Microbial food web responses to light and nutrients beneath the coastal Arctic Ocean sea ice during the winter–spring transition

We measured the abundance and biomass of phototrophic and heterotrophic microbes in the upper mixed layer of the water column in ice-covered Franklin Bay, Beaufort Sea, Canada, from December 2003 to May 2004, and evaluated the influence of light and nutrients on these communities by way of a shipboard enrichment experiment. Bacterial cell concentrations showed no consistent trends throughout the sampling period, averaging (± SD) 2.4 (0.9) × 10⁸ cells L^− 1; integrated bacterial biomass for the upper mixed layer ranged from 1.33 mg C m^− 3 to 3.60 mg C m^− 3. Small cells numerically dominated the heterotrophic protist community in both winter and spring, but in terms of biomass, protists with a diameter > 10 µm generally dominated the standing stocks. Heterotrophic protist biomass integrated over the upper mixed layer ranged from 1.23 mg C m^− 3 to 6.56 mg C m^− 3. Phytoplankton biomass was low and variable, but persisted during the winter period. The standing stock of pigment-containing protists ranged from a minimum value of 0.38 mg C m^− 3 in winter to a maximal value of 6.09 mg C m^− 3 in spring and the most abundant taxa were Micromonas-like cells. These picoprasinophytes began to increase under the ice in February and their population size was positively correlated with surface irradiance. Despite the continuing presence of sea ice, phytoplankton biomass rose by more than an order of magnitude in the upper mixed layer by May. The shipboard experiment in April showed that this phototrophic increase in the community was not responsive to pulsed nutrient enrichment, with all treatments showing a strong growth response to improved irradiance conditions. Molecular (DGGE) and microscopic analyses indicated that most components of the eukaryotic community responded positively to the light treatment. These results show the persistence of a phototrophic inoculum throughout winter darkness, and the strong seasonal response by arctic microbial food webs to sub-ice irradiance in early spring.     

Microbial community structure in the marginal ice zone of the Bellingshausen Sea

Phytoplankton, bacteria and microzooplankton were investigated on a transect in the Bellingshausen Sea during the ice melt period in November–December 1992. The transect along the 85°W meridian comprised seven stations that progressed from solid pack-ice (70°S), through melting ice into open water (67°S). The abundance, biomass and taxonomic composition were determined for each component of the microbial community. The phytoplankton was mostly dominated by diatoms, particularly small (<20 μm) species. Diatom abundance ranged from 66 000 cells l⁻¹ under the ice to 410 000 cells l⁻¹ in open water. Phytoplankton biomass varied from <1 to 167 mg C m⁻³, with diatoms comprising 89–95% of the total biomass in open water and autotrophic nanoflagellates comprising 57% under the ice. The standing stocks of autotrophs in the mixed layer ranged from 95 mg C m⁻² under the pack-ice to 9478 mg C m⁻² in open waters. Bacterial abundance in ice-covered and open water stations varied from 1.1 to 5.5×10⁸ cells l⁻¹. Bacterial biomass ranged from 2.4 mg C m⁻³ under pack-ice to an average of 14 mg C m⁻³ in open water. The microzooplankton consisted mainly of aloricate oligotrich ciliates and heterotrophic dinoflagellates and these were most abundant in open waters. Their biomass varied between 0.2 and 54 mg C m⁻³ with a minimum at depth under the ice and maximum in open surface waters. Microheterotrophic standing stocks varied between 396 mg C m⁻² under pack-ice and 3677 mg C m⁻² in the open waters. The standing stocks of the total microbial community increased consistently from 491 mg C m⁻² at the ice station to 13 155 mg C m⁻² in open waters, reflecting the productive response of the community to ice-melt. The composition of the microbial community also shifted markedly from one dominated by heterotrophs (82% of microbial stocks) at the ice station to one dominated by autotrophs (73% of microbial stocks) in the open water. Our estimates suggest that the microbial community comprised >100% of the total particulate organic carbon (POC) under the ice and 62–66% of the measured POC in the open waters.     

Bacterial production and microbial food web structure in a large arctic river and the coastal Arctic Ocean

Globally significant quantities of organic carbon are stored in northern permafrost soils, but little is known about how this carbon is processed by microbial communities once it enters rivers and is transported to the coastal Arctic Ocean. As part of the Arctic River-Delta Experiment (ARDEX), we measured environmental and microbiological variables along a 300 km transect in the Mackenzie River and coastal Beaufort Sea, in July–August 2004. Surface bacterial concentrations averaged 6.7 × 10⁵ cells mL^− 1 with no significant differences between sampling zones. Picocyanobacteria were abundant in the river, and mostly observed as cell colonies. Their concentrations in the surface waters decreased across the salinity gradient, dropping from 51,000 (river) to 30 (sea) cells mL^− 1. There were accompanying shifts in protist community structure, from diatoms, cryptophytes, heterotrophic protists and chrysophytes in the river, to dinoflagellates, prymnesiophytes, chrysophytes, prasinophytes, diatoms and heterotrophic protists in the Beaufort Sea.Size-fractionated bacterial production, as measured by ³H–leucine uptake, varied from 76 to 416 ng C L^− 1 h^− 1. The contribution of particle-attached bacteria (> 3 µm fraction) to total bacterial production decreased from > 90% at the Mackenzie River stations to < 20% at an offshore marine site, and the relative importance of this particle-based fraction was inversely correlated with salinity and positively correlated with particulate organic carbon concentrations. Glucose enrichment experiments indicated that bacterial metabolism was carbon limited in the Mackenzie River but not in the coastal ocean. Prior exposure of water samples to full sunlight increased the biolability of dissolved organic carbon (DOC) in the Mackenzie River but decreased it in the Beaufort Sea.Estimated depth-integrated bacterial respiration rates in the Mackenzie River were higher than depth-integrated primary production rates, while at the marine stations bacterial respiration rates were near or below the integrated primary production rates. Consistent with these results, P_CO2 measurements showed surface water supersaturation in the river (mean of 146% of air equilibrium values) and subsaturation or near-saturation in the coastal sea. These results show a well-developed microbial food web in the Mackenzie River system that will likely convert tundra carbon to atmospheric CO₂ at increasing rates as the arctic climate continues to warm.     

Variations of the abundance and nucleic acid content of heterotrophic bacteria in Beaufort Shelf waters during winter and spring

Depth profiles of heterotrophic bacteria abundance were measured weekly over a 6-month period from December to May in Franklin Bay, a 230 m-deep coastal Arctic Ocean site of the southeastern Beaufort Sea. Total bacteria, low nucleic acid (LNA) and high nucleic acid (HNA) bacteria abundances were measured using flow cytometry after SYBR Green I staining. The HNA bacteria abundance in surface waters started to increase 5–6 weeks after phytoplankton growth resumed in spring, increasing from 1 × 10⁵ to 3 × 10⁵ cells mL^− 1 over an 8-week period, with a net growth rate of 0.018 d^− 1. LNA bacteria response was delayed by more than two months relative to the beginning of the phytoplankton biomass accumulation and had a lower net growth rate of 0.013 d^− 1. The marked increase in bacterial abundance occurred before any significant increase in organic matter input from river discharge (as indicated by the unchanged surface water salinity and DOC concentrations), and in the absence of water temperature increase. The abundance of bacteria below the halocline was relatively high until January (up to 5 × 10⁵ cells mL^− 1) but then decreased to values close to 2 × 10⁵ cells mL^− 1. The three-fold bacterial abundance increase observed in surface waters in spring was mostly due to HNA bacteria, supporting the idea that these cells are the most active.     

Picophytoplankton and nanophytoplankton abundance and distribution in the southeastern Beaufort Sea (Mackenzie Shelf and Amundsen Gulf) during Fall 2002

The distribution of picophytoplankton (0.2–2 µm) and nanophytoplankton (2–20 µm) in the Beaufort Sea–Mackenzie Shelf and Amundsen Gulf regions during autumn, 2002 is examined relative to their ambient water mass properties (salinity, temperature and nutrients: nitrate + nitrite, phosphate, and silicate) and to the ratio of variable to maximum fluorescence, Fv/Fm. Total phytoplankton and cell abundances (< 20 µm) were mainly correlated with salinity. Significant differences in picophytoplankton cell numbers were found among waters near the mouth of the Mackenzie River, ice melt waters and the underlying halocline water masses of Pacific origin. Picophytoplankton was the most abundant phytoplankton fraction during the autumnal season, probably reflecting low nitrate concentrations (surface waters average ~ 0.65 µM). The ratio Fv/Fm averaged 0.44, indicating that cells were still physiologically active, even though their concentrations were low (max Chl a = 0.9 mg m^− 3). No significant differences in Fv/Fm were evident in the different water masses, indicating that rate limiting conditions for photosynthesis and growth were uniform across the whole system, which was in a pre-winter stage, and was probably already experiencing light limitation as a result of shortening day lengths.     

Seasonality of picophytoplankton chlorophyll a and biomass in the central Cantabrian Sea, southern Bay of Biscay

Seasonal changes in the abundance and biomass of cyanobacteria (Synechococcus and Prochlorococcus) and picoeukaryotes were studied by flow cytometry in the upper layers of the central Cantabrian Sea continental shelf, from April 2002 to April 2006. The study area displayed the typical hydrographic conditions of temperate coastal zones. A marked seasonality of the relative contribution of prokaryotes and eukaryotes was found. While cyanobacteria were generally more abundant for most of the year (up to 2.4 10⁵ cells mL^− 1), picoeukaryotes dominated the community (up to 10⁴ cells mL^− 1) from February to May. The disappearance of Prochlorococcus from spring through summer is likely related to shifts in the prevailing current regime. The maximum total abundance of picophytoplankton was consistently found in late summer–early autumn. Mean photic-layer picoplanktonic chlorophyll a ranged from 0.06 to 0.53 µg L^− 1 with a relatively high mean contribution to total values (33 ± 2% SE), showing maxima around autumn and minima in spring. Biomass (range 0.58–40.16 mg C m^− 3) was generally dominated by picoeukaryotes (mean ± SE, 4.28 ± 0.27 mg C m^− 3) with an average contribution of cyanobacteria of 30 ± 2%. Different seasonality of pigment and biomass values resulted in a clear temporal pattern of picophytoplanktonic carbon to chlorophyll a ratio, which ranged from 10 (winter) to 140 (summer). This study highlights the important contribution of picoplanktonic chlorophyll a and carbon biomass in this coastal ecosystem.     

Atmospheric CO2 measurements and error analysis on seasonal air–sea CO2 fluxes in the Bay of Biscay

Atmospheric molar fraction of CO₂ (xCO₂^atm) measurements obtained on board of ships of opportunity are used to parameterize the seasonal cycle of atmospheric xCO₂ (xCO₂^atm) in three regions of the eastern North Atlantic (Galician and French offshore and Bay of Biscay). Three selection criteria are established to eliminate spurious values and identify xCO₂^atm data representative of atmospheric background values. The filtered data set is fitted to seasonal curve, consisting of an annual trend plus a seasonal cycle. Although the fitted curves are consistent with the seasonal evolution of xCO₂^atm data series from land meteorological stations, only ship-board measurements can report the presence of winter xCO₂^atm minimum on Bay of Biscay. Weekly air–sea CO₂ flux differences (mmol C·m^− 2 day^− 1) produced by the several options of xCO₂^atm usually used (ship-board measurements, data from land meteorological stations and annually averaged values) were calculated in Bay of Biscay throughout 2003. Flux error using fitted seasonal curve relative to on board measurements was minimal, whereas land stations and annual means yielded random (− 0.2 ± 0.3 mmol C·m^− 2·day^− 1) and systematic (− 0.1 ± 0.4 mmol C·m^− 2 day^− 1), respectively. The effect of different available sources of sea level pressure, wind speed and transfer velocity were also evaluated. Wind speed and transfer velocity parameters are found as the most critical choice in the estimate of CO₂ fluxes reaching a flux uncertainty of 7 mmol C·m^− 2·day^− 1 during springtime. The atmospheric pressure shows a notable relative effect during summertime although its influence is quantitatively slight on annual scale (0.3 ± 0.2 mmol C·m^− 2·day^− 1). All results confirms the role of the Bay of Biscay as CO₂ sink for the 2003 with an annual mean CO₂ flux around − 5 ± 5 mmol C m^− 2 day^− 1.     

Effect of wave boundary layer on sea-to-air dimethylsulfide transfer velocity during typhoon passage

A full-spectral third-generation ocean wind–wave model (Wavewatch-III) implemented in the South China Sea is used to investigate the effects of the wave boundary layer on the drag coefficient and the sea-to-air transfer velocity of dimethylsulfide (DMS) during passage of Typhoon Wukong (September 5–11, 2000) with a maximum sustained wind speed of 38 m s^− 1. The model is driven by the reanalyzed surface winds (1° × 1°, four times daily) from the National Centers for Environmental Prediction. It is found that the wave boundary layer evidently enhances (16.5%) the drag coefficient (in turn increases the momentum flux across the air–sea interface), and reduces (13.1%) the sea-to-air DMS transfer velocity (in turn decreases the sea-to-air DMS flux). This indicates the possibility of important roles of wave boundary layer in atmospheric DMS contents and global climate system.     

Diatom stratigraphy and long-term dissolved silica concentrations in the Baltic Sea

In many parts of the world coastal waters with anthropogenic eutrophication have experienced a gradual depletion of dissolved silica (DSi) stocks. This could put pressure on spring bloom diatom populations, e.g. by limiting the intensity of blooms or by causing shifts in species composition. In addition, eutrophication driven enhanced diatom growth is responsible for the redistribution of DSi from the water phase to the sediments, and changes in the growth conditions may be reflected in the sediment diatom stratigraphy.To test for changes in diatom communities we have analyzed four sediment cores from the Baltic Sea covering approximately the last 100 years. The sediment cores originate from the western Gulf of Finland, the Kattegat, the Baltic Proper and the Gulf of Riga. Three out of the four cores reveal only minor changes in composition of diatom assemblages, while the Gulf of Riga core contains major changes, occurring after the second World War. This area is set apart from the other Baltic Sea basins by a high frequency of low after spring bloom DSi concentrations (< 2 µmol L^− 1) during a relatively well defined time period from 1991–1998. In 1991 to 1993 a rapid decline of DSi spring concentrations and winter stocks (down to 5 µmol L^− 1) in the Gulf was preceded by exceptionally intense diatom spring blooms dominated by the heavily silicified species Thalassiosira baltica (1991–1992; up to 5.5 mg ww L^− 1). T. baltica has been the principal spring bloom diatom in the Gulf of Riga since records began in 1975. DSi consumption and biomass yield experiments with cultured T. baltica suggest that intense blooms can potentially exhaust the DSi stock of the water column and exceed the annual Si dissolution in the Gulf of Riga. The phytoplankton time series reveals another exceptional T. baltica bloom period in 1981–1983 (up to 8 mg L^− 1), which, however, took place before the regular DSi measurements. These periods may be reflected in the conspicuous accumulation of T. baltica frustules in the sediment core corresponding to ca. 1975–1985.     

On the distribution of dissolved lead in the Loire estuary and the North Biscay continental shelf, France

The dissolved lead was studied in the whole salinity gradient of the system composed of the Loire estuary and the North Biscay continental shelf. About 130 samples were collected in winter 2001 and spring 2002 during Nutrigas and Gasprod campaigns (Programme PNEC-Golfe de Gascogne, RV Thalassa) and metal measurements were conducted on board by Potentiometric Stripping Analysis. In the Loire estuary, levels of dissolved lead ranged from 0.15 to 0.24 nM and from 0.04 to 0.26 nM in winter and spring, respectively. Compared to the concentrations reported in 1987 and 1990 (0.4–1.7 nM; Boutier, B., Chiffoleau, J.F., Auger, D., Truquet, I., 1993. Influence of the Loire river on dissolved lead and cadmium concentrations in coastal waters of Brittany. Estuar. Coast. Shelf S., 36:133–143, Estuarine, Coastal and Shelf Science 36, 133–143) our study indicated much lower values. The fall in concentration in the estuary could be attributed to the stopping of activity of Octel, a big manufacturer of tetra alkyl lead. Discharge in dissolved metal to the continental shelf by the Loire river was assessed as 7.5 and 1.9 kg day^− 1 for winter and spring, respectively. On the continental shelf, levels of dissolved lead varied within 0.06 and 0.27 nM in winter (0.15 ± 0.06 nM, sd = 1.96, n = 49), whereas concentrations measured in spring were in the range 0.06–0.17 nM (0.09 ± 0.03 nM, sd = 1.96, n = 60). This difference in metal concentration was related to the amounts of rainfall that have fallen over the continental shelf: estimations of inputs by this way (74 and 32 kg day^− 1 in winter and spring, respectively) appeared to be significantly higher than inputs from the Loire river (7.5 and 1.9 kg day^− 1 in winter and spring, respectively). The distributions of dissolved metal in the surface waters highlighted the role of suspended particular matter (SPM) for a rapid “trapping” of lead near the mouth of the estuary. The vertical distributions showed, in the stratified area, a biological transfer of lead between winter and spring from surface waters to the halocline.     

Physical and biological correlates of virus dynamics in the southern Beaufort Sea and Amundsen Gulf

As part of the Canadian Arctic Shelf Exchange Study (CASES), we investigated the spatial and seasonal distributions of viruses in relation to biotic (bacteria, chlorophyll-a (chl a)) and abiotic variables (temperature, salinity and depth). Sampling occurred in the southern Beaufort Sea Shelf in the region of the Amundsen Gulf and Mackenzie Shelf, between November 2003 and August 2004. Bacterial and viral abundances estimated by epifluorescence microscopy (EFM) and flow cytometry (FC) were highly correlated (r² = 0.89 and r² = 0.87, respectively), although estimates by EFM were slightly higher (FC = 1.08 × EFM + 0.12 and FC = 1.07 × EFM + 0.43, respectively). Viral abundances ranged from 0.13 × 10⁶ to 23 × 10⁶ ml^− 1, and in surface waters were ~ 2-fold higher during the spring bloom in May and June and ~ 1.5-fold higher during July and August, relative to winter abundances. These increases were coincident with a ~ 6-fold increase in chl a during spring and a ~ 4-fold increase in bacteria during summer. Surface viral abundances near the Mackenzie River were ~ 2-fold higher than in the Mackenzie Shelf and Amundsen Gulf regions during the peak summer discharge, concomitant with a ~ 5.5-fold increase in chl a (up to 2.4 μg l^− 1) and a ~ 2-fold increase in bacterial abundance (up to 22 × 10⁵ ml^− 1). Using FC, two subgroups of viruses and heterotrophic bacteria were defined. A low SYBR-green fluorescence virus subgroup (V2) representing ~ 71% of the total viral abundance, was linked to the abundance of high nucleic acid fluorescence (HNA) bacteria (a proxy for bacterial activity), which represented 42 to 72% of the bacteria in surface layers. A high SYBR-green fluorescence viral subgroup (V1) was more related to high chl a concentrations that occurred in surface waters during spring and at stations near the Mackenzie River plume during the summer discharge. These results suggest that V1 infect phytoplankton, while most V2 are bacteriophages. On the Beaufort Sea shelf, viral abundance displayed seasonal and spatial variations in conjunction with chl a concentration, bacterial abundance and composition, temperature, salinity and depth. The highly dynamic nature of viral abundance and its correlation with increases in chl a concentration and bacterial abundance implies that viruses are important agents of microbial mortality in Arctic shelf waters.     

Spatial heterogeneity and temporal dynamics of particles, bacteria, and pEPS in Arctic winter sea ice

Abundances of particles, total bacteria, and particulate extracellular polymeric substances (pEPS) in Arctic sea ice were tracked through a winter season to examine the impact of combined extremes of low temperature and high salinity on the prokaryotic microbial community. Three horizons, centered at depths of 25, 45, and 65 cm from the ice surface, with mean seasonal temperatures of − 20, − 17, and − 13 °C, respectively, were sampled 16 times over the course of 12 weeks. Microscopic counts of bacteria (stained with DAPI) and particles (stained with acridine orange) reflected the dynamic conditions of the growing ice sheet, with greater abundances and variability in the upper ice horizons compared to the lower. The trend of higher particle and bacterial abundances in the upper ice was corroborated by several full-depth profiles taken during the expedition, which also displayed significantly decreasing cell abundance with depth. Bacterial abundance declined slowly and significantly with time in the upper and middle ice horizons, but not in the lowest, suggesting that much of the prokaryotic microbial community is resilient to extreme environmental conditions. We found that pEPS concentrations increased significantly with time and with decreasing temperatures in all depth horizons, which may lend support to the argument that sea ice bacteria produce EPS in situ as a cryoprotectant.     

Pore water phosphate and ammonia below the permanent halocline in the south-eastern Baltic Sea and their benthic fluxes under anoxic conditions

In this paper the results of a study on the distribution of pore water phosphates and ammonia, and their fluxes under anoxic condition in a deep (> 70 m) accumulation-type bottom of the south-eastern Baltic Sea, namely in the Gdańsk Deep and the adjacent areas, are presented. All measurements were taken during the growth period, i.e. in September 2000, April 2001 and June 2002. Benthic phosphate and ammonia fluxes were estimated using Fick's First Law. Phosphate and ammonia concentrations ranged from 7.5 to 266.3 μmol dm^− 3 and from 53.6 to 1248.3 μmol dm^− 3, respectively. The values recorded in the central part of the Gdańsk Deep were lower than those found both on its slopes and on the SW slope of the Gotland Deep. The lowest phosphate contents were typical of the Oblique Sill which separates the Gdańsk and Gotland Deeps.In 1993–2002, as a result of anoxia the sediments in the Gdańsk Deep released about 5.1 × 10³ t P and 22.8 × 10³ t N. These loads supplied on average 1.5% and 0.9% of phytoplankton's demand for P and N, respectively. In comparison to the total external load of nutrients discharged to the Gulf of Gdańsk (i.e. 8.79 × 10³ t year^− 1 P_tot and 130.79 × 10³ t year^− 1 N_tot; [Witek, Z., Humborg, Ch., Savchuk, O., Grelowski, A. and Łysiak-Pastuszak, E., 2003. Nitrogen and phosphorus budgets of the Gulf of Gdańsk (Baltic Sea). Est. Coast. Shelf Sci., 57:239–248.]), the return flux of P and N from the anoxic sediments to the water column in the Gdańsk Deep was a minor source of these elements.     

The biogeochemical cycling of methane in Ria de Vigo, NW Spain: Sediment processing and sea–air exchange

Methane (CH₄) concentrations were measured in the water column, in sediment porewaters, and in atmospheric air, in the Ría de Vigo, NW Spain, during both the onset (April 2003) and at the end of (September 2004) seasonal upwelling. In addition, CH₄ concentration and stable isotopic signatures (δ¹³CH₄) were measured in porewaters, and sediment methanogenesis and aerobic oxidation of CH₄ were determined in sediment incubations. Surface water column CH₄ (2 m depth) was in the range 3–180 nmol l^− 1 (110–8500% saturation) and followed a generally landward increase but with localised maxima in both the inner and middle Ría. These maxima were consistent with CH₄ inputs from underlying porewaters in which CH₄ concentrations were up to 3 orders of magnitude higher (maximum 350 μmol l^− 1). Surface water CH₄ concentrations were approximately three times higher in September than in April, consistent with a significant benthic CH₄ flux driven by enhanced sediment methanogenesis following the summer productivity maximum. CH₄ and δ¹³CH₄ in sediment porewaters and in incubated sediment slurries (20 °C) revealed significant sediment CH₄ oxidation, with an apparent isotopic fractionation factor (r_c) of  1.004. Using turbulent diffusion models of air–sea exchange we estimate an annual emission of atmospheric CH₄ from the Ría de Vigo of 18–44 × 10⁶ g (1.1–2.7 × 10⁶ mol). This estimate is approximately 1–2 orders of magnitude lower than a previous estimate based on a bubble transport model.     

Th-derived particulate organic carbon fluxes in the northern Barents Sea with comparison to drifting sediment trap fluxes

Large-volume sampling of ²³⁴Th was conducted to estimate particulate organic carbon (POC) export in conjunction with drifting sediment trap deployments in the northern Barents Sea in July 2003 and May 2005. ²³⁴Th-derived POC fluxes averaged 42.3 ± 39.7 mmol C m^− 2 d^− 1 in 2003 and 47.1 ± 30.6 mmol C m^− 2 d^− 1 in 2005. Sediment trap POC fluxes averaged 13.1 ± 8.2 mmol C m^− 2 d^− 1 in 2003 and 17.3 ± 11.4 mmol C m^− 2 d^− 1 in 2005, but better reflected the transient bloom conditions that were observed at each station within a season. Although ²³⁴Th fluxes agreed within a factor 2 at most stations and depths sampled, sediment trap POC fluxes were lower than large-volume POC flux estimates at almost every station. This may represent an under-collection of POC by the drifting sediment traps or, conversely, an over-collection of POC by the large-volume sampling of ²³⁴Th. It is hypothesized that the offset between the two methods is partly due to the presence of the prymnesiophyte Phaeocystis pouchetii, which potentially causes a large variation in > 53-μm POC/²³⁴Th ratios. Due to the large proportion of dissolved carbon or mucilage released by P. pouchetii, and because it is thought that P. pouchetii does not contribute significantly to the vertical export of biogenic matter in the Barents Sea, the application of large-volume sampling of ²³⁴Th may yield relatively high, and possibly inaccurate POC/²³⁴Th ratios. Hence, POC fluxes derived from ²³⁴Th sampling may be inappropriate and drifting sediment traps might be a more reliable method to measure the vertical export of biogenic matter in regions that have recurrent P. pouchetii blooms, such as the Barents Sea.     

Suspended sediment and erosion dynamics in Kugmallit Bay and Beaufort Sea during ice-free conditions

The Mackenzie River is the largest river on the North American side of the Arctic and its huge freshwater and sediment load impacts the Canadian Beaufort Shelf. Huge quantities of sediment and associated organic carbon are transported in the Mackenzie plume into the interior of the Arctic Ocean mainly during the freshet (May to September). Changing climate scenarios portend increased coastal erosion and resuspension that lead to altered river-shelf-slope particle budgets. We measured sedimentation rates, suspended particulate matter (SPM), particle size and settling rates during ice-free conditions in Kugmallit Bay (3–5 m depth). Additionally, measurements of erosion rate, critical shear stress, particle size distribution and resuspension threshold of bottom sediments were examined at four regionally contrasting sites (33–523 m depth) on the Canadian Beaufort Shelf using a new method for assessing sediment erosion. Wind induced resuspension was evidenced by a strong relationship between SPM and wind speed in Kugmallit Bay. Deployment of sediment traps showed decreasing sedimentation rates at sites along an inshore–offshore transect ranging from 5400 to 3700 g m^− 2 day^− 1. Particle settling rates and size distributions measured using a Perspex settling chamber showed strong relationships between equivalent spherical diameter (ESD) and particle settling rates (r² = 0.91). Mean settling rates were 0.72 cm s^− 1 with corresponding ESD values of 0.9 mm. Undisturbed sediment cores were exposed to shear stress in an attempt to compare differences in sediment stability across the shelf during September to October 2003. Shear was generated by vertically oscillating a perforated disc at controlled frequencies corresponding to calibrated shear velocity using a piston grid erosion device. Critical (Type I) erosion thresholds (u) varied between 1.1 and 1.3 cm s^− 1 with no obvious differences in location. Sediments at the deepest site Amundsen Gulf displayed the highest erosion rates (22–54 g m^− 2 min^− 1) with resuspended particle sizes ranging from 100 to 930 µm for all sites. There was no indication of biotic influence on sediment stability, although our cores did not display a fluff layer of unconsolidated sediment. Concurrent studies in the delta and shelf region suggest the importance of a nepheloid layer which transports suspended particles to the slope. Continuous cycles of resuspension, deposition, and horizontal advection may intensify with reduction of sea ice in this region. Our measurements coupled with studies of circulation and cross-shelf exchange allow parameterization and modeling of particle dynamics and carbon fluxes under various climate change scenarios.     

Turbulent mixing and internal tides in Gaoping (Kaoping) Submarine Canyon, Taiwan

Turbulent overturning on scales greater than 10 m is observed near the bottom and in mid-depth layers within the Gaoping (formerly spelled Kaoping) Submarine Canyon (KPSC) in southern Taiwan. Bursts of strong turbulence coexist with bursts of strong sediment concentrations in mid-depth layers. The turbulence kinetic energy dissipation rate in some turbulence bursts exceeds 10^− 4 W kg^− 1, and the eddy diffusivity exceeds 10^− 1 m² s^− 1. Within the canyon, the depth averaged turbulence kinetic energy dissipation rate is ~ 7 × 10^− 6 W kg^− 1, and the depth averaged eddy diffusivity is ~ 10^− 2 m² s^− 1. These are more than two orders of magnitude greater than typical values in the open ocean, and are much larger than those found in the Monterey Canyon where the strong turbulent mixing has also been. The interaction of tidal currents with the complex topography in Gaoping Submarine Canyon is presumably responsible for the observed turbulent overturning via shear instability and the breaking of internal tides and internal waves at critical frequencies. Strong 1st-mode internal tides exist in KPSC. The depth averaged internal tidal energy near the canyon mouth is ~ 0.17 m² s^− 2. The depth integrated internal tidal energy flux at the mouth of the canyon is ~ 14 kW m^− 1, propagating along the axis of the canyon toward the canyon head. The internal tidal energy flux in the canyon is 3–7 times greater than that found in Monterey Canyon, presumably due to the more than 10 times larger barotropic tide in the canyon. Simple energy budget calculations conclude that internal tides alone may provide energy sufficient to explain the turbulent mixing estimated within the canyon. Further experiments are needed in order to quantify the seasonal and geographical distributions of internal tides in Gaoping Submarine Canyon and their effects on the sediment flux in the canyon.     

Appendicularian ecophysiology I: Food concentration dependent clearance rate, assimilation efficiency, growth and reproduction of Oikopleura dioica

Three aspects of the appendicularian O. dioica's ecophysiology were measured here: 1) morphological parameters over a wide range of appendicularian sizes, including mature animals in order to document the morphological characteristics inducing reproduction; 2) clearance rate and assimilation efficiency using feeding incubations with different algal concentrations and 3) the effect of food concentration on growth, mortality and reproduction.The relationship between the body carbon weight and the clearance rate follows a power function, with an exponent of 0.91 (± 0.07). The rate of particles retention increases with the food concentration following a Michaelis–Menten relationship (half-saturation constant = 151 ± 22 µg C l^− 1, maximum clearance rate = 12 ± 1 µg C µg C^− 1 d^− 1). The carbon assimilation efficiency decreases with the increasing food concentration. As a result, appendicularian growth which is limited in concentrations lower than 50 µg C l^− 1 is saturated above 100 µg C l^− 1.In immature animals the gonad represents less than 30% of the body volume whereas in mature individuals, its volume varies between 50% and 87% (mean 63%) suggesting that gonad/total volume ratio can be used as indicator of the maturation stages. The gonad weight in mature animals represents 70.3 (± 4.6)% of the total body carbon weight. Two major maturity stages can explain the changes in energy allocation: i) the somatic growth, when less energy is invested in gonad growth when compared to the rest of the body and ii) the maturation phase where most of the assimilated matter is invested in gonad maturation. This process is rapid, lasting only few hours. For this reason we measured completely mature organisms that are generally not measured during the experimental work with appendicularians. In food-limited conditions, the gonad maturation process starts with smaller individuals and ends with smaller reproductive animals having the same gonad to total volume ratio than in unlimited food conditions. The results obtained in this study were used to model the life cycle of O. dioica (see Lombard, F., Sciandra, A. and Gorsky, G., 2009-this volume. Appendicularian ecophysiology. II. Modeling nutrition, metabolism, growth and reproduction of the appendicularian Oikopleura dioica.).     

Spatial and temporal variability of size-fractionated biomass and primary production in the Ross Sea (Antarctica) during austral spring and summer

The results of a study on the spatial and temporal dynamics of size-fractionated biomass and production of phytoplankton in the Ross Sea during the austral spring and summer are reported. The spring cruise took place in the offshore Ross Sea from 14 November to 14 December 1994. Sampling was carried out on a transect of 27 stations distributed from 76.5 to 72.0°S along 175°E, and covered the three main Antarctic environments of the polynya open waters, the marginal ice zone and the pack ice area. Three subsystems were identified. The subsystem of the polynya was characterised by the predominance of the micro- and nano-planktonic fractions, chlorophyll (Chl a) concentrations from 69.6 to 164.7 mg m⁻² and production rates from 0.68 to 1.14 g C m⁻² day⁻¹. The second subsystem, the marginal ice zone, showed a relative increase of the micro-planktonic fraction, high biomass levels (from 99.64 to 220 mg Chl m⁻²) and production rates from 0.99 to 2.7 g C m⁻² day⁻¹. The subsystem of the pack ice area had a phytoplankton community dominated by the pico-planktonic fraction and showed low biomasses (from 19.4 to 37.7 mg Chl m⁻²) and production rates (0.28 to 0.60 g C m⁻² day⁻¹). Selective grazing by krill is considered an important factor in determining the size structure of the phytoplankton communities. The summer study consisted of a time series carried out in inshore waters of Terra Nova Bay from 12 January to 8 February 1990. In a well stabilised water column and with high levels of PAR always available, the primary production rates of a community dominated by micro-plankton varied from 0.52 to 1.2 g C m⁻² day⁻¹ (average 0.84). A high P/B ratio, up to 3, and a remarkably elevated mean phaeopigment (Phaeo)/Chl a ratio of 2.4 indicated an active removal of biomass by grazing, confirmed by the presence of faecal pellets in quantities reaching 6000 m⁻³ in the upper 50 m. The peculiarities of the inshore versus offshore environments in terms of community size structure, production processes and their implications as regards the food web are discussed.     

The use of polycyclic aromatic hydrocarbons as a particulate tracer in the water column of Gaoping (Kaoping) Submarine Canyon

Time-series samples of settling particles were collected in the water column of Gaoping (formerly spelled Kaoping) Submarine Canyon (KPSC) with two sediment traps on taut-line moorings deployed at two different depths (60 and 280 m) between May 26 and June 27, 2004. Average total polycyclic aromatic hydrocarbon (PAH) concentrations of upper and lower trap array samples were 310 ± 61 ng g^− 1 dw (range: 200–440) and 240 ± 36 ng g^− 1 dw (range: 180–290), respectively. Principal component analysis results suggest that PAH sources in the trap-collected particles included diesel vehicle/coal burning, diagenetic sources, and petroleum release. PAH downward fluxes based on settling particles were estimated to be 12–44 μg m^− 2 d^− 1. These values are higher than those reported in the literature for most coastal areas. During the sampling period, both traps were significantly tilted by tidal current and fluctuated vertically. The upper traps experienced greater vertical movements, thus their particle characteristics (e.g., POC, particle mass, and fine particle fraction) varied more than those of the lower traps. Hourly depth variations of the tilted sediment trap array were echoed by the corresponding total PAH concentrations. Moreover, the PAH composition of the collected particles was related to the flow direction and speed. These observations suggest that PAHs can be used as an effective chemical tracer for the transport of terrestrial and marine particulates in a complex aquatic environment like Gaoping (Kaoping) Submarine Canyon.