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 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.     

Characterisation of water masses and phytoplankton nutrient limitation in the East Australian Current separation zone during spring 2008

This study focuses on the comparison of oceanic and coastal cold-core eddies with inner-shelf and East Australian Current (EAC) waters at the time of the spring bloom (October 2008). The surface water was biologically characterised by the phytoplankton biomass, composition, photo-physiology, carbon fixation and by nutrient-enrichment experiments. Marked differences in phytoplankton biomass and composition were observed. Contrasted biomarker composition suggests that biomarkers could be used to track water masses in this area. Divinyl chlorophyll a, a biomarker for tropical Prochlorophytes, was found only in the EAC. Zeaxanthin a biomarker for Cyanophytes, was found only within the oceanic eddy and in the EAC, whereas chlorophyll b (Chlorophytes) was only present in the coastal eddy and at the front between the inner-shelf and EAC waters.This study showed that cold-core eddies can affect phytoplankton, biomass, biodiversity and productivity. Inside the oceanic eddy, greater phytoplankton biomass and a more complex phytoplankton community were observed relative to adjacent water masses (including the EAC). In fact, phytoplankton communities inside the oceanic eddy more closely resembled the community observed in the inner-shelf waters. At a light level close to half-saturation, phytoplankton carbon fixation (gC d⁻¹) in the oceanic eddy was 13-times greater than at the frontal zone between the eddy and the EAC and 3-times greater than in the inner-shelf water. Nutrient-enrichment experiments demonstrated that nitrogen was the major macronutrient limiting phytoplankton growth in water masses associated with the oceanic eddy. Although the effective quantum yield values demonstrate healthy phytoplankton communities, the phytoplankton community bloomed and shifted in response to nitrogen enrichments inside the oceanic eddy and in the frontal zone between this eddy and the EAC. An effect of Si enrichment was only observed at the frontal zone between the eddy and the EAC. No response to nutrient enrichment was observed in the inner-shelf water where ambient NO_x, Si and PO₄ concentrations were up to 14, 4 and 3-times greater than in the EAC and oceanic eddy. Although results from the nutrient-enrichment experiments suggest that nutrients can affect biomass and the composition of the phytoplankton community, the comparison of all sites sampled showed no direct relationship between phytoplankton biomass, nutrients and the depth of the mixed layer. This is probably due to the different timeframe between the rapidly changing physical and chemical oceanography in the separation zone of the EAC.     

Distribution and sinking rates of phytoplankton, detritus, and particulate biogenic silica in the Laptev Sea and Lena River (Arctic Siberia)

Concentrations and sinking rates of particulate biogenic silica (BSi), chlorophyll a (chl a) and phaeopigments (phae) (< 3 μm, 3–10 μm, > 10 μm and total), as well as the abundances of the major phytoplankton species, were studied during September 1991 in the Eastern Laptev Sea and the lower Lena River (Siberian Arctic). The highest chl a concentrations were found in two major “new” production regimes of the study area: (1) a deep chl a maximum (5.8 mg chl a m⁻³) (formed by the diatom Chaetoceros socialis) at 30 m depth on the outer shelf of the northern Laptev Sea, and (2) in the Lena River, where the phytoplankton community was dominated by fresh water diatoms (1.5 to 4.5 mg chl a m⁻³). Elevated chl a concentrations were also found in the river plume phytoplankton community (dominated by brackish water diatoms), NE of the Lena delta. In the Laptev Sea, the low chl a (0.1 to 3 mg chl a m⁻³) and high phae concentrations (0.5 to 14 mg phae m⁻³) indicated that the phytoplankton community (dominated by picoplanktic algae and nanoflagellates) was already senescent and affected by grazing losses. Biogenic silica values were highest in the Lena River (4 to 17 μM) as compared to the low values found in the Laptev Sea (0.3 to 4 μM). The large chl a size fraction, phae and BSi in the Lena River samples revealed the highest measured sinking rates (1.4, 2.3, and 1.5 m d⁻¹, respectively). The formation of a strong halocline, decreasing turbulence, and possible nutrient deficiency resulted in death, disintegration and rapid sedimentation of fresh water diatoms. This was accompanied by a decrease in the BSi concentration and growth of the picoplanktic size fraction (< 3 μm) in the estuarine mixing zone (Gulf of Buorkhaya). Only a minor part of BSi was bound to intact diatom cells (< 3%) in the surface layer, most of which being apparently associated with detrital particles. In the Lena River, approximately 12% of the total silica was bound to BSi fraction, yet elsewhere in the Laptev Sea and in the estuarine mixing zone the BSi:total silica ratio was ≤ 5%. Thus, the results reflected the successional stage of a late summer phytoplankton community, characterized by dominance of small autotrophs and patchy distribution of senescent diatoms no longer able to affect the relative high levels of dissolved silica supplied by the Lena River.     

Photosynthetic pigments as indicators of phytoplankton development during spring and summer in Adventfjorden (Spitsbergen)

The study was carried out from April 30 until July 13 of 1997 in Adventfjorden (Spitsbergen). Formation of less saline and warmer surface water (～1 m thick) caused by melting of the fast ice was observed in the fjord during the first days of May. In summer a less saline surface layer was about 3 m thick. Euphotic depth measured under ice sheet reached 12 m, whereas load of mineral matter brought with riverine discharge in summer (the content of total particulate matter in the fjord reached 1.66 kg m^?2) dramatically reduced euphotic zone depth to 0.35 m. By pigment measurement three phases of phytoplankton development in Adventfjorden were distinguished: (1) spring bloom that has started under fast ice and reached maximum in the mid of May, (2) stagnation period in June, (3) increase of pigment concentration in July, what could indicate a start of the next algae bloom. Analyses of chlorophylls and carotenoids revealed that diatoms (chl c, fucoxanthin), and green algae (chl b, lutein) dominated phytoplankton community in the fjord. Moreover, the presence of peridinin indicates the presence of Dinophyta and alloxanthin—the occurence of Cryptophyta. In May and June 1997 phytoplankton appeared mainly in the surface of water, while in July, as a result of inflow of turbulent riverine waters into Adventfjorden, algae cells were pushed down and the highest numbers were observed at the depth ～20 m. Great phaeopigments to chl a ratio (= 0.54) found in the fjord seston in June and July probably shows strong impact of zooplankton grazing on phytoplankton development. High contribution of chlorophyllide a in porphyrin a poll in samples collected under fast ice (chlorophyllide a/chl a ratio = 0.18) reflects the final stage of algal communitie succession in ice, just before spring ice melt and release of biota to oceanic water. Chloropyllide a content during summer was minor or not detectable, demonstrating that diatom cells were in good physiological condition. High chl a allomer/chl a ratio (average = 0.11 for the period investigated) confirms high oxygen concentration in environment of Adventfjorden.     

Impact of a cyclonic eddy on phytoplankton community structure and photosynthetic competency in the subtropical North Pacific Ocean

A synoptic spatial examination of the eddy Haulani (17–20 November 2000) revealed a structure typical of Hawaiian cyclonic eddies with divergent surface flow forcing the upward displacement of deep waters. Hydrographic surveys revealed that surface water in the eddy center was ca. 3.5°C cooler, 0.5 saltier, and 1.4 kg m⁻³ denser than surface waters outside the eddy. Vertically integrated concentrations of nitrate+nitrite, phosphate and silicate were enhanced over out-eddy values by about 2-fold, and nitrate+nitrite concentrations were ca. 8× greater within the euphotic zone inside the eddy than outside. Si:N ratios were lower within the upper mixed layer of the eddy, indicating an enhanced Si uptake relative to nitrate+nitrite. Chlorophyll a concentrations were higher within the eddy compared to control stations outside, when integrated over the upper 150 m, but were not significantly different when integrated over the depth of the euphotic zone. Photosynthetic competency, assessed using fast repetition-rate fluorometry, varied with the doming of the isopycnals and the supply of macro-nutrients to the euphotic zone. The physical and chemical environment of the eddy selected for the accumulation of larger phytoplankton species. Photosynthetic bacteria (Prochlorococcus and Synechococcus) and small (<3 μm diameter) photosynthetic eukaryotes were 3.6-fold more numerically abundant outside the eddy as compared to inside. Large photosynthetic eukaryotes (>3 μm diameter) were more abundant inside the eddy than outside. Diatoms of the genera Rhizosolenia and Hemiaulus outside the eddy contained diazotrophic endosymbiontic cyanobacteria, but these endosymbionts were absent from the cells of these species inside the eddy. The increase in cell numbers of large photosynthetic eukaryotes with hard silica or calcite cell walls is likely to have a profound impact on the proportion of the organic carbon production that is exported to deep water by sinking of senescent cells and cells grazed by herbivorous zooplankton and repackaged as large fecal pellets.     

Distribution of algal chlorophyll and carotenoid pigments in a stratified estuary: the Krka River, Adriatic Sea

The detailed distribution of algal chlorophyll and carotenoid pigments was determined around the halocline (freshwater-seawater interface) in the Krka Estuary on the east coast of the Adriatic Sea; in May 1988. After collection of water along the estuary, particulate matter was extracted and analyzed for pigments by high-performance liquid chromatography coupled with absorbance and fluorescence detection. Bottom marine waters were characterized by lower chlorophyll a (chl a) concentrations than encountered in surface waters, decreasing downstream from 0.50 μg l⁻¹ to 0.16 μg l⁻¹ at the marine end-member. The highest concentrations of chl α (up to 26.34 μg l⁻¹) were found in the interfacial layer, an particularly at one station located off the city of ibenik, where high inputs of nutrients supported the accumulation of living algae at the halocline. Fucoxanthin was the most abundant carotenoid, which indicates a euryhaline dominance of diatoms in the estuary, whereas the dinoflagellate-derived carotenoid peridinin was confined to the interfacial and bottom saline waters of the inner estuary. High concentrations of alloxanthin and chl b were found in the interfacial layer, which also suggests an accumulation of Cryptophyceae and green algae in the inner estuary. Phaeophorbides showed higher concentrations in bottom waters than in surface waters, whereas the highest concentrations occurred in the interfacial layer. These high levels could reflect a density trapping of dead cells in an early degradation state, as suggested by the importance of allomerized chl a and chlorophyllide a vs. total chl a, or of faecal pellets originating from zooplankton grazing in the interfacial layer.     

Enhanced chlorophyll associated with island-induced cyclonic eddies in the eastern channel of the Tsushima Straits

The relationship between island-induced cyclonic eddies and chlorophyll a (chl-a) was investigated using field data and satellite images in the eastern channel of the Tsushima Straits. The maximum chl-a concentration around the leeward eddy of the Tsushima Islands was two or three times greater than that of outside the eddy. Two different mechanisms of chl-a enhancement associated with island-induced cyclonic eddies were found in the post-bloom periods. In summer, when nutrients were depleted in the surface layer, eddy pumping increased the nutrient supply in the euphotic zone, resulting in enhanced chl-a around the shallow thermocline near the eddy core. In late autumn, when the mixed layer deepened over the euphotic zone, the mixed layer depth became shallow due to the doming effect of the cyclonic eddy, therefore, the improved irradiance condition led to an increase in the chl-a concentration in the surface mixed layer. Nighttime satellite visible images showed a number of fishing vessels in the lee region of the Tsushima Islands, implying that the enhanced phytoplankton biomass may have resulted in good feeding conditions for fishes and squids in the Tsushima Straits.     

Structure and photosynthetic properties of phytoplankton assemblages in a highly dynamic system, the Northern Adriatic Sea

The photosynthetic properties of phytoplankton populations as related to physical–chemical variations on small temporal and spatial scales and to phytoplankton size structure and pigment spectra were investigated in the Northern Adriatic Sea off the Po River delta in late winter 1997. Large diatoms (fucoxanthin) dominated the phytoplankton in the coastal area whereas small phytoflagellates (mainly 19′-hexanoyloxyfucoxanthin, chlorophyll b, 19′-butanoyloxyfucoxanthin) occurred outside the front. The front was defined by the steep gradient in density in the surface layer separating low-salinity coastal waters from the offshore waters.Physical features of the area strongly influenced phytoplankton biomass distributions, composition and size structure. After high volumes of Po River discharge several gyres and meanders occurred in the area off the river delta in February. Decreasing river discharge and the subsequent disappearance of the gyres and the spreading dilution of the river plume was observed in March. The dynamic circulation of February resulted in high photosynthetic capacity of the abundant phytoplankton population (>3.40 mg m⁻³). In March, the slow circulation and an upper low-salinity water layer, segregated from the deeper layers, resulted in lack of renewal of this water mass. The huge phytoplankton biomass, up to 15.77 mg chl a m⁻³, became nutrient depleted and showed low photosynthetic capacity. In February, an exceptionally high P_max^B, 20.11 mg C (mg chl a)⁻¹ h⁻¹ was recorded in the Po River plume area and average P_max^B was three-fold in February as compared to the March recordings, 10.50 mg C (mg chl a)⁻¹ h⁻¹ and 3.22 mg C (mg chl a)⁻¹ h⁻¹, respectively.The extreme variability and values of phytoplankton biomass in the innermost plume area was not always reflected in primary production. Modeling of circulation patterns and water mass resilience in the area will help to predict phytoplankton response and biomass distributions. In the frontal area, despite a considerable variability in environmental conditions, our findings have shown that the phytoplankton assemblages will compensate for nutrient depression and hydrographic constraints, by means of size and taxonomic composition and, as a result, the variability in the photosynthetic capacity was much less pronounced than that observed for other parameters.     

SeaWiFS retrievals of chlorophyll in Chesapeake Bay and the mid-Atlantic bight

This paper presents three years (1998–2000) of chlorophyll a (chl a) data from the Sea-viewing Wide Field-of-view Sensor (SeaWiFS) for Case 2 waters of Chesapeake Bay and the middle Atlantic bight (MAB) to describe phytoplankton dynamics on seasonal to interannual time scales. We used extensive data on inherent and apparent optical properties in conjunction with satellite retrievals to: (1) characterize the bio-optical properties of the study area relevant to processing and interpreting SeaWiFS data; (2) test the applicability of the SeaWiFS bio-optical algorithm (OC4v.4) for the estuarine and coastal waters; (3) evaluate the accuracy of the SeaWiFS remote sensing reflectance (R_RS) and chl a products on regional and seasonal bases using in situ observations. The characteristically strong absorption by chromophoric dissolved organic matter (a_cdom) and non-pigmented particulate matter (a_d) in estuarine and coastal waters contributed to overestimates of chl a using OC4v.4 applied to in situ radiances for the Bay (mean ratio 1.42±1.20) and the MAB (2.60±1.36). Values of R_RS from SeaWiFS in the blue region of the spectrum were low compared to in situ R_RS, suggesting that uncertainties remain in atmospheric correction. Direct comparisons of SeaWiFS retrievals of chl a with in situ chl a for the Bay showed larger biases and uncertainties (mean ratio 1.97±1.85) than for chl a estimated from OC4v.4 applied to in situ R_RS. The larger biases were attributed to errors in SeaWiFS radiances and the larger uncertainties to time-space “aliasing” of satellite observations and in situ measurements. To reduce the time differences between SeaWiFS and in situ data, we compared chl a obtained from continuous underway fluorometric measurements on selected ship tracks to SeaWiFS chl a and showed that SeaWiFS captured phytoplankton dynamics in much of the Bay. The agreement of SeaWiFS chl a with in situ chl a was strongest in the mid- (regions 3, 4) to lower Bay (regions 1, 2), and deteriorated toward the upper Bay (regions 5, 6), in part due to a reduction of sensitivity and an increase of noise for SeaWiFS products in the highly absorbing, low R_RS waters of the upper Bay. A three-year time-series of SeaWiFS and in situ data showed that SeaWiFS accurately and reliably captured seasonal and interannual variability of chl a associated with variations of freshwater flow. Significant short-term variability of chl a in summer that was unresolved with shipboard data was detected in the SeaWiFS time-series and the implications are discussed. The overall performance of SeaWiFS in the mid- to lower Bay and the MAB, combined with high spatial (∼1 km²) and temporal (∼100 clear scenes per year) resolution, indicate current SeaWiFS products are valuable for quantifying seasonal to interannual variability of chl a in estuarine and coastal waters.     

Microbial dynamics in cyclonic and anticyclonic mode-water eddies in the northwestern Sargasso Sea

The EDdy Dynamics, mixing, Export, and Species composition (EDDIES) project provided a unique opportunity to evaluate the response of the microbial community and further understand the biological and biogeochemical consequences of mesoscale perturbation events in an oligotrophic system. In order to characterize microbial dynamics, we performed measurements of bacterial biomass (BB) and production (BP) and phytoplankton pigment analyses in two upwelling eddies in the Sargasso Sea sampled in 2004 and 2005. We also observed a 3-fold increase in BP at the Bermuda Atlantic Time-series Study (BATS) site during the passage of a cyclonic eddy in 2003. Although the integrated BB and BP over 140 m in 2004 and 2005 eddies remained within the climatological range measured at the BATS site, there was systematic variability in bacterioplankton dynamics across both eddies. Cyclonic eddy C1 demonstrated decreased BP at the feature's center relative to its periphery, and BP was not correlated with total chlorophyll a (TChl a) variability. However, BP correlated with prymnesiophyte pigments throughout the feature. In contrast, mode-water eddy A4 showed an enhancement in BP at the eddy center (EC) relative to its edges and was coincident with elevated TChl a, high primary production measurements, and a high concentration of diatoms. In eddy A4, the tight relationship between enhanced BP, TChl a and specific phytoplankton taxa implies that the phytoplankton community structure was an important factor influencing BP variability. While the heterotrophic bacterial response in C1 and A4 was not enhanced relative to BATS summer climatology, these data and the presence of similar nutrient fields across both eddies suggest that BP and BB were influenced by the eddy perturbations and responded to changes in the phytoplankton community.     

Remote sensing of phytoplankton groups in case 1 waters from global SeaWiFS imagery

Ocean color sensors enable a quasi-permanent monitoring of the chlorophyll a concentration (Chl a) in surface waters. This ubiquitous photosynthetic pigment cannot, however, be used to distinguish between phytoplankton species. Distinguishing phytoplankton groups from space is nevertheless necessary to better study some biochemical processes such as carbon fixation at the global scale, and is thus one of the major challenges of ocean color research. In situ data have shown that the water-leaving radiances (nLw), measured by ocean color sensors at different wavelengths in the visible spectrum, vary significantly for a given Chl a. This natural variability is due partly to differences in optical properties of phytoplankton species. Here, we derive relationships between nLw and phytoplankton species by using a large set of quantitative inventories of phytoplankton pigments collected during nine cruises from Le Havre (France) to Nouméa (New Caledonia) in the framework of the GeP&CO program. Coincident SeaWiFS nLw data between 412 and 555 nm are extracted and normalized to remove the effect of Chl a. These normalized spectra vary significantly with in situ pigment composition, so that four major phytoplankton groups, i.e., haptophytes, Prochlorococcus, Synechococcus-like cyanobacteria and diatoms, can be distinguished. This classification (PHYSAT) is applied to the global SeaWiFS dataset for year 2001, and global maps of phytoplankton groups are presented. Haptophytes and diatoms are found mostly in high latitudes and in eutrophic regions. Diatoms show a strong seasonal cycle with large-scale blooms during spring and summer. These results, obtained with only five channels in the visible spectrum, demonstrate that ocean color measurements can be used to discriminate between dominant phytoplankton groups provided that sufficient data are available to establish the necessary empirical relationships.     

Strong CO2 outgassing from high nutrient low chlorophyll coastal waters off central Chile (30°S): The role of dissolved iron

Carbonate system parameters (pH and alkalinity) were used to estimate the coastal water CO₂ fluxes off central Chile (30°S) during September 2007. Coastal waters rich in nitrate and silicate were strongly CO₂ supersaturated and normally poor in chlorophyll a. MODIS satellite chlorophyll a data suggest that phytoplankton biomass remained particularly low during September 2007 although coastal waters were highly fertilized with nitrate and silicate. The phytoplankton gross primary productivity in macronutrient-rich waters was very low with the exception of shallow waters (e.g. within or near bays). Several iron-enrichment bottle experiments show that fCO₂ rapidly decreases during iron-enrichment treatments compared to controls. This suggests that iron limitation of phytoplankton growth (mainly diatoms) plays a role in maintaining high-CO₂ outgassing by preventing rapid interception of upwelled CO₂.     

Phytoplankton composition and its ecological effect in subsurface cold pool of the northern Bering Sea in summer as revealed by HPLC derived pigment signatures

CHEMTAX analysis of high-performance liquid chromatography(HPLC) pigment was conducted to study phytoplankton community structure in the northern Bering Sea shelf, where a seasonal subsurface cold pool emerges. The results showed that fucoxanthin(Fuco) and chlorophyll a(Chl a) were the most abundant diagnostic pigments, with the integrated water column values ranging from 141 to 2 160 μg/m2 and 477 to 5 535 μg/m2, respectively. Moreover, a diatom bloom was identified at Sta. BB06 with the standing stock of Fuco up to 9 214 μg/m3. The results of CHEMTAX suggested that the phytoplankton community in the northern Bering Sea shelf was dominated by diatoms and chrysophytes with an average relative contribution to Chl a of 80% and 12%, respectively, followed by chlorophytes, dinoflagellates, and cryptophytes. Diatoms were the absolutely dominant algae in the subsurface cold pool with a relative contribution exceeding 90%, while the contribution of chrysophytes was generally higher in oligotrophic upper water. Additionally, the presence of a cold pool would tend to favor accumulation of diatom biomass and a bloom that occurred beneath the halocline would be beneficial to organic matter sinks, which suggests that a large part of the phytoplankton biomass would settle to the seabed and support a rich benthic biomass.     

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.     

Phytoplankton pigments,functional types,and absorption properties in the Delagoa and Natal Bights of the Agulhas ecosystem

An investigation of pigments, phytoplankton types and absorption characteristics was conducted in the Delagoa and Natal Bights during late winter and spring in the southwest Indian Ocean. The study demonstrated that small flagellates dominated the phytoplankton communities in both bights and were ubiquitous across a temperature range of 18–24 °C. Diatoms were dominant in patches of cool water (<22 °C) related to upwelling processes and were associated with elevated levels of phytoplankton biomass, while prokaryotes were observed to increase in warm waters >22 °C. Absorption coefficients varied closely with variations in chlorophyll a and specific coefficients were lower for diatoms compared to flagellates. Chlorophyll-specific coefficients also provided useful information on the level of pigment packaging and were related to the proportion of chlorophylls and carotenoids in the pigment pool.     

Depth-stratified phytoplankton dynamics in Cyclone Opal, a subtropical mesoscale eddy

As part of E-Flux III cruise studies in March 2005, we investigated phytoplankton community dynamics in a cyclonic cold-core eddy (Cyclone Opal) in the lee of the Hawaiian Islands. Experimental incubations were conducted under in situ temperature and light conditions on a drift array using a two-treatment dilution technique. Taxon-specific estimates of growth, grazing and production rates were obtained from analyses of incubation results based on phytoplankton pigments, flow cytometry and microscopy. Cyclone Opal was sampled at a biologically and physically mature state, with an 80–100 m doming of isopycnal surfaces in its central region and a deep biomass maximum of large diatoms. Depth-profile experimentation defined three main zones. The upper (mixed) zone (0–40 m), showed little compositional or biomass response to eddy nutrient enrichment, but growth, grazing and production rates were significantly enhanced in this layer relative to the ambient community outside of the eddy. Prochlorococcus spp. dominated the upper mixed layer, accounting for 50–60% of its estimated primary production both inside and outside of Opal. In contrast, the deep zone of 70–90 m showed little evidence of growth rate enhancement and was principally defined by a 100-fold increase of large (>20-μm) diatoms and a shift from Prochlorococcus to diatom dominance (80%) of production. The intermediate layer of 50–60 m marked the transition between the upper and lower extremes but also contained an elevated biomass of physiologically unhealthy diatoms with significantly depressed growth rates and proportionately greater grazing losses relative to diatoms above or below. Microzooplankton grazers consumed 58%, 65% and 55%, respectively, of the production of diatoms, Prochlorococcus and the total phytoplankton community in Cyclone Opal. The substantial grazing impact on diatoms suggests that efficient recycling was the major primary fate of diatom organic production, consistent with the low export fluxes and selective export of biogenic silica, as empty diatom frustules, in Cyclone Opal.     

Phytoplankton characteristics and hydrological conditions in the western part of the Sea of Okhotsk in the spring of 1999 and 2000 based on expeditionary and satellite data

Using the data obtained in 1999–2000 during the spring bloom of phytoplankton (late May–early June), the variability of the pigment concentrations, the phytoplankton biomass and species compositions, and the hydrological conditions on the eastern shelf of Sakhalin Island was studied. The study resulted in revealing 135 microalgae species belonging to eight divisions. The most diversely presented were the Dinophyta dinoflagellates and Bacillariophyta diatoms (70 and 53 species, respectively). The concentration of chlorophyll a in the euphotic zone amounted, on average, to 3.8 mg/m³ in 1999 and 2.4 mg/m³ in 2000. It was shown that, in the northern and southern parts of the coastal zone, the concentration of chlorophyll a and the phytoplankton density in the spring were considerably different and depended on the hydrological conditions. In the north, their maximum values were found in the area of the depth break and were determined by the tidal mixing. The increased algae concentrations and temperature inversions at depths of 400–600 m confirm the downslope sliding of the near-bottom shelf waters. In the southern part, the high phytoplankton concentrations in the surface layer in 1999 confirmed by the monthly averaged estimates from the SeaWiFS satellite color scanner were caused by the abnormal northward propagation of the Soya Current waters and by intense tidal mixing.     

Seasonal pigment patterns of surface phytoplankton in the subtropical southern hemisphere

Pigment indices were used to characterise the seasonal succession of phytoplankton, and associated changes in chlorophyll a and accessory pigments, in subtropical waters of the three ocean basins in the southern hemisphere. Diagnostic indices revealed the dominance of small flagellates and elevated biomass during winter–spring in the Pacific, mixed flagellate–prokaryote communities and intermediate biomass during early summer in the Atlantic, and prokaryote dominance with low biomass in mid-summer in the Indian Ocean. Photo-pigment indices indicated only a small variation in the chlorophyll a proportion of total pigments across the ocean basins, but the accessory pigments varied considerably. Under low temperature and irradiance conditions, the photosynthetic carotenoids were prominent, but as temperatures and irradiance increased and nutrients declined, there was a significant increase in the proportion of photoprotective carotenoids. At high temperatures and irradiances, the photoprotective carotenoids were the largest component of the pigment pool, exceeding the proportion of chlorophyll a. These variations in phytoplankton composition, and their photoacclimation status, could be explained according to environmental changes and have implications for satellite estimations of biomass and primary production.     

Trophic Status of the South-Eastern Baltic Sea: A Comparison of Coastal and Open Areas

Primary production, nutrient concentrations, phytoplankton biomass (incl. chlorophyll a) and water transparency (Secchi depth), are important indicators of eutrophication. Earlier basin-wide primary production estimates for the Baltic Sea, a shallow shelf sea, were based mainly on open-sea data, neglecting the fundamentally different conditions in the large river plumes, which might have substantially higher production. Mean values of the period 1993–1997 of nutrient concentrations (phosphate, nitrate, ammonium and silicate), phytoplankton biomass, chlorophyll a (chl a) concentration, turbidity and primary production were calculated in the plumes of the rivers Oder, Vistula and Daugava and Klaipeda Strait as well as the open waters of the Arkona Sea, Bornholm Sea, eastern Gotland Sea and the Gulf of Riga. In the plumes, these values, except for primary production, were significantly higher than in the open waters. N:P ratios in the plumes were >16 (with some exceptions in summer and autumn), indicating potential P-limitation of phytoplankton growth, whereas they were <16 in the open Baltic Proper, indicating potential N-limitation. On the basis of in situ phytoplankton primary production, phytoplankton biomass and nutrient concentrations, the large river plumes and the Gulf of Riga could be characterized as eutrophic and the outer parts of the coastal waters and the open sea as mesotrophic. Using salinity to define the border of the plumes, their mean extension was calculated by means of a circulation model. Taking into account the contribution of coastal waters, the primary production in the Baltic Proper and the Gulf of Riga was 42·6 and 4·3×10⁶ t C yr⁻¹, respectively. Hence, an annual phytoplankton primary production in the whole Baltic Sea was estimated at 62×10⁶ t C yr⁻¹. The separate consideration of the plumes had only a minor effect on the estimation of total primary production in comparison with an estimate based on open sea data only. There is evidence for a doubling of primary production in the last two decades. Moreover, a replacement of diatoms by dinoflagellates during the spring bloom was noticed in the open sea but not in the coastal waters. A scheme for trophic classification of the Baltic Sea, based on phytoplankton primary production and biomass, chl a and nutrient concentrations, is proposed.