A reduction in BL density was found downstream of the Palinuro seamount from 300 to 600 m. No effect on BL density in the overlying water column was detected from the presence of cold water corals. Higher BL densities were detected over the W Madonna dello Ionio mud volcano than at other sites sampled in the NW Ionian Sea. We find surface Chl a to be a good predictor of BL density in the mesopelagic zone; below this depth we hypothesise that processes affecting the efficiency of particle export to deep water may exert greater influence on BL density.     

Molecular and isotopic constraints on the sources of suspended particulate organic carbon on the northwestern Atlantic margin

The abundance, carbon isotopic composition (Δ¹⁴C and δ¹³C), and lipid biomarker (alkenones and saturated fatty acids) distributions of suspended particulate organic matter were investigated at three stations centered on the 2000, 3000, and 3500 m isobaths over the New England slope in order to assess particulate carbon sources and dynamics in this highly productive and energetic region. Transmissometry profiles reveal that particle abundances exhibit considerable fine structure, with several distinct layers of elevated suspended particulate matter concentration at intermediate water depths in addition to the presence of a thick bottom nepheloid layer at each station. Excluding surface water samples, the Δ¹⁴C values of particulate organic carbon (POC) indicated the presence of a pre-aged component in the suspended POC pool (Δ¹⁴C<+38‰). The Δ¹⁴C values at the 3000 m station exhibited greater variability and generally were lower than those at the other two stations where the values decreased in a more systematic matter with increasing sampling depth. These lower Δ¹⁴C values were consistent with higher relative abundances of terrigenous long-chain fatty acids at this station than at the other two stations. Two scenarios were considered regarding the potential provenances of laterally transported POC: cross-shelf transport of shelf sediment (Δ¹⁴C=?140‰) and along-slope transport of the slope sediment proximal to the sampling locations (Δ¹⁴C=?260‰). Depending on the scenario, isotopic mass balance calculations indicate allochthonous POC contributions ranging between 15% and 54% in the meso- and bathy-pelagic zone, with the highest proportions at the 3000 m station. Alkenone-derived temperatures recorded on suspended particles from surface waters closely matched in-situ temperatures at each station. However, alkenone-derived temperatures recorded on particles from the subsurface layer down to 250 m were lower than those of overlying surface waters, especially at the 3000 m station, implying supply of phytoplankton organic matter originally produced in cooler surface waters. AVHRR images and temperature profiles indicate that the stations were under the influence of a warm-core ring during the sampling period. The low alkenone-derived temperatures in the subsurface layer coupled with the lower Δ¹⁴C values for the corresponding POC suggests supply of OC on resuspended sediments underlying cooler surface waters distal to the study area, possibly further north or west. Taken together, variations in Δ¹⁴C values, terrigenous fatty acid abundances, and alkenone-derived temperatures among the stations suggest that input of laterally advected OC is a prominent feature of POC dynamics on the NW Atlantic margin, and is spatially heterogeneous on a scale smaller than the distance between the stations (<150 km).     

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.     

Spatial and temporal variability of POC in the northeast Subarctic Pacific

Particulate organic carbon (POC) concentrations from 0 to 1000 m were quantified in size-fractionated particulate matter samples obtained by the multiple unit large volume in situ filtration system (MULVFS) in 1996 and 1997 along the 1600 km long “line P” transect from continental slope waters near southern Vancouver Island to Ocean Station PAPA (OSP, 50°N, 145°W). Regression of in situ POC vs. beam attenuation coefficient, c, from a simultaneously deployed 1-m pathlength SeaTech transmissometer gave slope, intercept and r² values of 6.15±0.19×10⁻⁵ m⁻¹ (nmol C l⁻¹)⁻¹, 0.363±0.003 m⁻¹, and 0.951 (n=145), respectively. This result agreed within several percent of calibrations obtained from two 2600-km-long transects of the equatorial Pacific in 1992 (Bishop, 1999). Data from other, more frequently deployed transmissometers were standardized against the 1-m instrument, and the combined optical data set was used to document POC variability at finer spatial and temporal scales than could be sampled directly using either conventional water bottle casts or MULVFS. Published bottle POC vs. c relationships show much more variability and remain problematic. Along the line P transect in the salinity-stratified upper 100 m, POC isolines shoaled from winter to summer in concert with seasonal stratification. At the same time, POC was progressively enriched in subeuphotic zone waters to depths greater than 500 m. Near-surface POC fields sampled in the winter time showed strong temporal POC variability over time scales of days as well as between years. POC concentrations at OSP in February 1996 were higher than those found at any other time of year. Less variability was found along line P in other seasons. In May 1996, kilometer-scale spatial variability of POC at OSP was small; dawn vs. dusk variations of c were used to calculate 0–100 m POC turnover times shorter than 6 d. Calculations also suggest that 25–50% of primary productivity was expressed as dissolved organic carbon at OSP in May 1996.     

Seasonal POC fluxes at BATS estimated from 210Po deficits

In this study at the Bermuda Atlantic Time-series Study (BATS) site we demonstrate that the polonium–lead disequilibrium system may perform better as a tracer of organic carbon export under low-flux conditions (in this case, <2.5 mmol C m^?2 d^?1) than under bloom conditions in an oligotrophic setting. With very few exceptions, the POC flux predictions calculated from the water-column ²¹⁰Po deficit were within a factor of 2 of the POC flux caught in surface-tethered sediment traps. However, we found higher correlation between size-fractionated particulate ²¹⁰Po activity and POC concentration in November 2006 (r=0.93) than in January (r=0.79) and during the spring bloom in March 2007 (r=0.80). We suggest that this is due to the ability of polonium to distinguish between bulk mass flux and organic carbon export under oligotrophic and lithogenic-driven flux regimes. Further, we found that the POC/Po ratio on particles was largely independent of size class between 10 and 100 μm (P=0.13) during each season, supporting the notion that export in this oligotrophic system is driven by sinking aggregates of smaller cells and not by large, individual cells.     

Biogeochemical variability in the southern Ross Sea as observed by a glider deployment

High-resolution autonomous glider data (including temperature, salinity, fluorescence, and optical backscatter) collected during the 2010–2011 austral summer identified variations in phytoplankton biomass along two glider sections near 76°40′S. Sea surface temperatures were warmer during the latter, westward section, while mixed layer depths were deeper. Substantial quantities of Modified Circumpolar Deep Water, identified by neutral density criteria, were located within both sections. Chlorophyll (Chl) concentrations computed from fluorescence exhibited daily quenching near the surface, and deep chlorophyll concentrations at 200 m became periodically elevated, suggesting substantial export on small space and time scales. The concentrations of particulate organic carbon (POC) computed from backscatter increased abruptly during the latter, westward section, concurrent with a decrease in chlorophyll. These higher POC:Chl ratios were not strongly correlated with presence of MCDW or with shallower mixed layer depths, but were strongly associated with higher surface temperatures and wind speed. The observed POC:Chl increase suggests a marked spatial and temporal transition between a Phaeocystis antarctica-dominated assemblage characterized by modest POC:Chl ratios to a diatom-dominated assemblage. Finally, a subsampling analysis highlights the capability of high-resolution glider data to resolve these biological/physical parameter correlations that are not discernible from lower frequency data typical of traditional cruise stations.     

Dissolved organic carbon and apparent oxygen utilization in the Atlantic Ocean

Dissolved organic carbon (DOC) distributions along two Atlantic Meridional Transects conducted in 2005 in the region between 47°N and 34°S showed clear latitudinal patterns. The DOC concentrations in the epipelagic zone (0–100 m) were the highest (70–90 µM) in tropical and subtropical waters with stable mixed layers, and lowest (50–55 µM) at the poleward extremities of the transects due to deep convective mixing supplying low DOC waters to the surface. A decrease in DOC occurred with depth, and lowest DOC concentrations (41–45 µM) in the 100–300 m depth range were observed in the equatorial region due to upwelling of low DOC waters. A strong relationship between DOC and AOU was observed in the σ–t 26–26.5 isopycnal layer which underlies the euphotic zone and outcrops at the poleward extremities of the North and South Atlantic Subtropical Gyres (NASG and SASG) in the region ventilating the thermocline waters. Our observations reveal significant north–south variability in the DOC–AOU relationship. The gradient of the relationship suggests that 52% of the AOU in the σ–t 26–26.5 density range was driven by DOC degradation in the NASG and 36% in the SASG, with the remainder due to the remineralisation of sinking particulate material. We assess possible causes for the greater contribution of DOC remineralisation in the NASG compared to the SASG.     

A phytoplankton class-specific primary production model applied to the Kerguelen Islands region (Southern Ocean)

As part of the KErguelen: compared study of the Ocean and the Plateau in Surface water (KEOPS) project in late summer 2005, we examine the phytoplankton community composition and associated primary production in the waters surrounding the Kerguelen Archipelago, with the emphasis on two contrasted environments: (i) the Kerguelen Plateau, where a large bloom occurs annually, and (ii) the high-nutrient low-chlorophyll (HNLC) offshore waters. A biomarker pigment approach was used to assess the community composition in terms of chlorophyll biomass of three phytoplankton size classes, namely micro-, nano-, and picophytoplankton. The second objective was to evaluate a global class-specific approach for estimating the contribution of the three pigment-based size classes to the primary production in the study area. To do so, primary production rates associated with each phytoplankton class were computed from the class-specific chlorophyll biomass coupled to a class-specific primary production model, and compared with in situ measurements of size-fractionated ¹³C-based primary production. The iron-enriched bloom region was dominated by microphytoplankton (diatoms), which contributed 80–90% to the total primary production (of ≈1 g C m^?2 d^?1). In the HNLC area, the primary production was about 0.30 g C m^?2 d^?1, mainly (65%) achieved by small diatoms and nanoflagellates. The model results show a good overall agreement between predicted and measured total primary production rates. In terms of size classes, agreements were higher for the bloom region than for the HNLC waters. Discrepancies in this complex iron-limited area may be explained essentially by the smaller size of diatoms, or a different set of photophysiological properties.     

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.     

Vertical distribution of phytoplankton biomass,production and growth in the Atlantic subtropical gyres

Ninety-four stations were sampled in the Atlantic subtropical gyres during 10 cruises carried out between 1995 and 2001, mainly in boreal spring and autumn. Chlorophyll a (Chl-a) and primary production were measured during all cruises, and phytoplankton biomass was estimated in part of them. Picoplankton (<2 μm) represented >60% of total Chl-a concentration measured at the surface, and their contribution to this variable increased with depth. Phytoplankton carbon concentrations were higher in the upper metres of the water column, whereas Chl-a showed a deep maximum (DCM). At each station, the water column was divided into the upper mixed layer (ML) and the DCM layer (DCML). The boundary between the two layers was calculated as the depth where Chl-a concentration was 50% of the maximum Chl-a concentration. On average DCML extends from 67 to 126 m depth. Carbon to Chl-a (C:Chl-a) ratios were used to estimate phytoplankton carbon content from Chl-a in order to obtain a large phytoplankton carbon dataset. Total C:Chl-a ratios averaged (±s.e.) 103±7 (n=22) in the ML and 24±4 (n=12) in the DCML and were higher in larger cells than in picoplankton. Using these ratios and primary production measurements, we derived mean specific growth rates of 0.17±0.01 d⁻¹ (n=173) in the ML and 0.20±0.01 d⁻¹ (n=165) in the DCML although the differences were not significant (t-test, p>0.05). Our results suggest a moderate contribution of the DCML (43%) to both phytoplankton biomass and primary production in the Atlantic subtropical gyres.     

Abundance and size distribution of transparent exopolymer particles (TEP) in a coccolithophorid bloom in the northern Bay of Biscay

The distribution of transparent exopolymer particles (TEP) was investigated during a coccolithophorid bloom in the northern Bay of Biscay (North Atlantic Ocean) in early June 2006. MODIS chlorophyll-a (Chl-a) and reflectance images before and during the cruise were used to localize areas of important biological activity and high reflectance (HR). TEP profiles along the continental margin, determined using microscopic (TEP_micro) and colorimetric (TEP_color) methods, showed abundant (6.1×10⁶–4.4×10⁷ L^?1) and relatively small (0.5–20 μm) particles, leading to a low total volume fraction (0.05–2.2 ppm) of TEP_micro and similar vertical profiles of TEP_color. Estimates of carbon content in TEP (TEP-C) derived from the microscopic approach yielded surface concentration of 1.50 μmol C L^?1. The contribution of TEP-C to particulate organic carbon (POC) was estimated to be 12% (molar C ratio) during this survey. Our results suggest that TEP formation is a probable first step to rapid and efficient export of C during declining coccolithophorid blooms.