Bio-Optical Properties of Seawater in the Western Subarctic Gyre and Alaskan Gyre in the Subarctic North Pacific and the Southern Bering Sea during the Summer of 1997

Chlorophyll a concentrations (chla) and the absorption coefficients of total particulate matter [a _p()], phytoplankton [a _ph()], detritus [a _d()], and colored dissolved organic matter: CDOM [a _CDOM()] were measured in seawater samples collected in the subarctic North Pacific and the southern Bering Sea during the summer of 1997. We examined the specific spectral properties of absorption for each material, and compared the light fields in the Western subarctic Gyre (area WSG) with those in the Alaskan Gyre (area AG), and the southern Bering Sea (area SB). In the area WSG, the irradiance in the surface layer decreased markedly, indicating high absorption. In the area AG, the radiant energy penetrated deeply, and the chl a and absorption values were low throughout the water column. In the area SB, light absorption was high in the surface layer on the shelf edge and decreased with increasing depth; on the other hand, light absorption was low in the surface layer in the shelf area and increased with increasing depth.     

Summer phytoplankton blooms in the oligotrophic North Pacific Subtropical Gyre: Historical perspective and recent observations

The export of organic matter from the oceanic euphotic zone is a critical process in the global biogeochemical cycling of bioelements (C, N, P, Si). Much of this export occurs in the form of sinking particles, which rain down into the unlit waters of the deep sea. Classical models of oceanic production and export balance this gravitational loss of particulate bioelements with an upward flux of dissolved nutrients, and they describe reasonably well those areas of the ocean where deep winter mixing occurs. The surface waters of the North Pacific Subtropical Gyre (NPSG), however, are strongly stratified and chronically nutrient-depleted, especially in summer. Nevertheless, there is ample evidence that blooms of phytoplankton and subsequent pulses of particle export occur during the height of summer stratification in these waters, especially to the northeast of the Hawaiian Islands. These blooms impact regional bioelemental cycling and act as a food source to the deep-sea benthos. We review here numerous published observations of these events in the NPSG, and present new data collected at Station ALOHA (22.75°N, 158°W) during the first 176 cruises of the Hawaii Ocean Time-series program (1988-2005), along with results from transect cruises conducted in the region in 1996 and 2005. We suggest that the summer phytoplankton bloom can be considered a frequent, perhaps annual feature in the northeastern NPSG, and that its perceived stochastic nature is a manifestation of chronic undersampling in time and space. The bloom is typically dominated by only a few genera of large diatoms and the cyanobacterium Trichodesmium. It appears to be consistently supported by dinitrogen fixation, but the fate of the organic matter produced during the summer depends critically on the species composition of the responsible diazotrophs. We estimate that the summer bloom is responsible for up to 38% of N₂ fixation and up to 18% of N-based new production annually at Station ALOHA. We hypothesize that the spatial distribution, timing and magnitude of the bloom may be determined largely by the physical and biological processes controlling new phosphorus delivery into the euphotic zone during the summer and the preceding winter.     

Depth Distribution of the Subtropical Gyre in the North Pacific

Large-scale aspects of the North Pacific subtropical gyre have been investigated using a climatology of temperature and salinity (World Ocean Atlas 1998). In the central and eastern parts of the basin, the axis of the subtropical gyre, defined as the meridional maximum of dynamic height, tends to move poleward from about 25°N near the surface to about 40°N in the upper intermediate layers. In the western part of the basin, the axis is seen at about 30°N, remaining almost unchanged with depth. Striking features associated with this vertical distribution include a northward shift of the bifurcation latitude of the North Equatorial Current at increasing depth and a barotropic nature of the confluence point between the Kuroshio and Oyashio at their respective western boundaries. The former occurs at about 14°N near the surface and extends north of 20°N at depths around 800 m. The latter, situated at about 36.4°N off Japan, does not appear to have a strong signature of depth-dependence. While some of these results are already known from sporadic hydrographic observations, they have not hitherto been represented in a three-dimensional climatology. This revised version was published online in July 2006 with corrections to the Cover Date.     

Spatial Heterogeneity in Distributions of Chlorophyll a Derivatives in the Subarctic North Pacific during Summer

The distribution of chlorophyll a derivatives was examined in samples collected from the subarctic North Pacific during July to September 1997. Pheophorbide a, pheophytin a and pyropheophorbide a as determined by high performance liquid chromatography (HPLC) were the major derivatives recorded. The distribution patterns of chlorophyll a and its derivatives showed a strong vertical and horizontal heterogeneity. Patches with high concentration of derivatives seemed to be associated with high concentration of chlorophyll a. A clear east-west gradient was observed in both chlorophyll a and pheophorbide a integrated from the surface to 100 m depth with significantly higher amounts of both the pigments in the Western Subarctic Gyre and in the Bering Sea than in the Alaskan Gyre. In contrast, no apparent gradient was observed in the integrated pyropheophorbide a and pheophytin a. Grazing experiments conducted with the copepod (Neocalanus cristatus) and salp (Cyclosalpa bakeri) fed on five species of phytoplankton cultures, showed a marked difference in the composition of the derivatives in their fecal pellets. Pyropheophorbide a was dominant in the copepod fecal pellet regardless of the phytoplankton species fed on. In the salp, however, pheophytin a and pheophorbide a were found in the fecal pellets, the relative concentrations varying with the algal food. Spatial heterogeneity in the distribution of the derivatives is considered to reflect local variations in dominant herbivorous processes.     

校园网蠕虫综合防御系统的设计和应用

在蠕虫综合防御思想的基础上,应用各种防御技术,设计1个校园网蠕虫综合防御系统。采用分布式架构,不同类型的Agent独立运行,互相协作配合,共同进行蠕虫防御。系统中采用疫苗更新机制,提高了整体效率。仿真实验和实际应用表明,该系统对提高校园网络的安全性能够起到良好作用。     

北太平洋副热带海洋环流气候变化研究

北太平洋副热带环流的变化在全球气候变化和热量的经向输送中占重要地位。本文对近10年有关北太平洋副热带海洋环流气候变化的研究进行了综述。主要研究成果有：用卫星高度计首次观测到全球海洋Rossby波的传播特征；确定了气候意义下北太平洋副热带逆流为2支．揭示了其中一支与北太平洋模态水的存在有关，另一支是夏威夷群岛附近海洋．大气-陆地相互作用的结果；首次发现了台湾以东黑潮流量有显著的准100天振荡等。本文还提出了在北太平洋副热带环流研究中目前存在的新科学问题。     

北太平洋环流振荡形成机理的数值研究

利用大洋环流模式,探讨海洋对大气强迫的响应及北太平洋环流振荡模态(NPGO)形成的直接原因。对控制试验模拟的海表温度异常(SSTA)进行经验正交函数(EOF)分解,发现第二模态类似于经典NPGO 模态,说明采用该模式研究海洋对大气强迫的响应是可行的。在控制试验基础上,通过改变大气强迫场设计了一系列敏感试验,发现大气强迫场为NPGO 模态正强年的合成场时,所得SST异常场能较好再现NPGO 空间特征,说明海洋状态强烈依赖于大气强迫,大气强迫是造成NPGO 的直接原因;对大气强迫场中的动力强迫、热力强迫等物理量进行不同配置进行试验,发现风场动力强迫对NPGO 的影响最大,是形成NPGO 的关键强迫,其中又以纬向风应力的影响居首。     

Export stoichiometry and migrant-mediated flux of phosphorus in the North Pacific Subtropical Gyre

Export processes play a major role in regulating global marine primary production by reducing the efficiency of nutrient cycling and turnover in surface waters. Most studies of euphotic zone export focus on passive fluxes, that is, sinking particles. However, active transport, the vertical transfer of material by migrating zooplankton, can also be an important component of carbon (C) and nitrogen (N) removal from the surface ocean. Here we demonstrate that active transport is an especially important mechanism for phosphorus (P) removal from the euphotic zone at Station ALOHA (Hawaii Ocean Time-series program; 22°45′N, 158°W), a P-stressed site in the North Pacific Subtropical Gyre. Migrant excretions in this region are P-rich (C₅₁:N₁₂:P₁) relative to sinking particles (C₂₅₀:N₃₁:P₁), and migrant-mediated P fluxes are almost equal in magnitude (82%) to P fluxes from sediment traps. Migrant zooplankton biomass and therefore the importance of this P removal pathway relative to sinking fluxes has increased significantly over the past 12 years, suggesting that active transport may be a major driving force for enhanced P-limitation of biological production in the NPSG. We further assess the C:N:P composition of zooplankton size fractions at Station ALOHA (C₈₈:N₁₈:P₁, on average) and discuss migrant-mediated P export in light of the balance between zooplankton and suspended particle stoichiometries. We conclude that, because active transport is such a large component of the total P flux and significantly impacts ecosystem stoichiometry, export processes involving migrant zooplankton must be included in large-scale efforts to understand biogeochemical cycles.     

Spatial Variability of Living Coccolithophore Distribution in the Western Subarctic Pacific and Western Bering Sea

Vertical distributions of coccolithophores were observed in the depth range 0–50 m in the western subarctic Pacific and western Bering Sea in summer, 1997. Thirty-five species of coccolithophores were collected. Overall, Emiliania huxleyi var. huxleyi was the most abundant taxon, accounting for 82.8% of all coccolithophores, although it was less abundant in the western Bering Sea. Maximum abundance of this species was found in an area south of 41°N and east of 175°E (Transition Zone) reaching >10,000 cells L⁻¹ in the water column. In addition to this species, Coccolithus pelagicus f. pelagicus, which accounted for 4.2% of the assemblage, was representative of the coccolithophore standing crop in the western part of the subarctic Pacific. Coccolithus pelagicus f. hyalinus was relatively abundant in the Bering Sea, accounting for 2.6% of the assemblage. Coccolithophore standing crops in the top 50 m were high south of 41°N (>241 × 10⁶ cells m⁻²) and east of 170°E (542 × 10⁶ cells m⁻²) where temperatures were higher than 12°C and salinities were greater than 34.2. The lowest standing crop was observed in the Bering Sea and Oyashio areas where temperatures were lower than 6–10°C and salinities were less than 33.0. From the coccolithophore volumes, the calcite stocks in the Transition, Subarctic, and the Bering Sea regions were estimated to be 73.0, 9.7, and 6.9 mg m⁻², respectively, corresponding to calcite fluxes of 3.6, 0.5, and 0.3 mg m⁻²d⁻¹ using Stoke's Law. This revised version was published online in July 2006 with corrections to the Cover Date.     

Biogeochemical Variation in Dimethylsulfide, Phytoplankton Pigments and Heterotrophic Bacterial Production in the Subarctic North Pacific during Summer

Dimethylsulfide (DMS), chlorophyll a (Chl-a), accessory pigments (fucoxanthin, peridinin and 19-hexanoyloxyfucoxanthin), and bacterial production (BP) were measured in the surface layer (0–100 m) of the subarctic North Pacific, including the Bering Sea, during summer (14 July–5 September, 1997). In surface sewater, the concentrations of DMS and Chl-a varied widely from 1.3 to 13.2 nM (5.1 ± 3.0 nM, mean ± S.D., n = 48) and from 0.1 to 2.4 µg L^–1 (0.6 ± 0.6 µg L^–1, n = 24), respectively. In the subarctic North Pacific, DMS to Chl-a ratios (DMS/Chl-a) were higher on the eastern side than the western side (p < 0.0001). Below the euphotic zone, DMS/Chl-a ratios were law and the correlation between DMS and Chl-a was relatively strong (r ² = 0.700, n = 27, p < 0.0001). In the euphotic zone, DMS/Chl-a ratios were higher and the correlation between DMS and Chl-a was weak (r ² = 0.128, n = 50, p = 0.01). The wide variation in DMS/Chl-a ratios would be at least partially explained by the geographic variation in the taxonomic composition of phytoplankton, because of the negative correlation between DMS/Chl-a and fucoxanthin-to-Chl-a ratios (Fuc/Chl-a) (r ² = 0.476, n = 26, p = 0.0001). Furthermore, there was a positive correlation between DMS and BP (r ² = 0.380, n = 19, p = 0.005). This suggests that BP did not represent DMS and dimethylsulfoniopropionate (DMSP) removal by bacterial consumption but rather DMSP degradation to DMS by bacterial enzyme.     

北太平洋亚热带环流区与西太平洋暖池区浮游动物群落垂直分布的差异及其对碳通量的意义

The mesozooplankton in both epipelagic and mesopelagic zones is essentially important for the study of ecosystem and biological carbon pump. Previous studies showed that the diel vertical migration(DVM) pattern of mesozooplankton varied among ecosystems. However, that pattern was largely unknown in the Western Pacific Warm Pool(WPWP). The vertical distribution, DVM and community structure of mesozooplankton from the surface to 1 000 m were compared at Stas JL7K(WPWP) and MA(North Pacific Subtropical Gyre, NPSG). Two sites showed similarly low biomass in both epipelagic and mesopelagic zones, which were in accordance with oligotrophic conditions of these two ecosystems. Stronger DVM(night/day ratio) was found at JL7K(1.31) than that at MA(1.09) on surface 0–100 m, and an obvious night increase of mesopelagic biomass was observed at JL7K, which was probably due to migrators from bathypelagic zone. Active carbon flux by DVM of zooplankton was estimated to be 0.23 mmol/(m2·d) at JL7K and 0.16 mmol/(m~2·d) at MA. The community structure analysis showed that calanoid copepods, cnidarians and appendicularians were the main contributors to DVM of mesozooplankton at both sites. We also compared the present result with previous studies of the two ecosystems,and suggested that the DVM of mesozooplankton was more homogeneous within the WPWP and more variable within the NPSG, though both ecosystems showed typically extremely oligotrophic conditions. The different diel vertical migration strength of mesozooplankton between NPSG and WPWP implied different efficiency of carbon pump in these two ecosystems.     

Behavior of particulate materials during iron fertilization experiments in the Western Subarctic Pacific (SEEDS and SEEDS II)

During two mesoscale iron-enrichment studies in the northwestern subarctic Pacific (SEEDS in 2001 summer and SEEDS II in 2004 summer), particulate materials from the iron-induced phytoplankton bloom in the upper water column were monitored to analyze the export processes beneath the upper mixed layer, mainly with drifting sediment traps. We could not observe the total downward export process of the high accumulation of particulate organic carbon from the mixed layer induced by the large diatom bloom of SEEDS [e.g., Tsuda, A., Takeda, S., Saito, H., Nishioka, J., Nojiri, Y., Kudo, I., Kiyosawa, H., Shiomoto, A., Imai, K., Ono, T., Shimamoto, A., Tsumune, D., Yoshimura, T., Aono, T., Hinuma, A., Kinugasa, M., Suzuki, K., Sohrin, Y., Noiri, Y., Tani, H., Deguchi, Y., Tsurushima, N., Ogawa, H., Fukami, K., Kuma, K., Saino, T., 2003. A mesoscale iron enrichment in the western subarctic Pacific induces large centric diatom bloom. Science 300, 958–961] because the 2-week observation period was too short to examine the decline phase of the bloom. In contrast, in SEEDS II, the particulate organic carbon and particulate organic nitrogen were accumulated 123 and 23 mmol m⁻², respectively, in the mixed layer until day-15 (days from iron-enrichment), and then ca. 90% were removed from the mixed layer by day-25. The sediment traps at 40 m depth between day-15 and day-25 accounted for at least more than 35% of these particles. There was no large variation in chemical composition in settling particles above 100 m depth throughout the experimental periods both in SEEDS and SEEDS II. The content of biogenic opal remained more than 50% of all settling particles during SEEDS, while the content of biogenic calcium carbonate was relatively high, with a low biogenic opal content of consistently less than 30% during SEEDS II. These results suggest that high standing stock of seed population of diatoms before the iron fertilization, indicated by low C/Si ratio of particulate matter, is an important factor to induce the large diatom bloom in SEEDS.     

Long-Term Variability of North Pacific Subtropical Mode Water in Response to Spin-Up of the Subtropical Gyre

The Meteorological Research Institute's ocean general circulation model (MRI-OGCM) has been used to investigate the temperature variability of the North Pacific Subtropical Mode Water (NPSTMW) over a time series longer than 5 years via the spin-up of the subtropical gyre. Besides an interannual variation, the wintertime sea surface temperature in the area where the NPSTMW is formed, and the temperature of the NPSTMW itself, both change remarkably in a >5-year time scale. An analysis of heat budgets showed that the long-term changes in NPSTMW temperature are due mainly to a leading advection of heat by the Kuroshio Extension and compensating surface heat flux. As a result of a dynamical adjustment to the wind stress fields, the transports of the Kuroshio and the Kuroshio Extension increased in the mid 1970s with a lag of 3 years after the wind stress curl in the central North Pacific. The increased heat advection by the Kuroshio Extension induces a warming in the mixed layer in the NPSTMW formation area, followed by a warming of the NPSTMW itself. Both these warming actions increase the heat release to the atmosphere. These results imply that the surface heat flux over the Kuroshio Extension area varies in response to the change in the ocean circulation through the spin-up of the subtropical gyre. This revised version was published online in August 2006 with corrections to the Cover Date.     

Community Structure and Dynamics of Phytoplankton in the Western Subarctic Pacific Ocean: A Synthesis

The phytoplankton community in the western subarctic Pacific (WSP) is composed mostly of pico- and nanophytoplankton. Chlorophyll a (Chl a) in the <2 μm size fraction accounted for more than half of the total Chl a in all seasons, with higher contributions of up to 75% of the total Chl a in summer and fall. The exception is the western boundary along the Kamchatka Peninsula and Kuril Islands and the Oyashio region where diatoms make up the majority of total Chl a during the spring bloom. Among the picophytoplankton, picoeukaryotes and Synechococcus are approximately equally abundant, but the former is more important in term of carbon biomass. Despite the lack of a clear seasonal variation in Chl a concentration, primary productivity showed a large seasonal variation, and was lowest in winter and highest in spring. Seasonal succession in the phytoplankton community is also evident with the abundance of diatoms peaking in May, followed by picoeukaryotes and Synechococcus in summer. The growth of phytoplankton (especially >10 μm cell size) in the western subarctic Pacific is often limited by iron bioavailability, and microzooplankton grazing keeps the standing stock of pico- and nano-phytoplankton low. Compared to the other HNLC regions (the eastern equatorial Pacific, the Southern Ocean, and the eastern subarctic Pacific), iron limitation in the Western Subarctic Gyre (WSG) may be less severe probably due to higher iron concentrations. The Oyashio region has similar physical condition, macronutrient supply and phytoplankton species compositions to the WSG, but much higher phytoplankton biomass and primary productivity. The difference between the Oyashio region and the WSG is also believed to be the results of difference in iron bioavailability in both regions. This revised version was published online in July 2006 with corrections to the Cover Date.     

Benthic community responses to pulses in pelagic food supply: North Pacific Subtropical Gyre

Time-series measurements of particulate organic carbon (POC) and particulate nitrogen (PN) fluxes, sediment community composition, and sediment community oxygen consumption (SCOC) were made at the Hawaii Ocean Time-series station (Sta. ALOHA, 4730 m depth) between December 1997 and January 1999. POC and PN fluxes, estimated from sediment trap collections made at 4000 m depth (730 m above bottom), peaked in late August and early September 1998. SCOC was measured in situ using a free vehicle grab respirometer that also recovered sediments for chemical and biological analyses on six cruises during the 1-year study. Surface sediment organic carbon, total nitrogen and phaeopigments significantly increased in September, corresponding to the pulses in particulate matter fluxes. Bacterial abundance in the surface sediment was highest in September with a subsurface high in November. Sediment macrofauna were numerically dominated by agglutinating Foraminifera fragments with highest density in September. Metazoan abundance, dominated by nematodes was also highest in September. SCOC significantly increased from a low in February to a high in September. POC and PN fluxes at 730 m above bottom were significantly correlated with SCOC with a lag time of ⩽14 days, linking pelagic food supply with benthic processes in the oligotrophic North Pacific gyre. The annual supply of POC into the abyss compared to the estimated annual demand by the sediment community (POC:SCOC) indicates that only 65% of the food demand is met by the supply of organic carbon.     

Balance of Volume Transports between Horizontal Circulation and Meridional Overturn in the North Pacific Subarctic Region

Horizontal and meridional volume transports on timescales from intra-seasonal to interannual in the North Pacific subarctic region were investigated using a reanalysis dataset for 1993–2001 that was constructed from an assimilation of the TOPEX altimeter and in situ data into an eddy-permitting North Pacific ocean general circulation model. The barotropic flow is excited along east of the Emperor Seamounts by the western intensification dynamics. The volume transport of this flow compensates for that across the interior region east of the Seamounts below the summit depth of the Seamounts. The Oyashio, which is also considered as a compensation flow for the transport in the whole interior region, includes baroclinic as well as barotropic components. Baroclinic transports in the whole interior region exceed those in the western boundary region in the upper (200–1000 m) and lower (2000–5000 m) layers, and the total transport is northward (southward) in the upper (lower) layer. These excesses of the baroclinic transport are balanced by a vertical transport of the meridional overturn. The meridional overturn has a complementary relation to the basin-scale baroclinic circulation in the North Pacific subactic region. This revised version was published online in July 2006 with corrections to the Cover Date.     

Diatom Assemblage and Productivity Changes during the Last 340,000 Years in the Subarctic Pacific

Diatoms represent the major part of the microfossils preserved in the subarctic Pacific sediments. During the warm climate intervals the diatom accumulation rate (DAR) tended to increase, whereas it tended to decrease during the cold intervals. Principal component (PC) analysis of the fossil diatom assemblages in Piston Core KH99-3 ES samples from the subarctic Pacific was carried out to investigate the paleoceanographic conditions of the area. PC1 (59.9% of the total variance) was represented by Neodenticula seminae, a characteristic species representing the high nutrient concentrations, which dominated during the interglacial periods. The DAR and PC changes can be attributed to a global origin that is influenced by the nutrient change due to intermediate water change and to a local one that is explained by the change of upwelling region. The analysis of PC2 (17.1% of the total variance) indicates the change of water mass in the western subarctic Pacific. During Marine Isotope Stages (MIS) 2 and 3, the western subarctic Pacific region was significantly influenced by the sea-ice, which may have been derived from the coastal region or a proximal marginal sea. This revised version was published online in July 2006 with corrections to the Cover Date.     

Influence of non-photosynthetic pigments on light absorption and quantum yield of photosynthesis in the western equatorial Pacific and the Subarctic North Pacific

Regional variations in the contribution of non-photosynthetic pigments (ā _np^*) to the total light absorption of phytoplankton (ā _ph^*) and its influence on the maximum quantum yield of photosynthesis (φ _m) were investigated. In the western equatorial Pacific, the surface ā _np^* : ā _ph^* ratio was higher in the western warm pool than that in the upwelling region. This difference appears to be attributable to severe nitrate depletion and higher percentage of prokaryotes, which can accumulate very high concentrations of zeaxanthin in the western warm pool. In the subarctic North Pacific, the ā _np^* : ā _ph^* ratio was expected to be higher in the Alaskan Gyre where the thermocline is sharper and iron limitation may possibly be more severe than in the Western Subarctic Gyre. However, the ratio was actually higher in the Western Subarctic Gyre, contradictory to our expectations. This east-west variation appears to be attributable to changes in the taxonomic composition; cyanobacteria were more abundant in the Western Subarctic Gyre. The values of ā _np^* : ā _ph^* and its vertical variations were relatively small in the subarctic North Pacific compared to those in the western equatorial Pacific. These inter-regional variations appear to be attributable to the lower solar radiation intensity, smaller percentage of cyanobacteria, and relatively strong vertical mixing in the subarctic North Pacific. The spatial variations in ā _np^* : ā _ph^* significantly influence φ _m. In comparison with φ _m based on the total light absorption (φ _{m ph}), the values corrected for the contribution of non-photosynthetic pigments (φ _{m ps}) showed an increase in both the western equatorial Pacific and the subarctic North Pacific.     

Variability of chlorophyll and primary production in the Eastern North Atlantic Subtropical Gyre: potential factors affecting phytoplankton activity

Size-fractionated chlorophyll-a and carbon incorporation rates were determined on a series of 13 cruises carried out from 1992 to 2001with the aim of investigating the patterns and causes of variability in phytoplankton chlorophyll and production in the Eastern North Atlantic Subtropical Gyral Province (NASE). Averaged (±SE) integrated chlorophyll-a concentration and primary production rate were 17±1 mg m⁻² and 253±22 mg C m⁻² d⁻¹. Small-sized cells (<2 μm) formed the bulk of phytoplankton biomass (71%) and accounted for 54% of total primary production. A clear latitudinal gradient in these variables was not detected. By contrast, large seasonal variability was detected in terms of primary production, although integrated phytoplankton biomass, as estimated from chlorophyll-a concentration, remained rather constant and did not display significant changes with time. Variability in primary production (PP) was related mainly to variability in surface temperature and surface chlorophyll-a concentration. The control exerted by surface temperature was related to nutrient availability. By contrary, euphotic-zone depth, depth of maximum concentration of chlorophyll-a and integrated chlorophyll-a did not contribute significantly to the high variability in primary production observed in this oligotrophic region.