Physical-biological coupling in the Algerian Basin (SW Mediterranean): Influence of mesoscale instabilities on the biomass and production of phytoplankton and bacterioplankton

The biomass and production of phytoplankton and bacterioplankton was investigated in relation to the mesoscale structures found in the Algerian Current during the ALGERS'96 cruise (October 1996). Biological determinations were carried out in three transects between 0° and 2°E aimed at crossing a so-called event, formed by a coastal anticyclonic eddy associated with an offshore cyclonic eddy to the west. The concentration of chlorophyll a (Chl) was maximum (>1.2 mg m⁻³) within the cyclonic eddy and at the frontal zones between the Modified Atlantic Water (MAW) of the Algerian Current and the Mediterranean waters further north. Chl (total and >2 μm) was significantly correlated with proxies of nutrient flux into the upper layers. Autotrophic picoplankton and heterotrophic bacterial abundance and production presented clear differences between MAW and Mediterranean water, with higher values at those stations under the influence of the Algerian Current. In general, greater differences were observed in production than in biomass variables. The photosynthetic parameters (derived from P–E relationships) and integrated primary production (range 189–645 mg m⁻² d⁻¹) responded greatly to the different hydrological conditions. The mesoscale phenomena inducing fertilization caused a 2 to 3-fold increase in primary production rates. The relatively high values found within the cyclonic eddy suggest that, although short-lived in comparison with anticyclonic eddies, these eddies may produce episodic increases of biological production not accounted for in previous surveys in the region.     

Impacts of the wintertime mesozooplankton community to downward carbon flux in the subarctic and subtropical Pacific Oceans

We compared wintertime depth distributions of the mesozooplankton community and dominant copepods between the subtropical (S1) and subarctic (K2) Pacific Oceans to evaluate the relative importance of actively transported carbon by vertical migrants to sinking particulate organic carbon flux. Primary production was higher and the ratio of sinking particulate organic carbon flux to primary production was lower at S1 compared with those at K2. The mesozooplankton community was lower in abundance and biomass at S1 compared to K2. Copepods were the dominant group among both mesozooplankton abundance and biomass throughout the water column down to 1000 m at both sites. The depth distribution showed that diel vertical migration was obvious for the mesozooplankton abundance and biomass at S1 but was not apparent for the abundance at K2, because the dominant component was diurnally migrating species at S1 and overwintering species residing at mesopelagic depths at K2. The major components of diel migrants were copepods and euphausiids at S1 and only euphausiids at K2. Respiratory flux by the diurnally migrating mesozooplankton was estimated to be 2 mgC m⁻² day⁻¹ at S1 and 7 mgC m⁻² day⁻¹ at K2. The respiratory flux was equivalent to 131% of sedimentary fecal pellet flux at S1 and 136% of that at K2. Because pathways of downward carbon flux are facilitated by the mesozooplankton community, the actively transported carbon (respiration of dissolved inorganic carbon, excretion of dissolved organic carbon and egestion of fecal pellets at depth) might be larger during winter than the flux of sinking fecal pellets.     

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.     

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

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

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.     

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

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

Seasonal dynamics of primary and new production in the northern South China Sea: The significance of river discharge and nutrient advection

Biochemical and productivity measurements and nutrient enrichment experiments were conducted on three cruises in summer and two cruises in winter on the shelf and the basin of the northern South China Sea (SCS) between 2001 and 2004. Phytoplankton production, in terms of depth-integrated new production (INP) or depth-integrated primary production (IPP), was higher in winter than in summer and on the shelf than in the basin. In winter, with deepening of the mixed layer, nitrate from the shallow nitracline that characterized the SCS waters was made available in the surface and supported the highest production of the year. Averaged INP measured in winter (0.25 g C m⁻² d⁻¹) was about twice the summer average (0.12 g C m⁻² d⁻¹) and was 0.19 g C m⁻² d⁻¹ on the shelf compared with 0.15 g C m⁻² d⁻¹ in the basin. In winter, average INP on the shelf was higher than the basin (0.34 versus 0.21 g C m⁻² d⁻¹); whereas in summer, averaged INP on the shelf (0.13 g C m⁻² d⁻¹) and the basin (0.11 g C m⁻² d⁻¹) were similar. While averaged IPP measured in the basin was higher in winter than in summer (0.53 versus 0.35 g C m⁻² d⁻¹), IPP on the shelf showed little temporal variation (0.82 in winter versus 0.84 g C m⁻² d⁻¹ in summer). Considerable spatial and inter-annual variation in production was measured in the shelf waters during summer, which could be linked to discharge volume and plume flow direction of the Zhujiang River. While the shelf waters in summer were mostly nitrogen starved or nitrogen and phosphorus co-limited, excessive river runoff may cause the nutritive state to shift to phosphorus deficiency. Waters with low surface salinities and high fluorescence from riverine mixing could be found extending from the Zhujiang mouth to as far as offshore southern Taiwan after a typhoon passed the northern SCS and brought heavy rainfall. Overall, both nutrient advection in winter and river discharge from the China coast in summer made new nitrogen available and shaped the dynamics of phytoplankton production in these oligotrophic waters.     

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%).     

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

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

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.     

Vertical flux of particulate Al,Fe, Pb,and Ba from the upper ocean estimated from 234Th/238U disequilibria

The vertical sinking flux of particulate Al, Fe, Pb, and Ba from the upper 250 m of the Labrador Sea has been estimated from measurements of ²³⁴Th/²³⁸U disequilibrium and the respective metal/²³⁴Th ratios in >53 μm size particles. ²³⁴Th-derived particulate metal fluxes include in situ scavenged metals, labile lithogenic metals, and metals derived from external input (e.g., atmospheric supply). In contrast to the POC/²³⁴Th ratio, particle size-fractionated (0.4–10 μm, 10–53 μm, and >53 μm) Al/²³⁴Th, Fe/²³⁴Th and Pb/²³⁴Th, and Ba/²³⁴Th ratios generally increase with depth and exhibit no systematic change with particle diameter. Sinking fluxes of particulate Al (2.47–22.3 μmol m⁻² d⁻¹), Fe (2.69–16.3 μmol m⁻² d⁻¹), Pb (2.85–70 nmol m⁻² d⁻¹), and Ba (0.13–2.1 μmol m⁻² d⁻¹) at 50 m (base of the euphotic zone) and 100 m (base of the mixed layer) are largely within the range of previous sediment trap results from other ocean basins. Estimates of the upper ocean residence time of Al (0.07–0.28 yr) and Pb (0.8–2.9 yr) are short compared to previously reported values. The settling rate of >53 μm particles calculated from the ²³⁴Th data ranges from 14 to 38 m d⁻¹.     

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.     

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

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

High-frequency fluxes of labile compounds in the central Ligurian Sea,northwestern Mediterranean

Sinking particles were collected every 4 h with drifting sediment traps deployed at 200 m depth in May 1995 in a 1-D vertical system during the DYNAPROC observations in the northwestern Mediterranean sea. POC, proteins, glucosamine and lipid classes were used as indicators of the intensity and quality of the particle flux. The roles of day/night cycle and wind on the particle flux were examined. The transient regime of production from late spring bloom to pre-oligotrophy determined the flux intensity and quality. POC fluxes decreased from, on average, 34 to 11 mg m⁻² d⁻¹, representing 6–14% of the primary production under late spring bloom conditions to 1–2% under pre-oligotrophic conditions. Total protein and chloroplast lipid fluxes correlated with POC and reflected the input of algal biomass into the traps. As the season proceeded, changes in the biochemical composition of the exported material were observed. The C/N ratio rose from 7.8 to 12. Increases of serine (10–28% of total proteins), total lipids (7–9 to 14–28% of POC) and reserve lipids (1–5 to 5–22% of total lipids) were noticeable, whereas total protein content in POC decreased (20–27 to 18–7%). N-acetyl glucosamine, a tracer of fecal pellet flux, showed that zooplankton grazing was a major vector of downward export during the decaying bloom. Against this background pattern, episodic events specifically increased the flux, modifying the quality and the settling velocity of particles. Day/night signals in biotracers (POC, N-acetyl glucosamine, protein and chloroplast lipids) showed that zooplankton migrations were responsible for sedimentation of fresh material through fast sinking particles (V=170–180 m d⁻¹) at night. Periodic signatures of re-processed material (high lipolysis and bacterial biomass indices) suggested that other zooplankton fecal pellets or small aggregates, probably of lower settling velocities (V<170 m d⁻¹), contributed to the flux during calm periods. At the beginning of the experiment, during the development of a prymnesiophyte bloom in the upper layers, the sterol signal with no periodicity enabled us to estimate high particle settling velocities (⩾600 m d⁻¹) likely related to large aggregate formation. A wind event increased biotracer fluxes (POC, protein, chloroplast lipids). The rapid transmission of surface signals through extremely fast sinking particles could be a general feature of particle fluxes in marine areas unaffected by horizontal advection.     

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.     

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.     

Zooplankton associated with the oxygen minimum zone system in the northern upwelling region of Chile during March 2000

Zooplankton in the coastal upwelling region off northern Chile may play a significant biogeochemical role by promoting carbon flux into the subsurface OMZ (oxygen minimum zone). This work identifies the dominant zooplankton species inhabiting the area influenced by the OMZ in March 2000 off Iquique (20°S, northern Chile). Abundance and vertical distribution studies revealed 17 copepod and 9 euphausiid species distributed between the surface and 600 m at four stations sampled both by day and by night. Some abundant species remained in the well-oxygenated upper layer (30 m), with no evidence of diel vertical migration, apparently restricted by a shallow (40–60 m) oxycline. Other species, however, were found closely associated with the OMZ. The large-sized copepod Eucalanus inermis was found below the oxycline and performed diel vertical migrations into the OMZ, whereas the very abundant Euphausia mucronata performed extensive diel vertical migrations between the surface waters and the core of the OMZ (200 m), even crossing it. A complete assessment of copepods and euphausiids revealed that the whole sampled water column (0–600 m) is occupied by distinct species having well-defined habitats, some of them within the OMZ. Ontogenetic migrations were evident in Eucalanidae and E. mucronata. Estimates of species biomass showed a substantial (>75% of total zooplankton biomass) daily exchange of C between the photic layer and the OMZ. Both E. inermis and E. mucronata can actively exchange about 37.8 g C m⁻² d⁻¹ between the upper well-oxygenated (0–60 m) layer and the deeper (60–600 m) OMZ layer. This migrant biomass may contribute about 7.2 g C m⁻² d⁻¹ to the OMZ system through respiration, mortality, and production of fecal pellets within the OMZ. This movement of zooplankton in and out of the OMZ, mainly as a result of the migratory behavior of E. mucronata, suggests a very efficient mechanism for introducing large amounts of freshly produced carbon into the OMZ system and should, therefore, be considered when establishing C budgets for coastal upwelling systems.     

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⁺.     

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.     

Lithogenic and biogenic particle fluxes on the Lomonosov Ridge (central Arctic Ocean) and their relevance for sediment accumulation: Vertical vs. lateral transport

Investigations of lithogenic and biogenic particle fluxes using long-term sediment traps are still very rare in the northern high latitudes and are restricted to the arctic marginal seas and sub-arctic regions. Here data on the variability of fluxes of lithogenic matter, CaCO₃, opal, and organic carbon and biomarker composition from the central Arctic Ocean are presented for a 1-year period. The study was carried out on material obtained from a long-term mooring system equipped with two multi-sampling traps, at 150 and 1550 m depth, and deployed on the southern Lomonosov Ridge close to the Laptev Sea continental margin from September 1995 to August 1996. In addition, data from surface sediments were included in the study. Annual fluxes of lithogenic matter, CaCO₃, opal, and particulate organic carbon were 3.9, 0.8, 2.6, and 1.5 g m⁻² y⁻¹, respectively, in the shallow trap and 11.3, 0.5, 2.9, and 1.05 g m⁻² y⁻¹, respectively, in the deep trap.Both the shallow and the deep trap showed significant variations in vertical flux over the year. Higher values were found from mid-July to the end of October (total mass flux of 75–130 mg m⁻² d⁻¹ in the shallow trap and 40–225 mg m⁻² d⁻¹ in the deep trap). During all other months, fluxes were fairly low in both traps (most total mass flux values <10 mg m⁻² d⁻¹). The interval of increased fluxes can be separated into (1) a mid-July/August maximum caused by increased primary production as documented in high abundances of marine biomarkers and diatoms and (2) a September/October maximum caused by increased influence of Lena River discharge indicated by maximum lithogenic flux and large amounts of terrigenous/fluvial biomarkers in both traps. During September/October, total mass fluxes in the deep trap were significantly higher than in the shallow trap, suggesting a lateral sediment flux at greater depth. The lithogenic flux data also support the importance of sediment input from the Laptev Sea for the sediment accumulation on the Lomonosov Ridge on geological time scales, as indicated in sedimentary records from this region.