The metabolic balance between autotrophy and heterotrophy in the western Arctic Ocean

The goal of this study was to explore how net community production (NCP) is influenced by the relationship between primary production and community respiration in the western Arctic Ocean. Plankton NCP and respiration were determined by measuring changes in oxygen in light and dark bottle incubations, respectively. Rates of NCP averaged over shelf, slope and basin waters were positive in summer 2002 (57±191 mmol O₂ m⁻² d⁻¹) and spring 2004 (85±86 mmol O₂ m⁻² d⁻¹) and negative in summer 2004 (−25±176 mmol O₂ m⁻² d⁻¹). Determinations of NCP obtained from bottle incubations were similar to rates inferred from in situ changes in dissolved inorganic carbon. An examination of the spatial variability of primary production and community respiration indicated that respiration is distributed more uniformly than primary production. A spatial offset between photosynthesis and respiration from the shelf to the Arctic basin was present in spring 2004, but was not seen at other times. NCP and the potential for export appear to be dependent on an uncoupling of primary production and community respiration. NCP continued into the summer after the stock of NO₃⁻ had been depleted. Our data suggest that the uniform distribution of respiration relative to primary production is an important factor influencing NCP and the potential for export in the western Arctic.     

Microzooplankton grazing impact in the Western Arctic Ocean

Microzooplankton grazing impact on phytoplankton was assessed using the Landry–Hassett dilution technique in the Western Arctic Ocean during spring and summer 2002 and 2004. Forty experiments were completed in a region encompassing productive shelf regions of the Chukchi Sea, mesotrophic slope regions of the Beaufort Sea off the North Slope of Alaska, and oligotrophic deep-water sites in the Canada Basin. A variety of conditions were encountered, from heavy sea-ice cover during both spring cruises, moderate sea-ice cover during summer of 2002, and light to no sea ice during summer of 2004, with a concomitant range of trophic conditions, from low chlorophyll-a (Chl-a; <0.5 μg L⁻¹) during heavy ice cover in spring and in the open basin, to late spring and summer shelf and slope open-water diatom blooms with Chl-a >5 μg L⁻¹. The microzooplankton community was dominated by large naked ciliates and heterotrophic gymnodinoid dinoflagellates. Significant, but low, rates of microzooplankton herbivory were found in half of the experiments. The maximum grazing rate was 0.16 d⁻¹ and average grazing rate, including experiments with no significant grazing, was 0.04±0.06 d⁻¹. Phytoplankton intrinsic growth rates varied from the highest values of about 0.4 d⁻¹ to the lowest values of zero to slightly negative growth, on average 0.16±0.15 d⁻¹. Light limitation in spring and post-bloom senescence during summer were likely explanations of observed low phytoplankton growth rates. Microzooplankton grazing consumed 0–120% (average 22±26%) of phytoplankton daily growth. Grazing and growth rates found in this study were low compared to rates reported in another Arctic system, the Barents Sea, and in major geographic regions of the world ocean.     

Standing stocks,production, and respiration of phytoplankton and heterotrophic bacteria in the western Arctic Ocean

Standing stocks and production rates for phytoplankton and heterotrophic bacteria were examined during four expeditions in the western Arctic Ocean (Chukchi Sea and Canada Basin) in the spring and summer of 2002 and 2004. Rates of primary production (PP) and bacterial production (BP) were higher in the summer than in spring and in shelf waters than in the basin. Most surprisingly, PP was 3-fold higher in 2004 than in 2002; ice-corrected rates were 1581 and 458 mg C m⁻² d⁻¹, respectively, for the entire region. The difference between years was mainly due to low ice coverage in the summer of 2004. The spatial and temporal variation in PP led to comparable variation in BP. Although temperature explained as much variability in BP as did PP or phytoplankton biomass, there was no relationship between temperature and bacterial growth rates above about 0 °C. The average ratio of BP to PP was 0.06 and 0.79 when ice-corrected PP rates were greater than and less than 100 mg C m⁻² d⁻¹, respectively; the overall average was 0.34. Bacteria accounted for a highly variable fraction of total respiration, from 3% to over 60% with a mean of 25%. Likewise, the fraction of PP consumed by bacterial respiration, when calculated from growth efficiency (average of 6.9%) and BP estimates, varied greatly over time and space (7% to >500%). The apparent uncoupling between respiration and PP has several implications for carbon export and storage in the western Arctic Ocean.     

Seasonal and spatial patterns of sedimentary denitrification rates in the Chukchi sea

Recent constructions of the global nitrogen budget estimate that at least half of the ocean's fixed nitrogen is lost by sedimentary denitrification, the majority of which occurs in continental shelves. The Arctic contains approximately 20% of the world's continental shelf, suggesting it is a substantial contributor to the global sedimentary denitrification rate. During two cruises in the summer and spring of 2002 and 2004, respectively, denitrification rates were calculated from the downward diffusive flux of nitrate in the shelf and slope sediments of the Chukchi Sea in the western Arctic. Additionally, in the spring of 2004, denitrification rates were determined by whole-core incubations in which the flux of nitrogen gas out of the sediments was measured. Measurements were made along three transects crossing the shelf and slope (50–3000 m), each transect having different overlying water characteristics. Denitrification rates generally decreased with increasing water depth: rates varied from about 1.6 mmol N m⁻² d⁻¹ for the shallow-water sediments to undetectable in deep-water sediments. Rates showed little variation between the two seasons. However, rates were found to correspond with differences in annual overlying primary productivities and particulate organic carbon export fluxes. An extrapolation to the whole Arctic yielded an average Arctic sedimentary denitrification rate of 13 Tg N yr⁻¹. Taken in the context of the global nitrogen budget, it is about 4–13% of the total sink of fixed nitrogen in the ocean.     

Phytoplankton of the western Arctic in the spring and summer of 2002: Structure and seasonal changes

We analyzed the taxonomic structure and spatial variability of phytoplankton abundance and biomass in the Chukchi and Beaufort Seas during spring and summer seasons of the SBI program. Phytoplankton samples were collected during two surveys from May 10 to June 13 and from July 19 to August 21 of 2002. In May and June, ice cover exceeded 80% over most of the study area and there was no vertical stratification, indicating that the successional state of the phytoplankton corresponded to the end of the winter biological season. The phytoplankton abundance ranged from a few tens to a few thousands of cells per liter, while biomass varied from 0.1 to 3.0 mg C m⁻³. Small areas of high phytoplankton abundance (0.13–1.3×10⁶ cells L⁻¹) and biomass (22–536 mg C m⁻³), dominated by early spring diatoms Pauliella taeniata and Fragilariopsis oceanica in the surface waters, which indicated the beginning of the spring bloom, were observed only in the southeastern part of the Chukchi shelf and off Point Barrow. In July and August summer period, more than a half of the study area had <50% ice cover and the water column was stratified by temperature and salinity. Over the Chukchi shelf and continental slope of the Beaufort Sea, the phytoplankton abundance and biomass were an order of magnitude higher in July–August than in May–June. The taxonomic diversity of algae also increased due to the appearance of late-spring and summer diatoms, dinoflagellates, and coccolithophorids (Emiliania huxleyi). Interestingly, the seasonal differences between phytoplankton abundance and taxonomic composition in the spring and summer periods varied the least over the Chukchi Sea slope and in the deep-water area of the Arctic Ocean. High algae concentrations in summer were located in the lower layers of the euphotic zone, suggesting that the spring bloom on both the Chukchi shelf and in the western part of the Beaufort Sea occurred in late June/early July. In the spring and summer, the microalgal community was characterized by a high abundance of 4–10 μm flagellates, which exceeded the abundance of all other taxonomic groups. In both seasons studied, phytoplankton reached its maximum abundance within restricted areas in the southern part of the Chukchi Sea southwest of Point Hope, in the northern part of the Chukchi shelf between the 50- and 100-m isobaths, on the shelf northwest of Point Barrow, and over the continental slope in the Beaufort Sea. The pronounced spatial difference in the seasonal state was a characteristic feature of the phytoplankton community in the western Arctic.     

Spatio-temporal distribution of dissolved inorganic carbon and net community production in the Chukchi and Beaufort Seas

As part of the 2002 Western Arctic Shelf–Basin Interactions (SBI) project, spatio-temporal variability of dissolved inorganic carbon (DIC) was employed to determine rates of net community production (NCP) for the Chukchi and western Beaufort Sea shelf and slope, and Canada Basin of the Arctic Ocean. Seasonal and spatial distributions of DIC were characterized for all water masses (e.g., mixed layer, halocline waters, Atlantic layer, and deep Arctic Ocean) of the Chukchi Sea region during field investigations in spring (5 May–15 June 2002) and summer (15 July–25 August 2002). Between these periods, high rates of phytoplankton production resulted in large drawdown of inorganic nutrients and DIC in the Polar Mixed Layer (PML) and in the shallow depths of the Upper Halocline Layer (UHL). The highest rates of NCP (1000–2850 mg C m⁻² d⁻¹) occurred on the shelf in the Barrow Canyon region of the Chukchi Sea and east of Barrow in the western Beaufort Sea. A total NCP rate of 8.9–17.8×10¹² g for the growing season was estimated for the eastern Chukchi Sea shelf and slope region. Very low inorganic nutrient concentrations and low rates of NCP (<15–25 mg C m⁻² d⁻¹) estimated for the mixed layer of the adjacent Arctic Ocean basin indicate that this area is perennially oligotrophic.     

Mesozooplankton prey preference and grazing impact in the western Arctic Ocean

The role of mesozooplankton as consumers and transformers of primary and secondary production in the Beaufort and Chukchi Seas was examined during four cruises in spring and summer of both 2002 and 2004 as part of the western Arctic Shelf–Basin Interactions (SBI) program. Forty-seven grazing experiments using dominant mesozooplankton species and life stages were conducted at locations across the shelf, slope, and basin of the Chukchi and Beaufort Seas to measure feeding rates on both chlorophyll and microzooplankton and to determine mesozooplankton prey preferences.Mesozooplankton biomass was at all times dominated by life stages of four copepod taxa: Calanus glacialis, Calanus hyperboreus, Metridia longa, and Pseudocalanus spp. Significant interannual, seasonal, regional, between species and within species differences in grazing rates were observed. Overall, the dominant zooplankton exhibited typical feeding behavior in response to chlorophyll concentration that could be modeled using species and life-stage specific Ivlev functions. Microzooplankton were preferred prey at almost all times, with the strength of the preference positively related to the proportion of microzooplankton prey availability. Average mesozooplankton grazing impacts on both chlorophyll standing stock (0.6±0.5% d⁻¹ in spring, 5.1±6.3% d⁻¹ in summer) and primary production (12.8±11.8% d⁻¹ in spring, 27.6±24.5% d⁻¹ in summer) were quite low and varied between shelf, slope, and basin. Coincident microzooplankton grazing experiments [Sherr, E.B., Sherr, B.F., Hartz, A.J., 2009. Microzooplankton grazing impact in the Western Arctic Ocean. Deep-Sea Research II] were conducted at most stations. Together, microzooplankton–mesozooplankton grazing consumed only 44% of the total water-column primary production, leaving more than half directly available for local export to the benthos or for offshore transport into the adjacent basin.     

Community respiration/production and bacterial activity in the upper water column of the central Arctic Ocean

Community metabolism (respiration and production) and bacterial activity were assessed in the upper water column of the central Arctic Ocean during the SHEBA/JOIS ice camp experiment, October 1997–September 1998. In the upper 50 m, decrease in integrated dissolved oxygen (DO) stocks over a period of 124 d in mid-winter suggested a respiration rate of ∼3.3 nM O₂ h⁻¹ and a carbon demand of ∼4.5 gC m⁻². Increase in 0–50 m integrated stocks of DO during summer implied a net community production of ∼20 gC m⁻². Community respiration rates were directly measured via rate of decrease in DO in whole seawater during 72-h dark incubation experiments. Incubation-based respiration rates were on average 3-fold lower during winter (11.0±10.6 nM O₂ h⁻¹) compared to summer (35.3±24.8 nM O₂ h⁻¹). Bacterial heterotrophic activity responded strongly, without noticeable lag, to phytoplankton growth. Rate of leucine incorporation by bacteria (a proxy for protein synthesis and cell growth) increased ∼10-fold, and the cell-specific rate of leucine incorporation ∼5-fold, from winter to summer. Rates of production of bacterial biomass in the upper 50 m were, however, low compared to other oceanic regions, averaging 0.52±0.47 ngC l⁻¹ h⁻¹ during winter and 5.1±3.1 ngC l⁻¹ h⁻¹ during summer. Total carbon demand based on respiration experiments averaged 2.4±2.3 mgC m⁻³ d⁻¹ in winter and 7.8±5.5 mgC m⁻³ d⁻¹ in summer. Estimated bacterial carbon demand based on bacterial productivity and an assumed 10% gross growth efficiency was much lower, averaging about 0.12±0.12 mgC m⁻³ d⁻¹ in winter and 1.3±0.7 mgC m⁻³ d⁻¹ in summer. Our estimates of bacterial activity during summer were an order of magnitude less than rates reported from a summer 1994 study in the central Arctic Ocean, implying significant inter-annual variability of microbial processes in this region.     

Seasonal dynamics of primary and new production in the northern South China Sea: The significance of river discharge and nutrient advection

Biochemical and productivity measurements and nutrient enrichment experiments were conducted on three cruises in summer and two cruises in winter on the shelf and the basin of the northern South China Sea (SCS) between 2001 and 2004. Phytoplankton production, in terms of depth-integrated new production (INP) or depth-integrated primary production (IPP), was higher in winter than in summer and on the shelf than in the basin. In winter, with deepening of the mixed layer, nitrate from the shallow nitracline that characterized the SCS waters was made available in the surface and supported the highest production of the year. Averaged INP measured in winter (0.25 g C m⁻² d⁻¹) was about twice the summer average (0.12 g C m⁻² d⁻¹) and was 0.19 g C m⁻² d⁻¹ on the shelf compared with 0.15 g C m⁻² d⁻¹ in the basin. In winter, average INP on the shelf was higher than the basin (0.34 versus 0.21 g C m⁻² d⁻¹); whereas in summer, averaged INP on the shelf (0.13 g C m⁻² d⁻¹) and the basin (0.11 g C m⁻² d⁻¹) were similar. While averaged IPP measured in the basin was higher in winter than in summer (0.53 versus 0.35 g C m⁻² d⁻¹), IPP on the shelf showed little temporal variation (0.82 in winter versus 0.84 g C m⁻² d⁻¹ in summer). Considerable spatial and inter-annual variation in production was measured in the shelf waters during summer, which could be linked to discharge volume and plume flow direction of the Zhujiang River. While the shelf waters in summer were mostly nitrogen starved or nitrogen and phosphorus co-limited, excessive river runoff may cause the nutritive state to shift to phosphorus deficiency. Waters with low surface salinities and high fluorescence from riverine mixing could be found extending from the Zhujiang mouth to as far as offshore southern Taiwan after a typhoon passed the northern SCS and brought heavy rainfall. Overall, both nutrient advection in winter and river discharge from the China coast in summer made new nitrogen available and shaped the dynamics of phytoplankton production in these oligotrophic waters.     

Rapid removal of terrigenous dissolved organic carbon over the Eurasian shelves of the Arctic Ocean

The fate of terrigenous dissolved organic carbon (tDOC) delivered to the Arctic Ocean by rivers remains poorly constrained on both spatial and temporal scales. Early reports suggested Arctic tDOC was refractory to degradation, while recent studies have shown tDOC removal to be an active but slow process. Here we present observations of DOC, salinity, δ¹⁸O, and ²²⁸Ra/²²⁶Ra in the Polar Surface Layer (PSL) over the outer East Siberian/Chukchi shelf and the adjacent Makarov and Eurasian basins of the eastern Arctic Ocean. This off-shelf system receives meteoric water, introduced by rivers, after a few years residence on the shelf. Elevated concentrations of DOC (> 120 μM C) were observed in low salinity (~ 27) water over the Makarov Basin, suggesting inputs of tDOC-enriched river water to the source waters of the Transpolar Drift. The regression of DOC against salinity indicated an apparent tDOC concentration of 315 ± 7 μM C in the river water fraction, which is significantly lower than the estimated DOC concentration in the riverine sources to the region (724 ± 55 μM C). To obtain the timescale of removal, estimates of shelf residence were coupled with measurements of dissolved ²²⁸Ra/²²⁶Ra, an isotopic tracer of time since shelf residence. Shelf residence time coupled with DOC distributions indicates a first order tDOC removal rate constant, λ = 0.24 ± 0.07 yr^-1, for the eastern Arctic, 2.5–4 times higher than rates previously observed in the western Arctic. The observed removal of tDOC in the eastern Arctic occurs over the expansive shelf area, highlighting the initial lability of tDOC upon delivery to the Arctic Ocean, and suggests that tDOC is composed of multiple compartments defined by reactivity. The relatively rapid remineralization of tDOC on the shelves may mitigate the strength of the Arctic Ocean atmospheric CO₂ sink if a projected increase in labile tDOC flux occurs.     

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

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

Zooplankton inside an Arctic Ocean cold-core eddy: Probable origin and fate

In September 2004, an extensive survey of a cold-core eddy in the Canada Basin, western Arctic was carried out with high-horizontal-resolution physical and chemical sampling and lower-horizontal-resolution biological sampling. The eddy was located over the continental slope north of the Chukchi Shelf and had a radius of ∼8 km. Its core was centered at a depth of ∼160 m. Water mass characteristics and the presence of copepods from the North Pacific Ocean (Neocalanus flemingeri and Metridia pacifica) demonstrated that the core contained water of Pacific origin. Vertical distributions of zooplankton were associated with the physical structure of the water column. For most taxa, concentrations in the eddy core were elevated compared with those in similar density water in the surrounding Basin. Based on tracer-age estimates and previous observations of eddy formation, the eddy is believed to have been formed during the previous spring/summer from the Chukchi shelfbreak jet. Surprisingly, the eddy also contained elevated abundances of Arctic-origin copepods (Metridia longa and Calanus glacialis). Analysis of a shelf–basin transect occupied in the region in August 2004 showed that these species were present in high abundances in relatively shallow water (50 m) inshore of the shelfbreak due to upwelling of deeper basin water, and copepods, onto the shelf in response to easterly winds. If the formation of the observed eddy occurred during, or shortly after, a period of such winds, upwelled Arctic-origin copepods on the shelf might have been entrained into the feature. Our observations suggest that formation and subsequent migration of such eddies may provide a mechanism for transporting zooplankton from the Chukchi Shelf into the interior Canada Basin. The periodic input of high abundances of zooplankton from productive shelf areas could affect food webs in the less productive basin.     

Dissolved inorganic carbon,alkalinity, nutrient and oxygen seasonal and interannual variations at the Antarctic Ocean JGOFS-KERFIX site

JGOFS-KERFIX (KERguelen point FIXe) time-series station, located south of the polar front in the Indian sector of the Antarctic Ocean, was occupied monthly between January 1990 and March 1995. Annual cycles of dissolved inorganic carbon (DIC), total alkalinity (TALK), oxygen (O₂) and nutrients (nitrate, silicate, phosphate and ammonia) in the upper ocean are presented for this site. From seasonal drawdown of nutrients and DIC, we estimate a spring–summer net community production of 3.2±0.5 mol m⁻² and C/N/P ratios of 100/16/1. The Si/N ratio varies between 1.8 and 3, suggesting low iron concentrations. The spring–summer biogenic silicon export derived from silicate drawdown is 1.18 mol m⁻², consistent with model estimates of silicate export at this site. Seasonal and interannual variations of oxygen, nitrate and DIC due to physical and biological processes are quantified using a simple month-to-month budget formulation. From these budgets, an annual net community production of 5.7±3.3 mol m⁻² yr⁻¹ is estimated, about twice the averaged spring–summer production, indicating that, at KERFIX, there is a positive net community production throughout the year. Air–sea CO₂ fluxes show that KERFIX is a strong CO₂ sink for the atmosphere of 2.4–5.1 mol m⁻² yr⁻¹ in 1993, depending on the gas exchange formulation used. A 2.1–3.3 mol m⁻² yr⁻¹ outgassing of O₂ is observed at KERFIX except in 1993 and 1994 where a decreasing trend of temperature induces an increase of O₂ solubility.     

Primary productivity,differential size fraction and pigment composition responses in two Southern Ocean in situ iron enrichments

Two in situ iron-enrichment experiments were conducted in the Pacific sector of the Southern Ocean during summer 2002 (SOFeX). The “north patch,” established within the Subantarctic Zone (∼56°S), was characterized by high nitrate (∼21 mmol m⁻³) but low silicic acid (2 mmol m⁻³) concentrations. North patch iron enrichment increased chlorophyll (Chl) by 12-fold to 2.1 mg m⁻³ and primary productivity (PP_EU) by 8-fold to 188 mmol C m⁻² d⁻¹. Surprisingly, despite low silicic acid concentrations, diagnostic pigment and size-fraction composition changes indicated an assemblage shift from prymnesiophytes toward diatoms. The “south patch,” poleward of the Southern Boundary of the Antarctic Circumpolar Current (SBACC) (∼66°S), had high concentrations of nitrate (∼27 mmol m⁻³) and silicic acid (64 mmol m⁻³). South patch iron enrichment increased Chl by 9-fold to 3.8 mg m⁻³ and PP_EU 5-fold to 161 mmol C m⁻² d⁻¹ but, notably, did not alter the phytoplankton assemblage from the initial composition of ∼50% diatoms. South patch iron addition also reduced total particulate organic carbon:Chl from ∼300 to 100; enhanced the presence of novel non-photosynthetic, but fluorescent, compounds; and counteracted a decrease in photosynthetic performance as photoperiod decreased. These experiments show unambiguously that in the contemporary, high nitrate Southern Ocean increasing iron supply increases primary productivity, confirming the initial premise of the Martin Iron Hypothesis. However, despite a 5-fold increase in PP_EU under iron-replete conditions in late summer, the effect of iron on annual productivity in the Southern Ocean poleward of the SBACC is limited by seasonal ice coverage and the dark of polar winter.     

Nutrient and phytoplankton dynamics off the west coast of Vancouver Island during the 1997/98 ENSO event

Six research cruises were conducted off the west coast of Vancouver Island between April and October of 1997 and 1998 as part of the Canadian GLOBEC project to compare nutrient and phytoplankton dynamics between ENSO (1997) and non-ENSO (1998) years. Limited sampling also was conducted during three cruises in 1999. During the 1997 ENSO period, there was a shallow thermocline (∼10 m) that resulted in a shallower mixed layer, lower salinity and density, and stronger summer stratification. In general on the shelf, the 1997 growing season was characterized by higher nitrate (7.5 μM) and silicic acid (17 μM) concentrations, lower total chlorophyll (∼76 mg m⁻²), lower phytoplankton carbon biomass (0.2 mg C L⁻¹), and lower diatom abundance and biomass than in 1998. Phytoplankton assemblages were dominated by nanoplankton in 1997 and by diatoms in 1998. These results suggest that the 1997 ENSO was responsible for a reduction in the growth and biomass of larger phytoplankton cells. In mid-1998, the hydrographic characteristics off the west coast of Vancouver Island changed suddenly. The 1997 poleward transport of warm water reversed to an equatorward transport of coastal water in July 1998, which was accompanied by normal summer upwelling. During 1998, a large diatom bloom (mainly dominated by Chaetoceros debilis, Leptocylindrus danicus and to a lesser extent by Skeletomema and Pseudo-nitzschia sp.) was observed in July over the continental shelf. This large bloom resulted in chlorophyll concentrations of up to 400 mg m⁻², primary productivity of up to 11 g C m⁻² d⁻¹, and near undetectable dissolved nitrogen concentrations at some of the shelf stations in 1998. In contrast, during 1997, the sub-tropical waters that were advected over the slope, resulted in low chlorophyll a and primary productivity (generally <1 g C m⁻² d⁻¹). Therefore, there was a sharp contrast between the very high primary productivity on the shelf in July 1998, due to normal nutrient replenishment from summer upwelling and outflow from the Strait of Juan de Fuca, and the lower primary productivity during the 1997 ENSO year. During 1998, non-ENSO conditions resulted in phytoplankton biomass that was twice as high on the shelf as that measured in regions beyond the continental shelf of the west coast of Vancouver Island.     

Dry atmospheric deposition and diazotrophy as sources of new nitrogen to northwestern Mediterranean oligotrophic surface waters

Atmospheric dry deposition of nitrogen (N) and dinitrogen (N₂) fixation rates were assessed in 2004 at the time-series DYFAMED station (northwestern Mediterranean, 43°25′N, 7°52′E). The atmospheric input was monitored over the whole year. Dinitrogen fixation was measured during different seasonal trophic states (from mesotrophy to oligotrophy) sampled during nine cruises. The bioavailability of atmospherically deposited nutrients was estimated by apparent solubility after 96 h. The solubility of dry atmospheric N deposition was highly variable (from ∼18% to more than 96% of total N). New N supplied to surface waters by the dry atmospheric deposition was mainly nitrate (NO₃⁻) (∼57% of total N, compared to ∼6% released as ammonium (NH₄⁺)). The mean bioavailable dry flux of total N was estimated to be ∼112 μmol m⁻² d⁻¹ over the whole year. The NO₃⁻ contribution (70 μmol NO₃⁻ m⁻² d⁻¹) was much higher than the NH₄⁺ contribution (1.2 μmol NH₄⁺ m⁻² d⁻¹). The N:P ratios in the bioavailable fraction of atmospheric inputs (122.5–1340) were always much higher than the Redfield N:P ratio (16). Insoluble N in atmospheric dry deposition (referred to as “organic” and believed to be strongly related to anthropogenic emissions) was ∼40 μmol m⁻² d⁻¹. N₂ fixation rates ranged from 2 to 7.5 nmol L⁻¹ d⁻¹. The highest values were found in August, during the oligotrophic period (7.5 nmol L⁻¹ at 10 m depth), and in April, during the productive period (4 nmol L⁻¹ d⁻¹ at 10 m depth). Daily integrated values of N₂ fixation ranged from 22 to 100 μmol N m⁻² d⁻¹, with a maximum of 245 μmol N m⁻² d⁻¹ in August. No relationship was found between the availability of phosphorus or iron and the observed temporal variability of N₂ fixation rates. The atmospheric dry deposition and N₂ fixation represented 0.5–6% and 1–20% of the total biological nitrogen demand, respectively. Their contribution to new production was more significant: 1–28% and 2–55% for atmospheric dry deposition and N₂ fixation, respectively. The dry atmospheric input was particularly significant in conditions of water column stratification (16–28% of new production), while N₂ fixation reached its highest values in June (46% of new production) and in August (55%).     

Sea-ice algae: Major contributors to primary production and algal biomass in the Chukchi and Beaufort Seas during May/June 2002

Sea-ice and water samples were collected at 14 stations on the shelves and slope regions of the Chukchi and Beaufort Seas during the spring 2002 expedition as part of the Shelf–Basin Interaction Studies. Algal pigment content, particulate organic carbon and nitrogen, and primary productivity were estimated for both habitats based on ice cores, brine collection and water samples from 5-m depth. The pigment content (0.2–304.3 mg pigments m⁻²) and primary productivity (0.1–23.0 mg C m⁻³ h⁻¹) of the sea-ice algae significantly exceeded water-column parameters (0.2 and 1.0 mg pigments m⁻³; <0.1–0.4 mg C m⁻³ h⁻¹), making sea ice the habitat with the highest food availability for herbivores in early spring in the Chukchi and Beaufort Seas. Stable isotope signatures for ice and water samples did not differ significantly for δ¹⁵N, but for δ¹³C (ice: −25.1‰ to −14.2‰; water: −26.1‰ to −22.4‰). The analysis of nutrient concentrations and the pulse-amplitude-modulated fluorescence signal of ice algae and phytoplankton indicate that nutrients were the prime limiting factor for sea-ice algal productivity. The estimated spring primary production of about 1–2 g C m⁻² of sea-ice algae on the shelves requires the use of substantial nutrient reservoirs from the water column.     

Sulfate reduction in Black Sea sediments: in situ and laboratory radiotracer measurements from the shelf to 2000 m depth

Sulfate reduction rate measurements by the ³⁵SO₄²⁻ core injection method were carried out in situ with a benthic lander, LUISE, and in parallel by shipboard incubations in sediments of the Black Sea. Eight stations were studied along a transect from the Romanian shelf to the deep western anoxic basin. The highest rates measured on an areal basis for the upper 0–15 cm were 1.97 mmol m⁻² d⁻¹ on the shelf and 1.54 mmol m⁻² d⁻¹ at 181 m water depth just below the chemocline. At all stations sulfate reduction rates decreased to values <3 nmol cm⁻³ d⁻¹ below 15 cm depth in the sediment. The importance of sulfate reduction relative to the total mineralization of organic matter was very low, 6%, on the inner shelf, which was paved with mussels, and increased to 47% on the outer shelf at 100 m depth. Where the oxic–anoxic interface of the water column impinged on the sea floor at around 150 m depth, the contribution of sulfate reduction increased from >50% just above the chemocline to 100% just below. In the deep sea, mean sulfate reduction rates were 0.6 mmol m⁻² d⁻¹ corresponding to an organic carbon oxidation of 1.3 mmol m⁻² d⁻¹. This is close to the mean sedimentation rate of organic carbon over the year in the western basin. A comparison with published data on sulfate reduction in Black Sea sediments showed that the present results tend to be higher in shelf sediments and lower in the deep-sea than most other data. Based on the present water column H₂S inventory and the H₂S flux out of the sediment, the calculated turnover time of H₂S below the chemocline is 2100 years.     

Fate of biogenic carbon in the upper 200 m of the central Greenland Sea

Sedimentation of particulate carbon from the upper 200–300 m in the central Greenland Sea from August 1993 to June 1995 was less than 2 g C m⁻² yr⁻¹. Daily rates of sedimentation of particulate organic carbon reached highest values of about 18 mg m⁻² d⁻¹ in fall 1994. For total particulate material, maximum rates of sedimentation of about 250 mg m⁻² d⁻¹ were recorded in spring and fall 1994. For chlorophyll equivalent, highest rates of sedimentation of about 140 μg m⁻² d⁻¹ were recorded in spring 1994. As reported in related investigations, the transient accumulation of DOC in surface waters during summer, as well as respiration and mortality of deep overwintering zooplankton stocks, appeared to dominate the fate of photosynthetically fixed organic carbon. The above processes may account for roughly 43 g C m⁻² in the upper 200 m of the central Greenland Sea. For comparison, the seasonal deficit in dissolved inorganic carbon was reported to be about 23 g C m⁻² in the upper 20 m of surface water, and estimates for new annual production were reported to be about 57 g C m⁻². In our investigation, the biological carbon pump was not unusually effective in transporting carbon out of the productive surface layer.     

Dissolved organic carbon export and subsequent remineralization in the mesopelagic and bathypelagic realms of the North Atlantic basin

Dissolved organic carbon (DOC) data are presented from three meridional transects conducted in the North Atlantic as part of the US Climate Variability (CLIVAR) Repeat Hydrography program in 2003. The hydrographic sections covered a latitudinal range of 6°S to 63°N along longitudes 20°W (CLIVAR line A16), 52°W (A20) and 66°W (A22). Over 3700 individual measurements reveal unprecedented detail in the DOC distribution and systematic variations in the mesopelagic and bathypelagic zones of the North Atlantic basin. Latitudinal gradients in DOC concentrations combined with published estimates of ventilation rates for the main thermocline and North Atlantic Deep Water (NADW) indicate a net DOC export rate of 0.081 Pg C yr⁻¹ from the epipelagic zone into the mesopelagic and bathypelagic zones. Model II regression and multiple linear regression models applied to pairwise measures of DOC and chlorofluorocarbon (CFC-12) ventilation age, retrieved from major water masses within the main thermocline and NADW, indicate decay rates for exported DOC ranging from 0.13 to 0.94 μmol kg⁻¹ yr⁻¹, with higher DOC concentrations driving higher rates. The contribution of DOC oxidation to oxygen consumption ranged from 5 to 29% while mineralization of sinking biogenic particles drove the balance of the apparent oxygen utilization.