Viral abundance and distribution in mesopelagic and bathypelagic waters of the Mediterranean Sea

Despite the fact that marine viruses have been increasingly investigated in the last decade, knowledge on virus abundance, biomass and distribution in mesopelagic and bathypelagic waters is limited. We report here the results of a large-spatial-scale study (covering more than 3000 km) on the virioplankton distribution in epi-, meso- and bathypelagic waters in 19 areas of the Mediterranean Sea, from the Alboran Sea and Western Mediterranean, to the Tyrrhenian Sea, Sicily Channel and Ionian Sea. Integrated viral abundance in epipelagic waters was significantly higher than in deep-sea waters (on average, 2.4 vs. 0.5×10¹² viruses m⁻³). However, abundance of viruses in the deep-Mediterranean waters was the highest reported so far for deep seas worldwide (7.0 and 3.1×10¹¹ viruses m⁻³ in mesopelagic and bathypelagic waters, respectively) and their biomass accounted for 13–18% of total prokaryotic C biomass. The significant relationship between viral abundance and prokaryotic abundance and production in deep waters suggests that also deep-sea viruses are closely dependent on the abundance and metabolism of their hosts. Moreover, virus to prokaryote (and nucleoid-containing cell (NuCC)) abundance ratio increased with increasing depths suggesting that deep waters may represent optimal environments for viral survival or proliferation. Overall, our results indicate that deep waters may represent a significant reservoir of viruses and open new perspectives for future investigations of viral impact on the functioning of meso-bathypelagic ecosystems.     

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

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

Vertical changes in abundance,biomass and community structure of copepods down to 3000 m in the southern Bering Sea

Vertical changes in abundance, biomass and community structure of copepods down to 3000 m depth were studied at a single station of the Aleutian Basin of the Bering Sea (53°28′N, 177°00′W, depth 3779 m) on the 14th June 2006. Both abundance and biomass of copepods were greatest near the surface layer and decreased with increase in depth. Abundance and biomass of copepods integrated over 0–3000 m were 1,390,000 inds. m^?2 and 5056 mg C m^?2, respectively. Copepod carcasses occurred throughout the layer, and the carcass:living specimen ratio was the greatest in the oxygen minimum layer (750–100 m, the ratio was 2.3). A total of 72 calanoid copepod species belonging to 34 genera and 15 families occurred in the 0–3000 m water column (Cyclopoida, Harpacticoida and Poecilostomatoida were not identified to species level). Cluster analysis separated calanoid copepod communities into 5 groups (A–E). Each group was separated by depth, and the depth range of each group was at 0–75 m (A), 75–500 m (B), 500–750 m (C), 750–1500 m (D) and 1500–3000 m (E). Copepods were divided into four types based on the feeding pattern: suspension feeders, suspension feeders in diapause, detritivores and carnivores. In terms of abundance the most dominant group was suspension feeders (mainly Cyclopoida) in the epipelagic zone, and detritivores (mainly Poecilostomatoida) were dominant in the meso- and bathypelagic zones. In terms of biomass, suspension feeders in diapause (calanoid copepods Neocalanus spp. and Eucalanus bungii) were the major component (ca. 10–45%), especially in the 250–3000 m depth. These results are compared with the previous studies in the same region and that down to greater depths in the worldwide oceans.     

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

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

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.     

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).     

Full-depth profiles of prokaryotes,heterotrophic nanoflagellates,and ciliates along a transect from the equatorial to the subarctic central Pacific Ocean

Studies in epipelagic waters report higher heterotrophic microbial biomass in the productive high latitudes than in the oligotrophic low latitudes; however, biogeographical data are scarce in the deep ocean. To examine the hypothesis that the observed latitudinal differences in heterotrophic microbial biomass in the epipelagic zone also occur at depth, abundance and biomass of heterotrophic prokaryotes, nanoflagellates (HNF), and ciliates were determined at depths of 5–5000 m in the central Pacific between August and September of 2005. Heterotrophic microbial biomass increased from the tropical to the subarctic region over the full water column, with latitudinal differences in prokaryotic biomass increasing from 2.3-fold in the epipelagic zone to 4.4-fold in the bathypelagic zone. However, the latitudinal difference in HNF and ciliate biomass decreased with depth. In the mesopelagic zone, the vertical attenuation rate of prokaryotic abundance, which was calculated as the linear regression slope of log-log plot of abundance versus depth, ranged from –0.55 to –1.26 and was more pronounced (steeper slope) in the lower latitudes. In contrast, the vertical attenuation rate of HNF in the mesopelagic zone (–1.06 to –1.27) did not differ with latitude. In the subarctic, the attenuation rate of HNF was 1.7 times steeper than for prokaryotes. These results suggest the accumulation of prokaryotes in the deep subarctic Pacific, possibly due to low grazing pressure. Although the vertical attenuation rate of ciliates was steepest in the bathypelagic zone, HNF abundance did not further decrease at depths below 1000 m, except for at 2000 m where HNF was lowest across the study area. Ciliate abundance ranged 0.3–0.8 cells l^–1 at 4000 m, and were below the detection limit (<0.1 cells l ^–1) at 5000 m. To our knowledge, this study presents the first data for ciliates below 2000 m.     

Spatial and temporal variability of planktonic archaeal abundance in the Humboldt Current System off Chile

The latest advances in the field of microbial ecology have shown that planktonic Archaea are one of the most abundant unicellular microorganisms of the oceans. However, no information is available on the contribution this group makes to the prokaryote assemblages that inhabit the eastern South Pacific Ocean. Here, we describe the relative abundance and vertical distribution of planktonic Archaea off northern and central-southern Chile. Data come from several cruises and a 45-month time series at a station located on the shelf off central-southern Chile. Both the taxonomic composition of the prokaryote community and its relative abundance were determined using quantitative dot blot 16S-rRNA hybridizations. Total Archaea in central-southern Chile made up 6–87% of the prokaryote rRNA in the water column and did not present evidence of any seasonal pattern. Crenarchaea were the most abundant archaeal group at this site and were significantly associated with the ammonium concentration (r²=0.16, p=0.0003, n=80). Archaeal abundance in the time series was usually greater in the deeper layer (>50 m), with contributions reaching up to ∼90% of the prokaryote rRNA on certain occasions, and decreasing towards the surface. Important increments in the relative abundance of total Archaea were observed on given dates at the surface of the time-series station off central-southern Chile. Off northern Chile, total Archaea normally contributed from ∼10% to 50% of the prokaryote rRNA found between 10 and 1000 m, and were generally important in the mesopelagic realm. Our results indicate that Archaea constitute an important fraction of the prokaryote assemblage in the water column of the Humboldt Current System, especially in the oxygen minimum zone.     

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.     

Characterization of the solar light field within the ocean mesopelagic zone based on radiative transfer simulations

The solar light field within the ocean from the sea surface to the bottom of the mesopelagic zone was simulated with a radiative transfer model that accounts for the presence of inelastic radiative processes associated with Raman scattering by water molecules, fluorescence of colored dissolved organic matter (CDOM), and fluorescence of chlorophyll-a contained in phytoplankton. The simulation results provide a comprehensive characterization of the ambient light field and apparent optical properties (AOPs) across the entire visible spectral range within the depth range 200–1000 m of the entire mesopelagic zone for varying chlorophyll-a concentration and seawater optical properties in the mixed surface layer of the ocean. With increasing depth in the mesopelagic zone, the solar irradiance is reduced by ~9–10 orders of magnitude and exhibits a major spectral maximum in the blue, typically centered around a light wavelength of 475 nm. In the green and red spectral regions, the light levels are significantly lower but still important owing to local generation of photons via inelastic processes, mostly Raman scattering and to a lesser extent CDOM fluorescence. The Raman scattering produces a distinct secondary maximum in irradiance spectra centered around 565 nm. Comparisons of our results with light produced by the radioactive decay of the unstable potassium isotope contained in sea salt (⁴⁰K) indicates that the solar irradiance dominates over the ⁴⁰K-produced irradiance within the majority of the mesopelagic zone for most scenarios considered in our simulations. The angular distribution of radiance indicates the dominance of downward propagation of light in the blue and approach to uniform distribution in the red throughout the mesopelagic zone. Below the approximate depth range 400–500 m, the shape of the angular distribution is nearly invariant with increasing depth in the green and red and varies weakly in the blue. The AOPs at any light wavelength also assume nearly constant values within the deeper portion of the mesopelagic zone. These results indicate that the mesopelagic light field reaches a nearly-asymptotic regime at depths exceeding ~400–500 m.     

DOC dynamics in the meso and bathypelagic layers of the Mediterranean Sea

Seven years (2001–2008) of dissolved organic carbon (DOC) vertical profiles were examined in order to assess the main processes determining DOC concentration and distribution in the meso- and bathypelagic layers of the Mediterranean Sea. As expected, DOC showed high and highly variable concentrations in the surface layer of 57–68 μM (average values between 0 and 100 m), with a decrease to 44–53 μM between 200 and 500 m. Deep DOC distribution was strongly affected by deep-water formation, with a significant increase to values of 76 μM in recently ventilated deep waters, and low concentrations, comparable to those observed in the open oceanic waters (34–45 μM), where the oldest, deep waters occurred. In winter 2004/2005 a deep-water formation event was observed and the consequent DOC export at depth was estimated to range between 0.76–3.02 Tg C month^–1. In the intermediate layer, the main path of the Levantine Intermediate Water (LIW) was followed in order to estimate the DOC consumption rate in its core. Multiple regression between DOC, apparent oxygen utilization (AOU), and salinity indicated that 38% of the oxygen consumption was related to DOC mineralization when the effect of mixing was removed. In deep waters of the southern Adriatic Sea a DOC decrease of 6 μM, together with an AOU increase of 9 μM, was observed between the end of January 2008 and the end of June 2008 (5 months). These data indicate a rate of microbial utilization of DOC of about 1.2 μM C month⁻¹, with 92% of the oxygen consumption due to DOC mineralization. These values are surprisingly high for the deep sea and represent a peculiarity of the Mediterranean Sea.     

Cross-slope zonation of erosion and deposition in the Faeroe-Shetland Channel,North Atlantic Ocean

Boundary currents and internal waves determine cross-slope zonation of erosion and deposition in the Faeroe-Shetland Channel. Currents were measured at 8 and 34–50 m above the bottom at three mooring sites (502, 595 and 708 m depth) for 14 days. The structure of the water column was evaluated from CTD sections, and included nepheloid layers and particulate matter concentrations. Indicators for recent deposition in the sediment (organic carbon, phytopigments, ²¹⁰Pb) were measured at eight stations across the slope. Strong near-bottom currents at the upper slope sustain down-slope particle transport in a benthic nepheloid layer, which is eroded under the influence of critically reflecting M₂ internal tidal waves at 350–550 m, where the major pycnocline meets the sloping bottom. Beam attenuation profiles confirmed the presence of intermediate nepheloid layers intruding into the Channel along the major pycnocline, and elevated concentrations of particulate matter and chlorophyll-a were measured at this depth. Near-bottom currents decreased with depth, thus allowing particle deposition down the slope. Inventories of excess ²¹⁰Pb activity in the sediment deeper than 600 m were higher than what was expected on the basis of atmospheric input of ²¹⁰Pb and production in the water column, thus indicating additional lateral inputs. Simple calculations showed that off-slope input of particles from areas shallower than 600 m may be responsible for the enhanced deposition at greater depths.     

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.     

Bioluminescence in the deep sea: Free-fall lander observations in the Atlantic Ocean off Cape Verde

A novel autonomous free-fall lander vehicle, with a capability down to 6000 m, was deployed off Cape Verde for studies on bioluminescence in the deep sea. The system was equipped with a high-sensitivity Intensified Silicon Intensified Target (ISIT) video camera, a programmable control-recording unit and an acoustic current meter with depth and temperature sensors. The ISIT lander was used in three modes: (1) free falling at 34 m min⁻¹, with the camera looking downwards at a mesh screen, recording impacts of luminescent organisms to obtain a vertical profile down to the abyssal sea floor, sampling at >100 l s⁻¹; (2) rotating, with the lander on the sea floor and the camera orienting to the bottom current using a servo-controlled turntable, impacts of luminescent organisms carried by the bottom current onto a mesh screen mounted 0.5 m in front of the camera were recorded to estimate abundance in the benthic boundary layer; (3) baited, with the camera focused on a bait placed on the sea floor.Profiles recorded abundance of luminescent organisms as 26.7 m⁻³ at 500–999 m depth, decreasing to 1.6 m⁻³ at 2000–2499 m and 0.5 m⁻³ between 2500 m and the sea floor at 4046 m, with no further detectable significant change with depth. Rotator measurements at a 0.5 m height above the sea floor gave a mean abundance of 0.47 m⁻³ in the benthic boundary layer at 4046 m and of 2.04 m⁻³ at 3200 m. Thirty five minutes after the bait was placed on the sea floor at 3200 m, bioluminescent fauna apparently arrived at the bait and produced luminescent displays at a rate of 2 min⁻¹. Moving, flashing light sources were observed and luminescent material was released into the bottom current.     

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.     

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.     

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.     

Nitrogen sources for new production in the NE Arabian Sea

New productivity measurements using the ¹⁵N tracer technique were conducted in the north-eastern (NE) Arabian Sea during six expeditions from 2003 to 2007, mostly in winter. Our results indicate that the NE Arabian Sea has a potential for higher new productivity during blooms. Nitrate uptake by plankton is the highest during late winter. New productivity and f-ratios in the NE Arabian Sea are mainly controlled by hydrodynamic and meteorological parameters such as wind strength, sea surface temperature (SST), mixed layer depth (MLD) and mixed layer nitrate. Deepening of the mixed layer supplies nitrate from below, which supports the observed nitrogen uptake. Higher f-ratios during blooms indicate the strong coupling between surface layers and sub-surface layers. Deepening of mixed layer below 100 m (from its inter-monsoon value between 30 and 40 m) transferred often more than 100 mmol N–NO₃ m^? 2 into the surface layers from below. The observed winter blooms in the region are supported by such input and are sustained for more than a month. Higher new productivity has been found in late winter, whereas transport of nitrate is maximum in early winter. In general, new production varies progressively during winter. Diurnal cycling of the mixed layer could be the reason for the under utilization of entrained nitrate during early winter. New productivity values and wind strength show significant differences during Feb–Mar 03 and Feb–Mar 04. These differences indicate that the winter cooling and parameters related the biological productivity also vary inter-annually. However, the difference between the new productivity values between Feb–Mar 03 and Feb–Mar 04 is much lower than the difference between Jan 03 and Feb–Mar 03. The results suggest that amplitude of seasonal variation is higher than the inter-annual variation in the region. During spring, Fickian diffusive fluxes of nitrate into the surface layer range from 0.51 to 1.38 mmol N–NO₃ m^? 2 day^? 1, and can account for 67% and 78% of the observed nitrogen uptake in the coastal and open ocean regions, respectively. We document the intra-seasonal and inter-annual variations in new productivity during winter and identify sources of nitrate which support the observed productivity during spring.     

Primary production export flux in Marguerite Bay (Antarctic Peninsula): Linking upper water-column production to sediment trap flux

A study was carried out to assess primary production and associated export flux in the coastal waters of the western Antarctic Peninsula at an oceanographic time-series site. New, i.e., exportable, primary production in the upper water-column was estimated in two ways; by nutrient deficit measurements, and by primary production rate measurements using separate ¹⁴C-labelled radioisotope and ¹⁵N-labelled stable isotope uptake incubations. The resulting average annual exportable primary production estimates at the time-series site from nutrient deficit and primary production rates were 13 and 16 mol C m⁻², respectively. Regenerated primary production was measured using ¹⁵N-labelled ammonium and urea uptake, and was low throughout the sampling period.The exportable primary production measurements were compared with sediment trap flux measurements from 2 locations; the time-series site and at a site 40 km away in deeper water. Results showed ∼1% of the upper mixed layer exportable primary production was exported to traps at 200 m depth at the time-series site (total water column depth 520 m). The maximum particle flux rate to sediment traps at the deeper offshore site (total water column depth 820 m) was lower than the flux at the coastal time-series site. Flux of particulate organic carbon was similar throughout the spring–summer high flux period for both sites. Remineralisation of particulate organic matter predominantly occurred in the upper water-column (<200 m depth), with minimal remineralisation below 200 m, at both sites. This highly productive region on the Western Antarctic Peninsula is therefore best characterised as ‘high recycling, low export’.