Diel Changes in Vertical Distribution and Feeding Conditions of the Chaetognath Parasagitta elegans (Verill) in the Subarctic Pacific in Summer

Diel changes in vertical distribution and feeding conditions of the chaetognath Parasagitta elegans (Verill) were observed in three regions of the subarctic North Pacific in the summer of 1997. Samples were collected by repeated vertical hauls with a Vertical Multiple Plankton Sampler (VMPS) for 15–45 hours by demarcating the 0–500 m water column into four sampling layers. Integrated abundance through the entire water column and the proportion of juveniles were higher in the Bering Sea than the western and eastern subarctic Pacific. Juveniles always inhabited the surface layer in the western subarctic Pacific and Bering Sea, but they inhabited the underlying layer in the eastern subarctic Pacific. Stages I–III concentrated into the upper 150 m in the western subarctic Pacific but were distributed widely from 20–300 m in the Bering Sea. Among them, Stages II and III migrated rather synchronously over a wide vertical range in the eastern subarctic Pacific. The feeding rate of P. elegans was calculated to be 0.18 prey/chaetognath/day in the western subarctic Pacific, 0.27 prey/chaetognath/day in the Bering Sea and 0.07 prey/chaetognath/day in the eastern subarctic Pacific.     

Nutrient variability in the mixed layer of the subarctic Pacific Ocean, 1987–2010

Data collected primarily from commercial ships between 1987 and 2010 are used to provide details of seasonal, interannual and bidecadal variability in nutrient supply and removal in the surface ocean mixed layer across the subarctic Pacific. Linear trend analyses are used to look for impacts of climate change in oceanic domains (geographic regions) representing the entire subarctic ocean. Trends are mixed and weak (generally not significant) in both winter and summer despite evidence that the upper ocean is becoming more stratified. Overall, these data suggest little change in the winter resupply of the mixed layer with nutrients over the past 23 years. The few significant trends indicate a winter increase in nitrate (~0.16 μM year⁻¹) in the Bering Sea and in waters off the British Columbia coast, and a decline in summer phosphate (0.018 μM year⁻¹) in the Bering. An oscillation in Bering winter nutrient maxima matches the lunar nodal cycle (18.6 years) suggesting variability in tidal mixing intensity in the Aleutian Islands affects nutrient transport. Nitrate removal from the seasonal mixed layer varies between 6 μM along the subarctic–subtropical boundary and 18 μM off the north coast of Japan, with April to September new production rates in the subarctic Pacific being estimated at 2 and 6 moles C m⁻². Changes in nutrient removal in the Bering and western subarctic Pacific (WSP) suggest either the summer mixed layer is thinning with little change in new production or new production is increasing which would require an increase in iron transport to these high-nutrient low-chlorophyll (HNLC) waters. Si/N and N/P removal ratios of 2.1 and 19.7 are sufficient to push waters into Si then N limitation with sufficient iron supply. Because ~3 μM winter nitrate is transferred to reduced N pools in summer, new production calculated from seasonal nutrient drawdown should not be directly equated to export production.     

Distribution of Trace Bioelements in the Subarctic North Pacific Ocean and the Bering Sea (the R/V Hakuho Maru Cruise KH-97-2)

A column concentration-high resolution inductively coupled plasma mass spectrometry (ICP-MS) determination was applied to measure the total dissolved concentrations of Fe, Co, Ni, Cu and Zn in seawater collected from the subarctic North Pacific (~45°N) and the Bering Sea in July–September 1997. Total adsorbable Mn was determined on board by column electrolysis preconcentration and chemiluminescence detection. The vertical profiles for Fe, Ni and Zn were nutrient-like. The deep water concentration of Fe was ~0.5 nM in the northeast Pacific (18°-140°W) and increased to ~1 nM in the northwest Pacific (161°E) and ~2 nM in the Bering Sea (57°N, 180°E). The deep water concentrations for Ni and Zn in the Bering Sea were also 1.3–2 times higher than in the North Pacific. The profiles for Co and Cu were examined in the subarctic North Pacific, and results obtained were consistent with previous reports. There was a significant correlation between the concentrations of Co and Mn except for surface mixed layer. The profiles for total adsorbable Mn were similar to the reported profiles for total dissolvable Mn. The deep water concentration of Mn in the Bering Sea was also 4 times higher than in the North Pacific. Iron and zinc were depleted in surface water of the subarctic North Pacific. The relationship between these trace elements and nutrients suggests that these elements could be a limiting factor of phytoplankton productivity. In the Bering Sea, surface water contained ~0.3 nM of Fe. The Zn concentration, which was less than the detection limit in surface water, increased at shallower depths (~30 m) compared with the subarctic North Pacific. These results imply a higher flux of Fe and Zn to surface water in the Bering Sea. This in turn may cause the ecosystem in the Bering Sea characterized by a dominance of diatoms and high regenerated production.     

Basin-Scale Geographic Patterns of Bacterioplankton Biomass and Production in the Subarctic Pacific, July–September 1997

Bacterial biomass and production rate were measured in the surface (0–100 m) and mesopelagic layers (100–1,000 m) in the subarctic Pacific and the Bering Sea between July–September, 1997. Depth profiles were determined at stations occupied in oceanic domains including the subarctic gyres (western, Bering Sea, and Gulf of Alaska) and a boundary region south of the gyres. In the surface layer (0–100 m), both bacterial biomass and production were generally high in the western and Bering Sea gyres, with the tendency of decrease toward east. This geographic pattern was consistent with the dominant regime of phytoplankton biomass at the time of our survey. A significant portion of variation in bacterial production was explained by the concentration of chlorophyll a (r ² = 0.340, n = 60, P < 0.001) and, to the greater extent, by the concentration of semilabile total organic carbon (SL-TOC = TOC at a given depth—TOC at 1,000 m, r ² = 0.488, n = 59, P < 0.0001). Temperature significantly improved the regression model: temperature and chlorophyll jointly explained 60% of variation in bacterial production. These results support the hypothesis that bacteiral growth is largely regulated by the combination of temperature and the supply of dissolved organic carbon in subarctic surface waters. In the mesopelagic layer (100–1,000 m), the geographic pattern of bacterial production was strikingly different from the surface phytoplankton distribution: the production was high in the boundary region where the phytoplankton biomass was lowest. Bacterial growth appeared to be largely controlled by the supply of organic carbon, as indicated by the strong dependency of bacterial production on SL-TOC (r ² = 0.753, n = 75, P < 0.0001). The spatial uncoupling between surface phytoplankton and mesopelagic bacterial production suggests that the supply rate of labile dissolved organic carbon in the mesopelagic zone does not simply reflect the magnitude of the particulate organic carbon flux in the subarctic Pacific.     

Differences in the structure of the lower trophic levels of pelagic ecosystems in the eastern and western subarctic Pacific

To illustrate areal differences in the structure of lower trophic levels of the pelagic ecosystems in the subarctic Pacific, data collected in the quasi-steady state summer/fall conditions were analysed for five areas, i.e. the Bering Basin, Western Subarctic Gyre, the area south of the Aleutians, the Gulf of Alaska, and the Oyashio Region. Average values of stock size of four components of the lower trophic levels showed a clear difference between areas with ranges of 7.5-fold for nitrate, 3.0 for chlorophyll a, 9.9 for microzooplankton, and 2.4 for mesozooplankton. Such differences were more striking when the structure of the lower trophic levels was expressed as a biomass pyramid. In the Gulf of Alaska, Western Subarctic Gyre, and south of the Aleutians, the relative biomass of microzooplankton to phytoplankton is large and large amounts of nitrate remained unused. In addition to possible iron limitation, grazing control by the microzooplankton on small phytoplankton must be substantial in these areas. Conversely, in the Oyashio Region, the nitrate stock is very small indicating higher efficiency of nitrate consumption by phytoplankton. However, since phytoplankton and zooplankton stocks are not particularly large, their products are likely to be transferred, also efficiently, to the higher trophic levels such as planktivorous pelagic fish. The situation in the Bering Basin is intermediate between the Oyashio Region and the other three areas. Inter-annual fluctuations in stock size of the planktivorous fish which migrate into the Oyashio Region in summer/fall were quite large. However, the inter-annual variation of mesozooplankton biomass was small, suggesting the existence of certain mechanisms to stabilize plankton abundance under increasing predation pressure. As a result, the increasing fish stocks likely keep the transfer efficiency from nitrate through to fish higher, at least in the Oyashio Region.     

Standing Crops of Planktonic Ciliates and Copepod Nauplii in the Subarctic North Pacific and the Bering Sea in Summer

Abundances and biomasses of planktonic ciliates and copepod nauplii, major components of the microzooplankton community, were investigated in the subarctic North Pacific and the Bering Sea in summer of 1997. Their regional variation was illustrated by demarcating the entire area into five regions. Ciliates always predominated both in abundance (>94%) and biomass (>78%) over nauplii. Regional means of ciliates in the water column were higher in the Alaskan Gyre (120 × 10⁶ cells/m²) and the Western Subarctic Gyre (110 × 10⁶ cells/m²) in terms of abundance, and rich in the Bering Sea Gyre (360 mgC/m²) and the Western Subarctic Gyre (340 mgC/m²) in terms of biomass. By contrast, standing crops of ciliates were poor in the Oyashio Region (67 × 10⁶ cells/m²; 170 mgC/m²) and the Transition Region (64 × 10⁶ cells/m²; 160 mgC/m²). The values of biomass reported here are generally in agreement with the values reported previously from the Bering Sea Gyre and the Alaskan Gyre but are considerably higher than the previous value found in the Western Subarctic Gyre. No significant correlations could be found between chlorophyll a crop and standing crops of ciliates and copepod nauplii over the entire subarctic North Pacific and the Bering Sea during this summer.     

Mesopelagic fishes of the Bering Sea and adjacent northern North Pacific Ocean

Fourteen midwater trawl collections to depths of 450 m to 1,400 m were taken at eleven stations in the Bering Sea and adjoining regions of the northern North Pacific by the R/V Hakuho Maru during the summer of 1975. A total of 29 kinds of fishes were identified. Mesopelagic fishes of the families Myctophidae, Gonostomatidae and Bathylagidae predominated in the catches, contributing 14 species (94%) of the fishes caught.Seventeen species of fishes were caught in the Bering Sea, and all of these are known from nearby areas. The mesopelagic fish fauna of the Bering Sea is similar to that in adjoining regions of the northern North Pacific Ocean: endemic species are rare or absent. Stenobrachius nannochir was usually the most common mesopelagic fish in our catches.Stenobrachius leucopsarus is a diel vertical migrant that is usually the dominant mesopelagic fish in modified Subarctic waters of the northeastern Pacific. The change in dominance fromS. nannochir in the western Bering Sea toS. leucopsarus in the eastern Bering Sea is related to differences in oceanographic conditions.     

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.     

A 9-year time-series of planktonic foraminifer fluxes and environmental change in the Bering sea and the central subarctic Pacific Ocean, 1990–1999

A 9-year study of planktonic foraminifer fluxes was conducted in the Bering Sea (Station AB) and in the central subarctic Pacific (Station SA). Results clearly reflected variations of the water mass characteristics in the upper layers. The 9-year means of total foraminifer fluxes were the same (1400 shells m⁻² d⁻¹) at both stations. However, total foraminifer flux at Station AB tended to show its primary maximum during fall (October–December) and its secondary maximum in spring (April–June), whereas the primary maximum appeared in spring and the secondary maximum in fall at Station SA. Seasonal variation was more apparent at hemipelagic Station AB than at pelagic Station SA. Planktonic foraminifers found at both stations were of six species: Neogloboquadrina pachyderma, Globigerina umbilicata, Globigerinita glutinata, Globigerina quinqueloba, Globorotalia wilesi, and Orbulina universa. The foraminifer assemblages at the two stations reflected the temperature difference in the surface waters. The variable %G. umbilicata tended to be high in the warm surface waters during the summers. The temporal and geographical variation of %G. quinqueloba indicated that this taxon prefers regions with relatively low diatom fluxes. A notable appearance of O. universa occurred in 1997 at Station SA. During this period, other measured biogenic particle fluxes, such as those of diatoms, were low. This unusual 1997 event may be a reflection of global climatic change that happened to be observed in the central subarctic Pacific Ocean.     

Seasonal and Interannual Variability in the Distribution of Surface Nutrients and Dissolved Inorganic Carbon in the Northern North Pacific: Influence of El Niño

The seasonal and interannual changes in surface nutrients, dissolved inorganic carbon (DIC) and total alkalinity (TA) were recorded in the North Pacific (30–54°N) from 1995 to 2001. This study focuses on the region north of the subarctic boundary (∼40°N) where there was extensive monthly coverage of surface properties. The nutrient cycles showed large interannual variations in the eastern and western subarctic gyres. In the Alaska Gyre the seasonal depletion of nitrate (ΔNO₃) increased from 8–14 μmol kg⁻¹ in 1995–1999 to 21.5 μmol kg⁻¹ in 2000. In the western subarctic the shifts were similar in amplitude but more frequent. The large ΔNO₃ levels were associated with high silicate depletions, indicating enhanced diatom production. The seasonal DIC:NO₃ drawdown ratios were elevated in the eastern and central subarctic due to calcification. In the western subarctic and the central Bering Sea calcification was significant only during 1997 and/or 1998, two El Ni?o years. Regional C/N stoichiometric molar ratios of 5.7 to 7.0 (>40°N) were determined based on the years with negligible or no calcification. The annual new production (NP_a) based on ΔNO₃ and these C/N ratios showed large interannual variations. NP_a was usually higher in the western than in the eastern subarctic. However, values of 84 gC m⁻²yr⁻¹ were found in the Alaska Gyre in 2000 which is similar to that in the most productive provinces of the northern North Pacific. There were also large increases in NP_a around the Alaska Peninsula in 1997 and 1998. Finally, the net removal of carbon by the biological pump was estimated as 0.72 Gt C yr⁻¹ in the North Pacific (>30°N). This revised version was published online in August 2006 with corrections to the Cover Date.     

Impacts of elevated CO2 on particulate and dissolved organic matter production: microcosm experiments using iron-deficient plankton communities in open subarctic waters

Response of phytoplankton to increasing CO₂ in seawater in terms of physiology and ecology is key to predicting changes in marine ecosystems. However, responses of natural plankton communities especially in the open ocean to higher CO₂ levels have not been fully examined. We conducted CO₂ manipulation experiments in the Bering Sea and the central subarctic Pacific, known as high nutrient and low chlorophyll regions, in summer 2007 to investigate the response of organic matter production in iron-deficient plankton communities to CO₂ increases. During the 14-day incubations of surface waters with natural plankton assemblages in microcosms under multiple pCO₂ levels, the dynamics of particulate organic carbon (POC) and nitrogen (PN), and dissolved organic carbon (DOC) and phosphorus (DOP) were examined with the plankton community compositions. In the Bering site, net production of POC, PN, and DOP relative to net chlorophyll-a production decreased with increasing pCO₂. While net produced POC:PN did not show any CO₂-related variations, net produced DOC:DOP increased with increasing pCO₂. On the other hand, no apparent trends for these parameters were observed in the Pacific site. The contrasting results observed were probably due to the different plankton community compositions between the two sites, with plankton biomass dominated by large-sized diatoms in the Bering Sea versus ultra-eukaryotes in the Pacific Ocean. We conclude that the quantity and quality of the production of particulate and dissolved organic matter may be altered under future elevated CO₂ environments in some iron-deficient ecosystems, while the impacts may be negligible in some systems.     

Relationship between phytoplankton composition and environmental factors in the surface waters of southern South China Sea in early summer of 2009

Horizontal distributions of phytoplankton abundance,species composition as well as environmental factors were investigated in the surface waters of southern South China Sea(SCS) in early summer(May 16 to June 7) of 2009.In most areas of southern SCS,the concentrations of nitrogen and phosphorus were very low and DIN/DIP ratios usually were lower than the Redfield N/P Ratio of 16:1.Nitrogen nutrients were significant lower in the upwelling region off Vietnam.A total of 144 taxa of phytoplankton were identified in the study area.The dominant genera were Prorocentrum,Gonyaulax,Gyrodinium,Scrippsiella and Chaetoceros,respectively.Spatial patterns of early-summer phytoplankton abundance and species composition should be mainly controlled by the upwelling off Vietnam and the discharge of Mekong River in the southern SCS.Diatoms dominated in the nutritious Mekong River Estuary or upwelling region off Vietnam;while dinoflagellates dominated in the oligotrophic pelagic region.The canonical correspondence analysis(CCA) indicates that most of diatoms favor higher levels of silicate and phosphorus,as well as lower temperature;while most of dinoflagellates favor the lower silicate and phosphorous and higher temperature.Correlation and CCA results indicate that silicate,nitrate and temperature were the most relevant environmental factors to regulate the horizontal pattern of early-summer phytoplankton in the surface waters of southern SCS.     

2008年夏季白令海营养盐的分布及其结构状况

中国第3次北极考察对白令海营养盐的分布及结构状况进行了观测分析,结果表明,白令海营养盐分布和结构状况区域性特征明显。海盆区表层DIN、磷酸盐和硅酸盐平均浓度分别为9.73,0.94,11.06 μmol/dm3;陆架区表层DIN,磷酸盐和硅酸盐平均浓度分别为0.60, 0.43, 3.74 μmol/dm3。营养盐高值主要出现在白令海西南部的海盆区和海峡口西南侧水域,低值出现于陆架边缘的陆坡区和陆架东部水域。白令海盆区真光层DIN,磷酸盐、硅酸盐浓度普遍较高,叶绿素浓度则较低,具有典型的高营养盐、低叶绿素(HNLC)特征。海盆区生物作用不是营养盐空间分布的主要调控因子,而陆架区营养盐的分布变化不仅受控于物理海洋输运过程的变化,同时也受夏季浮游生物生长、营养盐吸收消耗所影响。陆架和陆坡区表层海水N/P,Si/P比值平均分别为1.8, 9.9和3.2, 2.2,呈明显的低N/P,Si/P比值结构特征,陆坡区缺硅明显,陆架区缺氮显著。在白令海水域磷酸盐浓度普遍较高,它不可能成为浮游植物光合作用限制因子。受硅限制水域主要限于陆坡区硅藻大量繁殖时期,属偶然性限制,在白令海陆架区绝大部分水域主要表现为氮限制。     

Long-term biogenic particle fluxes in the Bering Sea and the central subarctic Pacific Ocean, 1990–1995

Time-series sediment traps were deployed for five consecutive years in two distinctively different subarctic marine environments. The centrally located subarctic pelagic Station SA (49°N, 174°W; water depth 5406 m) was simultaneously studied along with the marginal sea Station AB (53.5°N, 177°W; water depth 3788 m) in the Aleutian Basin of the Bering Sea. A mooring system was tethered to the sea-floor with a PARFLUX type trap with 13 sample bottles, which was placed at 600 m above the sea-floor at each of the two stations. Sampling intervals were synchronized at the stations, and they were generally set for 20 days during highly productive seasons, spring through fall, and 56 days during winter months of low productivity. Total mass fluxes, which consisted of mainly biogenic phases, were significantly greater at the marginal sea Station AB than at the pelagic Station SA for the first four years and moderately greater for the last year of the observations. This reflects the generally recognized higher productivity in the Bering Sea. Temporal excursion patterns of the mass fluxes at the two stations generally were in parallel, implying that temporal changes in their biological productivity are strongly governed by a large-scale seasonal climatic variability over the region rather than local phenomena. The primary reason for the difference in total mass flux at the two stations stems mainly from varying contributions of siliceous and calcareous planktonic assemblages. A significantly higher opal contribution at Station AB than at Station SA was mainly due to diatoms. Diatom fluxes at the marginal sea station were about twice those observed at the pelagic station, resulting in a very high opal contribution at Station AB. In contrast to the opal fluxes, CaCO₃ fluxes at Station AB were slightly lower than at Station SA. The ratios of C_org/C_inorg were usually significantly greater than one in both regions, suggesting that preferentially greater organic carbon from cytoplasm than skeletal inorganic carbon was exported from the surface layers. Such a process, known as the biological pump, leads to a carbon sink which effectively lowers p CO₂ in the surface layers and then allows a net flux of atmospheric CO₂ into the surface layer. The efficiency of the biological pump is greater in the Bering Sea than at the open-ocean station.     

Comparison of factors controlling phytoplankton productivity in the NE and NW subarctic Pacific gyres

The subarctic North Pacific is one of the three major high nitrate low chlorophyll (HNLC) regions of the world. The two gyres, the NE and the NW subarctic Pacific gyres dominate this region; the NE subarctic Pacific gyre is also known as the Alaska Gyre. The NE subarctic Pacific has one of the longest time series of any open ocean station, primarily as a result of the biological sampling that began in 1956 on the weathership stationed at Stn P (50°N, 145°W; also known as Ocean Station Papa (OSP)). Sampling along Line P, a transect from the coast (south end of Vancouver Island) out to Stn P has provided valuable information on how various parameters change along this coastal to open ocean gradient. The NW subarctic Pacific gyre has been less well studied than the NE gyre. This review focuses mainly on the NE gyre because of the large and long term data set available, but makes a brief comparison with the NW gyre. The NE gyre has saturating NO₃ concentrations all year (winter = about 16 μM and summer = about 8 μM), constantly very low chlorophyll (chl) (usually <0.5 mg m⁻³) which is dominated by small cells (<5 μm). Primary productivity is low (about 300–600 mg C m⁻² d⁻¹ and varies little (2 times) seasonally. Annual primary productivity is 3 to 4 times higher than earlier estimates ranging from 140 to 215 g C m⁻² y⁻¹. Iron limits the utilization of nitrate and hence the primary productivity of large cells (especially diatoms) except in the winter when iron and light may be co-limiting. There are observations of episodic increases in chl above 1 mg m⁻³, suggesting episodic iron inputs, most likely from Asian dust in the spring/early summer, but possibly from horizontal advection from the Alaskan Gyre in summer/early fall. The small cells normally dominate the phytoplankton biomass and productivity, and utilize the ammonium produced by the micrograzers. They do not appear to be Fe-limited, but are controlled by microzooplankton grazers. The NW Subarctic Gyre has higher nutrient concentrations and a shallower summer mixed depth and photic zone than Stn P in the NE gyre. Chl concentrations tend to be higher (0.5 to 1.5 μg L⁻¹) than Stn P, but primary productivity in the summer is similar to Stn P (600 mg C m⁻² d⁻¹). There are no seasonal data from this gyre. Iron enrichment experiments in October, resulted in an increase in chl (mainly the centric diatom Thalassiosira sp.) and a draw down of nitrate, suggesting that large phytoplankton are Fe-limited, similar to Stn P.     

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.     

白令海大型底栖动物群落结构及其空间分布格局

Due to its unique geological location, the Bering Sea is an ideal place to investigate the water exchange and ecosystem connectivity of the Pacific Ocean–Arctic Ocean and subarctic–Arctic region. Based on a number of summer surveys(July to September, 2010, 2012 and 2014), macrobenthic communities and their spatial-temporal patterns are exhibited for the majority of the Bering Sea(53°59′–64°36′N). The results show that the macrobenthic communities were dominated by northern cold-water species and immigrant eurythermic species, and the communities assumed a dispersed and patchy distribution pattern. Polychaetes(Scoloplos armiger), crustaceans(Ceradocus capensis) and sea urchins(Echinarachnius parma) were the main dominant groups in the shallow shelves; the sea star(Ctenodiscus crispatus) and the brittle star(Ophiura sarsii) were the main dominant groups in the continental slope; whereas small polychaetes(Prionospio malmgreni) dominated the basin area. Sediment type, water depth, and currents were the major factors affecting the structure and spatial distribution of the macrobenthic communities. Compared with other seas, the shallow areas of the Bering Sea showed an extremely high-standing biomass. In particular, the northern shelf area(north of St. Lawrence Islands and west of 170°W),which is primarily controlled by Anadyr Water, is an undersea oasis. In contrast, a deficiency in the downward transport of particulate organic carbon has resulted in a desert-like seabed in the basin area. By comparing our results to previous studies, we found that macrobenthic communities of the Bering Sea have undergone significant structural changes in recent decades, resulting in a decrease in abundance and an increase in biomass.In addition, populations of amphipods and bivalves in the northern shelves have decreased significantly and have been gradually replaced by other species. These changes might be associated with advanced seasonal ice melting,changes in organic carbon input, and global warming, indicating that large-scale ecosystem changes have been occurring in the Bering Sea.     

Lethality of increasing CO2 levels on deep-sea copepods in the western North Pacific

The first CO₂ exposure experiments on several species of pelagic copepods inhabiting surface and deep layers in the western North Pacific were conducted. Living organisms were collected from two layers between the surface and 1,500 m between latitudes of 11 and 44°N, and they were exposed aboard ship to various pCO₂ up to about 98,000 μatm. Mortality of copepods from both shallow and deep layers in subarctic to subtropical regions increased with increasing pCO₂ and exposure time. Deep-living copepods showed higher tolerance to pCO₂ than shallow-living copepods. Furthermore, deep-living copepods from subarctic and transitional regions had higher tolerances than the subtropical copepods. The higher tolerances of the deep-living copepods from subarctic and transitional regions may be due to the adaptation to the natural pCO₂ conditions in the subarctic ocean.     

An Ecosystem Model Coupled with Nitrogen-Silicon-Carbon Cycles Applied to Station A7 in the Northwestern Pacific

A model based on that of Kishi et al. (2001) has been extended to 15 compartments including silicon and carbon cycles. This model was applied to Station A7 off Hokkaido, Japan, in the Northwestern Pacific. The model successfully simulated the observations of: 1. a spring bloom of diatoms; 2. large seasonal variations of nitrate and silicate concentrations in the surface water; and 3. large inter-annual variations in chlorophyll-a. It also reproduced the observed features of the seasonal variations of carbon dioxide partial pressure (pCO₂)—a peak in pCO₂ in winter resulting from deep winter convection, a rapid decrease in pCO₂ as a result of the spring bloom, and an almost constant pCO₂ from summer through fall (when the effect of increasing temperature cancels the effect of biological production). A comparison of cases with and without silicate limitation shows that including silicate limitation in the model results in: 1. decreased production by diatoms during summer; and 2. a transition in the dominant phytoplankton species, from diatoms to other species that do not take up silicate. Both of these phenomena are observed at Station A7, and our results support the hypothesis that they are caused by silicate limitation of diatom growth. This revised version was published online in July 2006 with corrections to the Cover Date.     

白令海颗粒有机物碳、氮同位素组成及其生物地球化学意义

Stable carbon and nitrogen isotopic composition of particulate organic matter(POM) were measured for samples collected from the Bering Sea in 2010 summer. Particulate organic carbon(POC) and particulate nitrogen(PN) showed high concentrations in the shelf and slope regions and decreased with depth in the slope and basin, indicating that biological processes play an important role on POM distribution. The low C/N ratio and heavy isotopic composition of POM, compared to those from the Alaska River, suggested a predominant contribution of marine biogenic organic matter in the Bering Sea. The fact that δ13C and δ15N generally increased with depth in the Bering Sea basin demonstrated that organic components with light carbon or nitrogen were decomposed preferentially during their transport to deep water. However, the high δ13C and δ15N observed in shelf bottom water were mostly resulted from sediment resuspension.