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.     

Downward export of carbon by diel migrant mesozooplankton in the central equatorial Pacific

Using data collected during cruises of the JGOFS equatorial Pacific Study in March/April and October of 1992 at the equator (140°W), we examine the downward transport of carbon by three size classes of die] migrant mesozooplankton (200–500 gm, 500–1000 μm and 1000–2000 gm). In addition to respiratory carbon flux, we consider the flux due to mortality of migrators below the euphotic zone. Diel migrant mesozooplankton biomass was estimated from the difference between nighttime and daytime biomass within the euphotic zone. Except for a four-day period early in the March/April cruise, mesozooplankton nighttime biomass was significantly larger than daytime biomass within the euphotic zone during both cruises. We estimate that the downward flux of carbon from the euphotic zone due to mesozooplankton die] vertical migrators was an average of 0.6 mmol Cm⁻² d⁻¹ and 1.1 mmol C m⁻² d⁻¹ during the March/April and October cruises, respectively. Addition of this flux to the gravitational particle sinking flux estimated from²³⁴Th measurements during the same period results in a 31 % increase in the carbon export flux from the euphotic zone in the equatorial Pacific during the March/April cruise and a 44% increase in the October cruise. The migratory flux is strongly dependent on whether feeding takes place below the euphoric zone, the length of time migrators spend in the deep waters, and the mortality rate of migrators.     

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.     

A time-series study of particulate matter export in the North Pacific Subtropical Gyre based on Th : U disequilibrium

Depth profiles of total ²³⁴Th (dissolved+particulate) were collected at Station ALOHA (22°45N, 158°00W) in the North Pacific Subtropical Gyre during 9 cruises from April 1999 to March 2000. Samples were collected and processed by a new 2 L technique that enables more detailed depth resolution then previous ²³⁴Th studies. Significant zones of particle export (²³⁴Th deficiency) and particle remineralization (²³⁴Th excess) were measured both temporally and with depth. ²³⁴Th derived particulate carbon (PC) and nitrogen (PN) fluxes were determined with steady-state and non-steady-state models and PC/²³⁴Th and PN/²³⁴Th ratios measured with both in situ pumps and free-drifting particle interceptor traps deployed at 150 m. ²³⁴Th based export estimates of 4.0±2.3 mmol C m⁻² d⁻¹ and 0.53±0.19 mmol N m⁻² d⁻¹, were approximately 60% higher than those measured in PIT style sediment traps from the same time period, 2.4±0.2 mmol C m⁻² d⁻¹ and 0.32±0.08 mmol N m⁻² d⁻¹. Most of this difference is attributable to two large export events that occurred during October and December 1999, when traps undercollected for ²³⁴Th by a factor of 2 to 4. ²³⁴Th export (ThE) ratios based on ²³⁴Th derived PC flux/¹⁴C based primary production ranged from 4% to 22% (average=8.8%). Our results confirm the recent estimates of C export by Emerson et al. (Nature 389 (1997) 951) and Sonnerup et al. (Deep-Sea Research I 46 (1999) 777) and indicate that C export from the oligotrophic ocean must be considered when discussing C sequestration in global climate change.     

Vertical flux of biogenic matter during a Lagrangian study off the NW Spanish continental margin

Lagrangian experiments with short-term, drifting sediment traps were conducted during a cruise on RRS Charles Darwin to the NW coast of Spain to study the vertical flux and composition of settling biogenic matter. The cruise was split into two legs corresponding to (i) a period of increased production following an upwelling event on the continental shelf (3–10 August 1998) and (ii) an evolution of a cold water filament originating from the upwelled water off the shelf (14–19 August). The export of particulate organic carbon (POC) from the upper layer (0–60m) on the shelf was 90–240mgC.m⁻².d⁻¹ and off the shelf was 60–180mgC.m⁻².d⁻¹. Off shelf the POC flux at 200m was 50–60mg.m⁻².d⁻¹. A modest sedimentation of diatoms (15–30mgC.m⁻².d⁻¹) after the upwelling was associated with increased vertical flux of chlorophyll a (1.8–2.1mg.m⁻².d⁻¹) and a decrease of the POC:PON molar ratio of the settled material from 9 to 6.4. Most of the pico-, nano-, and microplankton in the settled material were flagellates; diatoms were significant during the on shelf and dinoflagellates during the off shelf leg. Off shelf, the exponential attenuation of POC flux indicated a strong retention capacity of the plankton community between 40 and 75m. POC:PON ratio of the settled particulate matter decreased with depth and the relative portion of flagellates increased, suggesting a novel, flagellate and aggregate mediated particulate flux in these waters. Export of POC from the euphotic layer comprised 14–26% of the integrated primary production per day during the on shelf leg and 25–42% during the off shelf leg, which characterises the importance of sedimentation in the organic carbon budget of these waters.     

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.     

Microzooplankton and mesozooplankton in an upwelling filament off Galicia: modelling and sensitivity analysis of the linkages and their impact on the carbon dynamics

This paper reports estimates of trophic flows of carbon off the Galician coast from a 1D ecological model, which are compared with field data from a two week Lagrangian drift experiment. The model consists of 9 biological components: nitrate, ammonium, >5μm phytoplankton, <5μm phytoplankton, heterotrophic nanoflagellates/dinoflagellates (5–20 μm), heterotrophic dinoflagellates (>20 μm), ciliates, fast sinking detritus and slow sinking detritus. Calculations were made for the fluxes of carbon between biological components within the upper 45m of the water column. The temporal development of primary production during the simulation period of two weeks was in good agreement with field estimates, which varied between 248 and 436mgC.m⁻².d⁻¹. Heterotrophic nanoflagellates had the greatest impact on carbon flux, with a grazing rate of 168mgC.m⁻².d⁻¹. Herbivorous grazing by microzooplankton amounted to 215mgC.m⁻².d⁻¹, whereas grazing by copepods on phytoplankton was 35mgC.m⁻² d⁻¹. Copepods grazing on microzooplankton was minor (0.47mgC.m⁻².d⁻¹) and the export flux from the upper 45m was 302mgC.m⁻².d⁻¹. Sensitivity analyses, in which the grazing parameters (i.e the functional relationship between ingestion and food concentration) were changed, were carried out on the heterotrophic dinoflagellate, ciliate and heterotrophic nanoflagellates/dinoflagellate components of the model. These changes did not alter the temporal development of heterotrophic nanoflagellates/dinoflagellates biomass significantly, but ciliates and heterotrophic dinoflagellates were more sensitive to variations in the grazing parameters. The overall conclusion from this modelling study is that the coupling between small phytoplankton and heterotrophic nanoflagellates was the quantitatively most important process controlling carbon flow in this region.     

Seasonal changes and biochemical composition of the labile organic matter flux in the Cretan Sea

Downward fluxes of labile organic matter (lipids, proteins and carbohydrates) at 200 (trap A) and 1515 m depth (trap B), measured during a 12 months sediment trap experiment, are presented, together with estimates of the bacterial and cyanobacterial biomasses associated to the particles. The biochemical composition of the settling particles was determined in order to provide qualitative and quantitative information on the flux of readily available organic carbon supplying the deep-sea benthic communities of the Cretan Sea. Total mass flux and labile carbon fluxes were characterised by a clear seasonality. Higher labile organic fluxes were reported in trap B, indicating the presence of resuspended particles coming from lateral inputs. Particulate carbohydrates were the major component of the flux of labile compounds (on annual average about 66% of the total labile organic flux) followed by lipids (20%) and proteins (13%). The biopolymeric carbon flux was very low (on annual average 0.9 and 1.2 gC m⁻² y⁻¹, at trap A and B). Labile carbon accounted for most of the OC flux (on annual average 84% and 74% in trap A and B respectively). In trap A, highest carbohydrate and protein fluxes in April and September, corresponded to high faecal pellet fluxes. The qualitative composition of the organic fluxes indicated a strong protein depletion in trap B and a decrease of the bioavailability of the settling particles as a result of a higher degree of dilution with inorganic material. Quantity and quality of the food supply to the benthos displayed different temporal patterns. Bacterial biomass in the sediment traps (on average 122 and 229 μgC m⁻² d⁻¹ in trap A and B, respectively) was significantly correlated to the flux of labile organic carbon, and particularly to the protein and carbohydrate fluxes. Cyanobacterial flux (on average, 1.1 and 0.4 μgC m⁻² d⁻¹, in trap A and B, respectively) was significantly correlated with total mass and protein fluxes only in trap A. Bacterial carbon flux, equivalent to 84.2 and 156 mgC m⁻² y⁻¹, accounted for 5–6.5% of the labile carbon flux (in trap A and B respectively) and for 22–41% protein pool of the settling particles. These results suggest that in the Cretan Sea, bacteria attached to the settling particles represent a potential food source of primary importance for deep-sea benthic communities.     

Contrasts in sedimentation flux below the southern California Current in late 1996 and during the El Niño event of 1997–1998

The vertical flux of particulate matter at 330 m depth in San Lázaro Basin off southern Baja California ranged from 63 to 587 mg m⁻² d⁻¹ between August and November 1996. Organic carbon contents were between 5.6 and 14.8%, yielding flux rates of 9–40 mgC m⁻² d⁻¹. In December 1997 and January 1998, at the height of the strong El Niño event, the respective fluxes (47–202 mg m⁻² d⁻¹ and 3–8 mgC m⁻² d⁻¹) were comparable. The February–June 1998 records, however, revealed sharply reduced mass (1–6 mg m⁻² d⁻¹) and organic carbon (0.2–0.8 mgC m⁻² d⁻¹) fluxes. The organics collected in 1996 were predominantly autochthonous (δ¹³C=−22‰; C/N=8). The variations in δ¹⁵N (8.3–11.0‰) suggest an alternation of new and regenerated production, possibly associated with fluctuations in the intensity of deep mixing during that autumn. The relatively high organic matter fluxes in December 1997 appear to be associated with regenerated production. The average composition from February to June 1998 (δ¹³C=−23.6‰; ¹⁵N=11.7‰; C/N=10.5) indicates degraded material of marine origin. The maximum δ¹⁵N value found (14‰) suggests that deeper, denitrified waters were brought to the surface and possibly advected laterally. Regime changes in the waters of the basin occur at 6–10 week intervals, evidenced by concurrent shifts in most of the measured parameters, including fecal pellet types and metal chemistry. The marine snow-dominated detritus collected showed a shift from a mixed diatom-rich-radiolarian-coccolith assemblage in late 1996 to a coccolith-dominated assemblage, including the contents of fecal pellets, during the 1997–1998 El-Niño period. T–S profiles, plankton analysis and chlorophyll contents of the upper water column indicated that the strong phytoplankton bloom, normally associated with seasonal upwelling along the Pacific coast of Baja, did not occur during the spring of 1998. The persistence of oligotrophic conditions during the 1997–1998 El Niño event favored the dominance of nanoplankton and reduced the vertical flux of particles.     

Nitrogen cycling in deep-sea sediments of the Porcupine Abyssal Plain, NE Atlantic

Rates of transformation, recycling and burial of nitrogen and their temporal and spatial variability were investigated in deep-sea sediments of the Porcupine Abyssal Plain (PAP), NE Atlantic during eight cruises from 1996 to 2000. Benthic fluxes of ammonium (NH₄) and nitrate (NO₃) were measured in situ using a benthic lander. Fluxes of dissolved organic nitrogen (DON) and denitrification rates were calculated from pore water profiles of DON and NO₃, respectively. Burial of nitrogen was calculated from down core profiles of nitrogen in the solid phase together with ¹⁴C-based sediment accumulation rates and dry bulk density. Average NH₄ and NO₃-effluxes were 7.4 ± 19 μmol m⁻² d⁻¹ (n = 7) and 52 ± 30 μmol m⁻² d⁻¹ (n = 14), respectively, during the period 1996–2000. During the same period, the DON-flux was 11 ± 5.6 μmol m⁻² d⁻¹ (n = 5) and the denitrification rate was 5.1 ± 3.0 μmol m⁻² d⁻¹ (n = 22). Temporal and spatial variations were only found in the benthic NO₃ fluxes. The average burial rate was 4.6 ± 0.9 μmol m⁻² d⁻¹. On average over the sampling period, the recycling efficiency of the PON input to the sediment was 94% and the burial efficiency hence 6%. The DON flux constituted 14% of the nitrogen recycled, and it was of similar magnitude as the sum of burial and denitrification. By assuming the PAP is representative of all deep-sea areas, rates of denitrification, burial and DON efflux were extrapolated to the total area of the deep-sea floor (>2000 m) and integrated values of denitrification and burial of 8 ± 5 and 7 ± 1 Tg N year⁻¹, respectively, were obtained. This value of total deep-sea sediment denitrification corresponds to 3–12% of the global ocean benthic denitrification. Burial in deep-sea sediments makes up at least 25% of the global ocean nitrogen burial. The integrated DON flux from the deep-sea floor is comparable in magnitude to a reported global riverine input of DON suggesting that deep-sea sediments constitute an important source of DON to the world ocean.     

Elemental mercury concentrations and formation rates in the Scheldt estuary and the North Sea

Concentrations of Hg⁰ in surface waters and atmosphere of the Scheldt estuary and the North Sea are presented and their relationship with biological processes is discussed. Hg⁰ concentrations in the Scheldt estuary range from 0.1 to 0.38 pmol·l⁻¹ in the winter and from 0.24 to 0.65 pmol·l⁻¹ in the summer and show a positive relationship with phytoplankton pigments. In the North Sea Hg⁰ concentrations range from 0.06 to 0.8 pmol·l⁻¹ and are higher in coastal stations. Transfer velocities across the air–sea interface were calculated using a classical shear turbulence model. Volatilization fluxes of Hg⁰ were calculated for the Scheldt estuary and the North Sea. For the Scheldt estuary the fluxes range from 226–284 pmol·m⁻²·d⁻¹ in winter and 500–701 pmol·m⁻²·d⁻¹ in summer and for the North Sea the fluxes range from 59–1110 pmol·m⁻²·d⁻¹ for an average windspeed of 8.1 m·s⁻¹. These fluxes are comparable to the wet and dry depositional fluxes to the North Sea. Hg⁰ formation rates necessary to balance the volatilization fluxes vary from 0.2 to 4% d⁻¹.     

Effects of irradiance on benthic and water column processes in a Gulf of Mexico estuary: Pensacola Bay, Florida, USA

We examined the effect of light on water column and benthic fluxes in the Pensacola Bay estuary, a river-dominated system in the northeastern Gulf of Mexico. Measurements were made during the summers of 2003 and 2004 on 16 dates distributed along depth and salinity gradients. Dissolved oxygen fluxes were measured on replicate sediment and water column samples exposed to a gradient of photosynthetically active radiation. Sediment inorganic nutrient (NH₄⁺, NO₃⁻, PO₄³⁻) fluxes were measured. The response of dissolved oxygen fluxes to variation in light was fit to a photosynthesis–irradiance model and the parameter estimates were used to calculate daily integrated production in the water column and the benthos. The results suggest that shoal environments supported substantial benthic productivity, averaging 13.6 ± 4.7 mmol O₂ m⁻² d⁻¹, whereas channel environments supported low benthic productivity, averaging 0.5 ± 0.3 mmol O₂ m⁻² d⁻¹ (±SE). Estimates of baywide microphytobenthic productivity ranged from 8.1 to 16.5 mmol O₂ m⁻² d⁻¹, comprising about 16–32% of total system productivity. Benthic and water column dark respiration averaged 15.2 ± 3.2 and 33.6 ± 3.7 mmol O₂ m⁻² d⁻¹, respectively Inorganic nutrient fluxes were generally low compared to relevant estuarine literature values, and responded minimally to light exposure. Across all stations, nutrient fluxes from sediments to the water column averaged 1.11 ± 0.98 mmol m⁻² d⁻¹ for NH₄⁺, 0.58 ± 1.08 mmol m⁻² d⁻¹ for NO₃⁻, 0.01 ± 0.09 mmol m⁻² d⁻¹ for PO₄³⁻. The results of this study illustrate how light reaching the sediments is an important modulator of benthic nutrient and oxygen dynamics in shallow estuarine systems.     

Rapid sediment accumulation and microbial mineralization in forests of the mangrove Kandelia candel in the Jiulongjiang Estuary, China

Rates of sediment accumulation and microbial mineralization were examined at three Kandelia candel forests spanning the intertidal zone along the south coastline of the heavily urbanized Jiulongljiang Estuary, Fujian Province, China. Mass sediment accumulation rates were rapid (range: 10–62 kg m⁻² y⁻¹) but decreased from the low- to the high-intertidal zone. High levels of radionuclides suggest that these sediments originate from erosion of agricultural soils within the catchment. Mineralization of sediment carbon and nitrogen was correspondingly rapid, with total rate of mineralization ranging from 135 to 191 mol C m⁻² y⁻¹ and 9 to 11 mol N m⁻² y⁻¹; rates were faster in summer than in autumn/winter. Rates of mineralization efficiency (70–93% for C; 69–92% for N) increased, as burial efficiency (7–30% for C; 8–31% for N) decreased, from the low-to the high-intertidal mangroves. Sulphate reduction was the dominant metabolic pathway to a depth of 1 m, with rates (19–281 mmol S m⁻² d⁻¹) exceeding those measured in other intertidal deposits. There is some evidence that Fe and Mn reduction-oxidation cycles are coupled to the activities of live roots within the 0–40 cm depth horizon. Oxic respiration accounted for 5–12% of total carbon mineralization. Methane flux was slow and highly variable when detectable (range: 5–66 μmol CH₄ m⁻² d⁻¹). Nitrous oxide flux was also highly variable, but within the range (1.6–106.5 μmol N₂O m⁻² d⁻¹) measured in other intertidal sediments. Rates of denitrification were rapid, ranging from 1106 to 3780 μmol N₂ m⁻² d⁻¹, and equating to 11–20% of total sediment nitrogen inputs. Denitrification was supported by rapid NH₄ release within surface deposits (range: 3.6–6.1 mmol m⁻² d⁻¹). Our results support the notion that mangrove forests are net accumulation sites for sediment and associated elements within estuaries, especially Kandelia candel forests receiving significant inputs as a direct result of intense human activity along the south China coast.     

Nitrogen dynamics within a wind-driven eddy

Wind-driven cyclonic eddies are hypothesized to relieve nutrient stress and enhance primary production by the upward displacement of nutrient-rich deep waters into the euphotic zone. In this study, we measured nitrate (NO₃⁻), particulate carbon (PC), particulate nitrogen (PN), their stable isotope compositions (δ¹⁵N-NO₃⁻, δ¹³C-PC and δ¹⁵N-PN, respectively), and dissolved organic nitrogen (DON) within Cyclone Opal, a mature wind-driven eddy generated in the lee of the Hawaiian Islands. Sampling occurred in March 2005 as part of the multi-disciplinary E-Flux study, approximately 4–6 weeks after eddy formation. Integrated NO₃⁻ concentrations above 110 m were 4.8 times greater inside the eddy (85.8±6.4 mmol N m⁻²) compared to the surrounding water column (17.8±7.8 mmol N m⁻²). Using N-isotope derived estimates of NO₃⁻ assimilation, we estimated that 213±59 mmol m⁻² of NO₃⁻ was initially injected into the upper 110 m Cyclone Opal formation, implying that NO₃⁻ was assimilated at a rate of 3.75±0.5 mmol N m⁻² d⁻¹. This injected NO₃⁻ supported 68±19% and 66±9% of the phytoplankton N demand and export production, respectively. N isotope data suggest that 32±6% of the initial NO₃⁻ remained unassimilated. Self-shading, inefficiency in the transfer of N from dissolved to particulate export, or depletion of a specific nutrient other than N may have led to a lack of complete NO₃⁻ assimilation. Using a salt budget approach, we estimate that dissolved organic nitrogen (DON) concentrations increased from eddy formation (3.8±0.4 mmol N m⁻²) to the time of sampling (4.0±0.09 mmol N m⁻²), implying that DON accumulated at rate of 0.83±1.3 mmol N m⁻² d⁻¹, and accounted for 22±15% of the injected NO₃⁻. Interestingly, no significant increase in suspended PN and PC, or export production was observed inside Cyclone Opal relative to the surrounding water column. A simple N budget shows that if 22±15% of the injected NO₃⁻ was shunted into the DON pool, and 32±6% is unassimilated, then 46±16% of the injected NO₃⁻ remains undocumented. Alternative loss processes within the eddy include lateral exchange of injected NO₃⁻ along isopycnal surfaces, remineralization of PN at depth, as well as microzooplankton grazing. A 9-day time series within Cyclone Opal revealed a temporal depletion in δ¹⁵N-PN, implying a rapid change in the N source. A change in NO₃⁻ assimilation, or a shift from NO₃⁻ fueled growth to assimilation of a ¹⁵N-deplete N source, may be responsible for such observations.     

Effect of changes in water salinity on ammonium, calcium, dissolved inorganic carbon and influence on water/sediment dynamics

The effect of a sudden increase in salinity from 10 to 37 in porewater concentration and the benthic fluxes of ammonium, calcium and dissolved inorganic carbon were studied in sediments of a small coastal lagoon, the Albufera d'Es Grau (Minorca Island, Spain). The temporal effects of the changes in salinity were examined over 17 days using a single diffusion-reaction model and a mass-balance approach. After the salinity change, NH₄⁺-flux to the water and Ca-flux toward sediments increased (NH₄⁺-flux: 5000–3000 μmol m⁻² d⁻¹ in seawater and 600/250 μmol m⁻² d⁻¹ in brackish water; Ca-flux: −40/−76 meq m⁻² d⁻¹ at S=37 and −13/−10 meq m⁻² d⁻¹ at S=10); however, later NH₄⁺-flux decreased in seawater, reaching values lower than in brackish water. In contrast, Ca-flux presented similar values in both conditions. The fluxes of dissolved inorganic carbon, which were constant at S=10 (55/45 mmol m⁻² d⁻¹), increased during the experiment at S=37 (from 30 mmol m⁻² d⁻¹ immediately after salinity increase to 60 mmol m⁻² d⁻¹ after 17 days).In brackish conditions, NH₄⁺ and Ca²⁺ fluxes were consistent with a single diffusion-reaction model that assumes a zero-order reaction for NH₄⁺ production and a first-order reaction for Ca²⁺ production. In seawater, this model explained the Ca-flux observed, but did not account for the high initial flux of NH₄⁺.The mass balance for 17 days indicated a higher retention of NH₄⁺ in porewater in the littoral station in seawater conditions (9.5 mmol m⁻² at S=37 and 1.6 mmol m⁻² at S=10) and a significant reduction in the water consumption at both sites (5 mmol m⁻² at S=37; 35/23 mmol m⁻² at S=10). In contrast, accumulation of dissolved inorganic carbon in porewater was lower in seawater incubations (−10/−1 meq m⁻² at S=37; 50/90 meq m⁻² at S=10) and was linked to a higher efflux of CO₂ to the atmosphere, because of calcium carbonate precipitation in water (675/500 meq m⁻²). These results indicate that increased salinity in shallow coastal waters could play a major role in the global carbon cycle.     

Imbalance in the carbonate budget of surficial sediments in the North Atlantic Ocean: variations over the last millenium?

Fluxes contributing to the particulate carbonate system in deep-sea sediments were investigated at the BENGAL site in the Porcupine Abyssal Plain (Northeast Atlantic). Deposition fluxes were estimated using sediment traps at a nominal depth of 3000 m and amounted to 0.37±0.1 mmol C m⁻² d⁻¹. Dissolution of carbonate was determined using flux of total alkalinity from in situ benthic chambers, is 0.4±0.1 mmol C m⁻² d⁻¹. Burial of carbonate was calculated from data on the carbonate content of the sediment and sedimentation rates from a model age based on ¹⁴C dating on foraminifera (0.66±0.1 mmol C m⁻² d⁻¹). Burial plus dissolution was three times larger than particle deposition flux which indicates that steady-state is not achieved in these sediments. Mass balances for other components (BSi, ²¹⁰Pb), and calculations of the focusing factor using ²³⁰Th, show that lateral inputs play only a minor role in this imbalance. Decadal variations of annual particle fluxes are also within the uncertainty of our average. Long-term change in dissolution may contribute to the imbalance, but can not be the main reason because burial alone is greater than the input flux. The observed imbalance is thus the consequence of a large change of carbonate input flux which has occured in the recent past. A box model is used to check the response time of the solid carbonate system in these sediments and the time to reach a new steady-state is in the order of 3 kyr. Thus it is likely that the system has been perturbed recently and that large dissolution and burial rates reflect the previously larger particulate carbonate deposition rates. We estimate that particulate carbonate fluxes have certainly decreased by a factor of at least 3 and that this change has occurred during the last few centuries.     

New production in the subarctic Pacific Ocean: Net changes in nitrate concentrations, rates of nitrate assimilation and accumulation of particulate nitrogen

Changes in water column nitrate and particulate nitrogen (PN) concentrations and rates of nitrate assimilation at 50°N 145°W were measured over a four-month interval for 1984, 1987 and 1988. Rates of nitrate depletion in the upper 80m of the water column averaged 12mg N m⁻²d⁻¹, but most of the net depletion occurred during May when rates were high (75mg N m⁻²d⁻¹) compared to later in the year. Particulate nitrogen (collected on GF/F filters) increased 2- to 3-fold during the month of May and accounted for 30–60% of the net nitrate depletion for May. Mean rates of PN accumulation for the 4-month intervals were 2.4mg N m⁻²d⁻¹ and accounted for about 20% of the net nitrate depletion. Rates of nitrate assimilation (measured in incubation bottles with ¹⁵N) averaged 45.0±4.5mg N m⁻²d⁻¹ (mean±SD), and appeared to decrease between May and September. A good correspondence between in situ and incubation estimates of nitrate assimilation was found for the 4-month comparison, but not for the month of May when net changes in nitrate concentrations were greatest. Vertical and horizontal inputs of nitrate are about the same order of magnitude as biological removal, thus the high inout of nitrate into the euphotic zone contributed to the continuously high nitrate concentrations in this region. Seasonal changes in nitrate and PN were significant and need to be considered in comparisons of new and export production.     

Functional roles of interzonal migrating mesozooplankton in the western subarctic Pacific

Grazing experiments and production estimation based on life-history analysis of Neocalanus copepods (N. cristatus, N. plumchrus and N. flemingeri) were carried out in the Oyashio region to understand the carbon flows associated with the interzonal migrating copepods. These copepods, and also Eucalanus bungii, fed on nano- and micro-sized organisms non-selectively throughout the season. However, diatoms were the dominant food resource until May and organisms, such as ciliates were the major resource after May. Daily growth rate was estimated from the Ikeda–Motoda, Huntley–Lopez and Hirst–Sheader models. Since the growth rates were considered to be overestimates for the Huntley–Lopez model and underestimates for the other two models, we applied the weight-specific growth rates previously reported for these species in the Bering Shelf. Surface biomass of Neocalanus increased rapidly in June during the appearance of C5, and a successive increase of overwintering stock was evident in the deeper layer. The deep biomass decreased gradually from September to May during the dormant and reproduction period. N. cristatus has the largest annual mean biomass (2.3 gC m⁻²), followed by N. plumchrus (1.1) and N. flemingeri (0.4). Daily production rate of Neocalanus varied from 0.4 to 363.4 mgC m⁻² day⁻¹, to which N. cristatus was the largest contributor. Annual production was estimated as 11.5 gC m⁻² year⁻¹ for N. cristatus, 5.7 for N. plumchrus and 2.1 for N. flemingeri, yielding annual P/B ratio of 5 for each species. The annual production of Neocalanus accounted for 13.2% of the primary production in the Oyashio region. Their fecal pellets were estimated to account for 14.9% (0.7 gC m⁻² year⁻¹) of sinking flux of organic carbon at 1000-m depth. Moreover, their export flux by ontogenetic vertical migration, which is not measured by sediment trap observations, is estimated to be 91.5% (4.3 gC m⁻² year⁻¹) of carbon flux of sinking particles at 1000-m depth. These results suggest the important role of interzonal migrating copepods in the export flux of carbon.     

Upper ocean export of particulate organic carbon and biogenic silica in the Southern Ocean along 170°W

Upper-ocean fluxes of particulate organic carbon (POC) and biogenic silica (bSi) are calculated from four US JGOFS cruises along 170°W using a thorium-234 based approach. Both POC and bSi fluxes exhibit large variability vs. latitude during the seasonal progression of diatom dominated blooms. POC fluxes at 100 m of up to 50 mmol C m⁻² d⁻¹ are found late in the bloom, and farthest south near the Ross Sea Gyre. Biogenic Si fluxes also peak late in the bloom as high as 15 mmol Si m⁻² d⁻¹, but this flux peak occurs at a different latitude, just south of the Antarctic Polar Front (APF), which is centered around 60°S along this cruise track. The ratios of both POC and bSi export relative to their production rates are large, suggesting an efficient biological pump at these latitudes. The highest relative bSi/POC flux ratios at 100 m are found just south of the APF, coincident with a bSi/POC flux peak seen in 1000 m traps during this same program by Deep-Sea Research II (Honjo et al., Deep-Sea Research II 47, 3521–3548). These data suggest that efficient export at these latitudes can support the high accumulation rates of bSi found in the sediments under and south of the APF, despite the generally low biomass and productivity levels in this region.     

The carbonic system distribution and fluxes in the NE Atlantic during Spring 1991

The potential of the North Atlantic as a sink for atmospheric CO₂ was investigated by studying the carbonic system using data obtained during the spring of 1991. The air-sea flux of CO₂ was related to chlorophyll and other environmental variables, and the regeneration of carbon in the mid-ocean studied by examining vertical sections representative of the study area.Poor correlations were found between pCO₂ and chlorophyll throughout much of the study area, although a good correlation was found along 16°W. The highest air-sea fluxes of CO₂ were calculated for areas where chlorophyll was highest (45°13′N, 16°04′W), and where the greatest wind speeds occurred (47°51′N, 28°18′W). The mean CO₂ flux from the atmosphere to the ocean during the study period (May) was calculated as 0.65mmol m⁻²d⁻¹, which compares well with other studies. Regression equations were developed to predict total inorganic carbon from nutrients; errors were typically less than 1 μmol kg⁻¹. Regeneration of carbon in the mid-ocean occurred in two principal stages: 0–1000m and>2300m. Regeneration in the upper zone was dominated by soft tissue carbon (86%), with skeletal carbon (calcite) contributing only 14%. The fraction of regenerated carbon of skeletal origin increased to 51% in the>2300m zone.