Plankton copper requirements and uptake in the subarctic Northeast Pacific Ocean

We undertook the first measurements of metabolic Cu requirements (net Cu:C assimilation ratios) and steady-state Cu uptake rates (ρCu_ss) of natural plankton assemblages in the northeast subarctic Pacific using the short-lived radioisotope ⁶⁷Cu. Size-fractionated net Cu:C assimilation ratios varied ～3 fold (1.35–4.21 μmol Cu mol C^?1) among the stations along Line P, from high Fe coastal waters to the Fe-limited open ocean. The variability in Cu:C was comparable to biogenic Fe:C ratios in this region. As previously observed for Fe uptake, the bacterial size class accounted for half of the total particulate ρCu_ss. Interestingly, carbon biomass-normalized rates of Fe uptake from the siderophore desferrioxamine B (DFB) (ρFe_DFB; a physiological proxy for Fe-limitation) by the >20 μm size class were positively correlated with the intracellular net Cu:C assimilation ratios in this size class, suggesting that intracellular Cu requirements for large phytoplankton respond to increased Fe-limitation. At Fe-limited Ocean Station Papa (OSP), we performed short-term Cu uptake (ρCu_L) assays to determine the relative bioavailability of Cu bound to natural and synthetic ligands. Like the volumetric ρCu_ss measured along Line P, the bacterial size class was responsible for at least 50% of the total ρCu_L. Uptake rates of Cu from the various organic complexes suggest that Cu uptake was controlled by the oxidation state of the metal and by the metal:ligand concentration ratio, rather than the concentration of inorganic species of Cu in solution. Collectively, these data suggest that Cu likely plays an important role in the physiology of natural plankton communities beyond the toxicological effects studied previously.     

Early spring bloom phytoplankton-nutrient dynamics at the Celtic Sea Shelf Edge

Phytoplankton production was measured at the shelf edge region of the Celtic Sea in April/May 1994 at the beginning of the spring bloom. Size fractionated ¹⁴C uptake experiments showed that phytoplankton >2 μm dominated the bloom although, in the period immediately before the increase in phytoplankton biomass, picophytoplankton (<2 μm) was responsible for up to 42% of the production; in these late winter conditions, chlorophyll concentrations were generally <0.7 μg l^-1 and primary production was ca. 70 mmol C m^-2 d^-1. As the spring bloom developed, phytoplankton production rates of 120 mmol C m^-2 d^-1 were measured. Chlorophyll concentration increased to >2 μg l^-1 as a result of growth of larger phytoplankton, including diatoms, with large numbers of Nitzschia, Thalassionema and Chaetoceros dominating the assemblage. Picophytoplankton production declined as the spring bloom progressed. Nutrient concentrations were not depleted during the sampling period, and NO^-₃ concentrations were >6 μmol l^-1. Nutrient assimilation rates were measured at the same time as primary production was estimated. Before the development of any substantial phytoplankton biomass, the uptake rates for ammonium and nitrate were very similar, with f-ratios ranging from 0.5 to 0.6. Assimilation of ammonium remained relatively constant after the onset of stratification and bloom development, but nitrate uptake increased by a factor of 2 or more, resulting in f-ratios >0.8. There was significant phosphate uptake in the dark, which was generally ca. 50% of the rate in the light. The C : N : P assimilation ratios changed as the bloom developed; in the pre-bloom situation, when small phytoplankton cells dominated the assemblage, the C : N assimilation ratio was variable, with some stations having ratios less than (ca 2.5), and some higher than (ca. 9), the Redfield ratio. The most actively growing assemblages had N : P ratios close to the Redfield ratio, but the C : N ratios were consistently lower. New production was found to be closely correlated with the size of the species making up the phytoplankton assemblage, and high f ratios were measured when larger phytoplankton dominated the assemblage.     

Microzooplankton grazing and selective feeding during bloom periods in the Tolo Harbour area as revealed by HPLC pigment analysis

Dilution experiments were conducted to investigate microzooplankton grazing impact on phytoplankton of different taxonomic groups and size fractions (< 5, 5–20, 20–200 μm) during spring and summer bloom periods at two different sites (inner Tolo Harbour and Tolo Channel) in the Tolo Harbour area, the northeastern coastal area of Hong Kong. Experiments combined with HPLC pigment analysis in three phytoplankton size fractions measured pigment and size specific phytoplankton growth rates and microzooplankton grazing rates. Pigment-specific phytoplankton growth rates ranged between 0.08 and 3.53 d^− 1, while specific grazing rates of microzooplankton ranged between 0.07 and 2.82 d^− 1. Highest specific rates of phytoplankton growth and microzooplankton grazing were both measured in fucoxanthin in 5–20 μm size fraction in inner Tolo Harbour in summer, which coincided with the occurrence of diatom bloom. Results showed significant correlations between phytoplankton growth and microzooplankton grazing rates. Microzooplankton placed high grazing pressure on phytoplankton community. High microzooplankton grazing impact on alloxanthin (2.63–5.13) suggested strong selection toward cryptophytes. Our results provided no evidence for size selective grazing on phytoplankton by microzooplankton.     

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.     

Bacterivory in the northwestern Indian Ocean during the intermonsoon – northeast monsoon period

Bacterial grazing loss rates were studied by radioactive labeling of natural bacteria with L-(4,5-³H) leucine and from the rate of disappearance of bacterial cells in the northwestern Indian Ocean. Bacterivory was measured in a mixed sample that had been combined from various depths across the euphotic zone. Experiments were performed on 26 occasions at 19 stations in the Gulf of Oman and the Arabian Sea during the intermonsoon–northeast monsoon period (November–December 1994). Combined uptake of radiolabeled bacteria (ULB) in 1–8 and 8–100 μm size fractions was somewhat lower than loss of label (LBL) measured in the bacterial fraction (0.2–1.0 μm), suggesting loss of radioactivity from the grazers due to metabolism. The less sensitive rate of disappearance of bacterial cells (LBC) was on average 51% higher than LBL estimates. Results from ULB and LBL measurements revealed that bacterivory was higher in the Gulf of Oman (average loss rate 4.1% h^-1) than in the Arabian Sea where rates were slightly higher inshore (1.7% h^-1) than in the central gyre. Heterotrophic nanoflagellates in the 1–8 μm size fractions were identified as the primary bacterivores. Microzooplankton (8–100 μm) accounted for 33% of total bacterivory in the Gulf of Oman but only 16% in the central Arabian Sea. Time-course experiments conducted at two stations indicated that diel changes in bacterivory may be substantial in the northwestern Indian Ocean.     

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.     

Picophytoplankton dynamics and production in the Arabian Sea during the 1995 Southwest Monsoon

Phytoplankton community structure is expected to shift to larger cells (e.g., diatoms) with monsoonal forcing in the Arabian Sea, but recent studies suggest that small primary producers remain active and important, even in areas strongly influenced by coastal upwelling. To better understand the role of smaller phytoplankton in such systems, we investigated growth and grazing rates of picophytoplankton populations and their contributions to phytoplankton community biomass and primary productivity during the 1995 Southwest Monsoon (August–September). Environmental conditions at six study stations varied broadly from open-ocean oligotrophic to coastal eutrophic, with mixed-layer nitrate and chlorophyll concentrations ranging from 0.01 to 11.5 μM NO₃ and 0.16 to 1.5 μg Chl a. Picophytoplankton comprised up to 92% of phytoplankton carbon at the oceanic stations, 35% in the diatom-dominated coastal zone, and 26% in a declining Phaeocystis bloom. Concurrent in situ dilution and ¹⁴C-uptake experiments gave comparable ranges of community growth rates (0.53–1.05 d⁻¹ and 0.44–1.17 d⁻¹, to the 1% light level), but uncertainties in C:Chl a confounded agreement at individual stations. Microzooplankton grazing utilized 81% of community phytoplankton growth at the oligotrophic stations and 54% at high-nutrient coastal stations. Prochlorococcus (PRO) was present at two oligotrophic stations, where its maximum growth approached 1.4 d⁻¹ (two doublings per day) and depth-integrated growth varied from 0.2 to 0.8 d⁻¹. Synechococcus (SYN) growth ranged from 0.5 to 1.1 d⁻¹ at offshore stations and 0.6 to 0.7 d⁻¹ at coastal sites. Except for the most oligotrophic stations, growth rates of picoeukaryotic algae (PEUK) exceeded PRO and SYN, reaching 1.3 d⁻¹ offshore and decreasing to 0.8 d⁻¹ at the most coastal station. Microzooplankton grazing impact averaged 90, 70, and 86% of growth for PRO, SYN, and PEUK, respectively. Picoplankton as a group accounted for 64% of estimated gross carbon production for all stations, and 50% at high-nutrient, upwelling stations. Prokaryotes (PRO and SYN) contributed disproportionately to production relative to biomass at the most oligotrophic station, while PEUK were more important at the coastal stations. Even during intense monsoonal forcing in the Arabian Sea, picoeukaryotic algae appear to account for a large portion of primary production in the coastal upwelling regions, supporting an active community of protistan grazers and a high rate of carbon cycling in these areas.     

Factors controlling silicon and nitrogen biogeochemical cycles in high nutrient,low chlorophyll systems (the Southern Ocean and the North Pacific): Comparison with a mesotrophic system (the North Atlantic)

A 1-D coupled physical-biogeochemical model is used to study the seasonal cycles of silicon and nitrogen in two High Nutrient Low Chlorophyll (HNLC) systems, the Antarctic Circumpolar Current (ACC) and the North Pacific Ocean, and a mesotrophic system, the North Atlantic Ocean. The biological model consists of nine compartments (diatoms, nano-flagellates, microzooplankton, mesozooplankton, two types of detritus, nitrate, ammonium and silicic acid) forced by irradiance, temperature, mixing and deep nitrate and silicic acid concentrations. At all sites, nanophytoplankton standing crop variations are low, in spite of variations in primary production, because of a “top–down” control by microzooplankton. Although nanophytoplankton sustain more than 60% of the annual primary production in these areas, their contribution to the export production does not exceed 1% of the total. The differences in the seasonal plankton cycle among these regions come mainly from differences in the dynamics of large phytoplankton (here diatoms). In the ACC, the chlorophyll maximum remains <1.5 mg m⁻³, as an unfavourable light/mixing regime and a likely trace-metal limitation keep diatoms from blooming. In the northeast Pacific, trace-metal limitation seems to keep diatoms from blooming throughout the year. In both these systems, light or iron limitations induce high Si/N uptake ratios. Incidentally these high Si/N uptake ratios lead to a net excess of silicic acid utilization over nitrate, and to a subsequent silicic acid limitation during the summertime. In the North Atlantic, under favourable light/mixing regime and nutrient-replete conditions at the onset of the growing period, diatoms outburst and sustain a bloom >3.5 mg Chl-a m⁻³. Thereafter, mesozooplankton grazing pressure and silicic acid limitation induce the collapse of the chlorophyll maximum and the persistence of lower chlorophyll concentrations in summer. Although the ACC and the North Pacific show HNLC features, they support a high biogenic silica production (1.9 and 1.07 mol Si m⁻² yr⁻¹) and export flux (0.79 and 0.61 mol Si m⁻² yr⁻¹), compared to the North Atlantic (production: 0.23 mol Si m⁻² yr⁻¹, export: 0.12 mol Si m⁻² yr⁻¹). The differences in Si production and export between the HNLC systems and the mesotrophic North Atlantic come from both higher Si concentrations and Si/N uptake ratios in the HNLC areas compared to the North Atlantic. Also, the low dissolution rate of biogenic silica compared to nitrogen degradation rate, and the inhibition of nitrate uptake by ammonium, reinforce the net excess of silicic acid utilization over nitrate. As a result, the model also illustrates the efficiency of the silica pump for the three sites: about 50% of the biogenic silica synthesized in the euphotic layer is exported out of the first 100 m, while only 4–11% of the particulate organic nitrogen escapes recycling in the surface layer.     

A phytoplankton class-specific primary production model applied to the Kerguelen Islands region (Southern Ocean)

As part of the KErguelen: compared study of the Ocean and the Plateau in Surface water (KEOPS) project in late summer 2005, we examine the phytoplankton community composition and associated primary production in the waters surrounding the Kerguelen Archipelago, with the emphasis on two contrasted environments: (i) the Kerguelen Plateau, where a large bloom occurs annually, and (ii) the high-nutrient low-chlorophyll (HNLC) offshore waters. A biomarker pigment approach was used to assess the community composition in terms of chlorophyll biomass of three phytoplankton size classes, namely micro-, nano-, and picophytoplankton. The second objective was to evaluate a global class-specific approach for estimating the contribution of the three pigment-based size classes to the primary production in the study area. To do so, primary production rates associated with each phytoplankton class were computed from the class-specific chlorophyll biomass coupled to a class-specific primary production model, and compared with in situ measurements of size-fractionated ¹³C-based primary production. The iron-enriched bloom region was dominated by microphytoplankton (diatoms), which contributed 80–90% to the total primary production (of ≈1 g C m^?2 d^?1). In the HNLC area, the primary production was about 0.30 g C m^?2 d^?1, mainly (65%) achieved by small diatoms and nanoflagellates. The model results show a good overall agreement between predicted and measured total primary production rates. In terms of size classes, agreements were higher for the bloom region than for the HNLC waters. Discrepancies in this complex iron-limited area may be explained essentially by the smaller size of diatoms, or a different set of photophysiological properties.     

Phytoplankton distributions in the Shackleton Fracture Zone/Elephant Island region of the Drake Passage in February–March 2004

The Drake Passage region near Elephant Island in the Southern Ocean displays patchy phytoplankton blooms. To test the hypothesis that natural Fe addition from localized sources promoted phytoplankton growth here, a grid of stations (59°S to 62°S, 59°W to 53°W, as well as four stations in the eastern Bransfield Strait) were occupied from 12 February–24 March 2004. Phytoplankton abundance was measured using shipboard flow cytometry (70 stations), with abundances conservatively converted to biomass, and compared with measurements of dissolved iron (dFe) at a subset of stations (30 stations). Based on T–S property plots, stations were divided into Antarctic Circumpolar Current (ACC), Water On Shelf (WOS), Bransfield Strait (BS), and Mixed water stations, the latter representing locations with T–S properties intermediate between ACC and WOS stations. The highest integrated phytoplankton biomass was found at Mixed water stations, however, the highest integrated abundance was found at WOS stations, demonstrating that abundance and biomass do not necessarily show the same patterns. The distributions of nano- and micro-phytoplankton (<20 and >20 μm diameter cells, respectively) were also examined, with nano- and micro-plankton contributing equally to the total biomass at WOS and BS stations, but micro-plankton representing ∼2/3 of the biomass at Mixed and ACC stations. Increased inventories of dFe did not always correspond to increases in phytoplankton biomass – rather stations with lower mean light levels in the mixed layer (<110 μEinsteins m⁻² s⁻¹) had lower biomass despite higher ambient dFe concentrations. However, where the mean light levels in the mixed layer were >110 μEinsteins m⁻² s⁻¹, total biomass shows a positive trend with dFe, as does micro-phytoplankton biomass, but neither regression is significant at the 95% level. In contrast, if just nano-phytoplankton biomass is considered as a function of dFe, there is a significant correlation (r²=0.62). These data suggest a dual mechanism for the patterns observed in biomass: an increasing reservoir of dFe allows increased phytoplankton biomass, but biomass can only accumulate where the light levels are relatively high, such that light is not limiting to growth.     

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.     

Effects of mesoscale phytoplankton variability on the copepods Neocalanus flemingeri and N. plumchrus in the coastal Gulf of Alaska

The copepods Neocalanus flemingeri and N. plumchrus are major components of the mesozooplankton on the shelf of the Gulf of Alaska, where they feed, grow and develop during April–June, the period encompassing the spring phytoplankton bloom. Satellite imagery indicates high mesoscale variability in phytoplankton concentration during this time. Because copepod ingestion is related to food concentration, we hypothesized that phytoplankton ingestion by N. flemingeri and N. plumchrus would vary in response to mesoscale variability of phytoplankton. We proposed that copepods on the inner shelf, where the phytoplankton bloom is most pronounced, would be larger and have more lipid stores than animals collected from the outer shelf, where phytoplankton concentrations are typically low. Shipboard feeding experiments with both copepods were done in spring of 2001 and 2003 using natural water as food medium. Chlorophyll concentration ranged widely, between 0.32 and 11.44 μg l⁻¹ and ingestion rates varied accordingly, between 6.0 and 627.0 ng chl cop⁻¹ d⁻¹. At chlorophyll concentrations<0.50 μg l⁻¹, ingestion is always low, <40 ng cop⁻¹ d⁻¹. Intermediate ingestion rates were observed at chlorophyll concentrations between 0.5 and 1.5 μg l⁻¹, and maximum rates at chlorophyll concentrations>1.5 μg l⁻¹. Application of these feeding rates to the phytoplankton distribution on the shelf allowed locations and time periods of low, intermediate and high daily feeding to be calculated for 2001 and 2003. A detailed cross-shelf survey of body size and lipid store in these copepods, however, indicated they were indistinguishable regardless of collection site. Although the daily ingestion of phytoplankton by N. flemingeri and N. plumchrus varied widely because of mesoscale variability in phytoplankton, these daily differences did not result in differences in final body size or lipid storage of these copepods. These copepods efficiently dealt with small and mesoscale variations in their food environment such that mesoscale structure in phytoplankton did not affect their final body size.     

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.     

Microzooplankton composition,biomass and grazing rates along the WOCE SR3 line between Tasmania and Antarctica

Microzooplankton species composition and grazing rates on phytoplankton were investigated along a transect between ∼46 and 67°S, and between 140 and 145°E. Experiments were conducted in summer between November 2nd and December 14th in 2001. The structure of the microbial food web changed considerably along the transect and was associated with marked differences in the physical and chemical environment encountered in the different water masses and frontal regions. On average microzooplankton grazing experiments indicated that 91%, 102%, and 157%, (see results) of the phytoplankton production would be grazed in the <200, <20 and <2 μm size fractions, respectively, indicating microzooplankton grazing was potentially constraining phytoplankton populations (<200 μm) along most of the transect. Small ciliates in general and especially oligotrich species declined in importance from the relatively warm, Southern Subtropical Front waters (6.8 μg C/L) to the colder waters of the southern branch of the Polar Front (S-PF), (∼0.5 μg C/L) before increasing again near the Antarctic landmass. Large changes in microzooplankton dominance were observed, with heterotrophic nanoflagellates (HNF), ciliates and larger dinoflagellates having significant biomass in different water masses. HNF were the dominant grazers when chlorophyll a was low in areas such as the Inter-Polar Frontal Zone (IPFZ), while in areas of elevated biomass such as the S-PF and Southern Antarctic Circumpolar Current (SACC), a mix of copepod nauplii and large heterotrophic and mixotrophic dinoflagellates tended to dominate the grazing community. In the S-PF and SACC water masses the tight coupling observed between the microzooplankton grazers and phytoplankton populations over most of the rest of the transect was relaxed. In these regions grazing was low on the >20 μm size fraction of chlorophyll a, which dominated the biomass, while smaller diatoms and nanoplankton in the <20 μm size fraction were still heavily grazed. The lack of grazing pressure on large phytoplankton contributes to this region's potential to export carbon with larger cells known to have higher sinking rates.     

Atmospheric iron deposition and sea-surface dissolved iron concentrations in the eastern Atlantic Ocean

Atmospheric iron and underway sea-surface dissolved (<0.2 μm) iron (DFe) concentrations were investigated along a north–south transect in the eastern Atlantic Ocean (27°N/16°W–19°S/5°E). Fe concentrations in aerosols and dry deposition fluxes of soluble Fe were at least two orders of magnitude higher in the Saharan dust plume than at the equator or at the extreme south of the transect. A weaker source of atmospheric Fe was also observed in the South Atlantic, possibly originating in southern Africa via the north-easterly outflow of the Angolan plume. Estimations of total atmospheric deposition fluxes (dry plus wet) of soluble Fe suggested that wet deposition dominated in the intertropical convergence zone, due to the very high amount of precipitation and to the fact that a substantial part of Fe was delivered in dissolved form. On the other hand, dry deposition dominated in the other regions of the transect (73–97%), where rainfall rates were much lower. Underway sea-surface DFe concentrations ranged 0.02–1.1 nM. Such low values (0.02 nM) are reported for the first time in the Atlantic Ocean and may be (co)-limiting for primary production. A significant correlation (Spearman's rho=0.862, p<0.01) was observed between mean DFe concentrations and total atmospheric deposition fluxes, confirming the importance of atmospheric deposition on the iron cycle in the Atlantic. Residence time of DFe in the surface waters relative to atmospheric inputs were estimated in the northern part of our study area (17±8 to 28±16 d). These values confirmed the rapid removal of Fe from the surface waters, possibly by colloidal aggregation.     

Biogeochemical cycling of N and Si during the mesoscale iron-enrichment experiment in the western subarctic Pacific (SEEDS-II)

Biogeochemical cycles of N and Si were examined in the surface mixed layer during the mesoscale iron-enrichment (IE) experiment in the high-nutrient low-chlorophyll (HNLC) western subarctic Pacific (SEEDS-II). Although the IEs increased nitrate uptake, silicic acid utilization was not stimulated. The nitrate drawdown in the iron-patch (IN-patch, 140.3 mmol m⁻² in the surface mixed layer, 0–30 m) was only 25% of the initial inventory, which was 1/3–2/5 of the previous IE experiments in the subarctic Pacific. This relatively weak response of nutrient drawdown to IEs was due to the high biomass of mesozooplankton (MZ) dominated by copepod Neocalanus plumchrus. Feeding of MZ (247.2 mmol m⁻² during Day 0–21 from the first IE) in the IN-patch was higher than the nitrate drawdown and prevented further development of the phytoplankton bloom. In the later period of the experiment (Day 14–21), the increase in the feeding activity and resultant decrease in phytoplankton biomass induced the accumulation of dissolved organic nitrogen (DON) and ammonium. Among total growth of MZ (81.6 mmol N m⁻²), 89% (72.8 mmol N m⁻²) was transported to the depth by the ontogenetic downward migration of N. plumchrus. Although silicic acid drawdown was not increased by the IEs, Si export flux increased by 2.7 times. The increase in Si export was also due to the increase in MZ, which egested faecal pellets with higher Si:N ratio and faster sinking speed than diatoms. The export efficiency (78% of new production) and total amount of export flux (143.8 mmol N m⁻², 1392 mmol C m⁻²) were highest records within the IE experiments despite weak responses of nutrient drawdown to the IE. During SEEDS-II, the high biomass of MZ reduced the phytoplankton response and nutrient drawdown to the IEs but via grazing and ontogenetic vertical migration accelerated the export flux as well as accumulations of dissolved forms of N. Results of the present and previous IE experiments indicate that the ecosystem and biogeochemical responses to IEs in the HNLC region are quite sensitive to the ecosystem components, especially for grazers of diatoms such as copepods and heterotrophic dinoflagellates. More attention needs to be paid to the ecosystem components and their biogeochemical functions as well as physical and chemical properties of the ecosystems in order to hindcast or forecast the impacts of changes in atmospheric iron deposition.     

Phytoplankton growth and mortality during the 1995 Northeast Monsoon and Spring Intermonsoon in the Arabian Sea

Phytoplankton growth rates and mortality rates were experimentally examined at eight stations in the Arabian Sea along the U.S. JGOFS cruise track during the 1995 Northeast Monsoon (January) and Spring Intermonsoon (March–April). Instantaneous growth rates averaged over an entire cruise were approximately twice as high during the NE Monsoon than during the Spring Intermonsoon period (overall averages of 0.84±0.29 (s.d.) versus 0.44±0.19 d⁻¹). Average herbivore grazing (mortality) rates, however, were quite similar for the two seasons (overall averages of 0.35±0.18 and 0.30±0.17 d⁻¹ for the NE Monsoon and Spring Intermonsoon, respectively). The absolute amounts of phytoplankton biomass consumed during each season also were similar (29 and 25% of standing stock consumed d⁻¹ for the January and March–April cruises, respectively), as were the geographical trends of this removal. These seasonal trends in growth and removal rates resulted in net phytoplankton growth rates that were considerably higher during the January cruise (0.48 d⁻¹) than during the March–April cruise (0.14 d⁻¹). That is, phytoplankton production was more closely balanced during the Spring Intermonsoon season (87% of daily primary production consumed) relative to the NE Monsoon season (49% of daily primary production consumed). Station-to-station variability was high for rate measurements during either cruise. Nevertheless, there was a clear onshore–offshore trend in the absolute rate of removal of phytoplankton biomass (μg chlorophyll consumed l⁻¹ d⁻¹) during both cruises. Coastal stations had removal rates that were typically 2–4 times higher than removal rates at oceanic stations.     

Environmental controls on phytoplankton community composition in the Thames plume,U.K.

The aim of this study was to investigate controls on the phytoplankton community composition and biogeochemistry of the estuarine plume zone of the River Thames, U.K. using an instrumented moored buoy for in situ measurements and preserved sample collection, and laboratory-based measurements from samples collected at the same site. Instrumentation on the moored buoy enabled high frequency measurements of a suite of environmental variables including in situ chlorophyll, water-column integrated irradiance, macronutrients throughout an annual cycle for 2001 e.g. nitrate and silicate, and phytoplankton biomass and species composition. The Thames plume region acts as a conduit for fluvial nutrients into the wider southern North Sea with typical winter concentrations of 45 μM nitrate, 17 μM silicate and 2 μM phosphate measured. The spring bloom resulted from water-column integrated irradiance increasing above 60 W h m^− 2 d^− 1 and was initially dominated by a diatom bloom mainly composed of Nitzschia sp. and Odontella sinesis. The spring bloom then switched after ∼ 30 days to become dominated by the flagellate Phaeocystis reaching a maximum chlorophyll concentration of 37.8 μg L^− 1. During the spring bloom there were high numbers of the heterotrophic dinoflagellates Gyrodinium spirale and Katodinium glaucum that potentially grazed the phytoplankton bloom. This diatom–flagellate switch was predicted to be due to a combination of further increasing water-column integrated irradiance > 100 W h m^− 2 d^− 1 and/or silicate reaching potentially limiting concentrations (< 1 μM). Post spring bloom, diatom dominance of the lower continuous summer phytoplankton biomass occurred despite the low silicate concentrations (Av. 0.7 μM from June–August). Summer diatom dominance, generally due to Guinardia delicatula, was expected to be as a result of microzooplankton grazing, dominated by the heterotrophic dinoflagellate Noctiluca scintillans, controlling 0.7–5.0 μm ‘flagellate’ fraction of the phytoplankton community with grazing rates up to 178% of ‘flagellate’ growth rate. The Thames plume region was therefore shown to be an active region of nutrient and phytoplankton processing and transport to the southern North Sea. The use of a combination of moorings and ship-based sampling was essential in understanding the factors influencing nutrient transport, phytoplankton biomass and species composition in this shelf sea plume region.     

Effects of solar radiation on dimethylsulfide cycling in the western Atlantic Ocean

The influence of solar radiation on springtime rates of photochemical and biological consumption of dimethylsulfide (DMS) in surface waters from the western Atlantic Ocean was examined by exposing 0.2 μm filtered and unfiltered surface seawater to natural sunlight at five depths in the upper 30 m. Parallel deck incubations of 0.2 μm filtered seawater under various long-pass optical filters were also carried out to aid in assessing the wavelength dependence of DMS photolysis. DMS photolysis rate constants for mid-day exposure (∼10:30–17:30 local time) to surface irradiance ranged from 0.026 to 0.086 h⁻¹ and were highest in coastal and shelf waters. Photolysis rate constants decreased with increasing irradiation depth, in accordance with the attenuation of ultraviolet radiation (UVR, 280–400 nm). Total DMS consumption rates (photochemical+biological) in unfiltered surface samples also decreased with increasing incubation depth and were larger than photolysis rates at nearly all depths and all stations. The decrease in photolysis rate constants with exposure depth was mirrored by biological DMS consumption rate constants that were severely inhibited at surface irradiances, and approached or exceeded dark rate constants at deeper exposure depths. Photolysis rates were 2–19 times greater than estimated biological consumption rates in the surface light exposed samples, while biological consumption rates were significantly larger than photolysis rates at incubation depths below the 1% light level for UV–B radiation (280–320 nm). Total DMS loss rates increased up to nine-fold with UVR exposure, but changes in DMS concentrations were not strongly correlated to light dose, presumably due to parallel, light-mediated DMS production processes. The primary loss process for DMS depended mainly on the depth interval considered and the attenuation of UVR; in general, photochemical removal dominated shallow layers characterized by high UV–B intensities, whereas biological removal dominated in deeper layers where UV–B was absent, but UV–A (320–400 nm) and visible (400–700 nm) light fluxes were still relatively high. These results demonstrate that UVR exposure significantly influences the spatial and temporal pattern of DMS production and loss processes, and ultimately the DMS flux to the atmosphere.     

Abundance and biomass of nano- and microplankton during the 1995 Northeast Monsoon and Spring Intermonsoon in the Arabian Sea

Phototrophic and heterotrophic nanoplankton (PNAN, HNAN; 2–20 μm protists) and microplankton (PMIC, HMIC; 20–200 μm protists and micrometazoa) are major components of the producer and consumer assemblages in oceanic plankton communities. Abundances and biomasses of these microorganisms were determined from samples collected along two transects during the Northeast Monsoon and Spring Intermonsoon process cruises of the US JGOFS Arabian Sea Program in 1995. Vertical profiles of these assemblages were strongly affected by the presence of a subsurface oxygen minimum layer. Abundances of all four assemblages decreased dramatically below the top of this layer. Depth-integrated (0–160 m) abundances and biomasses of nanoplankton and microplankton were of similar magnitude for most samples. Exceptions to this rule were primarily due to PMIC (mostly diatom) species which dominated phytoplankton assemblages at a few stations during each season. Depth-integrated biomasses for the combined nano- and microplankton averaged over all stations for each cruise were surprisingly similar for the Northeast Monsoon and Spring Intermonsoon seasons in this ecosystem (2.0 and 1.8 g C m⁻² [170 and 150 m moles C m⁻²] for the two seasons, respectively). Nano- and microplankton biomass for these two time periods constituted a signficant portion of the total amount of the particulate organic carbon (POC) in the water column. Summed over all stations, these assemblages constituted approximately 25–35% of the POC in the top 160 m of the northern Arabian Sea.