天鹅洲长江段浮游植物群落季节演替及环境因子研究

通过研究长江天鹅洲白鱀豚国家级自然保护区长江段浮游动植物群落组成及其与环境因子之间的关系,揭示长江干流浮游植物群落的结构特征,以期为该区域生态系统保护与研究工作提供基础资料和科学依据。2014—2015年按季节分四次在干流沿途设置的10个采样点开展了浮游植物调查,并同步开展水温、溶解氧、总磷等水质因子监测,并采用典范对应分析(canonical correspondence analysis,CCA)研究了不同季节浮游植物群落结构变化及其与环境因子的关系。结果表明:浮游植物种类以硅藻、绿藻和蓝藻为主;全年来看生物量组成主要是硅藻和隐藻,不过春秋季时蓝绿藻密度组成占优。CCA分析则进一步显示浮游植物群落组成四季变化明显,其中磷酸盐、溶解氧和氧化还原位等水质因子对春季浮游植物群落结构存在显著的影响,但是夏冬季的浮游植物群落变化则可能是水文情势等条件起到了重要作用。     

南流江河口区春季浮游植物群落结构组成与分布特征

通过2016年3月底现场航次11个站点的调查,应用反相高效液色谱（RP-HPLC）并结合二极管阵列检测器分析技术,分析了春季广西南流江河口区浮游植物光合色素组成,进而由CHEMTAX软件估算全粒级浮游植物的群落结构。结果表明:春季含量较高的浮游植物特征光合色素含量以叶绿素b最高,其次为岩黄藻素;浮游植物的优势类群为隐藻,其次为绿藻和硅藻,它们分别平均占据了浮游植物生物量的54.95%、23.36%和17.37%,其他藻类所占比例很低。南流江河口区浮游植物群落结构东西部入海分支有较大差异:东部分支营养盐较西部分支低,隐藻所占生物量比例最高,其次为绿藻和硅藻,浮游植物群结构与分布受营养盐因素影响较大;西部分支营养盐含量明显比东部分支高,绿藻和硅藻的所占比例有所提升,隐藻的生物量所占比则有所下降,浮游植物群落结构与分布受非营养盐因素的影响较大。南流江河口区浮游植物生物量和群落结构除了受营养盐影响外,还与浊度、盐度等密切相关,表明南流江浊度增加已明显影响着生态系统结构与功能,需要密切关注和进一步研究。     

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.     

Long-term changes in summer phytoplankton communities of the open northern Baltic Sea

Changes in the biomass and species composition of phytoplankton may reflect major shifts in environmental conditions. We investigated relationships between the late summer biomass of different phytoplankton taxa and environmental factors, and their long-term (1979–2003) trends in two areas of the Baltic Sea, the northern Baltic proper (NBP) and the Gulf of Finland (GF), with statistical analyses. An increasing trend was found in late summer temperature and chlorophyll a of the surface water layer (0–10 m) in both areas. There was also a significant decrease in summer salinity and an increase in winter dissolved inorganic nitrogen to phosphorus (DIN:DIP) ratio in the NBP, as well as increases in winter DIN concentrations and DIN:SiO₄ ratio in the GF. Simultaneously, the biomass of chrysophytes and chlorophytes increased in both areas. In the NBP, also the biomass of dinophytes increased and that of euglenophytes decreased, whereas in the GF, cyanobacteria increased and cryptophytes decreased. Redundancy analysis (RDA) indicated that summer temperature and winter DIN concentration were the most important factors with respect to changes in the phytoplankton community structure. Thus, the phytoplankton communities seem to reflect both hydrographic changes and the ongoing eutrophication process in the northern Baltic Sea.     

Environmental influence on phytoplankton communities in the northern Benguela ecosystem

An investigation of surface phytoplankton communities was undertaken on the shelf of the northern Benguela upwelling ecosystem during austral autumn (May) and spring (September), along latitudinal transects at 20° S and 23° S, from 2 to 70 nautical miles offshore, as well as on a zigzag grid located between these transects. Microscopic identification of the phytoplankton and CHEMTAX analysis of pigment biomarkers were used to characterise the community composition. During May 2014, warmer, more-saline water with a shallower upper mixed layer corresponding to periods of less-intense offshore Ekman transport was encountered on the shelf. Satellite imagery indicated high phytoplankton biomass extending for a considerable distance from the coast, and CHEMTAX indicated diatoms as dominant at most of the stations (52–92%), although dinoflagellates were dominant at some inshore localities (57–74%). Species of Chaetoceros, Bacteriastrum and Cylindrotheca were the most abundant, with abundance of the Pseudo-nitzschia ‘seriata-group’ being particularly high at a number of stations. In September 2014, more-intense wind-forcing resulted in a deeper upper mixed layer and stronger upwelling of colder, less-saline water. Elevated phytoplankton biomass was confined close to the coast, where diatoms accounted for most of the population (54–87%), whereas small flagellates, such as prasinophytes, haptophytes and cryptophytes, as well as the cyanobacterium Synechococcus, dominated the communities (58–90%) farther from the coast. It is hypothesised that stronger upwelling and deeper vertical mixing in September of that year were not conducive for widespread diatom growth, and that small flagellates populated the water column by being entrained from offshore onto the shelf in the upwelled water that moved in towards the coast.     

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

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

Evaluation of the performance of HPLC–CHEMTAX analysis for determining phytoplankton biomass and composition in a turbid estuary (Schelde, Belgium)

In the upper Schelde estuary in 2002, phytoplankton biomass and community composition were studied using microscopic and pigment analyses. Chlorophyll a concentration was a good predictor of phytoplankton biomass estimated from cell counts and biovolume measurements. The phytoplankton carbon to chlorophyll a ratio, however, was often unrealistically low (<10). CHEMTAX was used to estimate the contribution of the major algal groups to total chlorophyll a. The dominant algal groups were diatoms and chlorophytes. While diatom equivalents in chlorophyll a predicted diatom biomass relatively well, chlorophyte equivalents in chlorophyll a were only weakly related to chlorophyte biomass. The pigment-based approach to study phytoplankton overestimated phytoplankton biomass in general and chlorophyte biomass in particular in late autumn and winter, when phytoplankton biomass was low. A possible explanation for this overestimation may be the presence of large amounts of vascular plant detritus in the upper Schelde estuary. Residual chlorophyll a, chlorophyll b and lutein in this detritus may result in an overestimation of total phytoplankton and chlorophyte biomass when the contribution of phytoplankton to total particulate organic matter is low.     

Seasonal dynamics of the phytoplankton community in Sendai Bay,northern Japan

Sendai Bay is located on the Pacific coast of northern Japan and suffered serious damage following the 2011 off the Pacific coast of Tohoku earthquake and tsunami in March 2011. To assess the impact on the marine ecosystem, information was needed on the phytoplankton communities and their seasonal variation. However, such information was limited. Therefore, an intensive monitoring of the phytoplankton was carried out from March 2012 to April 2014. Seasonal variation of the phytoplankton community was similar at coastal and offshore stations. Total phytoplankton biomass, based on Chl a concentration, peaked in spring and then decreased to a minimum in summer, before gradually increasing during early winter and peaking again in the following spring. This seasonal pattern was consistent with previous studies conducted before the earthquake and tsunami. Also, size structure of the phytoplankton community and its four main groups was estimated from the size-fractioned samples of Chl a. Our results also showed that the spring bloom consisted of large diatoms, with their growth ceasing due to nitrogen depletion. The bloom was followed by a summer period where cyanobacteria and picoeukaryote became dominant, with high cell densities in spite of low nutrient concentrations. In addition, sporadic environmental changes, such as those following typhoons, were observed. These resulted in large increases/decreases in individual phytoplankton groups.     

Temporal variations in lower trophic level biological environments in the northwestern North Pacific Subtropical Gyre from 1950 to 1997

An examination of large archives (1950–1997) of the oceanographic and atmospheric data from the northwestern North Pacific Subtropical Gyre has revealed clear linkages between atmospheric forcing factors, physical processes and biological events. Large changes in the winter and spring biomass of phytoplankton and macroplankton observed over annual, decadal and inter-decadal time scales could clearly be attributed to climate-related changes in oceanographic processes. Interannual changes in the intensity of the winter-time East Asian Monsoon had a significant impact on the extent of convective overturning, on nitrate inputs into the euphotic zone and the concentrations of chlorophyll a in winter and during the following spring. A prolonged period of deeper winter mixed layers observed from the mid-1970s to the mid-1980s led to a sizeable increase in winter mixed-layer nitrate concentrations. This change resulted in a decrease in winter-time phytoplankton biomass. Spring-time chlorophyll a, in contrast, showed a steady increase during this period. The decline in winter phytoplankton biomass could be attributed to the depths of mixed layer. A deeper mixed layer prevents phytoplankton from remaining in the euphotic zone for long enough to photosynthesize and grow, leaving substantial amounts of nutrients unutilised. However, as a result of stratification of the water column in spring following each of these winters, phytoplankton could take advantage of the enhanced ambient concentrations of nutrients and increase its biomass. Another noteworthy observation for the period from the mid-1970s to the early 1980s is that the western subtropical gyre progressively became phosphate limited. The period of diminishing mixed-layer phosphate concentrations was observed in our study area from the early 1990s onwards was consistent with recent observations at Station ALOHA in the eastern subtropical gyre.     

Grazing impact of microzooplankton on phytoplankton in the Xiamen Bay using pigment-specific dilution technique

Phytoplankton group-specific growth and microzooplankton grazing were determined seasonally using the dilution technique with high-performance liquid chromatography （HPLC） in the Xiamen Bay, a subtropical bay in southeast China, between May 2003 and February 2004. The results showed that growth rates of phytoplankton ranged from 0.71 to 2.2 d^-1 with the highest value occurred in the inner bay in May. Mierozooplankton grazing rates ranged from 0.5 to 3.1 d^-1 with the highest value occurred in the inner bay in August. Microzooplankton grazing impact ranged from 39% to 95% on total phytoplankton Chl a biomass, and 65% to 181% on primary production. The growth and grazing rates of each phytoplankton group varied, the highest growth rate （up to 3.3 d^-1 ） was recorded for diatoms in August, while the maximum grazing rate （ up to 2.1 d ^-1 ） was recorded for chlorophytes in February in the inner bay. Among main phytoplankton groups, grazing pressure of microzooplankton ranged from 10% to 83% on Chl a biomass, and from 14% to 151% on primary production. The highest grazing pressure on biomass was observed for cryptophytes （83%） in August, while the maximum grazing pressure on primary production was observed for eyanobacteria （up to 151% ） in December in the inner bay. Net growth rates of larger phytoplanktons （diatoms and dinoflagellates） were higher than those of smaller groups （ prasinophytes, chlorophytes and cyanobacteria）. Relative preference index showed that microzooplankton grazed preferentially on prasinophytes and avoided to harvest diatoms in cold seasons （December and February）.     

Spring and summer phytoplankton communities in the Chukchi and Eastern Beaufort Seas

Phytoplankton pigments and size-fractionated biomass in the Chukchi and Beaufort Seas showed spatial and temporal variation during the spring and summer of 2002. Cluster analysis of pigment ratios revealed different assemblages over the shelf, slope and basin regions. In spring, phytoplankton with particle sizes greater than 5 μm, identified as diatoms and/or haptophytes, dominated over the shelf. Smaller (<5 μm) phytoplankton containing chlorophyll b, most likely prasinophytes, were more abundant over the slope and basin. Due to extensive ice cover at this time, phytoplankton experienced low irradiance, but nutrients were near maximal for the year. By summer, small prasinophytes and larger haptophytes and diatoms co-dominated in near-surface assemblages in largely ice-free waters when nitrate was mostly depleted. Deeper in the water column at 1–15% of the surface irradiance larger sized diatoms were still abundant in the upper nutricline. Phytoplankton from the shelf appeared to be advected through Barrow Canyon to the adjacent basin, explaining similar composition between the two areas in spring and summer. Off-shelf advection was much less pronounced for other slope and basin areas, which are influenced by the low-nutrient Beaufort gyre circulation, leading to a dominance of smaller prasinophytes and chlorophytes. The correlation of large-sized fucoxanthin containing phytoplankton with the higher primary production measurements shows promise for trophic status to be estimated using accessory pigment ratios.     

大亚湾大鹏澳牡蛎养殖临近海域自养微微型浮游生物种群分布特征

于2013年5月到2014年6月,在大亚湾大鹏澳牡蛎区及邻近海域开展了为期14个月的采样调查,利用高效液相色谱(HPLC)法对表层水体中微微型浮游植物(0.7—2.7μm)光合色素进行测定,并应用色素化学分类软件CHEMTAX对自养微微型浮游生物(aototrophicpicoplankton,APP)功能类群进行分析。结果表明,该海域APP中共检出了15种光合色素,其中青绿藻素(Pras)和玉米黄素(Zea)是微微型色素中浓度最高的2种特征色素,均具有明显的季节变化特征:Pras主要出现在低温季节(牡蛎养殖期),而Zea主要出现在高温季节(非牡蛎养殖期)。CHEMTAX分析表明,大鹏澳海域APP最主要的类群是硅藻、蓝藻和青绿藻,而甲藻、隐藻、定鞭金藻、绿藻和金藻生物量较低。温度和营养盐浓度是影响大鹏澳海域APP的时空分布的重要因素,青绿藻主要出现在低温季节(主要在冬季牡蛎养殖期间),且其生物量与溶解无机氮呈显著正相关;而蓝藻则主要出现在高温季节,与温度呈显著正相关。另外,贝类养殖也是能够影响APP空间分布的重要因素,在大鹏澳海域牡蛎养殖期间,青绿藻生物量在养殖区明显高于非养殖海域。     

Pigment signatures of phytoplankton composition in the northern Benguela ecosystem during spring

Pigment indices were used to investigate the distribution and composition of phytoplankton in the northern Benguela during the austral spring of 2000, with sampling being conducted on five transect lines between 19°S and 25°S and at other inshore stations on the shelf. Total chlorophyll a concentrations (TChla) at the surface ranged from 18.4mg m^?3 at shallow inshore stations to 0.58mg ^m?3 at offshore localities in deeper water. Phytoplankton communities in the inshore environment were characterised by elevated biomass levels and the dominance of diatoms (high Fuc_DP indices). These diatom populations were associated with cool, higher density, nutrient-rich waters near the coast. Phytoplankton biomass declined with increasing distance offshore and the communities changed to a mixed population where small flagellates were generally dominant (high Flag_DP indices). These flagellate communities were associated with warmer, lower density waters. Whereas dinoflagellates (Per_DP Index) and prokaryotes (Zea_DP Index) were observed throughout the region, their contribution to the communities was of secondary importance.     

Contrasting oceanographic conditions and phytoplankton communities on the east and west coasts of Australia

The composition and dynamics of the phytoplankton communities and hydrographic factors that control them are described for eastern and western Australia with a focus on the Eastern Australian Current (EAC) and Leeuwin Current (LC) between 27.5° and 34.5°S latitude. A total of 1685 samples collected from 1996 to 2010 and analysed for pigments by high performance liquid chromatography (HPLC) showed the average TChla (monovinyl+divinyl chlorophyll a) concentration on the west coast to be 0.28±0.16 ??g L⁻¹ while it was 0.58±1.4 ??g L⁻¹ on the east coast. Both coasts showed significant decreases in the proportions of picoplankton and relatively more nanoplankton and microplankton with increasing latitude. On both coasts the phytoplankton biomass (by SeaWiFS) increased with the onset of winter. At higher latitudes (>27.5°S) the southeast coast developed a spring bloom (September) when the mean monthly, surface chlorophyll a (chla) concentration (by SeaWiFS) was 48% greater than on the south west coast. In this southern region (27.5-34.5°S) Synechococcus was the dominant taxon with 60% of the total biomass in the southeast (SE) and 43% in the southwest (SW). Both the SE and SW regions had similar proportions of haptophytes; ∼14% of the phytoplankton community. The SW coast had relatively more pelagophytes, prasinophytes, cryptophytes, chlorophytes and less bacillariophytes and dinophytes. These differences in phytoplankton biomass and community composition reflect the differences in seasonality of the 2 major boundary currents, the influence this has on the vertical stability of the water column and the average availability of nutrients in the euphotic zone. Seasonal variation in mixed layer depth and upwelling on the west coast appears to be suppressed by the Leeuwin Current. The long-term depth averaged (0-100 m) nitrate concentration on the west coast was only 14% of the average concentration on the east coast. Redfield ratios for NO₃:SiO₂:PO₄ were 6.5:11.9:1 on the east coast and 2.2:16.2:1 on the west coast. Thus new production (nitrate based) on the west coast was likely to be substantially more limited than on the eastcoast. Short term (hourly) rates of vertical mixing were greater on the east coast. The more stable water column on the west coast produced deeper subsurface chlorophyll a maxima with a 25% greater proportion of picoeukaryotes.     

Temporal variation in phytoplankton composition related to water mass properties in the central part of Sagami Bay

Temporal variations in water mass properties and the composition of phytoplankton pigments in the central part of Sagami Bay were investigated by monthly observations from June 2002 to May 2004. Eleven pigments were quantified using high-performance liquid chromatography (HPLC) from 100%, 20%, and 5% light depths relative to the surface; the class-specific composition of phytoplankton community was then obtained by CHEMTAX analysis. The study area was influenced by the Kuroshio water for most of the observation period. The mean contribution of diatoms in all samples was relatively low (29%), while that of flagellates, mainly chlorophytes or cryptophytes, was quite high (60%). The phytoplankton composition at the three depths was uniform throughout the observation period, indicating that the vertical structure of the phytoplankton community did not develop significantly over time. A distinct temporal pattern was observed: flagellates dominated during the summer of 2002 and the winters of 2002–2003 and 2003–2004, while diatoms dominated during the summer of 2003. This pattern was associated with water mass changes. The community in the summer of 2003 was influenced by coastal water. While no distinct spring bloom of phytoplankton was observed, a weak increase in chlorophyll a was observed during the spring of 2004. Ocean color satellite data showed that fluctuations in chlorophyll a concentrations at time scales much shorter than a month occurred during the spring of 2003 and that the elevations in chlorophyll a levels were not continuous. The fluctuations were probably associated with rapid flushing by the Kuroshio water, which has low chlorophyll a content.     

Chemotaxonomic investigation of phytoplankton in the shelf ecosystem of the KwaZulu-Natal Bight,South Africa

Phytoplankton community structure was elucidated during summer and winter in the KwaZulu-Natal Bight using pigment and CHEMTAX analyses. The surface pattern in January 2010 indicated that diatoms, haptophytes and prasinophytes tended to be the most prominent groups inshore and in the southern sector of the bight, whereas Prochlorococcus and Synechococcus were more dominant in the north and towards the offshore region. At the deep chlorophyll maximum (DCM), diatoms were dominant in the inner bight and in the south, with prasinophytes being the prominent flagellates, but this changed to Prochlorococcus and pelagophytes being the important groups in the northern sector and outer part of the bight. A different pattern was observed at the surface in July 2010, where diatoms, haptophytes, prasinophytes and cryptophytes in varying proportions comprised most of the community in the inner half of the bight, whereas Synechococcus and haptophytes were the main groups in the outer sector. A similar pattern occurred at the DCM, except that Synechococcus was less prominent and pelagophytes were distributed across the bight in both the inshore and offshore zones. Observations and relationships between phytoplankton groups and environmental parameters indicated that the groups were most closely related to temperature, accounting for 24–64% of the deviance. The influence of nutrients on phytoplankton was less clear but nitrate and silicate seemed to account for some of the patchy distribution patterns.     

Annually recurrent phytoplanktonic assemblages during summer in the seasonal ice zone west of the Antarctic Peninsula (Southern Ocean)

The distribution of phytoplankton composition, cell abundance and biomass from an area along the Western Antarctic Peninsula was studied during three summers, with the aim of understanding its dynamics over spatial and interannual scales. The studied area is characterized by seasonal sea-ice retreat and advance. Algae composition and concentration were found to be highly variable through the area as well as from year to year. Small unidentified phytoflagellates, diatoms and cryptophytes were the main phytoplankton groups, contributing the major proportion of total phytoplankton cell abundance and biomass concentration. Three annually recurrent phytoplankton assemblages were recognized in the area according to the algae composition and abundance: a diatom bloom associated with the sea-ice edge, an assemblage dominated by small unidentified phytoflagellates and cryptophytes, and a diatom-enriched assemblage in open waters. The distribution of these assemblages varied from year-to-year. During the summers preceded by early sea-ice retreat, the diatom bloom was spatially restricted and the other two assemblages occupied extended regions, whereas during the late sea-ice retreat year, the diatom bloom extended over a larger region and the other assemblages occupied smaller regions or were just absent. It was detected that these assemblages resemble different stages of the phytoplankton seasonal cycle, and that their distribution through the area can be related to a latitudinal and longitudinal gradient in the phytoplankton growth onset timing, associated with the progressive sea-ice retreat during spring. The local environmental conditions associated with each assemblage were also analyzed, but further study is needed for understanding the causes of the replacement of one assemblage by another through the area. On the other hand, the interannual variability in the distribution of the assemblages can be related to year-to-year differences in the timing of phytoplankton growth onset, associated with variations in the timing of the sea-ice retreat.     

Seasonal variations in nutrients and chlorophyll-a concentrations in the Northern east China sea

Nutrients, chlorophyll-a, particulate organic carbon (POC), and environmental conditions were extensively investigated in the northern East China Sea (ECS) near Cheju Island during three seasonal cruises from 2003 to 2005. In spring and autumn, relatively high concentrations of nitrate (2.6~12.4 μmol kg^-1) and phosphate (0.17~0.61 μmol kg^-1) were observed in the surface waters in the western part of the study area because of the large supply of nutrients from deep waters by vertical mixing. The surface concentrations of nitrate and phosphate in summer were much lower than those in spring and autumn, which is ascribed to a reduced nutrient supply from the deep waters in summer because of surface layer stratification. While previous studies indicate that upwellings of the Kuroshio Current and the Changjiang (Yangtze River) are main sources of nutrients in the ECS, these two inputs seem not to have contributed significantly to the build-up of nutrients in the northern ECS during the time of this study. The lower nitrate:phosphate (N:P) ratio in the surface waters and the positive correlation between the surface N:P ratio and nitrate concentration indicate that nitrate acts as a main nutrient limiting phytoplankton growth in the northern ECS, contrary to previous reports of phosphate-limited phytoplankton growth in the ECS. This difference arises because most surface water nutrients are supplied by vertical mixing from deep waters with low N:P ratios and are not directly influenced by the Changjiang, which has a high N:P ratio. Surface chlorophyll-a levels showed large seasonal variation, with high concentrations (0.38~4.14 mg m^-3) in spring and autumn and low concentrations (0.22~1.05 mg m^-3) in summer. The surface distribution of chlorophyll-a coincided fairly well with that of nitrate in the northern ECS, implying that nitrate is an important nutrient controlling phytoplankton biomass. The POC:chlorophyll-a ratio was 4~6 times higher in summer than in spring and autumn, presumably because of the high summer phytoplankton death rate caused by nutrient depletion in the surface waters.     

The seasonal cycle of surface chlorophyll in the Peruvian upwelling system: A modelling study

The seasonal variability of surface chlorophyll in the northern Humboldt Current System is studied using satellite data, in situ observations and model simulations. The data show that surface chlorophyll concentration is highest in austral summer and decreases during austral winter, in phase opposition with coastal upwelling intensity. A regional model coupling ocean dynamics and biogeochemical cycles is used to investigate the processes which control this apparently paradoxical seasonal cycle. Model results suggest that the seasonal variability of the mixed layer depth is the main controlling factor of the seasonality. In winter, the mixed layer deepening reduces the surface chlorophyll accumulation because of a dilution effect and light limitation. In summer, biomass concentrates near the surface in the shallow mixed layer and nitrate limitation occurs, resulting in a biomass decrease in the middle of summer. Intense blooms occur during the spring restratification period, when winter light limitation relaxes, and during the fall destratification period, when the surface layer is supplied with new nutrients. Model sensitivity experiments show that the seasonal variations in insolation and surface temperature have little impact on the surface chlorophyll variability.     

Modeling of spring bloom in the western subarctic Pacific (off Japan) with observed vertical density structure

Effects of vertical stability on spring blooms of phytoplankton were investigated for the western subarctic Pacific ocean using a one-dimensional (depth) ecosystem model. In the model, vertical stability was expressed by diffusion constants calculated from observed density distribution. Dynamics of phytoplankton in blooms was calculated by the model using the vertical diffusion. Then, the calculated results were compared with the Coastal Zone Color Scanner (CZCS) data. The comparison shows that the shallow surface mixed layer causes early start days of spring blooms at inshore (northern) stations. In addition, spring blooms continue long at inshore (northern) stations since a water column has weak stability. This is because weak stability of a water column causes large nutrient supply from a deep layer and large diffusive transport of phytoplankton biomass from the subsurface maximum.