Intrusions of eastern North Atlantic central waters and phytoplankton in the north and northwestern Iberian shelf during spring

The influence of intrusions of eastern North Atlantic central water (ENACW) in the north and northwestern Iberian shelf on phytoplankton composition and abundance and on particle-size distributions of seston was analyzed using data collected on three extensive cruises during spring 1991 and 1992. Water with temperature and salinity values between 12.20 and 13.86 °C and between 35.66 and 35.98 psu, respectively, characteristics of the subtropical type of ENACW (ENACW_t), was detected in the upper 100 m of the water-column in all cruises, but particularly in the western coast in 1992. The highest salinity values of this water were found near the surface (0–100-m depth) and in early spring 1992, while minimum salinity values, and also minimum geographical extension, were found in late spring in both years. Phytoplankton blooms concentrated in frontal areas between different water types, with maximum intensity and extension in early spring.Using temperature and salinity characteristics, samples were classified in four groups corresponding to the major water types found in the region: Bay of Biscay central water (BBCW), two segments of ENACW of different salinity and surface water influenced by continental runoff. This classification was significantly confirmed by three independent discriminant analyses using hydrographic and chemical (dissolved nutrients and chlorophyll) variables, phytoplankton species abundance variables and particle-size concentration of seston variables. Phytoplankton blooms related to the presence of saline waters were characterized by the dominance of either chain-forming diatoms or a mixture of diatoms and phytoflagellates and high concentrations of seston. The diatom species dominating in saline waters were typical of upwelling-induced blooms occurring generally during summer. Blooms occurring in waters influenced by runoff also contained diatoms but in lower numbers than those of saline waters. Nutrients were not exhausted in the region, suggesting that phytoplankton populations were still in active growth. These results are interpreted taking into account the known variability in water-mass formation and in the poleward current driving ENACW_t along the shelf, and indicate that saline intrusions are a major feature affecting the distribution and composition of plankton in the spring in the southern Bay of Biscay, thus enlarging to a wider spatial scale their reported influence on the pelagic ecosystem.     

A generalized model of pelagic biogeochemistry for the global ocean ecosystem. Part II: Numerical simulations

This paper presents a global ocean implementation of a multi-component model of marine pelagic biogeochemistry coupled on-line with an ocean general circulation model forced with climatological surface fields (PELAgic biogeochemistry for Global Ocean Simulations, PELAGOS). The final objective is the inclusion of this model as a component in an Earth System model for climate studies. The pelagic model is based on a functional stoichiometric representation of marine biogeochemical cycles and allows simulating the dynamics of C, N, P, Si, O and Fe taking into account the variation of their elemental ratios in the functional groups. The model also includes a parameterization of variable chlorophyll/carbon ratio in phytoplankton, carrying chl as a prognostic variable. The first part of the paper analyzes the contribution of non-local advective–diffusive terms and local vertical processes to the simulated chl distributions. The comparison of the three experiments shows that the mean chl distribution at higher latitudes is largely determined by mixing processes, while vertical advection controls the distribution in the equatorial upwelling regions. Horizontal advective and diffusive processes are necessary mechanisms for the shape of chl distribution in the sub-tropical Pacific. In the second part, the results have been compared with existing datasets of satellite-derived chlorophyll, surface nutrients, estimates of phytoplankton community composition and primary production data. The agreement is reasonable both in terms of the spatial distribution of annual means and of the seasonal variability in different dynamical oceanographic regions. Results indicate that some of the model biases in chl and surface nutrients distributions can be related to deficiencies in the simulation of physical processes such as advection and mixing. Other discrepancies are attributed to inadequate parameterizations of phytoplankton functional groups. The model has skill in reproducing the overall distribution of large and small phytoplankton but tends to underestimate diatoms in the northern higher latitudes and overestimate nanophytoplankton with respect to picoautotrophs in oligotrophic regions. The performance of the model is discussed in the context of its use in climate studies and an approach for improving the parameterization of functional groups in deterministic models is outlined.     

Biologically induced modification of seawater viscosity in the Eastern English Channel during a Phaeocystis globosa spring bloom

To identify the potential relationship between Pheaocystis globosa bloom conditions and seawater properties, a hydrobiological survey was performed in the inshore waters of the Eastern English Channel over the course of the phytoplankton spring bloom. Chlorophyll concentration, auto- and hetero/mixotrophic composition of protists and standing stock, and seawater viscosity were measured weekly from March to June 2004. The decline of the bloom is characterized by a massive foam formation in the turbulent surf zone. Before foam formation, seawater viscosity significantly increased, showing a significant positive correlation with chlorophyll concentration. In contrast, after foam formation this correlation was negative, seawater viscosity kept increasing despite a sharp decrease in chlorophyll concentrations. No significant correlation has been found between seawater viscosity and the composition of the phytoplankton assemblages observed during the survey. However, significant positive correlations have been found between seawater viscosity and both the size and the abundance of P. globosa colonies. From the correlation patterns observed between chlorophyll concentration and seawater viscosity, we suggest that the rheological properties of seawater are mainly driven by extracellular materials associated with colony formation and maintenance rather than by cell composition and standing stock.     

Sequential data assimilation in an upwelling influenced estuary

We have evaluated the impact of assimilating chlorophyll, nitrate, phosphate, silicate and ammonium into a coupled 1D hydrodynamic ecosystem model (GOTM-ERSEM) in an upwelling influenced estuary. The assimilation method chosen is the Ensemble Kalman Filter (EnKF), which has been demonstrated to improve field estimates of key variables (chlorophyll, nutrients) for bulk algal bloom prediction. The 1D model has been set up for a central station inside the Ría de Vigo (Spain). Data from bi-weekly surveys are used to constrain the model. Temperature and salinity profiles are used to ensure the correct representation of the water structure through a relaxation scheme. Chlorophyll extracts and nutrients at three depths are assimilated sequentially during 1 year simulation (1991). The assimilation period includes episodes of active upwelling and downwelling. All five assimilated variables are successfully constrained and represent a large improvement on the reference simulation (without assimilation). Small divergences can be related to poorly resolved physical processes in the model. The assimilation was further evaluated by comparing observed biomass partitioning with model results. Diatoms accounted for the largest biomass update and the largest improvement in terms of percentage of variance explained (R²). This is particularly significant as they represent the 46% of the yearly integrated observed biomass of the planktonic autotrophs. Nonetheless the R² value was low for all phytoplankton groups. Bacteria and nanoflagellates showed an improvement with respect to their yearly Root Mean Square (RMS), while the other functional groups worsen or remained unaffected. Chlorophyll assimilation was responsible for most of the impact on the phytoplankton biomass with small contributions from the silicate. It had minor impact on the updates of nutrients which in turn corrected the state variables related to the detrital pool. In this current setting, combined assimilation of chlorophyll and nutrients is not sufficient to produce a skillful simulation of the phytoplankton succession.     

Primary productivity of the Palmer Long Term Ecological Research Area and the Southern Ocean

A major objective of the Palmer Long Term Ecological Research (Palmer LTER) project is to obtain a comprehensive understanding of the various components of the Antarctic marine ecosystem. Phytoplankton production plays a key role in this so-called high nutrient, low chlorophyll environment, and factors that regulate production include those that control cell growth (light, temperature, and nutrients) and those that control cell accumulation rate and hence population growth (water column stability, grazing, and sinking). Sea ice mediates several of these factors and frequently conditions the water column for a spring bloom which is characterized by a pulse of production restricted in both time and space. This study models the spatial and temporal variability of primary production within the Palmer LTER area west of the Antarctic Peninsula and discusses this production in the context of historical data for the Southern Ocean. Primary production for the Southern Ocean and the Palmer LTER area have been computed using both light-pigment production models [Smith, R.C., Bidigare, R.R., Prézelin, B.B., Baker, K.S., Brooks, J.M., 1987. Optical characterization of primary productivity across a coastal front. Mar. Biol. (96), 575–591; Bidigare, R.R., Smith, R.C., Baker, K.S., Marra, J., 1987. Oceanic primary production estimates from measurements of spectral irradiance and pigment concentrations. Global Biogeochem. Cycles (1), 171–186; Morel, A., Berthon, J.F., 1989. Surface pigments, algal biomass profiles and potential production of the euphotic layer—relationships reinvestigated in view of remote-sensing applications. Limnol. Oceanogr. (34), 1545–1562] and an ice edge production model [Nelson, D.M., Smith, W.O., 1986. Phytoplankton bloom dynamics of the western Ross Sea ice edge: II. Mesoscale cycling of nitrogen and silicon. Deep-Sea Res. (33), 1389–1412; Wilson, D.L., Smith, W.O., Nelson, D.M., 1986. Phytoplankton bloom dynamics of the Western Ross Sea ice edge: I. primary productivity and species-specific production. Deep-Sea Res., 33, 1375–1387; Smith, W.O., Nelson, D.M., 1986. Importance of ice edge phytoplankton production in the Southern Ocean. BioScience (36), 251–257]. Chlorophyll concentrations, total photosynthetically available radiation (PAR) and sea ice concentrations were derived from satellite data. These same parameters, in addition to hydrodynamic conditions, have also been determined from shipboard and Palmer Station observations during the LTER program. Model results are compared, sensitivity studies evaluated, and productivity of the Palmer LTER region is discussed in terms of its space time distribution, seasonal and interannual variability, and overall contribution to the marine ecology of the Southern Ocean.     

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

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

An analysis of the characteristics of chlorophyll in the Sulu Sea

The seasonal and spatial variations of chlorophyll concentrations, Sea Surface Temperature (SST), wind fields and wind-induced Ekman pumping in the Sulu Sea are investigated using a set of new remote sensing measurements from October 1997 to December 2004. The results show the seasonality of chlorophyll, wind fields and SST and reveal the phytoplankton blooming events in the Sulu Sea basin during the northeast monsoon season. In summer, chlorophyll concentrations were relatively low (< 0.2 mg/m³) and distributed uniformly throughout the basin with a narrow belt of high chlorophyll concentrations along the coastal waters, particularly the coasts of Borneo and of the Sulu Archipelago. In winter, chlorophyll concentrations increased (> 0.2 mg/m³) throughout the entire basin, and phytoplankton bloomed southward to the central basin, while chlorophyll concentrations reached high levels (1 mg/m³) in the center of the blooms. One peak was observed during the northeast monsoon season each year. SSTs have significant negative correlations with chlorophyll concentrations; i.e., high and uniformly distributed in summer but lower with an obvious tongue of cold waters southward to the central basin in winter. The seasonal variation of chlorophyll concentrations and SST distribution were associated with the seasonally reversing monsoon. The winter phytoplankton blooming and the tongue of the cold waters were correlated to the vertical upwelling cold and nutrient-rich waters drawn by the northeast wind, with the center of the blooms and the location of cold tongues coinciding with the maximum of the wind speeds and the Ekman pumping velocities.     

Phytoplankton size classes competitions at sub-mesoscale in a frontal oceanic region

The effects of mesoscale and sub-mesoscale dynamics on the competition between two different phytoplankton size classes are investigated with a 3D primitive equations model. The model reproduces realistic simulations of mesoscale turbulence generated by a westward current in the southern hemisphere at statistical equilibrium in a summer situation. Effects of two different grazing pressures on phytoplankton competitions are compared and the role of eddy variability is quantified comparing high and low resolution simulations.High resolution simulations reveal a filamentary distribution of biomass and nutrients induced by the combination of vertical advection and horizontal stirring. This fine scale variability is observed not only on the horizontal but also on the vertical into the subsurface chlorophyll maximum.One of the key results is that such a dynamics induces a spatial segregation of the phytoplankton in the southern part of the frontal region that is mainly filamentary. This spatial segregation consists in biomass maxima for large phytoplankton in rich nutrients filaments and maxima for small phytoplankton outside these filaments. This anti-correlation is particularly strong when grazing pressure is low and is confirmed by statistical analysis. In the central frontal region, dominated by mesoscale dynamics, the two phytoplankton classes are strongly correlated together and biomass maxima are located close to downwelling regions that are poor in nutrients.It is shown that the effect of grazing is significantly amplified by the fine scale dynamics and that the combination of these two mechanisms is responsible of a switch of the ecosystem dominance in the surface layers.In addition, the effect of frontal dynamics on the detritus export is very sensitive to grazing pressure: increasing grazing induces a significant decrease of the export in the presence of frontal dynamics whereas it induces an increase of the export without small-scale variability.     

Validation of the 3D biogeochemical model MIRO&CO with field nutrient and phytoplankton data and MERIS-derived surface chlorophyll a images

This paper presents results obtained with MIRO&CO-3D, a biogeochemical model dedicated to the study of eutrophication and applied to the Channel and Southern Bight of the North Sea (48.5°N–52.5°N). The model results from coupling of the COHERENS-3D hydrodynamic model and the biogeochemical model MIRO, which was previously calibrated in a multi-box implementation. MIRO&CO-3D is run to simulate the annual cycle of inorganic and organic carbon and nutrients (nitrogen, phosphorus and silica), phytoplankton (diatoms, nanoflagellates and Phaeocystis), bacteria and zooplankton (microzooplankton and copepods) with realistic forcing (meteorological conditions and river loads) for the period 1991–2003. Model validation is first shown by comparing time series of model concentrations of nutrients, chlorophyll a, diatom and Phaeocystis with in situ data from station 330 (51°26.00′N, 2°48.50′E) located in the centre of the Belgian coastal zone. This comparison shows the model's ability to represent the seasonal dynamics of nutrients and phytoplankton in Belgian waters. However the model fails to simulate correctly the dissolved silica cycle, especially during the beginning of spring, due to the late onset (in the model) of the early spring diatom bloom. As a general trend the chlorophyll a spring maximum is underestimated in simulations. A comparison between the seasonal average of surface winter nutrients and spring chlorophyll a concentrations simulated with in situ data for different stations is used to assess the accuracy of the simulated spatial distribution. At a seasonal scale, the spatial distribution of surface winter nutrients is in general well reproduced by the model with nevertheless a small overestimation for a few stations close to the Rhine/Meuse mouth and a tendency to underestimation in the coastal zone from Belgium to France. PO₄ was simulated best; silica was simulated with less success. Spring chlorophyll a concentration is in general underestimated by the model. The accuracy of the simulated phytoplankton spatial distribution is further evaluated by comparing simulated surface chlorophyll a with that derived from the satellite sensor MERIS for the year 2003. Reasonable agreement is found between simulated and satellite-derived regions of high chlorophyll a with nevertheless discrepancies close to the boundaries.     

Steady-state single cell model simulations of photoacclimation in a vertically mixed layer: implications for biological tracer studies and primary productivity

A numerical single-cell photoacclimation–diffusion model was constructed and used to develop criteria regarding the use of individual phytoplankton cells as tracers for vertical mixing and to illustrate how rates of vertical mixing might affect phytoplankton physiology. Both first-order and logistic representations of photoacclimation kinetics were used. Steady state was assumed for simplicity and to provide a starting point for further investigations. The modeled variance and higher moments (within a phytoplankton population) of a generic photoacclimative parameter all show trends, which are diagnostic of mixing rates and/or boundary effects. This allowed the establishment of criteria by which frequency distributions of phytoplankton physiological properties (e.g., cell fluorescence) might be used as indicators of vertical mixing. The same model can be used to predict the effects of vertical mixing on phytoplankton productivity and growth. Application of the model to both photosynthesis and carbon to chlorophyll ratios suggested that a combination of vertical mixing and hysteresis (as represented in the logistic model of photoacclimation) in acclimation kinetics can enhance specific growth rates of phytoplankton. This enhanced growth occurred as a result of mixing-induced variation in carbon to chlorophyll ratios and is in contrast to chlorophyll-specific productivity, which was maximal at low mixing rates. Differential rates of photoacclimation to upward vs. downward shifts in irradiance, may enable phytoplankton cells to better survive in a turbulent environment.     

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.     

Wind-triggered events of phytoplankton downward flux in the Northeast Water Polynya

Phytoplankton carbon fluxes were studied in the Northeast Water (NEW) Polynya, off the eastern coast of Greenland (79° to 81°N, 6° to 17°W), during summer 1993. The downward flux of organic particles was determined during 54 days using a sediment trap moored at a fixed location, below the pycnocline (130 m). The hypothesis of the present study is that wind events were ultimately responsible for the events of diatoms downward flux recorded in the trap.Wind conditions can influence the vertical transport of phytoplankton by affecting (1) the environmental conditions (e.g. hydrostatic pressure, nutrient concentrations, and irradiance) encountered by phytoplankton during their vertical excursion, and (2) the aggregation and disaggregation of phytoplankton flocs. The first mechanism affects the physiological regulation of buoyancy, whereas the second one affects the size and shape of settling particles.Using field data (wind velocity, density profiles and phytoplankton abundance), we assessed the potential aggregation and the vertical excursion of phytoplankton in surface waters. The results show that, upstream from the trap, wind and hydrodynamic conditions were sometimes favourable to the downward export of phytoplankton. Lag-correlation between time series of wind and phytoplankton downward flux shows that flux events lagged wind events by ca. 16 days. Given that the average current velocity in the top 100 m was ca. 10 cm s⁻¹, a lag of 16 days corresponded to a lateral transport of ca. 130 km, upstream from the sediment trap, where phytoplankton production was lower than at the location of the trap. According to that scenario, 21% to 60% of primary production was exported to depth during wind events. If we had assumed instead a tight spatial coupling between the material collected in the trap and the relatively high phytoplankton production at the location of the trap, we would have concluded that <7% of primary production was exported to depth. The difference between the two scenarios has great implications for the fate of phytoplankton. Our results stress the importance of investigating the spatial coupling between surface and trap data before assessing the pathways of phytoplankton carbon cycling.     

Seasonal phytoplankton responses to environmental factors in a shallow Mediterranean lagoon

Phytoplankton community structure was investigated in a 1-year study period from January to December 2006 in the Tunis North Lagoon (South Mediterranean). Twice a month, sampling was carried out from the whole water column. Phytoplankton species composition showed seasonal dynamics following the general environmental variable trends in the study area, with variation in species abundance levels within each season characterised by the presence of different phytoplankton communities. Analysis of environmental variables indicated that phytoplankton-dominant communities were associated with various water physicochemical characteristics, especially water temperature and salinity. Accordingly, significant correlation was recorded between water temperature and dinoflagellates (r = 0.35; p < 0.05) in summer and diatoms (r = 0.69; p < 0.05) in autumn, whereas euglenophytes, cyanophytes and chlorophytes were slightly correlated with temperature in autumn. Salinity was positively correlated with dichtyophytes (r = 0.41; p < 0.05) in winter and with diatoms (r = 0.65; p < 0.05) and euglenophytes (r = 0.57; p < 0.05) in autumn. On the other hand, relationships between high nitrogen nutrient concentration and phytoplankton concentration were recorded for diatoms (r = 0.43; p < 0.05 with NO₂; r = 0.49; p < 0.05 with NO₃) in winter. Silicate concentration supported proliferation of diatoms (r = 0.58; p < 0.05) in autumn in our study period. In contrast, increase of dinoflagellate concentration was associated with the decrease of these parameters in spring and summer.     

Relationship between phytoplankton bloom and wind stress in the sub-polar frontal area of the Japan/East Sea

The seasonal dynamics of phytoplankton blooms in the central Japan/East Sea (JES) show pronounced year-to-year variability based on Sea-viewing Wide Field-of-view Sensor (SeaWiFS; 19972003) and Moderate Resolution Imaging Spectroradiometer (MODIS)/Terra (20002003) observations. Wind seems to strongly influence this variability. To study the relationship between wind and bloom initiation, we analyzed daily, remotely sensed wind stress data (Active Microwave Instrument–wind [AMI–wind], NASA Scatterometer [NSCAT], and Quick Scatterometer [QuickSCAT]: 19972003) and daily chlorophyll concentrations based on ocean color data (SeaWiFS and MODIS). The results agreed well with the hypotheses; in spring, blooms began 615 days after wind stress weakened. Fall blooms started 39 days after wind strengthened. We also simulated seasonal changes using a simple light–nutrient model using two values for the respiration ratio: 10% and 20%. The use of 20% seemed to reproduce the timing of the spring bloom quite well but underestimated the absolute level of chlorophyll concentration. On the other hand, using 10% produced a better estimation of the chlorophyll concentration but failed to match the timing. Neither of the model runs reproduced the timing of the fall bloom well.     

Size-fractionated phytoplankton biomass and species composition in the Indian sector of the Southern Ocean during austral summer

During the late austral summer of 1994, Antarctic waters were characterized by low phytoplankton biomass. Along the 62°E meridian transect, between 49°S and 67°S, chlorophyll (Chl.) a concentration in the upper 150 m was on average 0.2 mg m⁻³. However, in the Seasonal Ice Zone (SIZ) chlorophyll a concentrations were higher, with a characteristic deep chlorophyll maximum. The highest value (0.6 mg Chl. a m⁻³) was measured at the Antarctic Divergence, 64°S, corresponding to the depth of the temperature minimum (100 m). This deep biomass maximum decreased from South to North, disappeared in the Permanently Open Ocean Zone (POOZ) and reappeared with less vigour in the vicinity of the Polar Front Zone (PFZ). In the SIZ, the upper mixed layer was shallow, biomass was higher and the >10 μm fraction was predominant. In this zone the >10 μm, 2–10 μm and <2 μm size fractions represented on the average 46%, 25.1% and 28.9% of the total integrated Chl. a stock in the upper 100 m, respectively. The phytoplankton assemblage was diverse, mainly composed of large diatoms and dinoflagellate cells which contributed 42.7% and 33.1% of the autotrophic carbon biomass, respectively. Moving northwards, in parallel with the decrease in biomass, the biomass of autotrophic pico- and nanoflagellates (mainly Cryptophytes) increased steadily. In the POOZ, the picoplanktonic size fraction contributed 47.4% of the total integrated Chl. a stock. A phytoplankton community structure with low biomass and picoplankton-dominated assemblage in the POOZ contrasted with the relatively rich, diverse and diatom-dominated assemblage in the SIZ. These differences reflect the spatial and temporal variations prevailing in the Southern Ocean pelagic ecosystem.     

Adapting the light · biomass (BZI) models of phytoplankton primary production to shallow marine ecosystems

Several authors have reported a strong linear relationship between daily phytoplankton production and the product of chlorophyll biomass, photic depth, and incident irradiance for a variety of estuaries. This “light · biomass” (BZ_pI_o) formulation has been proposed as an alternative to traditional mechanistic approaches for computing phytoplankton production in numerical estuarine models. One limitation to their application in shallow systems is that the BZ_pI_o models have been developed in relatively deep estuaries where light does not reach the bottom. We propose a nonlinear correction factor to adapt the BZ_pI_o relationship to shallow systems where light does reach the bottom. Our function takes into account variations in incident irradiance, attenuation coefficient for light, photosynthetic efficiency, and maximum rate of photosynthesis. A series of correction polynomials are proposed for various ranges of incident irradiance, and are integrated into a single multiple polynomial which applies across all irradiance levels. Our new correction factor was tested against a ¹⁴C-based productivity dataset from shallow stations in Narragansett Bay, RI and an O₂-based dataset from shallow (1.1 m) lagoon mesocosms at the University of Rhode Island. Results showed that our polynomials accurately correct BZ_pI_o-predicted rates of production in shallow water columns. Application of our correction factor to a series of shallow water productivity datasets from the literature together with theoretical calculations show how significant the shallow water correction can be, especially in very shallow water columns with low turbidity.     

Microscale variability in the distributions of large fluorescent particles observed in situ with a planar laser imaging fluorometer

It has been known for decades that particle-size and biomass spectra show regular patterns in the ocean, and that these patterns often show systematic variations with other properties such as total biomass, nutrient concentration, season, and distance (both vertical and horizontal). The recent finding of the ubiquitous nature of layers of phytoplankton < 1 m thick prompted us to explore the fine- and microscale vertical variations of size- and fluorescence-abundance spectra in the ocean. Using a two-dimensional planar laser imaging system mounted on a free-falling platform, we quantified the properties of large fluorescent particles ( 20 μm–2 cm) through the water column, obtaining images every 10–30 cm. These images showed systematic relationships of the spectral properties to total chlorophyll: increased proportions of the smallest particles at high chlorophyll concentrations, and a lengthening of the spectral size range at high total chlorophyll concentrations (more large particles at high chlorophyll concentrations). Further, we observed significant variations of the spectral properties over scales of 1 m and less, and recorded the frequent occurrence of unusual layers of large particles. Our new instrument, which is sensitive to thin layers of enhanced phytoplankton biomass, shows the planktonic community to be highly structured vertically on scales of 1–2 m, particularly within the DCM.     

Environmental changes and the responses of the ecosystems of the Yellow Sea during 1976–2000

The changes in the environmental features of the Yellow Sea during the last 25 years of the 20th century were studied using a set of seasonally monitored data along a transect (at 36°N) maintained by the State Oceanic Administration of China. The data included the ocean temperature (T), salinity (S) and biogenic elements, such as dissolved oxygen (DO), phosphorus (PO₄-P), silicon (SiO₃-Si) and dissolved inorganic nitrogen (DIN).The seasonal (summer and winter) values and the annual mean of these elements showed significant changes during the monitored period. Time series of T, S, DIN and N:P ratios exhibited positive trends, while those of DO, P and Si exhibited negative trends. During this period, the annual mean of T and DIN in the Yellow Sea increased by 1.7 °C and 2.95 μmol L⁻¹, respectively, while those of DO, P and Si decreased by 59.1, 0.1 and 3.93 μmol L⁻¹, respectively. In the 1980s, particularly in between 1985 and 1989, concentrations of P and Si dropped to near the ecological threshold for growth of diatoms. The N:P ratio increased from 4 in 1984 to over 16 in 2000. The climate trend coefficients, R_xt, for these time series are all above 0.43 with significance levels of 95%, except for salinity. The increases in T were consistent with the recent climate warming in northern China and the adjacent seas, i.e. the Bohai Sea and the East China Sea. The reduction of DO was probably attributable to the increase in T and decrease in primary production in these regions. The positive trend of DIN was mainly attributable to precipitation and partly to Changjiang River discharge. The negative concentration trends of P and Si were due to the decreases in their concentrations in seawater that flowed to the Yellow Sea from the Bohai Sea. As a result, N:P ratios greatly increased in the seawater of the Yellow Sea.Moreover, some important responses of the Yellow Sea ecosystems to the changes in physical variables and chemical biogenic elements were obviously displayed. These responses include strengthening nutrient limitation, decreasing chlorophyll a, primary production and phytoplankton abundance, succession of dominant phytoplankton species from diatoms to non-diatoms, changes in fish community structure and species diversity.     

The coastal edge of the Northeast Water polynya in spring 1993

Multidisciplinary, marine ecological observations were conducted at the shallow water edge of the Northeast Water in June, 1993. Although variable in size and shape, a small polynya was constantly present at Eskimonaes, at the fast-ice edge of Ingolfsfjord. A shallow stratified layer developed at the water sufface at negative water and air temperatures—an effect of sea ice melting in cold water early in the season. Nutrients were recorded in considerable quantities, although by mid July NO₃ had become depleted. The chlorophyll and phytoplankton maxima at 8–12 m depth had peak values of 2 mg chl a m⁻³, typical for Arctic algal blooms. The phytoplankton included over 90 species and was dominated by the Fragillariopsis group. Zooplankton was poor in biomass and density, but over 23 taxa were found, with the copepods Oithona similis and Pseudocalanus acuspes being numerically dominant. Sedimentation was approximately 0.2 g dry weight m⁻² d⁻¹ and suspended matter concentrations ranged from 4 to 19 mg l⁻¹. The benthos was represented by hard bottom forms only, with a surprisingly dense cover of macrophytes. Juvenile sea urchins (Strongylocentrotus droebachiensis), brittle stars (Ophiocten sericeum) and amphipods were dominant. Higher trophic levels were represented by benthic feeders, such as eiders and walruses. The area observed was more similar to high Arctic fjord ecosystems than to the offshore central Northeast Water polynya.     

Use of SeaWiFS data for light availability and parameter estimation of a phytoplankton production model of the Bay of Biscay

Processing SeaWiFS (Sea-viewing Wide Field-of-view Sensor) data provides useful information for the observation and modelling of the phytoplankton production of the Bay of Biscay. Empirical algorithms allow the retrieval of chlorophyll a and non-living Suspended Particulate Matter (SPM) concentrations. These data are used to constrain a coupled 3D physical–biogeochemical model of the Bay of Biscay continental shelf. Two issues are investigated, depending on the variable used, to constrain the winter to spring phytoplankton production for the year 2001. First, SPM data is used as forcing data to correct the corresponding state variable of our model. This allows the realistic simulation of the light limited bloom at the end of February 2001, as observed with SeaWiFS chlorophyll a images and from the NUTRIGAS field cruise. Second, chlorophyll a data is used for parameter estimation of the biogeochemical model. The ability of assimilating these data is tested to improve the simulation of strong blooms observed in late May 2001 in the Loire and Gironde plumes. A global optimization method (Evolutive Strategies) is adapted to the complete 3-D coupled model, in order to find the best set of parameters. The hydrological conditions during the bloom can be validated with data from the PEL01 field cruise. After selection of the most sensitive parameters, the method is tested with twin experiments. Then, the use of real SeaWiFS data reduces the model/data misfit by a factor of two, improving the simulation of bloom intensities and extensions. The sets of parameters retrieved in each plume are discussed.