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.     

Biomass size-spectra of macrobenthic communities in the oxygen minimum zone off Chile

Estimates of macrofaunal secondary production and normalized biomass size-spectra (NBSS) were constructed for macrobenthic communities associated with the oxygen minimum zone (OMZ) in four areas of the continental margin off Chile. The presence of low oxygen conditions in the Humboldt Current System (HCS) off Chile was shown to have important effects on the size structure and secondary production of the benthic communities living in this ecosystem. The distribution of normalized biomass by size was linear (log₂–log₂ scale) at all stations. The slope of the NBSS ranged from −0.481 to −0.908. There were significant differences between the slopes of the NBS-spectra from the stations located in the OMZ (slope = −0.837) and those located outside the OMZ (slope = −0.463) (p < 0.05). The results of this study suggest that low oxygen conditions (<0.5 ml L⁻¹) appear to influence biomass size-spectra, because small organisms are better able to satisfy their metabolic demands. The annual secondary production was higher off central Chile (6.8 g C m⁻² y⁻¹) than off northern Chile (2.02 g C m⁻² y⁻¹) and off southern Chile (0.83 g C m⁻² y⁻¹). A comparison with other studies suggests that secondary production in terms of carbon equivalents was higher than in other upwelling regions.     

Meiofaunal distributions on the Peru margin:: relationship to oxygen and organic matter availability

A quantitative study of metazoan meiofauna was carried out on bathyal sediments (305, 562, 830 and 1210 m) along a transect within and beneath the oxygen minimum zone (OMZ) in the southeastern Pacific off Callao, Peru (12°S). Meiobenthos densities ranged from 1517 (upper slope, middle of OMZ) to 440–548 ind. 10 cm⁻² (lower slope stations, beneath the OMZ). Nematodes were the numerically dominant meiofaunal taxon at every station, followed by copepods and nauplii. Increasing bottom-water oxygen concentration and decreasing organic matter availability downslope were correlated with observed changes in meiofaunal abundance. The 300-m site, located in the middle of the OMZ, differed significantly in meiofaunal abundance, dominance, and in vertical distribution pattern from the deeper sites. At 305 m, nematodes amounted to over 99% of total meiofauna; about 70% of nematodes were found in the 2–5 cm interval. At the deeper sites, about 50% were restricted to the top 1 cm. The importance of copepods and nauplii increased consistently with depth, reaching ∼12% of the total meiofauna at the deepest site. The observation of high nematode abundances at oxygen concentrations <0.02 ml l⁻¹ supports the hypothesis that densities are enhanced by an indirect positive effect of low oxygen involving (a) reduction of predators and competitors and (b) preservation of organic matter leading to high food availability and quality. Food input and quality, represented here by chloroplastic pigment equivalents (CPE) and sedimentary labile organic compounds (protein, carbohydrates and lipids), were strongly, positively correlated with nematode abundance. By way of contrast, oxygen exhibited a strong negative correlation, overriding food availability, with abundance of other meiofauna such as copepods and nauplii. These taxa were absent at the 300-m site. The high correlation of labile organic matter (C-LOM, sum of carbon contents in lipids, proteins and carbohydrates) with CPE (Pearson's r=0.99, p<0.01) suggests that most of the sedimentary organic material sampled was of phytodetrital origin. The fraction of sediment organic carbon potentially available to benthic heterotrophs, measured as C-LOM/Total organic carbon, was on average 17% at all stations. Thus, a residual, refractory fraction, constitutes the major portion of organic matter at the studied bathyal sites.     

Nitrous oxide and N-nutrient cycling in the oxygen minimum zone off northern Chile

Measurements of dissolved gases (O₂, N₂O), nutrients (NO₃⁻, NO₂⁻, PO₄³⁻), and oceanographic variables were performed off northern Chile (∼21°S) between March 2000 and July 2004, in order to characterize the existing oxygen minimum zone (OMZ) and identify processes involved in N₂O cycling. Both N₂O and NO₃⁻ displayed sharp, shallow peaks with concentrations of up to 124 nM (1370% saturation) and 26 μM, respectively, in association with a strong oxycline that impinges on the euphotic zone. NO₂⁻ accumulation below the oxycline's base reached up to 9 μM. The vertical distribution of physical and chemical parameters and the existing relationships between apparent oxygen utilization (AOU), apparent N₂O production (ΔN₂O), and NO₃⁻ revealed three main layers within the upper OMZ. The first layer, or the upper part of the oxycline, is located between the base of the mixed layer and the mid-point of the oxycline (around σ_t=25.5 kg m⁻³). There the O₂ declines from ∼250 to ∼50 μM, and strong (but opposing) O₂ and NO₃⁻ gradients and their associated AOU–ΔN₂O and AOU–NO₃⁻ relationships indicate that nitrification produces N₂O and NO₃⁻ in the presence of light. The second layer, or lower part of the oxycline, represents the upper OMZ boundary and is located between the middle and the base of the oxycline (25.9<σ_t<26.1 kg m⁻³). In this layer NO₃⁻ reduction begins at O₂ levels ranging from ∼50 to ∼11 μM and accumulation of 41–68% of the ΔN₂O pool occurs. The accumulation of N₂O (but not of NO₂⁻ or NH₄⁺) and the observed AOU–ΔN₂O and AOU–NO₃⁻ relationships (which are opposite to those of the overlying first layer) suggest that a coupling between nitrification and NO₃⁻ reduction is involved in N₂O cycling in this second layer. The third layer is the OMZ core, where the O₂ concentration remains constant (O₂<11 μM). It coincides with σ_t>26.2 kg m⁻³, which is typical of Equatorial Subsurface Water (ESSW). In this layer, N₂O and NO₃⁻ continue to decrease, but a large NO₂⁻ accumulation is observed. Considering all the data, a biogeochemical model for the upper OMZ off northern of Chile is proposed, in which nitrification and denitrification differentially mediate N₂O cycling in each layer.     

Aerobic ammonium oxidation in the oxycline and oxygen minimum zone of the eastern tropical South Pacific off northern Chile (∼20°S)

Aerobic NH₄⁺ oxidation rates were measured along the strong oxygen gradient associated with the oxygen minimum zone (OMZ) of the eastern tropical South Pacific off northern Chile (∼20°S) during 2000, 2003, and 2004. This process was examined by comparing NH₄⁺ rates of change during dark incubations, with and without the addition of allylthiourea, a classical inhibitor of the ammonia monooxygenase enzyme of ammonium-oxidizing bacteria. The contribution of aerobic NH₄⁺ oxidation in dark carbon fixation and NO₂⁻ rates of change were also explored. Thirteen samples were retrieved from the oxycline (252 to ⩽5 μM O₂; 15 to ∼65 m depth) and three from the oxygen minimum core (⩽5 μM O₂; 100–200 m depth). Aerobic NH₄⁺ oxidation rates were mainly detected in the upper part (15–30 m depth) of the oxycline, with rates ranging from 0.16 to 0.79 μM d⁻¹, but not towards the oxycline base (40–65 m depth). In the oxygen minimum core, aerobic NH₄⁺ oxidation was in the upper range and higher than in the upper part of the oxycline (0.70 and 1.0 μM d⁻¹). Carbon fixation rates through aerobic NH₄⁺ oxidation ranged from 0.18 to 0.43 μg C L⁻¹ d⁻¹ and contributed between 33% and 57% of the total dark carbon fixation, mainly towards the oxycline base and, in a single experiment, in the upper part of the oxycline. NO₂⁻ consumption was high (up to 10 μM d⁻¹) towards the oxycline base and OMZ core, but was significantly reduced in experiments amended with allylthiourea, indicating that aerobic NH₄⁺ oxidation could contribute between 8% and 76% of NO₂⁻ production, which in turn could be available for denitrifiers. Overall, these results support the important role of aerobic NH₄⁺ oxidizers in the nitrogen and carbon cycling in the OMZ and at its upper boundary.     

Distribution of bacterial abundance and cell-specific nucleic acid content in the Northeast Pacific Ocean

We tested the idea that bacterial cells with high nucleic acid content (HNA cells) are the active component of marine bacterioplankton assemblages, while bacteria with low nucleic acid content (LNA cells) are inactive, with a large data set (>1700 discrete samples) based on flow cytometric analysis of bacterioplankton in the Northeast Pacific Ocean off the coast of Oregon and northern California, USA. Samples were collected in the upper 150 m of the water column from the coast to 250 km offshore during 14 cruises from March 2001 to September 2003. During this period, a wide range of trophic states was encountered, from dense diatom blooms (chlorophyll-a concentrations up to 43 μg l⁻¹) at shelf stations during upwelling season (March–September) to lower chlorophyll-a concentrations (0.1–5 μg l⁻¹) during winter (November–February) and at basin stations (>1700 m depth). We found only weakly positive relations of log total bacterial abundance to log chlorophyll-a concentration (as a proxy for availability of organic substrate), and of HNA bacteria as a fraction of total bacteria to log chlorophyll-a. Abundance of HNA and LNA bacteria co-varied positively in all regions, although HNA bacteria were more responsive to high phytoplankton biomass in shelf waters than in slope and basin waters. Since LNA cell abundance in general showed responses similar to those of HNA cell abundance to changes in phytoplankton biomass, our data do not support the hypothesis that HNA cells are the sole active component of marine bacterioplankton.     

Export fluxes of particulate organic carbon in the Chukchi Sea: A comparative study using 234Th/238U disequilibria and drifting sediment traps

Large-volume sampling of ²³⁴Th and drifting sediment trap deployments were conducted as part of the 2004 Western Arctic Shelf–Basin Interactions (SBI) spring (May 15–June 23) and summer (July 17–August 26) process cruises in the Chukchi Sea. Measurements of ²³⁴Th and particulate organic carbon (POC) export fluxes were obtained at five stations during the spring cruise and four stations during the summer cruise along Barrow Canyon (BC) and along a parallel shelf-to-basin transect from East Hanna Shoal (EHS) to the Canada Basin. ²³⁴Th and POC fluxes obtained with in situ pumps and drifting sediment traps agreed to within a factor of 2 for 70% of the measurements. POC export fluxes measured with in situ pumps at 50 m along BC were similar in spring and summer (average = 14.0 ± 8.0 mmol C m^− 2 day^− 1 and 16.5 ± 6.5 mmol C m^− 2 day^− 1, respectively), but increased from spring to summer at the EHS transect (average = 1.9 ± 1.1 mmol C m^− 2 day^− 1 and 19.5 ± 3.3 mmol C m^− 2 day^− 1, respectively). POC fluxes measured with sediment traps at 50 m along BC were also similar in both seasons (31.3 ± 9.3 mmol C m^− 2 day^− 1 and 29.1 ± 14.2 mmol C m^− 2 day^− 1, respectively), but were approximately twice as high as POC fluxes measured with in situ pumps. Sediment trap POC fluxes measured along the EHS transect also increased from spring to summer (3.0 ± 1.9 mmol C m^− 2 day^− 1 and 13.0 ± 6.4 mmol C m^− 2 day^− 1, respectively), and these fluxes were similar to the POC fluxes obtained with in situ pumps. Discrepancies in POC export fluxes measured using in situ pumps and sediment traps may be reasonably explained by differences in the estimated POC/²³⁴Th ratios that arise from differences between the techniques, such as time-scale of measurement and size and composition of the collected particles. Despite this variability, in situ pump and sediment trap-derived POC fluxes were only significantly different at a highly productive station in BC during the spring.     

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.     

Polychaete community structure in the South Eastern Arabian Sea continental margin (200–1000 m)

Macrofaunal polychaete communities (>500 µm) in the South Eastern Arabian Sea (SEAS) continental margin (200–1000 m) are described, based on three systematic surveys carried out in 9 transects (at ~200 m, 500 m and 1000 m) between 7°00′and 14°30′N latitudes. A total of 7938 polychaetes belonging to 195 species were obtained in 136 grab samples collected at 27 sites. Three distinct assemblages were identified in the northern part of the SEAS margin (10–14°30′N), occupying the three sampled depth strata (shelf edge, upper and mid-slope) and two assemblages (shelf edge and slope) in the south (7–10°N). Highest density of polychaetes and dominance of a few species were observed in the shelf edge, where the Arabian Sea oxygen minimum zone (OMZ) impinged on the seafloor, particularly in the northern transects. The resident fauna in this region (Cossura coasta, Paraonis gracilis, Prionospio spp. and Tharyx spp.) were characteristically of smaller size, and well suited to thrive in the sandy sediments in OMZ settings. Densities were lowest along the most northerly transect (T9), where dissolved oxygen (DO) concentrations were extremely low (<0.15 ml l⁻¹, i.e.<6.7 μmol l⁻¹). Beyond the realm of influence of the OMZ (i.e. mid-slope, ~1000 m), the faunal density decreased while species diversity increased. The relative proportion of silt increased with depth, and the dominance of the aforementioned species decreased, giving way to forms such as Paraprionospio pinnata, Notomastus sp., Eunoe sp. and lumbrinerids. Relatively high species richness and diversity were observed in the sandy sediments of the southern sector (7–9°N), where influence of the OMZ was less intense. The area was also characterized by certain species (e.g. Aionidella cirrobranchiata, Isolda pulchella) that were nearly absent in the northern region. The gradients in DO concentration across the core and lower boundary of the OMZ, along with bathymetric and latitudinal variation in sediment texture, were responsible for differences in polychaete size and community structure on the SEAS margin. Spatial and temporal variations were observed in organic matter (OM) content of the sediment, but these were not reflected in the density, diversity or distribution pattern of the polychaetes.     

Compositional and fluorescence characteristics of the giant diatom Ethmodiscus along a 3000 km transect (28°N) in the central North Pacific gyre

The giant diatom Ethmodiscus was examined along an east–west transect at 28–30°N during 2002 and 2003 to determine if abundance, chemical composition or physiological status of this largest of diatoms varied on the scale of 100's–1000's of km in North Pacific gyre. Abundance ranged from <0.1–>2.0 cells m⁻³ and supported the notion of an abundance mosaic reported previously. However, there was only minimal support for the relationship between abundance and nutrient concentration at 125 m reported previously. Cellular chlorophyll varied little along the transect (7.3–10.9 ng chl cell⁻¹) except at the westernmost station. Cellular N and P quotas co-varied 3–4.5 fold (mean=50.8±3.7 and 3.7±0.8 nmol N and P cell⁻¹) and yielded N:P ratios that closely clustered around the Redfield ratio (average=14.6±1.1). Only low levels of chlorophyll-normalized alkaline phosphatase (APase) activity were observed (0.4–2.5 nmol P μg chl⁻¹ h⁻¹) with APase activity lower than that in either the bulk water, or co-occurring Trichodesmium spp. and Pyrocystis noctiluca. The active fluorescence parameter F_v:F_m, a property sensitive to Fe stress, was uniformly high at all stations (average=0.73±0.04 for 2003, and 0.69±0.05 for 2002), indicating sufficient Fe for optimum photosynthetic competence. These results contrasted sharply with results from Rhizosolenia mats reported along the same transect where there was a significant decline westward in F_v:F_m. Both ferredoxin (Fd) and flavodoxin accumulated in cells of Ethmodiscus, resulting in Fd Index values of<0.6. Iron cell quotas ranged from 0.7–5.1 pmol Fe cell⁻¹. When normalized to cytoplasmic volume, the Fe μm⁻³ was comparable to that of Escherichia coli. We note that the disproportionate contribution of the vacuole (with its high organic content) to total volume typical of large diatoms is a potentially significant source of error in Fe:C ratios and suggest that Fe should be normalized to cytoplasmic volume whenever possible to permit valid intercomparisons between studies. The composition, F_v:F_m data and Fe:C ratio suggest a relatively uniform population experiencing little N, P or Fe stress. The uncoupling of the Fd Index from these measures is consistent with previous findings showing that the expression of flavodoxin can be characterized as an early stress response and that its accumulation is not necessarily correlated with physiological deficit. Ethmodiscus appears to be well adapted to some of the most oligotrophic waters in the ocean. Because it is an important sedimentary marker, the biology of living Ethmodiscus provides insights into the source of extensive Ethmodiscus oozes. Mass sedimentation after frontal accumulation has been suggested as a source for these oozes. Our data contain no evidence that the flux is linked directly to Fe, N or P stress.     

Ectoenzymatic activity in surface waters: A transect from the Mediterranean Sea across the Indian Ocean to Australia

The activities of two hydrolytic enzymes (leucine aminopeptidase and β glucosidase), belonging to the particle-bound enzymatic fraction, were measured in open-sea surface waters. Samples were collected along a transect crossing the Indian Ocean during the early NW monsoon period (November and December 2001). The latitudinal pattern of the ectoenzymatic activities highlighted a generally increasing trend of glycolysis approaching the equator, with significantly higher β glucosidase activity (0.79–3.00 nmol l⁻¹ h⁻¹) within the latitudinal range from 12°N to 16°S. In this area, the surface waters coming from the Indonesian Throughflow and the Bay of Bengal carry a considerable quantity of carbohydrates (38.9–41.9 μg l⁻¹), which stimulated glycolytic activity and its cell-specific rates scaled to bacterial abundance. On the other hand, in the Central Indian Ocean, the proteolytic activity was considerable (0.91–2.03 nmol l⁻¹ h⁻¹), although the particulate proteins did not show significant increases and the dissolved proteinlike signal was one of the lowest of the entire transect (0.7 mg l⁻¹ on average compared to the 1.4–1.6 mg l⁻¹ of the adjacent areas). Therefore, in this area, the two ectoenzymes studied did not respond to the same stimulatory effect (namely the specific substrate concentrations). The time needed for the hydrolysis of macromolecules within the particulate and dissolved organic substrate fractions, although these measures are affected by a number of assumptions starting with the potential nature of the ectoenzymatic determinations, confirms these observations. The Central Indian Ocean displayed the lowest values, from 8 to 26 days for particulate and dissolved organic carbon, respectively. As observed in the equatorial areas of the Atlantic Ocean, the relevant degradation activity of the central area of the Indian Ocean Basin suggests a notable heterotrophy based on a faster turnover of organic substrates.     

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.     

Phytoplankton chemotaxonomy in the Atlantic sector of the Southern Ocean during late summer 2009

A chemotaxonomic investigation of surface phytoplankton was undertaken on a research cruise to the Atlantic sector of the Southern Ocean during late austral summer 2009. Based on pigment signatures, several distinct regions emerged that were delineated by physical features. CHEMTAX analysis of high performance liquid chromatography (HPLC) pigment data indicated that diatoms generally dominated communities south of the Antarctic Polar Front (APF), particularly in regions of elevated biomass where chlorophyll-a (chl-a) was >1.5 µg l⁻¹ and diatoms comprised >80% of biomass. Pigment signatures representative of haptophytes-8, indicative of Phaeocystis antarctica, were dominant near the ice shelf. Chl-a concentrations were 0.2–0.6 µg l⁻¹ between the APF and the Subtropical Front (STF) and outputs suggested that chlorophytes, haptophytes-8 and haptophyte-6, in the form of coccolithophores, were the major constituents. Very low chl-a levels (<0.2 µg l⁻¹) were observed north of the STF and the prokaryotes Synechococcus spp. and Prochlorococcus spp. were the dominant groups in these oligotrophic waters.     

Phytoplankton growth and mortality during the 1995 Northeast Monsoon and Spring Intermonsoon in the Arabian Sea

Phytoplankton growth rates and mortality rates were experimentally examined at eight stations in the Arabian Sea along the U.S. JGOFS cruise track during the 1995 Northeast Monsoon (January) and Spring Intermonsoon (March–April). Instantaneous growth rates averaged over an entire cruise were approximately twice as high during the NE Monsoon than during the Spring Intermonsoon period (overall averages of 0.84±0.29 (s.d.) versus 0.44±0.19 d⁻¹). Average herbivore grazing (mortality) rates, however, were quite similar for the two seasons (overall averages of 0.35±0.18 and 0.30±0.17 d⁻¹ for the NE Monsoon and Spring Intermonsoon, respectively). The absolute amounts of phytoplankton biomass consumed during each season also were similar (29 and 25% of standing stock consumed d⁻¹ for the January and March–April cruises, respectively), as were the geographical trends of this removal. These seasonal trends in growth and removal rates resulted in net phytoplankton growth rates that were considerably higher during the January cruise (0.48 d⁻¹) than during the March–April cruise (0.14 d⁻¹). That is, phytoplankton production was more closely balanced during the Spring Intermonsoon season (87% of daily primary production consumed) relative to the NE Monsoon season (49% of daily primary production consumed). Station-to-station variability was high for rate measurements during either cruise. Nevertheless, there was a clear onshore–offshore trend in the absolute rate of removal of phytoplankton biomass (μg chlorophyll consumed l⁻¹ d⁻¹) during both cruises. Coastal stations had removal rates that were typically 2–4 times higher than removal rates at oceanic stations.     

Picophytoplankton in the equatorial Pacific: vertical distributions in the warm pool and in the high nutrient low chlorophyll conditions

Abundance distribution and cellular characteristics of picophytoplankton were studied in two distinct regions of the equatorial Pacific: the western warm pool (0°, 167°E), where oligotrophic conditions prevail, and the equatorial upwelling at 150°W characterized by high-nutrient low-chlorophyll (HNLC) conditions. The study was done in September–October 1994 during abnormally warm conditions. Populations of Prochlorococcus, orange fluorescing Synechococcus and picoeukaryotes were enumerated by flow cytometry. Pigment concentrations were studied by spectrofluorometry. In the warm pool, Prochlorococcus were clearly the dominant organisms in terms of cell abundance, estimated carbon biomass and measured pigment concentration. Integrated concentrations of Prochlorococcus, Synechococcus and picoeukaryotes were 1.5×10¹³, 1.3×10¹¹ and 1.5×10¹¹ cells m⁻², respectively. Integrated estimated carbon biomass of picophytoplankton was 1 g m⁻², and the respective contributions of each group to the biomass were 69, 3 and 28%. In the HNLC waters, Prochlorococcus cells were slightly less numerous than in the warm pool, whereas the other groups were several times more abundant (from 3 to 5 times). Abundance of Prochlorococcus, Synechococcus and picoeukaryotes were 1.2×10¹³, 6.2×10¹¹ and 5.1×10¹¹ cells m⁻², respectively. The integrated biomass was 1.9 g C m⁻². Prochlorococcus was again the dominant group in terms of abundance and biomass (chlorophyll, carbon); the respective contributions of each group to the carbon biomass were 58, 7 and 35%. In the warm pool the total chlorophyll biomass was 28 mg m⁻², 57% of which was divinyl chlorophyll a. In the HNLC waters, the total chlorophyll biomass was 38 mg m⁻², 44% of which was divinyl chlorophyll a. Estimates of Prochlorococcus, Synechococcus and picoeukaryotes cell size were made in both hydrological conditions.     

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.     

Deep-sea macrofaunal assemblages within the Benthic Boundary Layer of the Cap-Ferret Canyon (Bay of Biscay,NE Atlantic)

The Benthic Boundary Layer (BBL) assemblages from the Cap-Ferret Canyon (Bay of Biscay) were quantitatively sampled at two sites located within its main channel near mooring deployments (Mooring Sites MS 1: ca. 2400 m; MS 2: ca. 3000 m) with a suprabenthic sled equipped with four nets fishing at different heights above the bottom. The macrofaunal abundance above the sea-floor was mainly represented by Isopoda (42.2%), Amphipoda (19.0%), Euphausiacea (17.3%), Cumacea (13.5%), Mysidacea (2.8%) and Tanaidacea (2.6%). At both sampling sites, the highest total densities were generally recorded in the immediate vicinity of the sea floor (10–40 cm water layer), and a drastic decrease occurred higher in the BBL community. The BBL assemblages from the two sampling sites were similar in their faunal composition (major taxa), and their mean density estimates were not statistically different (MS 1 : 525.3 ind. 100 m⁻²; MS 2 : 283.3 ind. m⁻²) although the recorded values during each cruise were always lower at the deeper site. The BBL macrofauna abundance showed obvious temporal fluctuations at both sites, probably linked with a seasonal organic input from the euphotic zone (vertical flux) via phytodetritus deposition on the sea bottom.     

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.     

The seasonal succession of zooplankton in the Southern Ocean south of Australia,part II: The Sub-Antarctic to Polar Frontal Zones

Between October 2001 and March 2002 six transects were completed at monthly intervals in the Sub-Antarctic Zone (SAZ) and Inter-Sub-Antarctic Front Zone (ISAFZ)/Polar Frontal Zone (PFZ) in the Southern Ocean south of Australia. Zooplankton were collected with a Continuous Plankton Recorder and NORPAC net and multivariate analysis was used to analyse the seasonal succession of communities. Despite strong, seasonally consistent, biogeographic differences between the SAZ and ISAFZ/PFZ, community structure in all zones was dominated by a suite of common taxa. These included the ubiquitous Oithona similis, foraminiferans and appendicularians (Core taxa), occurring in >97% of samples and contributing an average of 75% to total sample abundance, and Calanus simillimus, Rhincalanus gigas, Ctenocalanus citer, Clausocalanus brevipes, Clausocalanus laticeps, Oithona frigida, Limacina spp. and chaetognaths (Summer taxa), present in >57% of samples and occurring at seasonally high densities. Because of the dominance of the Core and Summer taxa, the seasonal succession was most clearly evident as a change in zooplankton densities. In October densities averaged <15 ind m⁻³, rising to 52 ind m⁻³ (max=92 ind m⁻³) in November, and subsequently increasing slowly through to January (ave=115 ind m⁻³; max=255 ind m⁻³). Densities peaked abruptly in February (ave=634 ind m⁻³; max=1593 ind m⁻³), and remained relatively high in March (ave=193 ind m⁻³; max=789 ind m⁻³). A latitudinal lag in seasonal development was observed with peak densities occurring first in the SAZ (February) and then in the ISAFZ/PFZ (March). The seasonal community succession was strongly influenced by species population cycles. The role of zooplankton in biogeochemical cycling in the SAZ and ISAFZ/PFZ was discussed in the light of past sediment trap data collected from the study area.     

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.