Consequences of winter upwelling events on biogeochemical and phytoplankton patterns in a western Galician ria (NW Iberian peninsula)

The consequences of two upwelling events in mid- (MW) and late (LW) winter on biogeochemical and phytoplankton patterns were studied in the Pontevedra Ria and compared with the patterns measured under typical winter conditions and under a summer upwelling event. Thermohaline patterns measured during the mid-winter upwelling event (MW-up) revealed the intrusion of saltier seawater (35.9) into the ria associated with the Iberian Poleward Current (IPC). During the late-winter upwelling event (LW-up), the seawater which had welled up into the ria showed characteristics of the Eastern North Atlantic Central Water mass (ENACW). In both cases the measured water residence time (4 days during MW-up and 10 days during LW-up) was related to both meteorological and fluvial forcing. This residence time contrasts with that of summer upwelling (7 days) and with that estimated under unfavorable upwelling atmospheric conditions (2–4 weeks). During MW-up, the ria became poor in nutrients due to continental freshwater dilution, associated with the shorter residence time of the water, and the intrusion of IPC, which is a water body poor in nutrient salts: 2.9 μM of nitrate, 0.1 μM of phosphate and 1.5 μM of silicate. During this event, the ria exported 3.4 mol_DIN s⁻¹, compared with 6.9 mol_DIN s⁻¹ in non-upwelling conditions. Phytoplankton showed a uniform distribution throughout the ria, as during unfavorable upwelling conditions, and was characterized by the dominance of diatoms, mainly Nitzschia longissima and Skeletonema costatum. During LW-up, a nutrient depletion in the photic layer also occurred, but as a result of a phytoplankton spring bloom developing at this time. The ria was a nutrient trap where 4.1 mol_DIN s⁻¹ were processed by photosynthesis. This budget is three times higher than the one under non-upwelling conditions. In contrast with the MW-up, which had no effect on primary production, during LW-up the ria became more productive, although not as productive as during a summer upwelling event (9.9 mol_DIN s⁻¹). The taxonomic composition of the phytoplankton community did not change noticeably during LW-up and the summer upwelling, with the same species present and changing only in relative proportions. Diatoms were always the dominant microphytoplankton community, with Pseudonitzschia pungens, Thalassionema nitzschioides and several species of Chaetoceros as characteristic taxons.     

Influence of upwelling and river runoff interaction on phytoplankton assemblages in a Middle Galician Ria and Comparison with northern and southern rias (NW Iberian Peninsula)

The first oceanographic research (hydrography, nutrient salts, chlorophyll, primary production and phytoplankton assemblages) in a Middle Galician Ria was carried out in Corme-Laxe during 2001, just a year before the Prestige oil spill, being the only reference to evaluate eventual changes in the phytoplankton community. Due to the small size of this ria (6.5 km²), oceanographic processes were driven by the continental water supplied by Anllons River during the wet season (20–30 m³ s⁻¹ in winter), and the strong oceanic influence from the nearby shelf during the dry season. The annual cycle showed a spring bloom with high levels of chlorophyll (up to 14 μg Chl-a L⁻¹) and primary production (3 g C m⁻² d⁻¹) and a summer upwelling bloom (up to 8 μg Chl-a L⁻¹ and 10 g C m⁻² d⁻¹) where the proximity of the Galician upwelling core (<13.5 °C at sea surface) favors the input of upwelled seawater (up to 9 μM of nitrate and silicate) to the bottom ria layer, even during summer stratification events (primary production around 2 g C m⁻² d⁻¹). Thus, phytoplankton assemblages form a “continuum” from spring to autumn with a predominance of diatoms and overlapping species between consecutive periods; only in autumn dinoflagellates and flagellates characterized the phytoplankton community. In the Middle Rias as Corme-Laxe, the nutrient values, Chl-a, primary production and phytoplankton abundance for productive periods were higher than those reported for the Northern (Ria of A Coruña) and Southern Rias (Ria of Arousa) for year 2001; this suggests the importance of the hydrographic events occurring in the zone of maximum upwelling intensity of the Western Iberian Shelf, where a lack of annual cycles studies exists.     

Microzooplankton feeding impact in a coastal upwelling system on the NW Iberian margin: The Ría de Vigo

The dilution technique, combined with identification and enumeration of pico-, nano- and micro-plankton by microscopy, was used to estimate microzooplankton impact on the microbial community in surface waters of a coastal embayment on the NW Iberian upwelling system. Microzooplankton were important consumers of autotrophic and heterotrophic plankton in this system, feeding up to 93% of standing stock and more than 100% of production of several groups. Heterotrophic bacteria and heterotrophic picoflagellates experienced the highest and constant impact, with 75–84% of their standing stocks and 85–102% of their production being channelled through the microbial food web. Pico- and nano-phytoplankton were also consumed, although maximum grazing occurred on diatoms during upwelling events, coinciding with highest primary production. Predation on pico-nano-heterotrophs was especially relevant under downwelling conditions, when consumption of total carbon and particularly autotrophic carbon was considerably lower than during upwelling. The results suggest that the existence of a multivorous food web, extending from the microbial loop to the herbivorous food web, could be a major feature in this coastal upwelling system. The microbial loop, which occurs as a permanent background in the system, would contribute to sustain the microbial food web during downwelling, whereas the herbivorous food web could coexist with a microbial food web based on large diatoms during upwelling. The multivorous food web would partially divert diatoms from sinking and hence favour the retention of organic matter in the water column. This could enhance the energy transfer to higher pelagic trophic levels in coastal upwelling systems.     

The reduction of plankton biomass induced by mesoscale stirring: A modeling study in the Benguela upwelling

Recent studies, both based on remote sensed data and coupled models, showed a reduction of biological productivity due to vigorous horizontal stirring in upwelling areas. In order to better understand this phenomenon, we consider a system of oceanic flow from the Benguela area coupled with a simple biogeochemical model of Nutrient-Phyto-Zooplankton (NPZ) type. For the flow three different surface velocity fields are considered: one derived from satellite altimetry data, and the other two from a regional numerical model at two different spatial resolutions. We compute horizontal particle dispersion in terms of Lyapunov exponents, and analyzed their correlations with phytoplankton concentrations. Our modeling approach confirms that in the south Benguela there is a reduction of biological activity when stirring is increased. Two-dimensional offshore advection and latitudinal difference in primary production, also mediated by the flow, seem to be the dominant processes involved. We estimate that mesoscale processes are responsible for 30–50% of the offshore fluxes of biological tracers. In the northern area, other factors not taken into account in our simulation are influencing the ecosystem. We suggest explanations for these results in the context of studies performed in other eastern boundary upwelling areas.     

2012年夏季楚科奇海浮游动物垂直迁移及上升流对其生物量的影响

The diel vertical migration(DVM) of zooplankton and the influence of upwelling on zooplankton biomass were examined using water column data of current velocity and mean volume backscattering strength(MVBS)collected by moored acoustic Doppler current profilers(ADCPs) deployed in the southeastern Chukchi Sea during the 5th Chinese National Arctic Research Expedition(CHINARE) in summer 2012, combined with the satellite observational data such as sea surface temperature(SST), wind, and chlorophyll a(Chl a). Hourly acoustic data were continuously collected for 49-d in the mooring site. Spectral analysis indicated that there were different migrating patterns of zooplankton, even though precisely classifying the zooplankton taxa was not available. The prevailing 24-h cycle corresponded to the normal DVM with zooplankton swimming upwards at sunrise and returning to deep waters at sunset. There was a clear DVM in the upper 17 m of the water column during the period with distinct day-night cycles, and no active DVM throughout the water column when the sun above the horizon(polar day), suggesting that light intensity was the trigger for DVM. Also there was a second migrating pattern with 12-h cycle. The upwelling event occurring in the northwest of Alaskan coastal area had important influence on zooplankton biomass at the mooring site. During the upwelling, the SST close to the mooring site dropped significantly from maximal 6.35°C to minimal 1.31°C within five days. Simultaneously, there was a rapid increase in the MVBS and Chl a level, suggesting the aggregation of zooplankton related to upwelling.     

Hydrographic and atmospheric analysis of an autumnal upwelling event in the Ria of Vigo (NW Iberian Peninsula)

An autumnal upwelling event was observed in the Ria of Vigo (NW Iberian Peninsula) on 15th November 2001. This event was analyzed by means of thermohaline variables measured at CTD stations located in the study area, satellite sea surface temperature and wind data provided by QuikSCAT. Salinity and temperature distributions revealed that the upwelled water mass was Eastern North Atlantic Central Water (ENACW), typically observed during summer upwelling events. However, previous to the upwelling event, the characteristic autumnal body of water was recorded on 31st October 2001. Subsequent sea surface temperature and upwelling index corroborated the presence of an autumnal upwelling event of approximately 15 days duration. The probability of upwelling-favorable winds (from 1999 to 2004) was lower during autumn–winter than during spring–summer, although they may occur at any time of the year. Probabilities of 45% were calculated for February and November, with the highest probability (65%) corresponding to July.     

春季长江口富营养化水域浮游植物群集对冲淡水和上升流的响应

A comprehensive study on the phytoplankton standing stocks, species composition and dominant species in the eutrophic Changjiang(Yangtze River) Estuary(CE) was conducted to reveal the response of phytoplankton assemblage to Changjiang Diluted Water(CDW) and upwelling in the spring. Phytoplankton presented peak standing stocks(13.03 μg/L of chlorophyll a, 984.5×103 cells/L of phytoplankton abundance) along the surface isohaline of 25. Sixty-six species in 41 genera of Bacillariophyta and 33 species in 19 genera of Pyrrophyta were identified, as well as 5 species in Chlorophyta and Chrysophyta. Karenia mikimotoi was the most dominant species, followed by Prorocentrum dentatum, Paralia sulcata, Pseudo-nitzschia delicatissima and Skeletonema costatum. A bloom of K. mikimotoi was observed in the stratified stations, where the water was characterized by low nitrate, low phosphate, low turbidity, and specific ranges of temperature(18–22 °C) and salinity(27–32). K.mikimotoi and P. dentatum accumulated densely in the upper layers along the isohaline of 25. S. costatum was distributed in the west of the isohaline of 20. Benthonic P. sulcata presented high abundance near the bottom,while spread upward at upwelling stations. CDW resulted in overt gradients of salinity, turbidity and nutritional condition, determining the spatial distribution of phytoplankton species. The restricted upwelling resulted in the upward transport of P. sulcata and exclusion of S. costatum, K. mikimotoi and P. dentatum. The results suggested that CDW and upwelling were of importance in regulating the structure and distribution of phytoplankton assemblage in the CE and the East China Sea.     

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.     

Diversity and distribution of bacterioplankton in the coastal upwelling waters off Hainan Island,China

The diversity, community composition and 16 S rRNA gene abundance of bacterioplankton along a transect across an upwelling area off the eastern coast of Hainan Island（the Qiongdong upwelling） were investigated in August of 2016 using high throughput sequencing and quantitative PCR assay of 16 S rRNA genes. Compared with the offshore stations, the inner-shelf stations had higher bacterial gene abundance（up to 3 fold） and operational taxonomic unit richness, a result of the influence of upwelled a...     

Seasonal and spatial distributions of phytoplankton biomass associated with monsoon and oceanic environments in the South China Sea

Seasonal variations of phytoplankton/chlorophyll-a （Chl-a） distribution, sea surface wind, sea height anomaly, sea surface temperature and other oceanic environments for long periods are analyzed in the South China Sea （SCS）, especially in the two typical regions off the east coast of Vietnam and off the northwest coast of Luzon, using remote sensing data and other oceanographic data. The results show that seasonal and spatial distributions of phytoplankton biomass in the SCS are primarily influenced by the monsoon winds and oceanic environments. Off the east coast of Vietnam, Chl-a concentration is a peak in August, a jet shape extending into the interior SCS, which is associated with strong southwesterly monsoon winds, the coastal upwetling induced by offshore Ekman transport and the strong offshore current in the western SCS. In December, high Chl-a concentration appears in the upwelling region off the northwest coast of Luzon and spreads southwestward. Strong mixing by the strong northeasterly monsoon winds, the cyclonic circulation, southwestward coastal currents and river discharge have impacts on distribution of phytoplankton, so that the high phytoplankton biomass extends from the coastal areas over the northern SCS to the entire SCS in winter. These research activities could be important for revealing spatial and temporal patterns of phytoplankton and their interactions with physical environments in the SCS.     

A numerical study on the role of wind forcing,bottom topography,and nonhydrostacy in coastal upwelling

The responses of coastal upwelling to different magnitudes of wind stress over a narrow and a wide shelf are studied using a 3-D primitive equation numerical model. The results show that the position of the upwelling front depends on both the strength and the duration of the wind forcing. The comparison between different shelf widths shows that wide shelf will limit the cold water intrusion, so that the corresponding decrease in sea surface temperature is less compared to narrow shelves. Besides, the difference between hydrostatic and nonhydrostatic model results shows that nonhydrostatic effects will enhance the growth of surface meandering, and can be more pronounced near steep fronts. Although difference does exist, our results show that the nonhydrostatic effects are very small at least in this idealized study case.     

Basin-scale variability of phytoplankton biomass,production and growth in the Atlantic Ocean

The latitudinal distributions of phytoplankton biomass, composition and production in the Atlantic Ocean were determined along a 10,000-km transect from 50°N to 50°S in October 1995, May 1996 and October 1996. Highest levels of euphotic layer-integrated chlorophyll a (Chl a) concentration (75–125 mg Chl m⁻²) were found in North Atlantic temperate waters and in the upwelling region off NW Africa, whereas typical Chl a concentrations in oligotrophic waters ranged from 20 to 40 mg Chl m⁻². The estimated concentration of surface phytoplankton carbon (C) biomass was 5–15 mg C m⁻² in the oligotrophic regions and increased over 40 mg C m⁻² in richer areas. The deep chlorophyll maximum did not seem to constitute a biomass or productivity maximum, but resulted mainly from an increase in the Chl a to C ratio and represented a relatively small contribution to total integrated productivity. Primary production rates varied from 50 mg C m⁻² d⁻¹ at the central gyres to 500–1000 mg C m⁻² d⁻¹ in upwelling and higher latitude regions, where faster growth rates (μ) of phytoplankton (>0.5 d⁻¹) were also measured. In oligotrophic waters, microalgal growth was consistently slow [surface μ averaged 0.21±0.02 d⁻¹ (mean±SE)], representing <20% of maximum expected growth. These results argue against the view that the subtropical gyres are characterized by high phytoplankton turnover rates. The latitudinal variations in μ were inversely correlated to the changes in the depth of the nitracline and positively correlated to those of the integrated nitrate concentration, supporting the case for the role of nutrients in controlling the large-scale distribution of phytoplankton growth rates. We observed a large degree of temporal variability in the phytoplankton dynamics in the oligotrophic regions: productivity and growth rates varied in excess of 8-fold, whereas microalgal biomass remained relatively constant. The observed spatial and temporal variability in the biomass specific rate of photosynthesis is at least three times larger than currently assumed in most satellite-based models of global productivity.     

南海北部东沙天然气水合物区浮游植物生物量和生产力粒级结构特征及其环境影响分析

本文讨论了2013年5月南海东沙天然气水合物区浮游植物生物量和生产力粒级结构特征及其环境影响因素。结果表明,研究海域表现出典型的低营养盐、低叶绿素a、低生产力特征,浮游植物叶绿素a和初级生产力具有明显的次表层最大值现象。东沙海域生物量和初级生产力粒级结构差异性显著,从生物量和生产力贡献度来看,表现为微微型浮游植物> 微型浮游植物> 小型浮游植物。生物量的垂直分布结果表明,春季不同粒级类群浮游植物在真光层内的分布存在明显不同,比如小型浮游植物在真光层内分布较均匀;微型浮游植物则主要分布于近表层或真光层中部,而微微型浮游植物则主要分布于真光层中部和底部。微微型浮游植物在纬度较低的热带贫营养海区之所以能够占主导优势,最主要的原因是其极小的细胞体积和较大的表面积使其有利于营养竞争。相关性分析表明,南海东沙浮游植物各粒级生物量与温度、pH显著正相关,与硅酸盐、磷酸盐显著负相关;浮游植物各粒级生产力与温度显著正相关,与盐度、磷酸盐显著负相关。磷酸盐含量是影响东沙海域浮游植物粒级结构差异的重要因素之一,同时,光辐照度和水体的真光层深度对东沙天然气水合物区不同粒径浮游植物的垂直分布起着更为重要的调控作用。     

Vertical distribution of phytoplankton biomass,production and growth in the Atlantic subtropical gyres

Ninety-four stations were sampled in the Atlantic subtropical gyres during 10 cruises carried out between 1995 and 2001, mainly in boreal spring and autumn. Chlorophyll a (Chl-a) and primary production were measured during all cruises, and phytoplankton biomass was estimated in part of them. Picoplankton (<2 μm) represented >60% of total Chl-a concentration measured at the surface, and their contribution to this variable increased with depth. Phytoplankton carbon concentrations were higher in the upper metres of the water column, whereas Chl-a showed a deep maximum (DCM). At each station, the water column was divided into the upper mixed layer (ML) and the DCM layer (DCML). The boundary between the two layers was calculated as the depth where Chl-a concentration was 50% of the maximum Chl-a concentration. On average DCML extends from 67 to 126 m depth. Carbon to Chl-a (C:Chl-a) ratios were used to estimate phytoplankton carbon content from Chl-a in order to obtain a large phytoplankton carbon dataset. Total C:Chl-a ratios averaged (±s.e.) 103±7 (n=22) in the ML and 24±4 (n=12) in the DCML and were higher in larger cells than in picoplankton. Using these ratios and primary production measurements, we derived mean specific growth rates of 0.17±0.01 d⁻¹ (n=173) in the ML and 0.20±0.01 d⁻¹ (n=165) in the DCML although the differences were not significant (t-test, p>0.05). Our results suggest a moderate contribution of the DCML (43%) to both phytoplankton biomass and primary production in the Atlantic subtropical gyres.     

Seasonal distribution of primary production,phytoplankton biomass and size distribution in the Greenland Sea

The study establishes an annual estimate for annual primary production of 81 g C m⁻² for the open Greenland Sea based on data from five cruises and literature data. This estimate agrees well with a model estimate based on nutrient utilisation but is a factor of 2–5 less than published primary production estimates made by remote sensing of this area. The seasonal distribution of particulate primary production in open Greenland Sea waters followed the seasonal distribution of surface irradiance with a peak in June, indicating that light is the primary factor governing primary production in the area. At stations along the ice edge, blooms were recorded in both June and August, suggesting a pattern of repeated blooms during the summer season at the ice edge. Subsurface phytoplankton peaks were a persistent feature in the open Greenland Sea from May to August. These peaks were consisted of actively photosynthesising phytoplankton and up to 90% of total water column particulate primary production was estimated to occur in association with these peaks. Diatoms dominated the phytoplankton community during the spring bloom and in the Polar Water during August. Size distribution analyses of the phytoplankton communities indicated that the relative abundance of large cells compared to small cells was greatest in May as compared to June and August. No significant differences were noted between June and August in the slope of the phytoplankton size distribution spectra. Inorganic nitrogen and phosphorus nutrients were measurable in surface waters on all cruises. Only in August were there some indications (altered Redfield ratios and higher nutrient concentrations in subsurface chlorophyll peaks than at the surface) of nutrient depletion of surface waters. Implications for food web structure and carbon flux of these patterns in phytoplankton activity and distribution are discussed.     

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.