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

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

Spatial and temporal variability of size-fractionated biomass and primary production in the Ross Sea (Antarctica) during austral spring and summer

The results of a study on the spatial and temporal dynamics of size-fractionated biomass and production of phytoplankton in the Ross Sea during the austral spring and summer are reported. The spring cruise took place in the offshore Ross Sea from 14 November to 14 December 1994. Sampling was carried out on a transect of 27 stations distributed from 76.5 to 72.0°S along 175°E, and covered the three main Antarctic environments of the polynya open waters, the marginal ice zone and the pack ice area. Three subsystems were identified. The subsystem of the polynya was characterised by the predominance of the micro- and nano-planktonic fractions, chlorophyll (Chl a) concentrations from 69.6 to 164.7 mg m⁻² and production rates from 0.68 to 1.14 g C m⁻² day⁻¹. The second subsystem, the marginal ice zone, showed a relative increase of the micro-planktonic fraction, high biomass levels (from 99.64 to 220 mg Chl m⁻²) and production rates from 0.99 to 2.7 g C m⁻² day⁻¹. The subsystem of the pack ice area had a phytoplankton community dominated by the pico-planktonic fraction and showed low biomasses (from 19.4 to 37.7 mg Chl m⁻²) and production rates (0.28 to 0.60 g C m⁻² day⁻¹). Selective grazing by krill is considered an important factor in determining the size structure of the phytoplankton communities. The summer study consisted of a time series carried out in inshore waters of Terra Nova Bay from 12 January to 8 February 1990. In a well stabilised water column and with high levels of PAR always available, the primary production rates of a community dominated by micro-plankton varied from 0.52 to 1.2 g C m⁻² day⁻¹ (average 0.84). A high P/B ratio, up to 3, and a remarkably elevated mean phaeopigment (Phaeo)/Chl a ratio of 2.4 indicated an active removal of biomass by grazing, confirmed by the presence of faecal pellets in quantities reaching 6000 m⁻³ in the upper 50 m. The peculiarities of the inshore versus offshore environments in terms of community size structure, production processes and their implications as regards the food web are discussed.     

Seasonal variability of fractionated phytoplankton, biomass and primary production in the Straits of Magellan

The results on the distribution of phytoplankton biomass (expressed as Chla) and primary production (¹⁴C assimilation), during three oceanographic cruises carried out during Austral spring and at the end of the summer and the autumn in the Straits of Magellan, suggest a strong variability of trophic levels for this ecosystem.Seasonal evolution of the biomass concentration goes from the spring maximum of 2.33 μg/l through a sharp decrease, 0.49 μg/l, observed at the end of summer, until the minimum of 0.24 μg/l measured during the autumn.The trophic conditions are dependent on hydrographic, meteo-climatic and geo-morphological characteristics: at the Atlantic entrance and between the two Angosturas the strong mixing of water column limit the development of phytoplankton; at the Western opening and along the Pacific arm the complex exchange mechanisms with the ocean, the glacio-fluvial contribution and the presence of a thermohaline front near the Isla Carlos III influence both biomass and primary production distributions. The maximum values are reached in the Central Zone (Paso Ancho) characterized by high stability of the water column.Primary production ranged from a minimum of 12.3 to a maximum of 125.9 mgC m⁻² h⁻¹. The overall trend seems to be a progressive and simultaneous increase from the Pacific and Atlantic openings to the Central Zone of Paso Ancho where the maximum value was reached. In general, biomass and primary production distributions correspond quite well except for the area of Isla Carlos III where biological and chemico-physical causes tend to limit ¹⁴C assimilation.Contribution of pico-phytoplankton (< 2 μm) to total biomass appears to be time dependent: in the blooms observed during spring a very modest incidence (< 6%) was observed whereas became more (> 50%) during the summer-autumn seasons when total biomass was decreasing.Within the Straits, at the end of summer, the contribution of pico-phytoplankton primary production is 59%, whereas nano and microplankton contribute 39% and 2%, respectively. At the oceanic external stations the photosynthetic activity of the bigger size-fraction (> 2 μm) is predominant (> 50%).These findings support the hypothesis that the pico-phytoplankton ( < 2 μm) is substantially constant, whereas temporal variations are due to the larger (> 10 μm) cells only.     

Distribution of phytoplankton in the southern Black Sea in summer 1996, spring and autumn 1998

The species composition, abundance, and biomass of micro- (>15 μm) and nano- (<15 μm) phytoplankton were studied along the southern Black Sea during June–July 1996 and March–April and September 1998. A total of 150 species were identified, 50% of them being dinoflagellates. The average total phytoplankton abundance changed from 77×10³ cells l⁻¹ in spring to 110×10³ cells l⁻¹ in autumn and biomass from 250 μg l⁻¹ in summer to 1370 μg l⁻¹ in spring. Based on the extensive sampling grid from June–July 1996, phytoplankton seemed to have a rather homogeneous biomass distribution in the southern Black Sea. In all periods, the coccolithophorid Emiliania huxleyi was the most abundant species, its contribution to the total abundance ranging from 73% in autumn to 43% in spring. However, in terms of biomass, diatoms made up the bulk of phytoplankton in spring (97%, majority being Proboscia alata) and autumn (73%, majority being Pseudosolenia calcar-avis), and dinoflagellates in summer (74%, Gymnodinium sp.). There was a remarkable similarity in the dominant species between the western and eastern regions of the southern Black Sea, indicating transport of phytoplankton within the basin.     

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.     

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.     

Seasonality of picophytoplankton chlorophyll a and biomass in the central Cantabrian Sea, southern Bay of Biscay

Seasonal changes in the abundance and biomass of cyanobacteria (Synechococcus and Prochlorococcus) and picoeukaryotes were studied by flow cytometry in the upper layers of the central Cantabrian Sea continental shelf, from April 2002 to April 2006. The study area displayed the typical hydrographic conditions of temperate coastal zones. A marked seasonality of the relative contribution of prokaryotes and eukaryotes was found. While cyanobacteria were generally more abundant for most of the year (up to 2.4 10⁵ cells mL^− 1), picoeukaryotes dominated the community (up to 10⁴ cells mL^− 1) from February to May. The disappearance of Prochlorococcus from spring through summer is likely related to shifts in the prevailing current regime. The maximum total abundance of picophytoplankton was consistently found in late summer–early autumn. Mean photic-layer picoplanktonic chlorophyll a ranged from 0.06 to 0.53 µg L^− 1 with a relatively high mean contribution to total values (33 ± 2% SE), showing maxima around autumn and minima in spring. Biomass (range 0.58–40.16 mg C m^− 3) was generally dominated by picoeukaryotes (mean ± SE, 4.28 ± 0.27 mg C m^− 3) with an average contribution of cyanobacteria of 30 ± 2%. Different seasonality of pigment and biomass values resulted in a clear temporal pattern of picophytoplanktonic carbon to chlorophyll a ratio, which ranged from 10 (winter) to 140 (summer). This study highlights the important contribution of picoplanktonic chlorophyll a and carbon biomass in this coastal ecosystem.     

Picophytoplankton and nanophytoplankton abundance and distribution in the southeastern Beaufort Sea (Mackenzie Shelf and Amundsen Gulf) during Fall 2002

The distribution of picophytoplankton (0.2–2 µm) and nanophytoplankton (2–20 µm) in the Beaufort Sea–Mackenzie Shelf and Amundsen Gulf regions during autumn, 2002 is examined relative to their ambient water mass properties (salinity, temperature and nutrients: nitrate + nitrite, phosphate, and silicate) and to the ratio of variable to maximum fluorescence, Fv/Fm. Total phytoplankton and cell abundances (< 20 µm) were mainly correlated with salinity. Significant differences in picophytoplankton cell numbers were found among waters near the mouth of the Mackenzie River, ice melt waters and the underlying halocline water masses of Pacific origin. Picophytoplankton was the most abundant phytoplankton fraction during the autumnal season, probably reflecting low nitrate concentrations (surface waters average ~ 0.65 µM). The ratio Fv/Fm averaged 0.44, indicating that cells were still physiologically active, even though their concentrations were low (max Chl a = 0.9 mg m^− 3). No significant differences in Fv/Fm were evident in the different water masses, indicating that rate limiting conditions for photosynthesis and growth were uniform across the whole system, which was in a pre-winter stage, and was probably already experiencing light limitation as a result of shortening day lengths.     

100-years-changes in the phytoplankton community of Kiel Bight (Baltic Sea)

Literature data from 1905/06, 1912/13 and 1949/50 were compared with recent data (2001–2003) from Kiel Bight in order to investigate changes in phytoplankton composition and biomass, which may serve as indicators of environmental changes. In terms of biomass, diatomophyceae and dinophyceae are by far the most important groups. Their ratio is still close to unity. The share of diatomophyceae increased strongly in years with exceptionally high summer blooms (2001) or exceptionally early spring blooms (2003). The summer and autumn blooms of Chaetoceros and Skeletonema, detected in the early 20th century, are replaced by other diatoms (Cerataulina pelagica, Dactyliosolen fragilissimus, Proboscia alata, Pseudo-nitzschia spp.). Chaetoceros and Skeletonema are still important components of the spring blooms. Now as before, the autumn blooms are dominated by Ceratium spp., sometimes also by diatoms. Newly appearing bloom-forming species are mostly potentially toxic (Dictyocha speculum, Prorocentrum minimum, Pseudo-nitzschia spp.). The total phytoplankton biomass has roughly doubled in the course of the last century. The reference condition for phytoplankton biomass in Kiel Bight in the sense of the Water Framework Directive was defined at 55 mg C m^− 3 (± 10%, annual mean). The mean annual biomass of diatomophyceae and dinophyceae was 25 mg C m^− 3 (± 40%) for each, indicating that the sum of their carbon biomass amounted to 90% (± 10%) of the total phytoplankton biomass on an annual average. Diatomophyceae represented at least 80% of carbon biomass in the spring bloom peak at the beginning of the 20th century.     

A study of the response of chlorophyll-a biomass to a winter upwelling event off Western Iberia using SeaWiFS and in situ data

Observations of a winter upwelling event off Western Iberia shelf/slope in the area of influence of the Western Iberia Buoyant Plume (WIBP) were conducted in February 2000. Spatial patterns and time evolution of the chlorophyll-a (chl-a) biomass are analysed, based on in situ and satellite data. SeaWiFS-derived chl-a concentration L2 products were used to track the chlorophyll front and estimate its westward migration velocity (maximum up to 29 km day⁻¹), as well as to characterize the frontal system and its evolution. A method associating the type of spectral signature of a pixel to the fraction of chlorophyll probed by SeaWiFS enabled the estimation of the chl-a biomass within error intervals. High chlorophyll concentrations (for wintertime) were observed over the shelf and slope, up to large distances to the coast. Due to the WIBP, a shallow Ekman layer developed, being nearly coincident with the stratified upper meters. The transport comprised westward advection and stretching of the plume, with little entrainment with the offshore deep mixed layer waters. The relative enlargement of the total area of the Inside-Front Zone (IFZ) during the upwelling event was roughly accompanied by the maintenance of the average biomass per unit of area, considering the water column up to depths of interest. This suggests that there was a net increase of chl-a biomass inside the water column associated with the IFZ, roughly proportional to the increase in the IFZ area. Retention of phytoplankton in the shallow stratified nutrient-rich waters of the WIBP was a key factor for this increase in chl-a biomass.     

The annual cycle of nanoflagellates in the Central Cantabrian Sea (Bay of Biscay)

The annual cycle of nanoflagellates (NF) including autotrophic (ANF), heterotrophic (HNF) and mixotropic (MNF) flagellates carried out in a temperate sea (Central Cantabrian Sea, southern Bay of Biscay) is presented. Three stations with characteristics ranging from coastal to oceanic conditions were analysed in order to compare NF response to this gradient. Samples were monthly collected at each station at three different depths between February 2002 and December 2002. CTD profiles were also taken at each station. NF were grouped according to their trophic status into ANF, HNF and MNF. Abundance and biomass were determined for each group. The annual cycle showed a general pattern consisting in a maximum in July with secondary maxima in March and October and minimum values in May. ANF were the most important fraction, making a major contribution (nearly 75%) to total NF biomass in all stations. HNF represented over 20% along the cycle, except for a peak in spring found in every station. MNF reached less than 5%, showing low seasonability. Small flagellates (2–5 µm) dominated throughout the cycle. Microplankton community was also analysed in terms of abundance and biomass. A significant positive correlation (r² = 0.49) was obtained between 2–5 µm NF and 10–20 µm HNF–MNF biomasses, suggesting a possible trophic relationship between these groups which should be cautiously taken. No significant relationships were found between microplankton and NF or between nutrients and ANF, indicating that the regulation of NF numbers is complex and probably implicates other groups. In addition to this, the unexpected 2002 Chl a concentration pattern and the misplacing of upwelling events render necessary to perform additional studies to fully understand the precise behaviour of NF in the Cantabrian Sea. To the best of our knowledge, this is the first study of a NF cycle in a temperate sea that considers all functional groups.     

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.     

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.     

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.     

Role of filtering and biodeposition by Adamussium colbecki in circulation of organic matter in Terra Nova Bay (Ross Sea, Antarctica)

At Terra Nova Bay, the scallop Adamussium colbecki (Smith, 1902) characterises the soft and hard bottoms from 20 to 80 m depth, constituting large beds and reaching high values of density (50–60 individuals/m²) and biomass (120 g/m² DW soft tissues). To assess its role in the organic matter recycling in the coastal ecosystem, its filtering and biodeposition rates were evaluated in laboratory experiments during the austral summer 1993/94. Filtration rates, measured in a flow-through system, were calculated from the difference in particulate organic carbon (POC), nitrogen (PON) and chlorophyll-a (Chl-a) concentration in inflow and outflow water. Experiments were performed using natural sea water with POC, PON and Chl-a concentrations of about 450 μg/l, 90 μg/l and 2 μg/l, respectively. The biodeposition rate and the biochemical composition of the biodeposits were studied in order to detect how the organic matter is transformed through feeding activity of A. colbecki. At +1°C temperature, the average filtering rate was about 1 l h⁻¹ g⁻¹ (DW soft tissues) in specimens ranging in body mass from 2 to 3 g (DW soft tissues) and 6–7 cm long. The biodeposition rate in 3–8 cm long specimens, ranging from 0.4 to 5.7 g (DW soft tissues), was about 5.65 mg DW/g DW/day, leading to an estimate of Corg flux, through biodeposition by A. colbecki, of about 21 mg C m⁻² day⁻¹ at in situ conditions. Comparison between the biochemical composition of seston and biodeposits shows a decrease of the labile compounds, of the Chl-a/phaeopigments ratio in the biodeposits. The recorded C/N ratio decrease suggests a microbial colonisation in the biodeposits. This study suggests that Adamussium colbecki plays an important role in coupling the material fluxes from the water column to the sea bed, processing about 14% of total Carbon flux from the water column to the sediments, with an assimilation efficiency of 36%.     

Microbial community structure in the marginal ice zone of the Bellingshausen Sea

Phytoplankton, bacteria and microzooplankton were investigated on a transect in the Bellingshausen Sea during the ice melt period in November–December 1992. The transect along the 85°W meridian comprised seven stations that progressed from solid pack-ice (70°S), through melting ice into open water (67°S). The abundance, biomass and taxonomic composition were determined for each component of the microbial community. The phytoplankton was mostly dominated by diatoms, particularly small (<20 μm) species. Diatom abundance ranged from 66 000 cells l⁻¹ under the ice to 410 000 cells l⁻¹ in open water. Phytoplankton biomass varied from <1 to 167 mg C m⁻³, with diatoms comprising 89–95% of the total biomass in open water and autotrophic nanoflagellates comprising 57% under the ice. The standing stocks of autotrophs in the mixed layer ranged from 95 mg C m⁻² under the pack-ice to 9478 mg C m⁻² in open waters. Bacterial abundance in ice-covered and open water stations varied from 1.1 to 5.5×10⁸ cells l⁻¹. Bacterial biomass ranged from 2.4 mg C m⁻³ under pack-ice to an average of 14 mg C m⁻³ in open water. The microzooplankton consisted mainly of aloricate oligotrich ciliates and heterotrophic dinoflagellates and these were most abundant in open waters. Their biomass varied between 0.2 and 54 mg C m⁻³ with a minimum at depth under the ice and maximum in open surface waters. Microheterotrophic standing stocks varied between 396 mg C m⁻² under pack-ice and 3677 mg C m⁻² in the open waters. The standing stocks of the total microbial community increased consistently from 491 mg C m⁻² at the ice station to 13 155 mg C m⁻² in open waters, reflecting the productive response of the community to ice-melt. The composition of the microbial community also shifted markedly from one dominated by heterotrophs (82% of microbial stocks) at the ice station to one dominated by autotrophs (73% of microbial stocks) in the open water. Our estimates suggest that the microbial community comprised >100% of the total particulate organic carbon (POC) under the ice and 62–66% of the measured POC in the open waters.     

Surface chlorophyll in the Black Sea over 1978–1986 derived from satellite and in situ data

Absolute values of chlorophyll a concentration and its spatial and seasonal variations in the Black Sea were assessed by using satellite CZCS and in situ data. Since the satellite CZCS had operated for the 1978–1986 period, CZCS data was used for assessing the past state of the Black Sea just before the onset of drastic changes observed in late 1980s. The approach used for the calculation of the absolute values of chlorophyll a concentration from CZCS data was based on the direct comparison of in situ chlorophyll a data and those of CZCS and by applying the algorithm developed for the transformation of CZCS data into chlorophyll a values. CZCS Level 2 data related with pigment concentration having a spatial resolution of 1 km at nadir were used. The daily Level 3 files were derived by binning Level 2 values into 4-km grid cells and the monthly and seasonal Level 3 files were created by averaging the daily Level 3 files over the corresponding period. In situ chlorophyll a data were obtained by spectrophotometric and fluorometric methods in 15 scientific cruises over the 1978–1986 period. Total number of ship-measured data used for the comparison with those CZCS values was 590.Chlorophyll a concentration (Chl) was derived from CZCS values (C) with regression equations Chl=kC; the coefficient of transformation k was calculated from six different data sets by taking into account distinctions between subregions and seasons. The reasons for difference in the k values have been analyzed.Statistical comparison of the chlorophyll a values measured in situ and those derived from CZCS data was based on log-transformed data and gave the following results: regression SLOPE=0.842, regression INTERCEPT=−0.081, coefficient of determination (R²)=0.806, root–mean–square ERROR=0.195. The mean monthly chlorophyll a distributions derived from CZCS data over 1978–1986 have been constructed and the mean seasonal chlorophyll a values in different regions have been calculated and analyzed. The significant difference in chlorophyll concentration between the western shelf regions and the open part of the Black Sea has been demonstrated, especially in warm season. At almost all seasons, the highest chlorophyll concentration is observed in the western interior shelf region which is under strong influence of Danube. The summer mean chlorophyll concentration in this region is 18 times higher than that in the open parts and about nine times higher than in the eastern shelf region. The greatest seasonal variations are observed in the open part of the Black Sea: chlorophyll concentration in cold season is four to six times higher than in summer and three to five times higher than in April and October. To the contrary, in the western interior shelf regions, the concentration is higher in May–October (about twice than that in November–March). Seasonal variations in the western outer shelf regions are smoothed out as compared with both the western interior shelf and the open regions.     

A numerical investigation of phytoplankton and Pseudocalanus elongatus dynamics in the spring bloom time in the Gda sk Gulf

A nutrient–phytoplankton–zooplankton–detritus (1D-NPZD) ‘phytoplankton {Phyt} and Pseudocalanus elongatus {Zoop} dynamics in the spring bloom time in the Gda sk Gulf. The 1D-NPZD model consists of three coupled, partial second-order differential equations of the diffusion type for phytoplankton {Phyt}, zooplankton {Zoop}, nutrients {Nutr} and one ordinary first-order differential equation for benthic detritus pool {Detr}, together with initial and boundary conditions. In this model, the {Zoop} is presented by only one species of copepod (P. elongatus) and {Zoop} is composed of six cohorts of copepods with weights (W_i) and numbers (Z_i); where . The calculations were made for 90 days (March, April, May) for two stations at Gda sk Gulf with a vertical space step of 0.5m and a time step of 900 s. The flow field and water temperature used as the inputs in the biological model 1D-NPZD were reproduced by the prognostic numerical simulation technique using hydrographic climatological data. The results of the numerical investigations described here were compared with the mean observed values of surface chlorophyll-a and depth integrated P. elongatus biomass for 10 years, 1980–1990. The slight differences between the calculated and mean observed values of surface chlorophyll-a and zooplankton biomass are ca. 10–60 mg C m⁻³ and ca. 5–23 mg C m⁻², respectively, depending on the location of the hydrographic station. The 1D-NPZD model with a high-resolution zooplankton module for P. elongatus can be used to describe the temporal patterns for phytoplankton biomass and P. elongatus in the centre of the Gda sk Gulf.     

CO2 partial pressure in Northeast Atlantic and adjacent shelf waters: Processes and seasonal variability

CO₂ partial pressure in surface water was measured in the Northeast Atlantic and in the Hebride Shelf/North Sea area during a cruise with R.V. Poseidon in June 1991. A mean p_CO2 of 303 μatm was found in the Atlantic between 50°N and 60°N. For an atmospheric CO₂ content of 357.5 ppm(v) this corresponds to a partial pressure difference of −55 μatm. This supports the view that the subarctic Atlantic is a significant sink within the CO₂ cycle between the ocean and the atmosphere. A comparison of our measurements with other data reveals that the p_CO2 distribution changes significantly during May/June. This explained by seasonal warming, CO₂ exchange with the atmosphere and biomass production. The contribution by each of these processes to the seasonal variations is calculated. It was found that during a plankton bloom the production of biomass is the dominating factor and may lower seawater p_CO2 by almost 100 μatm. The shelf areas are charactrized by strong p_CO2 gradients which are explained by water exchange with the Atlantic, temperature effects and biomass production.     

Observations on colony formation by the cosmopolitan phytoplankton genus Phaeocystis

Few marine phytoplankton have heteromorphic life cycles and also often dominate the ecosystems in which they occur. The class Prymnesiophyceae contains a notable exception: the genus Phaeocystis includes three species that form gelatinous colonies but also occur within their ranges as solitary cells. Phaeocystis antarctica and P. pouchetii are exclusively high latitude taxa, and are notable for regionally tremendous blooms of the colony stage. P. globosa occurs circumglobally, yet its colony blooms primarily are confined to colder waters within its range. Three additional species are warm water forms that have been reported only as solitary cells or loose aggregations that bear little resemblance to the organized colonies of the other taxa. Interpretation of existing data indicates that resource availability (light, temperature and nutrients) by itself is not sufficient to explain this distinction between cold-water colony-forming taxa and warm water solitary cell taxa, nor why colony development in P. globosa is essentially a spatially restricted phenomenon within a much broader geographic range. Colony development by P. globosa in situ has been observed at temperatures ≥20 °C, but only rarely and generally under conditions of seasonally or anthropogenically elevated nutrient supply. Data presented here demonstrate colony development at 20–22 °C in natural plankton communities from oligotrophic waters that were pre-screened through 63 μm mesh (i.e. lacking mesozooplankton and large microzooplankton), but not in unscreened communities containing microzooplankton and >63 μm zooplankton. Reduction of colony proliferation at higher temperatures by mesozooplankton grazing remains as an intriguing possibility that is consistent with available evidence to help explain differences in latitudinal extent of in situ colony development. These data are interpreted within a theoretical framework regarding the potential advantages and disadvantages of the two life cycle stages.