Phytoplankton dynamics in the NE subarctic Pacific

Ocean Station Papa (OSP, 50°N 145°W) in the NE subarctic Pacific is characterised as high nitrate low chlorophyll (HNLC). However, little is known about the spatial extent of these HNLC waters or the phytoplankton dynamics on the basin scale. Algal biomass, production and size-structure data are presented from winter, spring and summer between 1992 and 1997 for five stations ranging from coastal to open-ocean conditions. The inshore stations (P04–P16) are characterised by the classical seasonal cycle of spring and late summer blooms (production >3 g C m⁻² d⁻¹), diatoms are not Fe-stressed, and growth rate is probably controlled by macronutrient supply. The fate of the phytoplankton is likely sedimentation by diatom-dominated spring blooms, with a pelagic recycling system predominating at other times. The offshore stations (P20/OSP) display low seasonality in biomass and production (OSP, mean winter production 0.3 g C m⁻² d⁻¹, mean spring/summer production 0.85 g C m⁻² d⁻¹), and are dominated by small algal cells. Low Fe availability prevents the occurrence of diatom blooms observed inshore. The main fate of phytoplankton is probably recycling through the microbial food web, with relatively low sedimentation compared to inshore. However, the supply of macro- and micro-nutrients to the coastal and open ocean, respectively, may vary between years. Variability in macro-nutrient supply to the coastal ocean may result in decreased winter reserve nitrate, summer nitrate limitation, subsequent floristic shifts towards small cells, and reduced primary production. Offshore, higher diatom abundances are occasionally observed, perhaps indicating episodic Fe supply. The two distinct oceanic regimes have different phytoplankton dynamics resulting in different seasonality, community structure and fate of algal carbon. These differences will strongly influence the biogeochemical signatures of the coastal and open-oceanic NE subarctic Pacific.     

Microzooplankton grazing impact in the Western Arctic Ocean

Microzooplankton grazing impact on phytoplankton was assessed using the Landry–Hassett dilution technique in the Western Arctic Ocean during spring and summer 2002 and 2004. Forty experiments were completed in a region encompassing productive shelf regions of the Chukchi Sea, mesotrophic slope regions of the Beaufort Sea off the North Slope of Alaska, and oligotrophic deep-water sites in the Canada Basin. A variety of conditions were encountered, from heavy sea-ice cover during both spring cruises, moderate sea-ice cover during summer of 2002, and light to no sea ice during summer of 2004, with a concomitant range of trophic conditions, from low chlorophyll-a (Chl-a; <0.5 μg L⁻¹) during heavy ice cover in spring and in the open basin, to late spring and summer shelf and slope open-water diatom blooms with Chl-a >5 μg L⁻¹. The microzooplankton community was dominated by large naked ciliates and heterotrophic gymnodinoid dinoflagellates. Significant, but low, rates of microzooplankton herbivory were found in half of the experiments. The maximum grazing rate was 0.16 d⁻¹ and average grazing rate, including experiments with no significant grazing, was 0.04±0.06 d⁻¹. Phytoplankton intrinsic growth rates varied from the highest values of about 0.4 d⁻¹ to the lowest values of zero to slightly negative growth, on average 0.16±0.15 d⁻¹. Light limitation in spring and post-bloom senescence during summer were likely explanations of observed low phytoplankton growth rates. Microzooplankton grazing consumed 0–120% (average 22±26%) of phytoplankton daily growth. Grazing and growth rates found in this study were low compared to rates reported in another Arctic system, the Barents Sea, and in major geographic regions of the world ocean.     

Iron and mixing affect biological carbon uptake in SOIREE and EisenEx,two Southern Ocean iron fertilisation experiments

This study explores the changes in the surface water fugacity of carbon dioxide (fCO₂) and biological carbon uptake in two Southern Ocean iron fertilisation experiments with different hydrographic regimes. The Southern Ocean Iron Release Experiment (SOIREE) experiment was carried out south of the Antarctic Polar Front (APF) at 61°S, 141°E in February 1999 in a stable hydrographic setting. The EisenEx experiment was conducted in a cyclonic eddy north of the APF at 48°S, 21°E in November 2000 and was characterised by a rapid succession of low to storm-force wind speeds and dynamic hydrographic conditions. The iron additions promoted algal blooms in both studies. They alleviated algal iron limitation during the 13-day SOIREE experiment and probably during the first 12 days of EisenEx. The fCO₂ in surface water decreased at a constant rate of 3.8 μatm day⁻¹ from 4 to 5 days onwards in SOIREE. The fCO₂ reduction was 35 μatm after 13 days. The evolution of surface water fCO₂ in the iron-enriched waters (or ‘patch’) displayed a saw tooth pattern in EisenEx, in response to algal carbon uptake in calm conditions and deep mixing and horizontal dispersion during storms. The maximum fCO₂ reduction was 18–20 μatm after 12 and 21 days with lower values in between. The iron-enriched waters in EisenEx absorbed four times more atmospheric CO₂ than in SOIREE between 5 and 12 days, as a result of stronger winds. The total biological uptake of inorganic carbon across the patch was 1389 ton C (±10%) in SOIREE and 1433 ton C (±27%) in EisenEx after 12 days (1 ton=10⁶ g). This similarity probably reflects the comparable size of the iron additions, as well as algal growth at a similar near-maximum growth rate in these regions. The findings imply that the different mixing regimes had less effect on the overall biological carbon uptake across the iron-enriched waters than suggested by the evolution of fCO₂ in surface water.     

Dissolved organic matter in the subterranean estuary of a volcanic island,Jeju: Importance of dissolved organic nitrogen fluxes to the ocean

We observed the origin, behavior, and flux of dissolved organic carbon (DOC), dissolved organic nitrogen (DON), colored dissolved organic matter (CDOM), and dissolved inorganic nitrogen (DIN) in the subterranean estuary of a volcanic island, Jeju, Korea. The sampling of surface seawater and coastal groundwater was conducted in Hwasun Bay, Jeju, in three sampling campaigns (October 2010, January 2011, and June 2011). We observed conservative mixing of these components in this subterranean environment for a salinity range from 0 to 32. The fresh groundwater was characterized by relatively high DON, DIN, and CDOM, while the marine groundwater showed relatively high DOC. The DON and DIN fluxes through submarine groundwater discharge (SGD) in the groundwater of Hwasun Bay were estimated to be 1.3 × 10⁵ and 2.9 × 10⁵ mol d^− 1, respectively. In the seawater of Hwasun Bay, the groundwater-origin DON was almost conservative while about 91% of the groundwater-origin DIN was removed perhaps due to biological production. The DON flux from the entire Jeju was estimated to be 7.9 × 10⁸ mol yr^− 1, which is comparable to some of the world's large rivers. Thus, our study highlights that DON flux through SGD is potentially important for delivery of organic nitrogen to further offshore while DIN is readily utilized by marine plankton in near-shore waters under N-limited conditions.     

Isotopic and elemental indicators of nutrient sources and status of coastal habitats in the Caribbean Sea,Yucatan Peninsula,Mexico

Nutrient inputs associated with coastal population growth threaten the integrity of coastal ecosystems around the globe. In order to assess the threat posed by rapid growth in tourism, we analyzed the nutrient concentrations as well as the δ¹⁵N of NO₃⁻ and macrophytes to detect wastewater nitrogen (N) at 6 locations along a groundwater-dominated coastal seagrass bed on the Caribbean coast of Mexico. We predicted that locations with greater coastal development would have higher concentrations of dissolved inorganic nitrogen (DIN) and phosphorus (P), as well as δ¹⁵N of NO₃⁻, reflecting wastewater sources of N. However, concentrations of NO₃⁻ were not significantly different between developed (3.3 ± 5.3 μM NO₃⁻) and undeveloped (1.1 ± 0.7 μM) marine embayments. The most important control on DIN concentration appeared to be mixing of fresh and salt water, with DIN concentrations negatively correlated with salinity. The δ¹⁵N of NO₃⁻ was elevated at an inland pond (7.0 ± 0.42‰) and a hydrologically-connected tide pool (7.6 ± 0.57‰) approximately 1 km downstream of the pond. The elevated δ¹⁵N of NO₃⁻ at the pond was paralleled by high δ¹⁵N values of Cladophora sp., a ubiquitous green alga (10 ± 1‰). We hypothesize that inputs of nitrogen rich (NO₃⁻ > 30 μM) groundwater, characterized by ¹⁵N enriched signatures, flow through localized submarine groundwater discharges (SGD) and contribute to the elevated δ¹⁵N signatures observed in many benthic macrophytes. However, changes in nitrogen concentrations and isotope values over the salinity gradient suggest that other processes (e.g. denitrification) could also be contributing to the ¹⁵N enrichments observed in primary producers. More measurements are needed to determine the relative importance of nitrogen transformation processes as a source of ¹⁵N to groundwaters; however, it is clear that continued inputs of anthropogenic N via SGD have the potential to severely impact ecologically and economically valuable seagrass meadows and coral reefs along the Caribbean coast of Mexico.     

Biogeochemical study of a coccolithophore bloom in the northern Bay of Biscay (NE Atlantic Ocean) in June 2004

The present paper synthesizes data obtained during a multidisciplinary cruise carried out in June 2004 at the continental margin of the northern Bay of Biscay. The data-set allows to describe the different stages of a coccolithophore bloom dominated by Emiliania huxleyi. The cruise was carried out after the main spring phytoplankton bloom that started in mid-April and peaked in mid-May. Consequently, low phosphate (PO₄ < 0.2 μM) and silicate (DSi < 2.0 μM) concentrations, low partial pressure of carbon dioxide (pCO₂) and high calcite saturation degree in surface waters combined with thermal stratification, probably favoured the blooming of coccolithophores. During the period of the year our cruise was carried out, internal tides induce enhanced vertical mixing at the continental shelf break leading to the injection of inorganic nutrients to surface waters that probably trigger the bloom. The bloom developed as the water-column stratified and as the water mass was advected over the continental shelf, following the general residual circulation in the area. The most developed phase of the bloom was sampled in a remote sensed high reflectance (HR) patch over the continental shelf that was characterized by low chlorophyll-a (Chl-a) concentration in surface waters (<1.0 μg L^?1), high particulate inorganic carbon (PIC) concentration (～8 μmol L^?1) and coccolithophore abundance up to 57 × 10⁶ cells L^?1. Transparent exopolymer particles (TEP) concentrations ranged between 15 and 75 μg C L^?1 and carbon content of TEP represented up to 26% of the particulate organic carbon (POC; maximum concentration of 15.5 μmol L^?1 in the upper 40 m). Integrated primary production (PP) ranged between 210 and 680 mg C m^?2 d^?1 and integrated calcification (CAL) ranged between 14 and 140 mg C m^?2 d^?1, within the range of PP and CAL values previously reported during coccolithophore blooms in open and shelf waters of the North Atlantic Ocean. Bacterial protein production (BPP) measurements in surface waters (0.3–0.7 μg C L^?1 h^?1) were much higher than those reported during early phases of coccolithophore blooms in natural conditions, but similar to those during peak and declining coocolithophorid blooms reported in mesocosms. Total alkalinity anomalies with respect to conservative mixing (ΔTA) down to ?49 μmol kg^?1 are consistent with the occurrence of biogenic precipitation of calcite, while pCO₂ remained 15–107 μatm lower than atmospheric equilibrium (372 μatm). The correlation between ΔTA and pCO₂ suggested that pCO₂ increased in part due to calcification, but this increase was insufficient to overcome the background under-saturation of CO₂. This is related to the biogeochemical history of the water masses due to net carbon fixation by the successive phytoplankton blooms in the area prior to the cruise, hence, the investigated area remained a sink for atmospheric CO₂ despite calcification.     

Algal concentration profiles above mussel beds

The distribution of phytoplankton was measured in vertical profiles at three stations in Limfjorden (Denmark) within and above Mytilus edulis beds (−5 to 200 cm above the bed surface) and related to the filtration capacity of the mussels and hydrodynamic conditions. The density of phytoplankton was 2690 to 10 600 cells cm⁻³ at 200 cm above the bottom (∼1.4 to 3.2 mg Chl-a m⁻³), declining towards the bottom to about 1000 cells cm⁻³ at two of the stations. This is a threshold of minimum algal concentration below which mussels are reported to stop feeding. The results of the present study further demonstrate that the near-bed phytoplankton concentrations observed in the field often remained below the concentration level at which mussels reduce their maximal filtration activity.     

Export fluxes of particulate organic carbon in the Chukchi Sea: A comparative study using 234Th/238U disequilibria and drifting sediment traps

Large-volume sampling of ²³⁴Th and drifting sediment trap deployments were conducted as part of the 2004 Western Arctic Shelf–Basin Interactions (SBI) spring (May 15–June 23) and summer (July 17–August 26) process cruises in the Chukchi Sea. Measurements of ²³⁴Th and particulate organic carbon (POC) export fluxes were obtained at five stations during the spring cruise and four stations during the summer cruise along Barrow Canyon (BC) and along a parallel shelf-to-basin transect from East Hanna Shoal (EHS) to the Canada Basin. ²³⁴Th and POC fluxes obtained with in situ pumps and drifting sediment traps agreed to within a factor of 2 for 70% of the measurements. POC export fluxes measured with in situ pumps at 50 m along BC were similar in spring and summer (average = 14.0 ± 8.0 mmol C m^− 2 day^− 1 and 16.5 ± 6.5 mmol C m^− 2 day^− 1, respectively), but increased from spring to summer at the EHS transect (average = 1.9 ± 1.1 mmol C m^− 2 day^− 1 and 19.5 ± 3.3 mmol C m^− 2 day^− 1, respectively). POC fluxes measured with sediment traps at 50 m along BC were also similar in both seasons (31.3 ± 9.3 mmol C m^− 2 day^− 1 and 29.1 ± 14.2 mmol C m^− 2 day^− 1, respectively), but were approximately twice as high as POC fluxes measured with in situ pumps. Sediment trap POC fluxes measured along the EHS transect also increased from spring to summer (3.0 ± 1.9 mmol C m^− 2 day^− 1 and 13.0 ± 6.4 mmol C m^− 2 day^− 1, respectively), and these fluxes were similar to the POC fluxes obtained with in situ pumps. Discrepancies in POC export fluxes measured using in situ pumps and sediment traps may be reasonably explained by differences in the estimated POC/²³⁴Th ratios that arise from differences between the techniques, such as time-scale of measurement and size and composition of the collected particles. Despite this variability, in situ pump and sediment trap-derived POC fluxes were only significantly different at a highly productive station in BC during the spring.     

Nutrient and phytoplankton dynamics off the west coast of Vancouver Island during the 1997/98 ENSO event

Six research cruises were conducted off the west coast of Vancouver Island between April and October of 1997 and 1998 as part of the Canadian GLOBEC project to compare nutrient and phytoplankton dynamics between ENSO (1997) and non-ENSO (1998) years. Limited sampling also was conducted during three cruises in 1999. During the 1997 ENSO period, there was a shallow thermocline (∼10 m) that resulted in a shallower mixed layer, lower salinity and density, and stronger summer stratification. In general on the shelf, the 1997 growing season was characterized by higher nitrate (7.5 μM) and silicic acid (17 μM) concentrations, lower total chlorophyll (∼76 mg m⁻²), lower phytoplankton carbon biomass (0.2 mg C L⁻¹), and lower diatom abundance and biomass than in 1998. Phytoplankton assemblages were dominated by nanoplankton in 1997 and by diatoms in 1998. These results suggest that the 1997 ENSO was responsible for a reduction in the growth and biomass of larger phytoplankton cells. In mid-1998, the hydrographic characteristics off the west coast of Vancouver Island changed suddenly. The 1997 poleward transport of warm water reversed to an equatorward transport of coastal water in July 1998, which was accompanied by normal summer upwelling. During 1998, a large diatom bloom (mainly dominated by Chaetoceros debilis, Leptocylindrus danicus and to a lesser extent by Skeletomema and Pseudo-nitzschia sp.) was observed in July over the continental shelf. This large bloom resulted in chlorophyll concentrations of up to 400 mg m⁻², primary productivity of up to 11 g C m⁻² d⁻¹, and near undetectable dissolved nitrogen concentrations at some of the shelf stations in 1998. In contrast, during 1997, the sub-tropical waters that were advected over the slope, resulted in low chlorophyll a and primary productivity (generally <1 g C m⁻² d⁻¹). Therefore, there was a sharp contrast between the very high primary productivity on the shelf in July 1998, due to normal nutrient replenishment from summer upwelling and outflow from the Strait of Juan de Fuca, and the lower primary productivity during the 1997 ENSO year. During 1998, non-ENSO conditions resulted in phytoplankton biomass that was twice as high on the shelf as that measured in regions beyond the continental shelf of the west coast of Vancouver Island.     

Abundance,biomass and composition of the sea ice biota of the Greenland Sea pack ice

During two cruises to the Greenland Sea, we studied the abundance and biomass of the sea ice biota in summer and late autumn. The mean calculated biomass of the sympagic community was 0.2 g C m⁻² ice. Algae contributed on average 43% to total biomass, followed by bacteria (31%), heterotrophic flagellates (20%), and meiofauna (4%). Diatoms were the main primary producers (60% of total algal biomass), but flagellated cells contributed significantly to the algal biomass. Among the meiofauna, ciliates, nematodes, acoel turbellarians and crustaceans were dominant. Calculated potential ingestion rates of meiofauna (0.6 g C m⁻² (120 d)⁻¹) are on the same order of magnitude as annual primary production estimates for Arctic multi-year sea ice. We therefore assume that grazing can control biomass accumulation of primary producers inside the sea ice.     

Light absorption by phytoplankton,non-algal particles and dissolved organic matter at the Patagonia shelf-break in spring and summer

Satellite image studies and recent in situ sampling have identified conspicuous phytoplankton blooms during spring and summer along the Patagonia shelf-break front. The magnitudes and spectral characteristics of light absorption by total particulate matter (phytoplankton and detritus) and colored dissolved organic matter (CDOM) have been determined by spectrophotometry in that region for spring 2006 and late summer 2007 seasons. In spring, phytoplankton absorption was the dominant optical component of light absorption (60–85%), and CDOM showed variable and important contributions in summer (10–90%). However, there was a lack of correlation between phytoplankton biomass (chlorophyll-a concentration or [chl a]) and the non-algal compartment in both periods. A statistically significant difference was found between the two periods with respect to the CDOM spectral shape parameter (S_cdom), with means of 0.015 (spring) and 0.012 nm^?1 (summer). Nonetheless, the mean S_cdm values, which describe the slope of detritus plus CDOM spectra, did not differ between the periods (average of 0.013 nm^?1). Phytoplankton absorption values in this work showed deviations from mean parameterizations in previous studies, with respect to [chl a], as well as between the two study periods. In spring, despite the microplankton dominance, high specific absorption values and large dispersion were found (a*_ph(440)=0.04±0.03 m² mg [chl a]^?1), which could be attributed to an important influence of photo-protector accessory pigments. In summer, deviations from general trends, with values of a*_ph(440) even higher (0.09±0.02 m² mg [chl a]^?1), were due to the dominance of small cell sizes and also to accessory pigments. These results highlight the difficulty in deriving robust relationships between chlorophyll concentration and phytoplankton absorption coefficients regardless of the season period. The validity of a size parameter (S_f) derived from the absorption spectra has been demonstrated and was shown to describe the size structure of phytoplankton populations, independently of pigment concentration, with mean values of 0.41 in spring and 0.72 in summer. Our results emphasize the need for specific parameterization for the study region and seasonal sampling approach in order to model the inherent optical properties from water reflectance signatures.     

Toxic effects of harmful benthic dinoflagellate Ostreopsis ovata on invertebrate and vertebrate marine organisms

Harmful benthic microalgae blooms are an emerging phenomenon causing health and economic concern, especially in tourist areas. This is the case of the Mediterranean Sea, where Ostreopsis ovata blooms occur in summer, with increasing regularity. Ostreopsis species produce palytoxin (PTX) and analogues, and a number of deaths directly associated with the ingestion of PTX contaminated seafood have been reported. PTX is considered one of the most toxic molecules occurring in nature and can provoke severe and sometimes lethal intoxications in humans. So far in temperate areas, O. ovata blooms were reported to cause intoxications of humans by inhalation and irritations by contact.In addition, invertebrate mass mortalities have been reported, possibly linked to O. ovata blooms, although other causes cannot be ruled out, such as oxygen depletion or high seawater temperature. In order to improve our knowledge about the direct toxicity of this species on invertebrate and vertebrate marine organisms, we performed an ecotoxicological screening to investigate the toxic effects of different concentrations of O. ovata (cultured in the laboratory and sampled in the field during blooms) on crustaceans and fish as model organisms. Artemia salina, Tigriopus fulvus, and Amphibalanus amphitrite larvae and juveniles of the sea bass Dicentrarchus labrax were used as model species. Toxic effects associated with cultured O. ovata cells were investigated using a crossed design: testing two different temperatures (20 and 25 °C), four different cell concentrations, and four treatments (untreated O. ovata culture, filtered and resuspended algal cells, growth medium devoid of algal cells, and sonicated algal cells). The results indicate that the toxicity of cultured O. ovata is related to the presence of living O. ovata cells, and that this effect is amplified by temperature. Furthermore, both tests with laboratory cultured algae and field sampled cells pointed out that A. salina is the most sensitive species even at concentrations below the Environmental Alarm Threshold set by the Italian Ministry of Health.Some possible explanations of such sensitivity are discussed, taking into account evidence of O. ovata cells ingestion and the activity of its toxins on the Na⁺/K⁺-ATPase.     

Biological versus physical processes as drivers of large oscillations of the air–sea CO2 flux in the Antarctic marginal ice zone during summer

The fugacity of CO₂ and abundance of chlorophyll a (Chla) were determined in two long transects from the Polar Front to the Antarctic Continent in austral summer, December 1995–January 1996. Large undersaturations of CO₂ in the surface water were observed coinciding with high Chla content. In the major hydrographic regions the mean air–sea fluxes were found to range from −3 to +7 mmol m⁻² d⁻¹ making these regions act as a sink as well as a source for CO₂. In the total 40-d period, the summation of the several strong source and sink regions revealed an overall modest net source of 0.3 mmol m⁻² d⁻¹, this based on the Wanninkhof (J. Geophys. Res. 97 (1992) 7373) quadratic relationship at in situ windspeed. A simple budget approach was used to quantify the role of phytoplankton blooms in the inorganic carbonate system of the Antarctic seas in a time frame spanning several weeks. The major controlling physical factors such as air–sea flux, Ekman pumping and upwelling are included. Net community production varies between −9 and +7 mmol m⁻² d⁻¹, because of the large oscillations in the dominance of autotrophic (CO₂ fixation) versus heterotrophic (CO₂ respiration) activity. Here the mixed layer depth is the major controlling factor. When integrated over time the gross influx and efflux of CO₂ from air to sea is large, but the net residual air/sea exchange is a modest efflux from sea to atmosphere.     

The influence of phytoplankton assemblage composition on biogeochemical characteristics and cycles in the southern Ross Sea,Antarctica

To test the hypothesis that phytoplankton assemblages dominated by different taxa have distinct biogeochemical characteristics and cycles, the temporal and spatial variations in phytoplankton biomass and composition were studied within the Ross Sea polynya, where diatoms and the haptophyte Phaeocystis antarctica are thought to have spatially distinct distributions. Two cruises were completed, with the first conducted in spring, 1994, and the second in late spring–early summer, 1995/1996. Ice concentrations decreased substantially from spring to summer. Mixed layer depths for the region decreased markedly in early spring and were relatively invariant thereafter; the strength of the stratification varied both in time and space. Mixed layers were greater in spring in assemblages dominated by diatoms (as determined by HPLC pigment concentrations) than those dominated by Phaeocystis antarctica, whereas in summer no difference was observed. Nutrient concentrations were initially high and near winter values, but decreased throughout November and December. Nitrate : phosphate removal ratios varied widely, with ratios exceeding 20 in spring but decreasing below 14 in summer. N : P removal ratios at stations dominated by diatoms were less than the Redfield ratio in both spring and summer, and at those stations dominated by P. antarctica the N : P removal ratio was ca. 19 in both seasons. Chlorophyll and particulate matter concentrations increased as nutrients decreased. Spatial and temporal variations of phytoplankton pigments occurred, with 19′-hexanoylfucoxanthin, a pigment of P. antarctica, exceeding 3.9 μg l⁻¹ during spring in the south-central polynya, and fucoxanthin, an accessory pigment of diatoms, found in concentrations >1 μg l⁻¹ in the western Ross Sea. The distributions were not mutually exclusive, and concentrations of both pigments were greatest in spring. The early growth of P. antarctica appears to be related to earlier stratification and disappearance of ice from the south-central Ross Sea. Ratios of FUCO/CHL were relatively invariant, but substantial changes in the HEX/CHL and POC/CHL ratios were observed through time. A one-dimensional nitrogen budget for the spring–early summer period suggests that much of the surface production was partitioned into particles, most (53%) of which remained in the upper 200 m. The rest was partitioned into dissolved organic matter (14%), remineralized as ammonium (19%), or sank from the surface layer as particles (13%). The region may serve as a useful analog to other polar systems, and an understanding of the processes controlling assemblage composition, production, and biomass accumulation may provide insights into biogeochemical cycles of other Antarctic environments.     

Xanthophyll cycling in Phaeocystis globosa and Thalassiosira sp.: a possible mechanism for species succession

The ability of phytoplankton species to initiate photo-protective mechanisms and the rates by which they do so have been suggested to be partly responsible for species succession. To examine whether this is also true in the case of diatom spring blooms preceding Phaeocystis globosa, cultures of P. globosa and Thalassiosira sp. were investigated under controlled laboratory conditions for differences in their xanthophyll cycling rates and abilities. It was found that P. globosa exhibited active and rapid xanthophyll cycling when cultures photoacclimated to 10, 50 and 100 μmol quanta m⁻² s⁻¹, were shifted to 150 and 250 μmol quanta m⁻² s⁻¹. The early spring diatom Thalassiosira only exhibited xanthophyll cycling when acclimated to 10 μmol quanta m⁻² s⁻¹. P. globosa always exhibited faster xanthophyll cycling rates than Thalassiosira, giving P. globosa a possible competitive edge over Thalassiosira sp. It was also found that the magnitude of xanthophyll cycling correlates with the intensity of light changes during the one-hour xanthophyll cycling experiments, and thus may be regarded as related to photoacclimation.     

High short-term variability of CO2 fluxes during an upwelling event off the Chilean coast at 30°S

pH and alkalinity measurements from a coastal upwelling area located near 30°S (Coquimbo, Chile), are used to describe the short-term variations of CO₂ air–sea exchanges over a period of one week in summer 1996. A 180 km ocean–coastal transect, together with two almost-synoptic grid surveys off Coquimbo covering approximate 2500 km² each, showed that during and immediately after a 4 day long southwesterly wind event (24–28 January) a large area of cold surface water (≈14°C), highly supersaturated in CO₂ (fCO₂ up to 900 μatm), was located near the coast. Three days after the end of the event, the second grid survey showed that in most of the study area the surface temperature and pH had increased significantly (by 1–3°C and 0.05–0.2, respectively), and that the surface water was no longer supersaturated in CO₂. The CO₂-supersaturated water observed in the first grid survey was identified as upwelled subsurface equatorial water, a water mass with its core at about 200 m depth: the depth from which the water upwells is a major determinant of the surface water fCO₂. Integrated C fluxes within a 20 km wide coastal strip (1900 km²) indicate a strong outgassing of CO₂ from the ocean under upwelling conditions (Grid 1; 121 t C day^-1), while the net C exchange was directed to the ocean during the relaxation period (Grid 2; 19 t C day^-1). Estimates of CO₂ fluxes in upwelling areas based on surface water fCO₂ measurements must therefore take into account these short-term variations: reliance on longer-term averages and interpolation will lead to erroneous results.     

Annual and spatial variation in the abundance length and condition of juvenile turbot (Psetta maxima L.) on nursery grounds on the west coast of Ireland: 2000–2007

Turbot (Psetta maxima Linnaeus) is a high value commercially exploited marine flatfish which occurs in European waters, from the Northeast Atlantic to the Arctic Circle, the Baltic and Mediterranean Sea. In Ireland, turbot are the most valuable commercial non-quota species. Very little is known about their population dynamics in the wild, in particular during the sandy beach nursery phase of the life history. In 2000, a survey was established to assess flatfish species on nursery grounds on the west coast of Ireland. Eleven sandy beaches were assessed for 0+ turbot by beach seining, over an eight year period (2000–2007) during the months of August and September. The objective of the study was to estimate juvenile turbot abundance and size structure to determine if any spatial and annual trends existed. Large scale variability in the recruitment of fish to nursery grounds may be indicative of fluctuations in the adult stock. Turbot were found to recruit to five beaches consistently over the eight year period. Temporal and spatial variability in the relative abundance and length of turbot was discerned, with no apparent overall trend. However, certain nursery grounds were shown in most of the years examined to support higher abundances of turbot in comparison to other areas over the eight year period. Turbot abundances on nursery grounds were significantly correlated with mean spring sea temperatures during the pelagic stage. The condition of turbot did not significantly differ on an annual or spatial scale. Mean densities of 0+ turbot along the Irish coast were found to be similar and at times higher than other areas in Europe, ranging from 0.1 (± 0.3) individuals 1000 m^− 2 to 18.5 (± 6.9) individuals 1000 m^− 2. Mean turbot total length on beaches ranged from 3.8 cm (± 0.6) to 6.6 cm (± 4.3). The observed spatial and temporal variability in abundance and length highlights the need for long-term studies when assessing juvenile flatfish populations. Results from the present study have provided much needed baseline data on wild juvenile turbot populations which is severely lacking for this species both on an Irish and on a European scale.     

Winter and spring phytoplankton composition and production in a shallow eutrophic Baltic lagoon

Taxonomic composition and productivity of winter and spring phytoplankton in a eutrophic estuary have been investigated in order to elucidate the carbon flux under conditions of limitation by physical factors – light and temperature. In spite of the important differences in nutrients, solar radiation and water temperature between winter and spring season, mean concentrations of particulate organic carbon were equal to 13.2 and 13.0 mgC l⁻¹, respectively. Chlorophyll a averaged at 79 μgChl l⁻¹ in winter, that is 69% of spring. Although community respiration accounted for only 6–26% of light saturated photosynthesis, integrated net primary production of the 1.2 m deep water column was negative until April. High attenuation of the water body (K_o = 2.9 m⁻¹) lead to a negative carbon balance (net heterotrophy) below 35 cm for all sampling dates. Thus, the high winter POC and phytoplankton values can only originate from summer or autumn primary production. This assumption was supported by a carbon loss rate of just 3% of total organic carbon per day for the whole water column. The composition of phytoplankton was very constant through both seasons: 39% Chlorophyceae, 33% Cyanobacteria and 25% Bacillariophyceae. As expected, phytoplankton was low light acclimated, having high α values (slope of light limited photosynthesis), but moderate maximum photosynthesis rates at saturating irradiances, which were heavily affected by temperature. Calculation of net carbon flux yet showed net heterotrophy of the Bodden waters in winter and early spring were caused by external physical limitation (low surface irradiance and low temperature) in combination with a high light attenuation of the water body.     

Seasonal patterns in the fish and epibenthic crustaceans community of an intertidal zone with particular reference to the population dynamics of plaice and brown shrimp

The intertidal zone of a sandy beach located on the French coast of the Eastern Channel, was sampled during spring and summer 2000 to analyse the community structure of fish and epibenthic crustaceans. The presence of many juvenile fish (mainly O-group) and crustaceans indicated the important role played by the intertidal zone as a nursery ground. The brown shrimp, Crangon crangon and O-group plaice, Pleuronectes platessa are the two most abundant species of the intertidal ecosystem.Plaice settlement period extended from mid-March to early June. Over the survey period, densities increased to maximum numbers of about 27 ind. 10 m⁻² in mid-April. In the following week, density rapidly decreased due to mortality and migration into deeper waters. The mortality was attributed mainly to predation by brown shrimp (C. crangon) and to a lesser extent by the shore crab (Carcinus maenas). The mean size of 0-group plaice increased from 19 mm in mid-April to 58 mm in July. Growth of juvenile 0-group plaice is described by an exponential equation: total length (TL, mm) = 12.602 e^{0.022 (post-settlement age)}. Post-settlement growth rates, estimated by otolith microstructure analysis, were 0.38 mm d⁻¹ for plaice ≤30 mm and 0.61 mm d⁻¹ for plaice >30 mm. Settlement of juvenile brown shrimp started in mid-April, peaked in early June (93 ind. 10 m⁻²) and continued with fluctuating intensity throughout the summer. Growth rate of juvenile C. crangon, estimated after the settlement peak, was 0.163 mm d⁻¹. Growth conditions of juvenile plaice and C. crangon were analysed by comparing estimated growth in the field with predicted maximum growth according to temperature-growth rate models from experimental studies of growth with unlimited food supply. For plaice, the estimated growth rate was lower (plaice ≤30 mm) but similar (plaice >30 mm) to the predicted maximum growth suggesting a food limitation only for newly settled individual. The observed increase in mean length of juvenile C. crangon was lower than the maximum possible growth. The function of the intertidal zone in the early juvenile stages of marine species is discussed.     

Nitrogen sources for new production in the NE Arabian Sea

New productivity measurements using the ¹⁵N tracer technique were conducted in the north-eastern (NE) Arabian Sea during six expeditions from 2003 to 2007, mostly in winter. Our results indicate that the NE Arabian Sea has a potential for higher new productivity during blooms. Nitrate uptake by plankton is the highest during late winter. New productivity and f-ratios in the NE Arabian Sea are mainly controlled by hydrodynamic and meteorological parameters such as wind strength, sea surface temperature (SST), mixed layer depth (MLD) and mixed layer nitrate. Deepening of the mixed layer supplies nitrate from below, which supports the observed nitrogen uptake. Higher f-ratios during blooms indicate the strong coupling between surface layers and sub-surface layers. Deepening of mixed layer below 100 m (from its inter-monsoon value between 30 and 40 m) transferred often more than 100 mmol N–NO₃ m^? 2 into the surface layers from below. The observed winter blooms in the region are supported by such input and are sustained for more than a month. Higher new productivity has been found in late winter, whereas transport of nitrate is maximum in early winter. In general, new production varies progressively during winter. Diurnal cycling of the mixed layer could be the reason for the under utilization of entrained nitrate during early winter. New productivity values and wind strength show significant differences during Feb–Mar 03 and Feb–Mar 04. These differences indicate that the winter cooling and parameters related the biological productivity also vary inter-annually. However, the difference between the new productivity values between Feb–Mar 03 and Feb–Mar 04 is much lower than the difference between Jan 03 and Feb–Mar 03. The results suggest that amplitude of seasonal variation is higher than the inter-annual variation in the region. During spring, Fickian diffusive fluxes of nitrate into the surface layer range from 0.51 to 1.38 mmol N–NO₃ m^? 2 day^? 1, and can account for 67% and 78% of the observed nitrogen uptake in the coastal and open ocean regions, respectively. We document the intra-seasonal and inter-annual variations in new productivity during winter and identify sources of nitrate which support the observed productivity during spring.