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.     

Functional roles of interzonal migrating mesozooplankton in the western subarctic Pacific

Grazing experiments and production estimation based on life-history analysis of Neocalanus copepods (N. cristatus, N. plumchrus and N. flemingeri) were carried out in the Oyashio region to understand the carbon flows associated with the interzonal migrating copepods. These copepods, and also Eucalanus bungii, fed on nano- and micro-sized organisms non-selectively throughout the season. However, diatoms were the dominant food resource until May and organisms, such as ciliates were the major resource after May. Daily growth rate was estimated from the Ikeda–Motoda, Huntley–Lopez and Hirst–Sheader models. Since the growth rates were considered to be overestimates for the Huntley–Lopez model and underestimates for the other two models, we applied the weight-specific growth rates previously reported for these species in the Bering Shelf. Surface biomass of Neocalanus increased rapidly in June during the appearance of C5, and a successive increase of overwintering stock was evident in the deeper layer. The deep biomass decreased gradually from September to May during the dormant and reproduction period. N. cristatus has the largest annual mean biomass (2.3 gC m⁻²), followed by N. plumchrus (1.1) and N. flemingeri (0.4). Daily production rate of Neocalanus varied from 0.4 to 363.4 mgC m⁻² day⁻¹, to which N. cristatus was the largest contributor. Annual production was estimated as 11.5 gC m⁻² year⁻¹ for N. cristatus, 5.7 for N. plumchrus and 2.1 for N. flemingeri, yielding annual P/B ratio of 5 for each species. The annual production of Neocalanus accounted for 13.2% of the primary production in the Oyashio region. Their fecal pellets were estimated to account for 14.9% (0.7 gC m⁻² year⁻¹) of sinking flux of organic carbon at 1000-m depth. Moreover, their export flux by ontogenetic vertical migration, which is not measured by sediment trap observations, is estimated to be 91.5% (4.3 gC m⁻² year⁻¹) of carbon flux of sinking particles at 1000-m depth. These results suggest the important role of interzonal migrating copepods in the export flux of carbon.     

Geographical Variations in Prosome Length and Body Weight of Neocalanus Copepods in the North Pacific

Geographical variations in prosome length and body weight of Neocalanus copepods (N. cristatus, N. plumchrus and N. flemingeri) were investigated on samples from North-South and East-West transects in the North Pacific during spring to early summer in 1998 and 1999. Southward and eastward increasing patterns were pronounced for water temperature, although no significant pattern was observed for chlorophyll a concentrations. All Neocalanus species showed large geographical variations in prosome length and body weight, being smaller in the southern and eastern waters. Comparing the relationship between prosome length and body weight, large deviations (lower body weight at a given prosome length) were evident for the eastern specimens of N. cristatus and N. plumchrus. In stepwise regression analysis, the geographical variations of prosome length and body weight revealed a significantly negative correlation with temperature variations. These results suggest that temperature is a more important environmental factor than chlorophyll a concentration in its effect on geographical variations in prosome length and body weight of Neocalanus copepods in the North Pacific. This revised version was published online in July 2006 with corrections to the Cover Date.     

Seasonal variations in abundance,growth and mortality of heterotrophic bacteria in Kagoshima Bay

Seasonal changes in abundance, growth and mortality of heterotrophic bacteria were investigated monthly from collections and dilution experiments in Kagoshima Bay, the southernmost of Japan. Bacteria occurred abundantly with considerable variation in the surface layers where chlorophyll a concentrations were high, whereas seasonal variations were obscure below 100 m. Especially, bacteria showed a decline of cell density toward summer when heterotrophic nano-flagellates increased their abundance. Seasonal and vertical variations in bacterial cell number during the study period were positively correlated with those of water temperature and pico-sized chlorophyll a concentration. Maximum growth and mortality rates showed positive correlations with water temperature but no positive relationships to size-fractionated chlorophyll a. Net increase rate (i.e. in situ rate if abundance changes) was negatively correlated with cell density of heterotrophic nano-flagellates. It is concluded that bacterial abundance is controlled by the resultant (i.e. net growth rate) of the balance between maximum growth and predatory mortality by heterotrophic nano-flagelllates which are both dependent on ambient temperature.     

Biological organic carbon export estimated from the annual carbon budget observed in the surface waters of the western subarctic and subtropical North Pacific Ocean from 2004 to 2013

The annual flux of biologically produced organic carbon from surface waters is equivalent to annual net community production (NCP) at a steady state and equals the export of particulate and dissolved organic carbon (POC and DOC, respectively) to the ocean interior. NCP was estimated from carbon budgets of salinity-normalized dissolved inorganic carbon (nDIC) inventories at two time-series stations in the western subarctic (K2) and subtropical (S1) North Pacific Ocean. By using quasi-monthly biogeochemical observations from 2004 to 2013, monthly mean nDIC inventories were integrated from the surface to the annual maximum mixed layer depth and corrected for changes due to net air–sea CO₂ exchange, net CaCO₃ production, vertical diffusion from the upper thermocline, and horizontal advection. The annual organic carbon flux at K2 (1.49 ± 0.42 mol m^?2 year^?1) was lower than S1 (2.81 ± 0.53 mol m^?2 year^?1) (p < 0.001 based on t test). These fluxes consist of three components: vertically exported POC fluxes (K2: 1.43 mol m^?2 year^?1; S1: 2.49 mol m^?2 year^?1), vertical diffusive DOC fluxes (K2: 0.03 mol m^?2 year^?1; S1: 0.25 mol m^?2 year^?1), and suspended POC fluxes (K2: 0.03 mol m^?2 year^?1; S1: 0.07 mol m^?2 year^?1). The estimated POC export flux at K2 was comparable to the sum of the POC flux observed with drifting sediment traps and active carbon flux exported by migrating zooplankton. The export fluxes at both stations were higher than those reported at other time-series sites (ALOHA, the Bermuda Atlantic Time-series Study, and Ocean Station Papa).     

Comparison of sinking particles in the upper 200 m between subarctic station K2 and subtropical station S1 based on drifting sediment trap experiments

Drifting sediment trap experiments were conducted during various seasons to elucidate the characteristics of particles sinking through the upper 200 m of the water column in the western Pacific at subarctic station K2 and subtropical station S1. The sinking particle flux increased when primary productivity was high at each station, during June–July at K2 and during February at S1. Biogenic opal (Opal) and CaCO₃ were the major components of the fluxes at K2 and S1, respectively. Contrary to the expectation of a high flux at the eutrophic station K2 and low flux at the oligotrophic station S1, the annual average organic carbon fluxes at 100 m were comparable at both stations: 62.7 mg C m^?2 day^?1 at K2 and 56.1 mg C m^?2 day^?1 at S1. The similarity of the fluxes was perhaps a reflection of the unexpectedly high primary production at S1. At K2, the organic carbon export ratio (organic carbon flux/primary productivity) was significantly and negatively correlated with primary production and tended to decrease with depth. The magnitude of the rate of attenuation of the organic carbon flux with depth was larger at S1 than at K2. This rate of attenuation tended to decrease and increase with primary production at K2 and S1, respectively. The explanation for these patterns may be that the flux of labile organic carbon at relatively shallow depths decreased with increasing primary production at K2, and zooplankton grazing pressure increased with increasing primary productivity at S1.     

Response of Eucalanus bungii to oceanographic conditions in the western subarctic Pacific Ocean: Retrospective analysis of the Odate Collections

To evaluate their response to oceanographic conditions, interannual variations in seasonal abundance of Eucalanus bungii were investigated in zooplankton samples collected from the Oyashio Current system from 1960 to 2002. Large decadal changes were observed in seasonal timing and population age-structure. During the early 1970s and 1990s, E. bungii were abundant until mid-summer, but during the late 1970s and early 1980s, the season of maximum abundance was limited to spring and early summer. From the late 1970s to early 1980s, spring–summer abundance of newly recruited young copepodites (C1–C2) declined significantly, and an even more pronounced decline was observed for the abundance of the late copepodite stages (C3–C5). Monthly population structure showed that young of the year stopped development at C3 during the late 1970s to early 1980s, but molted into late copepodite stages in the other decades. Seasonal weakening of the Aleutian Low Pressure System estimated from North Pacific Index (NPI) was rapid during the late 1970s to early 1980s, and the NPI was positively correlated with phosphate concentrations at sea surface, spring–summer abundance of the young copepodites stages, and the extended duration of the season of high abundance. These results suggest that the decadal decline of copepod abundance originated at the early life stages, and was associated with a shift of atmospheric and oceanographic conditions. As possible biological mechanisms, we propose reduced egg production, lower survival for the portion of the annual cohort with late birth date, and overwintering of the survivors at younger stages.     

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.     

Effects of the copepod community structure on fecal pellet flux in Kagoshima bay,a deep,semi-enclosed embayment

Seasonal changes in the shape and size composition of fecal pellets were investigated with sediment trap samples from 50 and 150 m in Kagoshima Bay to evaluate how the mesozooplankton community affects fecal pellet flux. Deep vertical mixing was evident in March, and thermal stratification was developed above 50 m in June, August and November. Chlorophyll a, suspended particulate organic carbon (POC) and copepod abundance were uniform throughout the water column during the seasonal mixing and concentrated above 50 m in the stratified seasons. Calanoids were the most predominant copepods in March and poecilostomatoids composed more than 45% of the copepod community in June, August and November. Fecal pellet fluxes at 50 and 150 m were the highest in March, nearly half of POC flux. The relative contribution declined considerably in the other months, especially for less than 4% of POC flux in August. The decline was corresponded to the predominance of cyclopoids and poecilostomatoids. Cylindrical pellets dominated the fecal matters at both depths throughout the study period, while larger cylindrical pellets nearly disappeared at 150 m in June, August and November. Copepod incubation revealed that cylindrical and oval pellets were egested by calanoids and the other copepods, respectively. We suggest that cylindrical fecal pellets produced by calanoid copepods contribute to feces flux but the predominance of poecilostomatoids and/or cyclopoids decreases feces flux via the increase of oval pellets and fragmentation of larger cylindrical pellets.