Large-scale physical controls on phytoplankton growth in the Irminger Sea Part I: Hydrographic zones, mixing and stratification

Hydrographic surveys in three consecutive seasons in the Irminger Sea in 2001/2002 have revealed six physical regimes (zones) in which different surface mixing and spring re-stratification processes dominate. They are the South Irminger Current, the North Irminger Current, the Central Irminger Sea, the Polar-origin East Greenland Current, the Atlantic-origin East Greenland Current and the Reykjanes Ridge. The variations in restratification processes in particular have significant implications for the timing of shallow spring mixed layer development and therefore the timing and strength of the spring bloom. The relative roles of heat and freshwater in controlling re-stratification are examined for each hydrographic zone, and it is shown that the simplest concept of solar warming generating spring stratification is appropriate for the Irminger Current and the central Irminger Sea. However in the East Greenland Current and the Reykjanes Ridge zones, the springtime arrival of fresh or saline water at the surface dominates re-stratification and generates the earliest and strongest spring blooms of the region. In the cool fresh centre of the Irminger Sea the relatively low chlorophyll-a throughout the year cannot be wholly explained by stratification or nutrient concentrations. Details of the annual cycle in temperature, salinity, chlorophyll-a and nutrients are presented for each hydrographic zone.     

An eddy-permitting, coupled ecosystem-circulation model of the Arabian Sea: comparison with observations

A nitrogen-based, pelagic ecosystem model has been coupled with an eddy-permitting ocean general circulation model of the Arabian Sea, and the results are compared with observations. The seasonal variability simulated by the model is in good agreement with observations: during the southwest monsoon season, phytoplankton increases in the western Arabian Sea due to upwelling along the coast; during the northeast monsoon season, phytoplankton abundance is large in the northern Arabian Sea because of the enhanced nitrate entrained by relatively deep vertical mixing. Two major differences are, however, found in the basin-wide comparison between model results and observations: an unrealistic nitrate maximum in the subsurface layer of the northern Arabian Sea and too low primary production in oligotrophic regimes. The former may be attributed to the lack of denitrification in the model. Possible causes for the latter include the present model's underestimation of fast nutrient recycling, the neglect of carbon fixation decoupled from nitrogen uptake and of nitrogen fixation, and inadequate nitrate entrainment by mixed layer deepening. The rate at which simulated nitrate increases in the northern Arabian Sea is 11–24 TgN/year, and should correspond to the denitrification rate integrated over the northern Arabian Sea assuming that the loss of nitrogen through denitrification is balanced by advective input. The model does not reproduce the observed phytoplankton bloom in the late southwest monsoon season. Possible causes are that the mixed layer may be too shallow in summer and that the horizontal transport of nitrate from the coast of Oman may be too weak. Sensitivity experiments demonstrate a strong dependence of the simulated primary productivity on the vertical mixing scheme and on the inclusion of a fast recycling loop in the ecosystem model.     

Convection and the seeding of the North Atlantic bloom

Observations of vertical velocities in deep wintertime mixed layers using neutrally buoyant floats show that the convectively driven vertical velocities, roughly 1000 m per day, greatly exceed the sinking velocities of phytoplankton, 10 m or less per day. These velocities mix plankton effectively and uniformly across the convective layer and are therefore capable of returning those that have sunk to depth back into the euphotic zone. This mechanism cycles cells through the surface layer during the winter and provides a seed population for the spring bloom. A simple model of this mechanism applied to immortal phytoplankton in the subpolar Labrador Sea predicts that the seed population in early spring will be a few percent of the fall concentration if the plankton sink more slowly than the mean rate at which the surface well-mixed layer grows over the winter. Plankton that sink faster than this will mostly sink into the abyss with only a minute fraction remaining by spring. The shallower mixed layers of mid-latitudes are predicted to be much less effective at maintaining a seed population over the winter, limiting the ability of rapidly sinking cells to survive the winter.     

Ecophysiological growth characteristics and modeling of the onset of the spring bloom in the Baltic Sea

The onset of spring bloom in temperate areas is a transition period where the low productive, winter phytoplankton community is transformed into a high productive spring community. Downwelling irradiance, mixing depth and the ability of the phytoplankton community to utilize the light, are key parameters determining the timing of the onset of the spring bloom. Knowing these parameters would thus provide tools for modeling the spring bloom and enhance our knowledge of ecophysiological processes during this period.Our main objective with this study was to provide data for the growth characteristics of some key species forming the spring bloom in the Gulf of Finland, and to apply those results in a simple dynamic model for the onset of the spring bloom, in order to test if the timing of the spring bloom predicted by the models corresponds to field observations. We investigated the photosynthetic characteristics of three diatoms and two dinoflagellates (Chaetoceros wighamii, Melosira arctica, Thalassiosira baltica, Scrippsiella hangoei and Woloszynskia halophila), at low temperatures (4–5 °C). All of these species are common during spring bloom in the Baltic Sea.Cultures of these species were acclimated to different irradiance regimes prior to measurements of photosynthesis, respiration, pigment concentration and light absorption. We did not find a positive relationship between respiration and growth rate, and we hypothesize that this relationship, which is well established at higher temperatures, is negligible or absent at low temperatures (< 10 °C). Photosynthetic maximum (P_m), and maximum light utilization coefficient (α) was lowest and respiration (R) highest in the dinoflagellates.We made a model of the onset of the spring bloom in the western part of Gulf of Finland, using the obtained data together with monitoring data of mixing depth and water transparency from this area. Model results were compared to field observations of chlorophyll-a (Chl-a) concentration. There was a good agreement between the model predictions and the observed onset of the spring bloom for the diatoms. S. hangoei, however, was not able to reach positive production in the model, and W. halophila had the similar growth characteristics as S. hangoei. Consequently, these species must have other competition strategies enabling them to exist and grow during spring bloom.     

Seasonal variation of riverine nutrient inputs in the northern Bay of Biscay (France), and patterns of marine phytoplankton response

Seasonal variations in nutrient inputs are described for the main rivers (Loire and Vilaine) flowing into the northern Bay of Biscay. The river plumes are high in N/P ratio in late winter and spring, but not in the inner plume during the summer. Conservative behavior results in most nutrients entering the estuary and eventually reaching the coastal zone. Temporal and spatial aspects of phytoplankton growth and nutrient uptake in the northern Bay of Biscay distinguish the central area of salinity 34 from the plume area. The first diatom bloom appears offshore in late winter, at the edge of the river plumes, taking advantage of haline stratification and anticyclonic “weather windows.” In spring, when the central area of the northern shelf is phosphorus-limited, small cells predominate in the phytoplankton community and compete with bacteria for both mineral and organic phosphorus. At that period, river plumes are less extensive than in winter, but local nutrient enrichment at the river mouth allows diatom growth. In summer, phytoplankton become nitrogen-limited in the river plumes; the central area of the shelf is occupied by small forms of phytoplankton, which are located on the thermocline and use predominantly regenerated nutrients.     

Gulf of Gdansk spring bloom physical, bio-optical, biological modelling and contact data assimilation

In this study we propose a model of phytoplankton population dynamics in the marine ecosystem, which includes physical, biological and bio-optical parts. As an example we simulate the abnormal 1993 Gulf of Gdansk spring bloom, when extremely high chlorophyll concentrations were observed. For the one-dimensional model we use two different methods of contact chlorophyll observation assimilation to fit a model of “in situ” data. The results are compared with two-dimensional ecosystem modelling based on a barotropic model of wind-driven circulation without assimilation.     

Seasonal and interannual variation of physical and biological processes during 1994–2001 in the Sea of Japan/East Sea: A three-dimensional physical–biogeochemical modeling study

A Pacific basin-wide physical–biogeochemical model has been used to investigate the seasonal and interannual variation of physical and biological fields with analyses focusing on the Sea of Japan/East Sea (JES). The physical model is based on the Regional Ocean Model System (ROMS), and the biogeochemical model is based on the Carbon, Si(OH)₄, Nitrogen Ecosystem (CoSiNE) model. The coupled ROMS–CoSiNE model is forced with the daily air–sea fluxes derived from the National Centers for Environmental Prediction (NCEP) and the National Center for Atmospheric Research (NCAR) reanalysis for the period of 1994 to 2001, and the model results are used to evaluate climate impact on nutrient transport in Mixed Layer Depth (MLD) and phytoplankton spring bloom dynamics in the JES.The model reproduces several key features of sea surface temperature (SST) and surface currents, which are consistent with the previous modeling and observational results in the JES. The calculated volume transports through the three major straits show that the Korea Strait (KS) dominates the inflow to the JES with 2.46 Sv annually, and the Tsugaru Strait (TS) and the Soya Strait (SS) are major outflows with 1.85 Sv and 0.64 Sv, respectively. Domain-averaged phytoplankton biomass in the JES reaches its spring peak 1.8 mmol N m^− 3 in May and shows a relatively weak autumn increase in November. Strong summer stratification and intense consumption of nitrate by phytoplankton during the spring result in very low nitrate concentration at the upper layer, which limits phytoplankton growth in the JES during the summer. On the other hand, the higher grazer abundance likely contributes to the strong suppression of phytoplankton biomass after the spring bloom in the JES. The model results show strong interannual variability of SST, nutrients, and phytoplankton biomass with sudden changes in 1998, which correspond to large-scale changes of the Pacific Decadal Oscillation (PDO). Regional comparisons of interannual variations in springtime were made for the southern and northern JES. Variations of nutrients and phytoplankton biomass related to the PDO warm/cold phase changes were detected in both the southern and northern JES, and there were regional differences with respect to the mechanisms and timing. During the warm PDO, the nutrients integrated in the MLD increased in the south and decreased in the north in winter. Conversely, during the cold PDO, the nutrients integrated in the MLD decreased in the south and increased in the north. Wind divergence/convergence likely drives the differences in the southern and northern regions when northerly and northwesterly monsoon dominates in winter in the JES. Subjected to the nutrient change, the growth of phytoplankton biomass appears to be limited neither by nutrient nor by light consistently both in the southern and northern regions. Namely, the JES is at the transition zone of the lower trophic-level ecosystem between light-limited and nutrient-limited zones.     

Numerical experiments on the prediction of sea surface temperature anomalies in the Gulf of Mexico

The conservation of thermal energy equation applied to the mixed layer of the ocean, has been used to predict the sea surface temperature anomalies (SSTA) and the month-to-month changes in the Gulf of Mexico. The model includes the horizontal transport of heat by mean ocean currents and by turbulent eddies, as well as the heating by short and long wave radiation, evaporation and sensible heat given off to the atmosphere. A comparative study is carried out on the relative importance of the heating and transport terms. An objective verification of the skill of the predictions is presented for each season and for the whole period from March 1986 to February 1987. The predictions using only the heating terms have some skill over the control predictions (persistence and return to normal). The skill is substantially increased when the horizontal transport of heat by turbulent mixing is included in the model. The incorporation in the model of the Ekman wind drift current anomalies computed from the anomalous surface geostrophic wind improves appreciably the skill of the predictions in winter and fall.The mixed layer depth computed using the Kraus and Turner theory with dissipation, shows that the depths in summer and fall are shallower than in spring and winter. The effect of the shallow mixed layer depth in the model becomes apparent in summer and fall, improving the skill of the predictions in these seasons, with respect to the skill obtained using a constant mixed layer depth of 60 m.The incorporation in the model of the cooling in the mixed layer by turbulent entrainment of colder water from the thermocline, does not improve in an appreciable way the average skill of the predictions.     

Nutrient cycling and primary production in the marine systems of the Arctic and Antarctic

Primary production events in both the Arctic and the Antarctic are highly localized. Carbon-14 incubations that did not account for this caused antarctic primary production estimates to be revised too far downwards from the historic view of high productivity. The primary production regime in the Arctic is even more heterogeneous than in the Antarctic. Arctic primary production rates are in the process of being revised upwards because of a better spatial and temporal distribution of incubation experiments and a re-awakening of interest in estimating new production from the distribution of chemical variables. Similarly, recent examination of temporal changes in nitrate concentrations and recognition of the importance of ice-edge blooms has caused antarctic primary productivity to be revised upwards. In both the Arctic and the Antarctic, the ratio of “new” to total primary production is high, and neglect of this fact can lead to an underestimation of the potential that these regions have for influencing global cycles of bioactive chemicals. Some recent data on temporal changes in nitrate from Fram Strait emphasize the poor state of our knowledge by suggesting an unexpectedly high “new” production rate of 1 g C m⁻² d⁻¹ for a 35 day experiment that encountered an early Phaeocystis bloom. Chemical distributions suggest that new production over the shelf seas that border the Polar Basin is about 50 g Cm⁻² yr⁻¹.The shelves in the Arctic Ocean's marginal and adjacent seas comprise 25% of the total global continental shelf. These extensive shallow regions have much higher rates of primary production than the Polar Basin and may be globally significant sites of denitrification. Globally significant silica deposition could occur on these shelves or on the adjacent slopes.Because of the differences in geomorphology and stratification, global warming is likely to increase primary production in the Arctic and will probably decrease antarctic primary production.In addition to sharing high ratios of “new” to total primary production, high ammonium concentrations occur in the Arctic and Antarctic. It is possible that these accumulations arise from a strong repression of nitrification at low temperatures.     

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

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

Biological effects of dilution of Dead Sea brine with seawater: implications for the planning of the Red Sea–Dead Sea “Peace Conduit”

The Dead Sea is a severely disturbed ecosystem. Its water level has been decreasing at a rate of nearly 1 m per year during the last decade due to anthropogenic intervention in its water balance. Since the peace treaty between Israel and Jordan was established in 1994, a proposal for the construction of a water carrier, the “Peace Conduit,” between the Gulf of Aqaba (Red Sea) and the Dead Sea is being investigated. This water carrier is intended to mitigate damaging processes that currently occur in the Dead Sea and its surrounding area. The difference in elevation of about 416 m could be exploited for seawater desalination by reverse osmosis. To examine the possible effects of the mixing of Dead Sea brines with seawater and seawater concentrates on the Dead Sea as an ecosystem, we have set up simulation experiments under field conditions in experimental ponds at Sedom, in which Dead Sea water was diluted with Red Sea water. The main components of the Dead Sea biota are the unicellular green alga Dunaliella and several types of red halophilic Archaea. Phosphate is the limiting inorganic nutrient. Massive Dunaliella blooms developed, accompanied by dense communities of red halophilic Archaea, in some of the experimental ponds, imparting a brown-red coloration to the brines. The extent of biological development depended on the extent of dilution and on phosphate availability. The results of the simulation experiments show that biological phenomena and their impact on the Dead Sea ecosystem should be taken into consideration during the planning of the “Peace Conduit.”     

Simulations of biological production in the Rhodes and Ionian basins of the eastern Mediterranean

The biological production characteristics of the Rhodes and western Ionian basins of the eastern Mediterranean are studied by a one-dimensional, coupled physical–biological model. The biological model involves single aggregated compartments of phytoplankton, zooplankton, detritus as well as ammonium and nitrate forms of the inorganic nitrogen. It interacts with the physical model through the vertical eddy diffusivity which is calculated using the Mellor–Yamada level 2.5 turbulence parameterization. The model simulations demonstrate the importance of the contrasting physical oceanographic characteristics of these two basins on affecting their yearly planktonic structures. The annual primary production in the Rhodes basin is estimated as 97 g C m² yr⁻¹ which is comparable with the northwestern Mediterranean. The western Ionian basin, on the contrary, possesses only 10% of the Rhodes' productivity and therefore represent a most oligotrophic site in the eastern Mediterranean. The Rhodes basin reveals a strong bloom in early spring, typically in March, a weaker bloom in early winter, typically in January, and a subsurface production below the seasonal thermocline during summer. This structure is slightly modified in the western Ionian basin, and the early winter and early spring blooms are merged to cover the entire winter. These results are supported favorably by the available observations both in their magnitudes and timing.     

Material transport from the nearshore to the basinal environment in the southern Baltic Sea: I. Processes and mass estimates

Processes involved in erosion, transport and deposition of cohesive materials are studied in a transect from shallow (16 m) to deep (47 m) water of the SW Baltic Sea. The wave- and current-induced energy input to the seabed in shallow water is high with strong variability and suspended matter concentrations may double within a few hours. Primary settling fluxes (from sedimentation traps) are less than 10 g m⁻² day⁻¹, whereas resuspension fluxes (evaluated from sedimentation flux gradients) are 15–20 times higher and the residence time for suspended matter in the water column is 1–2 days. Settling velocities of aggregates are on average six times higher than for individual particles resulting in an enhanced downward transport of organic matter. Wave-induced resuspension (four to six times per month) takes place with higher shear stresses on the bottom than current-induced resuspension (three to five times per month). The short residence time in the water column and the frequent resuspension events provide a fast operating benthic–pelagic coupling. Due to the high-energy input, the shallow water areas are nondepositional on time scales longer than 1–2 weeks. The sediment is sand partly covered by a thin fluff layer during low-energy periods. The presence of the fluff layer keeps the resuspension threshold very low (<0.023 N m⁻²) throughout the year. Evaluated from 3-D sediment transport modeling, transport from shallow to deep water is episodic. The net main directions are towards the Arkona Basin (5.5×10⁵ t per year) and the Bornholm Basin (3.7×10⁵ t per year). Energy input to the bottom in deep water is low and takes place much less frequently. Wave-induced resuspension occurs on average once per month. Residence time of particles (based on radioactive isotopes) in the water column is half a year and the sediment accumulation rate is 2.2 mm year⁻¹ in the Arkona Basin.     

Analysis of the hydrophysical structure of the Sea of Azov in the period of the bottom anoxia development

The hydrophysical and hydrochemical structure of the Sea of Azov, with developed bottom anoxia, was studied during the RV “Akvanavt” cruise from July 31 to August 03, 2001. The anoxic zone with a thickness from 0.5 to 4 m above the bottom was found in all deep regions of the Sea. Concentrations of hydrochemical parameters were similar to the pronounced anoxic conditions (about 90 mmol m^− 3 of hydrogen sulfide, 17 mmol m^− 3 of ammonia, 6 mmol m^− 3 of phosphate, 7 mmol m^− 3 of total manganese). The hydrophysical structure was characterized by the uniform distribution of temperature in the upper 6–7 m mixed layer (UML). Below this a thin (0.4–0.8 m) thermocline layer was observed, just above the anoxic waters. Formation of this phenomenon was connected with that summer weather conditions. Intensive rains led to increased influx of river waters in June. That resulted in large input of allochtonous organic matter (OM) and inorganic nutrients; the latter were consumed on the additional autochthonous organic matter production. In July the weather was characterized by a significant rise in the daily averaged air temperature and large oscillations of temperature during the day. In this period a wind of constant direction was absent, but wind bursts were observed. The completed analyses showed that the formation of such a structure could be connected with the following factors: (i) positive growth trends of the daily averaged temperature and the daily oscillations of temperature, (ii) presence of wind bursts. The joint action of these factors resulted in the formation of the UML. The amplitude of wind bursts determined the depth of UML, and the value of trend determined the value of the temperature change in the thermocline. An initial presence of bottom halocline (caused by the Black Sea water influx to the bottom of the Sea of Azov) prevented the heating of the bottom layer and therefore led to an increase of vertical gradient of temperature in the thermocline. The spatial distribution of the turbulent exchange coefficient confirmed the existence of a “stagnation” area located above the anoxia zone, which is also, apparently, the reason for its occurrence.     

Vertical profiles of particulate organic matter and its relationship with chlorophyll-a in the upper layer of the NE Mediterranean Sea

Particulate organic matter (POM), nutrients, chlorophyll-a (CHL) and primary production measurements were performed in the upper layer of three different regions (cyclonic, anticyclonic and frontal+peripherial) of the NE Mediterranean Sea in 1991–1994. Depth profiles of bulk POM exhibited a subsurface maximum, coinciding with the deep chlorophyll maximum (DCM) established near the base of the euphotic zone of the Rhodes cyclone and its periphery, where the nutricline was situated just below the euphotic zone for most of the year. Moreover, the POM peaks were broader and situated at shallower depths in late winter–early spring as compared to its position in the summer–autumn period. Under prolonged winter conditions, as experienced in March 1992, the characteristic POM feature disappeared in the center of the Rhodes cyclone, where the upper layer was entirely occupied by nutrient-rich Levantine deep water. Deep convective processes in the cyclonic gyre led to the formation of vertically uniform POM profiles with low concentrations of particulate organic carbon (POC) (2.1 μM), nitrogen (0.21 μM), total particulate phosphorus (PP) (0.02 μM) and chlorophyll-a (0.5 μg/L) in the euphotic zone. Though the Levantine deep waters ascended up to the surface layer with the nitrate/phosphate molar ratios (28–29) in March 1992, the N/P molar ratio of bulk POM in the upper layer was low as 10–12, indicating luxury consumption of phosphate during algal production. Depth-integrated primary production in the euphotic zone ranged from 38.5 for oligotrophic autumn to 457 mg C m⁻² day⁻¹ for moderately mesotrophic cool winter conditions.     

Remotely-sensed chlorophyll a observations of the northern Red Sea indicate seasonal variability and influence of coastal reefs

The biological dynamics of the open northern Red Sea (21.5°–27.5° N, 33.5°–40° E) have not been studied extensively, due in part to both the inaccessibility of this desert region and political considerations. Remotely-sensed chlorophyll a data therefore provide a framework to investigate the primary patterns of biological activity in this oceanic basin. Monthly chlorophyll a data from the 8-year Sea-viewing Wide Field-of-View sensor (SeaWiFS) mission, and data from the Moderate Resolution Imaging Spectroradiometer (MODIS), were analyzed with the Goddard Earth Sciences Data and Information Services Center (GES DISC) online data analysis system “Giovanni”. The data indicate that despite the normal low chlorophyll a concentrations (0.1–0.2 mg m^− 3) in these oligotrophic waters, there is a characteristic seasonal bloom in March–April in the northernmost open Red Sea (24° to 27.5° N) concurrent with minimum sea surface temperature. The location of the highest chlorophyll concentrations is consistent with a linear box model [Eshel, G., and Naik, N.H., 1997. Climatological coastal jet collision, intermediate water formation, and the general circulation of the Red Sea. J. Phys. Oceanogr. 27(7), 1233–1257.] of Red Sea circulation. Two years in the data set exhibited a different seasonal cycle consisting of a relatively weak northern spring bloom and elevated chlorophyll concentrations to the south (21.5° to 24° N).The data also indicate that large coral reef complexes may be sources of either nutrients or chlorophyll-rich detritus and sediment, enhancing chlorophyll a concentration in waters adjacent to the reefs.     

Ability of a “minimum” microbial food web model to reproduce response patterns observed in mesocosms manipulated with N and P, glucose, and Si

We compared an idealised mathematical model of the lower part of the pelagic food web to experimental data from a mesocosm experiment in which the supplies of mineral nutrients (nitrogen and phosphorous), bioavailable dissolved organic carbon (BDOC, as glucose), and silicate were manipulated. The central hypothesis of the experiment was that bacterial consumption of BDOC depends on whether the growth rate of heterotrophic bacteria is limited by organic-C or by mineral nutrients. In previous work, this hypothesis was examined qualitatively using a conceptual food web model. Here we explore the extent to which a “simplest possible” mathematical version of this conceptual model can reproduce the observed dynamics. The model combines algal–bacterial competition for mineral nutrients (phosphorous) and accounts for alternative limitation of bacterial and diatom growth rates by organic carbon and by silicate, respectively. Due to a slower succession in the diatom–copepod, compared to the flagellate–ciliate link, silicate availability increases the magnitude and extends the duration of phytoplankton blooms induced by mineral nutrient addition. As a result, Si interferes negatively with bacterial consumption of BDOC consumption by increasing and prolonging algal–bacterial competition for mineral nutrients. In order to reproduce the difference in primary production between Si and non-Si amended treatments, we had to assume a carbon overflow mechanism in diatom C-fixation. This model satisfactorily reproduced central features observed in the mesocosm experiment, including the dynamics of glucose consumption, algal, bacterial, and mesozooplankton biomass. While the parameter set chosen allows the model to reproduce the pattern seen in bacterial production, we were not able to find a single set of parameters that simultaneously reproduces both the level and the pattern observed for bacterial production. Profound changes in bacterial morphology and stoichiometry were reported in glucose-amended mesocosms. Our “simplest possible” model with one bacterial population with fixed stoichiometry cannot reproduce this, and we suggest that a more elaborate representation of the bacterial community is required for more accurate reproduction of bacterial production.     

Inorganic carbon fluxes through the boundaries of the Greenland Sea Basin based on in situ observations and water transport estimates

A carbon budget for the exchange of total dissolved inorganic carbon C_T between the Greenland Sea and the surrounding seas has been constructed for winter and summer situations. An extensive data set of C_T collected over the years 1994–1997 within the European Sub-polar Ocean Programmes (ESOP1 and ESOP2) are used for the budget calculation. Based on these data, mean values of C_T in eight different boxes representing the inflow and outflow of water through the boundaries of the Greenland Sea Basin are estimated. The obtained values are then combined with simulated water transports taken from the ESOP2 version of the Miami Isopycnic Coordinate Ocean Model (MICOM). The fluxes of inorganic carbon are presented for three layers; a surface mixed layer, an intermediate layer and a deep layer, and the imbalance in the fluxes are attributed to air–sea exchange, biological fixation of inorganic carbon, and sedimentation. The main influx of carbon is found in the surface and the deep layers in the Fram Strait, and in the surface waters of direct Atlantic origin, whereas the main outflux is found in the surface layer over the Jan Mayen Fracture Zone and the Knipovich Ridge, transporting carbon into the Atlantic Ocean via the Denmark Strait and towards the Arctic Ocean via the Norwegian Sea, respectively. The flux calculation indicates that there is a net transport of carbon out of the Greenland Sea during wintertime. In the absence of biological activity, this imbalance is attributed to air sea exchange, and requires an oceanic uptake of CO₂ of 0.024±0.006 Gt C yr⁻¹. The flux calculations from the summer period are complicated by biological fixation of inorganic carbon, and show that data on organic carbon is required in order to estimate the air–sea exchange in the area.     

Modelling the silica pump in the Permanently Open Ocean Zone of the Southern Ocean

A coupled 1D physical–biogeochemical model has been built to simulate the cycles of silicon and of nitrogen in the Indian sector of the Permanently Open Ocean Zone of the Southern Ocean. Based on a simplified trophic network, that includes two size classes of phytoplankton and of zooplankton, and a microbial loop, it has been calibrated by reference to surface physical, chemical and biological data sets collected at the KERFIX time-series station (50°40′S–68°25′E). The model correctly reproduces the high nutrient low chlorophyll features typical of the studied area. In a region where the spring–summer mixed layer depth is usually deeper than 60 m, the maximum of chlorophyll never exceeds 1.5 mg m⁻³, and the annual primary production is only 68 g C m⁻² year⁻¹. In the surface layer nitrate is never exhausted (range 27–23.5 mmoles m⁻³) while silicic acid shows strong seasonal variations (range 5–20 mmoles m⁻³). On an annual basis 71% of the primary production sustained by nanophytoplankton is grazed by microzooplankton. Compared to North Atlantic, siliceous microphytoplankton is mainly prevented from blooming because of an unfavourable spring–summer light-mixing regime. Silicic acid limitation (high half saturation constant for Si uptake: 8 mmoles m⁻³) also plays a major role on diatom growth. Mesozooplankton grazing pressure excerpts its influence especially in late spring. The model illustrates the efficiency of the silica pump in the Southern Ocean: up to 63% of the biogenic silica that has been synthetized in the photic layer is exported towards the deep ocean, while only 11% of the particulate organic nitrogen escapes recycling in the surface layer.