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.     

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.     

Zooplankton metabolism and carbon demand at two seamounts in the NE Atlantic

Zooplankton metabolic rates, determined from electron transfer system (ETS) activity, were studied at two seamounts (Seine: 34°N, 14°W, summit depth ∼170 m; Sedlo: 40°N, 27°W, summit depth ∼750 m) in the northeast (NE) Atlantic during three cruises in November 2003, April 2004 and July 2004. ETS activity and respiratory carbon demand were measured for samples taken at seamount and open-ocean locations in order to probe the hypothesis of locally enhanced seamount productivity. ETS activity and biomass revealed no consistent diel patterns of feeding activity and vertical migration at Seine and Sedlo Seamounts. Spatial differences of biomass-specific ETS activity were observed at both seamounts and coincided with differences in food abundance and quality. At Seine Seamount in April 2004, biomass-specific ETS activity was on average higher at the seamount locations compared to the open ocean, though the enhancement was of a lower magnitude than spatial and temporal variability and had no apparent influence on zooplankton respiratory carbon demand or biomass. A persistent pattern of reduced zooplankton biomass above the summit location at Seine Seamount in April 2004 and July 2004 resulted in a local reduction of respiratory carbon demand. At Sedlo Seamount in November 2003, large spatial differences in biomass-specific ETS activity observed at the seamount locations resulted in a large range of respiratory carbon demand at the seamount, but were not reflected in zooplankton biomass. The depth-integrated (0–150 m) median respiratory carbon demand of the zooplankton community estimated from day and night hauls was 2.1 mg C m⁻² d⁻¹ at Seine Seamount (range: 0.3–6.3) and 2.9 mg C m⁻² d⁻¹ at Sedlo Seamount (range: 1.6–12.0). The sporadic nature and low magnitude of locally higher zooplankton respiration rates at the seamounts, which did not result in locally higher zooplankton standing stock biomass, lead us to reject the hypothesis that locally enhanced seamount productivity provides an autochthonous food supply to the resident faunas at Seine and Sedlo Seamounts. Instead, we conclude that the faunas at both seamounts are more likely supported by advection of food from the surrounding ocean.     

Nutrient and phytoplankton dynamics off the west coast of Vancouver Island during the 1997/98 ENSO event

Six research cruises were conducted off the west coast of Vancouver Island between April and October of 1997 and 1998 as part of the Canadian GLOBEC project to compare nutrient and phytoplankton dynamics between ENSO (1997) and non-ENSO (1998) years. Limited sampling also was conducted during three cruises in 1999. During the 1997 ENSO period, there was a shallow thermocline (∼10 m) that resulted in a shallower mixed layer, lower salinity and density, and stronger summer stratification. In general on the shelf, the 1997 growing season was characterized by higher nitrate (7.5 μM) and silicic acid (17 μM) concentrations, lower total chlorophyll (∼76 mg m⁻²), lower phytoplankton carbon biomass (0.2 mg C L⁻¹), and lower diatom abundance and biomass than in 1998. Phytoplankton assemblages were dominated by nanoplankton in 1997 and by diatoms in 1998. These results suggest that the 1997 ENSO was responsible for a reduction in the growth and biomass of larger phytoplankton cells. In mid-1998, the hydrographic characteristics off the west coast of Vancouver Island changed suddenly. The 1997 poleward transport of warm water reversed to an equatorward transport of coastal water in July 1998, which was accompanied by normal summer upwelling. During 1998, a large diatom bloom (mainly dominated by Chaetoceros debilis, Leptocylindrus danicus and to a lesser extent by Skeletomema and Pseudo-nitzschia sp.) was observed in July over the continental shelf. This large bloom resulted in chlorophyll concentrations of up to 400 mg m⁻², primary productivity of up to 11 g C m⁻² d⁻¹, and near undetectable dissolved nitrogen concentrations at some of the shelf stations in 1998. In contrast, during 1997, the sub-tropical waters that were advected over the slope, resulted in low chlorophyll a and primary productivity (generally <1 g C m⁻² d⁻¹). Therefore, there was a sharp contrast between the very high primary productivity on the shelf in July 1998, due to normal nutrient replenishment from summer upwelling and outflow from the Strait of Juan de Fuca, and the lower primary productivity during the 1997 ENSO year. During 1998, non-ENSO conditions resulted in phytoplankton biomass that was twice as high on the shelf as that measured in regions beyond the continental shelf of the west coast of Vancouver Island.     

Episodic particulate fluxes at southern temperate mid-latitudes (42–45°S) in the Subtropical Front region,east of New Zealand

Sediment traps were deployed for almost 1 yr at two sites near 178°40′E in 1996–1997 on Chatham Rise (New Zealand). These sites were either side of the Subtropical Front (STF), which is a biologically productive zone, characterised by moderate atmospheric CO₂ uptake. At each site, PARFLUX sediment traps (Mk 7G–21) were deployed at 300 and 1000 m in 1500 m water depth. At 42°42′S, north of the STF, approximately 80% of the integrated total mass, POC and biogenic silica flux at 300 m occurred in a 7-day pulse in austral mid-spring (1064, 141 and 6 mg m⁻² d⁻¹, respectively, in early October). This pulse was recorded a week later in the 1000 m trap, indicating a particle sinking rate of 100 m d⁻¹. In contrast, at 44°37′S, south of the STF, the main flux of total mass and biogenic silica occurred 3 weeks later in late spring (289 and 3 mg m⁻² d⁻¹, respectively, in early November). Organic carbon, nitrogen and phosphorus fluxes were persistently high over spring at the southern site, although total POC flux integrated over 3 months was only 60 mg m⁻² d⁻¹. Thus, up to 2–3 times more material was exported north of the STF, compared with fluxes measured <200 km away to the south. As an integrated proportion of the annual total mass flux, however, more organic carbon was exported south of the STF (17% cf. 5–14%). Furthermore, organic material exported in spring from southern waters was labile and protein-rich (C : N — 8–16, C : P — 200–450, N : P — 13–36), compared to the more refractory, diatom-dominated material sinking out north of the STF in spring (C : N 9–22, C : P 50–230, N : P 5–19). These observations are consistent with anomalously high benthic biomass and diversity observed on south Chatham Rise. Resuspension and differential particle settling are probable causes for depth increases in particulate flux. Estimated particle source areas may be up to 120 km away due to high levels of mesoscale activity and mean flow in the STF region.     

Atmospheric input of dissolved inorganic phosphorus and silicon to the coastal northwestern Mediterranean Sea: Fluxes,variability and possible impact on phytoplankton dynamics

One hundred twelve rainwater samples collected from 1986 to 2003 at the signal station of Cap Ferrat (France, NW Mediterranean coast) were analysed for phosphate and silicate contents. This sampling site is affected by a European urban-dominated background material, with episodic Saharan dust inputs. The input of dissolved inorganic phosphorus (DIP) and dissolved inorganic silicon (DISi) was calculated. The most significant loadings of DIP and DISi were selected in order to assess their potential impact on phytoplankton dynamics, particularly in oligotrophic conditions, when surface waters are nutrient-depleted. The theoretical new production triggered by DIP and DISi inputs (NP_atmo) was estimated through Redfield calculations. The maximum theoretical DIP-triggered NP_atmo was up to 670 mg C m⁻² in October, at the end of the oligotrophic period (135 mg C m⁻³ in the 5 m-thick surface layer). During the same period, the daily integrated primary production measured at the DYFAMED site (NW Mediterranean Sea) was on average 219 mg C m⁻² d⁻¹ within the 0–100 m depth water column, while the mean daily primary production in the 5 m-thick surface layer was 1.6 mg C m⁻³ d⁻¹. However, high NP_atmo due to high DIP inputs might be episodically limited by lower DISi inputs, which may consequently lead to episodic preferential growth of non-siliceous phytoplanktonic species.     

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.     

Physical drivers of phytoplankton production in the southern Benguela upwelling system

Investigations of primary production (PP) were undertaken in the southern Benguela ecosystem during two research surveys in October 2006 and May 2007. Significant differences in environmental conditions, as well as biomass and PP, were observed between October and May. During October, integrated biomass and PP were significantly higher, ranging from 20.43 to 355.01 mg m⁻², and 0.71 to 6.98 g C m⁻² d⁻¹, respectively, than in May, where the range was 47.92–141.79 mg m⁻², and 0.70–3.35 g C m⁻² d⁻¹, respectively. Distribution patterns indicated low biomass and PP in newly upwelled water along the coast, higher biomass and PP in the mid-shelf region, while lower values were observed at and beyond the shelf edge. Latitudinal variations showed consistently higher biomass and PP in the St. Helena Bay region compared to biomass and PP south of Cape Town. During both surveys, phytoplankton communities were comprised primarily of diatoms and small flagellates, with no significant differences. Phytoplankton adaptation to environmental variability was characterised by increased P_m^B and E_k under elevated temperatures and irradiance, while no clear relationships were evident for α^B. Generalised Additive Models (GAMs) showed that photosynthetic parameters were all significant predictors of photosynthesis rates (P_z), with P_m^B being the most important, accounting for 36.97% of the deviance in P_z. However, biomass levels and environmental conditions exerted a much greater influence on P_z, with irradiance explaining the largest proportion (68.24%) of the deviance. Multiple predictor GAMs revealed that 96.26% of the deviance in P_z could be explained by a model which included nitrate, chlorophyll a, and irradiance.     

Organic carbon flux and remineralization in surface sediments from the northern North Atlantic derived from pore-water oxygen microprofiles

Organic carbon fluxes through the sediment/water interface in the high-latitude North Atlantic were calculated from oxygen microprofiles. A wire-operated in situ oxygen bottom profiler was deployed, and oxygen profiles were also measured onboard (ex situ). Diffusive oxygen fluxes, obtained by fitting exponential functions to the oxygen profiles, were translated into organic carbon fluxes and organic carbon degradation rates. The mean C_org input to the abyssal plain sediments of the Norwegian and Greenland Seas was found to be 1.9 mg C m⁻² d⁻¹. Typical values at the seasonally ice-covered East Greenland continental margin are between 1.3 and 10.9 mg C m⁻² d⁻¹ (mean 3.7 mg C m⁻² d⁻¹), whereas fluxes on the East Greenland shelf are considerably higher, 9.1–22.5 mg C m⁻² d⁻¹. On the Norwegian continental slope C_org fluxes of 3.3–13.9 mg C m⁻² d⁻¹ (mean 6.5 mg C m⁻² d⁻¹) were found. Fluxes are considerably higher here compared to stations on the East Greenland slope at similar water depths. By repeated occupation of three sites off southern Norway in 1997 the temporal variability of diffusive O₂ fluxes was found to be quite low. The seasonal signal of primary and export production from the upper water column appears to be strongly damped at the seafloor. Degradation rates of 0.004–1.1 mg C cm⁻³ a⁻¹ at the sediment surface were calculated from the oxygen profiles. First-order degradation constants, obtained from C_org degradation rates and sediment organic carbon content, are in the range 0.03–0.6 a⁻¹. Thus, the corresponding mean lifetime of organic carbon lies between 1.7 and 33.2 years, which also suggests that seasonal variations in C_org flux are small. The data presented here characterize the Norwegian and Greenland Seas as oligotrophic and relatively low organic carbon deep-sea environments.     

High primary productivity and f-ratio in summer in the Ulleung basin of the East/Japan Sea

To better understand the cause of high summer primary productivity in the Ulleung Basin located in the southwest part of the East/Japan Sea, the spatial dynamics of primary, new, and regenerated productivities (PP, NP, and RP) were examined along the path of the Tsushima Warm Current system in summer 2008. We compared hydrographic and chemical parameters in the Ulleung Basin with those of the Kuroshio Current in the Western Pacific Ocean and the East China Sea. In summer, integrated primary productivity (IPP, 0.37–0.96 g C m⁻² d⁻¹) and integrated new productivity (INP, 26–221 mg N m⁻² d⁻¹) within the euphotic zone in the Ulleung Basin were higher than those in the East China Sea and the Western Pacific Ocean (0.17–0.28 g C m⁻² d⁻¹, 2−5 mg N m⁻² d⁻¹, respectively). In contrast, there was no pronounced spatial variation in integrated regenerated productivity (IRP, 43–824 mg N m⁻² d⁻¹). Strong positive correlations between IPP and INP (also the f-ratio), and between nitrate uptake rate in the mixed layer and nitrate upward flux through the top of pycnocline in summer in the Ulleung Basin imply that the high IPP was mainly supported by supply of nitrate from the underlying water in the euphotic zone. Shallowing of the pycnocline depth as the current enters the East/Japan Sea facilitates nitrate supply from the nutrient-replete cold water immediately below the pycnocline through nitrate upward flux. A subsurface maximum in PP at or above the pycnocline and a high f-ratio further support the importance of this source of nitrate for maintaining the high summer PP in the Ulleung Basin. In comparison, the high PP layer was observed at the surface in the following fall and spring in the Ulleung Basin. Our results demonstrate the importance of hydrographic features in enhancing PP in this oligotrophic Tsushima Warm Current system.     

Downward transport of organic carbon by diel migratory micronekton in the western equatorial Pacific:: its quantitative and qualitative importance

Using simultaneous sampling with a commercial-sized trawl, a zooplankton net, and a sediment trap, we evaluated the contribution of vertically migrating micronekton to vertical material transport (biological pump) at two stations (3°00′N, 146°00′E and 3°30′N, 145°20′E) in the western equatorial North Pacific. The gravitational sinking particulate organic carbon flux out of the euphotic zone was 54.8 mg C m⁻² day⁻¹. The downward active carbon flux by diel migrant mesozooplankton was 23.53 and 9.97 mg C m⁻² day⁻¹, and by micronekton 4.40 and 2.26mg C m⁻² day⁻¹ at the two stations. Assuming that the micronekton sampling efficiency of the trawl was 14%, we corrected the downward carbon flux due to micronekton respiration to 29.9 and 15.2mg C m⁻² day⁻¹, or 54.6 and 27.7% of the sinking particle flux at the two stations. The corrected micronekton gut fluxes were 1.53 and 0.97mg C m⁻² day⁻¹. The role of myctophid fish fecal matter as a possible food resource for deep-sea organisms, based on its fatty acid and amino acid analysis, is discussed.     

Sea-ice algae: Major contributors to primary production and algal biomass in the Chukchi and Beaufort Seas during May/June 2002

Sea-ice and water samples were collected at 14 stations on the shelves and slope regions of the Chukchi and Beaufort Seas during the spring 2002 expedition as part of the Shelf–Basin Interaction Studies. Algal pigment content, particulate organic carbon and nitrogen, and primary productivity were estimated for both habitats based on ice cores, brine collection and water samples from 5-m depth. The pigment content (0.2–304.3 mg pigments m⁻²) and primary productivity (0.1–23.0 mg C m⁻³ h⁻¹) of the sea-ice algae significantly exceeded water-column parameters (0.2 and 1.0 mg pigments m⁻³; <0.1–0.4 mg C m⁻³ h⁻¹), making sea ice the habitat with the highest food availability for herbivores in early spring in the Chukchi and Beaufort Seas. Stable isotope signatures for ice and water samples did not differ significantly for δ¹⁵N, but for δ¹³C (ice: −25.1‰ to −14.2‰; water: −26.1‰ to −22.4‰). The analysis of nutrient concentrations and the pulse-amplitude-modulated fluorescence signal of ice algae and phytoplankton indicate that nutrients were the prime limiting factor for sea-ice algal productivity. The estimated spring primary production of about 1–2 g C m⁻² of sea-ice algae on the shelves requires the use of substantial nutrient reservoirs from the water column.     

Standing stocks,production, and respiration of phytoplankton and heterotrophic bacteria in the western Arctic Ocean

Standing stocks and production rates for phytoplankton and heterotrophic bacteria were examined during four expeditions in the western Arctic Ocean (Chukchi Sea and Canada Basin) in the spring and summer of 2002 and 2004. Rates of primary production (PP) and bacterial production (BP) were higher in the summer than in spring and in shelf waters than in the basin. Most surprisingly, PP was 3-fold higher in 2004 than in 2002; ice-corrected rates were 1581 and 458 mg C m⁻² d⁻¹, respectively, for the entire region. The difference between years was mainly due to low ice coverage in the summer of 2004. The spatial and temporal variation in PP led to comparable variation in BP. Although temperature explained as much variability in BP as did PP or phytoplankton biomass, there was no relationship between temperature and bacterial growth rates above about 0 °C. The average ratio of BP to PP was 0.06 and 0.79 when ice-corrected PP rates were greater than and less than 100 mg C m⁻² d⁻¹, respectively; the overall average was 0.34. Bacteria accounted for a highly variable fraction of total respiration, from 3% to over 60% with a mean of 25%. Likewise, the fraction of PP consumed by bacterial respiration, when calculated from growth efficiency (average of 6.9%) and BP estimates, varied greatly over time and space (7% to >500%). The apparent uncoupling between respiration and PP has several implications for carbon export and storage in the western Arctic Ocean.     

Ascending and descending particle flux from hydrothermal plumes at Endeavour Segment,Juan de Fuca Ridge

Bio-acoustic surveys and associated zooplankton net tows have documented anomalously high concentrations of zooplankton within a 100 m layer above the hydrothermal plumes at Endeavour Segment, Juan de Fuca Ridge. These and other data suggest that congregating epi-plume zooplankton are exploiting a food substrate associated with the hydrothermal plume. Ascending, organic-rich particles could provide a connection. Consequently, two paired sequentially sampling ascending and descending particle flux traps and a current meter were deployed on each of three moorings from July 1994 to May 1995. Mooring sites included an on-axis site (OAS; 47°57.0′N, 129°05.7′W) near the main Endeavour vent field, a “down-current” site 3 km west of the main vent field (WS), and a third background station 43 km northeast of the vent field (ES). Significant ascending and descending particle fluxes were measured at all sites and depths. Lipid analyses indicated that ascending POC was derived from mid-depth and deep zooplankton whereas descending POC also contained a component of photosynthetically derived products from the sea surface. Highest ascending POC fluxes were found at the hydrothermal plume-swept sites (OAS and WS). The limited data available, however, precludes an unequivocal conclusion that hydrothermal processes contribute to the ascending flux of organic carbon at each site. Highest ascending to descending POC flux ratios were also found at WS. Observed trends in POC, PMn/PTi, and PFe/PTi clearly support a hydrothermal component to the descending flux at the plume-swept WS site (no descending data was recovered at OAS) but not at the background ES site. Alternative explanations for ascending particle data are discussed. First-order calculations for the organic carbon input (5–22 mg C m⁻² d⁻¹) required to sustain observed epi-plume zooplankton anomalies at Endeavour are comparable both to measured total POC flux to epi-plume depths (2–5 mg C m⁻² d⁻¹: combined hydrothermal and surface derived organic carbon) and to estimates of the total potential in situ organic carbon production (2–9 mg C m⁻² d⁻¹) from microbial oxidation of hydrothermal plume H₂, CH₄ and NH₄⁺.     

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.     

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.     

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.     

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.     

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.