Phytoplankton community structure in the Arabian Sea during and after the SW monsoon, 1994

Diatoms, dinoflagellates, coccolithophores, nanoflagellates, picophytoplankton and procaryote algae (Synechococcus spp. and prochlorophytes) were quantified by microscopy and flow cytometry, and their biomass determined, at 12 stations along a 1600 km transect across the Arabian Sea at the end of the SW monsoon in September, and during the inter-monsoon period of November/December 1994. The transect spanned contrasting oceanic conditions that varied from seasonally eutrophic, upwelling waters through mesotrophic, downwelling waters to permanently oligotrophic, stratified waters. The overall diversity of diatoms, dinoflagellates and coccolithophores along the transect was not significantly different between the SW monsoon and inter-monsoon. However, diatoms showed greatest diversity during the SW monsoon and coccolithophores were most diverse during the inter-monsoon. Integrated phytoplankton standing stocks during the SW monsoon ranged from 3 to 9 g C m^-2 in the upwelling eutrophic waters, from 3 to 5 g C m^-2 in downwelling waters, and from 1 to 2 g C m^-2 in oligotrophic waters. Similar phytoplankton standing stocks were found in oligotrophic waters during the inter-monsoon, but were ca. 40% lower compared to the SW monsoon in the more physically dynamic waters. Phytoplankton abundance and biomass was dominated by procaryote taxa. Synechococcus spp. were abundant (often >10⁸ cells l^-1) during both the SW monsoon and inter-monsoon, where the nitrate concentration was ⩾0.1 μ mol l^-1, and often dominated the phytoplankton standing stocks. Prochlorophytes were restricted to oligotrophic stratified waters during the SW monsoon period but were found at all stations along the transect during the inter-monsoon, dominating the phytoplankton standing stocks (>40%) in the oligotrophic region during this period. Of the nano- and micro-phytoplankton, only diatoms contributed significantly to phytoplankton standing stocks, and then only in near-shore upwelling waters during the SW monsoon. There were significant changes in the temporal composition of the phytoplankton community. In nearshore waters a mixed community of diatoms and Synechococcus spp. dominated during the SW monsoon. This gave way to a community dominated by Synechococcus spp. in the intermonsoon. In the downwelling zone, a Synechococcus spp. dominated community was replaced by a mixed procaryote community of Synechococcus spp. and prochlorophytes. In the oligotrophic stratified waters, the mix of procaryote algae was replaced by one dominated by prochlorophytes alone.     

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.     

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.     

Monsoonal influence on the distribution of phytoplankton pigments in the Arabian Sea

Variations in the distribution of chemotaxonomic pigments were monitored in the Arabian Sea and the Gulf of Oman at the end of the SW monsoon in September 1994 and during the inter-monsoon period in November/December 1994 to determine the seasonal changes in phytoplankton composition. The Gulf of Oman was characterized by sub-surface chlorophyll maxima at 20-40 m during both seasons, and low levels of divinyl chlorophyll a indicated that prochlorophytes did not contribute significantly to the total chlorophyll a. Prymnesiophytes (19′-hexanoyloxyfucoxanthin), diatoms (fucoxanthin) and chlorophyll b containing organisms accounted for most of the phytoplankton biomass in September, while prymnesiophytes dominated in November/December. In the Arabian Sea in September, high total chlorophyll a concentrations up to 1742 ng l^-1 were measured in the coastal upwelling region and a progressive decline was monitored along the 1670 km offshore transect to oligotrophic waters at 8°N. Divinyl chlorophyll a was not detected along this transect except at the two most southerly stations where prochlorophytes were estimated to contribute 25–30% to the total chlorophyll a. Inshore, the dominance of fucoxanthin and/or hexanoyloxyfucoxanthin indicated that diatoms and prymnesiophytes generally dominated the patchy phytoplankton community, with zeaxanthin-containing Synechococcus also being important, especially in surface waters. At the southern oligotrophic localities, Synechococcus and prochlorophytes dominated the upper 40 m and prymnesiophytes were the most prominent at the deep chlorophyll maximum. During the inter-monsoon season, total chlorophyll a concentrations were generally half those measured in September and highest levels were found on the shelf (1170 ng l^-1). Divinyl chlorophyll a was detected at all stations along the Arabian Sea transect, and we estimated that prochlorophytes contributed between 3 and 28% to the total chlorophyll a, while at the two oligotrophic stations this proportion increased to 51–52%. While procaryotes were more important in November/December than September, eucaryotes still accounted for >50% of the total chlorophyll a. Pigment/total chlorophyll a ratios indicated that 19′-hexanoyloxyfucoxanthin-containing prymnesiophytes were the dominant group, although procaryotes accounted for 65% at the two southerly oligotrophic stations.     

Mesoscale decrease of surface phosphate and associated phytoplankton dynamics in the vicinity of the subtropical South Pacific islands

Surface distributions of nutrients and phytoplankton were investigated in the vicinity of the subtropical South Pacific islands by using a continuous underway system with a highly sensitive nutrient analysis. A total of 17 transects, whose lengths ranged between 42 and 271 km, were sampled for continuous nutrient measurements. The study area was characterized by an overall depletion of nitrate+nitrite (<15 nM), but phosphate varied from 7 to 192 nM. The transects were grouped into 4 patterns according to distribution of phosphate concentrations. In 7 transects, a mesoscale decrease in phosphate occurred, coinciding with an elevation of in vivo chlorophyll fluorescence, which was accompanied by an increase in phytoplankton abundance as revealed by microscopy, flow cytometry, and accessory pigments. This mirror–image relationship between the phosphate concentration and phytoplankton abundance was most apparent on both a 99-km transect east of Tonga, where the phosphate concentration ranged from 17 to 125 nM, and on a 98-km transect west of Fiji, where the phosphate concentration ranged from 23 to 136 nM. Both these transects contained distinct blooms of Trichodesmium in areas with the lowest concentrations of phosphate. In other 2 transects, fluctuations in phosphate concentrations showed no distinct relationship with chlorophyll fluorescence. Other patterns that emerged included consistently high concentrations, ranging from 109 to 192 nM, in 5 transects and consistently low phosphate concentrations, ranging from 7 to 50 nM, in 3 transects. Abundance of Trichodesmium, Prochlorococcus, Synechococcus, and all accessory pigments examined tended to be higher in the low phosphate transects than in the high phosphate ones. In particular, Trichodesmium occurred in low phosphate water (<25 nM). There was no significant relationship between phosphate concentrations and nanoplanktonic unicellular cyanobacteria. Our observations suggest that surface phosphate decreases are associated with phytoplankton utilization of phosphate, and that nitrogen supply from Trichodesmium may contributes to this utilization.     

Responses of pico- and nanophytoplankton to artificial iron infusions observed during the second iron enrichment experiment in the western subarctic Pacific (SEEDS II)

How pico- and nanophytoplankton responded to artificial iron infusions was investigated using flow cytometry during SEEDS II, the second mesoscale in situ iron enrichment experiment in the western subarctic North Pacific. Two iron infusions on days 0 and 7 caused a remarkable increase in cellular chlorophyll fluorescence and cell size of all the four phytoplankton groups investigated: Synechococcus, cryptophytes, picoeucaryotes and nanoeucaryotes other than cryptophytes. After the second infusion, the abundance of three phytoplankton groups, excluding Synechococcus, also started to increase. After surface dissolved iron concentration decreased to <0.2 nM on day 11, chlorophyll fluorescence of all the four groups returned to a level observed before the iron infusions, suggesting that pico- and nanophytoplankton were physiologically stressed by iron deficiency. Cell concentrations of pico- and nanoeucaryotes decreased to the pre-infusion level by day 23, while that of cryptophytes remained high until day 25. Flow cytometric diagnosis showed that cryptophytes were physiologically limited during this period and effective iron uptake from suspended particles, as reported for freshwater cryptophytes, was not observed. Thus their prosperity may have been due to alleviation from grazing. Cell concentration of Synechococcus started to increase at a net specific growth rate of 0.13 d⁻¹ after day 12, reaching more than 6 times that of the pre-infusion level on day 24. This may have been due to the elevation of surface water temperature observed during the survey period, together with trophic cascading effects of increased copepod grazing.     

Microbial biomass and viral infections of heterotrophic prokaryotes in the sub-surface layer of the central Arctic Ocean

Seawater samples were collected for microbial analyses between 55 and 235 m depth across the Arctic Ocean during the SCICEX 97 expedition (03 September–02 October 1997) using a nuclear submarine as a research platform. Abundances of prokaryotes (range 0.043–0.47×10⁹ dm⁻³) and viruses (range 0.68–11×10⁹ dm⁻³) were correlated (r=0.66, n=150) with an average virus:prokaryote ratio of 26 (range 5–70). Biomass of prokaryotes integrated from 55 to 235 m ranged from 0.27 to 0.85 g C m⁻² exceeding that of phytoplankton (0.005–0.2 g C m⁻²) or viruses (0.02–0.05 g C m⁻²) over the same depth range by an order of magnitude on average. Using transmission electron microscopy (TEM), we estimated that 0.5% of the prokaryote community on average (range 0–1.4%) was visibly infected with viruses, which suggests that very little of prokaryotic secondary production was lost due to viral lysis. Intracellular viruses ranged from 5 to >200/cell, with an average apparent burst size of 45±38 (mean±s.d.; n=45). TEM also revealed the presence of putative metal-precipitating bacteria in 8 of 13 samples, which averaged 0.3% of the total prokaryote community (range 0–1%). If these prokaryotes are accessible to protistan grazers, the Fe and Mn associated with their capsules might be an important source of trace metals to the planktonic food web. After combining our abundance and mortality data with data from the literature, we conclude that the biomass of prokaryoplankton exceeds that of phytoplankton when averaged over the upper 250 m of the central Arctic Ocean and that the fate of this biomass is poorly understood.     

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.     

Picophytoplankton in the equatorial Pacific: vertical distributions in the warm pool and in the high nutrient low chlorophyll conditions

Abundance distribution and cellular characteristics of picophytoplankton were studied in two distinct regions of the equatorial Pacific: the western warm pool (0°, 167°E), where oligotrophic conditions prevail, and the equatorial upwelling at 150°W characterized by high-nutrient low-chlorophyll (HNLC) conditions. The study was done in September–October 1994 during abnormally warm conditions. Populations of Prochlorococcus, orange fluorescing Synechococcus and picoeukaryotes were enumerated by flow cytometry. Pigment concentrations were studied by spectrofluorometry. In the warm pool, Prochlorococcus were clearly the dominant organisms in terms of cell abundance, estimated carbon biomass and measured pigment concentration. Integrated concentrations of Prochlorococcus, Synechococcus and picoeukaryotes were 1.5×10¹³, 1.3×10¹¹ and 1.5×10¹¹ cells m⁻², respectively. Integrated estimated carbon biomass of picophytoplankton was 1 g m⁻², and the respective contributions of each group to the biomass were 69, 3 and 28%. In the HNLC waters, Prochlorococcus cells were slightly less numerous than in the warm pool, whereas the other groups were several times more abundant (from 3 to 5 times). Abundance of Prochlorococcus, Synechococcus and picoeukaryotes were 1.2×10¹³, 6.2×10¹¹ and 5.1×10¹¹ cells m⁻², respectively. The integrated biomass was 1.9 g C m⁻². Prochlorococcus was again the dominant group in terms of abundance and biomass (chlorophyll, carbon); the respective contributions of each group to the carbon biomass were 58, 7 and 35%. In the warm pool the total chlorophyll biomass was 28 mg m⁻², 57% of which was divinyl chlorophyll a. In the HNLC waters, the total chlorophyll biomass was 38 mg m⁻², 44% of which was divinyl chlorophyll a. Estimates of Prochlorococcus, Synechococcus and picoeukaryotes cell size were made in both hydrological conditions.     

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.     

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.     

Carbon fluxes through major phytoplankton groups during the spring bloom and post-bloom in the Northwestern Mediterranean Sea

The carbon flux through major phytoplankton groups, defined by their pigment markers, was estimated in two contrasting conditions of the Northwestern Mediterranean open ocean ecosystem: the spring bloom and post-bloom situations (hereafter Bloom and Post-bloom, respectively). During Bloom, surface chlorophyll a (Chl a) concentration was higher and dominated by diatoms (53% of Chl a), while during Post-bloom Synechococcus (42%) and Prymnesiophyceae (29%) became dominant. The seawater dilution technique, coupled to high pressure liquid chromatography (HPLC) analysis of pigments and flow cytometry (FCM), was used to estimate growth and grazing rates of major phytoplankton groups in surface waters. Estimated growth rates were corrected for photoacclimation based on FCM-detected changes in red fluorescence per cell. Given the 30% average decrease in the pigment content per cell between the beginning and the end of the incubations, overlooking photoacclimation would have resulted in a 0.40 d^?1 underestimation of phytoplankton growth rates. Corrected average growth rates (μ_o) were 0.90±0.20 (SD) and 0.40±0.14 d^?1 for Bloom and Post-bloom phytoplankton, respectively. Diatoms, Cryptophyceae and Synechococcus were identified as fast-growing groups and Prymnesiophyceae and Prasinophyceae as slow-growing groups across Bloom and Post-bloom conditions. The higher growth rate during Bloom was due to dominance of phytoplankton groups with higher growth rates than those dominating in Post-bloom. Average grazing rates (m) were 0.58±0.20 d^?1 (SD) and 0.31±0.07 d^?1. The proportion of phytoplankton growth consumed by microzooplankton grazing (m/μ_o) tended to be lower in Bloom (0.69±0.34) than in Post-bloom (0.80±0.08). The intensity of nutrient limitation experienced by phytoplankton indicated by μ_o/μ_n (where μ_n is the nutrient-amended growth rate), was similar during Bloom (0.78) and Post-bloom (0.73). Primary production from surface water (PP) was estimated with ¹⁴C incubations. A combination of PP and Chl a synthesis rate yielded C/Chl a ratios of 34±21 and 168±75 (g:g) for Bloom and Post-bloom, respectively. Transformation of group-specific Chl a fluxes into carbon equivalents confirmed the dominant role of diatoms during Bloom and Synechococcus and Prymnesiophyceae during Post-bloom.     

The chemical speciation of iron in the north-east Atlantic Ocean

The distribution of dissolved iron and its chemical speciation (organic complexation and redox speciation) were studied in the northeastern Atlantic Ocean along 23°W between 37 and 42°N at depths between 0 and 2000 m, and in the upper-water column (upper 200 m) at two stations further east at 45°N10°W and 40°N17°W in the early spring of 1998. The iron speciation data are here combined with phytoplankton data to suggest cyanobacteria as a possible source for the iron binding ligands. The organic Fe-binding ligand concentrations were greater than that of dissolved iron by a factor of 1.5–5, thus maintaining iron in solution at levels well above it solubility. The water column distribution of the organic ligand indicates in-situ production of organic ligands by the plankton (consisting mainly of the cyanobacteria Synechococcus sp.) in the euphotic layer and a remineralisation from sinking biogenic particles in deeper waters. Fe(II) concentrations varied from below the detection limit (<0.1 nM) up to 0.55 nM but represented only a minor fraction of 0% to occasionally 35% of the dissolved iron throughout the water column. The water column distribution of the Fe(II) suggests biologically mediated production in the deep waters and photochemical production in the euphotic layer. Although there was no evidence of iron limitation in these waters, the aeolian iron input probably contributed to a shift in the phytoplankton assemblage towards increased Synechococcus growth.     

Distribution of bacterial abundance and cell-specific nucleic acid content in the Northeast Pacific Ocean

We tested the idea that bacterial cells with high nucleic acid content (HNA cells) are the active component of marine bacterioplankton assemblages, while bacteria with low nucleic acid content (LNA cells) are inactive, with a large data set (>1700 discrete samples) based on flow cytometric analysis of bacterioplankton in the Northeast Pacific Ocean off the coast of Oregon and northern California, USA. Samples were collected in the upper 150 m of the water column from the coast to 250 km offshore during 14 cruises from March 2001 to September 2003. During this period, a wide range of trophic states was encountered, from dense diatom blooms (chlorophyll-a concentrations up to 43 μg l⁻¹) at shelf stations during upwelling season (March–September) to lower chlorophyll-a concentrations (0.1–5 μg l⁻¹) during winter (November–February) and at basin stations (>1700 m depth). We found only weakly positive relations of log total bacterial abundance to log chlorophyll-a concentration (as a proxy for availability of organic substrate), and of HNA bacteria as a fraction of total bacteria to log chlorophyll-a. Abundance of HNA and LNA bacteria co-varied positively in all regions, although HNA bacteria were more responsive to high phytoplankton biomass in shelf waters than in slope and basin waters. Since LNA cell abundance in general showed responses similar to those of HNA cell abundance to changes in phytoplankton biomass, our data do not support the hypothesis that HNA cells are the sole active component of marine bacterioplankton.     

Size-fractionated biological iron and carbon uptake along a coastal to offshore transect in the NE Pacific

Iron has been shown to limit phytoplankton growth in high-nutrient low-chlorophyll (HNLC) regions such as the NE subarctic Pacific. We report size-fractionated Fe-uptake rates by the entire plankton community in short (6–8 h) light and dark incubations along an E–W transect from P04 (a coastal ocean station) to OSP (an open-ocean HNLC station) during August–September 1997. Size-fractionated primary productivity and chl a were measured to monitor algal Fe : C uptake ratios and Fe-uptake relative to phytoplankton biomass. The >5.0 μm size-class, which consisted mostly of large diatoms, had the highest Fe-uptake rate at nearshore stations (P04 and P8), but Fe-uptake rates for this size class decreased despite increases in biomass and primary productivity when transecting westwards to HNLC waters. Fe-uptake rates of the small size class (0.2–1.0 μm, including heterotrophic bacteria and autotrophs) were inversely related to the >5.0 μm size-class uptake rates, in that stations with high dissolved Fe (DFe) concentrations had relatively low uptake rates compared to those in the low-Fe offshore region. The 1.0–5.0 μm size-class Fe-uptake rates were low, relatively invariant along the transect, and differed little between light and dark incubations. Dark Fe-uptake rates averaged 10–20% less than those in the light for the >5.0 μm size class. Dark uptake rates however, were higher than light uptake rates for the 0.2–1.0 μm size class at all stations. Fe : C uptake ratios were high for all size classes at P04, but decreased as DFe concentrations decreased offshore. The prokaryote-dominated 0.2–1.0 μm size class had the highest Fe : C uptake ratios at all stations. These data suggest that prokaryotic organisms make an important contribution to biological Fe uptake in this region. Our experiments support the results of previous culture work, suggesting higher Fe : C ratios in coastal phytoplankton compared to open-ocean species, and demonstrate that light can have a large effect on Fe partitioning between size classes in subarctic Pacific HNLC waters.     

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.     

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.     

Factors controlling the temporal and spatial variations in Synechococcus abundance in a monsoonal estuary

Temporal and spatial variations in Synechococcus abundance were investigated over an annual cycle (February'10–January'11) along a salinity gradient (0–35) in the tropical Zuari estuary, influenced by south-west monsoons. Synechococcus exhibited salinity preferences with phycoerythrin-rich cells at salinities >2 (Synechococcus-PEI), >20 (Synechococcus-PEII) and <1 (Synechococcus-PEIII) whereas phycocyanin-rich (Synechococcus-PC) dominant at lower salinities. Downstream stratification during monsoon caused Synechococcus group segregation in the surface and near-bottom waters. During monsoon-break and non-monsoon period stabilized waters, increased salinity, temperature, solar radiation and low rainfall favored high Synechococcus abundance whereas unstable waters, increased turbidity and low solar radiation during active monsoon lowered abundance. SYN-PC positively co-related with nitrate and phosphate and SYN-PEI with phosphate. Synechococcus contribution to phytoplankton carbon biomass ranged from 9 to 29%. In monsoonal estuaries, rainfall intensity regulates freshwater runoff which modulates the estuarine environment, creating temporal–spatial niche segregation of Synechococcus groups thereby serving as indicator organisms of the estuarine hydrodynamics.     

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.     

The distribution and vertical flux of fecal pellets from large zooplankton in Monterey bay and coastal California

We sampled zooplankton and fecal pellets in the upper 200 m of Monterey Bay and nearby coastal regions in California, USA. On several occasions, we observed high concentrations of large pellets that appeared to be produced during night-time by dielly migrating euphausiids. High concentrations of pellets were found in near-surface waters only when euphausiids co-occurred with high concentrations of large (>10 μm) phytoplankton. Peak concentrations of pellets at mid-depth (100 or 150 m) during the day were consistent with the calculated sinking speeds of pellets produced near the surface at night. At these high flux locations (HI group), pellet concentrations declined below mid-depth. In contrast, at locations where the phytoplankton assemblage was dominated by small phytoplankton cells (<10 μm), pellet production and flux were low (LO group) whether or not euphausiid populations were high. Protozooplankton concentrations did not affect this pattern. We concluded that the day and night differences in pellet concentration and flux in the HI profiles were mostly due to sinking of dielly-pulsed inputs in the surface layer, and that small zooplankton (Oithona, Oncaea), heterotrophic dinoflagellates, and bacterial activity probably caused some pellet degradation or consumption below 100 m. We estimated that consumption of sinking pellets by large copepods was insignificant. High fluxes of pellets were episodic because they required both high concentrations of large phytoplankton and large stocks of euphausiids. Under these conditions, flux events overwhelmed retention mechanisms, resulting in large exports of organic matter from the upper 200 m.