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.     

Dissolved DMSO production via biological and photochemical oxidation of dissolved DMS in the Ross Sea,Antarctica

Dimethylsulfoxide (DMSO) is an important degradation product of the climate-influencing gas dimethylsulfide (DMS). In the Ross Sea, Antarctica, dissolved DMSO (DMSOd) concentrations exhibited substantial seasonal and vertical variations. Surface water DMSOd concentrations in pre-bloom waters were very low (<1 nM) but increased rapidly up to 41 nM during the spring Phaeocystis antarctica bloom (late November). Increases in DMSOd concentrations lagged by several days increases in DMS concentrations. Although DMSOd concentrations reached relatively high levels during the spring bloom, concentrations were generally higher (36.3–60.6 nM) during summer (January), even though phytoplankton biomass and DMS concentrations had decreased by that time. During both seasons, DMSOd concentrations were substantially higher within the surface mixed layer than below it. DMSOd production from biological DMS consumption (BDMSC) was higher during late November (3.4–5.2 nM d^?1) than during the summer (0.7–2.4 nM d^?1); therefore, production via BDMSC alone could not explain the higher DMSOd concentrations encountered during the summer. Mixed layer-integrated DMSOd production from BDMSC was 2.5–13.7 times greater than production from dissolved DMS photolysis during the P. antarctica bloom, while photolysis contributed 1.3 times more DMSO than BDMSC before the bloom. The DMSO yield from BDMSC was consistently higher within the upper mixed layer than at depths below. Experimental incubations with water from the mixed layer showed that exposure to full spectrum sunlight for 72 h caused an increase in the DMSO yield whereas exposure to only photosynthetically active radiation did not. This suggests that ultraviolet radiation is a potential factor shifting the fate of biologically consumed DMS toward DMSO. In general, the highest DMSO yields from BDMSC were in samples with slow biological DMS turnover, whereas fast turnover favored sulfate rather than DMSO as a major end-product. This study provides the first detailed information about DMSOd distribution and production in the Ross Sea and points to DMSOd as an important biological and photochemical degradation product of DMS and a major reservoir of methylated sulfur in these polar surface waters.     

Phytoplankton chemotaxonomy in the Atlantic sector of the Southern Ocean during late summer 2009

A chemotaxonomic investigation of surface phytoplankton was undertaken on a research cruise to the Atlantic sector of the Southern Ocean during late austral summer 2009. Based on pigment signatures, several distinct regions emerged that were delineated by physical features. CHEMTAX analysis of high performance liquid chromatography (HPLC) pigment data indicated that diatoms generally dominated communities south of the Antarctic Polar Front (APF), particularly in regions of elevated biomass where chlorophyll-a (chl-a) was >1.5 µg l⁻¹ and diatoms comprised >80% of biomass. Pigment signatures representative of haptophytes-8, indicative of Phaeocystis antarctica, were dominant near the ice shelf. Chl-a concentrations were 0.2–0.6 µg l⁻¹ between the APF and the Subtropical Front (STF) and outputs suggested that chlorophytes, haptophytes-8 and haptophyte-6, in the form of coccolithophores, were the major constituents. Very low chl-a levels (<0.2 µg l⁻¹) were observed north of the STF and the prokaryotes Synechococcus spp. and Prochlorococcus spp. were the dominant groups in these oligotrophic waters.     

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.     

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.     

Occurrence and turnover of DMSP and DMS in deep waters of the Ross Sea,Antarctica

High concentrations of the phytoplankton metabolite dimethylsulfoniopropionate (DMSP) and its degradation product dimethylsulfide (DMS) are associated with blooms of Phaeocystis antarctica in the Ross Sea, Antarctica. Episodic and rapid vertical export of Phaeocystis biomass to deep water has been reported for the Ross Sea, therefore we examined the distribution and microbial consumption rates of DMSP and DMS throughout the sub-euphotic water column. Total DMSP (dissolved+particulate; DMSPt) was present at 0.5–22 nM at depths between 70 and 690 m during both the early bloom (November) and the late bloom (January). Sub-euphotic peaks of DMSP were sometimes associated with mid-water temperature maxima, and elevated DMSP below 70 m was found mainly in water masses characterized as Modified Circumpolar Deep Water or Antarctic Shelf Water. Overall, 50–94% of the integrated water-column DMSPt was found below the euphotic zone. At one station during the early bloom, local maxima of DMSPt (14 nM) and DMS (20 nM) were observed between 113 and 240 m and these maxima corresponded with high chlorophyll a concentrations, P. antarctica cell numbers, and Fv/Fm (the quantum yield of photosystem II). During the late bloom, a sub-euphotic maximum of DMSPt (15.8 nM) at 250 m cooccurred with peaks of chlorophyll a concentration, DMSP lyase activity, bacterial production and dissolved DMSP consumption rates. DMSP turnover contributed ～12% of the bacterial carbon demand between 200 and 400 m. DMS concentrations peaked at 286 m but the maximum concentration (0.42 nM) was far lower than observed during the early bloom, probably because of relatively rapid biological consumption of DMS (1–3 turnovers per day) which, in turn, contributed to elevated dissolved dimethylsulfoxide (DMSO) concentrations. Relatively stable DMSPt distributions at some sites suggest that rapid sinking of Phaeocystis biomass is probably not the major mechanism responsible for mesopelagic DMSP accumulations. Rather, subduction of near-surface water masses, lateral advective transport or trapping of slowly sinking P. antarctica biomass in intermediate water masses are more likely mechanisms. We found that a culture of P. antarctica maintained cellular integrity during 34 days of darkness, therefore the presence of intact cells (and DMSP) at depth can be explained even under a slow sinking/advection scenario. Whatever the mechanism, the large pools of DMSP and DMS below the euphotic zone suggest that export exerts a control on potential DMS emission from the surface waters of the Ross Sea.     

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.     

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.     

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

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

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.     

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.     

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

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

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.     

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

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

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.     

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.     

Spatial variability and temporal dynamics of surface water pCO2, ��O2/Ar and dimethylsulfide in the Ross Sea, Antarctica

We report results from two surveys of pCO₂, biological O₂ saturation (??O₂/Ar) and dimethylsulfide (DMS) in surface waters of the Ross Sea polynya. Measurements were made during early spring (November 2006-December 2006) and mid-summer (December 2005-January 2006) using ship-board membrane inlet mass spectrometry (MIMS) for high spatial resolution (i.e. sub-km) analysis. During the early spring survey, the polynya was in the initial stages of development and exhibited a rapid increase in open water area and phytoplankton biomass over the course of our ∼3 week occupation. We observed a rapid transition from a net heterotrophic ice-covered system (supersaturated pCO₂ and undersaturated O₂) to a high productivity regime associated with a Phaeocystis-dominated phytoplankton bloom. The timing of the early spring phytoplankton bloom was closely tied to increasing sea surface temperature across the polynya, as well as reduced wind speeds and ice cover, leading to enhanced vertical stratification. There was a strong correlation between pCO₂, ??O₂/Ar, DMS and chlorophyll a (Chl a) during the spring phytoplankton bloom, indicating a strong biological imprint on gas distributions. Box model calculations suggest that pCO₂ drawdown was largely attributable to net community production, while gas exchange and shoaling mixed layers also exerted a strong control on the re-equilibration of mixed layer ??₂ with the overlying atmosphere. DMS concentrations were closely coupled to Phaeocystis biomass across the early spring polynya, with maximum concentrations exceeding 100 nM.During the summer cruise, we sampled a large net autotrophic polynya, shortly after the seasonal peak in phytoplankton productivity. Both diatoms and Phaeocystis were abundant in the phytoplankton assemblages during this time. Minimum pCO₂ was less than 100 ppm, while ??O₂/Ar exceeded 30% in some regions. Mean DMS concentrations were ∼2-fold lower than during the spring, although the range of concentrations was similar between the two surveys. There was a significant correlation between pCO₂, ??O₂/Ar and Chl a across the summer polynya, but the strength of these correlations and the slope of O₂ vs. CO₂ relationship were significantly lower than during the early spring. Summertime DMS concentrations were not significantly correlated to phytoplankton biomass (Chl a), pCO₂ or ??O₂/Ar. In contrast to the early spring time, there were no clear temporal trends in summertime gas concentrations. Rather, small-scale spatial variability, likely resulting from mixing and localized sea-ice melt, was clearly evident in surface gas distributions across the polynya. Analysis of length-scale dependent variability demonstrated that much of the spatial variance in surface water gases occurred at scales of <20 km, suggesting that high resolution analysis is needed to fully capture biogeochemical heterogeneity in this system.     

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.     

Phytoplankton pigment and absorption characteristics along meridional transects in the Atlantic Ocean

Pigment patterns and associated absorption properties of phytoplankton were investigated in the euphotic zone along two meridional transects in the Atlantic Ocean, between the UK and the Falkland Islands, and between South Africa and the UK. Total chlorophyll a (TChla=MVChla+DVChla+chlorophyllide a) concentrations and the biomarker pigments for diatoms (fucoxanthin), nanoflagellates and cyanobacteria (zeaxanthin) appeared to have similar distribution patterns in the spring and in the autumn in the temperate NE Atlantic and the northern oligotrophic gyre. Divinyl chlorophyll a levels (prochlorophytes) were greater in spring at the deep chlorophyll maximum in the oligotrophic gyre, however. Marked seasonal differences were observed in the NW African upwelling region. TChla concentrations were twice as high in the upper mixed layer in the spring, with the community dominated by diatoms and prymnesiophytes (19′-hexanoyloxyfucoxanthin). A layered structure was prevalent in the autumn where cyanobacteria, diatoms and prymnesiophytes were located in the upper water column and diatoms and mixed nanoflagellates at the sub-surface maximum. In the South Atlantic, the Benguela upwelling ecosystem and the Brazil-Falklands Current Confluence Zone (BFCCZ) were the most productive regions with the TChla levels being twice as high in the Benguela. Diatoms dominated the Benguela system, while nanoflagellates were the most ubiquitous group in the BFCCZ. Pigment concentrations were greater along the eastern boundary of the southern oligotrophic gyre and distributed at shallower depths. Deep chlorophyll maxima were a feature of the western boundary oligotrophic waters, and cyanobacteria tended to dominate the upper water column along both transects with a mixed group of nanoflagellates at the chlorophyll maximum.Absorption coefficients were estimated from spectra reconstructed from pigment data. Although absorption was greater in the productive areas, the TChla-specific coefficients were higher in oligotrophic regions. In communities that were dominated by diatoms or nanoflagellates, pigment absorption was generally uniform with depth and attenuating irradiance, with TChla being the major absorbing pigment at 440 nm and photosynthetic carotenoids (PSC) at 490 nm. Absorption by chlorophyll c and photoprotective carotenoids (PPC) was much lower. Populations where cyanobacteria were prevalent were characterized by high PPC absorption, particularly at 490 nm, throughout most of the euphotic zone. The data suggested that the effect of pigments on the variability of phytoplankton absorption was due primarily to the variations in absorption by PPC.     

Modelling phytoplankton deposition to Chesapeake Bay sediments during winter–spring: interannual variability in relation to river flow

The often-rapid deposition of phytoplankton to sediments at the end of the spring phytoplankton bloom is an important component of benthic–pelagic coupling in temperate and high latitude estuaries and other aquatic systems. However, quantifying the flux is difficult, particularly in spatially heterogeneous environments. Surficial sediment chlorophyll-a, which can be measured quickly at many locations, has been used effectively by previous studies as an indicator of phytoplankton deposition to estuarine sediments. In this study, surficial sediment chlorophyll-a was quantified in late spring at 20–50 locations throughout Chesapeake Bay for 8 years (1993–2000). A model was developed to estimate chlorophyll-a deposition to sediments using these measurements, while accounting for chlorophyll-a degradation during the time between deposition and sampling. Carbon flux was derived from these estimates via C:chl-a = 75.Bay-wide, the accumulation of chlorophyll-a on sediments by late spring averaged 171 mg m⁻², from which the chlorophyll-a and carbon sinking fluxes, respectively, were estimated to be 353 mg m⁻² and 26.5 gC m⁻². These deposition estimates were ∼50% of estimates based on a sediment trap study in the mid-Bay. During 1993–2000, the highest average chlorophyll-a flux was in the mid-Bay (248 mg m⁻²), while the lowest was in the lower Bay (191 mg m⁻²). Winter–spring average river flow was positively correlated with phytoplankton biomass in the lower Bay water column, while phytoplankton biomass in that same region of the Bay was correlated with increased chlorophyll-a deposition to sediments. Responses in other regions of the Bay were less clear and suggested that the concept that nutrient enrichment in high flow years leads to greater phytoplankton deposition to sediments may be an oversimplification. A comparison of the carbon flux associated with the deposition of the spring bloom with annual benthic carbon budgets indicated that the spring bloom did not contribute a disproportionately large fraction of annual carbon inputs to Chesapeake Bay sediments. Regional patterns in chlorophyll-a deposition did not correspond with the strong regional patterns that have been found for plankton net community metabolism during spring.