Chemical composition and energy content of deep-sea calanoid copepods in the Western North Pacific Ocean

Condition factor index [CFI=1000×DW/(PL)³; DW: dry weight, PL: prosome length], water content, carbon (C), nitrogen (N), ash and energy content were determined on a total of 69 copepod species caught from the mesopelagic (500–1000 m), upper-bathypelagic (1000–2000 m), lower-bathypelagic (2000–3000 m) and abyssopelagic (3000–5000 m) zones of the western subarctic Pacific. Resultant data were grouped into these four sampling zones, four developmental stage/sex categories (C4, C5 and C6 females and males), three feeding types (carnivore, detritivore and suspension feeder), or two reaction speed groups by the presence/absence of myelinated sheath enveloping axons (fast and slow reacting species). Zone-structured data showed the overall ranges were 3.8–4.6 mm for PL, 1.6–2.6 mg for DW, 21.4–25.0 for CFI, 75.0–78.6% of wet weight (WW) for water, 51.3–53.7% of DW for C, 7.7–8.8% of DW for N, 6.2–7.0 (by weight) for C/N, 6.9–9.6% of DW for ash and 25.3–27.4 J mg⁻¹ DW for energy. Among these components, N and ash exhibited significant between-zone differences characterized by gradual decrease downward for the former, and only the upper-bathypelagic zone>abyssopelagic zone for the latter. Stage/sex-structured data showed no significant differences among them, but energy content of C5 was higher than that of C6 females. From the analyses of feeding type-structured data, carnivores were shown to have lower water, N, ash, but higher C, C/N and energy contents than suspension feeders do. Reaction speed-structured data indicated that slow-reacting species have significantly higher water but lower CFI, C, N and energy contents than fast-reacting species. Designating these grouping criteria, PL and DW as independent variables, the attributes of these variables to the CFI, chemical composition or energy contents were evaluated by stepwise-multiple regression analysis, showing the most pronounced effect of suspension-feeder, followed by the presence of myelinated sheath, DW, C6 females and the abyssopelagic zone. Further analysis of zone-structured data, by adding epipelagic copepod data from identical thermal habitats (Arctic/Antarctic waters), revealed a more marked decline in N content from the epipelagic zone to the abyssopelagic zone, accompanied by an increase in C/N ratios downward. The decline in N (=protein or muscle) contents with depth cannot be explained by the “visual interactions” hypotheses being proposed for the metabolism of pelagic visual predators, but is consistent with the “predation-mediated selection” hypothesis for the metabolism of pelagic copepods.     

Patterns of bathymetric zonation of bivalves in the Porcupine Seabight and adjacent Abyssal plain,NE Atlantic

Although the organization patterns of fauna in the deep sea have been broadly documented, most studies have focused on the megafauna. Bivalves represent about 10% of the deep-sea macrobenthic fauna, being the third taxon in abundance after polychaetes and peracarid crustaceans. This study, based on a large data set, examined the bathymetric distribution, patterns of zonation and diversity–depth trends of bivalves from the Porcupine Seabight and adjacent Abyssal Plain (NE Atlantic). A total of 131,334 individuals belonging to 76 species were collected between 500 and 4866 m. Most of the species showed broad depth ranges with some ranges extending over more than 3000 m. Furthermore, many species overlapped in their depth distributions. Patterns of zonation were not very strong and faunal change was gradual. Nevertheless, four bathymetric discontinuities, more or less clearly delimited, occurred at about 750, 1900, 2900 and 4100 m. These boundaries indicated five faunistic zones: (1) a zone above ∼750 m marking the change from shelf species to bathyal species; (2) a zone from ∼750 to 1900 m that corresponds to the upper and mid-bathyal zones taken together; (3) a lower bathyal zone from ∼1900 to 2900 m; (4) a transition zone from ∼2900 to 4100 m where the bathyal fauna meets and overlaps with the abyssal fauna and (5) a truly abyssal zone from approximately 4100–4900 m (the lower depth limit of this study), characterized by the presence of abyssal species with restricted depth ranges and a few specimens of some bathyal species with very broad distributions. The ∼4100 m boundary marked the lower limit of distribution of many bathyal species. There was a pattern of increasing diversity downslope from ∼500 to 1600 m, followed by a decrease to minimum values at about 2700 m. This drop in diversity was followed by an increase up to maximum values at ∼4100 m and then again, a fall to ∼4900 m (the lower depth limit in this study).     

Zooplankton distribution across Fram Strait in autumn: Are small copepods and protozooplankton important?

We investigated zooplankton distribution in September 2006/2007 at eight stations across Fram Strait in contrasting water masses ranging from cold Polar water to warm Atlantic water. Our main objectives were: (1) to describe the plankton community in the upper 200 m during autumn, and (2) to investigate the importance of small-sized copepods and protozooplankton in an arctic ecosystem when the majority of the large Calanus species had entered diapause. We sampled both with a WP-2 net and Go-Flo bottle and show that small copepods <1 mm are significantly undersampled using a WP-2 net with 90 μm mesh.Small copepods and protozooplankton made a significant contribution both in terms of abundance and total zooplankton biomass at all stations in September, when the large calanoid copepods had left the upper 200 m. The dominating group in the upper 60 m at all stations was Oithona spp. nauplii and their daily estimated grazing potential on the <10 μm phytoplankton ranged from 0.1% to 82% of the standing stock. Both Oithona copepodites and nauplii biomass showed a significantly positive relation with temperature, but not with potential food. Heterotrophic protozooplankton, on the other hand, were most likely bottom-up regulated by the availability of phytoplankton <10 μm. We hypothesise that Oithona nauplii and protozooplankton compete for food and conclude that there was a strong link between the zooplankton community and the microbial food web in Fram Strait.     

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 prokaryote production and abundance in the mesopelagic and bathypelagic layers of the Canada Basin,western Arctic: Implications for the mode and extent of organic carbon delivery

The vertical distributions of prokaryote heterotrophic production (³H-leucine incorporation rate) and abundance were investigated in the meso- and bathy-pelagic layers of the Canada Basin, western Arctic Ocean, during September 2009. Prokaryote production and abundance were high in the Pacific-origin water mass located in the upper mesopelagic layer (depth, 100–200 m). Below the halocline layer (depth, 300–3000 m), both the production and abundance decreased with depth, with log–log regression slopes of −1.33 and −0.77, respectively. Depth-integrated production and biomass in the meso- and bathy-pelagic layers was three- to five-fold lower than the corresponding values reported in the subpolar regions, whereas they were close to or lower than the corresponding values in oligotrophic subtropical regions. Prokaryote turnover times were estimated to be 1.1 and 6.1 years for meso- and bathy-pelagic layers, respectively, with the latter being among the longest turnover times reported for oceanic basins. We estimated prokaryote carbon demand in the water column (100–3000 m) to be on the order of 11 mg C m⁻² d⁻¹, which largely exceeds (by 38-fold) the sinking particulate organic carbon flux at depths of 120–200 m reported in the literature. This large carbon imbalance may be partly explained by organic carbon delivery by lateral intrusion of the Pacific-origin water mass into the upper mesopelagic layer.     

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.     

Distribution and relative importance of jellyfish in a region of hydrothermal venting

Net sampling to 3000 m depth at Endeavour Ridge in the northeast Pacific in July 1991–1994 shows that medusae in the immediate vicinity of the hydrothermal vent fields often make up a larger proportion of the total zooplankton abundance and biomass from mesopelagic to bathypelagic depths than in the surrounding waters. This was particularly evident in the dominant Trachymedusae, and least evident in the siphonophores. In addition, the large red Scyphomedusa Stygiomedusa gigantea was a major biomass component in the region of the deep (1000–1800 m depth) migrating scattering layers at the vent field, but was not found in any net tows greater than 10 km away from vents. There is no concurrent increase in relative or percent biomass of fish or chaetognaths, which are the other major predators in the community. We hypothesize that predaceous medusae respond opportunistically to the enhanced zooplankton biomass throughout the water column around vents in spring to early summer, in a way that other predators do not.     

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.     

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.     

Mesozooplankton in the Arctic Ocean in summer

The biomass, species and chemical composition of the mesozooplankton and their impact on lower food levels were estimated along a transect across the Arctic Ocean. Mesozooplankton biomass in the upper 200 m of the water column was significantly higher (19–42 mg DW m^-3) than has previously been reported for the Arctic Ocean, and it reached a maximum at ca. 87°N in the Amundsen Basin. The lowest values were recorded in the Chukchi Sea and Nansen Basin, where ice cover was lower (50–80%) than in the central Arctic Ocean. In the deeper strata (200–500 m) of the Canadian and Eurasian Basins, the biomass was always much lower (4.35–16.44 mg DW m^-3). The C/N (g/g) ratio for the mesozooplankton population was high (6.5–8.5) but within the documented range. These high values (when compared to 4.5 at lower latitudes) may be explained by the high lipid content. Mesozooplankton accounted for approximately 40% of the total particulate organic carbon in the upper 100 m of the water column. Mesozooplankton species composition was homogeneous along the transect, consisting mainly of copepods (70–90% of the total number). It was dominated by four large copepod species (Calanus hyperboreus, C. glacialis, C. finmarchicus and Metridia longa), which together accounted for more than 80% of the total biomass. According to measurements of gut pigment and gut turnover rates, the mesozooplankton on average ingested between 6 and 30% of their body carbon per day as phytoplankton. Microzooplankton may have provided an additional source of energy for the mesozooplankton community. These data emphasize the importance of mesozooplankton in the arctic food web and reinforce the idea that the Arctic Ocean should no longer be considered to be a “biological desert”.     

Seasonal changes in zooplankton abundance,biomass, size structure and dominant copepods in the Oyashio region analysed by an optical plankton counter

To identify seasonal patterns of change in zooplankton communities, an optical plankton counter (OPC) and microscopic analysis were utilised to characterise zooplankton samples collected from 0 to 150 m and 0 to 500 m in the Oyashio region every one to three months from 2002 to 2007. Based on the OPC measurements, the abundance and biomass of zooplankton peaked in June (0–150 m) or August (150–500 m), depending on the depth stratum. The peak periods of the copepod species that were dominant in terms of abundance and biomass indicated species-specific patterns. Three Neocalanus species (Neocalanus cristatus, Neocalanus flemingeri and Neocalanus plumchrus) exhibited abundance peaks that occurred before their biomass peaks, whereas Eucalanus bungii and Metridia pacifica experienced biomass peaks before their abundance peaks. The abundance peaks corresponded to the recruitment periods of early copepodid stages, whereas the biomass peaks corresponded to the periods when the dominant populations reached the late copepodid stages (C5 or C6). Because the reproduction of Neocalanus spp. occurred in the deep layer (>500 m), their biomass peaks were observed when the major populations reached stage C5 after the abundance peaks of the early copepodid stages. The reproduction of E. bungii and M. pacifica occurred near the surface layer. These species first formed biomass peaks of C6 and later developed abundance peaks of newly recruited early copepodid stages. From the comparison between OPC measurements and microscopic analyses, seasonal changes in zooplankton biomass at depths of 0–150 m were governed primarily by E. bungii and M. pacifica, whereas those at depths of 150–500 m were primarily caused by the three Neocalanus species.     

The natural diet of a hexactinellid sponge: Benthic–pelagic coupling in a deep-sea microbial food web

Dense communities of shallow-water suspension feeders are known to sidestep the microbial loop by grazing on ultraplankton at its base. We quantified the diet, rates of water processing, and abundance of the deep-sea hexactinellid sponge Sericolophus hawaiicus, and found that, like their demosponge relatives in shallow water, hexactinellids are a significant sink for ultraplankton. S. hawaiicus forms a dense bed of sponges on the Big Island of Hawaii between 360 and 460 m depth, with a mean density of 4.7 sponges m⁻². Grazing of S. hawaiicus on ultraplankton was quantified from in situ samples using flow cytometry, and was found to be unselective. Rates of water processing were determined with dye visualization and ranged from 1.62 to 3.57 cm s⁻¹, resulting in a processing rate of 7.9±2.4 ml sponge⁻¹ s⁻¹. The large amount of water processed by these benthic suspension feeders results in the transfer of approximately 55 mg carbon and 7.3 mg N d⁻¹ m⁻² from the water column to the benthos. The magnitude of this flux places S. hawaiicus squarely within the functional group of organisms that link the pelagic microbial food web to the benthos.     

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.     

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.     

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.     

Commercially-cultured oysters (Crassostrea gigas) exert top-down control on intertidal pelagic resources in Willapa Bay,Washington, USA

The capacity of filter feeders to reduce seston and phytoplankton concentrations in the water column has important implications for restoration and management of coastal ecosystems. We directly measured changes in chlorophyll a concentration on commercially stocked intertidal oyster beds (Crassostrea gigas) in Willapa Bay, Washington, USA by recording water properties near small drifters as they tracked parcels of water across tide flats. Chlorophyll declined 9.6% per half hour in water passing on-bottom adult oysters and 41% for longline adult oysters, whereas chlorophyll concentrations increased as water flowed across tide flats without adult oysters. Field filtration rates, which were fit to exponential declines in chlorophyll and accounted for oyster density and water depth, averaged 0.35 L g^− 1 h^− 1 (shucked dry weight) for on-bottom aquaculture and 0.73 L g^− 1 h^− 1 for longline culture, compared to values of 2.5–12 L g^− 1 h^− 1 reported from laboratory studies of C. gigas. Field filtration rates may be lower than laboratory rates due to unfavorable field conditions (e.g., low initial chlorophyll concentrations) or masked by resuspension of benthic microalgae. In addition to distinctions among on-bottom, longline, and no-oyster habitats, Akaike's Information Criterion analysis showed temperature, initial chlorophyll concentration, and depth related to chlorophyll decline. This research corroborates mathematical models suggesting that benthic suspension feeders are exerting top-down control of pelagic production in this estuary, with strong patterns in chlorophyll emerging across extensive tideflats populated by C. gigas despite low field filtration rates.     

Biogeochemical cycling of N and Si during the mesoscale iron-enrichment experiment in the western subarctic Pacific (SEEDS-II)

Biogeochemical cycles of N and Si were examined in the surface mixed layer during the mesoscale iron-enrichment (IE) experiment in the high-nutrient low-chlorophyll (HNLC) western subarctic Pacific (SEEDS-II). Although the IEs increased nitrate uptake, silicic acid utilization was not stimulated. The nitrate drawdown in the iron-patch (IN-patch, 140.3 mmol m⁻² in the surface mixed layer, 0–30 m) was only 25% of the initial inventory, which was 1/3–2/5 of the previous IE experiments in the subarctic Pacific. This relatively weak response of nutrient drawdown to IEs was due to the high biomass of mesozooplankton (MZ) dominated by copepod Neocalanus plumchrus. Feeding of MZ (247.2 mmol m⁻² during Day 0–21 from the first IE) in the IN-patch was higher than the nitrate drawdown and prevented further development of the phytoplankton bloom. In the later period of the experiment (Day 14–21), the increase in the feeding activity and resultant decrease in phytoplankton biomass induced the accumulation of dissolved organic nitrogen (DON) and ammonium. Among total growth of MZ (81.6 mmol N m⁻²), 89% (72.8 mmol N m⁻²) was transported to the depth by the ontogenetic downward migration of N. plumchrus. Although silicic acid drawdown was not increased by the IEs, Si export flux increased by 2.7 times. The increase in Si export was also due to the increase in MZ, which egested faecal pellets with higher Si:N ratio and faster sinking speed than diatoms. The export efficiency (78% of new production) and total amount of export flux (143.8 mmol N m⁻², 1392 mmol C m⁻²) were highest records within the IE experiments despite weak responses of nutrient drawdown to the IE. During SEEDS-II, the high biomass of MZ reduced the phytoplankton response and nutrient drawdown to the IEs but via grazing and ontogenetic vertical migration accelerated the export flux as well as accumulations of dissolved forms of N. Results of the present and previous IE experiments indicate that the ecosystem and biogeochemical responses to IEs in the HNLC region are quite sensitive to the ecosystem components, especially for grazers of diatoms such as copepods and heterotrophic dinoflagellates. More attention needs to be paid to the ecosystem components and their biogeochemical functions as well as physical and chemical properties of the ecosystems in order to hindcast or forecast the impacts of changes in atmospheric iron deposition.     

Spring distribution of larger (>64 μm) protozoans in the Atlantic sector of the Southern Ocean

Distribution of larger protozoans (armoured dinoflagellates, tintinnids, heliozoans, radiolarians and foraminiferans >64 μm) is presented for three major water masses of the Southern Ocean: the Polar Front region (PFr), the southern Antarctic Circumpolar Current (southern ACC) and the northern Weddell Gyre. Sampling took place during the SO-JGOFS cruise ANT X/6 of R/V Polarstern (October–November 1992) along a meridional transect at 6°W between 48°00′S and 59°30′S. Multinet samples (64 μm mesh size) were taken at six stations from the surface down to 500 m depth at five different depth intervals. In the upper 100 m of the water column abundances of larger protozoans varied between 94 and 10,930 ind. m^–3, with highest abundances in the PFr, where phytoplankton blooms occurred, and lowest values in the Antarctic Circumpolar Current–Weddell Gyre Boundary (AWB). Foraminiferans and polycystine and smaller (<300 μm) phaeodarian radiolarians dominated larger protozoan assemblages in the PFr. In open water of the southern ACC, tintinnids, armoured dinoflagellates, foraminiferans and smaller (<300 μm) phaeodarian radiolarians were equally important. The heliozoans Sticholonche spp. and nassellarian radiolarians dominated assemblages in the Weddell Gyre and AWB. Larger protozoan biomasses ranged between 2 and 674 μg C m⁻³ and were always dominated by larger (>300 μm) phaeodarians. Highest biomasses were found in the AWB between 200 and 500 m depth. Standing stocks of larger protozoans constituted a negligible fraction of zooplankton biomass in the upper 200 m of the water column. In deeper layers of the ice-covered Weddell Gyre and AWB their biomasses, dominated by larger (>300 μm) phaeodarians, was significant contributing up to 45% to total larger protozoan and metazoan biomass. Analysis of correlation between distribution patterns and environmental conditions at the stations sampled indicate that spring distribution patterns of heterotrophic armoured dinoflagellates, polycystine radiolarians and foraminiferans follow productivity in the water column. Of the protozoan groups studied the smaller (<300 μm) phaeodarian radiolarians also showed a significant correlation with productivity during spring, however, results from previous studies do not suggest a consistent pattern. Spring distribution patterns of other larger protozoans were not related to differences in productivity in the water column, and effects such as ice-cover, grazing or silica limitation might be determining. Dead radiolarian skeletons constituted on average 27, 8 and 11% of the population of nassellarians, spumellarians and smaller (<300 μm) phaeodarians, respectively. The contribution of dead radiolarian skeletons to total radiolarian stocks varied with depth and water mass. Differences between live and skeleton assemblages composition were observed. These differences should be taken into consideration when interpreting the geological record.     

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.