Seasonal variability in carbon demand and flux by mesozooplankton communities at subarctic and subtropical sites in the western North Pacific Ocean

We investigated seasonal changes in carbon demand and flux by mesozooplankton communities at subtropical (S1) and subarctic sites (K2) in the western North Pacific Ocean to compare the impact of mesozooplankton communities on the carbon budget in surface and mesopelagic layers. Fecal pellet fluxes were one order higher at K2 than at S1, and seemed to be enhanced by copepod and euphausiid egestion under high chlorophyll a concentrations. The decrease in pellet volume and the lack of any substantial change in shape composition during sink suggest a decline in fecal pellet flux due to coprorhexy and coprophagy. While respiratory and excretory carbon by diel migrants at depth (i.e., active carbon flux) was similar between the two sites, the actively transported carbon exceeded sinking fecal pellets at S1. Mesozooplankton carbon demand in surface and mesopelagic layers was higher at K2 than S1, and an excess of demand to primary production and sinking POC flux was found during some seasons at K2. We propose that this demand was met by supplementary carbon sources such as feeding on protozoans and fecal pellets at the surface and carnivory of migrants at mesopelagic depths.     

Impact of mesoscale eddies on particulate organic carbon flux in the western subarctic North Pacific

The mechanism that controls particulate organic carbon (POC) flux in the deep sea differs depending on the season and sea. The POC produced in the western subarctic North Pacific are known to be transported to the deep sea efficiently, but the direct relationship between the POC flux and physical processes is still unclear. In this study, we evaluated the effect of mesoscale eddies on POC flux in the western subarctic North Pacific. The seasonal and interannual variabilities of POC flux were investigated using data from a time-series sediment trap deployed at 4810 m at station K2 (47°N, 160°E) from 2005 to 2018. POC flux was high during May–November, appearing to reflect spring and fall blooms at the ocean surface. POC flux also showed interannual variability, with twelve peaks that were mostly affected by enhanced bloom just before the peak. Nine peaks of the twelve peaks were affected by mesoscale eddies, which enhanced bloom around K2 by extending the area with a high chlorophyll-a concentration along the coastal region into the offshore region, suggesting that mesoscale eddies strongly impact the interannual variability of POC flux at K2.

     

Enhancement of particulate organic carbon export flux induced by atmospheric forcing in the subtropical oligotrophic northwest Pacific Ocean

The effects of extreme atmospheric forcing on the export flux of particulate organic carbon (POC) in the warm oligotrophic nitrogen-limited northwest Pacific Ocean were examined in 2007 during the spring Asian dust storm period. Several strong northeast monsoon events (maximum sustained wind speeds approaching 16.7 m s^? 1, and gusts up to 19.0 m s^? 1) accompanied by dust storms occurred during a 1-month period. The cold stormy events decreased surface water temperature and induced strong wind-driven vertical mixing of the water column, resulting in nutrient entrainment into the mixed layer from subsurface waters. As a result, the export flux of POC ranged from 49 to 98 (average value = 71 ± 16) mg m^? 2 day^? 1, approximately 2–3 times greater than average values in other seasons. As dry and wet deposition of nitrogen attributable to Asian dust storm events does not account for the associated increases in POC stocks in this N-limited oligotrophic oceanic region, the enhancement of POC flux must have been caused by nutrient entrainment from subsurface waters because of the high winds accompanying the dust storm events.     

Feeding ecology of mesopelagic zooplankton of the subtropical and subarctic North Pacific Ocean determined with fatty acid biomarkers

Mesopelagic zooplankton may meet their nutritional and metabolic requirements in a number of ways including consumption of sinking particles, carnivory, and vertical migration. How these feeding modes change with depth or location, however, is poorly known. We analyzed fatty acid (FA) profiles to characterize zooplankton diet and large particle (>51 μm) composition in the mesopelagic zone (base of euphotic zone ?1000 m) at two contrasting time-series sites in the subarctic (station K2) and subtropical (station ALOHA) Pacific Ocean. Total FA concentration was 15.5 times higher in zooplankton tissue at K2, largely due to FA storage by seasonal vertical migrators such as Neocalanus and Eucalanus. FA biomarkers specific to herbivory implied a higher plant-derived food source at mesotrophic K2 than at oligotrophic ALOHA. Zooplankton FA biomarkers specific to dinoflagellates and diatoms indicated that diatoms, and to a lesser extent, dinoflagellates were important food sources at K2. At ALOHA, dinoflagellate FAs were more prominent. Bacteria-specific FA biomarkers in zooplankton tissue were used as an indicator of particle feeding, and peaks were recorded at depths where known particle feeders were present at ALOHA (e.g., ostracods at 100–300 m). In contrast, depth profiles of bacterial FA were relatively constant with depth at K2. Diatom, dinoflagellate, and bacterial biomarkers were found in similar proportions in both zooplankton and particles with depth at both locations, providing additional evidence that mesopelagic zooplankton consume sinking particles. Carnivory indices were higher and increased significantly with depth at ALOHA, and exhibited distinct peaks at K2, representing an increase in dependence on other zooplankton for food in deep waters. Our results indicate that feeding ecology changes with depth as well as by location. These changes in zooplankton feeding ecology from the surface through the mesopelagic zone, and between contrasting environments, have important consequences for the quality and quantity of organic material available to deeper pelagic and benthic food webs, and for organic matter sequestration.     

Seasonal changes in the mesozooplankton biomass and community structure in subarctic and subtropical time-series stations in the western North Pacific

Seasonal changes in mesozooplankton biomass and their community structures were observed at time-series stations K2 (subarctic) and S1 (subtropical) in the western North Pacific Ocean. At K2, the maximum biomass was observed during the spring when primary productivity was still low. The annual mean biomasses in the euphotic and 200- to 1000-m layers were 1.39 (day) and 2.49 (night) g C m^?2 and 4.00 (day) and 3.63 (night) g C m^?2, respectively. Mesozooplankton vertical distribution was bimodal and mesopelagic peak was observed in a 200- to 300-m layer; it mainly comprised dormant copepods. Copepods predominated in most sampling layers, but euphausiids were dominant at the surface during the night. At S1, the maximum biomass was observed during the spring and the peak timing of biomass followed those of chlorophyll a and primary productivity. The annual mean biomasses in the euphotic and 200- to 1000-m layers were 0.10 (day) and 0.21 (night) g C m^?2 and 0.47 (day) and 0.26 (night) g C m^?2, respectively. Copepods were dominant in most sampling layers, but their mean proportion was lower than that in K2. Mesozooplankton community characteristics at both sites were compared with those at other time-series stations in the North Pacific and with each other. The annual mean primary productivities and sinking POC fluxes were equivalent at both sites; however, mesozooplankton biomasses were higher at K2 than at S1. The difference of biomasses was probably caused by differences of individual carbon losses, population turnover rates, and trophic structures of communities between the two sites.     

The diffusive component of particulate organic carbon export in the North Atlantic estimated from SeaWiFS ocean color

The diffusive component of the particulate organic carbon (POC) export from the ocean's surface layer has been estimated using a combination of the mixed layer model and SeaWiFS ocean color data. The calculations were carried out for several example sites located in the North Atlantic over a 10-year time period (1998–2007). Satellite estimates of surface POC derived from ocean color were applied as an input to the model driven by local surface heat and momentum fluxes. For each year of the examined period, the diffusive POC flux was estimated at a 200 m depth. The highest flux is generally observed in the spring and fall seasons, when surface waters are weakly stratified. In addition, the model results demonstrate significant interannual and geographical variability of the flux. The highest diffusive POC flux occurs in the northern North Atlantic and the lowest in the subtropical region. The interannual variability of the diffusive POC flux is associated with mixed layer dynamics and underscores the importance of atmospheric forcing for POC export from the surface layer to the ocean's interior.     

Splitting of the subtropical gyre in the western North Pacific

Examined here is a hypothetical idea of the splitting of the subtropical gyre in the western North Pacific on the basis of two independent sources of data,i.e., the long-term mean geopotential-anomaly data compiled by the Japanese Oceanographic Data Center and the synoptic hydrographic (STD) data taken by the Hakuho Maru in the source region of the Kuroshio and the Subtropical Countercurrent in the period February and March 1974. Both of the synoptic and the long-term mean dynamic-topographic maps reveal three major ridges, which indicate that the western subtropical gyre is split into three subgyres. Each subgyre is made up of the pair of currents, the Kuroshio and the Kuroshio Countercurrent, the Subtropical Countercurrent and a westward flow lying just south of the Countercurrent (18°N–21°N), and the northern part of the North Fquatorial Current and an eastward flow at around 18°N. The subgyres are more or less composed of a train of anticyclonic eddies with meridional scales of between 300 and 600 km, so that the volume transport of the subgyres varies by a factor of two or more from section to section. The upper-water characteristics also support the splitting of the subtropical gyre; the water characteristics are fairly uniform within each subgyre, but markedly different between them. The northern rim of each subgyre appears as a sharp density front accompanied by an eastward flow. The bifurcations of the sharp density fronts across the western boundary current indicate that the major part of the surface waters in the North Equatorial Countercurrent is not brought into the Kuroshio. The western boundary current appears as a continuous feature of high speed, but the waters transported change discontinuously at some places.     

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.     

Light scattering and size distribution of particles in the surface waters of the North Pacific Ocean

The volume scattering function and size distribution of suspended particles in the surface water were determined in the North Pacific. The relationship between the scattering coefficient estimated from observed volume scattering function and cross-section concentration of the particles greater than 2.4m in diameter was found to be linear in both northern and southern regions of central North Pacific. Difference in the constant of proportionality between two regions, however, was very great. Moreover the constant in the southern region was too large compared with the values obtained by the Mie theory. This is considered to be due to the fact that particles smaller than 2.4m which were not measureable by the Coulter Counter, were neglected in the calculation of cross-section concentration. If small particles are taken into consideration, total cross-section concentration and scattering coefficient in the two regions tend to follow a linear relation. From the correlation between the scattering coefficient computed from size distribution and the volume scattering function, the refractive index of particles was estimated to be 1.03–1.05. By the same procedure, the refractive index of particles in Tateyama Harbour where the water was very turbid, was estimated to be also 1.03–1.05. This is in contrast to the result for the refractive index of particles originating from the river which flows into the harbour. This index was found to be 1.10–1.20.     

Comparison of community structures between particle-associated and free-living prokaryotes in tropical and subtropical Pacific Ocean surface waters

The subtropical and tropical regions of the Pacific Ocean are less productive than other oceanic regions. Although particle association should be an important strategy for heterotrophic prokaryotes to survive in such environments, we have little information on particle-associated (PA) prokaryotes in these regions. The specific aim of this study was to determine bacterial and archaeal community structures in the PA assemblage in comparison to the free-living (FL) assemblage in the North Pacific Subtropical Gyre, the South Pacific Subtropical Gyre, and an eastern equatorial region of the Pacific Ocean. Community profiles and phylogenetic identities were obtained by denaturing gradient gel electrophoresis, 454-pyrosequencing, and cloning followed by Sanger sequencing of 16Sr RNA gene amplicons. The distribution patterns of some abundant groups in three regions and two lifestyles (PA and FL) are shown in this study. Also, the PA community structures of bacteria differed from the FL ones and exhibited higher diversity than the FL ones, while the archaeal community structures did not show significant differences between PA and FL assemblages. We found that specific phylotypes of Gammaproteobacteria and Flavobacteria were abundant in PA bacterial assemblages, suggesting that they prefer to attach and consume particulate organic matter. In summary, the surface seawater PA assemblages represent very different bacterial and archaeal community structures between three different oceanic regions, each of which had distinct PA and FL community structures. These results imply that environmental factors determine microbial community structures.     

Impacts of the wintertime mesozooplankton community to downward carbon flux in the subarctic and subtropical Pacific Oceans

We compared wintertime depth distributions of the mesozooplankton community and dominant copepods between the subtropical (S1) and subarctic (K2) Pacific Oceans to evaluate the relative importance of actively transported carbon by vertical migrants to sinking particulate organic carbon flux. Primary production was higher and the ratio of sinking particulate organic carbon flux to primary production was lower at S1 compared with those at K2. The mesozooplankton community was lower in abundance and biomass at S1 compared to K2. Copepods were the dominant group among both mesozooplankton abundance and biomass throughout the water column down to 1000 m at both sites. The depth distribution showed that diel vertical migration was obvious for the mesozooplankton abundance and biomass at S1 but was not apparent for the abundance at K2, because the dominant component was diurnally migrating species at S1 and overwintering species residing at mesopelagic depths at K2. The major components of diel migrants were copepods and euphausiids at S1 and only euphausiids at K2. Respiratory flux by the diurnally migrating mesozooplankton was estimated to be 2 mgC m⁻² day⁻¹ at S1 and 7 mgC m⁻² day⁻¹ at K2. The respiratory flux was equivalent to 131% of sedimentary fecal pellet flux at S1 and 136% of that at K2. Because pathways of downward carbon flux are facilitated by the mesozooplankton community, the actively transported carbon (respiration of dissolved inorganic carbon, excretion of dissolved organic carbon and egestion of fecal pellets at depth) might be larger during winter than the flux of sinking fecal pellets.     

Distributions of particulate and dissolved organic and inorganic phosphorus in North Pacific surface waters

Several operationally defined fractions of phosphorus (P) were measured along a surface water transect in the North Pacific. The P content in all fractions was found to increase northward from the edge of the subtropical to the subarctic region. Particulate organic P (POP) concentrations increased from 9 to 110 nmol L^− 1, whereas the particulate inorganic P (PIP) concentrations increased from 1 to 13 nmol L^− 1. A significant correlation between POP, PIP and chlorophyll a suggested that these P pools are associated directly or indirectly with phytoplankton cells. PIP comprised 10–20% of the total particulate P pool across the transect, indicating it is an important component of the marine P cycle in this region. Dissolved non-reactive P (assumed to consist predominantly of non-reactive organic P compounds, thus referred to as DOP) concentration increased from 0.10 to 0.22 μmol L^− 1, whereas soluble reactive P (SRP) concentration increased from 0.01 to 1.42 μmol L^− 1 along the transect. The proportion of DOP and SRP varied widely, with a large proportion of DOP in areas with low total dissolved P concentrations in lower latitudes and a large proportion of SRP in areas with high total dissolved P concentrations in higher latitudes. High demand for DOP in the lower latitudinal region would diminish the concentration of this pool relative to higher latitudinal regions where SRP is more abundant and would be preferentially utilized. The availability of SRP could have a significant impact on the concentration and probably on the composition of DOP. We show that P fractionation provides an important insight for discussing the marine P cycle.     

北太平洋西部和印度洋东部及其邻近海域表层水体中137Cs的分布

对北太平洋西部海域、苏禄海及印尼海、中国南海、印度洋东部海域、孟加拉湾及安达曼海等表层水体中放射性核素137Cs的活度进行了测定。结果表明,上述海域表层水体中137Cs活度显示了较大的变化范围,最低值出现在南极附近的南大洋(1.1Bqm-3),较高的活度值则出现在北太平洋西部海域及中国南海(3Bqm-3)。在所研究水域范围内,137Cs活度的纬度分布特征并没有完全有效地反映出137Cs的全球理论大气沉降趋势及其纬度效应。综合本研究及Miyake等人(1988)的测定结果,我们计算出137Cs自表层海水中的析出速率在苏禄海及印尼海约为0.016/a,在孟加拉湾及安达曼海约为0.033/a,在中国南海约为0.029/a,这一结果明显低于西北太平洋日本沿海表层水体中137Cs的析出速率。这可能是因为在这些海域,横向及纵向的水体混合过程相对都较慢,而且颗粒物对137Cs的吸附析出过程也比较弱所致。     

Distribution of particulate organic carbon and nitrogen in the Bering Sea and northern North Pacific Ocean

Particulate matter was collected in the Bering Sea and the northern North Pacific Ocean during the cruise of R. V. Hakuho-maru, Ocean Research Institute of Tokyo University in summer of 1975. The particulate matter was analyzed for organic carbon and nitrogen, chlorophylla and amino acids.The concentrations of particulate organic carbon and nitrogen were measured with the range of 16–422gC l^–1 and 1–85gN l^–1, 19–186gC l^–1 and 1–26gN l^–1, 46–1,038gC l^–1 and 6–79gN l^–1 and 19–246gC l^–1 and 2–25gN l^–1 in the Oyashio, the Deep Bering Sea, the continental shelf of Bering Sea and the northern North Pacific, respectively. Particulate organic carbon and nitrogen decreased with depth throughout the areas. The average concentrations of organic carbon and nitrogen in the entire water column tended to decrease in the following order; the continental shelf > Oyashio > northern North Pacific > Deep Bering Sea.C/N of particulate matter varied in the range of 3–15 (7 on average) in surface waters throughout the areas and these values tended to increase with depth to 5–20 (11 on average) in deep waters without significant regional variability.Linear regressions between chlorophylla and particulate organic carbon in the euphotic layers indicate that detrital organic carbon accounted for 34.2, 44.9, 49.1 and 25.2 % of particulate organic carbon in the Oyashio, the Deep Bering Sea, the continental shelf and the northern North Pacific, respectively.Particulate amino acid was determined in the range of 10.3–78.0g l^–1, 104–156g l^–1 and 10.4–96.4g l^–1 in the Deep Bering Sea, the continental shelf and the northern North Pacific, respectively. Aspartic acid, glutamic acid, serine, glycine and alanine were found as dominant species of amino acid of particulate matter.     

Comparison of carbon cycle between the western Pacific subarctic and subtropical time-series stations: highlights of the K2S1 project

A comparative study of ecosystems and biogeochemistry at time-series stations in the subarctic gyre (K2) and subtropical region (S1) of the western North Pacific Ocean (K2S1 project) was conducted between 2010 and 2013 to collect essential data about the ecosystem and biological pump in each area and to provide a baseline of information for predicting changes in biologically mediated material cycles in the future. From seasonal chemical and biological observations, general oceanographic settings were verified and annual carbon budgets at both stations were determined. Annual mean of phytoplankton biomass and primary productivity at the oligotrophic station S1 were comparable to that at the eutrophic station K2. Based on chemical/physical observations and numerical simulations, the likely “missing nutrient source” was suggested to include regeneration, meso-scale eddy driven upwelling, meteorological events, and eolian inputs in addition to winter vertical mixing. Time-series observation of carbonate chemistry revealed that ocean acidification (OA) was ongoing at both stations, and that the rate of OA was faster at S1 than at K2 although OA at K2 is more critical for calcifying organisms.