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.     

Iron deficiency in micro-sized diatoms in the Oyashio region of the Western subarctic Pacific during spring

In order to detect iron (Fe) stress in micro-sized (20–200 μm) diatoms in the Oyashio region, western subarctic Pacific during spring, immunological ferredoxin/flavodoxin assays were applied to samples collected from the surface layer in May 2005. Concomitantly, the community composition of the micro-sized phytoplankton and hydrographic conditions, including dissolved Fe and macronutrient concentrations, were also examined. Chlorophyll (Chl) a concentrations were <2 mg m⁻³ at all sampling stations, except at a station where the Chl a level was 9.0 mg m⁻³ and a micro-sized diatom bloom occurred. A high abundance of ferredoxin in micro-sized diatoms was detected only at a rather near-shore station where dissolved Fe and macronutrient concentrations were higher, indicating that the micro-sized diatoms did not suffer from iron deficiency. On the other hand, flavodoxin in micro-sized diatoms was often observed at the other stations, including the bloom station, where macronutrients were replete but dissolved Fe concentration was low (0.31 nM). A significant amount of chlorophyllide a, a degradation product of Chl a, was also observed at the bloom station, suggesting a decline of the diatom bloom. The micro-sized phytoplankton species at all the stations were mainly composed of the diatoms Thalassiosira, Chaetoceros, and Fragilariopsis spp. Our study indicates that micro-sized diatoms were stressed by Fe bioavailability during the spring season in the Oyashio region     

Carbon to nitrogen (C:N) stoichiometry of the spring–summer phytoplankton bloom in the North Water Polynya (NOW)

The carbon to nitrogen (C:N) stoichiometry of phytoplankton production varied significantly during the spring–summer bloom in the North Water Polynya (NOW), from April through July 1998. The molar ratio of particulate organic carbon (POC) to nitrogen (PON) production by phytoplankton (ΔPOC:ΔPON) increased from 5.8 during April through early June to 8.9 in late June and July. The molar dissolved inorganic carbon (DIC) to nitrate+nitrite (NO₃) drawdown ratio (ΔDIC: ΔNO₃) increased from 6.7 in April and May, to 11.9 in June (no estimate for July because of ice melting). The discrepancy between ΔPOC:ΔPON and ΔDIC:ΔNO₃ was likely due to dissolved organic carbon (DOC) production. Increased ΔPOC:ΔPON of phytoplankton and surface water ΔDIC:ΔNO₃ throughout the phytoplankton blooms resulted from changes in physical properties of the upper water column, such as reduced thickness of the surface mixed layer that exposed phytoplankton to increased photosynthetically available radiation (PAR), accompanied by NO₃ depletion. This is expected to have significant effects on the cycling of carbon (C) and nitrogen (N) in pelagic ecosystems, as the increased C:N ratio of organic matter decreases its quality as substrate for grazers and microbial communities. Based on ΔPOC:ΔPON, the ratio of POC to chlorophyll a (Chl) production (ΔPOC:ΔChl) and the relationship between Chl yields and NO₃ depletion, we estimate that 71±17% and 46±20% of the depleted NO₃ went to PON production in the euphotic zone over the polynya from April to early June, and late June to July, respectively. The remaining NO₃ was likely channelled to dissolved organic nitrogen (DON) and heterotrophic bacteria, which were not returned to the dissolved inorganic nitrogen (DIN) pool through recycling during the course of the study. Hence, the autotrophic production of organic N and its recycling by the microbial food web were not coupled temporally.     

Simulation of upper-ocean biogeochemistry with a flexible-composition phytoplankton model: C, N and Si cycling in the western Sargasso Sea

We report the first application of a biogeochemical model in which the major elemental composition of the phytoplankton is flexible, and responds to changing light and nutrient conditions. The model includes two phytoplankton groups: diatoms and non-siliceous picoplankton. Both fix C in accordance with photosynthesis-irradiance relationships used in other models and take up NO₃⁻ and NH₄⁺ (and Si(OH)₄ for diatoms) following Michaelis-Menten kinetics. The model allows for light dependence of photosynthesis and NO₃⁻ uptake, and for the observed near-total light independence of NH₄⁺ uptake and Si(OH)₄ uptake. It tracks the resulting C/N ratios of both phytoplankton groups and Si/N ratio of diatoms, and permits uptake of C, N and Si to proceed independently of one another when those ratios are close to those of nutrient-replete phytoplankton. When the C/N or Si/N ratio of either phytoplankton group indicates that its growth is limited by N, Si or light, uptake of non-limiting elements is controlled by the content of the limiting element in accordance with the cell-quota formulation of Droop (J. Mar. Biol. Ass. U.K 54 (1974) 825).We applied this model to the Bermuda Atlantic Time-series Study (BATS) site in the western Sargasso Sea. The model was tuned to produce vertical profiles and time courses of [NO₃⁻], [NH₄⁺] and [Si(OH)₄] that are consistent with the data, by adjusting the kinetic parameters for N and Si uptake and the rate of nitrification. The model then reproduces the observed time courses of chlorophyll-a, particulate organic carbon and nitrogen, biogenic silica, primary productivity, biogenic silica production and POC export with no further tuning. Simulated C/N and Si/N ratios of the phytoplankton indicate that N is the main growth-limiting nutrient throughout the thermally stratified period and that [Si(OH)₄], although always limiting to the rate of Si uptake by diatoms, seldom limits their growth rate. The model requires significant nitrification in the upper 200 m to yield realistic time courses and vertical profiles of [NH₄⁺] and [NO₃⁻], suggesting that NO₃⁻ is not supplied to the upper water column entirely by physical processes. A nitrification-corrected f-ratio (f_NC), calculated for the upper 200 m as: (NO₃⁻ uptake—nitrification)/(NO₃⁻ uptake+NH₄⁺ uptake) has annual values ranging from only 0.05–0.09, implying that 90–95% of the N taken up annually by phytoplankton is supplied by biological regeneration (including nitrification) in the upper 200 m. Reported discrepancies between estimates of organic C export based on seasonal chemical changes and POC export measured at the BATS site can be almost completely resolved if there is significant regeneration of NO₃⁻ via organic-matter decomposition in the upper 200 m.     

A nitrate and silicate budget in the equatorial Pacific Ocean: a coupled physical–biological model study

A coupled physical–biological model was developed to simulate the low-silicate, high-nitrate, and low-chlorophyll (LSHNLC) conditions in the equatorial Pacific Ocean and used to compute a detailed budget in the Wyrtki box (5°N–5°S, 180–90°W) for the major sources and cycling of nitrogen and silicon in the equatorial Pacific. With the incorporation of biogenic silicon dissolution, NH₄ regeneration from organic nitrogen and nitrification of ammonia in the model, we show that silicon recycling in the upper ocean is less efficient than nitrogen. As the major source of nutrients to the equatorial Pacific, the Equatorial Undercurrent provides slightly less Si(OH)₄ than NO₃ to the upwelling zone, which is defined as 2.5°N–2.5°S. As a result, the equatorial upwelling supplies less Si(OH)₄ than NO₃ into the euphotic zone in the Wyrtki box, having a Si/N supply ratio of about 0.85 (2.5 vs. 2.96 mmolm⁻² day⁻¹). More Si(OH)₄ than NO₃ is taken up with a Si/N ratio of 1.17 (2.72 vs. 2.33 mmolm⁻² day⁻¹) within the euphotic zone. The difference between upwelling supply and biological uptake is balanced by nutrient regeneration and horizontal advection. Excluding regeneration, the net silicate and nitrate uptakes are nearly equal (1.76 vs. 1.84 mmolm⁻² day⁻¹). However, biogenic silica export production is slightly higher than organic nitrogen (1.74 vs. 1.59 mmolm⁻² day⁻¹) following a 1.1 Si/N ratio. In the central equatorial Pacific, low silicate concentrations limit diatom growth; therefore non-diatom new production accounts for most of the new production. Higher silicate supply in the east maintains elevated diatom growth rates and new production associated with diatoms dominate upwelling zone. In contrast, the new production associated with small phytoplankton is nearly constant or decreases eastward along the equator. The total new production has a higher rate in the east than in the west, following the pattern of surface silicate. This suggests that silicate regulates the diatom production, total new production, and thereby carbon cycle in this area. The modeled mean primary production is 48.4 mmolCm⁻² day⁻¹, representing the lower end of direct field measurements, while new production is 15.0 mmolCm⁻² day⁻¹, which compares well with previous estimates.     

An Ecosystem Model Coupled with Nitrogen-Silicon-Carbon Cycles Applied to Station A7 in the Northwestern Pacific

A model based on that of Kishi et al. (2001) has been extended to 15 compartments including silicon and carbon cycles. This model was applied to Station A7 off Hokkaido, Japan, in the Northwestern Pacific. The model successfully simulated the observations of: 1. a spring bloom of diatoms; 2. large seasonal variations of nitrate and silicate concentrations in the surface water; and 3. large inter-annual variations in chlorophyll-a. It also reproduced the observed features of the seasonal variations of carbon dioxide partial pressure (pCO₂)—a peak in pCO₂ in winter resulting from deep winter convection, a rapid decrease in pCO₂ as a result of the spring bloom, and an almost constant pCO₂ from summer through fall (when the effect of increasing temperature cancels the effect of biological production). A comparison of cases with and without silicate limitation shows that including silicate limitation in the model results in: 1. decreased production by diatoms during summer; and 2. a transition in the dominant phytoplankton species, from diatoms to other species that do not take up silicate. Both of these phenomena are observed at Station A7, and our results support the hypothesis that they are caused by silicate limitation of diatom growth. This revised version was published online in July 2006 with corrections to the Cover Date.     

Selective depressions of surface silicic acid within cyclonic mesoscale eddies in the oligotrophic western North Pacific

To reveal spatial dynamics of silicic acid [Si(OH)₄] in the poorly sampled oligotrophic western North Pacific, we investigated the surface distribution of Si(OH)₄ and associated biogeochemical parameters by using an underway survey system with a highly sensitive nutrient analyzer along the 138°E transect (between 30 and 34°N) and the 155°E transect (between 10 and 35°N) during the summers of 2007 and 2008. Surface Si(OH)₄ concentrations ranged from the detection limit (11 nmol L⁻¹) to 2462 nmol L⁻¹. High Si(OH)₄ concentrations (>1000 nmol L⁻¹) and dynamic fluctuations were generally observed north of 23°N, while consistently stable low concentrations of 415–751 nmol L⁻¹ were observed south of 23°N. Surface nitrate+nitrite (N+N) and phosphate (PO₄³⁻) were typically depleted to <20 nmol L⁻¹, except for PO₄³⁻ in the area south of 16°N. The majority of the study area was characterized by high-Si(OH)₄ and low-N+N and PO₄³⁻. However, submesoscale/mesoscale depressions of Si(OH)₄ were locally observed in the cyclonic eddy fields north of 23°N. Among a total of six Si(OH)₄ depressions within the eddies, a complete Si(OH)₄ depletion (<11 nmol L⁻¹) was observed on the cyclonic side near the Kuroshio axis (33.1°N, 138°E). This depletion was closely coupled with a diatom bloom, suggesting that Si(OH)₄ was exhausted by diatoms. All of the Si(OH)₄ depressions were selective and not accompanied by local depressions of N+N and PO₄³⁻. This unique phenomenon might be driven by biogeochemical processes such as selective Si export (Si pump), anomalous Si uptake associated with diatom physiology, and/or Si uptake supported by N₂ fixation.     

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.     

Utility of an automatic water sampler to observe seasonal variability in nutrients and DIC in the Northwestern North Pacific

Concentrations of nutrients (NO₃, NO₂, Si(OH)₄, PO₄ and dissolved inorganic carbon (DIC) were measured in a series of seawater samples collected over approximately 15 months in 2005 and 2006 by an automatic water sampler (Remote Access Sampler, RAS) in the Northwestern North Pacific. Seasonal variability and concentrations of NO₃ + NO₂ (NO_x and Si(OH)₄ were comparable to previous shipboard observations, although there were small errors associated with measurements of PO₄ and DIC. Concentrations of these nutrients began to decrease in late April. After the end of June, NO_x and Si(OH)₄ decreased rapidly, with large fluctuations. After October, these nutrients increased again until late spring 2006. The ratio of the decrease of Si(OH)₄ to that of NO_x suggests that numbers of biogenic opal-producing creatures, such as diatoms, increased after the end of June. This conclusion was supported by a rapid increase in biogenic opal flux recorded in a sediment trap at 150 m. The relationship between NO_x concentrations at the RAS depth of 35 m and NO_x integrated over the upper 100 m was determined using previous shipboard hydrocast data. This relationship was used to estimate integrated mixed layer NO_x concentration from RAS data. Estimated new production based on seasonal drawdown of integrated NO_x averaged approximately 156 mg-C m⁻²day⁻¹ annually, which agrees with previous estimates. Thus, an automatic seawater sampler that documents annual maximum and minimum nutrient concentrations and episodic events such as storms and spring blooms, which might be missed by an ordinary research vessel, will contribute to time-series observations of nutrients and, by extension, biological pump activity.     

Seasonal dynamics of inorganic nutrients and phytoplankton biomass in the NW Alboran Sea

The seasonal dynamics of inorganic nutrients and phytoplankton biomass (chlorophyll a), and its relation with hydrological features, was studied in the NW Alboran Sea during four cruises conducted in February, April, July and October 2002. In the upper layers, the seasonal pattern of nutrient concentrations and their molar ratios (N:Si:P) was greatly influenced by hydrological conditions. The higher nutrient concentrations were observed during the spring cruise (2.54 μM NO₃⁻, 0.21 μM PO₄³⁻ and 1.55 μM Si(OH)₄, on average), coinciding with the increase of salinity due to upwelling induced by westerlies. The lowest nutrient concentrations were observed during summer (<0.54 μM NO₃⁻, 0.13 μM PO₄³⁻ and 0.75 μM Si(OH)₄, on average), when the lower salinities were detected. Nutrient molar ratios (N:Si:P) followed the same seasonal pattern as nutrient distribution. During all the cruises, the ratio N:P in the top 20 m was lower than 16:1, indicating a NO₃⁻ deficiency relative to PO₄³⁻. The N:P ratio increased with depth, reaching values higher than 16:1 in the deeper layers (200–300 m). The N:Si ratio in the top 20 m was lower than 1:1, excepting during spring when N:Si ratios higher than 1:1 were observed in some stations due to the upwelling event. The N:Si ratio increased with depth, showing a maximum at 50–100 m (>1.5:1), which indicates a shift towards Si-deficiency in these layers. The Si:P ratio was much lower than 16:1 throughout the water column during the four cruises. In general, the spatial and seasonal variation of phytoplankton biomass showed a strong coupling with hydrological and chemical fields. The higher chlorophyll a concentrations at the depth of the chlorophyll maximum were found in April (2.57 mg m⁻³ on average), while the lowest phytoplankton biomass corresponded to the winter cruise (0.74 mg m⁻³ on average). The low nitrate concentrations together with the low N:P ratios found in the upper layers (top 20 m) during the winter, summer and autumn cruises suggest that N-limitation could occur in these layers during great part of the year. However, N-limitation during the spring cruise was temporally overcome by nutrient enrichment caused by an intense wind-driven upwelling event.     

Silicon dynamics within an intense open-ocean diatom bloom in the Pacific sector of the Southern Ocean

An intense diatom bloom developed within a strong meridional silicic acid gradient across the Antarctic Polar Front at 61°S, 170°W following stratification of the water column in late October/early November 1997. The region of high diatom biomass and the silicic acid gradient propogated southward across the Seasonal Ice Zone through time, with the maximum diatom biomass tracking the center of the silicic acid gradient. High diatom biomass and high rates of silica production persisted within the silicic acid gradient until the end of January 1998 (ca. 70 d) driving the gradient over 500 km to the south of its original position at the Polar Front. The bloom consumed 30 to >40 μM Si(OH)₄ in the euphotic zone between about 60 and 66°S leaving near surface concentrations <2.5 μM and occasionally <1.0 μM in its wake. Integrated biogenic silica concentrations within the bloom averaged 410 mmol Si m⁻² (range 162–793 mmol Si m⁻²). Average integrated silica production on two consecutive cruises in December 1997 and January 1998 that sampled the bloom while it was well developed were 27.5±6.9 and 22.6±20 mmol Si m⁻² d⁻¹, respectively. Those levels of siliceous biomass and silica production are similar in magnitude to those reported for ice-edge diatom blooms in the Ross Sea, Antarctica, which is considered to be among the most productive regions in the Southern Ocean. Net silica production (production minus dissolution) in surface waters during the bloom was 16–21 mmol Si m⁻² d⁻¹, which is sufficient for diatom growth to be the cause of the southward displacement of the silicic acid gradient. A strong seasonal change in silica dissolution : silica production rate ratios was observed. Integrated silica dissolution rates in the upper 100–150 m during the low biomass period before stratification averaged 64% of integrated production. During the bloom integrated dissolution rates averaged only 23% of integrated silica production, making 77% of the opal produced available for export to depth. The bloom ended in late January apparently due to a mixing event. Dissolution : production rate ratios increased to an average of 0.67 during that period indicating a return to a predominantly regenerative system.Our observations indicate that high diatom biomass and high silica production rates previously observed in the marginal seas around Antarctica also occur in the deep ocean near the Polar Front. The bloom we observed propagated across the latitudinal band overlying the sedimentary opal belt which encircles most of Antarctica implying a role for such blooms in the formation of those sediments. Comparison of our surface silica production rates with new estimates of opal accumulation rates in the abyssal sediments of the Southern Ocean, which have been corrected for sediment focusing, indicate a burial efficiency of 4.6% for biogenic silica. That efficiency is considerably lower than previous estimates for the Southern Ocean.     

Factors controlling the timing of major spring bloom events in an UK south coast estuary

Factors controlling the timing of major (>10 mg chlorophyll a m⁻³) spring bloom events in the estuarine waters of the Solent, on the south coast of the UK, have been investigated. Winter to summer variations in chlorophyll a concentration together with relevant meteorological and hydrographical data have been analysed for 5 years (1988, 1992, 2001, 2002 and 2003). Mean water column irradiance is demonstrated to be the main factor controlling the timing of the first major spring bloom event, usually dominated by large chain-forming diatoms. When chlorophyll a concentration first exceeds 10 mg m⁻³ in spring (usually in May) the mean water column photosynthetic active radiation (PAR) averaged for one week prior to the sampling date was always >380 W h m⁻² d⁻¹. Prior to the main spring bloom event surface incident radiation and water turbidity combine to limit chlorophyll a concentration to levels <10 mg m⁻³. Chlorophyll a concentrations >10 mg m⁻³ do not occur in the Solent until almost the entire 10 m water column is within the euphotic zone (i.e. above 1% light level) and light extinction coefficient (k) is ca. ≤0.5 m⁻¹. Statistically, river flow explains the largest percentage of the variations in k and the delayed bloom in June 2002 is due to increased cloud cover and high levels of rainfall in May, which caused a reduction in surface incident irradiance and increased turbidity. Chlorophyll a peaks during these major bloom events generally occur on spring tides when increased mixing rates favour net growth of diatoms.     

Seasonal and interannual variability in phytoplankton and nutrient dynamics along Line P in the NE subarctic Pacific

We analysed mixed-layer seasonal and interannual variability in phytoplankton biomass and macronutrient (NO₃ and Si(OH)₄) concentrations from three decades of observations, and nitrogen uptake rates from the 1990s along Line P in the NE subarctic Pacific. Chlorophyll a concentrations near 0.35 mg m⁻³ were observed year-round along Line P except at the nearshore station (P4) where chlorophyll a concentrations in spring were on average 2.4 times the winter values. In contrast, the temporal variability in carbon-to-chlorophyll ratios at the two main end members of Line P (P4 and OSP) was high. Large seasonal and interannual variability in NO₃ and Si(OH)₄ concentration were observed along Line P. Highest upper mixed-layer (top 15 m) nutrient concentrations occurred on the continental shelf in late summer and early fall due to seasonal coastal upwelling. Beyond the shelf, maximum nutrient concentrations increased gradually offshore, and were highest in late winter and early spring due to mixing by winter storms. Interannual variations in upper mixed-layer nutrient concentrations beyond the shelf (>128°W) were correlated with E-W winds and the PDO since 1988 but were not correlated with either climate index between 1973 and 1981. Despite differences in nutrient concentration, nutrient utilization (ΔNO₃ and ΔSi(OH)₄) during the growing season were about 7.5 μM at all offshore stations. Variations in ΔNO₃ were correlated with those of ΔSi(OH)₄. The annual cycle of absolute NO₃ uptake (ρNO₃) and NH₄ uptake (ρNH₄) rates by phytoplankton in the upper mixed-layer showed a weak increasing trend from winter to spring/summer for the period 1992-1997. Rates were more variable at the nearshore station (P4). Rates of ρNO₃ were low along the entire line despite abundant NO₃ and low iron (Fe), at the offshore portion of Line P and sufficient Fe at the nearshore station (P4). As a result, new production contributed on average to only 32 ± 15% of the total nitrogen (N) uptake along Line P. NO₃ utilization in the NE subarctic Pacific is probably controlled by a combination of environmental variables, including Fe, light and ambient NH₄ levels. Elevated ambient NH₄ concentrations seem to decrease the rates of new production (and f-ratios) in surface waters of the oceanic subarctic NE Pacific. Contrary to expectation, phytoplankton biomass, nutrient utilization (ΔNO₃ and ΔSi(OH)₄), and nitrogen uptake (ρNO₃ + ρNH₄) varied relatively little along Line P, despite significant differences in the factors controlling phytoplankton composition assemblages and production. Future studies would benefit from including other variables, especially light limitation, to improve our understanding of the seasonal and interannual variability in phytoplankton biomass and nutrients in this region.     

The 1980/1981 phytoplankton cycle in the coastal waters off Connemara,Ireland

The results of a phytoplankton survey conducted in coastal waters off western Ireland in 1980/1981 are reported. Surface values of temperature, salinity, NO₃N, PO₄P, Si, total N, total P and chlorophyll a (Chla) collected at 6 stations during 14 cruises are presented along with the species composition of the net phytoplankton.The spring bloom occurred in late April and was dominated by diatoms. Between April and July further diatom blooms occurred. In July and August dinoflagellates were dominant. This change was associated with the stratification of offshore water and low concentrations of Chla and nutrients. In Autumn large blooms of Ceratium tripos were found; it is suggested that the cause was heavy freshwater runoff. In autumn and winter some warm water oceanic species (e.g. Oxytoxum scolopax) occurred.     

Nutrient and carbon removal ratios and fluxes in the Ross Sea, Antarctica

Net community production (NCP) and nutrient deficits (Def(X)) were calculated using decreases in dissolved CO₂ and nutrient concentrations due to biological removal in the upper 200 m of the water column during four cruises in the Ross Sea, Antarctica along 76°30′S in 1996 and 1997. A comparison to excess dissolved and particulate organic carbon showed close agreement between surplus total organic carbon (TOC) and NCP during bloom initiation and productivity maximum; however, when TOC values had returned to low wintertime values NCP was still significantly above zero. This seasonal NCP, 3.9±1 mol C m⁻², must be equivalent to the particle export to depths greater than 200 m over the whole productive season. We estimate that the annual export was 55±22% of the seasonal maximum in NCP. The fraction of the seasonal maximum NCP that is exported through 200 m is significantly higher than that measured by moored sediment traps at a depth of 206 m. The removal of carbon, nitrate and phosphate (based on nutrient disappearance since early spring) and their ratios showed significant differences between regions dominated by diatoms and regions dominated by the haptophyte Phaeocystis antarctica. While the ΔC/ΔN removal ratio was similar (7.8±0.2 for diatoms and 7.2±0.1 for P. antarctica), the ΔN/ΔP and ΔC/ΔP removal ratios for diatoms (10.1±0.3 and 80.5±2.3) were significantly smaller than those of P. antarctica (18.6±0.4 and 134.0±4.7). The similarity in ΔC/ΔN removal ratios of the two assemblages suggests that preferential uptake of phosphate by diatoms caused the dramatic differences in ΔC/ΔP and ΔN/ΔP removal ratios. In contrast to low ΔC/ΔP and ΔN/ΔP removal ratio in diatom-dominated areas early in the growing season, deficit N/P and C/P ratios in late autumn indicate that the elemental stochiometry of exported organic matter did not deviate significantly from traditional Redfield ratios. Changes in biologically utilized nutrient and carbon ratios over the course of the growing season indicated either a substantial remineralization of phosphate or a decrease in phosphate removal relative to carbon and total inorganic nitrogen over the bloom period. The species dependence in C/P ratios, and the relative constancy in the C/N ratios, makes N a better proxy of biological utilization of CO₂.     

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.     

长江口及邻近海区营养盐结构与限制

通过研究长江口及邻近海域溶解无机氮(DIN=NO₃-+NO₂-+NH₄+)、磷酸盐(PO₄3-)、硅酸盐(SiO₃2-)所表征的营养盐区域结构特征及影响因素,在分析营养盐绝对限制情况的基础上,划分了潜在相对营养限制区域。结果表明,123°E以西近岸表层区域DIN/P比值全年均高于16,而Si/DIN除秋季外基本小于1,显示出长江冲淡水影响下"过量氮"的特征。春夏季河口锋面区(31°~32.5°N,122.5°~124°E)硅藻的大量生长可使DIN/P异常升高和Si/DIN异常降低。秋季研究区域北部DIN/P西低东高且Si/DIN西高东低是由于在高DIN、低PO₄3-的长江冲淡水影响下,近岸受相对低DIN、高SiO₃2-的苏北沿岸流南下入侵影响而被分割而成。冬季长江口门东北部存在的高DIN/P和低Si/DIN区则主要由于寡营养盐的黑潮水深入陆架,向东北输送的部分长江冲淡水和增强的苏北沿岸流共同作用造成DIN升高所致。利用Redfield比值进行了不同站位表层潜在相对营养限制情况的区分。近岸123°E以西受高DIN、SiO₃2-长江冲淡水影响,四季多呈现PO₄3-潜在相对限制,而在春夏季由于浮游植物的大量吸收PO₄3-,造成局部PO₄3-绝对限制及潜在相对限制。春夏季氮限(DIN潜在相对限制)一般发生在外海部分站位,但较为零散。秋季除了东南外海大部分站位外,受苏北沿岸流影响在长江口北部近岸也存在氮限。随着低DIN/P的黑潮表层水(KSW)的入侵加强,冬季外海氮限站位增多。硅限(SiO₃2-潜在相对限制)在夏季发生在赤潮高发区,而冬季南部存在较多硅限站位表明KSW中SiO₃2-相对较为缺乏。     

Effect of nutrient amendments on the summer phytoplankton dynamics on the Bering Sea shelf under experimental conditions

Experiments on nutrient and iron amendments were performed with phytoplankton on the eastern shelf of the Bering Sea in June 2000 and August 2001. The nutrient amendments (NO₃, NH₄, SiO₄, NO₃ + SiO₄, NH₄ + SiO₄, and Fe + NO₃) increasing their initial concentrations by ～20 μM were put into test bottles 10 l in volume each. With iron addition (Fe or Fe + NO₃), its concentration increased by 5 nM. The experiments performed showed that the main nutrient that limited the phytoplankton development was nitrogen. Regardless of the composition of the dominant algae in the background community, the amendments caused massive development of diatoms. The intense growth was characteristic for diatoms of both the spring and spring-summer assemblages. At high abundances of Phaeocystis pouchetii or of the coccolithophore Emiliania huxleyi in the natural water, nitrogen-containing amendments caused an intense growth of these species, along with the massive development of diatoms. In the case of the diatom prevalence in the initial sample, the intensities of the utilization of NO₃ and NH₄ in combination with SiO₄ in the course of the experiment were 1.7 and 3 times as high as their intensities with no silicon amendments. Likewise, NO₃ + SiO₄ and NH₄ + SiO₄ mixed amendments caused an increase in the silicon assimilation by a factor of 4–5 as compared to pure silicon amendments. During one of the experimental series in which both diatoms and Phaeocystis pouchetii actively developed, virtually complete nitrogen utilization (90–99.8%) in 4–5 days was observed for both the NO₃ and NH₄. The addition of silicon and iron only caused no significant growth of the phytoplankton abundance. It was assumed that the destruction of the seasonal thermocline and the supply of nutrients into the surface layer as a result of strong wind forcing might cause a phytoplankton bloom in the summer time and result in the much pronounced qualitative and quantitative spatial heterogeneity of the phytoplankton characteristic of the eastern shelf of the Bering Sea.     

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.     

Nitrogen uptake kinetics in the Ross Sea, Antarctica

The first estimates of uptake kinetic parameters for NH₄⁺, NO₃⁻, and urea in the Ross Sea, Antarctica were measured on three cruises during austral late winter–early spring 1996 (pre-bloom), late spring 1997 (bloom development), and summer 1997 (bloom decline). Nitrogen (N) uptake experiments were conducted with water collected at the 50% light penetration depth using trace-metal clean protocols and ¹⁵N tracer techniques. At all sites, ambient NO₃⁻ concentrations ranged from 5.8 to 30.5 μg-at N l⁻¹ and silicic acid concentrations were greater than 62.0 μg-at Si l⁻¹. The following trends were observed. First, based on maximum uptake rates (V_max), apparent N utilization followed the order NO₃⁻>NH₄⁺>urea during the pre-bloom and bloom development cruises. During the summer cruise, as the bloom was declining, the apparent order of utilization was NH₄⁺>NO₃⁻>urea. Second, evidence for possible repression of NO₃⁻ uptake by elevated NH₄⁺ concentrations was only observed at one site. Third, the kinetic parameters of NH₄⁺ uptake rates corrected for isotope dilution were compared with the kinetic parameters determined from uncorrected rates. In this comparison, the measure of substrate affinity, α (α=V_max/K_s) increased by an average of 4.6-fold when rates were corrected for isotope dilution, but values of V_max remained unchanged. Fourth, using bacterial production data, the magnitude of bacterial N uptake was estimated. Assuming that all bacterial N demands were met with NH₄⁺, the estimated bacterial portion of NH₄⁺ uptake ranged from <1%, when the ratio of bacteria to autotrophic biomass was low, to 35%, when bacterial abundance and biomass were highest. Finally, dramatic changes in NH₄⁺ uptake capacity were observed at one station (Stn. O), where kinetic parameters were measured during all three cruises. We hypothesize that a mutualistic relationship exists between phytoplankton and heterotrophic bacteria, and that the creation of microzones of high NH₄⁺ concentrations contributed to the changes seen at this station.