Population dynamics of phytoplankton, heterotrophic bacteria, and viruses during the spring bloom in the western subarctic Pacific

We characterized the community composition of phytoplankton in the western subarctic Pacific from the pre-bloom to the decline phase of the spring bloom with special reference to decreases in the silicic acid concentration in surface waters as an index for diatom bloom development. Furthermore, responses of heterotrophic bacteria and viruses to the spring bloom were also concomitantly investigated. Under pre-bloom conditions when nutrients were abundant but the surface mixed layer depth was relatively deep, chlorophyll (Chl) a concentrations were consistently low and green algae (chlorophytes and prasinophytes), cryptophytes, and diatoms were predominant in the phytoplankton assemblages as estimated by algal pigment signatures. Together with the shallowing of the mixed layer depth and the decrease in silicic acid concentration, diatoms bloomed remarkably in the Oyashio region, though the magnitude of the bloom in the Kuroshio-Oyashio transition (hereafter Transition) region was relatively small. A total of 77 diatom species were identified, with the bloom-forming diatoms mainly consisting of Thalassiosira, Chaetoceros, and Fragilariopsis species. It has become evident that the carotenoid fucoxanthin can serve as a strong indicator of the diatom carbon biomass during the spring diatom bloom. Differences in the species richness of diatoms among stations generally enabled us to separate the Oyashio bloom stations from the Transition and the Oyashio pre-bloom stations. Relatively high values of the Shannon-Wiener index for the diatom species were also maintained during the Oyashio bloom, indicating that a wide variety of species then shared dominance. In the decline phase of the Oyashio bloom when surface nutrient concentrations decreased, senescent diatom cells increased, as inferred from the levels of chlorophyllide a. Although the cell density of heterotrophic bacteria changed little with the development of the diatom bloom, viral abundance increased toward the end of the bloom, suggesting an increased likelihood of mortality among organisms including diatoms resulting from viral infection. This is the first report on the microbial trophodynamics, including viruses, during the spring diatom bloom in the western subarctic Pacific.     

Importance of intracellular Fe pools on growth of marine diatoms by using unialgal cultures and on the Oyashio region phytoplankton community during spring

We report on the ability for luxury Fe uptake and the potential for growth utilizing intracellular Fe pools for 4 coastal centric diatom isolates and in situ phytoplankton assemblages, mainly composed of diatoms. Iron uptake of the diatom isolates and natural phytoplankton assemblages in the Oyashio region during spring blooms were prevented by adding hydroxamate siderophore desferrioxamine B (DFB). After the addition of DFB, intracellular Fe in the diatom isolates supported 2.4–4.2 cell divisions with 1.2–2.6 Chl a doublings. The intracellular Fe was primarily used for cell generation rather than Chl a production, leading to a reduction in the Chl a cell quota in the Fe-starved cells with time. The metabolic properties of the Fe-starved cells with their cell morphologies were different among species or genera. An on-deck incubation experiment also exhibited 1.9 cell divisions and 0.81 Chl a doublings of phytoplankton after the addition of DFB, also indicating the preference of cell generation over Chl a production. A decrease in the level of cellular Chl a, a main light-harvesting pigment in Fe-starved diatoms, may become a superior survival strategy to protect the cells from high irradiance that can cause photo-oxidative damages through photosynthesis. Such relatively low-Fe with high-light conditions could often occur in surface waters of the Oyashio region from spring to summer.     

Community Structure and Dynamics of Phytoplankton in the Western Subarctic Pacific Ocean: A Synthesis

The phytoplankton community in the western subarctic Pacific (WSP) is composed mostly of pico- and nanophytoplankton. Chlorophyll a (Chl a) in the <2 μm size fraction accounted for more than half of the total Chl a in all seasons, with higher contributions of up to 75% of the total Chl a in summer and fall. The exception is the western boundary along the Kamchatka Peninsula and Kuril Islands and the Oyashio region where diatoms make up the majority of total Chl a during the spring bloom. Among the picophytoplankton, picoeukaryotes and Synechococcus are approximately equally abundant, but the former is more important in term of carbon biomass. Despite the lack of a clear seasonal variation in Chl a concentration, primary productivity showed a large seasonal variation, and was lowest in winter and highest in spring. Seasonal succession in the phytoplankton community is also evident with the abundance of diatoms peaking in May, followed by picoeukaryotes and Synechococcus in summer. The growth of phytoplankton (especially >10 μm cell size) in the western subarctic Pacific is often limited by iron bioavailability, and microzooplankton grazing keeps the standing stock of pico- and nano-phytoplankton low. Compared to the other HNLC regions (the eastern equatorial Pacific, the Southern Ocean, and the eastern subarctic Pacific), iron limitation in the Western Subarctic Gyre (WSG) may be less severe probably due to higher iron concentrations. The Oyashio region has similar physical condition, macronutrient supply and phytoplankton species compositions to the WSG, but much higher phytoplankton biomass and primary productivity. The difference between the Oyashio region and the WSG is also believed to be the results of difference in iron bioavailability in both regions. This revised version was published online in July 2006 with corrections to the Cover Date.     

Community structure and photosynthetic physiology of phytoplankton in the northwest subarctic Pacific during an in situ iron fertilization experiment (SEEDS-II)

Temporal changes in the abundance, community composition, and photosynthetic physiology of phytoplankton in surface waters were investigated during the second in situ iron (Fe) fertilization experiment in the NW subarctic Pacific (SEEDS-II). Surface chlorophyll a concentration was 0.75 mg m⁻³ on the day before the first Fe enrichment (i.e. Day 0), increased ca. 3-fold until Day 13 after two Fe additions, and thereafter declined with time. The photochemical quantum efficiency (F_v/F_m) and functional absorption cross-section (σPSII) of photosystem II for total phytoplankton in surface waters increased and decreased inside the Fe-enriched patch through Day 13, respectively. These results indicate that the photosynthetic physiological condition of the phytoplankton improved after the Fe infusions. However, the maximum F_v/F_m value of 0.43 and the maximum quantum yield of carbon fixation (φ_max) of 0.041 mol C (mol photon)⁻¹ during the development phase of the bloom were rather low, compared to their theoretical maximum of ca. 0.65 and 0.10 mol C (mol photon)⁻¹, respectively. Diatoms, which were mainly composed of oceanic species, did not bloom, and autotrophic nanoflagellates such as cryptophytes and prasinophytes became predominant in the phytoplankton community inside the Fe-enriched patch. In ferredoxin/flavodoxin assays for micro-sized (20–200 μm in cell length) diatoms, ferredoxin was not detected but flavodoxin expressions consistently occurred with similar levels both inside and outside the Fe-enriched patch, indicating that the large-sized diatoms were stressed by Fe bioavailability inside the Fe-enriched patch even after the Fe enrichments. Our data suggest that the absence of a Fe-induced large-sized diatom bloom could be partly due to their Fe stress throughout SEEDS-II.     

Seasonal changes in size-fractionated primary production and nutrient concentrations in the temperate neritic water of Funka Bay,Japan

Size-fractionated primary productivity and chlorophylla concentration were studied at two stations in the temperate neritic water of Funka Bay, Japan, from April 1984 to May 1985. Size distributions of phytoplankton were discussed in relation to nutrient availability. In the central part of the bay, 66% of the annual primary production occurred during the spring phytoplankton bloom with 95% of the spring production being accounted for by the greater than 10µm size fraction, which was dominated by diatoms. The increase in this large fraction was enhanced at both stations when nutrient concentrations increased in the bay's upper layer. Under low nutrient concentrations during summer, small phytoplankters (<2µm) accounted for 40 to 75% (average 60%) of the total¹⁴C uptake at the central station, and from 25 to 59% (average, 45%) at the coastal station. However, a sudden nutrient enrichment at the coastal station during the summer triggered the growth of the large size fraction. These seasonal and regional changes in total¹⁴C uptake were attributed to the large size fraction, composed mainly of diatoms. From the decreases in various nutrients during diatom blooms, it was further suggested that the predominance of diatoms was determined, not only by nutrient concentrations, but also by their relative availability.Contribution No. 205 from the Research Institute of North Pacific Fisheries, Faculty of Fisheries, Hokkaido University.     

长江口南部赤潮区春季表层沉积物中色素组成、含量与分布状况

为剖析长江口邻近海域春季硅藻藻华后期藻类沉降与底层水体缺氧现象之间的关系,作者于2011年春季,在长江口南部赤潮区采集了表层沉积物样品,并通过高效液相色谱法(HPLC),对浮游植物色素进行了分析。结果表明,硅藻藻华发生后,表层沉积物中叶绿素a(Chl a)、岩藻黄素(Fuco)和19’-丁酰氧基岩藻黄素(But-Fuco)含量有显著增加,高值区主要分布在调查海域东南侧50 m等深线外侧,与底层低氧水体分布区基本吻合。因此,硅藻藻华后沉降的藻类对于该海域夏季缺氧区的形成应具有一定作用,其具体过程和机制仍有待于研究。     

Low reproductive success for copepods during a bloom of the non‐aldehyde‐producing diatom Cerataulina pelagica in the North Adriatic Sea

Egg production rates and/or hatching success in the copepods Acartia clausi, Calanus helgolandicus and Temora longicornis were negatively affected by a late spring (May–June 2003) phytoplankton bloom in the North Adriatic Sea, dominated mainly by the large diatom Cerataulina pelagica. Highest total concentrations of 3.3·10⁴ cells·ml^?1 were located in the vicinity of the Po River, which also corresponded to the area where the highest numbers of phaeophorbides were measured (0.779, 0.528 and 0.419 μg·l^?1, respectively, compared to an average of the remaining stations of 0.183 ± 0.049 SD), suggesting some grazing on the bloom. Phytoplankton biomass in terms of carbon was dominated by diatoms, representing on average 42% of total phytoplankton carbon and more than 80% at several stations. Cerataulina pelagica, Cyclotella spp., Chaetoceros spp. and small unidentified centric diatoms dominated the diatom community numerically but C. pelagica was by far the dominant diatom in terms of carbon due to its large cell size. This species represented more than 60% of the diatom biomass at nine of the 14 stations sampled, and was absent only at one station, which was the most offshore station sampled during the cruise. Although polyunsaturated aldehydes (PUAs) were not detected, other oxylipins which are hydroxy and keto derivatives of eicosapentaenoic and docosahexaenoic acids that affect copepod reproduction were found in these samples. Hence, we can attribute the negative impact of diatoms not only to PUAs, as previously believed, but also to these compounds. This is the first direct evidence of the presence of oxylipins other than PUAs in marine blooms dominated by diatoms.     

Phytoplankton growth, microzooplankton herbivory and community structure in the southeast Bering Sea: insight into the formation and temporal persistence of an Emiliania huxleyi bloom

Using the seawater dilution technique, we measured phytoplankton growth and microzooplankton grazing rates within and outside of the 1999 Bering Sea coccolithophorid bloom. We found that reduced microzooplankton grazing mortality is a key component in the formation and temporal persistence of the Emiliania huxleyi bloom that continues to proliferate in the southeast Bering Sea. Total chlorophyll a (Chl a) at the study sites ranged from 0.40 to 4.45 μg C l⁻¹. Highest phytoplankton biomass was found within the bloom, which was a mixed assemblage of diatoms and E. huxleyi. Here, 75% of the Chl a came from cells >10 μm and was attributed primarily to the high abundance of the diatom Nitzschia spp. Nutrient-enhanced total phytoplankton growth rates averaged 0.53 d⁻¹ across all experimental stations. Average growth rates for >10 μm and <10 μm cells were nearly equal, while microzooplankton grazing varied among stations and size fractions. Grazing on phytoplankton cells >10 μm ranged from 0.19 to 1.14 d⁻¹. Grazing on cells <10 μm ranged from 0.02 to 1.07 d⁻¹, and was significantly higher at non-bloom (avg. 0.71 d⁻¹) than at bloom (avg. 0.14 d⁻¹) stations. Averaged across all stations, grazing by microzooplankton accounted for 110% and 81% of phytoplankton growth for >10 and <10 μm cells, respectively. These findings contradict the paradigm that microzooplankton are constrained to diets of nanophytoplankton and strongly suggests that their grazing capability extends beyond boundaries assumed by size-based models. Dinoflagellates and oligotrich ciliates dominated the microzooplankton community. Estimates of abundance and biomass for microzooplankton >10 μm were higher than previously reported for the region, ranging from 22,000 to 227,430 cells l⁻¹ and 18 to 164 μg C l⁻¹. Highest abundance and biomass occurred in the bloom and corresponded with increased abundance of the large ciliate Laboea, and the heterotrophic dinoflagellates Protoperidinium and Gyrodinium spp. Despite low grazing rates on phytoplankton <10 μm within the bloom, the abundance and biomass of small microzooplankton (<20 μm) capable of grazing E. huxleyi was relatively high at bloom stations. This body of evidence, coupled with observed high grazing rates on large phytoplankton cells, suggests the phytoplankton community composition was strongly regulated by herbivorous activity of microzooplankton. Because grazing behavior deviated from size-based model predictions and was not proportional to microzooplankton biomass, alternate mechanisms that dictate levels of grazing activity were in effect in the southeastern Bering Sea. We hypothesize that these mechanisms included morphological or chemical signaling between phytoplankton and micrograzers, which led to selective grazing pressure.     

Chlorophyll-specific absorption coefficients and pigments of phytoplankton off Sanriku, northwestern North Pacific

The variety in shape and magnitude of thein vivo chlorophyll-specific absorption spectra of phytoplankton was investigated in relation to differences in pigment composition off Sanriku, northwestern North Pacific. Site-to-site variations of the absorption coefficients,a _ph ^* (λ), and pigment composition were clearly observed. At warm-streamer stations, higher values ofa _ph ^* (440) anda _ph ^* (650) were found with relatively high concentrations of chlorophyllb (a green algae marker). At stations located in the Oyashio water (cold streamer),a _ph ^* (440) values were lower and fucoxanthin (a diatom marker) concentrations were higher, compared to the other stations. The peak in the absorption spectra at the Oyashio stations was shifted toward shorter wavelengths, which was probably due to the presence of phaeopigments. In a Kuroshio warm-core ring, the magnitude ofa _ph ^* (440) was in between those at the warm-streamer and Oyashio stations, and the diagnostic pigment was peridinin (a dinoflagellate marker). These findings indicated that major differences in phytoplankton absorption spectra of each water mass were a result of differences in the phytoplankton pigment composition of each water mass, which was probably related to the phytoplankton community.     

Responses of diatoms to iron-enrichment (SEEDS) in the western subarctic Pacific, temporal and spatial comparisons

Phytoplankton species composition was analyzed inside and outside of the iron-enriched patch during the SEEDS experiment. Before the iron-enrichment, the phytoplankton community consisted of similar proportions of pico-, nano- and micro-sized phytoplankton, and the micro-phytoplankton was dominated by the pennate diatom Pseudo-nitzschia turgidula. Although all the diatoms, except the nano-sized Fragilariopsis sp., increased during the two weeks of the observation period, the flora in the patch dramatically changed with the increase of phytoplankton biomass to a centric diatom-dominated community. Neritic diatoms, especially Chaetoceros debilis, showed higher growth rates than other diatoms, without any delay in the initiation of growth after the enrichment, and accounted for 90% of the micro-phytoplankton after day 9. In contrast, the oceanic diatoms showed distinct delays in the initiation of growth. We conclude that the responses of the diatoms to the manipulation of iron concentration were different by species, and the fast and intensive response of the phytoplankton to iron-enrichment resulted from the presence of a small amount of neritic diatoms at the study site. The important factors that determine the dominant species in the bloom are the potential growth rates under an iron-replete condition and the growth lag. Abundant species in the patch are widely distributed in the North Pacific and their relative contributions in the Oyashio area and at Stn KNOT are high from spring to summer. However, a characteristic difference of species composition between the SEEDS bloom and natural blooms was the lack of Thalassiosira and Coscinodiscus species in the patch, which usually account for a major part of the phytoplankton community under blooming conditions in the western North Pacific.     

Influence of light intensity on diatom physiology and nutrient dynamics in the Oyashio region

The spring diatom bloom characterizes the plankton and nutrient dynamics in the Oyashio region, the westernmost part of the subarctic Pacific. Previous studies have shown that NO₃ was not depleted during the spring bloom, and an increase in the consumption ratio of Si(OH)₄ to NO₃ (ΔSi(OH)₄:ΔNO₃) was observed as the spring bloom progressed. The increase in ΔSi(OH)₄:ΔNO₃ has been suggested to be caused by growth stresses of diatoms, e.g. light limitation by self-shading. In the present study, incubation experiments of sea-surface water from the Oyashio region under saturated irradiance showed that NO₃ was depleted first and ΔSi(OH)₄:ΔNO₃ was more or less constant until the NO₃ depletion occurred. The increase in ΔSi(OH)₄:ΔNO₃ was observed after the NO₃ depletion had occurred in contrast with the field observation. This result of the increase in ΔSi(OH)₄:ΔNO₃ under saturated irradiance after NO₃ depletion suggests that the in situ increase in ΔSi(OH)₄:ΔNO₃ before the NO₃ depletion might be caused by light limitation for diatoms. Responses to a reduction in irradiance were examined using diatom species isolated from the Oyashio region. Variable responses to a reduced irradiance were observed for cell specific C, N, Si and chlorophyll a (Chl) contents. However, the examined diatom species showed similar tendencies for increases in Si:C and Si:N and decreases in C:Chl ratios with the reduction in irradiance. We conclude that light limitation changes the uptake ratio of nutrients and the elemental composition of diatoms and that light limitation is one of the factors influencing the physiology of diatoms and nutrient dynamics in the Oyashio region during the spring bloom.     

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.     

DMSP and DMS dynamics and microzooplankton grazing in the Labrador Sea: application of the dilution technique

We adapted the dilution technique to study microzooplankton grazing of algal dimethylsulfoniopropionate (DMSP) vs. Chl a, and to estimate the impact of microzooplankton grazing on dimethyl sulfide (DMS) production in the Labrador Sea. Phytoplankton numbers were dominated by autotrophic nanoflagellates in the Labrador basin, but diatoms and colonial Phaeocystis pouchetii contributed significantly to phytomass at several high chlorophyll stations and on the Newfoundland and Greenland shelfs. Throughout the region, growth of algal Chl a and DMSP was generally high (0.2–1 d⁻¹), but grazing rates were lower and more variable, characteristic of the early spring bloom period. Production and consumption of Chl a vs. DMSP followed no clear pattern, and sometimes diverged greatly, likely because of their differing distributions among algal prey taxa and size class. In several experiments where Phaeocystis was abundant, we observed DMS production proportional to grazing rate, and we found clear evidence of DMS production by this haptophyte following physical stress such as sparging or filtration. It is possible that grazing-activated DMSP cleavage by Phaeocystis contributes to grazer deterrence: protozoa and copepods apparently avoided healthy colonies (as judged by relative growth and grazing rates of Chl a and DMSP), and grazing of Phaeocystis was significant only at one station where cells were in poor condition. Although we hoped to examine selective grazing on or against DMSP-containing algal prey, the dilution technique cannot differentiate selective ingestion and varying digestion rates of Chl a and DMSP. We also found that the dilution method alone was poorly suited for assessing the impact of grazing on dissolved sulfur pools, because of rapid microbial consumption and the artifactual release of DMSP and DMS during filtration. Measuring and understanding the many processes affecting organosulfur cycling by the microbial food web in natural populations remain a technical challenge that will likely require a combination of techniques to address.     

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.     

Estimation of standing stock of chlorophylla and primary production from remote-sensed ocean color in the Oyashio region, the Western subarctic Pacific, during the spring bloom in 1997

In order to examine the applicability of remotely-sensed ocean color for the estimation of phytoplankton biomass and primary production in the Oyashio region, the western subarctic Pacific, vertical distributions of chlorophylla concentration and primary production were observed in April and May 1997. Spring bloom was observed in both April and May, and the surface concentration of chlorophylla exceeded 40 mg m⁻³. The relationship between the standing stocks of chlorophylla within the layer from the sea surface to one optical depth (0–1/k layer) and the surface chlorophylla concentration is expressed as a Michaelis-Menten equation. The mean ratio of the standing stock of chlorophylla in the euphotic layer to that in the 0–1/k layer was 4.41, this ratio did not significantly differ from 4.61 which was obtained at homogeneous distribution of chlorophylla within the euphotic layer. These facts suggest that the distribution of chlorophylla could be assumed to be homogeneous in the euphotic layer during the spring bloom. Results of primary production measurements by simulatedin situ method were compared with those by an algorithm with two variables; chlorphylla and non-spectral PAR. Daily primary production in the euphotic layer estimated by the algorithm varied in a range of 38–274% of that estimated by incubation, although the primary productions by the algorithm agreed with those by the incubation at a half of stations. Primary production within the euphotic layer calculated using simply the surface data was the same as that estimated using vertical distribution of chlorophylla. These results show that the primary production in the euphotic layer may be estimated from the remote sensed measurements during the spring bloom in the Oyashio region.     

Spatial variability of the species composition,abundance, and productivity of the phytoplankton in the white sea in the late summer period

The species composition, cell concentration (N), and biomass (B) of the phytoplankton, as well as the chlorophyll a (Chl a) concentration, primary production (PP), and the concentrations of the dissolved inorganic micronutrients (phosphorus, silica, nitrogen as nitrite), were estimated for Kandalaksha Bay (KB), Dvina Bay (DB), and the basin (Bas) of the White Sea in August of 2004. The micronutrient concentrations were lower compared to the average long-term values for the summer period. The Chl a concentration varies from 0.9 to 2.0 mg/m³ for most of the studied areas, reaching up to 7.5 mg/m³ in the Northern Dvina River estuary. The surface water layer of the DB was the most productive area, where the PP reached up to 270–375 mg C/(m³ day). The phytoplankton biomass varied from 11 to 205 mg C/m³ with the highest values observed in the Bas and DB. Three groups of stations were defined during the analysis of the phytoplankton’s species composition similarity. The dinoflagellates Dinophysis norvegica and Ceratium fusus were particular to the phytoplankton assemblages in the KB; the diatom Ditylum brightwellii was particular to the upper and central parts of the DB. These three phytoplankton species were less abundant in the Bas.     

Nutrient dynamics and seasonal succession of phytoplankton assemblages in a Southern European Estuary: Ria de Aveiro, Portugal

The spatial and temporal dynamics of dissolved inorganic nitrogen, dissolved phosphate, dissolved silica and chlorophyll a were measured seasonally at eight stations in the Ria de Aveiro. Between December 2000 and September 2001, the seasonal succession of phytoplankton assemblages, inferred after the spatial and seasonal variation of silica and of chlorophyll a concentrations, showed that diatoms (μmol Si L⁻¹) dominated from late autumn until early spring, while chlorophytes (μg Chl a L⁻¹) bloomed during late spring and summer. The Si:N:P ratios and Si concentrations indicated no seasonal depletion in dissolved silica, as in other temperate systems, possibly because of abnormal precipitation and flood events prolonging the supply of dissolved Si to the system. The Si:N:P ratios suggested P limitation at the system level. Despite the relative proportions of available nutrients, the mean phosphorus concentration (5.3 μmol L⁻¹) was above the reported half-saturation constants for P uptake by phytoplankton. Thus, in Ria de Aveiro, the seasonal succession of phytoplankton assemblages may also be dependent on the grazing capacity of the pelagic community through top-down regulation.     

Role of dissolved silicate in the occurrence of a phytoplankton bloom

The spring bloom of phytoplankton was studied in March in Funka Bay, Japan, to test the Tsunogai (1979)'s hypothesis regarding the role of silicate in the bloom. The hypothesis comprises two parts. 1) Diatoms are predominant when all the physical and chemical conditions are adequate for plankton growth. 2) Since the Si:P ratio of the diatom body is usually much larger than that of sea water, flagellates (non-siliceous phytoplankton) replace diatoms after dissolved silicate in the sea water has been almost completely consumed by diatoms. At the end of the bloom in late March phosphate still remained in the water but silicate was exhausted and the main species of phytoplankton changed from diatoms to flagellates. Grazing pressure by zooplankton at this time was not so great. A model using the data on assimilation rates of silicate showed a dramatic change of silicate uptake in late March. Poison in scallops caused byProtogonyaulux sp. (dinoflagellates) rapidly increased from mid-April at all stations along the coast of Funka Bay. All of these findings support Tsunogai's hypothesis.     

Grazing impact of the copepod community in the Oyashio region of the western subarctic Pacific Ocean

The role of copepod grazing on the ecosystem dynamics in the Oyashio region, western subarctic Pacific was investigated during six cruises from June 2001 to June 2002. In situ grazing rates of the copepod community (CGR) were measured by the gut fluorescence method in respect to developmental stages of dominant species. In terms of biomass, more than 80% of the copepod community was dominated by six large calanoid species (Neocalanus cristatus, Neocalanus flemingeri, Neocalanus plumchrus, Eucalanus bungii, Metridia pacifica and Metridia okhotensis) throughout the year. Resulting from the observed pattern of the interzonal migrating copepods, the CGR in the Oyashio region was divided into three phases, i.e. spring (bloom), summer (post-bloom) and autumn-winter phase. During the spring bloom, late copepodites of the interzonal migrating species, N. cristatus, N. flemingeri and E. bungii appeared in the surface layer (0-50 m) to consume the production of the bloom, resulting in a high grazing rate of the copepod community (7.9 mg Chl m⁻² d⁻¹), though its impact on phytoplankton community was low due to the high primary productivity. During the post-bloom period, although the copepod community which was dominated by N. cristatus, N. plumchrus, M. pacifica and newly recruited E. bungii still maintained a high biomass, the CGR was generally lower (1.8-2.6 mg Chl m⁻² d⁻¹ for June and August 2001), probably due to the lower availability of phytoplankton. Nevertheless, the highest CGR was also observed during this period (10.5 mg Chl m⁻² d⁻¹ in June 2002). The high CGR on autotrophic carbon accounted for 69% of the primary production, suggesting that the copepod community in the Oyashio region potentially terminates the phytoplankton bloom. Abundant occurrence of young E. bungii, which is a characteristic phenomenon in the Oyashio region, was largely responsible for the high grazing pressure in June 2002 suggesting that success of reproduction, growth, and survival in E. bungii during the spring bloom is an important factor in controlling phytoplankton abundance during the post-bloom season. During autumn and winter, CGR was the lowest in the year (0.29-0.38 mg Chl. m⁻² d⁻¹) due to the disappearance of the interzonal migrating copepods from the surface layer. Diel migrant M. pacifica was the most important grazer during this period. The annual ingestion of the copepod community is estimated as 37.7 gC m⁻² on autotrophic carbon (converted using C:Chl ratio of 30) or 137.9 gC m⁻² on suspended particles (using C:Chl ratio of in situ value, 58-191), accounting for 13% and 46% of annual primary production, respectively. This study confirms that copepod grazing is an important pathway in carbon flow in the Oyashio region and in particular their role in the phytoplankton dynamics is significant for the termination of the spring bloom.     

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.