Seasonal pigment patterns of surface phytoplankton in the subtropical southern hemisphere

Pigment indices were used to characterise the seasonal succession of phytoplankton, and associated changes in chlorophyll a and accessory pigments, in subtropical waters of the three ocean basins in the southern hemisphere. Diagnostic indices revealed the dominance of small flagellates and elevated biomass during winter–spring in the Pacific, mixed flagellate–prokaryote communities and intermediate biomass during early summer in the Atlantic, and prokaryote dominance with low biomass in mid-summer in the Indian Ocean. Photo-pigment indices indicated only a small variation in the chlorophyll a proportion of total pigments across the ocean basins, but the accessory pigments varied considerably. Under low temperature and irradiance conditions, the photosynthetic carotenoids were prominent, but as temperatures and irradiance increased and nutrients declined, there was a significant increase in the proportion of photoprotective carotenoids. At high temperatures and irradiances, the photoprotective carotenoids were the largest component of the pigment pool, exceeding the proportion of chlorophyll a. These variations in phytoplankton composition, and their photoacclimation status, could be explained according to environmental changes and have implications for satellite estimations of biomass and primary production.     

2010-2011年胶州湾叶绿素a与环境因子的时空变化特征

2010年4、6、8、10月和2011年1、3月在胶州湾开展了6个航次的综合调查,研究了表层海水温度、盐度、营养盐和叶绿素a浓度的时空变化特征。调查期间,总无机氮(DIN)、磷酸盐(PO4)和硅酸盐(SiO3)多呈现东北部湾边缘高,而湾内和湾口低的空间分布特征。季节变化表明,DIN和PO4主要受养殖排放、河流径流输入和浮游植物生长消耗的影响,呈现初夏和秋季高,夏末和冬季低的特点;而SiO3主要受河流径流输入和浮游植物消耗的影响,呈现夏、秋高,而冬、春低的特点。营养盐浓度和结构分析表明,胶州湾存在PO4和SiO3的绝对和相对限制;SiO3限制尤其严重,是控制胶州湾浮游植物生长的主要环境因子。SiO3和PO4的限制主要表现在冬季,几乎遍布整个海湾;夏季降水可有效缓解海域的SiO3限制。叶绿素a浓度呈现春、夏季高,秋、冬季低的季节分布,温度、营养盐浓度与结构和季节性贝类养殖活动是控制胶州湾叶绿素a浓度时空分布的关键因素。     

High-resolution phytoplankton diel variations in the summer stratified central Yellow Sea

Vertical distributions of phytoplankton biomass and community structure were studied in the summer stratified central Yellow Sea (YS) using a submersible spectrofluorometer (Fluoroprobe, bbe Moldaenke, Germany), along with photosynthetic pigments analysis (HPLC-CHEMTAX), and microscope observation. Above all, the results of the dominating group obtained from these methods generally coincided with each other on the transect 35°N. The concentrations of brown algae, green algae and total chlorophyll a (Chl a) biomass were highly correlated between the results of Fluoroprobe observations and pigments analysis (r?=?0.79, 0.91 and 0.82, respectively, n?=?54, p?<?0.01). In the summer stratified central YS, significant differences in phytoplankton compositions on the vertical distribution was observed. On the basis of HPLC-CHEMTAX results, the dominating group of phytoplankton composition generally changed from cyanobacteria to chrysophytes and then to diatoms, from surface to bottom. Interestingly, on the basis of high-resolution observations using Fluoroprobe, a periodic fluctuation of the pycnocline presumably due to the semidiurnal internal tides was observed at an anchor station (35°N, 123°E). In addition, both nutrients and Chl a concentrations at the depth of the subsurface chlorophyll maximum (SCM) seemed to coincide with the rhythm of the pycnocline fluctuation, indicating the latter might have a potential impact on the dynamics of SCM phenomena in the summer stratified central YS.     

Estuarine phytoplankton dynamics and shift of limiting factors: A study in the Changjiang (Yangtze River) Estuary and adjacent area

Environmental factors in estuaries are highly variable in terms of both spatial and temporal dimensions and hence phytoplankton biomass, as well as community structure, is dynamic. Two cruises were carried out in the Changjiang (Yangtze River) Estuary and adjacent area in spring and summer. The result of CHEMTAX calculation suggests that in spring diatoms and chlorophytes contribute equally to phytoplankton biomass, while phytoplankton community structure is mainly composed of diatoms in summer. We encountered blooms in summer with chlorophyll a (CHLa) over 10 μg l⁻¹ off the Changjiang Estuary and they were mainly caused by diatoms (>90%). Based on the HPLC analysis of samples collected, phytoplankton pigments mainly concentrated beyond the front between 122.5°E and 123°E where nutrients and turbidity were best balanced. Euphotic depth (Z_eu, calculated from Secchi disk depth) to surface mixed layer depth (Z_mix) ratio (i.e. Z_eu/Z_mix) were comparable in spring (average value 1.2) and the ratio increased to 5.2 in summer. Variation of the ratio indicates an apparent shift of light and physical conditions from spring to summer. Correspondingly, CHLa was positively related to Z_eu/Z_mix ratio (r² = 0.83) in spring, indicating the light limitation over the whole investigation area. On the other hand, the relationship of CHLa and Z_eu/Z_mix ratio became unclear when Z_eu/Z_mix ratio >3 in summer. This is probably due to the combination of both light limitation before the front and nutrient limitation beyond the front. In addition, evidence was found that light condition can impact the diagnostic pigments in the Changjiang Estuary.     

基于遥感数据分析南海叶绿素与颗粒物的季节变化与相互关系

本文基于卫星遥感的叶绿素a浓度与颗粒物后向散射系数月平均数据以及其他海洋与气象参数,详细分析了两个生物光学参量在季节尺度上的相关性及其与物理参数的耦合关系,并运用光驯化模型分析了南海表层水体浮游植物的生理学季节变化特征。结果表明,受南海地形和风场等物理参量的变化,南海叶绿素a浓度与颗粒物后向散射系数存在显著的季节和空间分布特征,具有一定的共变性和差异性。在南海近岸及浅水区,叶绿素a浓度与颗粒物后向散射系数有很好的耦合关系;而在南海深水海盆区,叶绿素a浓度冬高夏低,其季节循环过程与颗粒物后向散射系数相反,这主要是受浮游植物生理学过程的影响。"生物量控制区"与"光驯化控制区"的分界在南海与陆架-海盆分界线一致,体现了水深条件对浮游植物生理状态的影响。此外本文还发现,在吕宋海峡西部海区,叶绿素a与颗粒物后向散射系数的关系表现出"生物量-光驯化共同控制"的特点。     

Algal class abundances,estimated from chlorophyll and carotenoid pigments,in the western Equatorial Pacific under El Niño and non-El Niño conditions

Phytoplankton samples were collected from the equatorial Pacific (10°S to 10°N along 155°E) in June 1992 as part of the Australian contribution to the JGOFS program. Chlorophyll and carotenoid pigments were measured by HPLC, and a PC-based computer program (CHEMTAX) was used to estimate the contribution of 9 algal classes to the total chlorophyll a concentration in 9 separate depth bands at each location. This cruise occurred in the middle of the prolonged 1991/1993 El Niño, and the results are compared with similar data from a cruise in October 1990 which occurred before this El Niño but after the 1988/1989 La Niña.Changes in the pigment : chlorophyll a ratios appeared consistent across algal classes and, apart from some minor exceptions, consistent between cruises. Pigments involved in light-harvesting generally increased relative to chlorophyll a with increasing depth, whereas the ratio for photoprotective pigments (e.g. diadinoxanthin) usually decreased with depth. The zeaxanthin concentration per cell for cyanobacteria decreased with depth in the surface 75 m during 1992 as would be expected for a photoprotective pigment.Based on their contribution to the total chlorophyll a concentration, haptophytes, prochlorophytes, cyanobacteria (Synechococcus) and chlorophytes were the dominant algal classes in 1992. The chlorophyte contribution to chlorophyll a in 1992 (14.8%) was almost double that in 1990 (7.8%). This increase was largely at the expense of the cyanobacteria and haptophytes, which both decreased significantly. The increase in chlorophytes in 1992 was particularly noticeable in the surface waters south of the equator at about 4°S, where there was evidence of upwelling.     

Light absorption by phytoplankton,non-algal particles and dissolved organic matter at the Patagonia shelf-break in spring and summer

Satellite image studies and recent in situ sampling have identified conspicuous phytoplankton blooms during spring and summer along the Patagonia shelf-break front. The magnitudes and spectral characteristics of light absorption by total particulate matter (phytoplankton and detritus) and colored dissolved organic matter (CDOM) have been determined by spectrophotometry in that region for spring 2006 and late summer 2007 seasons. In spring, phytoplankton absorption was the dominant optical component of light absorption (60–85%), and CDOM showed variable and important contributions in summer (10–90%). However, there was a lack of correlation between phytoplankton biomass (chlorophyll-a concentration or [chl a]) and the non-algal compartment in both periods. A statistically significant difference was found between the two periods with respect to the CDOM spectral shape parameter (S_cdom), with means of 0.015 (spring) and 0.012 nm^?1 (summer). Nonetheless, the mean S_cdm values, which describe the slope of detritus plus CDOM spectra, did not differ between the periods (average of 0.013 nm^?1). Phytoplankton absorption values in this work showed deviations from mean parameterizations in previous studies, with respect to [chl a], as well as between the two study periods. In spring, despite the microplankton dominance, high specific absorption values and large dispersion were found (a*_ph(440)=0.04±0.03 m² mg [chl a]^?1), which could be attributed to an important influence of photo-protector accessory pigments. In summer, deviations from general trends, with values of a*_ph(440) even higher (0.09±0.02 m² mg [chl a]^?1), were due to the dominance of small cell sizes and also to accessory pigments. These results highlight the difficulty in deriving robust relationships between chlorophyll concentration and phytoplankton absorption coefficients regardless of the season period. The validity of a size parameter (S_f) derived from the absorption spectra has been demonstrated and was shown to describe the size structure of phytoplankton populations, independently of pigment concentration, with mean values of 0.41 in spring and 0.72 in summer. Our results emphasize the need for specific parameterization for the study region and seasonal sampling approach in order to model the inherent optical properties from water reflectance signatures.     

Depth habitats and seasonal distributions of recent planktic foraminifers in the Canary Islands region (29°N) based on oxygen isotopes

Seasonal depth stratified plankton tows, sediment traps and core tops taken from the same stations along a transect at 29°N off NW Africa are used to describe the seasonal succession, the depth habitats and the oxygen isotope ratios (δ¹⁸O_shell) of five planktic foraminiferal species. Both the δ¹⁸O_shell and shell concentration profiles show variations in seasonal depth habitats of individual species. None of the species maintain a specific habitat depth exclusively within the surface mixed layer (SML), within the thermocline, or beneath the thermocline. Globigerinoides ruber (white) and (pink) occur with moderate abundance throughout the year along the transect, with highest abundances in the winter and summer/fall season, respectively. The average δ¹⁸O_shell of G. ruber (w) from surface sediments is similar to the δ¹⁸O_shell values measured from the sediment-trap samples during winter. However, the δ¹⁸O_shell of G. ruber (w) underestimates sea surface temperature (SST) by 2 °C in winter and by 4 °C during summer/fall indicating an extension of the calcification/depth habitat into colder thermocline waters. Globigerinoides ruber (p) continues to calcify below the SML as well, particularly in summer/fall when the chlorophyll maximum is found within the thermocline. Its vertical distribution results in δ¹⁸O_shell values that underestimate SST by 2 °C. Shell fluxes of Globigerina bulloides are highest in summer/fall, where it lives and calcifies in association with the deep chlorophyll maximum found within the thermocline. Pulleniatina obliquiloculata and Globorotalia truncatulinoides, dwelling and calcifying a part of their lives in the winter SML, record winter thermocline (～180 m) and deep surface water (～350 m) temperatures, respectively. Our observations define the seasonal and vertical distribution of multiple species of foraminifera and the acquisition of their δ¹⁸O_shell.     

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.     

Assemblages of phytoplankton pigments along a shipping line through the North Atlantic and tropical Pacific

A set of phytoplankton pigment measurements collected on eight quarterly transects from France to New Caledonia is analyzed in order to identify the main assemblages of phytoplankton and to relate their occurrence to oceanic conditions. Pigment concentrations are first divided by the sum [monovinyl chlorophyll a plus divinyl chlorophyll a] to remove the effect of biomass, and second are normalized to give an equal weight to all pigments. The resulting 17 pigments × 799 observations matrix is then classified into 10 clusters using neural methodology. Eight out of these 10 clusters have a well marked regional or seasonal character, thus evidencing adapted responses of the phytoplankton communities. The main gradient opposes two clusters with high fucoxanthin and chlorophyll c₁₊₂ in the North Atlantic in January, April and July, to three clusters in the South Pacific Subtropical Gyre with high divinyl chlorophyll a, zeaxanthin and phycoerythrin. One of the clusters in the South Pacific Subtropical Gyre has relatively high zeaxanthin and phycoerythrin contents and dominates in November and February (austral summer), while another with relatively high divinyl chlorophylls a and b dominates in May and August (austral winter). The third one in the South Pacific is characterized by high carotene concentration and its occurrence peaks in February and May. In the equatorial current system, one cluster, rich in chlorophylls b and c₁₊₂, is strictly located in a narrow zone centred at the equator, while another with relatively high violaxanthin concentration is restricted to the high nutrient - low chlorophyll waters in only the southern part of the South Equatorial Current. One cluster with relatively high prasinoxanthin content has a spatial distribution spanning the entire South Equatorial Current. Two clusters have a ubiquitous distribution: one in the equatorial Pacific, the Carribbean Sea and the North Atlantic during summer has pigment concentrations close to the average of the entire dataset, and the other in the South Pacific Subtropical Gyre, the Carribbean Sea and the North Atlantic during autumn clearly has an oligotrophic character. Many of the differences between clusters are caused by diagnostic pigments of nano- or picoflagellates. While the space and time characteristics of the clusters are well marked and might correspond to differences in physical and chemical forcing, knowledge of the ecological requirements of these flagellates is generally lacking to explain how the variability of the environment triggers these clusters.     

Diel vertical distribution of planktonic ciliates within the surface layer of the NW Mediterranean (May 1995)

The composition and vertical distribution of planktonic ciliates within the surface layer was monitored over four diel cycles in May 95, during the JGOFS-France DYNAPROC cruise in the Ligurian Sea (NW Mediterranean). Ciliates were placed into size and trophic categories: micro- and nano-heterotrophic ciliates, mixotrophic ciliates, tintinnids and the autotrophic Mesodinium rubrum. Mixotrophic ciliates (micro and nano) represented an average of 46% of oligotrich abundance and 39% of oligotrich biomass; nano-ciliates (hetero and mixotrophic) were abundant, representing about 60 and 17% of oligotrich abundance and biomass, respectively. Tintinnid ciliates were a minor part of heterotrophic ciliates. The estimated contribution of mixotrophs to chlorophyll a concentration was modest, never exceeding 9% in discrete samples. Vertical profiles of ciliates showed that chlorophyll-containing ciliates (mixotrophs and autotrophs) were mainly concentrated and remained at the chlorophyll a maximum depth. In contrast, among heterotrophic ciliates, a portion of the population appeared to migrate from 20–30 m depth during the day to the surface at night or in the early morning. Correlation analyses of ciliate groups and phytoplankton pigments showed a strong relationship between nano-ciliates and zeaxanthin, and between chlorophyll-containing ciliates and chlorophyll a, as well as other pigments that were maximal at the chlorophyll a maximum depth. Total surface layer concentrations showed minima of ciliates during nightime/early morning hours.     

Seasonal dynamics of light absorption by chromophoric dissolved organic matter (CDOM) in the NW Mediterranean Sea (BOUSSOLE site)

We analyze a two-year time-series of chromophoric dissolved organic matter (CDOM) light absorption measurements in the upper 400 m of the water column at the BOUSSOLE site in the NW Mediterranean Sea. The seasonal dynamics of the CDOM light absorption coefficients at 440 nm (a_cdom(440)) is essentially characterized by (i) subsurface maxima forming in spring and progressively reinforcing throughout summer, (ii) impoverishment in the surface layer throughout summer and (iii) vertical homogeneity in winter. Seasonal variations of the spectral dependence of CDOM absorption, as described by the exponential slope value (S_cdom), are characterized by highest values in summer and autumn at the surface and low values at the depths of a_cdom(440) subsurface maxima or just below them. Variations of a_cdom(440) are likely controlled by microbial digestion of phytoplankton cells, which leads to CDOM production, and by photochemical destruction (photobleaching), which leads to CDOM degradation. Photobleaching is also the main driver of S_cdom variations. Consistently with previous observations, a_cdom(440) for a given chlorophyll a concentration is higher than expected from Case I waters bio-optical models. The total non-water light absorption budget shows that surface waters at the BOUSSOLE site are largely dominated by CDOM during all seasons but the algal bloom in March and April. These results improve the knowledge of CDOM absorption dynamics in the Mediterranean Sea, which is scarcely documented. In addition, they open the way to improved algorithms for the retrieval of CDOM absorption from field or satellite radiometric measurements.     

Phytoplankton pigment and absorption characteristics along meridional transects in the Atlantic Ocean

Pigment patterns and associated absorption properties of phytoplankton were investigated in the euphotic zone along two meridional transects in the Atlantic Ocean, between the UK and the Falkland Islands, and between South Africa and the UK. Total chlorophyll a (TChla=MVChla+DVChla+chlorophyllide a) concentrations and the biomarker pigments for diatoms (fucoxanthin), nanoflagellates and cyanobacteria (zeaxanthin) appeared to have similar distribution patterns in the spring and in the autumn in the temperate NE Atlantic and the northern oligotrophic gyre. Divinyl chlorophyll a levels (prochlorophytes) were greater in spring at the deep chlorophyll maximum in the oligotrophic gyre, however. Marked seasonal differences were observed in the NW African upwelling region. TChla concentrations were twice as high in the upper mixed layer in the spring, with the community dominated by diatoms and prymnesiophytes (19′-hexanoyloxyfucoxanthin). A layered structure was prevalent in the autumn where cyanobacteria, diatoms and prymnesiophytes were located in the upper water column and diatoms and mixed nanoflagellates at the sub-surface maximum. In the South Atlantic, the Benguela upwelling ecosystem and the Brazil-Falklands Current Confluence Zone (BFCCZ) were the most productive regions with the TChla levels being twice as high in the Benguela. Diatoms dominated the Benguela system, while nanoflagellates were the most ubiquitous group in the BFCCZ. Pigment concentrations were greater along the eastern boundary of the southern oligotrophic gyre and distributed at shallower depths. Deep chlorophyll maxima were a feature of the western boundary oligotrophic waters, and cyanobacteria tended to dominate the upper water column along both transects with a mixed group of nanoflagellates at the chlorophyll maximum.Absorption coefficients were estimated from spectra reconstructed from pigment data. Although absorption was greater in the productive areas, the TChla-specific coefficients were higher in oligotrophic regions. In communities that were dominated by diatoms or nanoflagellates, pigment absorption was generally uniform with depth and attenuating irradiance, with TChla being the major absorbing pigment at 440 nm and photosynthetic carotenoids (PSC) at 490 nm. Absorption by chlorophyll c and photoprotective carotenoids (PPC) was much lower. Populations where cyanobacteria were prevalent were characterized by high PPC absorption, particularly at 490 nm, throughout most of the euphotic zone. The data suggested that the effect of pigments on the variability of phytoplankton absorption was due primarily to the variations in absorption by PPC.     

Physiological response and photoacclimation capacity of Caulerpa prolifera (Forsskål) J.V. Lamouroux and Cymodocea nodosa (Ucria) Ascherson meadows in the Mar Menor lagoon (SE Spain)

The macroalga Caulerpa prolifera colonized the Mar Menor coastal lagoon after the enlargement of the main inlet in 1972, coexisting now with the previous Cymodocea nodosa meadows. The physiological response and the photoacclimation capacity of both species were studied. For this purpose in vivo chlorophyll a fluorescence, photoprotective mechanisms and oxidative stress were measured in both species in summer 2010 and exposure-recovery experiments were conducted to determine the acclimation capacity of both species. The results suggest that C. prolifera behaves as a shade-adapted species with a low photoprotective capacity, light being one of the main factors governing its distribution in the lagoon. The high photosynthetic capacity and lack of photoinhibition found in C. nodosa suggest that this species is highly photoprotected. It also possesses a high concentration of lutein and a high de-epoxidation degree, related to a much higher NPQ_max value.     

Growth and grazing rate dynamics of major phytoplankton groups in an oligotrophic coastal site

There has been more attention to phytoplankton dynamics in nutrient-rich waters than in oligotrophic ones thus requiring the need to study the dynamics and responses in oligotrophic waters. Accordingly, phytoplankton community in Blanes Bay was overall dominated by Prymnesiophyceae, remarkably constant throughout the year (31 ± 13% Total chlorophyll a, Tchl a) and Bacillariophyta with a more episodic appearance (20 ± 23% Tchl a). Prasinophyceae and Synechococcus contribution became substantial in winter (Prasinophyceae = 30% Tchl a) and summer (Synechococcus = 35% Tchl a). Phytoplankton growth and grazing mortality rates for major groups were estimated by dilution experiments in combination with high pressure liquid chromatography and flow cytometry carried out monthly over two years. Growth rates of total phytoplankton (range = 0.30–1.91 d⁻¹) were significantly higher in spring and summer (μ > 1.3 d⁻¹) than in autumn and winter (μ ∼ 0.65 d⁻¹) and showed a weak dependence on temperature but a significant positive correlation with day length. Microzooplankton grazing (range = 0.03–1.4 d⁻¹) was closely coupled to phytoplankton growth. Grazing represented the main process for loss of phytoplankton, removing 60 ± 34% (±SD) of daily primary production and 70 ± 48% of Tchl a stock. Chla synthesis was highest during the Bacillarophyceae-dominated spring bloom (Chl a_synt = 2.3 ± 1.6 μg Chl a L⁻¹ d⁻¹) and lowest during the following post-bloom conditions dominated by Prymnesiophyceae (Chl a_synt = 0.23 ± 0.08 μg Chl a L⁻¹ d⁻¹). This variability was smoothed when expressed in carbon equivalents mainly due to the opposite dynamics of C:chl a (range = 11–135) and chl a concentration (range = 0.07–2.0 μg chl a L⁻¹). Bacillariophyta and Synechococcus contribution to C fluxes was higher than to biomass because of their fast-growth rate. The opposite was true for Prymnesiophyceae.     

Temporal variability in phytoplankton pigments,picoplankton and coccolithophores along a transect through the North Atlantic and tropical southwestern Pacific

Biogeochemical processes in the sea are triggered in various ways by chlorophyll-containing phytoplankton groups. While the variability of chlorophyll concentration at sea has been observed from satellites for several years, these groups are known only from cruises which are limited in space and time. The Geochemistry, Phytoplankton and Color of the Ocean programme (GeP&CO) was set up to describe and understand the variability of phytoplankton composition on large spatial scales under a multi-year sampling strategy. It was based on sea-surface sampling along the route of the merchant ship Contship London which travelled four times a year from Le Havre (France) to Nouméa (New Caledonia) via New York, Panama and Auckland. Observations included the measurement of photosynthetic pigments, counts of picoplanktonic cells by flow cytometry (Prochlorococcus, Synechococcus, and picoeucaryotes) and counting and identification of coccolithophores. The results confirmed that tropical areas have low seasonal variability and are characterized by relatively high divinyl-chlorophyll a and zeaxanthin concentration and that the variability is strongest at high latitudes where the phytoplankton biomass and population structure are found to have large seasonal cycles. Thus, the spring bloom in the North Atlantic and an austral winter bloom north of New Zealand are marked by chlorophyll concentrations which are often higher than 0.5 μg l⁻¹ and by high concentration of fucoxanthin (a pigment used as an indicator for diatoms), while summer populations are dominated by Prochlorococcus sp. and have low chlorophyll concentrations. Apart from this yearly bloom at temperate latitudes, fucoxanthin is scarce, except in the equatorial upwelling zone in the eastern Pacific Ocean, where it is found in moderate amounts. In this region, relatively high chlorophyll concentrations extend generally as far as 14°S and do not respond to the seasonal strengthening of the equatorial upwelling during the austral winter. Prochlorococcus, which is known to dominate in oligotrophic tropical seas and to disappear in cold conditions, in fact has its minimum during the spring bloom in the North Atlantic, rather than during the winter. Coccolithophores are ubiquitous, showing a succession of species in response to oceanic conditions and provinces. 19′Hexanoyloxyfucoxanthin, the pigment generally considered as an indicator of coccolithophores, is relatively abundant at all times and in all regions, but its abundance is generally not tightly correlated with that of coccolithophores. The regional differences revealed by these results are in overall agreement with Longhurst's division of the ocean into ecological provinces.     

Seasonal dynamics of sub-ice fauna below pack ice in the Arctic (Fram Strait)

Pronounced seasonality is a characteristic feature of polar ecosystems, but seasonal studies in the high-Arctic pack-ice zone are still scarce because of logistical constraints. During six expeditions (1994–2003) to the Fram Strait area between Greenland and Svalbard in winter, spring, early summer, late summer and autumn, the sub-ice habitat and fauna below the pack ice (0–1 m depth) were analyzed for seasonal patterns. Both environmental variables such as ice cover, temperature, salinity and chlorophyll a (chl a), as well as species composition, abundance and biomass of the sub-ice fauna showed distinct seasonal dynamics. Most species of the sub-ice fauna were found in early summer, followed by autumn, spring and late summer; the lowest number occurred in winter. The sub-ice fauna was dominated by copepod nauplii during all seasons. Next numerous was the small pelagic copepod Oithona similis, followed by occassional swarms of Pseudocalanus minutus and Calanus spp. Abundances of the sympagic fauna in the sub-ice water layer were much lower, with ectinosomatid copepods being usually the most numerous sympagic group. In the course of the year, total abundances of the sub-ice fauna showed a steep increase from the earliest sampling dates towards the end of winter/beginning of spring reaching maximum numbers then, and a decrease to minimum numbers in early summer. A second peak occurred in late summer, followed by a decrease towards autumn. This significant trend was due to the abundances of copepod nauplii and Oithona similis. Sympagic species were virtually absent during winter, and increased significantly in spring and early and late summer. A factor analysis revealed the variables ice cover and thickness, water temperature and salinity, as well as chl a as the major controlling factors for the seasonal patterns in different groups and species of the sub-ice fauna. Because of the special environmental conditions in the sub-ice habitat, and the unique species composition characterized by small taxa, young stages, and sympagic species, the seasonal dynamics of the Arctic sub-ice fauna differ substantially from those of the epipelagic zooplankton community in the Arctic Ocean.     

Cross-basin differences in particulate organic carbon export and flux attenuation in the subtropical North Atlantic gyre

We compared in-situ and satellite-derived measures of the biological carbon pump efficiency at the two seemingly similar subtropical North Atlantic gyre time series sites, the Bermuda time series (BATS, Bermuda Atlantic time-series study and OFP, ocean flux program) in the western gyre and the ESTOC time series (European station for time-series in the ocean, Canary Islands) in the eastern gyre. Satellite-derived surface chlorophyll a was slightly lower at Bermuda compared to ESTOC (annual average of 0.10±0.04 vs. 0.14±0.05-mg-m^?3), as was satellite-derived primary production (annual average of 380±77 vs. 440±80-mg C-m^?2 d^?1). However, export production normalized to primary production (export ratio) was higher at Bermuda by a factor of 2–3 when estimated using mesopelagic traps moored at 500-m depth and by a factor of 3–4 when estimated using surface-tethered drifting traps. When averaged seasonally, flux at BATS was highest in spring (March, April, May) at all depths followed by summer (June, July, August) and decreasing towards fall, but this seasonality was less visible at ESTOC. Seasonal comparison showed the fastest flux attenuation at Bermuda in winter and spring, coinciding with the highest POC flux. POC/PIC ratios derived from the moored traps were significantly higher at BATS than at ESTOC in fall and winter, but this difference was not significant in spring (p>0.05). This study shows that while the western and eastern Atlantic subtropical gyres have similar rates of primary production, the biological carbon pump differs between the two provinces. Higher new nutrient input observed at Bermuda compared to ESTOC might explain part of the difference in export ratio but alone is insufficient. Greater winter mixed-layer depths and higher mesoscale eddy activity at Bermuda resulting in pulsed production events of labile organic matter might explain both the higher export flux and export ratios found at Bermuda.     

东海区叶绿素a和初级生产力季节变化特征

依据2008年春季(5月)、夏季(8月)、秋季(11月)和2009年冬季(2月)的现场调查结果,分析了东海区叶绿素a、初级生产力的平面分布、垂直分布和季节变化的特征,并探讨了其影响因素。结果表明,四个航次叶绿素a浓度分别为1.33、0.93、1.61和0.65 mg/m3,秋季春季夏季冬季。春季、夏季和秋季最大值均出现在0—10m水层,冬季最大值出现在底层。叶绿素a浓度远海年季变化较小,近岸区和垂直分布年季变化较大。四个航次初级生产力平均为375.03、414.37、245.45和102.60 mg/(m3 h),夏季秋季春季冬季。叶绿素a浓度和初级生产力水平均高于历史同期值。鱼外渔场的年平均初级生产力最大,海州湾渔场最小。通过分析叶绿素a和环境因子的相关性表明,叶绿素a与浮游植物显著正相关;春季和秋季的低温以及春季和夏季的低盐比较适合浮游植物的生长;活性磷酸盐可能是限制春季和秋季叶绿素a的重要因素。     

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.