Bacterioplankton and phytoplankton biomass and production during summer stratification in the northwestern Mediterranean Sea

We examined bacterioplankton biomass and heterotrophic production (BHP) during summer stratification in the northwestern Mediterranean in four successive stratification seasons (June–July of 1993–1996). Values of phytoplankton biomass and primary production were determined simultaneously so that the data sets for autotrophic and heterotrophic microbial plankton could be compared. Three standard stations were set along a transect from Barcelona to the channel between Mallorca and Menorca, representing coastally influenced shelf waters, frontal waters over the slope front, and open sea waters. Conversion factors from ³H-leucine incorporation to BHP were empirically determined and varied between 0.29 and 3.25 kg C mol^-1. Bacterial biomass values were among the lowest found in any marine environment. BHP values (between 0.02 and 2.5 μg C L^-1 d^-1) were larger than those of low nutrient low chlorophyll areas such as the Sargasso Sea and lower than those from high nutrient low chlorophyll areas such as the equatorial Pacific. Growth rates of bacterioplankton were highest at the slope front (0.20 d^-1) and lowest at the open sea station (0.04 d^-1). Phytoplankton growth rates were similar at the three stations (∼0.50 d^-1). Integrated values of bacterioplankton biomass, BHP and bacterial growth rates did not show significant differences among years, but differences between the three stations were clearly significant. Phytoplankton biomass, primary production, and phytoplankton growth rates did not show significant differences either with year or with station. As a consequence the bacterioplankton to phytoplankton biomass (BB/BPHY) and production (BHP/PP) ratios varied from the coastal to the open sea stations. The BB/BPHY ratio was 0.98 at the coast and ∼0.70 at the other two stations. These ratios are similar to those found in other oligotrophic marine environments. The BHP/PP ratio was 0.83 at the coast, 0.36 at the slope and 0.09 at the open sea station. The last value is also similar to values found in other oligotrophic marine environments. Vertical distribution of these ratios was also examined.The comparison of microbial parameters at the three stations indicates a different kind of relationship between bacterioplankton and phytoplankton in oligotrophic open sea waters and in coastal, nutrient-richer waters. According to such parameters and to the values of the BB/BPHY and BHP/PP ratios, open waters in the northwestern Mediterranean (despite their relatively short distance from the shore) were intermediate between the extremely oligotrophic waters of the eastern Mediterranean or the Sargasso Sea and the more productive waters of the equatorial Pacific.     

The zooplankton community in the Greenland Sea: Composition and role in carbon turnover

The micro- and mesozooplankton communities in surface waters of the Greenland Sea are described based on data from five cruises covering an annual cycle. Special emphasis is given to the summer period (June and August), prior to and after the descent of Calanus spp. Calanus spp. dominated the copepod community during the spring bloom and in the beginning of the summer. However, during the summer, there was a pronounced shift in the zooplankton composition in the euphotic zone. In contrast to what has been observed in other Arctic systems, smaller genera such as Pseudocalanus spp., Oncaea spp. and Oithona spp. became abundant and the total copepod biomass remained high after the Calanus spp. descended for hibernation. The peak protozooplankton biomass in the Greenland Sea (June) co-occurred with the peak in Calanus spp. Protozooplankton biomass then decreased during the summer. Growth of protozooplankton and grazing rates of the two dominating non-Calanus genera, Oithona and Pseudocalanus, were measured. For both copepod genera, protozooplankton constituted 40% or more of the diet, and maximum clearance was on prey items with an equivalent spherical diameter between 15 and 30 μm. The non-Calanus components of the zooplankton community were responsible for 70–99% of the total zooplankton grazing on phytoplankton during summer and were crucial for the recycling and respiration of primary production.     

Impact of open-ocean convection on nutrients,phytoplankton biomass and activity

We describe the impact of an open-ocean convection event on nutrient budgets, carbon budget, elemental stoichiometry, phytoplankton biomass and activity in the Northwestern Mediterranean Sea (NWM). In the convective episode examined here we estimated an input of nutrients to the surface layer of 7.0, 8.0 and 0.4×10⁸ mol of silicate, nitrate and phosphate, respectively. These quantities correspond to the annual nutrient input by river discharges and atmospheric depositions in the Gulf of Lion. Such nutrient input is sufficient to sustain new primary production from 46 to 63 g C m⁻² y⁻¹, which is the same order of magnitude found in the NWM open waters. Our results together with satellite data analysis, propose new scenarios that explain the origin of the spring phytoplankton bloom occurring in NWM.     

Bacterioplankton activity in the surface waters of the Arabian Sea during and after the 1994 SW monsoon

Bacterial biomass and production were measured on two cruises to the northwestern Arabian Sea in 1994; the first cruise took place towards the end of the SW monsoon in September, and the second cruise during the inter-monsoon period in November and December. Although phytoplankton production was significantly higher during the monsoon, bacterial numbers showed little difference. Bacteria were most abundant in the euphotic zone and highest bacterial numbers were measured during the monsoon period in the Gulf of Oman and the shelf waters off southern Oman; in these regions, numbers ranged from 0.9 to 1.6×10⁹ bacteria l^-1. On both cruises, bacteria were less abundant in the euphotic zone of the central Arabian Sea and typically ca 0.8×10⁹ cells l^-1 were present. The majority of bacteria (80–95%) were small cocci that were larger (median diameter 0.40 μm) during the monsoon period than the inter-monsoon, when the cells had a diameter of 0.36 μm; there was no comparable change in cell dimensions of bacteria present as rods. Bacterial production was measured by the incorporation of ³H-thymidine and ³H-leucine. On both cruises, uptake rates were highest on the Omani shelf and decreased offshore. In the central Arabian Sea, thymidine incorporation rates were similar in the monsoon and inter-monsoon periods, but higher rates of leucine incorporation were measured during the monsoon period. Bacterial production was a relatively small proportion of phytoplankton production in both periods sampled; bacterial production was equivalent to between 10 and 30% of the daily primary production in the Arabian Sea.     

Seasonal variations in the distribution of Fe and Al in the surface waters of the Arabian Sea

Concentrations of dissolved Al and Fe in the surface mixed layer were measured during five cruises of the 1995 US JGOFS Arabian Sea Process Study, Concentrations of both Al and Fe were relatively uniform between January and April, the NE Monsoon and the Spring Intermonsoon period, ranging from 2 to 11 nM Al (mean 5.3 nM) and 0.5 to 2.4 nM Fe (mean 1.0 nM). In July/August, after the onset of the SW Monsoon, surface water Al and Fe concentrations increased significantly (Al range 4.5–20.1 nM; mean=10 nM, Fe range 0.57–2.4 nM; mean=1.3 nM), particularly in the NE part of the Arabian Sea, as the result of the input and partial dissolution of eolian dust. Using the enrichment of Al in the surface waters, we estimate this is the equivalent to the deposition of 2.2–7.4 g m⁻² dust, which is comparable to values previously estimated for this region. Approximately one month later (August/September), surface water concentrations of both Al and Fe were found to have decreased significantly (mean Al 7.4 nM, mean Fe 0.90 nM) particularly in the same NE region, as the result of export of particulate material from the euphotic zone. Fe supply to the surface waters is also affected by upwelling of sub-surface waters in the coastal region of the Arabian Sea during the SW Monsoon. Despite the proximity of high concentrations of Fe in the shallow sub-oxic layer, freshly upwelled water is not drawn from this layer and the NO₃/Fe ratio in the initially upwelled water is below the value at which Fe limitation is through to occur. Continued deposition of eolian Fe into the upwelled water as it advects offshore provides the Fe required to raise this ratio above the Fe limitation value.     

Diatom biomass and productivity in oceanic and plume-influenced waters of the western tropical Atlantic ocean

Whereas diatoms (class Bacillariophyceae) often dominate phytoplankton taxa in the Amazon estuary and shelf, their contribution to phytoplankton dynamics and impacts on regional biogeochemistry are poorly understood further offshore in the western tropical Atlantic Ocean (WTAO). Thus, relative contribution of diatoms to phytoplankton biomass and primary production rates and associated environmental conditions were quantified during three month-long cruises in January–February 2001, July–August 2001, and April–May 2003. The upper water column was sampled at 6 light depths (100%, 50%, 25%, 10%, 1% and 0.1% of surface irradiance) at 64 stations between 3° and 14°N latitude and 41° and 58°W longitude. Each station was categorized as ‘oceanic’ or ‘plumewater’, based on principal component analysis of eight physical, chemical and biological variables. All stations were within the North Brazil Current, and plumewater stations were characterized by shallower mixed layers with lower surface salinities and higher dissolved silicon (dSi) concentrations than oceanic stations. The major finding was a much greater role of diatoms in phytoplankton biomass and productivity at plumewater stations relative to oceanic stations. Mean depth-integrated bSi concentrations at the plumewater and oceanic stations were 14.2 and 3.7 mmol m⁻², respectively. Mean depth-integrated SiP rates at the plumewater and oceanic stations were 0.17 and 0.02 mmol m⁻² h⁻¹, respectively. Based on ratios of SiP and PP rates, and typical Si:C ratios, diatoms contributed on average 29% of primary productivity at plumewater stations and only 3% of primary productivity at oceanic stations. In contrast, phytoplankton biomass (as chlorophyll a concentrations) and primary production (PP) rates (as ¹⁴C uptake rates) integrated over the euphotic zone were not significantly different at plumewater and oceanic stations. Chlorophyll a concentrations ranged from 8.5 to 42.4 mg m⁻² and 4.0 to 38.0 mg m⁻² and PP rates ranged from 2.2 to 11.2 mmol m⁻² h⁻² and 1.8 to 10.8 mmol m⁻² h⁻² at plumewater and oceanic stations, respectively. A conservative estimate of annual integrated SiP in offshore waters of Amazon plume between April and August is 0.59 Tmol Si, based on mean SiP rates in plumewaters and satellite-derived estimates of the area of the Amazon plume. In conclusion, river plumewaters dramatically alter the silicon dynamics of the WTAO, forming extensive diatom-dominated phytoplankton blooms that may contribute significantly to the global Si budget as well as contributing to energy and matter flow off of the continental shelf.     

Variability of chlorophyll and primary production in the Eastern North Atlantic Subtropical Gyre: potential factors affecting phytoplankton activity

Size-fractionated chlorophyll-a and carbon incorporation rates were determined on a series of 13 cruises carried out from 1992 to 2001with the aim of investigating the patterns and causes of variability in phytoplankton chlorophyll and production in the Eastern North Atlantic Subtropical Gyral Province (NASE). Averaged (±SE) integrated chlorophyll-a concentration and primary production rate were 17±1 mg m⁻² and 253±22 mg C m⁻² d⁻¹. Small-sized cells (<2 μm) formed the bulk of phytoplankton biomass (71%) and accounted for 54% of total primary production. A clear latitudinal gradient in these variables was not detected. By contrast, large seasonal variability was detected in terms of primary production, although integrated phytoplankton biomass, as estimated from chlorophyll-a concentration, remained rather constant and did not display significant changes with time. Variability in primary production (PP) was related mainly to variability in surface temperature and surface chlorophyll-a concentration. The control exerted by surface temperature was related to nutrient availability. By contrary, euphotic-zone depth, depth of maximum concentration of chlorophyll-a and integrated chlorophyll-a did not contribute significantly to the high variability in primary production observed in this oligotrophic region.     

Seasonal dissolved inorganic carbon variations in the Greenland Sea and implications for atmospheric CO2 exchange

During the 1993–1995 period of minimal deep convection in the Greenland Sea, the dissolved inorganic carbon concentration within the surface waters varied dramatically on the seasonal time scale, with average summer and winter values of 2064 (±10) and 2150 (±5) μmol kg⁻¹, respectively, indicative of a vigorous annual carbon cycle. In contrast, there was very little interannual variability throughout these three years. While primary production largely depleted the surface nutrient supplies in spring and summer, generating a strong seasonal CO₂ drawdown, a combination of relatively shallow remineralization and mixed-layer deepening brought essentially all of the carbon consumed by photosynthesis back into contact with the atmosphere before winter. This re-release of the inorganic carbon that had been consumed by phytoplankton earlier in the year was more than sufficient to counteract the cooling-induced increase in the carbon carrying capacity of the water during fall and winter, reducing the potential for atmospheric carbon dioxide absorption by the Greenland Sea over the same period.     

Phytoplankton of the western Arctic in the spring and summer of 2002: Structure and seasonal changes

We analyzed the taxonomic structure and spatial variability of phytoplankton abundance and biomass in the Chukchi and Beaufort Seas during spring and summer seasons of the SBI program. Phytoplankton samples were collected during two surveys from May 10 to June 13 and from July 19 to August 21 of 2002. In May and June, ice cover exceeded 80% over most of the study area and there was no vertical stratification, indicating that the successional state of the phytoplankton corresponded to the end of the winter biological season. The phytoplankton abundance ranged from a few tens to a few thousands of cells per liter, while biomass varied from 0.1 to 3.0 mg C m⁻³. Small areas of high phytoplankton abundance (0.13–1.3×10⁶ cells L⁻¹) and biomass (22–536 mg C m⁻³), dominated by early spring diatoms Pauliella taeniata and Fragilariopsis oceanica in the surface waters, which indicated the beginning of the spring bloom, were observed only in the southeastern part of the Chukchi shelf and off Point Barrow. In July and August summer period, more than a half of the study area had <50% ice cover and the water column was stratified by temperature and salinity. Over the Chukchi shelf and continental slope of the Beaufort Sea, the phytoplankton abundance and biomass were an order of magnitude higher in July–August than in May–June. The taxonomic diversity of algae also increased due to the appearance of late-spring and summer diatoms, dinoflagellates, and coccolithophorids (Emiliania huxleyi). Interestingly, the seasonal differences between phytoplankton abundance and taxonomic composition in the spring and summer periods varied the least over the Chukchi Sea slope and in the deep-water area of the Arctic Ocean. High algae concentrations in summer were located in the lower layers of the euphotic zone, suggesting that the spring bloom on both the Chukchi shelf and in the western part of the Beaufort Sea occurred in late June/early July. In the spring and summer, the microalgal community was characterized by a high abundance of 4–10 μm flagellates, which exceeded the abundance of all other taxonomic groups. In both seasons studied, phytoplankton reached its maximum abundance within restricted areas in the southern part of the Chukchi Sea southwest of Point Hope, in the northern part of the Chukchi shelf between the 50- and 100-m isobaths, on the shelf northwest of Point Barrow, and over the continental slope in the Beaufort Sea. The pronounced spatial difference in the seasonal state was a characteristic feature of the phytoplankton community in the western Arctic.     

Seasonal variability of primary production in the Cap-Ferret Canyon area (Bay of Biscay) during the ECOFER cruises

Primary production measurements were carried out during the five ECOFER cruises, in order to estimate the autochtonous contribution to particulate fluxes collected at two mooring sites, in the Cap-Ferret Canyon, from June 1989 to August 1991. Primary production rates are reported in relation to levels of chlorophyll and nitrogenous nutrients in the euphotic zone. Except in early spring, the water column remains stratified until the beginning of autumn surface-water cooling. During the stratified period, maximal chlorophyll concentrations were recorded at the bottom of the photic zone, near the 1% light depth, close to the nitracline; concentrations in some profiles were greater than those measured in surface waters during the early spring bloom. From June to October, a mean daily production of 0.4 g C m⁻² may be used to estimate particulate fluxes, because the recorded variability was low. During April and May, mean daily production rates can be about 3–4 times higher. Extrapolation of our data gives an estimate of yearly production from 145 to 170 g C m⁻² yr⁻¹. The possibility of greater production rates, under stratified conditions, is discussed taking into account the occurrence of changes in the depth of the chlorophyll maximum during the day; such vertical movements could be caused by internal waves, particularly at the shelf break.     

Phytoplankton growth and microzooplankton grazing in the Sea of Okhotsk during late summer of 2006

The Sea of Okhotsk is one of the most productive marine basins in the world ocean and plays an important role in transport of organic carbon and iron to the western subarctic Pacific. We report the first measurements of phytoplankton growth and microzooplankton grazing rates in the Sea of Okhotsk, in late summer of 2006. The study area can be divided into two areas: nutrient-sufficient waters on the continental shelf along the east coast of Sakhalin Island and in the vicinity of Bussol Strait, and surface nutrient-depleted waters beyond the shelf break and in the vicinity of Sakhalin Bay. Phytoplankton growth rate in the studied area was strongly affected by nutrient availability, with high phytoplankton growth rate (0.55±0.14 d^?1) in the nutrient-replete region and severely depressed growth (0.03±0.05 d^?1) in the nutrient-depleted region. On the other hand, microzooplankton grazing rates in both the nutrient-replete and nutrient-depleted regions were approximately the same (0.26±0.20 d^?1 vs. 0.27±0.24 d^?1). Consequently, microzooplankton grazing consumed <50% of the phytoplankton growth in nutrient-rich waters but >3 times the phytoplankton growth in nutrient-depleted waters. Phytoplankton physiological condition as measured by the maximum photochemical quantum efficiency (F_v/F_m) of algal photosystem II (PS II) showed a general trend in agreement with the in situ growth rate of phytoplankton. In contrast to the phytoplankton community, picophytoplankton, especially the cyanobacteria Synechococcus, showed no nutrient effect on their growth, and the growth and mortality rates were well balanced, suggesting that they have a low nutrient requirement and their biomass was controlled principally by microzooplankton grazing.     

Latitudinal distribution of microbial plankton abundance,production, and respiration in the Equatorial Atlantic in autumn 2000

Phytoplankton and bacterial abundance, size-fractionated phytoplankton chlorophyll-a (Chl-a) and production together with bacterial production, microbial oxygen production and respiration rates were measured along a transect that crossed the Equatorial Atlantic Ocean (10°N–10°S) in September 2000, as part of the Atlantic Meridional Transect 11 (AMT 11) cruise. From 2°N to 5°S, the equatorial divergence resulted in a shallowing of the pycnocline and the presence of relatively high nitrate (>1 μM) concentrations in surface waters. In contrast, a typical tropical structure (TTS) was found near the ends of the transect. Photic zone integrated ¹⁴C primary production ranged from ∼200 mg C m⁻² d⁻¹ in the TTS region to ∼1300 mg C m⁻² d⁻¹ in the equatorial divergence area. In spite of the relatively high primary production rates measured in the equatorial upwelling region, only a moderate rise in phytoplankton biomass was observed as compared to nearby nutrient-depleted areas (22 vs. 18 mg Chl-a m⁻², respectively). Picophytoplankton were the main contributors (>60%) to both Chl-a biomass and primary production throughout the region. The equatorial upwelling did not alter the phytoplankton size structure typically found in the tropical open ocean, which suggests a strong top-down control of primary producers by zooplankton. However, the impact of nutrient supply on net microbial community metabolism, integrated over the euphotic layer, was evidenced by an average net microbial community production within the equatorial divergence (1130 mg C m⁻² d⁻¹) three-fold larger than net production measured in the TTS region (370 mg C m⁻² d⁻¹). The entire region under study showed net autotrophic community metabolism, since respiration accounted on average for 51% of gross primary production integrated over the euphotic layer.     

Fate of biogenic carbon in the upper 200 m of the central Greenland Sea

Sedimentation of particulate carbon from the upper 200–300 m in the central Greenland Sea from August 1993 to June 1995 was less than 2 g C m⁻² yr⁻¹. Daily rates of sedimentation of particulate organic carbon reached highest values of about 18 mg m⁻² d⁻¹ in fall 1994. For total particulate material, maximum rates of sedimentation of about 250 mg m⁻² d⁻¹ were recorded in spring and fall 1994. For chlorophyll equivalent, highest rates of sedimentation of about 140 μg m⁻² d⁻¹ were recorded in spring 1994. As reported in related investigations, the transient accumulation of DOC in surface waters during summer, as well as respiration and mortality of deep overwintering zooplankton stocks, appeared to dominate the fate of photosynthetically fixed organic carbon. The above processes may account for roughly 43 g C m⁻² in the upper 200 m of the central Greenland Sea. For comparison, the seasonal deficit in dissolved inorganic carbon was reported to be about 23 g C m⁻² in the upper 20 m of surface water, and estimates for new annual production were reported to be about 57 g C m⁻². In our investigation, the biological carbon pump was not unusually effective in transporting carbon out of the productive surface layer.     

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.     

Viral abundance and distribution in mesopelagic and bathypelagic waters of the Mediterranean Sea

Despite the fact that marine viruses have been increasingly investigated in the last decade, knowledge on virus abundance, biomass and distribution in mesopelagic and bathypelagic waters is limited. We report here the results of a large-spatial-scale study (covering more than 3000 km) on the virioplankton distribution in epi-, meso- and bathypelagic waters in 19 areas of the Mediterranean Sea, from the Alboran Sea and Western Mediterranean, to the Tyrrhenian Sea, Sicily Channel and Ionian Sea. Integrated viral abundance in epipelagic waters was significantly higher than in deep-sea waters (on average, 2.4 vs. 0.5×10¹² viruses m⁻³). However, abundance of viruses in the deep-Mediterranean waters was the highest reported so far for deep seas worldwide (7.0 and 3.1×10¹¹ viruses m⁻³ in mesopelagic and bathypelagic waters, respectively) and their biomass accounted for 13–18% of total prokaryotic C biomass. The significant relationship between viral abundance and prokaryotic abundance and production in deep waters suggests that also deep-sea viruses are closely dependent on the abundance and metabolism of their hosts. Moreover, virus to prokaryote (and nucleoid-containing cell (NuCC)) abundance ratio increased with increasing depths suggesting that deep waters may represent optimal environments for viral survival or proliferation. Overall, our results indicate that deep waters may represent a significant reservoir of viruses and open new perspectives for future investigations of viral impact on the functioning of meso-bathypelagic ecosystems.     

The influence of the south-west monsoon upon the nutrient biogeochemistry of the Arabian Sea

Variations in the nutrient concentrations were studied during two cruises to the Arabian Sea. The situation towards the end of the southwest monsoon season (September/October 1994) was compared with the inter-monsoonal season during November and December 1994. Underway surface transects showed the influence of an upwelling system during the first cruise with deep, colder, nutrient-rich water being advected into the surface mixed layer. During the southwesterly monsoon there was an area of coastal Ekman upwelling, bringing colder water (24.2°C) into the surface waters of the coastal margin. Further offshore at about 350 km there was an area of Ekman upwelling, as a result of wind-stress curl, north of the Findlater Jet axis; this area also had cooler surface water (24.6°C). Further offshore (>1000 km) the average surface temperatures increased to >27°C. These waters were oligotrophic with no evidence of the upwelling effects observed further inshore. In the upwelling regions nutrient concentrations in the close inshore coastal zone were elevated (NO₃=18 μmol l^-1, PO₄=1.48 μmol l^-1); higher concentrations also were measured at the region of offshore upwelling off the shelf, with a maximum nitrate concentration of 12.5 μmol l^-1 and a maximum phosphate concentration of 1.2 μmol l^-1. Nitrate and phosphate concentrations decreased with increasing distance offshore to the oligotrophic waters beyond 1400 km, where typical nitrate concentrations were 35.0 nmol l^-1 (0.035 μmol l^-1) in the surface mixed layer. A CTD section from the coastal shelf, to 1650 km offshore to the oligotrophic waters, clearly showed that during the monsoon season, upwelling is one of the major influences upon the nutrient concentrations in the surface waters of the Arabian Sea off the coast of Oman. Productivity of the water column was enhanced to a distance of over 800 km offshore. During the intermonsoon period a stable surface mixed layer was established, with a well-defined thermocline and nitracline. Surface temperature was between 26.8 and 27.4°C for the entire transect from the coast to 1650 km offshore. Nitrate concentrations were typically between 2.0 and 0.4 μmol l^-1 for the transect, to about 1200 km where the waters became oligotrophic, and nitrate concentrations were then typically 8–12 nmol l^-1. Ammonia concentrations for the oligotrophic waters were typically 130 nmol l^-1, and are reported for the first time in the Indian Ocean. The nitrogen/phosphorus (N/P) ratios suggest that phytoplankton production was potentially nitrogen-limited in all the surface waters of the Arabian Sea, with the greatest nitrogen limitation during the intermonsoon period.     

Vertical distribution of phytoplankton biomass,production and growth in the Atlantic subtropical gyres

Ninety-four stations were sampled in the Atlantic subtropical gyres during 10 cruises carried out between 1995 and 2001, mainly in boreal spring and autumn. Chlorophyll a (Chl-a) and primary production were measured during all cruises, and phytoplankton biomass was estimated in part of them. Picoplankton (<2 μm) represented >60% of total Chl-a concentration measured at the surface, and their contribution to this variable increased with depth. Phytoplankton carbon concentrations were higher in the upper metres of the water column, whereas Chl-a showed a deep maximum (DCM). At each station, the water column was divided into the upper mixed layer (ML) and the DCM layer (DCML). The boundary between the two layers was calculated as the depth where Chl-a concentration was 50% of the maximum Chl-a concentration. On average DCML extends from 67 to 126 m depth. Carbon to Chl-a (C:Chl-a) ratios were used to estimate phytoplankton carbon content from Chl-a in order to obtain a large phytoplankton carbon dataset. Total C:Chl-a ratios averaged (±s.e.) 103±7 (n=22) in the ML and 24±4 (n=12) in the DCML and were higher in larger cells than in picoplankton. Using these ratios and primary production measurements, we derived mean specific growth rates of 0.17±0.01 d⁻¹ (n=173) in the ML and 0.20±0.01 d⁻¹ (n=165) in the DCML although the differences were not significant (t-test, p>0.05). Our results suggest a moderate contribution of the DCML (43%) to both phytoplankton biomass and primary production in the Atlantic subtropical gyres.     

Seasonal dynamics of primary and new production in the northern South China Sea: The significance of river discharge and nutrient advection

Biochemical and productivity measurements and nutrient enrichment experiments were conducted on three cruises in summer and two cruises in winter on the shelf and the basin of the northern South China Sea (SCS) between 2001 and 2004. Phytoplankton production, in terms of depth-integrated new production (INP) or depth-integrated primary production (IPP), was higher in winter than in summer and on the shelf than in the basin. In winter, with deepening of the mixed layer, nitrate from the shallow nitracline that characterized the SCS waters was made available in the surface and supported the highest production of the year. Averaged INP measured in winter (0.25 g C m⁻² d⁻¹) was about twice the summer average (0.12 g C m⁻² d⁻¹) and was 0.19 g C m⁻² d⁻¹ on the shelf compared with 0.15 g C m⁻² d⁻¹ in the basin. In winter, average INP on the shelf was higher than the basin (0.34 versus 0.21 g C m⁻² d⁻¹); whereas in summer, averaged INP on the shelf (0.13 g C m⁻² d⁻¹) and the basin (0.11 g C m⁻² d⁻¹) were similar. While averaged IPP measured in the basin was higher in winter than in summer (0.53 versus 0.35 g C m⁻² d⁻¹), IPP on the shelf showed little temporal variation (0.82 in winter versus 0.84 g C m⁻² d⁻¹ in summer). Considerable spatial and inter-annual variation in production was measured in the shelf waters during summer, which could be linked to discharge volume and plume flow direction of the Zhujiang River. While the shelf waters in summer were mostly nitrogen starved or nitrogen and phosphorus co-limited, excessive river runoff may cause the nutritive state to shift to phosphorus deficiency. Waters with low surface salinities and high fluorescence from riverine mixing could be found extending from the Zhujiang mouth to as far as offshore southern Taiwan after a typhoon passed the northern SCS and brought heavy rainfall. Overall, both nutrient advection in winter and river discharge from the China coast in summer made new nitrogen available and shaped the dynamics of phytoplankton production in these oligotrophic waters.     

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.     

Top-down control of spring surface phytoplankton blooms by microzooplankton in the Central Yellow Sea,China

Phytoplankton growth and microzooplankton grazing were studied during the 2007 spring bloom in Central Yellow Sea. The surveyed stations were divided to pre-bloom phase (Chl a concentration less than 2 μg L⁻¹), and bloom phase (Chl a concentration greater than 2 μg L⁻¹). Shipboard dilution incubation experiments were carried out at 19 stations to determine the phytoplankton specific growth rates and the specific grazing rates of microzooplankton on phytoplankton. Diatoms dominated in the phytoplankton community in surface waters at most stations. For microzooplankton, Myrionecta rubra and tintinnids were dominant, and heterotrophic dinoflagellate was also important in the community. Phytoplankton-specific growth rates, with an average of 0.60±0.19 d⁻¹, were higher at pre-bloom stations (average 0.62±0.17 d⁻¹), and lower at the bloom stations (average 0.59±0.21 d⁻¹), but the difference of growth rates between bloom and pre-bloom stations was not statistically significant (t test, p=0.77). The phytoplankton mortality rate by microzooplankton grazing averaged 0.41±0.23 d⁻¹ at pre-bloom stations, and 0.58±0.31 d⁻¹ during the blooms. In contrast to the growth rates, the statistic difference of grazing rates between bloom and pre-bloom stations was significant (after removal of outliers, t test, p=0.04), indicating the importance of the top-down control in the phytoplankton bloom processes. Average potential grazing efficiency on primary productivity was 66% at pre-bloom stations and 98% at bloom stations, respectively. Based on our results, the biomass maximum phase (bloom phase) was not the maximum growth rate phase. Both phytoplankton specific growth rate and net growth rate were higher in the pre-bloom phase than during the bloom phase. Microzooplankton grazing mortality rate was positively correlated with phytoplankton growth rate during both phases, but growth and grazing were highly coupled during the booming phase. There was no correlation between phytoplankton growth rate and cell size during the blooms, but they were positive correlated during the pre-bloom phase. Our results indicate that microzooplankton grazing is an important process controlling the growth of phytoplankton in spring bloom period in the Central Yellow Sea, particularly in the “blooming” phase.