Trophic pathways and carbon flux patterns in the Laptev Sea

The Laptev Sea is a high-Arctic epicontinental sea north of Siberia (Russia) that is one of the least understood regions of the world’s ocean. It is characterized by a shallow and broad shelf plateau, high influx of river water, sediments and nutrients during summer, long-lasting sea-ice cover from October to May, and the formation of a narrow flaw-lead polynya off the fast-ice edge during winter.Here, we describe results of a German–Russian research project (1993-present), presenting the distribution patterns and dynamics of its marine flora and fauna, as well as pathways and processes of coupling between sea-ice, water-column and sea-floor biota.Three ecological zones are distinguished along a combined east–west and Lena-impact gradient, differing in the composition of pelagic and benthic communities. In general, high Chl a concentrations in the sediments indicate a tight coupling between sympagic and pelagic primary production and nutrient supply to the benthos throughout the entire Laptev Sea. However, there were pronounced regional differences between the ecological zones in magnitude of primary production and trophic dynamics. Primary production during the ice-free summer was highest in the estuarine zone most strongly influenced by the Lena River (210 mg C m⁻² day⁻¹). The western and northeastern Laptev Sea yielded 55 and 95 mg C m⁻² day⁻¹, respectively. Moreover, the zones differed in the partitioning of carbon flux between zooplankton and benthic food webs. In the Lena zone zooplankton carbon demand was about 31 mg C m⁻² day⁻¹ whereas in the western zone it was 21 mg C m⁻² day⁻¹ and in the eastern zone 4 mg C m⁻² day⁻¹. Total benthic carbon demand was 32 mg C m⁻² day⁻¹ for the Lena zone, 56 mg C m⁻² day⁻¹ in the western zone and 100 mg C m⁻² day⁻¹ in the northeastern zone.A carbon budget constructed for the Laptev Sea indicates that (1) a high proportion of primary production is channelled through the benthic trophic web, bypassing the pelagic trophic web, and (2) autochthonous primary production in the northeastern and western Laptev Sea might not be sufficient to fuel both pelagic and benthic secondary production and, hence, input of allochthonous organic carbon is required to balance the overall carbon demand.     

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.     

Interannual and geographical variability of the brood size of the euphausiids Euphausia pacifica and Thysanoessa spinifera along the Oregon coast (1999–2004)

Brood sizes of 1259 adult female Euphausia pacifica and Thysanoessa spinifera were measured during 48 h incubations (10 °C, ±0.5 °C) on 27 oceanographic cruises between July 1999 and September 2004. The data set includes measurements from several stations off Newport, Oregon (Newport Hydrographic line, 44°39′N) made over a 5-year period and measurements from 14 more extensive cruises at stations representative of continental shelf, slope, and oceanic waters off Oregon and California, USA. E. pacifica had similar brood sizes at inshore (<200 m) and offshore (>200 m) stations with an average of 151 and 139 eggs brood⁻¹ fem⁻¹, respectively. T. spinifera brood sizes were considerably higher at inshore stations—particularly at Heceta Bank (44°N) and south of Cape Blanco (42°50′N)—than at offshore stations, 155 and 107 eggs brood⁻¹ fem⁻¹, respectively. Average brood sizes of E. pacifica increased during the study period, from 125 (in 2000) to 171 eggs brood⁻¹ fem⁻¹ (in 2003). Average percentage of carbon weight invested in spawning (reproductive effort) was higher in E. pacifica (14%) than in T. spinifera (6%), because both species have similar brood size but T. spinifera females are larger than E. pacifica females and produce smaller eggs. Reproductive effort for both species was higher during summer 2002, probably associated with anomalous cool subarctic waters and high chl-a concentration observed during that summer. Brood sizes and chl-a values remained relatively high in 2003–2004 compared to the 1999–2001 period. Geographical and temporal variability in brood sizes for both species were significantly correlated with in situ measurements of chl-a concentration but not with sea surface temperature. No gravid females were collected during late autumn and winter cruises, thus the spawning season along the Oregon coast appears to extend from March through September for both species. However, T. spinifera usually starts reproductive activity earlier in the spring (March) than E. pacifica. Both species had their highest brood sizes in summer during the period of most intense upwelling, which is associated with an increase in regional phytoplankton standing stock.     

Metridia pacifica in Dabob Bay, Washington: The diatom effect and the discrepancy between high abundance and low egg production rates

Information on life cycle strategies and reproductive parameters of Metridia pacifica is scarce, despite its importance in the zooplankton of the subarctic Pacific. In many regions it occurs in high abundance, but reproductive rates, when reported, are usually low. This discrepancy was studied in Dabob Bay, Washington, USA, in the context of an investigation of the effect of diatom blooms on the reproductive success of copepod grazers.In situ egg production rates of M. pacifica were measured in spring and mid-summer with standard methods (multi-wells) and a new incubation chamber (spawning towers) that separates the spawning female from its eggs and allows the eggs to develop undisturbed. Many females did not produce eggs, possibly due to a high fraction of immature individuals. Egg production rates were variable, but clutch sizes were higher in spawning towers, and estimates of female egg cannibalism revealed that females consume many eggs shortly after their release. Thus, a separation of females and eggs is mandatory for accurate measurements of M. pacifica egg production rates. The maximum clutch sizes recorded in our study were comparable to measurements for other calanoids. However, unviable eggs were a large fraction of those spawned, independent of incubation method, especially in late winter and early spring. In order to assess whether the diatom effect may be responsible for low viability of embryos and nauplii, we also measured in situ grazing. Adult females were omnivorous, but they ingested some diatoms that rank among the strongest anti-mitotic toxin producers known so far. Although M. pacifica’s vertical migration behavior suggests opportunistic feeding on abundant food during their short stay in the phytoplankton-rich surface, they often ignored the food items that contributed most to microplankton carbon concentrations. Thus, their feeding strategy remains ambiguous. Due to severe reproductive failure early in the season, recruitment was impaired in spring, while the population increased, reaching high abundance in mid-summer when reproductive output was low. While advection and interannual variability in bloom conditions might compensate for the losses described here, the paradox of high Metridia abundance versus low reproductive success still requires further investigation, and methodological constraints need to be ruled out in future studies.     

Carbon flux in the northwest Mediterranean estimated from microbial production

Spring profiles of microbial production derived from the dark incorporation of tritiated leucine and tritiated thymidine in the northwest Mediterranean show an exponential decline with depth. Assuming this to represent a steady-state balance between microbial respiration and the downward flux of carbon, the downward flux is estimated as (1−/)p/b, where p is the microbial production, their gross growth efficiency and b the coefficient of exponential decline with depth. Summer profiles, ranging over about 3° of latitude and 4° of longitude, were well fitted by a two-component exponential decline, suggesting two distinct microbial substrates. Values of b for the more rapidly declining component varied between 0.01 and 0.06 m⁻¹ according to location. In the case of the slower component, b was estimated as 0.002 m⁻¹, and did not vary significantly over the region. Estimated fluxes of carbon at the surface are 123–335 mg m⁻² d⁻¹ for the fast and 95 mg m⁻² d⁻¹ for the slow component. Below about 200 m, carbon flux is dominated by the slow component. Flux estimates are compatible with flux estimates from sediment traps in the same region. The observed changes between the spring and summer profiles, combined with the horizontal homogeneity of the summer profiles below 200 m, are consistent with a downward transport of about 5–10 m d^–1, implying a significant dispersive component to the observed fluxes.     

Mesozooplankton dynamics in nearshore waters of the Great Barrier Reef

Zooplankton dynamics (community composition, juvenile somatic growth rate, adult egg production, secondary production) were studied in coastal waters of the Great Barrier Reef. Two sectors were compared, one adjacent to a catchment of near-pristine land use patterns, the other to a more intensively farmed catchment. Sampling was conducted in the austral winter (August) and summer (January–March) of two succeeding years. Gradients in zooplankton community composition were weak, with only moderate effects of season and sector. Overall, 37% of zooplankton biomass was in the 73–150 μm size fraction, 26% in the 150–350 μm fraction, and 38% was >350 μm. There was no biomass difference and only small differences in community composition between samples taken during the day and at night; ostracods and large calanoid copepods were occasionally more common at night. Carbon-specific growth rates averaged 0.29 d⁻¹ for cyclopoid copepods and 0.35 d⁻¹ for calanoid copepods, with no difference between sectors. Calanoid copepod growth showed a significant relationship to chlorophyll concentration, but cyclopoid copepods did not. Copepod egg production was low (7.9 ± 5.9 eggs female⁻¹ d⁻¹) and apparently food-limited. Copepod secondary production was lower in August (mean = 2.6, range 1.4–4.0 mg C m⁻² d⁻¹) than in January–March (mean = 8.5, range 2.4–15.5 mg C m⁻² d⁻¹). Secondary production by mesozooplankton in the 73–100 μm size range averaged 0.9% of total phytoplankton production.     

The recovery of microalgal production and biomass in a South African temporarily open/closed estuary, following mouth breaching

Mouth breaching is a recurrent event in temporarily open/closed estuaries (TOCEs). Such disturbances result in flushing and sediment scouring, reducing the microalgal biomass stock. The depletion of these microalgae may have negative repercussions in the form of depleted stocks of commercial fish, game fish, crustaceans and mollusks. The aim of this investigation was therefore: (1) to monitor the recovery of microalgal biomass and production following a breaching event; and (2) to determine the key environmental parameters influencing primary production during the open and recovery phases. Phytoplankton and benthic microalgal production was measured (¹⁴C-uptake method) successively during the closed, open and recovery phases of the Mdloti TOCE (South Africa). Upon breaching, 94–99% of microalgal biomass was washed out to sea through flushing and sediment scouring. A temporary recovery of phytoplankton and benthic microalgal biomass was observed during the open phase, but this was not sustained because of continual flushing and scouring of the sediment. During the re-closure (recovery phase), microalgal biomass immediately increased, reaching pre-breaching levels 35–40 days following the breaching event. In contrast to biomass, autochthonous pelagic primary production reached a maximum level (341 mg C m⁻² h⁻¹) during the open phase. Pelagic primary production normalized to biomass (P^B) significantly increased during the open phase. This is attributed to a favorable combination of optimum light conditions, high influx of macronutrients and high water temperatures (33 °C). Similarly, benthic primary production normalized to biomass (P^B) peaked during the open phase (35 mg C mg chl-a⁻¹ h⁻¹). Multivariate analysis showed that major variations in primary production were mainly controlled by temperature, dissolved inorganic nitrogen (DIN) to phosphorus (DIP) molar ratios (water-column and pore-water) and light extinction (K_d), all of which were regulated by the state of the mouth.     

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.     

Effect of iron enrichment on the dynamics of transparent exopolymer particles in the western subarctic Pacific

Dynamics of transparent exopolymer particles (TEP) was studied during the first in situ iron-enrichment experiment conducted in the western subarctic Pacific in July–August 2001, with the goal of evaluating the contribution of TEP to vertical flux as a result of increased primary production following iron enrichment in open ocean ecosystems. Subsequent to the enhancement of phytoplankton production, we observed increase in TEP concentration in the surface layer and sedimentation of organic matter beneath it. Vertical profiles of TEP, chlorophyll a (Chl a) and particulate organic carbon (POC) were obtained from six depths between 5 and 70 m, from a station each located inside and outside the enriched patch. TEP and total mass flux were estimated from the floating sediment traps deployed at 200 m depth. Chl a and TEP concentrations outside the patch varied from 0.2 to 1.9 μg L⁻¹ and 40–60 μg XG equiv. L⁻¹, respectively. Inside the patch, Chl a increased drastically from day 7 reaching the peak of 19.2 μg L⁻¹ on day 13, which coincided with the TEP peak of 189 μg XG equiv. L⁻¹. TEP flux in the sediment trap increased from 41 to 88 mg XG equiv. m⁻² d⁻¹, with 8–14% contribution of TEP to total mass flux. This forms the basic data set on ambient concentrations of TEP in the western subarctic Pacific, and evaluation of the effect of iron enrichment on TEP.     

Remotely sensed sea-surface chlorophyll and POC flux at Deep Gulf of Mexico Benthos sampling stations

Biweekly composite averages of the standing stock of sea-surface chlorophyll (SSC) were derived from SeaWiFS satellite ocean-color data at 44 benthic sampling stations occupied along the continental slope and rise by the Deep Gulf of Mexico Benthos (DGoMB) program. At the 22 DGoMB sites north of 26°N and west of 91°W in the NW Gulf of Mexico, annual average SSC was 0.19 mg m⁻³, ranging at most locations from annual highs of about 0.3 mg m⁻³ in November–February to lows of about 0.1 mg m⁻³ in May–August. Comparison of three years of SeaWiFS data (January 1998–December 2000) showed little inter-annual variation at these NW Gulf stations. In contrast, at the 22 NE Gulf sites north of 26°N and east of 91°W, SSC averaged 2.8 times higher than in the NW Gulf, showing also strong inter-annual variation. Maxima in the NE region occurred in November–February and also during summers. The summer maxima were associated with Mississippi River water transported offshore to the east and southward by anticyclonic eddies in the NE Gulf. The apparent increases in SSC in June–August at NE Gulf stations reached average monthly concentrations >50% greater than in November–February. Based on a primary productivity model and a vertical flux model, the calculated export of particulate organic carbon (POC flux reaching the seafloor) was estimated as 18 mg C m⁻² day⁻¹ at the 22 NE Gulf stations, and 9 mg C m⁻² day⁻¹ at the 22 NW Gulf stations. These estimates are comparable to fluxes measured by benthic lander by others in the DGoMB program, which may drive the differences in west versus east bathymetric zonation and community structure of macrobenthos that were sampled with large box corers by others in the DGoMB program.     

Seasonal variability of leaf litter removal by crabs in a Kandelia candel mangrove forest in Jiulongjiang Estuary, China

The seasonal variability of leaf litter removal by crabs was observed from May 2006 to April 2007 in a Kandelia candel mangrove forest in Jiulongjiang Estuary, China. Daily average quantities of leaf fall ranged 0.85–3.86 gDW m⁻² d⁻¹, with high values in May, August, October and November. The whole-year's leaf fall was 6.48 t ha⁻¹ yr⁻¹ (1.81 gDW m⁻² d⁻¹). The standing stock of leaf litter on the forest floor was 7.78 gDW m⁻² averaged from the whole year's data, with the lowest value in December (1.23 gDW m⁻²) and the highest in April (16.18 gDW m⁻²). Annually averaged removal (consumption on mangrove floor + burial in burrows) rate of leaf litter by crabs was 0.59 gDW m⁻² d⁻¹. High seasonal variability was observed in the removal rates of leaf litter by crabs. Removal rates in the winter months (December, January and February) were 0.07–0.09 gDW m⁻² d⁻¹, much lower than those in other months with values of 0.59–1.18 gDW m⁻² d⁻¹. Annually averaged percentage of leaf fall removed by crabs was 33%, with the highest values in September (reached 76%) and the lowest values in winter months. Of leaf litter removed by crabs, a large proportion was buried by crabs, and only 12% was consumed by crabs on the forest floor. Leaf litter removal rate, consumption rate on the forest floor, percentages of leaf fall and standing stock removed on the forest floor were significantly positively correlated with air temperature, indicating that leaf removal ability by crabs was higher in warm months than in cold months.     

Grazing impact of the copepod community in the Oyashio region of the western subarctic Pacific Ocean

The role of copepod grazing on the ecosystem dynamics in the Oyashio region, western subarctic Pacific was investigated during six cruises from June 2001 to June 2002. In situ grazing rates of the copepod community (CGR) were measured by the gut fluorescence method in respect to developmental stages of dominant species. In terms of biomass, more than 80% of the copepod community was dominated by six large calanoid species (Neocalanus cristatus, Neocalanus flemingeri, Neocalanus plumchrus, Eucalanus bungii, Metridia pacifica and Metridia okhotensis) throughout the year. Resulting from the observed pattern of the interzonal migrating copepods, the CGR in the Oyashio region was divided into three phases, i.e. spring (bloom), summer (post-bloom) and autumn-winter phase. During the spring bloom, late copepodites of the interzonal migrating species, N. cristatus, N. flemingeri and E. bungii appeared in the surface layer (0-50 m) to consume the production of the bloom, resulting in a high grazing rate of the copepod community (7.9 mg Chl m⁻² d⁻¹), though its impact on phytoplankton community was low due to the high primary productivity. During the post-bloom period, although the copepod community which was dominated by N. cristatus, N. plumchrus, M. pacifica and newly recruited E. bungii still maintained a high biomass, the CGR was generally lower (1.8-2.6 mg Chl m⁻² d⁻¹ for June and August 2001), probably due to the lower availability of phytoplankton. Nevertheless, the highest CGR was also observed during this period (10.5 mg Chl m⁻² d⁻¹ in June 2002). The high CGR on autotrophic carbon accounted for 69% of the primary production, suggesting that the copepod community in the Oyashio region potentially terminates the phytoplankton bloom. Abundant occurrence of young E. bungii, which is a characteristic phenomenon in the Oyashio region, was largely responsible for the high grazing pressure in June 2002 suggesting that success of reproduction, growth, and survival in E. bungii during the spring bloom is an important factor in controlling phytoplankton abundance during the post-bloom season. During autumn and winter, CGR was the lowest in the year (0.29-0.38 mg Chl. m⁻² d⁻¹) due to the disappearance of the interzonal migrating copepods from the surface layer. Diel migrant M. pacifica was the most important grazer during this period. The annual ingestion of the copepod community is estimated as 37.7 gC m⁻² on autotrophic carbon (converted using C:Chl ratio of 30) or 137.9 gC m⁻² on suspended particles (using C:Chl ratio of in situ value, 58-191), accounting for 13% and 46% of annual primary production, respectively. This study confirms that copepod grazing is an important pathway in carbon flow in the Oyashio region and in particular their role in the phytoplankton dynamics is significant for the termination of the spring bloom.     

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.     

Particle export within cyclonic Hawaiian lee eddies derived from Pb–Po disequilibrium

Particle export from the upper waters of the oligotrophic ocean may play a crucial role in the global carbon cycle. Mesoscale eddies have been hypothesized to inject new nutrients into oligotrophic surface waters, thereby increasing new production and particle export in otherwise nutrient deficient regimes. The E-Flux Program was a large multidisciplinary project designed to investigate the physical, biological and biogeochemical characteristics of cold-core cyclonic eddies that form in the lee of the Hawaiian Islands. There, we investigated particle dynamics using ²¹⁰Pb–²¹⁰Po disequilibrium. Seawater samples for ²¹⁰Pb and ²¹⁰Po were collected both within (IN) and outside (OUT) of two cyclones, Noah and Opal, at different stages of their evolution as well as from the eddy generation region. Particulate carbon (PC), particulate nitrogen (PN) and biogenic silica (bSiO₂) export fluxes were determined using water-column PC, PN, and bSiO₂ inventories and the residence times of ²¹⁰Po. PC and PN fluxes at 150 m ranged from 1.58±0.10 to 1.71±0.16 mmol C m⁻² d⁻¹ and 0.22±0.02 to 0.30±0.02 mmol N m⁻² d⁻¹ within Cyclones Opal and Noah. PC and PN fluxes at OUT stations sampled during both cruises were of similar magnitudes, 1.69±0.16 to 1.67±0.16 mmol C m⁻² d⁻¹ and 0.30±0.03 to 0.26±0.03 mmol N m⁻² d⁻¹. The bSiO₂ fluxes within Cyclone Opal were 0.157±0.010 mmol Si m⁻² d⁻¹ versus 0.025±0.002 mmol Si m⁻² d⁻¹ at OUT stations. These results of minimal PC and PN export, but significant eddy-induced bSiO₂ fluxes, agree very well with other studies that used a variety of direct and indirect methods. Thus, our results suggest that using elemental inventories and residence times of ²¹⁰Po is another independent and robust method for determining particle export and should be investigated more fully.     

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.     

Phytoplankton productivity in the western subarctic gyre of the North Pacific in early summer 2006

We deployed a profiling buoy system incorporating a fast repetition rate fluorometer in the western subarctic Pacific and carried out time-series observations of phytoplankton productivity from 9 June to 15 July 2006. The chlorophyll a (Chl a) biomass integrated over the euphotic layer was as high as 45–50 mg Chl a m⁻² in the middle of June and remained in the 30–40 mg Chl a m⁻² range during the rest of observation period; day-to-day variation in Chl a biomass was relatively small. The daily net primary productivity integrated over the euphotic layer ranged from 144 to 919 mg C m⁻² day⁻¹ and varied greatly, depending more on insolation rather than Chl a biomass. In addition, we found that part of primary production was exported to a 150-m depth within 2 days, indicating that the variations in primary productivity quickly influenced the organic carbon flux from the upper ocean. Our results suggest that the short-term variability in primary productivity is one of the key factors controlling the carbon cycle in the surface ocean in the western subarctic Pacific.     

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.     

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.     

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.     

A 9-year time-series of planktonic foraminifer fluxes and environmental change in the Bering sea and the central subarctic Pacific Ocean, 1990–1999

A 9-year study of planktonic foraminifer fluxes was conducted in the Bering Sea (Station AB) and in the central subarctic Pacific (Station SA). Results clearly reflected variations of the water mass characteristics in the upper layers. The 9-year means of total foraminifer fluxes were the same (1400 shells m⁻² d⁻¹) at both stations. However, total foraminifer flux at Station AB tended to show its primary maximum during fall (October–December) and its secondary maximum in spring (April–June), whereas the primary maximum appeared in spring and the secondary maximum in fall at Station SA. Seasonal variation was more apparent at hemipelagic Station AB than at pelagic Station SA. Planktonic foraminifers found at both stations were of six species: Neogloboquadrina pachyderma, Globigerina umbilicata, Globigerinita glutinata, Globigerina quinqueloba, Globorotalia wilesi, and Orbulina universa. The foraminifer assemblages at the two stations reflected the temperature difference in the surface waters. The variable %G. umbilicata tended to be high in the warm surface waters during the summers. The temporal and geographical variation of %G. quinqueloba indicated that this taxon prefers regions with relatively low diatom fluxes. A notable appearance of O. universa occurred in 1997 at Station SA. During this period, other measured biogenic particle fluxes, such as those of diatoms, were low. This unusual 1997 event may be a reflection of global climatic change that happened to be observed in the central subarctic Pacific Ocean.