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We have evaluated development and growth of the ontogenetically migrating copepods in the northwestern subarctic Pacific by frequent sampling and by molting rate experiments during the spring phytoplankton bloom. Since different water masses frequently moved into the surface layers at our sampling station, the copepod abundance and species composition fluctuated greatly. Early copepodite stages of Eucalanus bungii and Neocalanus plumchrus appeared abundantly when a warmer and more saline water mass was present. Despite the fluctuating temporal pattern, the population structure revealed that each copepod species had a different life history strategy during the bloom. Eucalanus bungii were in the stages of gonad maturation, spawning and naupliar development when chlorophyll a concentrations were high. The phytoplankton bloom was utilized for development and lipid accumulation by late copepodites of Neocalanus cristatus and Neocalanus flemingeri and for development of nauplii and early copepodites of N. plumchrus. Molting experiments showed that carbon weight of ‘molters’ in the incubations was greater than that of ‘non-molters’, indicating that primarily animals with heavier body weight are those that molt into the next stage. Mean stage durations estimated by the molting rate method were on the order of 9.7 (C3) to 16.6 days (C4) for N. flemingeri, 13.9 (C3) to 29.1 days (C4) for N. plumchrus, and 12.2 days (C2) for E. bungii. Large fluctuations were observed for stage duration estimates, suggesting different development histories (i.e. ages-within-stage distributions) among the replicate incubations. From these results, we discuss development and growth of the ontogenetically migrating copepods during strongly fluctuating hydrographic conditions in the Oyashio.

Temporal changes in abundance, growth and mortality of heterotrophic bacteria were investigated from frequent dilution experiments in the Oyashio region to evaluate their response to the spring phytoplankton bloom. Bacterial abundance fluctuated from 1.0 to 2.6×10 5 cells mL −1 . Cell density was not different between the pre-bloom and bloom periods, and showed no clear temporal pattern. Potential growth ( g ′) and mortality ( m ) varied from 1.11 to 2.39 day −1 and from 1.21 to 2.16 day −1 , respectively. Neither g ′ nor m was correlated with water temperature or chlorophyll a . Apparent growth rate in unenriched 100% seawater incubations ( g o ) was not limited by nutrients throughout the study period and was comparable to net growth rate ( g ′ −m ) estimates from dilution experiments. These results suggest that bacterial abundance is nearly steady-state in the pre-bloom and bloom period, because bacterial growth is nearly nutritionally saturated and balanced with high mortality.

Bottle incubations were conducted to investigate silica deposition and growth characteristics of siliceous phytoplankton in the Oyashio region using PDMPO, 2-(4-pyridyl)-5-((4-(2-dimethylaminoethyl-aminocarbamoyl)-methoxy)phenyl)oxazole, a fluorescence probe. This stain shows clear fluorescence in the position of newly produced silica cell walls during their vegetative cell growth. Parmales and diatoms dominated siliceous phytoplankton during the pre-bloom and bloom periods, and were successfully stained with PDMPO, indicating active silica deposition. Low PDMPO labeling of cells of the dominant diatoms relative to their initial abundance was often observed in the later half of the bloom period compared with high PDMPO labeling early in the bloom. That suggests that diatom silica deposition had slowed. Silica deposition in other siliceous plankton, i.e. silicoflagellates and choanoflagellates, and diatom resting spores was also detected. Our results show that the PDMPO technique is appropriate to demonstrate important biological processes in the silicon and carbon biogeochemical and cycles in the Oyashio region.

We investigated the feeding habits and grazing rates of the ontogenetically migrating copepods in the Oyashio region to evaluate their grazing impacts on the food web during the spring phytoplankton bloom. The bloom was in progress from early to late April, although chlorophyll a concentrations fluctuated considerably with the frequent exchange of different water masses. Biomass of the copepod community reached a maximum in mid-April when late copepodites of Neocalanus cristatus , Neocalanus flemingeri and Eucalanus bungii contributed to the biomass increase. Gut pigment contents of the predominant copepods were much higher during the bloom compared with the levels in March (pre-bloom). The temporal fluctuations were not correlated with those of mean chlorophyll a concentrations in the 0–50 m layer. Feeding experiments indicated that major food items for the copepods were centric diatoms and flagellates. During the period of lower ambient chlorophyll, the copepods changed their heterotrophic prey from naked ciliates to tintinnids. Apparent clearance rates were positive for naked ciliates and negative for heterotrophic nanoplankton, Cryptophyceae and bacteria when chlorophyll was high, suggesting trophic cascade effects from copepod feeding even during the phytoplankton bloom. The carbon demands of the copepod community were estimated to be 156 mgC m −2 day −1 in early March to 797 mgC m −2 day −1 in mid-April. The grazing rates on phytoplankton reached 480 mgC m −2 day −1 , equivalent to as much as 28% of primary production. Non-phytoplankton prey supported 40 to 71% of the copepod carbon requirement. These results suggest that the copepod community does not graze the phytoplankton bloom down, but it does have significant impacts on microbial food webs.

To evaluate the impacts of ontogenetically (seasonally) migrating copepods on carbon transport to the mesopelagic zone, we investigated depth distribution, population structure, and feeding activity of the ontogentic copepod community in the western subarctic Pacific Ocean from day–night pairs of zooplankton samples down to 1000 m during the VERtical Transport In the Global Ocean (VERTIGO) program. Over the 31 July–16 August 2005 study period, the biomass of Neocalanus cristatus and Neocalanus plumchrus predominated in the near surface waters, while Neocalanus flemingeri was already dormant at depth. We observed a strong diel migration for Metridia pacifica, and a seasonal downward migration for Eucalanus bungii. Based on gut pigment analysis, ingestion rate of the copepod community was 214–375 mg C m−2 day−1, which was equal to 26–37% of the concurrent primary production. However, comparison of grazing estimated from gut pigments to calculated carbon demand of the copepod community indicates that phytoplankton comprised 37–59% of the ingested carbon. Thus, the copepod community appears to have also relied on detritus and microzooplankton for their nutrition, likely because primary production during this time was dominated by picophytoplankton too small to be grazed by these large copepods. Fecal pellet flux by the copepod community was estimated to account for 141–223% of the sedimentary particulate organic carbon (POC) flux at 150 m, suggesting considerable fragmentation and consumption of pellets in the upper layers. Fecal pellets alone were adequate to meet copepod carbon demand in the surface 0–150 m layer. Active carbon flux by diel migration of M. pacifica (respiration, egestion, and mortality) was 4–17 mg C m−2 day−1, equal to 6–44% of sedimentary POC flux at 150 m. Active carbon flux by N. flemingeri ontogenetic migration (i.e., respiration and mortality at depth) contributed 246 mg C m−2 year−1, equal to 9% of sedimentary POC flux at 1000 m. The imminent downward migration of N. cristatus and N. plumchrus would lead to an additional ontogenetic carbon flux on the order of 1719 mg C m−2 year−1. Copepod fecal pellet transport and active transport by diel and ontogenetic migration are thus important carbon fluxes during a season dominated by small phytoplankton, and ontogenetic migrants in the subarctic Pacific Ocean play a relatively more important role in active carbon flux compared with other open-ocean regions.

Mesopelagic mesozooplankton communities of an oligotrophic (Hawaii Ocean Time series-HOT station ALOHA) and a mesotrophic (Japanese time-series station K2) environment in the North Pacific Ocean are compared as part of a research program investigating the factors that control the efficiency of particle export to the deep sea (VERtical Transport In the Global Ocean—VERTIGO). We analyzed zooplankton (>350 μm) collected from net tows taken between 0 and 1000 m at each site to investigate the biomass size structure and the abundance of the major taxonomic groups in discrete depth intervals throughout the water column. Biomass of zooplankton at K2 over all depths was approximately an order of a magnitude higher than at ALOHA, with a significantly higher proportion of the biomass at K2 in the larger (>2 mm) size classes. This difference was mostly due to the abundance at K2 of the large calanoid copepods Neocalanus spp. and Eucalanus bungii, which undergo ontogenetic (seasonal) vertical migration. The overall strength of diel vertical migration was higher at K2, with a mean night:day biomass ratio in the upper 150 m of 2.5, vs. a ratio of 1.7 at ALOHA. However, the amplitude of the diel migration (change in weighted mean depth between day and night) was higher at ALOHA for all biomass size classes, perhaps due to deeper light penetration causing deeper migration to avoid visual predators. A number of taxa known to feed on suspended or sinking detritus showed distinct peaks in the mesopelagic zone, which affects particle transport efficiency at both sites. These taxa include calanoid and poecilostomatoid (e.g., Oncaea spp.) copepods, salps, polychaetes, and phaeodarian radiolaria at K2, harpacticoid copepods at ALOHA, and ostracods at both sites. We found distinct layers of carnivores (mainly gelatinous zooplankton) in the mesopelagic at K2 including chaetognaths, hydrozoan medusae, polychaetes, and gymnosome pteropods, and, in the upper mesopelagic zone, of ctenophores and siphonophores; at both sites a mesopelagic layer of hyperiid amphipods was found. The large population of ontogenetically migrating calanoid copepods is likely supporting large carnivorous populations at depth at K2. The contrasting zooplankton taxonomic structure at the two sites helps explain the higher efficiency of the biological pump at K2. Factors responsible for increased transport efficiency at K2 include rapid transport of POC via larger fecal pellets produced by zooplankton at K2, and enhanced active carbon export at K2 vs. ALOHA, due to the greater strength of diel vertical migration and to additional ontogenetic migration at K2.

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.