Particle fluxes associated with mesoscale eddies in the Sargasso Sea

We examined the impact of a cyclonic eddy and mode-water eddy on particle flux in the Sargasso Sea. The primary method used to quantify flux was based on measurements of the natural radionuclide, ²³⁴Th, and these flux estimates were compared to results from sediment traps in both eddies, and a ²¹⁰Po/²¹⁰Pb flux method in the mode-water eddy. Particulate organic carbon (POC) fluxes at 150 m ranged 1–4 mmol C m⁻² d⁻¹ and were comparable between methods, especially considering differences in integration times scales of each approach. Our main conclusion is that relative to summer mean conditions at the Bermuda Atlantic Time-series Study (BATS) site, eddy-driven changes in biogeochemistry did not enhance local POC fluxes during this later, more mature stage of the eddy life cycle (>6 months old). The absence of an enhancement in POC flux puts a constraint on the timing of higher POC flux events, which are thought to have caused the local O₂ minima below each eddy, and must have taken place >2 months prior to our arrival. The mode-water eddy did enhance preferentially diatom biomass in its center, where we estimated a factor of three times higher biogenic Si flux than the BATS summer average. An unexpected finding in the highly depth-resolved ²³⁴Th data sets is narrow layers of particle export and remineralization within the eddy. In particular, a strong excess ²³⁴Th signal is seen below the deep chlorophyll maxima, which we attribute to remineralization of ²³⁴Th-bearing particles. At this depth below the euphotic zone, de novo particle production in the euphotic zone has stopped, yet particle remineralization continues via consumption of labile sinking material by bacteria and/or zooplankton. These data suggest that further study of processes in ocean layers is warranted not only within, but below the euphotic zone.     

Zooplankton community structure in a cyclonic and mode-water eddy in the Sargasso Sea

Mesoscale eddies are important suppliers of nutrients to the surface waters of oligotrophic gyres, but little is known about the biological response, particularly that of higher trophic levels, to these physical perturbations. During the summers of 2004 and 2005, we followed the development of a cyclonic eddy and an anti-cyclonic mode-water eddy in the Sargasso Sea. Zooplankton (>150 μm) were collected across both eddies in 9 discrete depth intervals between 0 and 700 m. Comparison of the abundance of major taxa of mesozooplankton in the upper 150 m at eddy center and outside the eddies (day and night) indicated that the cyclone and mode-water eddy supported similar mesozooplankton communities, with several taxa significantly higher in abundance inside than outside the eddies, when compared with the Bermuda Atlantic Time-series Study site as representative of mean conditions. In both eddies copepod peak abundance occurred in the 50-100 m depth interval, coincident with the chlorophyll a maximum, suggesting elevated food concentration in the eddies may have influenced zooplankton vertical distribution. The two eddies differed in the strength of diel vertical migration of zooplankton, as indicated by the ratio of night:day abundance in the epipelagic zone, which was higher at the center of the mode-water eddy for most taxa. Over the sampling interval of 1-2 months, abundance of the three most common taxa (copepods, chaetognaths, and ostracods) decreased in the cyclone and increased in the mode-water eddy. This further supports previous findings that over the sampling period the cyclone was in a decay phase, while the mode-water eddy was sustaining nutrient fluxes and high phytoplankton concentrations. A more detailed analysis of community structure in the mode-water eddy indicated the 0-700 m integrated abundance of doliolids was significantly higher inside the mode-water eddy than outside. The presence of a mesopelagic (200-700 m) layer of lepadid barnacle cyprids in this eddy highlights the potential of eddies to transport and disperse biota. We conclude that when compared with average ambient conditions (as measured at BATS), mesoscale eddies can influence zooplankton behavior and alter zooplankton community structure which can affect food-web interactions and biogeochemical cycling in the open ocean.     

Shear and Richardson number in a mode-water eddy

Measurements of stratification and shear were carried out as part of the EDDIES tracer release experiment in mode-water eddy A4 during the summer of 2005. These measurements were accomplished using both shipboard instrumentation and a drifting mooring. A strong relationship between shear intensity and distance from the center of the eddy A4 was observed with the shipboard ADCP. Diapycnal diffusivity at the SF₆ tracer isopycnal prior to and during the release was estimated from the drifting mooring to be 2.9×10⁻⁶ m² s⁻¹. Diffusivity increased by an order of magnitude to 3.2×10⁻⁵ m² s⁻¹ during the period of the final tracer survey in early September, which was similar to the value estimated from the tracer analysis for the whole experiment (3.5×10⁻⁵ m² s⁻¹, [Ledwell, J.R., McGillicuddy Jr., D.J., Anderson, L.A., 2008. Nutrient flux into an intense deep chlorophyll layer in a mode-water eddy. Deep-Sea Research II, this issue [doi:10.1016/j.dsr2.2008.02.005]].     

Influence of basin-scale and mesoscale physical processes on biological productivity in the Bay of Bengal during the summer monsoon

Physical forcing plays a major role in determining biological processes in the ocean across the full spectrum of spatial and temporal scales. Variability of biological production in the Bay of Bengal (BoB) based on basin-scale and mesoscale physical processes is presented using hydrographic data collected during the peak summer monsoon in July–August, 2003. Three different and spatially varying physical processes were identified in the upper 300 m: (I) anticyclonic warm gyre offshore in the southern Bay; (II) a cyclonic eddy in the northern Bay; and (III) an upwelling region adjacent to the southern coast. In the warm gyre (>28.8 °C), the low salinity (33.5) surface waters contained low concentrations of nutrients. These warm surface waters extended below the euphotic zone, which resulted in an oligotrophic environment with low surface chlorophyll a (0.12 mg m⁻³), low surface primary production (2.55 mg C m⁻³ day⁻¹) and low zooplankton biovolume (0.14 ml m⁻³). In the cyclonic eddy, the elevated isopycnals raised the nutricline upto the surface (NO₃–N > 8.2 μM, PO₄–P > 0.8 μM, SiO₄–Si > 3.5 μM). Despite the system being highly eutrophic, response in the biological activity was low. In the upwelling zone, although the nutrient concentrations were lower compared to the cyclonic eddy, the surface phytoplankton biomass and production were high (Chl a – 0.25 mg m⁻³, PP – 9.23 mg C m⁻³ day⁻¹), and mesozooplankton biovolume (1.12 ml m⁻³) was rich. Normally in oligotrophic, open ocean ecosystems, primary production is based on ‘regenerated’ nutrients, but during episodic events like eddies the ‘production’ switches over to ‘new production’. The switching over from ‘regenerated production’ to ‘new production’ in the open ocean (cyclonic eddy) and establishment of a new phytoplankton community will take longer than in the coastal system (upwelling). Despite the functioning of a cyclonic eddy and upwelling being divergent (transporting of nutrients from deeper waters to surface), the utilization of nutrients leading to enhanced biological production and its transfer to upper trophic levels in the upwelling region imply that the energy transfer from primary production to secondary production (mesozooplankton) is more efficient than in the cyclonic eddy of the open ocean. The results suggest that basin-scale and mesoscale processes influence the abundance and spatial heterogeneity of plankton populations across a wide spatial scale in the BoB. The multifaceted effects of these physical processes on primary productivity thus play a prominent role in structuring of zooplankton communities and could consecutively affect the recruitment of pelagic fisheries.     

Mesozooplankton biomass and grazing responses to Cyclone Opal, a subtropical mesoscale eddy

As part of E-Flux III cruise studies in March 2005, plankton net collections were made to assess the effects of a cyclonic cold-core eddy (Cyclone Opal) on the biomass and grazing of mesozooplankton. Mesozooplankton biomass in the central region of Cyclone Opal, an area of uplifted nutricline and a subsurface diatom bloom, averaged 0.80±0.24 and 1.51±0.59 g DW m⁻², for day and night tows, respectively. These biomass estimates were about 80% higher than control (OUT) stations, with increases more or less proportionately distributed among size classes from 0.2 to >5 mm. Though elevated relative to surrounding waters south of the Hawaiian Islands (Hawai’i lee), total biomass and size distribution in Cyclone Opal were almost exactly the same as contemporary measurements made at Stn. ALOHA, 100 km north of the islands, by the HOT (Hawaii Ocean Time-series) Program. Mesozooplankton biomass and community composition at the OUT stations were also similar to ALOHA values from 1994 to 1996, preceding a recent decadal increase. These comparisons may therefore provide insight into production characteristics or biomass gradients associated with decadal changes at Stn. ALOHA. Gut fluorescence estimates were higher in Opal than in ambient waters, translating to grazing impacts of 0.11±0.02 d⁻¹ (IN) versus 0.03±0.01 d⁻¹ (OUT). Over the depth-integrated euphotic zone, mesozooplankton accounted for 30% of the combined grazing losses of phytoplankton to micro- and meso-herbivores in Opal, as compared to 13% at control stations. Estimates of active export flux by migrating zooplankton averaged 0.81 mmol C m⁻² d⁻¹ in Cyclone Opal and 0.37 mmol C m⁻² d⁻¹ at OUT stations, 53% and 24%, respectively, of the carbon export measured by passive sediment traps. Migrants also exported 0.18 mmol N m⁻² d⁻¹ (117% of trap N flux) in Cyclone Opal compared to 0.08 mmol N m⁻² d⁻¹ (51% of trap flux) at control stations. Overall, the food-web importance of mesozooplankton increased in Cyclone Opal both in absolute and relative terms. Diel migrants provided evidence for enhanced export flux in the eddy that was missed by sediment trap and ²³⁴Th techniques, and migrant-mediated flux was the major export term in the observed bloom-perturbation response and N mass balance of the eddy.     

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.     

The influence of island generated eddies on the carbon dioxide system, south of the Canary Islands

Measurements of surface partial pressure of CO₂ and water column alkalinity, pH_T, nutrients, oxygen, fluorescence and hydrography were carried out, south of the Canary Islands during September 1998. Cyclonic and anticyclonic eddies were alternatively observed from the northwestern area to the central area of the Canary Islands. Nutrient pumping and vertical uplifting of the deep chlorophyll maximum by cyclonic eddies were also ascertained by upward displacement of dissolved inorganic carbon. A model was applied to determine the net inorganic carbon balance in the cyclonic eddy. The fluxes were determined considering both the diffusive and convective contributions from the upward pumping and the corresponding horizontal transport of water outside the area. An increase in the total inorganic carbon concentration in the upper layers inside the eddy field of 133 mmol C m^− 2 d^− 1 was determined. The upward flux of inorganic carbon decreased the effect of the increased primary production on the carbon dioxide chemistry. The reduced fCO₂ inside the cyclonic eddy, 15 μatm lower than that observed in non-affected surface water, was explained by thermodynamic aspects, biological activity, eddy upward pumping and diffusion and air–sea water exchange effects.     

Biogeochemical impacts due to mesoscale eddy activity in the Sargasso Sea as measured at the Bermuda Atlantic Time-series Study (BATS)

A comparison of monthly biogeochemical measurements made from 1993 to 1995, combined with hydrography and satellite altimetry, was used to assess the impacts of nine eddy events on primary productivity and particle flux in the Sargasso Sea. Measurements of primary production, thorium-234 flux, nitrate+nitrite, and photosynthetic pigments made at the US JGOFS Bermuda Atlantic Time-series Study (BATS) site were used. During the 3 years of this study, four out of six high thorium-234 flux events occurred during the passage of an eddy. Primary production nearly as high as the spring bloom maximum was observed in two mode-water eddies (May 1993 and July 1995). The 1994 spring bloom at BATS was suppressed by the passage of an anticyclone. Distinct phytoplankton community shifts were observed in mode-water eddies, which had an increased percentage of diatoms and dinoflagellates, and in cyclones, which had an increased percentage of Synechococcus. These variations in species composition within mode-water eddies and cyclones may be associated with the ages of the sampled eddies, and/or differences in physical, chemical, and biological factors in these two distinct eddy types. In general, eddies that were 1–2 months old elicited a large biological response; eddies that were 3 months old may show a biological response and were accompanied by high thorium flux; eddies that were 4 months old or older did not show a biological response or high thorium flux. A conceptual model depicting temporal changes during eddy upwelling, maturation, and decay can explain the observations in all seven upwelling eddies present in the time-series investigated herein.     

The influence of a mature cyclonic eddy on particle export in the lee of Hawaii

Mesoscale eddies may enhance primary production (PP) in the open ocean by bringing nutrient-rich deep waters into the euphotic zone, potentially leading to increased transport of particles to depth. This hypothesis remains controversial, however, due to a paucity of direct particle export measurements. In this study, we investigated particle dynamics using ²³⁴Th–²³⁸U disequilibria within a mesoscale cold-core eddy, Cyclone Opal, which formed in the lee of the Hawaiian Islands. ²³⁴Th samples were collected along two transects across Cyclone Opal as well as during a time-series within the eddy core during a decaying diatom bloom. Particulate carbon (PC), particulate nitrogen (PN) and biogenic silica (bSiO₂) fluxes at 150 m varied spatially and temporally within the eddy and strongly depended on the ²³⁴Th model formulation used (e.g., steady state versus non-steady state, inclusion of upwelling, etc.). Particle fluxes estimated from a steady state model assuming an upwelling rate of 2 m day⁻¹ yielded the best fit to sediment-trap data. These ²³⁴Th-derived particle fluxes ranged from 332±14 to 1719±53 μmol C m⁻² day⁻¹, 27±3 to 114±12 μmol N m⁻² day⁻¹, and 33±20 to 309±73 μmol Si m⁻² day⁻¹. Although PP rates within Cyclone Opal were elevated by a factor of 2–3, PC and PN fluxes were the same, within error, inside and outside of Cyclone Opal. The ratio of PC export to PP remained surprisingly low at <0.03 and similar to those measured in surrounding waters. In contrast, bSiO₂ fluxes within the eddy core were three times higher. Detailed analyses of ²³⁴Th depth profiles consistently showed excess ²³⁴Th at 100–175 m, associated with the remineralization and possible accumulation of suspended and dissolved organic matter from the surface. We suggest that strong microzooplankton grazing facilitated particulate organic matter recycling and resulted in the export of empty diatom frustules. Thus, while eddies may increase PP, they do not necessarily increase PC and PN export to deep waters. This may be a general characteristic of wind-driven cyclonic eddies of the North Pacific Subtropical Gyre and suggests that eddies may preferentially act as a silica pump, thereby playing an important role in promoting silicic-acid limitation in the region.     

Flux of Reduced Sulfur Gases Along a Salinity Gradient in Louisiana Coastal Marshes

Seasonal and diurnal reduced sulfur gas emissions were measured along a salinity gradient in Louisiana Gulf Coast salt, brackish and freshwater marshes. Reduced sulfur gas emission was strongly associated with habitat and salinity gradient. The dominant emission component was dimethyl sulfide (average: 57·3 μg S m⁻² h⁻¹) in saltmarsh with considerable seasonal (max: 144·03 μg S m⁻² h⁻¹; min: 1·47 μg S m⁻² h⁻¹) and diurnal (max: 83·58 μg S m⁻² h⁻¹; min: 69·59 μg S m⁻² h⁻¹) changes in flux rates. Hydrogen sulfide was dominant (average: 21·2 μg S m⁻² h⁻¹, max: 79·2 μg S m⁻² h⁻¹; min: 5·29 μg S m⁻² h⁻¹) form in brackishmarsh and carbonyl sulfide (average: 1·09 μg S m⁻² h⁻¹; max: 3·42 μg S m⁻² h⁻¹; min: 0·32 μg S m⁻² h⁻¹) was dominant form in freshwater marsh. A greater amount of H₂S was evolved from brackishmarsh (21·22 μg S m⁻² h⁻¹) as compared to the saltmarsh (2·46 μg S m⁻² h⁻¹) and freshwater marsh (0·30 μg S m⁻² h⁻¹). Emission of total reduced sulfur gases decreased with decrease in salinity and distance inland from the coast. Emission of total reduced sulfur gases over the study averaged 73·3 μg S m⁻² h⁻¹ for the saltmarsh, 32·1 μg S m⁻² h⁻¹ for brackishmarsh and 2·76 μg S m⁻² h⁻¹ for the freshwater marsh.     

Mesoscale physical processes and zooplankton transport-retention in the northern Norwegian shelf region

Northern Norwegian shelf regions are highly productive, supporting fisheries rich in commercially important species such as cod, herring and capelin. It has been long recognized that the mesoscale jets, meanders and eddies associated with interactions between the North Atlantic Current, Norwegian Coastal Current and regional bottom topographic features such as troughs, banks and shelfbreaks play important roles in transporting and retaining zooplankton. To investigate zooplankton distributions and their correspondence with the physical fields, three large-scale surveys with mesoscale resolutions on physical and biological fields were conducted in northern Norwegian shelf regions between latitudes 68°15′N and 70°15′N in springs of 2000–2002. Survey results provide insights into the relationships between zooplankton distributions and the physical features such as fronts, the Norwegian Coastal Current and eddies related to topographic features. The physical and biological data are integrated and analyzed focusing on water types, estimation of geostrophic currents from direct current measurements, along-shelf transport of zooplankton, and retention of zooplankton by the mesoscale meander–eddy over a typical bank area on the shelf. The estimated mean transport in the upper 100 m on the shelf in the survey region is approximately 6.4×10³ tonnes wet weight day⁻¹ northward. High zooplankton abundances were found over both Malangsgrunnen and Sveinsgrunnen banks. The specific accumulation rate from northward–southward transport in the upper 100 m over Malangsgrunnen was approximately 0.08 day⁻¹, while variable currents with an offshore gradient of zooplankton abundance over Sveinsgrunnen implies an offshore dispersion of coastal-originated zooplankton cohort.     

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.     

Biological components of Ross Sea short-term particle fluxes in the austral summer of 1995–1996

The Ross Sea, a region of high seasonal production in the Southern Ocean, is characterized by blooms of the haptophyte Phaeocystis antarctica and of diatoms. The different morphology, structural composition and consumption of these two phytoplankton by grazing zooplankton may result in different carbon cycling dynamics and carbon flux from the euphotic zone. We sampled short-term (2 days) particle flux at 5 sites from 177.6°W to 165°E along a transect at 76.5°S with traps placed below the euphotic zone at 200 m during December 1995–January 1996. We estimated carbon flux of as many eucaryotic organisms and fecal pellets as possible using microscopy for counts and measurements and applying volume:carbon conversions from the literature. Eucaryotic organisms contributed about 20–40% of the total organic carbon flux in both the central Ross Sea polynya and in the western polynya, and groups of organisms differed in contribution to the carbon flux at the different sites. Algal carbon flux ranged from 4.5 to 21.1 mg C m⁻² day⁻¹ and consisted primarily of P. antarctica (cell plus mucus) and diatom carbon at all sites. Different diatom species dominated the diatom flux at different sites. Carbon fluxes of small pennate diatoms may have been enhanced by scavenging, by sinking senescent P. antarctica colonies. Heterotrophic carbon flux ranged from 9.2 to 37.6 mg C m⁻² day⁻¹ and was dominated by athecate heterotrophic dinoflagellate carbon in general and by carbon flux of a particular large athecate dinoflagellate at two sites. Fecal pellet carbon flux ranged from 4.6 to 54.5 mg C m⁻² day⁻¹ and was dominated by carbon from ovoid/angular pellets at most sites. Analysis of fecal pellet contents suggested that large protozoans identified by light microscopy contributed to ovoid/angular fecal pellet fluxes. Carbon flux as a percentage of daily primary production was lowest at sites where P. antarctica predominated in the water column and was highest at sites where fecal pellet flux was highest. This indicates the importance of grazers in carbon export.     

Downward carbon transport by diel vertical migration of the copepods Metridia pacifica and Metridia okhotensis in the Oyashio region of the western subarctic Pacific Ocean

Seasonal change in the downward carbon transport due to respiration and mortality through diel vertical migration (DVM) of the calanoid copepods Metridia pacifica and Metridia okhotensis was estimated in the Oyashio region, western subarctic Pacific during six cruises from June 2001 to June 2002. M. pacifica (C4, C5 and adult females) was an active migratory species throughout the year though its DVM amplitude varied among seasons and stages. The mean distribution depths of adult females during the daytime were positively related with the illumination level in the water column, being shallowest in April and deepest in January. M. okhotensis generally showed less-extensive migrations than M. pacifica. Therefore, together with their lower abundance, this species is considered to be a less-important mechanism of downward transport of carbon except for April when their DVM was more active and descended deeper than M. pacifica, which remained in the upper 150 m even during the daytime. The mean migrating biomass of the two Metridia species was 558 mg C m⁻² d⁻¹ and was high during summer to winter (263–1676 mg C m⁻² d⁻¹) and low during spring (59–63 mg C m⁻² d⁻¹). Total downward flux through DVM fluctuated between 1.0 and 20.0 mg C m⁻² d⁻¹ with an annual mean of 8.0 mg C m⁻² d⁻¹. Contribution of the respiratory flux was greater than the mortality flux and accounted for 64–98% of total migratory flux throughout the year except for January when contribution of both fluxes was equal. Overall the annual carbon transport by DVM of Metridia spp. was estimated as 3.0 g C m⁻² year⁻¹, corresponding to 15% of the annual total POC flux at 150 m at the study site, suggesting that DVM is a significant process for carbon export in the subarctic region as well as that in tropical and subtropical oceanic regions. Since DVM in M. pacifica is more active during the non-bloom season when the gravitational flux of particulate matter is low, this species plays an important role in driving the biological pump in the subarctic Pacific during summer to winter.     

A regional analysis of new production on the northwest European shelf using oxygen fluxes and a ship-of-opportunity

Seasonal new production (g C m⁻²) estimates obtained from dissolved oxygen and nitrate concentrations in surface waters (5 m depth) along a track between the UK (Portsmouth) and northern Spain (Bilbao) are compared. An oxygen flux method, in combination with a ship of opportunity (SOO), was tested on the northwest European shelf for its value in distinguishing high production in frontal regions. Dissolved oxygen, nitrate and chlorophyll a samples were collected monthly from February to July 2004, alongside continuous autonomous measurements of salinity, temperature and chlorophyll fluorescence. Depth integrated new production estimates for all the individually analysed hydrographic regions of the route were produced.Results from three widely used gas-exchange parameterizations gave seasonal (February–July) new production estimates of 54–68 g C m⁻² for the Ushant region of the western English Channel and 31–40 g C m⁻² for the shelf slope, averaging 24–31 g C m⁻² for the route. This is double the route average obtained using the nitrate assimilation method (17 g C m⁻²) and within the ranges of previous estimates in the same region. The oxygen flux method gave a fivefold enhancement compared to the nitrate method in the Ushant frontal region and a threefold enhancement in the English Channel and shelf break regions. Determining oxygen fluxes to estimate new production may be more reliable than nitrate assimilation in active tidal or frontal regions of shelves where nitrate may be added to the system post-winter through advection or entrainment.     

Factors influencing benthic bacterial abundance, biomass, and activity on the northern continental margin and deep basin of the Gulf of Mexico

As part of a larger project on the deep benthos of the Gulf of Mexico, an extensive data set on benthic bacterial abundance (n>750), supplemented with cell-size and rate measurements, was acquired from 51 sites across a depth range of 212–3732 m on the northern continental slope and deep basin during the years 2000, 2001, and 2002. Bacterial abundance, determined by epifluorescence microscopy, was examined region-wide as a function of spatial and temporal variables, while subsets of the data were examined for sediment-based chemical or mineralogical correlates according to the availability of collaborative data sets. In the latter case, depth of oxygen penetration helped to explain bacterial depth profiles into the sediment, but only porewater DOC correlated significantly (inversely) with bacterial abundance (p<0.05, n=24). Other (positive) correlations were detected with TOC, C/N ratios, and % sand when the analysis was restricted to data from the easternmost stations (p<0.05, n=9–12). Region-wide, neither surface bacterial abundance (3.30–16.8×10⁸ bacteria cm⁻³ in 0–1 cm and 4–5 cm strata) nor depth-integrated abundance (4.84–17.5×10¹³ bacteria m⁻², 0–15 cm) could be explained by water depth, station location, sampling year, or vertical POC flux. In contrast, depth-integrated bacterial biomass, derived from measured cell sizes of 0.027–0.072 μm³, declined significantly with station depth (p<0.001, n=56). Steeper declines in biomass were observed for the cross-slope transects (when unusual topographic sites and abyssal stations were excluded). The importance of resource changes with depth was supported by the positive relationship observed between bacterial biomass and vertical POC flux, derived from measures of overlying productivity, a relationship that remained significant when depth was held constant (partial correlation analysis, p<0.05, df=50). Whole-sediment incubation experiments under simulated in situ conditions, using ³H-thymidine or ¹⁴C-amino acids, yielded low production rates (5–75 μg C m⁻² d⁻¹) and higher respiration rates (76–242 μg C m⁻² d⁻¹), with kinetics suggestive of resource limitation at abyssal depths. Compared to similarly examined deep regions of the open ocean, the semi-enclosed Gulf of Mexico (like the Arabian Sea) harbors in its abyssal sediments a greater biomass of bacteria per unit of vertically delivered POC, likely reflecting the greater input of laterally advected, often unreactive, material from its margins.     

Impact of a cyclonic eddy on phytoplankton community structure and photosynthetic competency in the subtropical North Pacific Ocean

A synoptic spatial examination of the eddy Haulani (17–20 November 2000) revealed a structure typical of Hawaiian cyclonic eddies with divergent surface flow forcing the upward displacement of deep waters. Hydrographic surveys revealed that surface water in the eddy center was ca. 3.5°C cooler, 0.5 saltier, and 1.4 kg m⁻³ denser than surface waters outside the eddy. Vertically integrated concentrations of nitrate+nitrite, phosphate and silicate were enhanced over out-eddy values by about 2-fold, and nitrate+nitrite concentrations were ca. 8× greater within the euphotic zone inside the eddy than outside. Si:N ratios were lower within the upper mixed layer of the eddy, indicating an enhanced Si uptake relative to nitrate+nitrite. Chlorophyll a concentrations were higher within the eddy compared to control stations outside, when integrated over the upper 150 m, but were not significantly different when integrated over the depth of the euphotic zone. Photosynthetic competency, assessed using fast repetition-rate fluorometry, varied with the doming of the isopycnals and the supply of macro-nutrients to the euphotic zone. The physical and chemical environment of the eddy selected for the accumulation of larger phytoplankton species. Photosynthetic bacteria (Prochlorococcus and Synechococcus) and small (<3 μm diameter) photosynthetic eukaryotes were 3.6-fold more numerically abundant outside the eddy as compared to inside. Large photosynthetic eukaryotes (>3 μm diameter) were more abundant inside the eddy than outside. Diatoms of the genera Rhizosolenia and Hemiaulus outside the eddy contained diazotrophic endosymbiontic cyanobacteria, but these endosymbionts were absent from the cells of these species inside the eddy. The increase in cell numbers of large photosynthetic eukaryotes with hard silica or calcite cell walls is likely to have a profound impact on the proportion of the organic carbon production that is exported to deep water by sinking of senescent cells and cells grazed by herbivorous zooplankton and repackaged as large fecal pellets.     

Microbial dynamics in cyclonic and anticyclonic mode-water eddies in the northwestern Sargasso Sea

The EDdy Dynamics, mixing, Export, and Species composition (EDDIES) project provided a unique opportunity to evaluate the response of the microbial community and further understand the biological and biogeochemical consequences of mesoscale perturbation events in an oligotrophic system. In order to characterize microbial dynamics, we performed measurements of bacterial biomass (BB) and production (BP) and phytoplankton pigment analyses in two upwelling eddies in the Sargasso Sea sampled in 2004 and 2005. We also observed a 3-fold increase in BP at the Bermuda Atlantic Time-series Study (BATS) site during the passage of a cyclonic eddy in 2003. Although the integrated BB and BP over 140 m in 2004 and 2005 eddies remained within the climatological range measured at the BATS site, there was systematic variability in bacterioplankton dynamics across both eddies. Cyclonic eddy C1 demonstrated decreased BP at the feature's center relative to its periphery, and BP was not correlated with total chlorophyll a (TChl a) variability. However, BP correlated with prymnesiophyte pigments throughout the feature. In contrast, mode-water eddy A4 showed an enhancement in BP at the eddy center (EC) relative to its edges and was coincident with elevated TChl a, high primary production measurements, and a high concentration of diatoms. In eddy A4, the tight relationship between enhanced BP, TChl a and specific phytoplankton taxa implies that the phytoplankton community structure was an important factor influencing BP variability. While the heterotrophic bacterial response in C1 and A4 was not enhanced relative to BATS summer climatology, these data and the presence of similar nutrient fields across both eddies suggest that BP and BB were influenced by the eddy perturbations and responded to changes in the phytoplankton community.     

Response of Nereis diversicolor (Polychaeta, Nereidae) populations to reduced wastewater discharge in the polluted estuary of Oued Souss, Bay of Agadir, Morocco

Field investigations on the population dynamics of Nereis diversicolor were carried out from January 2002 to December 2003 in the estuary of Oued Souss (southwestern Morocco) to determine the changes caused by setting up of a domestic and industrial wastewater purification plant (M'zar) before and after by the end of wastewater discharges in November 2002 on the structure of the ecosystem. Samples of N. diversicolor were collected monthly in the intertidal zone at low tide before (during 2002) and after (during 2003) the end of wastewater discharges.Separation of cohorts using the Algorithm EM method (McLachlan, G.J., Krishnan, T., 1997. The EM algorithm and extensions. Wiley Series in Probability and Statistics. Wiley, New York, 274 pp.) allowed determination of the growth rate (mm day⁻¹) by cohort and the annual production. The data showed significant differences between populations of Nereis diversicolor before and after the end of wastewater discharges. During the wastewater discharge period (2002), the population had a mean annual density of 1992 ind m⁻², a mean annual biomass of 75.52 g DW m⁻² and an annual secondary production of 141.3 g DW m⁻² with a P/B ratio of 1.87. After the end of discharges (2003), density, biomass and secondary production decreased significantly. The annual averages for these parameters were 740 ind m⁻², 14.16 g DW m⁻² and 23.83 g DW m⁻², respectively, with a P/B ratio of 1.68.The important decrease observed in density, biomass and secondary production of Nereis diversicolor may be attributed (a) to the environmental changes observed after the end of wastewater discharges in the estuary of Oued Souss, namely the increase of salinity and the decrease of organic matter content, and (b) to the migration of this species towards other areas.