Influence of the Mackenzie River plume on the sinking export of particulate material on the shelf

We examined the influence of the Mackenzie River plume on sinking fluxes of particulate organic and inorganic material on the Mackenzie Shelf, Canadian Arctic. Short-term particle interceptor traps were deployed under the halocline at 3 stations across the shelf during fall 2002 and at 3 stations along the shelf edge during summer 2004. During the two sampling periods, the horizontal patterns in sinking fluxes of particulate organic carbon (POC) and chlorophyll a (chl a) paralleled those in chl a biomass within the plume. Highest sinking fluxes of particulate organic material occurred at stations strongly influenced by the river plume (maximum POC sinking fluxes at 25 m of 98 mg C m^− 2 d^− 1 and 197 mg C m^− 2 d^− 1 in 2002 and 2004, respectively). The biogeochemical composition of the sinking material varied seasonally with phytoplankton and fecal pellets contributing considerably to the sinking flux in summer, while amorphous detritus dominated in the fall. Also, the sinking phytoplankton assemblage showed a seasonal succession from a dominance of diatoms in summer to flagellates and dinoflagellates in the fall. The presence of the freshwater diatom Eunotia sp. in the sinking assemblage directly underneath the river plume indicates the contribution of a phytoplankton community carried by the plume to the sinking export of organic material. Yet, increasing chl a and BioSi sinking fluxes with depth indicated an export of phytoplankton from the water column below the river plume during summer and fall. Grazing activity, mostly by copepods, and to a lesser extent by appendicularians, appeared to occur in a well-defined stratum underneath the river plume, particularly during summer. These results show that the Mackenzie River influences the magnitude and composition of the sinking material on the shelf in summer and fall, but does not constitute the only source of material sinking to depth at stations influenced by the river plume.     

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.     

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.     

Sea ice and the onshore–offshore gradient in pre-winter zooplankton assemblages in southeastern Beaufort Sea

Zooplankton communities were studied in southeastern Beaufort Sea (Arctic Ocean) in September–October 2002. Cluster analysis and non-metric multidimensional scaling revealed three distinct mesozooplankton assemblages. A neritic assemblage occurred on the Mackenzie Shelf and in Franklin Bay, while distinct off-shelf assemblages prevailed in the Cape Bathurst Polynya and on the Beaufort Slope respectively. Over 95% of the mesozooplankton was comprised of eight copepod taxa. Pseudocalanus spp. contributed predominantly to the discrimination of the three assemblages and was the only significant indicator of the Shelf assemblage. Oithona similis, Oncaea borealis, Metridia longa and Calanus hyperboreus were indicators of the Polynya assemblage. Cyclopina sp. and Microcalanus pygmaeus were indicative of the overall off-shelf community (Polynya and Slope assemblages). The importance of omnivores and carnivores increased from the shelf to the polynya and the slope. Station depth and duration of reduced ice conditions during summer (< 50% ice concentration) underpinned the distribution of the assemblages (r² = 0.71 and 0.45 respectively). The abundance of Pseudocalanus spp. was independent of depth and increased with the duration of reduced ice conditions (r_s = 0.438). The abundance of Cyclopina sp., M. pygmaeus and other indicators of the offshore assemblages followed the opposite trend (r_s = − 0.467 and − 0.5 respectively). Under continued climate warming, a reduction of the ice cover will affect the biogeography of mesozooplankton on and around the Mackenzie Shelf, to the potential advantage of Pseudocalanus spp. and other calanoid herbivores.     

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.     

Winter–spring dynamics in sea-ice carbon cycling in the coastal Arctic Ocean

An understanding of microbial interactions in first-year sea ice on Arctic shelves is essential for identifying potential responses of the Arctic Ocean carbon cycle to changing sea-ice conditions. This study assessed dissolved and particulate organic carbon (DOC, POC), exopolymeric substances (EPS), chlorophyll a, bacteria and protists, in a seasonal (24 February to 20 June 2004) investigation of first-year sea ice and associated surface waters on the Mackenzie Shelf. The dynamics of and relationships between different sea-ice carbon pools were investigated for the periods prior to, during and following the sea-ice-algal bloom, under high and low snow cover. A predominantly heterotrophic sea-ice community was observed prior to the ice-algal bloom under high snow cover only. However, the heterotrophic community persisted throughout the study with bacteria accounting for, on average, 44% of the non-diatom particulate carbon biomass overall the study period. There was an extensive accumulation of sea-ice organic carbon following the onset of the ice-algal bloom, with diatoms driving seasonal and spatial trends in particulate sea-ice biomass. DOC and EPS were also significant sea-ice carbon contributors such that sea-ice DOC concentrations were higher than, or equivalent to, sea-ice-algal carbon concentrations prior to and following the algal bloom, respectively. Sea-ice-algal carbon, DOC and EPS-carbon concentrations were significantly interrelated under high and low snow cover during the algal bloom (r values ≥ 0.74, p < 0.01). These relationships suggest that algae are primarily responsible for the large pools of DOC and EPS-carbon and that similar stressors and/or processes could be involved in regulating their release. This study demonstrates that DOC can play a major role in organic carbon cycling on Arctic shelves.     

Distribution of dinoflagellate cysts in surface sediments of the Mackenzie Shelf and Amundsen Gulf, Beaufort Sea (Canada)

In order to document long-term climate cycles and predict future climate trends for the Arctic, we need to look at the geological records to establish the link between historical and pre-historical sea-surface parameters. Dinoflagellate cysts (dinocysts) are used as proxy indicators of sea-surface parameters (temperature, salinity, sea-ice cover, primary productivity) jointly with transfer functions and a modern dinocyst reference database, to reconstruct the evolution of sea-surface conditions at decadal and millennial timescales. Here we present the surface distribution of recent dinocyst assemblages from 34 surface sediment samples collected on the Mackenzie Slope/Amundsen Gulf during the 2004 CASES (Canadian Arctic Shelf Exchange Study) cruise. Dinocyst concentrations in surface sediments are relatively high outside the Mackenzie plume area and increase gradually eastward toward Amundsen Gulf. The cysts of autotrophic dinoflagellates are dominant throughout the study area, while the maximum abundance of heterotrophic taxa is found within the Mackenzie plume. Hierarchical clustering analyses allowed defining two dinocyst assemblages. Assemblage I is located on the Mackenzie Slope and southern Amundsen Gulf, while Assemblage II is located within the Cape Bathurst Polynya area in northern Amundsen Gulf. Both assemblages are dominated by Operculodinium centrocarpum, but are distinguished on the basis of the relative abundance of Islandinium minutum, a taxon generally associated with sea ice. I. minutum is found in lower abundance in the Cape Bathurst Polynya.     

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.     

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.     

Estimation of air–sea CO2 fluxes in the Bay of Biscay based on empirical relationships and remotely sensed observations

An empirical algorithm has been developed to compute the sea surface CO₂ fugacity (fCO₂^sw) in the Bay of Biscay from remotely sensed sea surface temperature (SST_RS) and chlorophyll a (chl a_RS) retrieved from AVHRR and SeaWiFS sensors, respectively. Underway fCO₂^sw measurements recorded during 2003 were correlated with SST_RS and chl a_RS data yielding a regression error of 0.1 ± 7.5 µatm (mean ± standard deviation). The spatial and temporal variability of air–sea fCO₂ gradient (ΔfCO₂) and air–sea CO₂ flux (FCO₂) was analyzed using remotely sensed images from September 1997 to December 2004. An average FCO₂ of ? 1.9 ± 0.1 mol m^? 2 yr^? 1 characterized the Bay of Biscay as a CO₂ sink that is suffering a significant long-term decrease of 0.08 ± 0.05 mol m^? 2 yr^? 2 in its capacity to store atmospheric CO₂. The main parameter controlling the long-term variability of the CO₂ uptake from the atmosphere was the air–sea CO₂ transfer velocity (57%), followed by the SST_RS (10%) and the chl a_RS (2%).     

Distribution of phytoplankton in the southern Black Sea in summer 1996, spring and autumn 1998

The species composition, abundance, and biomass of micro- (>15 μm) and nano- (<15 μm) phytoplankton were studied along the southern Black Sea during June–July 1996 and March–April and September 1998. A total of 150 species were identified, 50% of them being dinoflagellates. The average total phytoplankton abundance changed from 77×10³ cells l⁻¹ in spring to 110×10³ cells l⁻¹ in autumn and biomass from 250 μg l⁻¹ in summer to 1370 μg l⁻¹ in spring. Based on the extensive sampling grid from June–July 1996, phytoplankton seemed to have a rather homogeneous biomass distribution in the southern Black Sea. In all periods, the coccolithophorid Emiliania huxleyi was the most abundant species, its contribution to the total abundance ranging from 73% in autumn to 43% in spring. However, in terms of biomass, diatoms made up the bulk of phytoplankton in spring (97%, majority being Proboscia alata) and autumn (73%, majority being Pseudosolenia calcar-avis), and dinoflagellates in summer (74%, Gymnodinium sp.). There was a remarkable similarity in the dominant species between the western and eastern regions of the southern Black Sea, indicating transport of phytoplankton within the basin.     

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.     

Nutrient and organic matter patterns across the Mackenzie River, estuary and shelf during the seasonal recession of sea-ice

Suspended material, nutrients and organic matter in Mackenzie River water were tracked along a 300 km transect from Inuvik (Northwest Territories, Canada), across the estuarine salinity gradient in Kugmallit Bay, to offshore marine stations on the adjacent Mackenzie Shelf. All particulates measured (SPM, POC, PN, PP) declined by 87–95% across the salinity gradient and levels were generally below conservative mixing. Organic carbon content of suspended material decreased from 3.1% in the river to 1.7% in shelf surface waters while particulate C:N concurrently decreased from 17.1 to 8.6. Nitrate and silicate concentrations declined by more than 90% across the salinity gradient, with nitrate concentrations often below the conservative mixing line. Phosphate concentrations increased from 0.03 μmol/L in the river to 0.27 μmol/L over shelf waters, thereby changing the inorganic nutrient regime downstream from P to N limitation. Dissolved organic carbon decreased conservatively offshore while dissolved organic N and P persisted at high levels in the Mackenzie plume relative to river water, increasing 2.7 and 25.3 times respectively. A deep chlorophyll-a maximum was observed at two offshore stations and showed increases in most nutrients, particulates and organic matter relative to the rest of the water column. During river passage through the Mackenzie estuary, particulate matter, dissolved organic carbon and inorganic nutrients showed sedimentation, dilution and biological uptake patterns common to other arctic and non-arctic estuaries. Alternatively, inorganic content of particles increased offshore and dissolved organic N and P increased substantially over the shelf, reaching concentrations among the highest reported for the Arctic Ocean. These observations are consistent with the presence of a remnant ice-constrained (‘stamukhi’) lake from the freshet period and a slow flushing river plume constrained by sea-ice in close proximity to shore. Nutrient limitation in surface shelf waters during the ARDEX cruise contributed to the striking deep chlorophyll-a maximum at 21 m where phytoplankton communities congregated at the margin of nutrient-rich deep ocean waters.     

Protist entrapment in newly formed sea ice in the Coastal Arctic Ocean

Protist abundance and taxonomic composition were determined in four development stages of newly formed sea ice (new ice, nilas, young ice and thin first-year ice) and in the underlying surface waters of the Canadian Beaufort Sea from 30 September to 19 November 2003. Pico- and nanoalgae were counted by flow cytometry whereas photosynthetic and heterotrophic protists ≥ 4 µm were identified and counted by inverted microscopy. Protists were always present in sea ice and surface water samples throughout the study period. The most abundant protists in sea ice and surface waters were cells < 4 µm. They were less abundant in sea ice (418–3051 × 10³ cells L^− 1) than in surface waters (1393–5373 × 10³ cells L^− 1). In contrast, larger protists (≥ 4 µm) were more abundant in sea ice (59–821 × 10³ cells L^− 1) than in surface waters (22–256 × 10³ cells L^− 1). These results suggest a selective incorporation of larger cells into sea ice. The ≥ 4 µm protist assemblage was composed of a total number of 73 taxa, including 12 centric diatom species, 7 pennate diatoms, 11 dinoflagellates and 16 flagellates. The taxonomic composition in the early stage of ice formation (i.e., new ice) was very similar to that observed in surface waters and was composed of a mixed population of nanoflagellates (Prasinophyceae and Prymnesiophyceae), diatoms (mainly Chaetoceros species) and dinoflagellates. In older stages of sea ice (i.e., young ice and thin first-year ice), the taxonomic composition became markedly different from that of the surface waters. These older ice samples contained relatively fewer Prasinophyceae and more unidentified nanoflagellates than the younger ice. Diatom resting spores and dinoflagellate cysts were generally more abundant in sea ice than in surface waters. However, further studies are needed to determine the importance of this winter survival strategy in Arctic sea ice. This study clearly shows the selective incorporation of large cells (≥ 4 µm) in newly formed sea ice and the change in the taxonomic composition of protists between sea ice and surface waters as the fall season progresses.     

Pigments, size and distribution of Synechococcus spp. in the Black Sea

Pigments, size and distribution of Phycoerythrin-containing unicellular cyanobacteria Synechococcus spp. within the euphotic zone were studied for the first time in April–May 1994 in the western and southwestern Black Sea by epifluorescence microscopy and flow-cytometry. Synechococcus was present in varying quantities at every station and depth studied. Surface spatial distribution of Synechococcus revealed that cells were much more abundant in offshore waters than near coastal regions under the direct influence of the Danube river. Minimum and maximum cell concentrations ranged between 9×10² and 1.45×10⁵ cells/ml at the surface, between 2×10³ and 1.23×10⁵ cells/ml at the chlorophyll sub-maximum layer, and between 1.3×10² and 3.5×10² at the nitrite maximum layer. Cells at the chlorophyll sub-maximum layer (based on in-situ fluorometer readings) fluoresce brighter and longer than the ones at the surface and lower depths. Spectral properties of chromophore pigment types of total 64 clonal isolates from different depths down to the lower layer of the euphotic zone (60 m) in the southern Black Sea coast revealed that all have type 2 phycoerythrobilin in common, lacking in phycourobilin. In vivo fluorescence emission maxima for the phycoerythrobilin were about the same (578 nm) for all isolates. All isolates examined showed in vivo absorption maxima at between 435 and 442 nm and at about 681 nm due to chlorophyll-a. Based on the flow cytometer mean forward light scatter data for size distribution, it could be concluded that cells at the surface mixed layer (0–10 m) were larger in cell size than the cells at lower depths (20–60 m).     

Importance of particle-associated bacterial heterotrophy in a coastal Arctic ecosystem

The large quantities of particles delivered by the Mackenzie River to the coastal Beaufort Sea (Arctic Ocean) have implications for the spatial distribution, composition and productivity of its bacterial communities. Our objectives in this study were: (1) to assess the contribution of particle-associated bacteria (fraction ≥ 3 µm) to total bacterial production and their relationships with changing environmental conditions along a surface water transect; (2) to examine how particle-based heterotrophy changes over the annual cycle (Nov 2003–Aug 2004); and (3) to determine whether particle-associated bacterial assemblages differ in composition from the free-living communities (fraction < 3 µm). Our transect results showed that particle-associated bacteria contributed a variable percentage of leucine-based (BP-Leu) and thymidine-based (BP-TdR) bacterial production, with values up to 98% at the inshore, low salinity stations. The relative contribution of particle-associated bacteria to total BP-Leu was positively correlated with temperature and particulate organic material (POM) concentration. The annual dataset showed low activities of particle-associated bacteria during late fall and most of the winter, and a period of high particle-associated activity in spring and summer, likely related to the seasonal inputs of riverine POM. Results from catalyzed reporter deposition for fluorescence in situ hybridization (CARD-FISH) confirmed the dominance of Bacteria and presence of Archaea (43–84% and 0.2–5.5% of DAPI counts, respectively), which were evenly distributed throughout the Mackenzie Shelf, and not significantly related to environmental variables. Denaturing gradient gel electrophoresis (DGGE) revealed changes in the bacterial community structure among riverine, estuarine and marine stations, with separation according to temperature and salinity. There was evidence of differences between the particle-associated and free-living bacterial assemblages at the estuarine stations with highest POM content. Particle-associated bacteria are an important functional component of this Arctic ecosystem. Under a warmer climate, they are likely to play an increasing role in coastal biogeochemistry and carbon fluxes as a result of permafrost melting and increased particle transport from the tundra to coastal waters.     

The trophic role of the tunicate Salpa thompsoni in the Antarctic marine ecosystem

During a repeat grid survey and drogue study carried out in austral summer 1994/95, the abundance and feeding activity of salps were estimated in the Lazarev Sea region from net tows and in situ measurements of gut fluorescence. Throughout the survey area, Salpa thompsoni accounted for >95% of the total salp stock while Ihlea racovitzai was consistently represented in very low abundances. Maximum densities of S. thompsoni, with ≈4000 ind. 1000 m⁻³, were recorded in the Marginal Ice Zone (MIZ) in December when chlorophyll-a concentrations were well below 1 mg m⁻³. A dramatic decrease in salp stock was observed at the beginning of January, when S. thompsoni virtually disappeared from the most productive area of the MIZ where chlorophyll-a concentrations had by then reached bloom levels of 1.5–3 mg (Chl-a) m⁻³. In situ grazing measurements showed that throughout the cruise S. thompsoni exhibited the highest ingestion rates per individual of any of the most abundant components of the grazing pelagic community, with maxima of ≈160 μg (pigm) ind. ⁻¹ d⁻¹. These feeding rates are 3 to 5 times higher than those previously obtained using in vitro incubations. The total daily consumption of the population of S. thompsoni varied from 0.3 to 108% of daily primary production. We suggest that competitive removal of food by S. thompsoni, rather than direct predation, is responsible for the low krill abundances generally associated with salp swarms.     

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.     

Production of Acartia omorii (Copepoda: Calanoida) in Ilkwang Bay, southeastern coast of Korea

Production of the marine calanoid copepod Acartia omorii was measured from 2 October 1991 to 8 October 1992 at a station in Ilkwang Bay on the southeastern coast of Korea. A. omorii (nauplii + copepodites + adults) were present in the plankton throughout the year, with seasonal variation in abundance. Biomass of A. omorii was averaged at 0.44 mgC m^− 3, with peaks in February and July, and relatively low biomass in late summer and fall. Egg production rate ranged from 2.4 to 151.9 μgC m^− 3 day^− 1, which was equivalent to 95–6075 eggs m^− 3 day^− 1. Fecundity of an adult female was averaged at 38 eggs female^− 1 day^− 1. Instantaneous growth rates of copepodites were higher than those of nauplii stages. Annual production of A. omorii ranged from 33.5 mgC m^− 3 year^− 1 to 221 mgC m^− 2 year^− 1, showing a seasonal variation of daily production rate with peaks in February and July. The daily production rate of A. omorii was significantly correlated with chlorophyll a concentration. These results suggest that standing stocks and/or productivity of phytoplankton are the major influencing factors, rather than water temperature for the seasonal variation of production of A. omorii in Ilkwang Bay.