Winter Mixed Depths Research Articles

Giant saltwater inflow in AD 1951 triggered Baltic Sea hypoxia

A marked sedimentological change in subsurface sediments from the entire Baltic Proper, the Baltic Sea, has been previously noted. Our detailed work on a variety of multi‐cores from basin‐wide transects indicates that this sedimentological change was caused by a large shift in environmental conditions during the 1950s. Until the 1950s, the water column was rather weakly stratified and winter‐time convection – although weakened during the post Little Ice Age warming – was still able to ventilate the bottom waters of the Baltic Proper. Therefore, complete sediment sequences only accumulated in calm waters deeper than 150–160 m. High‐resolution benthic foraminiferal records of subsurface sediments obtained along the saline water inflow pathway in combination with historical data indicate that the depositional environment changed drastically owing to the giant saline water inflow in AD 1951. The accompanied sharpening of the halo(pycno)cline triggered a collapse in the ventilation of the basin, resulting in oxygen‐deficient bottom waters. This deficiency, in turn, caused the onset of phosphate release from the sediments, which accelerated primary production. The ventilation collapse also enabled the onset of deposition of organic carbon‐rich sediments also in shallower water areas as calm conditions prevailed up to the modern winter mixing depth (60–70 m). A slight return to Little Ice Age‐type conditions was observed during the late 1980s when temperatures decreased and stratification weakened. These conditions gave rise to a reduction in hypoxic areas and to a bottom‐water ventilation, most pronounced in the north of the so‐called Baltic Sea Klint, a hydrographic and topographic barrier. However, the general environmental conditions essentially have not changed since the 1950s. Remarkably, external (temperature and stratification) in combination with internal factors (e.g. ventilation collapse and phosphate release) were able to change the redox conditions of the Baltic Proper from oxic to hypoxic within less than 10 years.

Inter‐Annual Variability in Phytoplankton and Nutrients in the Gulf of Elat/Aqaba

AbstractThe Gulf of Elat/Aqaba exhibits high inter‐annual variability in mixed layer depth. Observations from the northern Gulf show differences of hundreds of meters in winter mixing depth, which ranges between 300 m in years with shallow mixing and up to 700 m in years with deep mixing. Deep mixing events can occur in two consecutive years or after four consecutive years of shallow mixing. The mixing depth has an effect on the concentration of nutrients and chlorophyll (and other tracers) in the surface and deep water. Using a 3D coupled physical‐ecological model, we studied the effect of shallow versus deep mixing on the processes controlling the phytoplankton bloom and on nutrient accumulation in the deep water. We found that years with deep mixing are characterized by larger spatial variability in surface and integrated chlorophyll concentration during the mixing season. We found that horizontal advection is a dominant contributor for integrated phytoplankton concentration in years with deep mixing in the northern Gulf. Even when mixing was deep and nutrient limitation decreased, light limitation on phytoplankton integrated growth was enhanced in the north compared with the south. In addition, we showed that the nutrient accumulation in the deep water after a year with deep mixing in the northern Gulf was initially affected mostly by physical processes (such as advection and vertical mixing), and less from ecological regeneration, and switched to be dominated by ecological processes alone during the third year after mixing.

An Arctic Strait of Two Halves: The Changing Dynamics of Nutrient Uptake and Limitation Across the Fram Strait

AbstractThe hydrography of the Arctic Seas is being altered by ongoing climate change, with knock‐on effects to nutrient dynamics and primary production. As the major pathway of exchange between the Arctic and the Atlantic, the Fram Strait hosts two distinct water masses in the upper water column, northward flowing warm and saline Atlantic Waters in the east, and southward flowing cold and fresh Polar Surface Water in the west. Here, we assess how physical processes control nutrient dynamics in the Fram Strait using nitrogen isotope data collected during 2016 and 2018. In Atlantic Waters, a weakly stratified water column and a shallow nitracline reduce nitrogen limitation. To the west, in Polar Surface Water, nitrogen limitation is greater because stronger stratification inhibits nutrient resupply from deeper water and lateral nitrate supply from central Arctic waters is low. A historical hindcast simulation of ocean biogeochemistry from 1970 to 2019 corroborates these findings and highlights a strong link between nitrate supply to Atlantic Waters and the depth of winter mixing, which shoaled during the simulation in response to a local reduction in sea‐ice formation. Overall, we find that while the eastern Fram Strait currently experiences seasonal nutrient replenishment and high primary production, the loss of winter sea ice and continued atmospheric warming has the potential to inhibit deep winter mixing and limit primary production in the future.

Differences in Eastern North Pacific Stratification and Their Potential Impact on the Depth of Winter Mixing in CMIP5 Models

AbstractUpper ocean stratification is known to play an important role in the seasonal evolution of the ocean mixed layer depth. This study examines biases in climatological upper ocean stratification in the Coupled Model Intercomparison Project version 5 historical simulations in the North Pacific. To quantify the potential impact of these stratification differences on the depth of winter mixing in two regions (the eastern subarctic and subtropical gyres), regionally averaged August stratification profiles were advanced to March using common atmospheric forcing fields and a single‐column mixed layer model. This approach is supported by correlation between the climate models' March mixed layer depth fields and those predicted by the one‐dimensional model. Substantial observation/model and model/model differences in predicted winter mixed layer depth have broad implications for how the climate models regulate the ocean‐atmosphere interactions that set thermocline water properties and impact nutrient supply to the upper ocean.

Global warming affects nutrient upwelling in deep lakes

Measures to reduce lake phosphorus concentrations have been encouragingly successful in many parts of the world. After significant eutrophication in the twentieth century, nutrient concentrations have declined in many natural settings. In addition to these direct anthropogenic impacts, however, climate change is also altering various processes in lakes. Its effects on lacustrine nutrient budgets remain poorly understood. Here we investigate the total phosphorus (TP) concentrations in the epilimnion of the meromictic Lake Zug under present and future climatic conditions. Results are compared with those of other deep lakes. Data showed that TP transported from the hypolimnion by convective winter mixing was the most important source of TP for the epilimnion, reaching values more than ten times higher than the external input from the catchment. We found a logarithmic relationship between winter mixing depth (WMD) and epilimnetic TP content in spring. Warming climate affects WMD mainly due to its dependence on autumn stratification. Model simulations predict a reduction of average WMD from 78 (current) to 65 m in 2085 assuming IPCC scenario A2. Other scenarios show similar but smaller changes in the future. In scenario A2, climate change is predicted to reduce epilimnetic TP concentrations by up to 24% during warm winters and may consequently introduce significant year-to-year variability in primary productivity.

Effects of climate change on deepwater oxygen and winter mixing in a deep lake (Lake Geneva): Comparing observational findings and modeling

AbstractLow concentrations of dissolved oxygen remain a global concern regarding the ecological health of lakes and reservoirs. In addition to high nutrient loads, climate‐induced changes in lake stratification and mixing represent additional anthropogenic menace resulting in decreased deepwater oxygen levels. The analysis of 43 years of monitoring data from Lake Geneva shows no decreasing trend neither in the areal hypolimnetic mineralization rate nor in the extent of hypoxia. Instead, hypoxic conditions are predominantly controlled by deep mixing in winter and much less by the trophic variations over the past decades. To reproduce winter mixing, the one‐dimensional hydrodynamic model SIMSTRAT was specially adapted to deep lakes and run for several climate scenarios. The simulations predicted a decrease in the maximum winter mixing depth from an average of ∼172 m for 1981–2012 to ∼136 m and ∼127 m in response to predicted atmospheric temperatures between 2045–2076 and 2070–2101 according to Intergovernmental Panel on Climate Change scenarios. Concurrently, events with complete homogenization of temperature and oxygen in winter will decrease by ∼50%. Consequently, the hypolimnetic oxygen concentrations will significantly decrease. These results demonstrate that changes in deep mixing can have stronger impact than eutrophication on the deepwater oxygen levels of oligomictic lakes.

The North Atlantic subpolar circulation in an eddy-resolving global ocean model

The subpolar North Atlantic represents a key region for global climate, but most numerical models still have well-described limitations in correctly simulating the local circulation patterns. Here, we present the analysis of a 30-year run with a global eddy-resolving (1/12°) version of the NEMO ocean model. Compared to the 1° and 1/4° equivalent versions, this simulation more realistically represents the shape of the Subpolar Gyre, the position of the North Atlantic Current, and the Gulf Stream separation. Other key improvements are found in the representation of boundary currents, multi-year variability of temperature and depth of winter mixing in the Labrador Sea, and the transport of overflows at the Greenland–Scotland Ridge. However, the salinity, stratification and mean depth of winter mixing in the Labrador Sea, and the density and depth of overflow water south of the sill, still present challenges to the model. This simulation also provides further insight into the spatio-temporal development of the warming event observed in the Subpolar Gyre in the mid 1990s, which appears to coincide with a phase of increased eddy activity in the southernmost part of the gyre. This may have provided a gateway through which heat would have propagated into the gyre's interior.

Annual cycles of ecological disturbance and recovery underlying the subarctic Atlantic spring plankton bloom

Satellite measurements allow global assessments of phytoplankton concentrations and, from observed temporal changes in biomass, direct access to net biomass accumulation rates (r). For the subarctic Atlantic basin, analysis of annual cycles in r reveals that initiation of the annual blooming phase does not occur in spring after stratification surpasses a critical threshold but rather occurs in early winter when growth conditions for phytoplankton are deteriorating. This finding has been confirmed with in situ profiling float data. The objective of the current study was to test whether satellite‐based annual cycles in r are reproduced by the Biogeochemical Element Cycling–Community Climate System Model and, if so, to use the additional ecosystem properties resolved by the model to better understand factors controlling phytoplankton blooms. We find that the model gives a similar early onset time for the blooming phase, that this initiation is largely due to the physical disruption of phytoplankton‐grazer interactions during mixed layer deepening, and that parallel increases in phytoplankton‐specific division and loss rates during spring maintain the subtle disruption in food web equilibrium that ultimately yields the spring bloom climax. The link between winter mixing and bloom dynamics is illustrated by contrasting annual plankton cycles between regions with deeper and shallower mixing. We show that maximum water column inventories of phytoplankton vary in proportion to maximum winter mixing depth, implying that future reductions in winter mixing may dampen plankton cycles in the subarctic Atlantic. We propose that ecosystem disturbance‐recovery sequences are a unifying property of global ocean plankton blooms.

What controls primary production in the Arctic Ocean? Results from an intercomparison of five general circulation models with biogeochemistry

As a part of Arctic Ocean Intercomparison Project, results from five coupled physical and biological ocean models were compared for the Arctic domain, defined here as north of 66.6°N. The global and regional (Arctic Ocean (AO)–only) models included in the intercomparison show similar features in terms of the distribution of present‐day water column–integrated primary production and are broadly in agreement with in situ and satellite‐derived data. However, the physical factors controlling this distribution differ between the models. The intercomparison between models finds substantial variation in the depth of winter mixing, one of the main mechanisms supplying inorganic nutrients over the majority of the AO. Although all models manifest similar level of light limitation owing to general agreement on the ice distribution, the amount of nutrients available for plankton utilization is different between models. Thus the participating models disagree on a fundamental question: which factor, light or nutrients, controls present‐day Arctic productivity. These differences between models may not be detrimental in determining present‐day AO primary production since both light and nutrient limitation are tightly coupled to the presence of sea ice. Essentially, as long as at least one of the two limiting factors is reproduced correctly, simulated total primary production will be close to that observed. However, if the retreat of Arctic sea ice continues into the future as expected, a decoupling between sea ice and nutrient limitation will occur, and the predictive capabilities of the models may potentially diminish unless more effort is spent on verifying the mechanisms of nutrient supply. Our study once again emphasizes the importance of a realistic representation of ocean physics, in particular vertical mixing, as a necessary foundation for ecosystem modeling and predictions.

Modeling the formation and fate of the near‐surface temperature maximum in the Canadian Basin of the Arctic Ocean

A numerical model is used to investigate the time and space extent of the near‐surface temperature maximum (NSTM) of the Canadian Basin of the Arctic Ocean over the years 2000–2009. The NSTM is formed from local summertime absorption of solar radiation which, in some circumstances, descends through the fall and early winter to form a warm subsurface layer just below the winter mixed layer. We find that winter survival of this layer is confined largely to the Beaufort Gyre of the Canadian Basin, where Ekman convergence and downwelling push the summer warm layer down below the winter mixing depth. In recent years, summer stratification has increased, downwelling has accelerated, and the NSTM has warmed as the sea ice cover in the Beaufort Gyre has thinned. The result is a strengthening NSTM which contained enough heat by the end of winter 2007/2008 to melt about 20 cm of sea ice. Northwest of Alaska the model also simulates a second, deeper temperature maximum layer that forms from advection of saltier summer Pacific water. However, this layer is difficult to adequately resolve and maintain given the model's resolution.

Subduction over the Southern Indian Ocean in a High-Resolution Atmosphere–Ocean Coupled Model

Abstract This study examines the subduction of the Subantarctic Mode Water in the Indian Ocean in an ocean–atmosphere coupled model in which the ocean component is eddy permitting. The purpose is to assess how sensitive the simulated mode water is to the horizontal resolution in the ocean by comparing with a coarse-resolution ocean coupled model. Subduction of water mass is principally set by the depth of the winter mixed layer. It is found that the path of the Agulhas Current system in the model with an eddy-permitting ocean is different from that with a coarse-resolution ocean. This results in a greater surface heat loss over the Agulhas Return Current and a deeper winter mixed layer downstream in the eddy-permitting ocean coupled model. The winter mixed layer depth in the eddy-permitting ocean compares well to the observations, whereas the winter mixed layer depth in the coarse-resolution ocean coupled model is too shallow and has the wrong spatial structure. To quantify the impacts of different winter mixed depths on the subduction, a way to diagnose local subduction is proposed that includes eddy subduction. It shows that the subduction in the eddy-permitting model is closer to the observations in terms of the magnitudes and the locations. Eddies in the eddy-permitting ocean are found to 1) increase stratification and thus oppose the densification by northward Ekman flow and 2) increase subduction locally. These effects of eddies are not well reproduced by the eddy parameterization in the coarse-resolution ocean coupled model.

Simulated response of North Pacific Mode Waters to global warming

This study investigates the response of the Mode Waters in the North Pacific to global warming based on a set of Intergovernmental Panel on Climate Change (IPCC) Fourth Assessment Report (AR4) models. Solutions between a present‐day climate and a future, warmer climate are compared. Under the warmer climate scenario, the Mode Waters are produced on lighter isopycnal surfaces and are significantly weakened in terms of their formation and evolution. These changes are due to a more stratified upper ocean and thus a shoaling of the winter mixing depth resulting mainly from a reduction of the ocean‐to‐atmosphere heat loss over the subtropical region. The basin‐wide wind stress may adjust the Mode Waters indirectly through its impact on the surface heat flux and subduction process.

Southern Ocean deep-water carbon export enhanced by natural iron fertilization

The addition of iron to high-nutrient, low-chlorophyll regions induces phytoplankton blooms that take up carbon. Carbon export from the surface layer and, in particular, the ability of the ocean and sediments to sequester carbon for many years remains, however, poorly quantified. Here we report data from the CROZEX experiment in the Southern Ocean, which was conducted to test the hypothesis that the observed north-south gradient in phytoplankton concentrations in the vicinity of the Crozet Islands is induced by natural iron fertilization that results in enhanced organic carbon flux to the deep ocean. We report annual particulate carbon fluxes out of the surface layer, at three kilometres below the ocean surface and to the ocean floor. We find that carbon fluxes from a highly productive, naturally iron-fertilized region of the sub-Antarctic Southern Ocean are two to three times larger than the carbon fluxes from an adjacent high-nutrient, low-chlorophyll area not fertilized by iron. Our findings support the hypothesis that increased iron supply to the glacial sub-Antarctic may have directly enhanced carbon export to the deep ocean. The CROZEX sequestration efficiency (the amount of carbon sequestered below the depth of winter mixing for a given iron supply) of 8,600 mol mol(-1) was 18 times greater than that of a phytoplankton bloom induced artificially by adding iron, but 77 times smaller than that of another bloom initiated, like CROZEX, by a natural supply of iron. Large losses of purposefully added iron can explain the lower efficiency of the induced bloom(6). The discrepancy between the blooms naturally supplied with iron may result in part from an underestimate of horizontal iron supply.

Lake Van, Eastern Anatolia, Hydrochemistry and History

Saline, 450-m-deep Lake Van (Eastern Anatolia, Turkey) is, with 576 km3, the third largest closed lake on Earth and its largest soda lake. In 1989 and 1990, we investigated the hydrochemistry of the lake’s water column and of the tributary rivers. We also cored the Postglacial sediment column at various water depths. The sediment is varved throughout, allowing precise dating back to ca. 15 ka BP. Furthermore, lake terrace sediments provided a 606-year-long floating chronology of the Glacial high-stand of the lake dating to 21 cal. ka BP. The sediments were investigated for their general mineralogical composition, important geochemical parameters, and pore water chemistry as well. These data allow reconstructing the history of the lake level that has seen several regressions and transgressions since the high-stand at the end of the Last Glacial Maximum. Today, the lake is very alkaline, highly supersaturated with Ca-carbonate and has a salt content of about 22 g kg−1. In summer, the warmer epilimnion is diluted with river water and forms a stable surface layer. Depth of winter mixing differs from year to year but during time of investigation the lake was oxygenated down to its bottom. In general, the lake is characterized by an Na–CO3–Cl–(SO4)-chemistry that evolved from the continuous loss of calcium as carbonate and magnesium in the form of Mg-silica-rich mineral phases. The Mg cycle is closely related to that of silica which in turn is governed by the production and dissolution of diatoms as the dominant phytoplankton species in Lake Van. In addition to Ca and Mg, a mass balance approach based on the recent lake chemistry and river influx suggests a fractional loss of potassium, sodium, sulfur, and carbon in comparison to chloride in the compositional history of Lake Van. Within the last 3 ka, minor lake level changes seem to control the frequency of deep water renewal, the depth of stratification, and the redox state of the hypolimnion. Former major regressions are marked by Mg-carbonate occurrences in the otherwise Ca-carbonate dominated sediment record. Pore water data suggest that, subsequent to the major regression culminating at 10.7 ka BP, a brine layer formed in the deep basin that existed for about 7 ka. Final overturn of the lake, triggered by the last major regression starting at about 3.5 ka BP, may partly account for the relative depletion in sulfur and carbon due to rapid loss of accumulated gases. An even stronger desiccation phase is proposed for the time span between about 20 and 15 ka BP following the LGM, during which major salts could have been lost by precipitation of Na-carbonates and Na-sulfates.

Large-scale physical controls on phytoplankton growth in the Irminger Sea, Part II: Model study of the physical and meteorological preconditioning

The upper water column in the Irminger Sea is characterized by cold fresh arctic and subarctic waters and warm saline North Atlantic waters. In this study the local physical and meteorological preconditioning of the phytoplankton development over an annual cycle in the upper water column in four physical zones of the Irminger Sea is investigated. Data from four cruises of the UK's Marine Productivity programme are combined with results from a coupled biological–physical nitrogen–phytoplankton–zooplankton–detritus model run using realistic forcing. The observations and model predictions are compared and analyzed to identify the key parameters and processes which determine the observed heterogeneity in biological production in the Irminger Sea. The simulations show differences in the onset of the bloom, in the time of the occurrence of the maximum phytoplankton biomass and in the length of the bloom between the zones. The longest phytoplankton bloom of 90 days duration was predicted for the East Greenland Current of Atlantic origin zone. In contrast, for the Central Irminger Sea zone a phytoplankton bloom with a start at the beginning of May and the shortest duration of only 70 days was simulated. The latest onset of the phytoplankton bloom in mid May and the latest occurrence of the maximum biomass (end of July) were predicted for the Northern Irminger Current zone. Here the bloom lasted for 80 days. In contrast the phytoplankton bloom in the Southern Irminger Current zone started at the same time as in Central Irminger Sea, but peaked end of June and lasted for 80 days. For all four zones relatively low daily (0.3–0.5 g C m − 2 d − 1 ) and annual primary production was simulated, ranging between 35.6 g C m − 2 y − 1 in the East Greenland Current of Atlantic origin zone and 45.6 g C m − 2 y − 1 in the Northern Irminger Current zone. The model successfully simulated the observed regional and spatial differences in terms of the maximum depth of winter mixing, the onset of stratification and the development of the seasonal thermocline, and the differences in biological characteristics between the zones. The initial properties of the water column and the seasonal cycle of physical and meteorological forcing in each of the zones are responsible for the observed differences during the Marine Productivity cruises. The timing of the transition from mixing to stratification regime, and the different prevailing light levels in each zone are identified as the crucial processes/parameters for the understanding of the dynamics of the pelagic ecosystem in the Irminger Sea.

Supply and demand of nutrients and dissolved organic matter at and across the NW European shelf break in relation to hydrography and biogeochemical activity

As part of the OMEX I project, nutrient determinations were made on 17 cruises in the region of the Goban Spur and La Chapelle Bank between 46 and 51°N, in all seasons of the year, between 1993–1995. Over this period no change was detectable in the structure of the water masses below the deep winter mixed layer. The N : P (dissolved nitrate-to-phosphate) ratio changed from 16 at 100-m depth to less than 15 at 3300-m depth. At intermediate depths nutrient and oxygen data indicate the presence of Mediterranean Outflow water overlying Labrador Sea Water at its most eastern extension. Estimated maximum levels of production in the spring bloom are the total N-limited new primary production equivalent between 24 and 41 gC m−2, the equivalent maximum diatom production is 11 gC m−2. Measurements during the spring bloom suggest a conversion factor of 1 μM nitrate to 1 μg l−1 chlorophyll, at the shelf break, which is consistent with other recent measurements in European shelf seawaters. Sediment trap data suggest that 80% (5.4 g m−2) of the opal produced in the spring bloom dissolved before reaching the sediment trap at 600 m. A comparison of the winter and summer profiles for dissolved silicon suggests a similar dissolution of 9±3 g opal m−2 above 300-m depth. Measurements of dissolved organic carbon (DOC) in September 1994 show an enrichment of 7 μM-C above the seasonal thermocline relative to the winter values (52±4 μM). In winter dissolved organic nitrogen represents 40% of the pool of total dissolved nitrogen. There is no consistent evidence of an increase in the concentration of DON during summer. Measurements of nitrate in surface waters in January 1994 show that concentrations off-shelf vary with the temperature of the water and are related to the depth of winter mixing. Mixing in surface waters is discontinuous at the shelf break, demonstrating the degree to which exchange across the shelf break is limited even in winter. OMEX winter measurements of nitrate concentrations can be used to estimate the flow of water across the shelf break that would be required to maintain the nitrogen balance in the North Sea at a steady state. The estimate is 0.6 Sv (1 Sv=106 m3 s−1), which is similar to an earlier estimate of a total flow of 1.7 Sv based on salt budgets (cf. Huthnance, Deutsche Hydrographische Zeitschrift, 49 (1997) 153).

Particle flux across the mid-European continental margin

Results are presented from particle flux studies using sediment trap and current meter moorings along a transect at the European continental margin at 49°N within the EU-funded Ocean Margin Exchange (OMEX) project. Two moorings were placed, at the mid- and outer slope in water depths of 1500 and 3660 m, with traps at 600 and 1050 m and at 580, 1440 and 3220 m, respectively. Residual currents at the mid-slope follow the slope contour, whereas seasonal off-slope flow was registered at the outer slope. At 600 m on the slope fluxes are similar to those in the abyssal North Atlantic. The flux of all components (bulk dry weight, particulate organic and inorganic carbon, lithogenic matter and opal) increased with water depth. Highest fluxes were recorded at 1440 m at the outer slope, where off-slope residual currents mediate particle export. The injection of biogenic and lithogenic particles below the depth of winter mixing results in the export of particles from shallower waters. Calculated lateral fluxes of particulate organic carbon exceed the primary flux by over a factor of 2 at 1440 m on the outer slope. Estimated lateral fluxes of suspended particulate matter in the water column and intermediate nepheloid layers at the outer slope are potentially large compared to sinking fluxes measured by sediment traps. A comparison is made of particle flux at three continental margin sites and two sites in the adjacent open North Atlantic, from which it is seen that bulk and organic matter flux increases exponentially with proximity to the shelf break. The percentage contribution of particulate organic carbon to biogenic fluxes increases from a mean of 5.7% in the abyssal N. Atlantic to 13.9% at the continental margins.

The critical pressures of gas vesicles in Planktorhrix rubescens in relation tothe depth of winter mixing in Lake Zürich, Switzerland

The vertical distribution of the cyanobacterium Planktothrix (Oscillatoria) rubescens in Lake Zurich was investigated from March 1993 to June 1995 by collecting filaments on filters and measuring them by epifluorescence microscopy and computer image analysis. The initial population, which began to stratify in April, decreased by up to 99% by June. During the summer, the population peaked at depths of 8—15 m; it reached a maximum areal filament-volume concentration of -60 cm3 m~2 of lake surface in early September and was then entrained in the deepening surface layer. It became mixed progressively deeper, to the lake bottom in the cold winter of 1993-94, but less completely in the milder winter of 1994-95. Most of the filaments remained viable during the winter. At the end of the mild winter of 1994-5, 70% of filaments in the water column retained buoyancy, but after the cold winter of 1996-7 only 22% were buoyant. Few remained buoyant below 80 m, where the hydrostatic pressure caused gas vesicle collapse. The proportion that remain buoyant decreases with the depth and duration of winter mixing, and increases with the critical collapse pressure (pc) of the gas vesicles, which provide buoyancy. Strains of P.rubescens isolated from Lake Zurich differed in mean pc of their gas vesicles, from 0.9 to 1.1 MPa, the highest values in freshwater cyanobacteria. Allowing for a turgor pressure of 0.2 MPa, these strains would remain buoyant at depths down to 70 and 90 m, respectively. Natural selection for gas vesicles of high pc will operate by increasing the proportion of filaments that remain buoyant in the upper parts of the water column after circulation to various depths during the winter because only buoyant filaments will form the inoculum for the following season.

The influence of intermediate waters on the stability of the eastern North Atlantic

Changes in the quantities and proportions of various intermediate water masses in the eastern North Atlantic have important consequences for the climate of the region. Subarctic Intermediate Water (SAIW) is mostly found within the subpolar gyre and west of 20°W. Small quantities of this water mass, however, are found on the 20°W meridian at the southern end of the Rockall Channel and are observed to influence the vertical structure of the water column. Typically the comparatively fresh SAIW is a highly stratified water mass, and its subduction into the eastern North Atlantic over more saline Mediterranean waters results in a stable layer at intermediate depths that restricts the maximum depth of winter mixing by up to 150 m compared to stations north-west of the Rockall-Hatton Bank. Analysis of Ocean Weather Ship (OWS) data from this region shows temporal variability of the vertical structure during the 1960s and 1970s. During the 1970s the “Great Salinity Anomaly” (Dickson et al., 1988) resulted in increasing quantities of comparatively fresh SAIW being present. This in turn resulted in a weakening of the stable layer instead of the expected strengthening. Changes in the stratification of the SAIW source are implied. By influencing the depth of maximum winter mixing, changes in the properties of the intermediate water masses of the eastern North Atlantic have more immediate consequences for ocean-atmosphere heat exchanges west of Europe than do longer term changes to the source waters of Labrador Sea Water (LSW) or Norwegian Sea Deep Water (NSDW). Such variations in the intermediate water column structure should therefore be taken into account in ocean climate models.

The gas vesicles, buoyancy and vertical distribution of cyanobacteria in the Baltic Sea

The mean pressures required to collapse gas vesicles in turgid cells of cyanobacteria from the Baltic Sea were 0·91 MPa (9·1 bar) in Aphanizomenon flos-aquae, 0·83 MPa in Nodularia sp. collected from the main deep basins and 0·34 MPa in Nodularia from shallower coastal regions. The gas vesicles were strong enough to withstand the depth of winter mixing, down to the permanent halocline (60 m in the Bornholm Sea, 90 m in the Eastern Gotland Sea) or to the sea bottom (30 m or less in the shallow Arkona Sea and Mecklenburg Bight). The cyanobacteria had low cell turgor pressures, within the range 0·08–0·18 MPa. The colonies were highly buoyant: the Aphanizomenon colonies floated up at a mean velocity of 22 m per day and the Nodularia colonies at 36 m per day. The colonies remained floating when up to half of the gas vesicles had been collapsed. In summer the cyanobacteria were mostly restricted to the water above the thermocline and in calm conditions their concentration increased towards the top of the water ...