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.     

Microphytobenthos biomass at Kerguelen's Land (Subantarctic Indian Ocean): repartition and variability during austral summers

Microphytobenthos biomass has been measured at several coastal sites on the SE of the main island of the Kerguelen Archipelago (Indian Ocean), during several austral summers (1985–1992), using a conventional fluorometric method. Heterogeneity tests, conducted on two different intertidal sites (Port-Aux-Français, PAF; and Port-Raymond, PRA), showed low standard deviations, whereas the mean concentrations were highly different. Pigment concentrations showed a high variability related to the characteristics of the sediments: from low biomass in coarse intertidal sand, submitted to intense scouring (0.32±0.31 μg Chl a g⁻¹ dw, 0.29±0.14 μg Phaeo g⁻¹ dw) to high biomass in intertidal muddy sand in sheltered areas, particularly along estuaries (54 μg Chl a, 15 μg Phaeo g⁻¹ dw at Korrigan). The subtidal muddy sediments under a Macrocystis pyrifera (Linné) and Durvillaea antarctica (Chamisso in Choris) belt exhibited high concentrations in phaeopigment (Phaeo) (up to 136±83 μg g⁻¹ dw; PRA), while the concentration of chlorophyll a (Chl a) was relatively low. The dense macroalgal canopy supports an important epiphytic diatom biomass (mainly the genera Cocconeis Ehrenberg and Grammatophora Ehrenberg), which is sedimenting after degradation and is in part responsible of the high levels of Phaeo in all sediments. Macroalgal debris were observed, but diatom frustules were dominant in most surficial subtidal sediments. A circatidal mud, in the Morbihan Bay, made of a sponge spicule mat (50 m deep; 4.96 μg Chl a g⁻¹ dw), showed a very low Chl a/Phaeo ratio (0.1), while it reached up to 6 in intertidal sand. Surprisingly, a penguin rookery beach, at the east side of Courbet Peninsula, was characterized by a very low biomass (0.07±0.04 μg Chl a g⁻¹ dw), while it was nutrient enriched, particularly with nitrates.In comparison with the data at the similar latitude, but in temperate regions from the Northern Hemisphere, the microphytobenthos biomass, recorded at Kerguelen's Land, exhibited relatively high pigment concentrations, particularly the Phaeo, and supported a dense and diversified subtidal macrofauna composed of polychaetes (particularly Thelepus extensus Hutchings and Glasby), sea urchins, mytillids and gammarids. The exuberant macroalgal canopy, coastal indentations and low tidal amplitude must be in part responsible of these large benthic primary and secondary biomasses.     

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.     

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.     

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.     

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.     

Carbonate chemistry dynamics and carbon dioxide fluxes across the atmosphere–ice–water interfaces in the Arctic Ocean: Pacific sector of the Arctic

Climatic changes in the Northern Hemisphere have led to remarkable environmental changes in the Arctic Ocean, which is surrounded by permafrost. These changes include significant shrinking of sea-ice cover in summer, increased time between sea-ice break-up and freeze-up, and Arctic surface water freshening and warming associated with melting sea-ice, thawing permafrost, and increased runoff. These changes are commonly attributed to the greenhouse effect resulting from increased atmospheric carbon dioxide (CO₂) concentration and other non-CO₂ radiatively active gases (methane, nitrous oxide). The greenhouse effect should be most pronounced in the Arctic where the largest air CO₂ concentrations and winter–summer variations in the world for a clean background environment were detected. However, the air–land–shelf interaction in the Arctic has a substantial impact on the composition of the overlying atmosphere; as the permafrost thaws, a significant amount of old terrestrial carbon becomes available for biogeochemical cycling and oxidation to CO₂. The Arctic Ocean's role in determining regional CO₂ balance has been ignored, because of its small size (only  4% of the world ocean area) and because its continuous sea-ice cover is considered to impede gaseous exchange with the atmosphere so efficiently that no global climate models include CO₂ exchange over sea-ice. In this paper we show that: (1) the Arctic shelf seas (the Laptev and East-Siberian seas) may become a strong source of atmospheric CO₂ because of oxidation of bio-available eroded terrestrial carbon and river transport; (2) the Chukchi Sea shelf exhibits the strong uptake of atmospheric CO₂; (3) the sea-ice melt ponds and open brine channels form an important spring/summer air CO₂ sink that also must be included in any Arctic regional CO₂ budget. Both the direction and amount of CO₂ transfer between air and sea during open water season may be different from transfer during freezing and thawing, or during winter when CO₂ accumulates beneath Arctic sea-ice; (4) direct measurements beneath the sea ice gave two initial results. First, a drastic pCO₂ decrease from 410 μatm to 288 μatm, which was recorded in February–March beneath the fast ice near Barrow using the SAMI-CO₂ sensor, may reflect increased photosynthetic activity beneath sea-ice just after polar sunrise. Second, new measurements made in summer 2005 beneath the sea ice in the Central Basin show relatively high values of pCO₂ ranging between 425 μatm and 475 μatm, values, which are larger than the mean atmospheric value in the Arctic in summertime. The sources of those high values are supposed to be: high rates of bacterial respiration, import of the Upper Halocline Water (UHW) from the Chukchi Sea (CS) where values of pCO₂ range between 400 and 600 μatm, a contribution from the Lena river plume, or any combination of these sources.     

Dynamics of suprabenthos-zooplankton communities around the Balearic Islands (western Mediterranean): Influence of environmental variables and effects on the biological cycle of Aristeus antennatus

Dynamics of suprabenthos and zooplankton were analyzed in two areas located in the NW (off Sóller harbour) and S (off Cabrera Archipelago) of Mallorca (Balearic Islands, western Mediterranean) at depths ranging between 135–780 m. Four stations situated respectively at 150 m (shelf-slope break), and at bathyal depths of 350, 650 and 750 m were sampled at bi-monthly intervals during six cruises performed between August 2003 and June 2004. Suprabenthos showed maximum biomass in both areas from late spring to summer (April to August), while minimum biomass was found in autumn (September–November). Though variable, temporal dynamics of zooplankton showed peaks of biomass in late winter and summer (February and June), while minimals occurred in autumn (August–September) and, at bathyal depths, in April. Suprabenthos (abundance; MDS analyses) showed a sample aggregation as a function of depth (3 groups corresponding to the shelf-slope break, upper slope — over 350 m; and the middle, deeper part of the slope — over 650–750 m), without any separation of hauls by season. By contrast, zooplankton samples were separated by season and not by depth. There was evidence of three seasonal groups corresponding to summer (June 2004–August 2003), autumn–winter (September and November 2003, February 2004), and spring (April 2004), being especially well established off Sóller. In general, suprabenthos was significantly correlated with the sediment variables (e.g. total organic matter content (% OM), potential REDOX), whereas zooplankton was almost exclusively dependent on Chl a at the surface, which suggests two different food sources for suprabenthos and zooplankton. The increase of suprabenthos abundance in April–June was paralleled by a sharp increase (ca. 2.8 times) in the %OM on sediment during the same period, coupled ca. 1–2 months of delay with the peak of surface Chl a recorded in February–March (from satellite imagery data). Suprabenthos biomass was also correlated with salinity close to the bottom, suggesting a link between suprabenthos abundance and changes in the oceanographic condition of water masses close to the bottom. It is suggested that a higher suprabenthos biomass recorded off Sóller in comparison to that off Cabrera in June could, in turn, be related to a seasonal inflow of Levantine Intermediate Water (LIW) in April–June in this area at mid bathyal depths (350–650 m). This trend would be based on: 1) it was evident only at mid-slope depths between 350–750 m, coinciding with the LIW distribution, and 2) it was not recorded among zooplankton (collected throughout the water column). The possible effect of the fluctuations of suprabenthos and zooplankton on higher trophic levels has been explored studying the diet and food consumption rates of the red shrimp Aristeus antennatus, as indicator species by its dominance in bathyal communities. A. antennatus increased its food consumption from February to April–June 2004 off Sóller, which in the case of large (CL > 40 mm) specimens was found in both areas. In addition, there was a shift of diet from winter to spring–early summer. In this last period, A. antennatus preyed upon euphausiids and mesopelagic decapods and fish, while benthos (e.g. polychaetes and bivalves) decreased in the diet. This indicates an increase in the food consumption and probably in the caloric content of the diet in pre-spawning females in April–June 2004, which is synchronized with the period when gonad development begins in A. antennatus females (May–June). Anyway, macrozooplankton, and not suprabenthos, is crucial as a high energetic food source in the coupling between food intake and reproduction in the red shrimp.     

Weaving marine food webs from end to end under global change

Marine food web dynamics are determined by interactions within and between species and between species and their environment. Global change directly affects abiotic conditions and living organisms, impinging on all trophic levels in food webs. Different groups of marine researchers traditionally study different aspects of these changes. However, over medium to long time scales perturbations affecting food webs need to be considered across the full range from nutrients to top predators. Studies of end-to-end marine food webs not only span organism sizes and trophic levels, but should also help align multidisciplinary research to common goals and perspectives. Topics are described that bridge disciplinary gaps and are needed to develop new understanding of the reciprocal impacts of global change on marine food webs and ocean biogeochemistry. These include (1) the effects of nutrients on biomass and production, (2) the effects of varying element ratios on food web structure and food quality, (3) bulk flows of energy and material in food webs and their efficiencies of transfer, (4) the ecological effects of species richness and the roles of microbial organisms, (5) the role of feeding behaviour in food web dynamics and trophic controls, (6) the spatial dynamics of communities and links between different food webs, (7) the combined effects of body size and behaviour in determining dynamics of food webs, and (8) the extent to which the ability of marine organisms (and communities) to adapt will influence food web dynamics. An overriding issue that influences all topics concerns the time and space scales of ecosystem variability. Threads link different nodes of information among various topics, emphasizing the importance of tackling food web studies with a variety of modelling approaches and through a combination of field and experimental studies with a strong comparative approach.     

Methane release and coastal environment in the East Siberian Arctic shelf

In this paper we present 2 years of data obtained during the late summer period (September 2003 and September 2004) for the East Siberian Arctic shelf (ESAS). According to our data, the surface layer of shelf water was supersaturated up to 2500% relative to the present average atmospheric methane content of 1.85 ppm, pointing to the rivers as a strong source of dissolved methane which comes from watersheds which are underlain with permafrost. Anomalously high concentrations (up to 154 nM or 4400% supersaturation) of dissolved methane in the bottom layer of shelf water at a few sites suggest that the bottom layer is somehow affected by near-bottom sources. The net flux of methane from this area of the East Siberian Arctic shelf can reach up to 13.7 × 10⁴ g CH₄ km^− 2 from plume areas during the period of ice free water, and thus is in the upper range of the estimated global marine methane release. Ongoing environmental change might affect the methane marine cycle since significant changes in the thermal regime of bottom sediments within a few sites were registered. Correlation between calculated methane storage within the water column and both integrated salinity values (r = 0.61) and integrated values of dissolved inorganic carbon (DIC) (r = 0.62) suggest that higher concentrations of dissolved methane were mostly derived from the marine environment, likely due to in-situ production or release from decaying submarine gas hydrates deposits. The calculated late summer potential methane emissions tend to vary from year to year, reflecting most likely the effect of changing hydrological and meteorological conditions (temperature, wind) on the ESAS rather than riverine export of dissolved methane. We point out additional sources of methane in this region such as submarine taliks, ice complex retreat, submarine permafrost itself and decaying gas hydrates deposits.     

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.).     

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.     

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.     

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.     

Collaborative Integrated Management in Canada's North: The Role of Local and Traditional Knowledge and Community-Based Monitoring

The objective of this article is to take stock of integrated management in the Canadian North, assessing its contribution to the advancement of knowledge and practice regarding the role of indigenous knowledge and community-based monitoring. This is done in three steps. (1) The Beaufort Sea, designated a Large Ocean Management Area under Canada's Oceans Action Plan, is used as an example of a consultative planning process, with special attention to indigenous peoples. (2) How specifically can indigenous knowledge contribute to integrated management? The problem of Arctic marine food web contamination is used to illustrate the strengths and limitations of traditional ecological knowledge and its relationship to science. (3) The discussion of community-based monitoring relies on Voices From The Bay study involving the Inuit and Cree of Hudson and James Bay, and Inuit observations of climate change study in the Canadian western Arctic. The examples together address integrated coastal management and the health of ocean ecosystems, showing how stakeholder participation and knowledge helps widen the range of knowledge to understand and help monitor environmental change.     

Measuring the thicknesses of the freshwater-layer plume and sea ice in the land-fast ice region of the Mackenzie Delta using helicopter-borne sensors

Helicopter-borne sensors have been used since the early 1990s to monitor ice properties in support of winter marine transportation along the east coast of Canada. The observations are used in ice chart production and to validate ice hazard identification algorithms using satellite advanced synthetic aperture radar (ASAR) imagery. In this study we evaluated the sensors' additional capability to monitor the freshwater plume characteristic beneath land-fast ice. During the Canadian Arctic Shelf Exchange Study (CASES) data were collected over the Mackenzie Delta in the southern Beaufort Sea where a buoyant river plume exists. Results showed that the electromagnetic–laser system could describe not only the ice properties but also the horizontal distribution of the freshwater plume depths that decreased in depth stepwise offshore as the flow of the buoyant plume was restricted by a series of ridge-rubble fields running parallel to the coast. Relative to the 2 m mean ice thickness, the plume layer depth varied from zero under mobile offshore pack ice to 3 m inshore of the third set of ridge-rubble fields.     

Topological constraints on the dynamics of wasp-waist ecosystems

Small pelagic fish species like anchovy or sardines are of high ecological and economical importance. As marine food webs are fished down, these small pelagics tend to be more exploited and overfished. It is not yet very well known what the possible effects of their collapse can be, therefore there is an urgent need to outline a theoretical framework for understanding their dynamics. These fish occupy very special position in food webs, ensuring energy transfer between species lower and higher levels, while forming narrow “wasp-waists” poor in number of species (but very abundant). Our purpose was to quantify the interaction structure of model food webs of equal complexity but different levels of “wasp-waistedness”. We analysed the topological properties of the webs by characterising every direct and indirect interactions between individual species, as well as by assessing the relative positional importance of each species in each web. We found that (1) the shorter the interaction pathways considered, the weaker the predictive power of node degree for positional importance, (2) the importance of species varies more in wasp-waist food webs, (3) if longer indirect chain effects are considered, indirect effects can well be stronger than direct ones, (4) interactions between coexisting wasp-waist species are stronger than the average, and (5) the “self-regulatory” looping effects are also stronger for wasp-waist species. Based on the topological properties of the networks, our results describe constraints acting on the dynamical behaviour of wasp-waist ecosystems. We give explanations, from this viewpoint, for regime shifts in which one WW species replaces another, and for the unpredictable dynamics of these fish stocks. From a marine conservation viewpoint, we illustrate that as the abundance of wasp-waist species decreases, the architecture of energy flows becomes highly vulnerable and unreliable. We provide an approach for quantifying these structural changes.     

Environmental causes of the fluctuations of red shrimp (Aristeus antennatus) landings in the Catalan Sea

The fluctuations of catches of the deep-sea shrimp Aristeus antennatus (Risso, 1816) were analysed in the 6 ports of Catalonia where production is concentrated, based on monthly landings from 1988 to 2004. The 6 selected ports produced 300 t of red shrimp in 2004, or 80% of the total production in Catalonia. The series for each port showed clear interannual variability, with peaks of production in the early 1990s and more recently from 2001 to 2003. The time periods of the monthly data series, estimated by frequency analysis based on Fourier transform, varied around 7–8 years in the four central ports and 12–13 years in the two northern ports. Additionally, the different curves were not in phase: even in nearby ports, the maximum production is observed in different years. Since the North Atlantic Oscillation (NAO) index is an excellent proxy for long-term series of environmental variables, we aimed to explore relationships between the fluctuation of the NAO index and A. antennatus landings in the Catalan Sea. The correlation between the mean annual NAO index and the annual catches in each port was positive and significant with some time lags (from 1 to 3 years). The existence of clear patterns linking the NAO with marine ecological processes has been demonstrated in many studies, but the underlying ecological mechanisms are far from being well understood. The variations in environmental parameters linked to the NAO may act on biological organisms at different levels (individual, population) through physiology (metabolic and reproductive processes) or through trophic relationships, including ecological cascade effects. We propose that NAO-induced environmental variability may enhance food supply to A. antennatus and hence strengthen the reproductive potential of particular year classes, which result in increased catches 1 to 3 years later, although other possible effects of environmental variability on the population dynamics of this species are worth investigating.     

Number and phylogenetic affiliation of bacteria assimilating dimethylsulfoniopropionate and leucine in the ice-covered coastal Arctic Ocean

The ability of bacteria to assimilate sulfur from dimethylsulfoniopropionate (DMSP) was examined in the western Arctic Ocean by combining microautoradiography and fluorescence in situ hybridization (FISH). Assimilation of leucine was also measured for comparative purposes since leucine is considered a universal substrate for bacteria, which use it for protein synthesis. Samples were collected at 3 m depth, through a hole in the ice, in the CASES (Canadian Arctic Shelf Ecosystem Study) overwintering station in Franklin Bay (eastern Beaufort Sea) in March and May 2004 to compare two contrasting situations: winter and early spring. FISH counts indicated that the bacterial assemblage consisted of α- (up to 60% of the EUB positive cells), β- (up to 10%) and γ-proteobacteria (around 20%), and Bacteroidetes (up to 60%). The β-proteobacteria were not active with any of the two substrates tested. The remaining groups were much less efficient at assimilating DMSP-sulfur (5% of the cells) than leucine (20–35%) both in winter and in spring. Only the Roseobacter group of α-proteobacteria showed a similar assimilation of both substrates.