Variabilité spatio-temporelle de la biomasse microphytobenthique en rade de Brest et en Manche occidentale

Microphytobenthic biomass was estimated in the Bay of Brest (1994) and the western English Channel (Trezen Vraz, 1993) using spectrophotometry. Best results (42 % difference) were obtained with an instantaneous extraction procedure at room temperature, compared with the cold extraction procedure. Chlorophyll biomass in sediment was higher in the western English Channel (65–215 mg Chl a m⁻²) than in the Bay of Brest (10–113 mg Chl a m⁻²), in contrast to total pigment biomass (Chl a + pheo a : 88–254 mg m⁻² at Trezen Vraz vs. 131–934 mg m⁻² in the Bay of Brest). This study emphasized decreasing pigment biomass from the estuary to the open sea. Low benthic pigment biomass in the Bay, compared to eutrophic ecosystems, agrees with previous studies on the pelagic system, suggesting that there is no eutrophication in the Bay of Brest. Microphytobenthic blooms were strongly correlated with environmental factors, such as river floods, light intensity, water temperature and the spring/neap tidal cycle. Turbidity and light could be limiting factors at Roscanvel, i.e. the deepest site facing gyre circulation. The latter would favour sedimentation of silt particles at the sediment-water interface. Microphytobenthos population dynamics would be also dependent on the spring/neap tidal cycle, since maximum chlorophyll biomass occurred at neap tides at all three sites in the Bay of Brest, in contrast to the western English Channel. Lower temporal variations of pigment biomass in sediment were observed in the Channel, compared with the Bay of Brest; maximum chlorophyll biomass in sediment was found to occur three to four weeks after the phytoplankton blooms.     

Phytoplankton productivity in the western subarctic gyre of the North Pacific in early summer 2006

We deployed a profiling buoy system incorporating a fast repetition rate fluorometer in the western subarctic Pacific and carried out time-series observations of phytoplankton productivity from 9 June to 15 July 2006. The chlorophyll a (Chl a) biomass integrated over the euphotic layer was as high as 45–50 mg Chl a m⁻² in the middle of June and remained in the 30–40 mg Chl a m⁻² range during the rest of observation period; day-to-day variation in Chl a biomass was relatively small. The daily net primary productivity integrated over the euphotic layer ranged from 144 to 919 mg C m⁻² day⁻¹ and varied greatly, depending more on insolation rather than Chl a biomass. In addition, we found that part of primary production was exported to a 150-m depth within 2 days, indicating that the variations in primary productivity quickly influenced the organic carbon flux from the upper ocean. Our results suggest that the short-term variability in primary productivity is one of the key factors controlling the carbon cycle in the surface ocean in the western subarctic Pacific.     

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.     

Microbial biomass response to different quantities and sources of organic matter in Brazilian coastal sediments

This study investigates the benthic microbial responses to organic matter (OM) variations in quantity and sources in two shallow water bays (Fortaleza and Ubatuba Bays) on the SE coast of Brazil on six occasions during the year. The pelagic and benthic compartments of the bays were evaluated by: (i) nutrients and chlorophyll a (Chl a) in the water column; (ii) quantity and sources of OM in the sediment (Chl a, total organic carbon and total nitrogen and lipid biomarker composition); and (iii) microbial biomass in sediments as an indicator of active benthic response. Although there were changes in water‐column nutrients during the year, Chl a was fairly constant, suggesting a regular supply of microalgae‐derived OM to the sea bottom. Based on the composition of lipid biomarkers in sediments, OM sources were classified as mostly marine and with high contributions of labile (microalgae‐derived) OM. Labile OM composition varied from diatoms in the summer to phytoflagellates in the winter and tended to accumulate in areas protected by physical disturbances in one of the bays. Microbial biomass followed this trend and was 160% higher in protected than in exposed areas. This study suggests that the coupling between labile OM and benthic microbial biomass occurs primarily in protected areas, irrespective of the time of the year. Since meio‐ and macrofaunal assemblages depend upon secondary microbial production within the sediments, this coupling may have an important role for the benthic food‐web.     

Saldanha Bay,South Africa I: the use of ocean colour remote sensing to assess phytoplankton biomass

The efficacy of ocean colour remote sensing in assessing the variability of phytoplankton biomass within Saldanha Bay is examined. Satellite estimates of chlorophyll a (Chl a) were obtained using the maximum peak-height (MPH) algorithm on full-resolution (300?m) data from the Medium Resolution Imaging Spectrometer (MERIS). Subsurface Chl a maxima often occur within Saldanha Bay below the mean detection depth of the satellite (1.5?m) during periods of thermal stratification. Consequently, the MPH product was poorly correlated to in situ data from 4?m depth (r² and average relative percentage difference [RPD] of 0.094 and 53% respectively); however, the coefficient of determination was much improved if limited to in situ data collected under conditions of mixing (r² and RPD of 0.869 and 89%, respectively). Composites of monthly MPH Chl a data reveal mean concentrations consistent with in situ seasonal trends of phytoplankton biomass, confirming the capability of the MPH algorithm to qualitatively resolve surface Chl a distribution within the bay.     

Phytoplankton pigment change as a photoadaptive response to light variation caused by tidal cycle in Ariake Bay,Japan

Underwater light environment and photosynthetic accessory pigments were investigated in Ariake Bay in order to understand how change of the pigments occurs in response to the tidal-induced changes in underwater light conditions. We hypothesize that phytoplankton increases photo-protective pigments and decreases light-harvesting pigments under higher light condition in the mixed layer caused by tidal cycle. Contribution rates of non-phytoplankton particles (a _nph (400–700)) for light attenuation coefficient (K _d ) was highest (32–85%), and those of phytoplankton particles (a _ph (400–700)), dissolved organic matter (a _g (400–700)) and water were 6–32, 6–21 and 5–23%, respectively. Mean K _d was higher during the spring tide (0.55 ± 0.23 m⁻¹) than the neap tide (0.44 ± 0.16 m⁻¹), and the K _d difference was caused by the substances resuspension due to the tidal current. In contrast, ratios of photo-protective pigments (diadinoxanthin and diatoxanthin) per chlorophyll a ((DD+DT)/Chl a) were higher during the neap tide (0.10 ± 0.03 mg mg-Chl a ⁻¹) than the spring tide (0.08 ± 0.03 mg mg-Chl a ⁻¹). And there was significant positive correlation between (DD+DT)/Chl a and mean relative PAR in the mixed layer ($ \overline {I_{mix} } $ \overline {I_{mix} } ). Moreover, there was significant negative correlation between ratios of light-harvesting pigments (fucoxanthin) per Chl a (Fuco/Chl a) and $ \overline {I_{mix} } $ \overline {I_{mix} } . These results suggested that phytoplankton in Ariake Bay increase photo-protective pigments and decrease light-harvesting pigments in the higher light condition of less turbid, shallower mixed layer during neap tide than spring tide.     

Spatial variability of the primary production and chlorophyll <Emphasis Type="Italic">a</Emphasis> concentration in the drake passage in the austral spring

The spatial distribution of the primary production (PP) and the chlorophyll a concentration (Chl) were investigated during two research cruises in the Drake Passage area in October–November of 2007 and 2008. The algorithm evaluating the integral PP (PP_int) for the water column in this area was developed based on the data on the surface chlorophyll (Chl_s) and the incident solar irradiance obtained in 2004–2008 in the Atlantic Sector of the Southern Ocean. The results obtained both by the experimental and model approaches suggested that the Polar Front (PF) region of the Drake Passage was characterized by low values of both the PP_int (<100 mg C/m² per day) and Chl_s (0.08–0.20 mg/m³) in October–November. Low values of the Chl_s and relatively high phaeophytine a concentrations indicated the winter succession state of the phytoplankton community in the Antarctic Ocean and the southern Polar Frontal Zone (PFZ). The seasonal warming of the surface water layers and the developing pycnocline resulted in a phytoplankton bloom and a Chl_s concentration of more than 1 mg/m³ in mid-November in this area and the Subantarctic waters.     

Comparative study on microphytobenthic pigments of muddy and sandy intertidal sediments of the Tagus estuary

The abundance and distribution of microphytobenthic pigments determined by HPLC (chlorophylls and carotenoids) were compared between muddy and sandy sediments of the Tagus estuary (Portugal). In the two types of sediment, with similar periods of illuminated emersion, chlorophyll a concentrations on a per area basis (mg m⁻²) were comparable (down to 2 mm). Pigment analysis also revealed similar microphytobenthic communities in terms of algal classes. Diatoms were the dominant microalgae, but cyanophytes, euglenophytes and phanerogam debris were also present. For both muddy and sandy sediments, microphytobenthic biomass showed a high level of variability both within and between two consecutive years. Microphytobenthos was highly stratified in the mud, with most of the chlorophyll a occurring in the top 500 μm. In the sand, relatively constant concentrations were found throughout the sediment profile down to 3 mm. This is probably related to deeper light penetration in sandy sediment and/or increased physical mixing caused by invertebrate activity or overlying currents, leading to the burial of an important fraction of the microphytobenthic cells. Differences observed in the intensity of sediment coloration of muddy and sandy sediments might have resulted from the different vertical distribution of benthic biomass.     

Dynamics of microphytobenthic biomass in a coastal area of western Seto Inland Sea, Japan

This study focused on the causes of the variation in microphytobenthic biomass and the effects of this variation on macrobenthic animals in the western Seto Inland Sea, Japan, where the importance of microphytobenthos as the primary food source for benthic animals has been recently reported. We investigated the microphytobenthic biomass together with light attenuation of seawater, phytoplanktonic biomass, macrobenthic density and biomass at eight stations (water depth = 5–15 m) during four cruises in 1999–2000. The increased light attenuation coefficient of the water column associated with increased concentration of the phytoplanktonic Chl-a caused a decrease in light flux that reached the seafloor. The biomass of the microphytobenthos within the upper 1 cm of the sediment, 1.9–46.5 mg Chl-a m⁻², was inversely correlated with the phytoplanktonic biomass in the overlying water column, 10.9–65.0 mg Chl-a m⁻². Thus, interception of light by phytoplankton is considered to be a main cause of the variation in the microphytobenthic biomass. The microphytobenthos biomass showed a significant positive correlation with the macrobenthic density (78–9369 ind. m⁻²) and biomass (0.4–78.8 gWW m⁻²). It appears that the increase in oxygen production by the microphytobenthos allowed macrobenthic animals to become more abundant, as a consequence of oxygenation of the organically enriched muddy sediments (14.5 ± 2.69 mg TOC g⁻¹). This study suggests that the variation in the microphytobenthic biomass is influenced by the phytoplanktonic biomass due to shading effect, and the balance between these two functional groups might affect the variability in the macrobenthic density and biomass.     

Satellite-derived measurements of spatial and temporal chlorophyll-a variability in Funka Bay, southwestern Hokkaido, Japan

Chlorophyll-a (chl-a) concentration has an important economic effect in coastal and marine environments on fisheries resources and marine aquaculture development. Monthly climatologies the Sea-viewing Wide Field-of-view Sensor (SeaWiFS) derived chl-a from February 1998 to August 2004 around Funka Bay were used to investigate the spatial and temporal variability of chl-a concentrations. SeaWiFS-derived suspended sediment, MODIS derived sea surface temperature (SST), solar radiation and wind data were also analyzed. Results showed two distinct chlorophyll blooms in spring and autumn. Chl-a concentrations were relatively low (<0.3 mg m⁻³) in the bay during summer, with high concentrations occurring along the coast, particularly near Yakumo and Shiraoi. In spring, chl-a concentrations increased, and a large (>2 mg m⁻³) phytoplankton bloom occurred. The spatial and temporal patterns were further confirmed by empirical orthogonal function (EOF) analysis. About 83.94% of the variability could be explained by the first three modes. The first chl-a mode (77.93% of the total variance) explained the general seasonal cycle and quantified interannual variability in the bay. The spring condition was explained by the second mode (3.89% of the total variance), while the third mode (2.12% of the total variance) was associated with autumn condition. Local forcing such as the timing of intrusion of Oyashio water, wind condition and surface heating are the mechanisms that controlled the spatial and temporal variations of chlorophyll concentrations. Moreover, the variation of chlorophyll concentration along the coast seemed to be influenced by suspended sediment caused by resuspension or river discharge.     

2003-2016年北极净初级生产力时空变化

The area of Arctic sea ice has dramatically decreased, and the length of the open water season has increased;these patterns have been observed by satellite remote sensing since the 1970 s. In this paper, we calculate the net primary productivity(NPP, calculated by carbon) from 2003 to 2016 based on sea ice concentration products,chlorophyll a(Chl a) concentration, photosynthetically active radiation(PAR), sea surface temperature(SST), and sunshine duration data. We then analyse the spatiotemporal changes in the Chl a concentration and NPP and further investigate the relations among NPP, the open water area, and the length of the open water season. The results indicate that(1) the Chl a concentration increased by 0.025 mg/m~3 per year;(2) the NPP increased by 4.29 mg/(m~2·d) per year, reaching a maximum of 525.74 mg/(m~2·d) in 2016; and(3) the Arctic open water area increased by 57.23×10~3 km~2/a, with a growth rate of 1.53 d/a for the length of the open water season. The annual NPP was significantly positively related to the open water area, the length of the open water season and the SST.The daily NPP was also found to have a lag correlation with the open water area, with a lag time of two months.With global warming, NPP has maintained an increasing trend, with the most significant increase occurring in the Kara Sea. In summary, this study provides a macroscopic understanding of the distribution of phytoplankton in the Arctic, which is valuable information for the evaluation and management of marine ecological environments.     

基于长时序遥感影像的胶州湾秋季叶绿素a浓度变化及其影响因素研究

水体中叶绿素a的含量是研究浮游植物生物量、初级生产力的重要指标，因此对于河口海湾区域叶绿素a的长期监测具有重要的生态学意义。借助遥感技术能够获取长时序大范围观测数据的特点，本文利用1986-2015年的Landsat TM/ETM+影像，通过构建基于实测数据的定量反演模型，得到了30年来胶州湾海域秋季叶绿素a浓度的分布情况:叶绿素a浓度的空间分布规律稳定，呈现西北高东南低的特点，叶绿素a浓度均值呈波动略有上升的趋势。本文利用遥感反演的胶州湾海表面温度及胶州湾流域的土地利用分类和景观格局指数，从自然因素和人为因素两方面对胶州湾叶绿素a浓度时空分布的影响因素进行分析，发现胶州湾秋季叶绿素a浓度与海表面温度密切相关，并且受到近海养殖区破碎化程度、修建大型人工设施等人为因素的影响。     

The distribution of chlorophyll a and its influencing factors in different regions of the Bering Sea

The distribution of chlorophyll a(Chl a) and its relationships with physical and chemical parameters in different regions of the Bering Sea were discussed in July 2010. The results showed the seawater column Chl a concentrations were 13.41–553.89 mg/m2 and the average value was 118.15 mg/m2 in the study areas. The horizontal distribution of Chl a varied remarkably from basin to shelf in the Bering Sea. The regional order of Chl a concentrations from low to high was basin, slope, outer shelf, inner shelf, and middle shelf. The vertical distribution of Chl a was grouped mainly from single-peak type in basin, slope, outer shelf, and middle shelf, where the deep Chl a maxima(DCM) layer was observed at 25–50 m, 30–35 m, 36–44 m, and 37–47 m, respectively. The vertical distribution of Chl a mainly had three basic patterns: standard single-peak type, surface maximum type, and bottom maximum type in the inner shelf. The analysis also showed that the transportation of ocean currents may control the distribution of Chl a, and the effects were not simple in the basin of the Bering Sea. There was a positive correlation between Chl a and temperature, but no significant correlation between Chl a and nutrients. The Bering Sea slope was an area deeply influenced by slope current. Silicate was the factor that controlled the distribution of Chl a within parts of the water in the slope. Light intensity was an important environmental factor in controlling seawater column Chl a in the shelf, where Chl a was limited by nitrate rather than phosphate within the upper water. Meanwhile, there was a positive relationship between Chl a and salinity. Algal blooms broke out at Sta. B6 of the southwestern St. Lawrence Island and Stas F6 and F11 in the middle of the Bering Strait.     

Comparison of Chlorophyll a Concentrations Obtained with 90% Acetone and N,N-dimethylformamide Extraction in Coastal Seawater

Determinations of Chlorophyll a (Chl a) in eutrophic coastal marine waters were compared using N, N-dimethylformamide (DMF) and 90% acetone techniques. Measured Chl a concentrations ranged from 0.89 to 10.65 μg l₋₁ for 90% acetone extracts and from 0.97 to 12.92 μg l⁻¹ for DMF extracts, respectively, for 24 surface water samples from the Harima-nada, Seto Inland Sea, Japan. Chl a concentrations using DMF as a solvent were consistently higher than those found using 90% acetone (p < 0.001, n= 24). Chl a is thus likely to be underestimated (by 13%) when 90% acetone is used for eutrophic waters. This revised version was published online in July 2006 with corrections to the Cover Date.     

桑沟湾养殖生态系统浮游植物的粒径结构及浮游生物对栉孔扇贝碳需求的贡献

The biomass and size fraction of phytoplankton in terms of chlorophyll a(Chl a) was measured during four cruises conducted in April, July, October 2013 and January 2014 in mariculture area, the Sanggou Bay, China.Results show that total Chl a levels in the surface seawater of the Sanggou Bay generally range from 0.10 to 20.46μg/L, with an average value of 2.13 μg/L. Nano-phytoplankton was the most important size-fraction and accounted for about 65.1% of total Chl a. In order to evaluate the importance of the "protozoan trophic link" for energy transfer from the microbial loop to filter-feeding feeders, Zhikong scallop Chlamys farreri was then offered a natural planktonic community as potential prey. Results show that scallops obtained carbon source from natural plankton with the rate of 11 033.05 μg/(g·d). Protists(nanoflagellates and ciliates) were the dominant source of carbon retained by scallop(48.78%). The microbial loop provided 58.45% of the carbon source for farmed scallops. These results indicate that the microbial loop represent a valuable trophic resource in mariculture system of the Sanggou Bay.     

Carbon production and export from Biscayne Bay,Florida. I. Temporal patterns in primary production,seston and zooplankton

Five stations along a transect from the western shore of Biscayne Bay, Florida to the Florida Current were sampled monthly for one year. The variability and amount of seston particulate organic carbon, adenosine triphosphate, chlorophyll a, primary production and zooplankton decreased along the seaward transect. The greater inshore biomass and variability of seston were the result of the allochthonous input of detritus and inorganic nutrients via terrestrial runoff. Annual primary production in this subtropical coastal lagoon ranged from 13 to 46 g C m^?2 yr^?1. Chlorophyll a in the bay ranged from 1 to 3 mg chlorophyll a m^?2. In contrast, chlorophyll a in the surface centimetre of the sediment ranged from 50 to 300 mg chlorophyll a m^?2. In this clear, shallow (2 to 3 m), oligotrophic lagoon, over 90% of total primary production is by submerged macrophytes and benthic algae. The high zooplankton biomass in the bay is most likely sustained by macrophyte detritus and the resuspension of benthic diatoms by the high winds associated with summer squalls and winter cold fronts.     

茅尾海夏季海水与表层沉积物叶绿素a空间分布研究

根据2011年6月茅尾海生态环境调查资料,对该海域海水和表层沉积物中叶绿素a的空间分布进行了分析。结果表明,海水叶绿素a变化范围1.384~4.060 mg/m3,平均值为2.143 mg/m3,表层沉积物叶绿素a范围为0.006~0.740 mg/kg(湿重),均值为0.124 mg/kg;海水与表层沉积物叶绿素a均呈现自河口向南逐渐降低的空间分布特征。单位面积表层沉积物叶绿素a平均含量为上方水柱叶绿素a平均含量的129.44%,沉积物对该海域初级生产力有显著的潜在贡献。相关分析表明,海水和沉积物叶绿素a均与无机氮及底栖动物栖息密度呈显著或极显著的正相关关系(P0.05或P0.01)。     

2008年夏季西北冰洋表层水浮游植物群落结构的色素表征及其对不同水团的响应

本文依托2008年夏季中国第三次北极科学考察航次,对西北冰洋海盆区和楚科奇海陆架营养盐及光合色素进行了测定和分析。根据海水理化性质将研究海区分为5个区,并使用CHEMTAX软件（Mackery et al.,1996）讨论了西北冰洋不同海区浮游植物群落组成结构及其与环境因子之间的关系。结果显示在楚科奇海陆架区,太平洋入流显著影响浮游植物生物量和群落结构。高营养盐Anadyr水团以及白令陆架水控制海域,表现出高Chl a且浮游植物以硅藻为主,相反,低营养盐如阿拉斯加沿岸流控制海域,Chl a生物量低且以微型,微微型浮游植物为主。在外陆架海区,海冰覆盖情况影响着水团的物理特征及营养盐浓度水平,相应地显著影响浮游植物群落结构。在海冰覆盖区域,硅藻生物量站到总Chl a生物量的75%以上;在靠近门捷列夫深海平原海区,受相对高盐的冰融水影响（MW-HS）,营养盐浓度和Chl a浓度相对海冰覆盖区略高,浮游植物结构中微型、微微型藻类比重增加,硅藻比例则降至33%;南加拿大海盆无冰海区（IfB）,表层水盐度最淡,营养盐浓度最低,相应地显示出低Chl a生物量,表明海冰消退,开阔大洋持续时间延长,将导致低生物量及激发更小型浮游植物的生长,并不有利于有机碳向深海的有效输出。     

Phytoplankton biomass and size fractions in surface waters of the Australian sector of the southern ocean

We collected surface water along the 142nd E meridian from Tasmania to Antarctica in December 1999. We measured temperature, salinity and total chlorophyll a; additionally, we collected suspended particle size fractions and used fluorometric analysis to determine the quantity of chlorophyll a in each of four cell size classes: picoplankton (<3 μm), two nanoplankton fractions (3–10 μm and 10–20 μm) and microplankton (> 20 μm). Changes in temperature and salinity show that we crossed 6 water masses separated by 5 fronts. We found low abundance (<0.2 mg m⁻³) of chlorophyll in all size classes, with the exception of higher values near the continent (0.2 to 0.4 mg m⁻³). Lowest chlorophyll values (<0.1 mg m⁻³) were found in the Polar Frontal Zone (51° to 54°S). Microplankton made up the largest portion of total chlorophyll throughout most of the region. We conclude that biomass of all phytoplankton fractions, especially pico-and nanoplankton, was constrained by limiting factors, most probably iron, throughout the region and that ecosystem dynamics within a zone are not circumpolar but are regionalized within sectors.     

中尺度涡影响下的南海西部活性铝分布

To understand the distribution of aluminum(Al) under the influence of mesocale eddies in the western South China Sea(SCS), sea level anomaly, geostrophic current, environmental parameters and reactive Al were investigated in the western SCS in August 2013. The highest reactive Al concentration((180±64) nmol/L) was observed in the surface waters, indicating a substantial atmospheric input. Vertically, the reactive Al decreased from the surface high concentration to the subsurface minima at the depth of chlorophyll a(Chl a) maxima and then increased again with depth at most of the stations. The average concentration of reactive Al in the upper 100 m water column was significantly lower in the cyclonic eddy((137±6) nmol/L) as compared with that in the noneddy waters((180±21) nmol/L). By contrast, the average concentrations of Chl a and silicate in the upper 100 m water column were higher in the cyclonic eddy and lower in the anticyclonic eddy. There was a significant negative correlation between the average concentrations of reactive Al and Chl a in the upper 100 m water column. The vertical distribution of reactive Al and the negative correlation between reactive Al and Chl a both suggest that the reactive Al in the upper water column was significantly influenced by biological removal processes. Our results indicate that mesoscale eddies could regulate the distribution of reactive Al by influencing the primary production and phytoplankton community structure in the western SCS.