Stratified Fjord Research Articles

Dinophysis acuminata or Dinophysis acuta: What Makes the Difference in Highly Stratified Fjords?

Dinophysis acuminata and D. acuta, which follows it seasonally, are the main producers of lipophilic toxins in temperate coastal waters, including Southern Chile. Strains of the two species differ in their toxin profiles and impacts on shellfish resources. D. acuta is considered the major cause of diarrhetic shellfish poisoning (DSP) outbreaks in Southern Chile, but there is uncertainty about the toxicity of D. acuminata, and little information on microscale oceanographic conditions promoting their blooms. During the austral summer of 2020, intensive sampling was carried out in two northern Patagonian fjords, Puyuhuapi (PUY) and Pitipalena (PIT), sharing D. acuminata dominance and D. acuta near detection levels. Dinophysistoxin 1 (DTX 1) and pectenotoxin 2 (PTX 2) were present in all net tow samples but OA was not detected. Although differing in hydrodynamics and sampling dates, D. acuminata shared behavioural traits in the two fjords: cell maxima (>103 cells L-1) in the interface (S ~ 21) between the estuarine freshwater (EFW)) and saline water (ESW) layers; and phased-cell division (µ = 0.3-0.4 d-1) peaking after dawn, and abundance of ciliate prey. Niche analysis (Outlying Mean Index, OMI) of D. acuta with a high marginality and much lower tolerance than D. acuminata indicated an unfavourable physical environment for D. acuta (bloom failure). Comparison of toxin profiles and Dinophysis niches in three contrasting years in PUY-2020 (D. acuminata bloom), 2018 (exceptional bloom of D. acuta), and 2019 (bloom co-occurrence of the two species)-shed light on the vertical gradients which promote each species. The presence of FW (S < 11) and thermal inversion may be used to provide short-term forecasts of no risk of D. acuta blooms and OA occurrence, but D. acuminata associated with DTX 1 pose a risk of DSP events in North Patagonian fjords.

Wind influence on residual circulation in Patagonian channels and fjords

Circulation in estuaries is typically considered to be driven by horizontal density gradients, called gravitational circulation, but can be modified by other forcing such as the wind. Studies have shown that wind influence can either augment or counteract density-driven residual flows in estuaries, but the temporal and spatial extent of this influence remains unknown, particularly in deep and highly stratified fjords. This study aims to fill this gap by investigating the seasonal variability in residual circulation, wind forcing, and stratification in an extensive fjord and channel system in Chilean Patagonia. Observations of current velocity and wind are supplemented by atmospheric and hydrodynamic circulation models in the Moraleda Channel, a system that connects the fjord system of Northern Patagonia to the Pacific Ocean. Results show a marked seasonality, given that the surface outflow is reversed by wind forcing during the winter, a season characterized by strong southward (up-channel) winds. This produces a near-surface flow directed up-channel in the upper ∼40 m of the water column, in opposition to the horizontal pressure gradient. Cross-channel transects revealed that the vertical structure of the along-channel residual velocity was three-layer during the winter. The near-surface (<50 m) along-channel current on the eastern side of the channel followed the wind direction (southward). Between 50 and 100 m depth the flow was out-channel and below 100 m depth the flow was again in-channel. On the western side, the along-channel residual flow was two-layer, with near surface flow directed northward (against the wind) with a return flow below (deeper than 50 m). During the spring, northward (out-channel) winds dominated and the vertical structure of the along-channel residual velocity was two-layer, augmenting the horizontal pressure gradient, which stabilized the water column, enhancing stratification, inhibiting mixing and elevated gravitational circulation.

Oxygen availability driven trends in DOM molecular composition and reactivity in a seasonally stratified fjord

Ocean deoxygenation could potentially trigger substantial changes in the composition and reactivity of dissolved organic matter (DOM) pool, which plays an important role in the global carbon cycle. To evaluate links between DOM dynamics and oxygen availability, we investigated the DOM composition under varying levels of oxygen in a seasonally hypoxic fjord through a monthly time-series over two years. We used ultrahigh-resolution Fourier transform ion cyclotron resonance mass spectrometry (FT-ICR MS) to characterize DOM on a molecular level. We find a clear trend both in diversity and molecular composition of the DOM along the oxygen gradient. As oxygen decreased, the chemodiversity was significantly increased, along with accumulation of relatively high-molecular-weight, reduced and unsaturated compounds enriched with carboxyl-group structures, which were also thermodynamically less favorable to biodegradation. Our results suggested that oxygen depletion selectively protected otherwise bioavailable compounds from decomposition and may promote the accumulation of a larger recalcitrant DOM pool in the global ocean, which could provide negative feedback to the ocean carbon sequestration and climate change.

Taxon-Specific Shifts in Bacterial and Archaeal Transcription of Dissolved Organic Matter Cycling Genes in a Stratified Fjord

ABSTRACTA considerable fraction of organic matter derived from photosynthesis in the euphotic zone settles into the ocean’s interior and, as it progresses, is degraded by diverse microbial consortia that utilize a suite of extracellular enzymes and membrane transporters. Still, the molecular details that regulate carbon cycling across depths remain little explored. As stratification in fjords has made them attractive models to explore patterns in biological oceanography, we here analyzed bacterial and archaeal transcription in samples from five depth layers in the Gullmar Fjord, Sweden. Transcriptional variation over depth correlated with gradients in chlorophyll a and nutrient concentrations. Differences in transcription between sampling dates (summer and early autumn) were strongly correlated with ammonium concentrations, which potentially was linked with a stronger influence of (micro-)zooplankton grazing in summer. Transcriptional investment in carbohydrate-active enzymes (CAZymes) decreased with depth and shifted toward peptidases, partly a result of elevated CAZyme transcription by Flavobacteriales, Cellvibrionales, and Synechococcales at 2 to 25 m and a dominance of peptidase transcription by Alteromonadales and Rhodobacterales from 50 m down. In particular, CAZymes for chitin, laminarin, and glycogen were important. High levels of transcription of ammonium transporter genes by Thaumarchaeota at depth (up to 18% of total transcription), along with the genes for ammonia oxidation and CO2 fixation, indicated that chemolithoautotrophy contributed to the carbon flux in the fjord. The taxon-specific expression of functional genes for processing of the marine pool of dissolved organic matter and inorganic nutrients across depths emphasizes the importance of different microbial foraging mechanisms over spatiotemporal scales for shaping biogeochemical cycles.IMPORTANCE It is generally recognized that stratification in the ocean strongly influences both the community composition and the distribution of ecological functions of microbial communities, which in turn are expected to shape the biogeochemical cycling of essential elements over depth. Here, we used metatranscriptomics analysis to infer molecular detail on the distribution of gene systems central to the utilization of organic matter in a stratified marine system. We thereby uncovered that pronounced shifts in the transcription of genes encoding CAZymes, peptidases, and membrane transporters occurred over depth among key prokaryotic orders. This implies that sequential utilization and transformation of organic matter through the water column is a key feature that ultimately influences the efficiency of the biological carbon pump.

Vertical Mixing in Stratified Fjords Near Tidewater Outlet Glaciers Along Northwest Greenland

AbstractVertical mixing of upper water masses in Arctic fjords is important for circulation and transport of nutrients and heat. However, the distribution of turbulent mixing is poorly known. Here we present hydrographic and microscale turbulence measurements from six fjords in Northwestern (69°N–75°N) Greenland. Water mass distributions showed the presence of warm bottom water of Atlantic origin in all fjord transects and a significant modification of upper water masses in fjords with tidewater outlet glaciers. Spatial and temporal distribution of turbulence in the fjords showed, in general, low rates of turbulent kinetic energy, ε (∼10−9 W kg−1), and strong stratification implies that vertical exchange in the upper ∼200 m was relatively weak. However, measurements within 2 km of the terminus of a tidewater outlet glacier showed values of ε and inferred turbulent diffusion coefficients &gt;10−4 m2 s−1 that were about two orders of magnitude larger than elsewhere in the deep fjords. This elevated mixing corresponded to comparatively large heat and salt fluxes. These observations suggest that vertical mixing near tidewater outlet glaciers results in localized mixing hot spots, likely due to subglacial discharge and near‐glacial current shear, that contribute significantly to vertical exchange in the deep fjords.

Bacterial and archaeal lipids trace chemo(auto)trophy along the redoxcline in Vancouver Island fjords.

Marine oxygen minimum zones play a crucial role in the global oceanic carbon, nitrogen, and sulfur cycles as they harbor microbial communities that are adapted to the water column chemistry and redox zonation, and in turn control the water column chemistry and greenhouse gas release. These micro-organisms have metabolisms that rely on terminal electron acceptors other than O2 and often benefit from syntrophic relationships (metabolic coupling). Here, we study chemo(auto)trophy along the redoxcline in two stratified fjords on Vancouver Island (Canada) using bacterial bacteriohopanepolyols and archaeal ether lipids. We analyze the distribution of these lipid classes in suspended particulate matter (SPM) to trace ammonia oxidation, anaerobic ammonium oxidation (anammox), sulfate reduction/sulfur oxidation, methanogenesis, and methane oxidation, and investigate ecological niches to evaluate potential links between their respective bacterial and archaeal sources. Our results show an unparalleled BHP and ether lipid structural diversity that allows tracing the major redox-driven metabolic processes at the time of sampling: Both fjords are dominated by archaeal ammonia oxidation and anammox; sulfate-reducing bacteria may be present in Deer Bay, but absent from Effingham Inlet; methanogenic Euryarchaeota and archaeal and bacterial methanotrophs are detectable at low abundance. Correlation analysis reveals distinct biomarker clusters that provide constraints on the biogeochemical niches of some orphan BHP and ether lipids such as in situ-produced adenosyl-BHPs or unsaturated archaeols.

Niche differentiation of Dinophysis acuta and D. acuminata in a stratified fjord

Dinophysis acuta and D. acuminata are associated with lipophilic toxins in Southern Chile. Blooms of the two species coincided during summer 2019 in a highly stratified fjord system (Puyuhuapi, Chilean Patagonia). High vertical resolution measurements of physical parameters were carried out during 48 h sampling to i) explore physiological status (e.g., division rates, toxin content) and ii) illustrate the fine scale distribution of D. acuta and D. acuminata populations with a focus on water column structure and co-occurring plastid-bearing ciliates. The species-specific resources and regulators defining the realized niches (sensu Hutchinson) of the two species were identified. Differences in vertical distribution, daily vertical migration and in situ division rates (with record values, 0.76 d−1, in D. acuta), in response to the environmental conditions and potential prey availability, revealed their niche differences. The Outlying Mean Index (OMI) analysis showed that the realized niche of D. acuta (cell maximum 7 × 103 cells L−1 within the pycnocline) was characterized by sub-surface estuarine waters (salinity 23 – 25), lower values of turbulence and PAR, and a narrow niche breath. In contrast, the realized niche of D. acuminata (cell maximum 6.8 × 103 cells L−1 just above the pycnocline) was characterized by fresher (salinity 17 – 20) outflowing surface waters, with higher turbulence and light intensity and a wider niche breadth. Results from OMI and PERMANOVA analyses of co-occurring microplanktonic ciliates were compatible with the hypothesis of species such as those from genera Pseudotontonia and Strombidium constituting an alternative ciliate prey to Mesodinium. The D. acuta cell maximum was associated with DSP (OA and DTX-1) toxins and pectenotoxins; that of D. acuminata only with pectenotoxins. Results presented here contribute to a better understanding of the environmental drivers of species-specific blooms of Dinophysis and management of their distinct effects in Southern Chile.

Drivers of Megabenthic Community Structure in One of the World’s Deepest Silled-Fjords, Sognefjord (Western Norway)

The Sognefjord is the longest (205 km) and deepest (1308 m) fjord in Norway, and the second-longest in the world. Coast-fjord exchange in Sognefjord is limited by a seaward sill at 170 m water depth, which causes a clear stratification between water masses as the dense oxygen-poor basin water mixes slowly with the well-oxygenated water directly above from the coastal ocean. Due to the homogeneity and limited variability in the deep-water, the deep slopes of Sognefjord represent the ideal setting to study how abiotic factors influence the deep-water benthic community structure. During the summer of 2017, two remotely operated vehicle (ROV) video transects were performed to compare the megabenthic community behind the sill (water depth: 1230 to 55 m; transect length: 1.39 km; distance from sill: ∼17 km) and within the central fjord (water depth: 1155–85 m; transect length: 2.43 km; distance from sill: ∼79 km). Accompanying conductivity–temperature–depth (CTD) deployments were made to measure the in situ abiotic factors and nutrient concentrations at each transect location, while the substrate characteristics (percent cover of soft and hard exposed substrate) were documented from the video footage. Here, Sognefjord’s megabenthic community composition, distribution, and species richness were analyzed in relation to abiotic factors (e.g., depth, salinity, dissolved oxygen, chlorophyll a concentration, and percent cover of hard and soft substrata) within the fjord. Basin communities were homogeneous and characterized by sponges, echinoderms, and crustaceans, whereas the shallower regions were dominated by mobile scavengers. Contrary to other fjord-based studies, species richness and diversity were stable in the fjord basin and decreased with proximity to the sill, decreasing water depth, and at the boundary between intermediate and basin water. The findings demonstrate that highly stratified fjords support stable communities in their basins; however, further research is needed to investigate the influence water mass dynamics have on silled-fjord megafauna communities.

Gas transfer velocities in Norwegian fjords and the adjacent North Atlantic waters

Summary We investigated air-sea carbon dioxide (CO2) transfer in situ to determine the role of wind and turbulence in forcing gas transfer. In situ gas transfer velocities of CO2 were measured with a floating chamber technique along the Norwegian coast and inside the Sogne- and Trondheimsfjord. Gas transfer velocities were related to wind speed and turbulence, but neither wind speed nor turbulence can satisfactorily predict gas transfer velocity. However, comparison to existing wind-based parameterizations showed that the data from this study have a similar trend. Generally, we measured higher transfer velocities than the parameterizations predict. In the North Atlantic, we measured transfer velocities of up to 54.9 cm h−1 versus predicted transfer velocities of 6.3 cm h−1 at a wind speed of 3.7 m s−1. In addition, we observed that measurements of transfer velocities at wind speeds below 4 m s−1 are higher than predictions. Wind-based parameterizations are lacking data in the low wind regime for validation, and we provide 25 data points for this critical wind speed range. Overall, results indicate that Norwegian fjords and the adjacent North Atlantic are sinks for atmospheric CO2 during summer, with uptake rates of −9.6 ± 7.6 μmol m−2 min−1 and −4.1 ± 1.7 μmol m−2 min−1, respectively. Due to the low partial pressure of CO2 in the upper water layer of the stratified fjords (down to 150.7 μatm), the Sogne- and Trondheimsfjord absorb 196 tons of carbon per day during the summer.

Interannual Variability of Dinophysis acuminata and Protoceratium reticulatum in a Chilean Fjord: Insights from the Realized Niche Analysis.

Here, we present the interannual distribution of Dinophysis acuminata and Protoceratium reticulatum over a 10-year period in the Reloncaví Fjord, a highly stratified fjord in southern Chile. A realized subniche approach based on the Within Outlying Mean Index (WitOMI) was used to decompose the species’ realized niche into realized subniches (found within subsets of environmental conditions). The interannual distribution of both D. acuminata and P. reticulatum summer blooms was strongly influenced by climatological regional events, i.e., El Niño Southern Oscillation (ENSO) and the Southern Annual Mode (SAM). The two species showed distinct niche preferences, with blooms of D. acuminata occurring under La Niña conditions (cold years) and low river streamflow whereas P. reticulatum blooms were observed in years of El Niño conditions and positive SAM phase. The biological constraint exerted on the species was further estimated based on the difference between the existing fundamental subniche and the realized subniche. The observed patterns suggested that D. acuminata was subject to strong biological constraint during the studied period, probably as a result of low cell densities of its putative prey (the mixotrophic ciliate Mesodinium cf. rubrum) usually observed in the studied area.

Effect of Topography on Subglacial Discharge and Submarine Melting During Tidewater Glacier Retreat

To first order, subglacial discharge depends on climate, which determines precipitation fluxes and glacier mass balance, and the rate of glacier volume change. For tidewater glaciers, large and rapid changes in glacier volume can occur independent of climate change due to strong glacier dynamic feedbacks. Using an idealized tidewater glacier model, we show that these feedbacks produce secular variations in subglacial discharge that are influenced by subglacial topography. Retreat along retrograde bed slopes (into deep water) results in rapid surface lowering and coincident increases in subglacial discharge. Consequently, submarine melting of glacier termini, which depends on subglacial discharge and ocean thermal forcing, also increases during retreat into deep water. Both subglacial discharge and submarine melting subsequently decrease as glacier termini retreat out of deep water and approach new steady state equilibria. In our simulations, subglacial discharge reached peaks that were 6–17% higher than preretreat values, with the highest values occurring during retreat from narrow sills, and submarine melting increased by 14% for unstratified fjords and 51% for highly stratified fjords. Our results therefore indicate that submarine melting acts in concert with iceberg calving to cause tidewater glacier termini to be unstable on retrograde beds. The full impact of submarine melting on tidewater glacier stability remains uncertain, however, due to poor understanding of the coupling between submarine melting and iceberg calving.

Seiche excitation in a highly stratified fjord of southern Chile: the Reloncaví fjord

Abstract. We describe a seiche process based on current, temperature, and sea-level data obtained from the Reloncaví fjord (41.6° S, 72.5° W) in southern Chile. We combined 4 months of acoustic Doppler current profiler (ADCP) data with sea-level, temperature, and wind time series to analyze the dynamics of low-frequency (periods &gt; 1 day) internal oscillations in the fjord. Additionally, seasonal conductivity, temperature, and depth (CTD) data from 19 along-fjord stations were used to characterize the seasonality of the density field. The density profiles were used to estimate the internal long-wave phase speed (c) using two approximations: (1) a simple reduced gravity model (RGM) and (2) a continuously stratified model (CSM). No major seasonal changes in c were observed using either approximation (e.g., the CSM yielded 0.73 &lt; c &lt; 0.87 m s−1 for mode 1). The natural internal periods (TN) were estimated using Merian's formula for a simple fjord-like basin and the above phase speeds. Estimated values of TN varied between 2.9 and 3.5 days and were highly consistent with spectral peaks observed in the along-fjord currents and temperature time series. We conclude that these oscillations were forced by the wind stress, despite the moderate wind energy. Wind conditions at the end of winter gave us an excellent opportunity to explore the damping process. The observed damping time (Td) was relatively long (Td = 9.1 days).

Metaproteomics reveals differential modes of metabolic coupling among ubiquitous oxygen minimum zone microbes.

Marine oxygen minimum zones (OMZs) are intrinsic water column features arising from respiratory oxygen demand during organic matter degradation in stratified waters. Currently OMZs are expanding due to global climate change with resulting feedback on marine ecosystem function. Here we use metaproteomics to chart spatial and temporal patterns of gene expression along defined redox gradients in a seasonally stratified fjord to better understand microbial community responses to OMZ expansion. The expression of metabolic pathway components for nitrification, anaerobic ammonium oxidation (anammox), denitrification, and inorganic carbon fixation were differentially expressed across the redoxcline and covaried with distribution patterns of ubiquitous OMZ microbes including Thaumarchaeota, Nitrospina, Nitrospira, Planctomycetes, and SUP05/ARCTIC96BD-19 Gammaproteobacteria. Nitrification and inorganic carbon fixation pathways affiliated with Thaumarchaeota dominated dysoxic waters, and denitrification, sulfur oxidation, and inorganic carbon fixation pathways affiliated with the SUP05 group of nitrate-reducing sulfur oxidizers dominated suboxic and anoxic waters. Nitrifier nitrite oxidation and anammox pathways affiliated with Nirospina, Nitrospira, and Planctomycetes, respectively, also exhibited redox partitioning between dysoxic and suboxic waters. The numerical abundance of SUP05 proteins mediating inorganic carbon fixation under anoxic conditions suggests that SUP05 will become increasingly important in global ocean carbon and nutrient cycling as OMZs expand.

Environmental change in the Limfjord, Denmark (ca 7500–1500 cal yrs BP): a multiproxy study

The Limfjord region of northern Jutland, Denmark, supports a rich archaeological record dating back to the Mesolithic, which documents long-term change in human practices and utilisation of marine resources since approximately 7500 BP. The presence and availability of marine resources in the Limfjord is sensitively regulated by environmental parameters such as salinity, sedimentary regime, nutrient status and primary productivity, but long-term changes in these parameters are currently poorly understood. In this study a multiproxy approach (including sedimentary parameters, diatoms, molluscs, foraminifera, sedimentary pigments, C and O stable isotopes and plant macrofossils) has been adopted to assess environmental change over the period ca 7500–1500 cal yrs BP at Kilen, a coastal fjord (before AD 1856) situated in the Western Limfjord. A diatom-based salinity transfer function based on a pan-Baltic training set has been applied to the fossil diatom dataset for quantitative assessment of salinity change over the study period. This study demonstrates that large-scale shifts in salinity are a common feature of the Limfjord's long-term history and are driven by the level of connection with the North Sea and the Skagerrak respectively, which in turn is likely driven by the complex interplay between climate, sea-level change, current velocity and rates of erosion/sedimentary accretion. Three shifts in state at Kilen are identified over the study period: a deep, periodically stratified fjord with medium–high salinity (and high productivity) between ca 7500–5000 BP, followed by a gradual transition to a shallow benthic system with more oceanic conditions (i.e. higher salinity, lower productivity, slower sedimentary accumulation rate and poorer fossil preservation) after ca 5000 BP and no stratification after ca 4400 BP, and lastly, within this shallow phase, an abrupt shift to brackish conditions around 2000 BP. Environmental–societal interactions are discussed on the basis of the data presented in this study and current environmental hypotheses for cultural change are challenged.

Seasonal variability of Dinophysis spp. and Protoceratium reticulatum associated to lipophilic shellfish toxins in a strongly stratified Chilean fjord

The fine scale vertical distribution of Dinophysis spp. and Protoceratium reticulatum (potential producers of lipophilic shellfish toxins, LSTs) and its relation with LSTs in shellfish was studied in Reloncaví fjord, a strongly stratified system in Southern Chile. Samples were taken over two years from late spring to early autumn (2007–2008 period) and from early spring to late summer (2008–2009 period). Dinophysis spp., in particular Dinophysis acuminata, were always detected, often forming thin layers in the region of the salinity driven pycnocline, with cell maxima for D. acuminata of 28.5×103cellsL−1 in March 2008 and 17.1×103cellsL−1 in November 2008. During the 2008–2009 sampling period, blooms of D. acuminata co-occurred with high densities of cryptophyceans and the ciliate Mesodinium spp. The highest levels of pectenotoxin-2 (PTX-2; 2.2ng L−1) were found in the plankton in February 2009, associated with moderate densities of D. acuminata, Dinophysis tripos and Dinophysis subcircularis (0.1–0.6×103cellsL−1). However, only trace levels of PTX-2 were observed in bivalves at that time. Dinophysistoxin (DTX-1 and DTX-3) levels in bivalves and densities of Dinophysis spp. were not well correlated. Low DTX levels in bivalves observed during a major bloom of D. acuminata in March 2008 suggested that there is a large seasonal intraspecific variability in toxin content of Dinophysis spp. driven by changes in population structure associated with distinct LST toxin profiles in Reloncaví fjord during the study period. A heterogeneous vertical distribution was also observed for P. reticulatum, whose presence was restricted to summer months. A bloom of this species of 2.2×103cellsL−1 at 14m depth in February 2009 was positively correlated with high concentrations of yessotoxins in bivalves (51–496ngg−1) and plankton samples (3.2ngL−1). Our results suggest that a review of monitoring strategies for Dinophysis spp. in strongly stratified fjord systems should be carried out. They also indicate that early warning of LST events based on Dinophysis cell numbers are not reliable for seafood control.

Artificial upwelling to stimulate growth of non-toxic algae in a habitat for mussel farming

Large-scale artificial upwelling was tested as a method to enhance the environmental conditions for the growth of non-toxic algae in a Norwegian fjord (61°0′N, 6°22′E). The experiment was designed to evaluate if nutrient-rich seawater, brought up from below the mixed zone of a stratified fjord to the euphotic zone by air bubbling, would stimulate the growth of non-toxic relative to toxic algae. Pumping 44 m3 min−1 of air at 1 atm through a pipe diffuser submerged at 40 m depth formed a buoyancy flux that lifted 60 m3 s−1 of deep water to the upper 17 m over a period of 21 days. The supply of silicate, inorganic nitrogen and phosphate to the upper 10 m in the fjord increased, and a significant increase in the biomass of non-toxic algae was observed. The upwelling gave an increased growth of the non-toxic dinoflagellates Ceratium furca and C. tripos. After termination of the experiment, the phytoplankton biomass decreased significantly, whereas a distinct increase occurred in the relative biomass of the potentially toxic Dinophysis spp. The result is considered promising when it comes to creating controlled geographical areas with non-toxic food for mussel production.

Temperature-dependent feed requirements in farmed blue mussels (Mytilus edulisL.) estimated from soft tissue growth and oxygen consumption and ammonia-N excretion

Norwegian salmon production doubled from 0.43 million tons in 1999 to 0.86 million tons in 2009, with further growth expected. A considerable amount of the feed used is released into the surrounding waters as respiratory products, faeces and uneaten feed, and there is an increasing concern regarding the potentially negative impacts that this nutrient load may have. One of the major challenges for the sustainable development of salmon aquaculture is therefore to minimize waste discharges that may lead to degradation of the local marine environment. For this purpose, it has been suggested to cultivate extractive and filter feeding species, e.g. seaweed and mussels, close to fish farms in integrated multi-trophic aquaculture (IMTA), thereby contributing to a more ecologically balanced ecosystem approach in marine aquaculture.The primary objectives of this thesis were to investigate whether blue mussels (Mytilus edulis) can incorporate and utilize components of salmon fish feed and faeces particles for growth. The secondary objectives were to assess the ambient conditions for mussel cultivation in the coastal areas of Central Norway, and to test for the possibility of using land-based storage or creating non-toxic areas for storage of mussels at sea to meet a possible increase in mussel production from a development of IMTA in Norway. Mussels cleared salmon feed and faeces particles out of suspension with a high efficiency, suggesting that mussels can remove particulate wastes from salmon farming. In combination with a better growth for mussels fed salmon feed than faeces, a more pronounced incorporation of salmon feed compared to salmon faeces components in mussel tissues indicated that mussels will utilize salmon fish feed more efficiently than faeces particles in an integrated production with salmon.A one-year case study further revealed the incorporation of salmon fish feed in mussel tissues and five months with a higher soft tissue weight of mussels co-produced with salmon compared to control mussels, particularly during autumn and winter when phytoplankton concentrations were low, while control mussels demonstrated a higher soft tissue weight after peak phytoplankton levels in early summer. Mussels at the salmon farm showed a faster growth in length during the spring, while control mussels grew faster during the summer, thus resulting in equal growth rates for the fastest growing mussels co-produced with salmon and control mussels for the entire year. The results suggest that the combined production of mussels and salmon can be seen as a strategy to mitigate environmental effects of particulate nutrient wastes from salmon farming, and to maintain a higher soft tissue content of mussels during autumn and winter.Ambient Chl a concentrations in the coastal areas of Central Norway ranged bellow 2 μg L−1 after the spring bloom and were considered low. Food availability measured as suspended particulate matter (SPM) was, however, consistently above the threshold level of 4 mg L-1 for pseudo-faeces production in mussels of 1 g soft tissue dry matter, with an organic content of 32-44%, and did not appear to restrict mussel growth. SPM may therefore be an important food source to sustain mussel growth when phytoplankton concentrations are low.Temperature-dependent feed requirements were evident from significantly higher oxygen consumption and ammonia-N excretion rates at 14°C compared to 7°C at landbased storage conditions. Minimum feed requirements for the weight maintenance of mussels with 500 mg soft tissue dry matter is estimated at 240 and 570 μg C ind-1 h-1 at 7°C and 14°C, respectively. Mussels kept at land-based storage conditions maintained their soft tissue content and thus a high quality in early summer (May-June) while a significant decrease in soft tissue matter was evident among farmed mussels at sea in the same period. The results suggest that land-based storage can be used for obtaining a continuous mussel production in Norway independent of harvesting problems related to toxic algae blooms and extreme weather.Artificial upwelling in a stratified fjord resulted in an increased nutrient supply to euphotic waters and a correspondingly increase in phytoplankton biomass with a relative reduction of toxic algae. The increase in phytoplankton biomass was mainly represented by non-toxic dinoflagellates, and not diatoms, which was expected from an increased input of silicate from deep water. Nevertheless, the result is promising when it comes to creating controlled geographical areas with non-toxic food for storage of mussels and a continuous mussel production.

Trophic interactions of pelagic crustaceans in Comau Fjord (Chile): their role in the food web structure

Grazing experiments were used to study the trophic interactions among plankton components in a stratified fjord of southern Chile. A strong pycnocline (7–10 m) separating the low-salinity layer from the deeper, high-salinity, marine layer, affected the composition, abundance and distribution of phytoplankton and zooplankton every season. Nanoflagellates averaging 59% were consistently the main components of the plankton biomass in both layers. The microphytoplankton contribution was always higher within the marine layer. Cladocerans and some small copepods showed a vertical gradient with greater abundances in the low-salinity layer, whereas medium-large copepods and euphausiids were more abundant below the pycnocline. Podon leuckarti averaged an ingestion rate of 3.1 µg C ind.−1 day−1, consisting mostly of particles from the nanoplankton size spectrum. Given significant clearance on ciliates and athecate dinoflagellates plus high ingestion rates of heterotrophic nanoflagellates it seems that this species selects for microzooplankton and that heterotrophic components are important in their diet. Calanus australis ingested 15 µg C ind.−1 day−1 on average, covering a wide spectrum of prey sizes (5–65 µm equivalent spherical diameter) and diversity, with active selection for ciliates. The average ingestion rate for Euphausia vallentini was 78.7 µg C ind.−1 day−1, including wide variety of prey sizes but no apparent selectivity. The potential grazing impact (11%) on primary productivity, together with high abundances, feeding selectivity on nanoplankton and production of unconsolidated fecal material indicate that cladocerans could have a double positive feedback effect on the microbial loop, thereby having an important impact by modifying the composition and structure of the planktonic community in Comau Fjord in spring-summer.

Impact of ocean stratification on submarine melting of a major Greenland outlet glacier

AbstractSubmarine melting is an important balance term for tidewater glaciers1,2 and recent observations point to a change in the submarine melt rate as a potential trigger for the widespread acceleration of outlet glaciers in Greenland3-5. Our understanding of the dynamics involved, and hence our ability to interpret past and predict future variability of the Greenland Ice Sheet, however, is severely impeded by the lack of measurements at the ice/ocean interface. To fill this gap, attempts to quantify the submarine melt rate and its variability have relied on a paradigm developed for tidewater glaciers terminating in fjords with shallow sills. In this case, the fjords’ waters are mostly homogeneous and the heat transport to the terminus, and hence the melt rate, is controlled by a single overturning cell in which glacially modified water upwells at the ice edge, driving an inflow at depth and a fresh outflow at the surface1. Greenland’s fjords, however, have deep sills which allow both cold, fresh Arctic and warm, salty Atlantic waters, circulating around Greenland, to reach the ice sheet margin3,6,7. Thus, Greenland’s glaciers flow into strongly stratified fjords and the generic tidewater glacier paradigm is not applicable. Here, using new summer data collected at the margins of Helheim Glacier, East Greenland, we show that melting is driven by both Atlantic and Arctic waters and that the circulation at the ice edge is organized in multiple, overturning cells that arise from their different properties. Multiple cells with different characteristics are also observed in winter, when glacial run off is at a minimum and there is little surface outflow. These results indicate that stratification in the fjord waters has a profound impact on the melting dynamics and suggest that the shape and stability of Greenland’s glaciers are strongly influenced by layering and variability in the Arctic and Atlantic waters.

Flow and Hydraulics near the Sill of Hood Canal, a Strongly Sheared, Continuously Stratified Fjord

Abstract Hood Canal, a long fjord in Washington State, has strong tides but limited deep-water renewal landward of a complex constriction. Tide-resolving hydrographic and velocity observations at the constriction, with a depth-cycling towed body, varied markedly during three consecutive years, partly because of stratification variations. To determine whether hydraulic control is generally important and to interpret observations of lee waves, blocking, and other features, hydraulic criticality is estimated over full tidal cycles for channel wide internal wave modes 1, 2, and 3, at five cross-channel sections, using mode speeds from the extended Taylor–Goldstein equation. These modes were strongly supercritical during most of ebb and flood on the gentle seaward sill face and for part of flood at the base of the steep landward side. Examining local criticality along the thalweg found repeated changes between mode 1 being critical and supercritical approaching the sill crest during flood, unsurprising given local minima and maxima in the cross-sectional area, with the sill crest near a maximum. Density crossing the sill sometimes resembled an overflow with an internal hydraulic control at the sill, followed by a hydraulic jump or lee wave. Long-wave speeds, however, suggest cross waves, particularly along the shallower gentler side, where flow downstream of a large-amplitude wave was uniformly supercritical. Supercritical approaching the sill, peak ebb was critical to mode 1 and supercritical to modes 2 and 3 at the base while forming a sluggish dome of dense water over the sill. Full interpretation exceeds observations and existing theory.