Plankton response to El Niño 1997–1998 and La Niña 1999 in the southern region of the California Current

The IMECOCAL Program began in 1997, with the objective of sampling plankton systematically in the Mexican region of the California Current. We present results of chlorophyll a concentrations and zooplankton displacement volumes for the eight cruises from September 1997 to October 1999. The abundance of 22 zooplankton groups was also analyzed for the first four cruises. The response of plankton to the 1997–1998 El Niño was atypical. From September 1997 to January 1998, chlorophyll a and zooplankton volume were at typical values (median integrated chlorophyll was 27 mg/m² and zooplankton 100 ml/1000 m³ in 9801/02). After the peak of El Niño, the system shifted to cooler conditions. Integrated chlorophyll gradually increased to a median of 77 mg/m² in April 1999. In contrast, zooplankton volumes decreased from October 1998 onward, despite favorable phytoplankton availability in 1999. Zooplankton structure was dominated by copepods and chaetognaths through the ENSO cycle, but interannual changes were evident. In the fall of 1997 there was a higher proportion of copepods, chaetognaths, and other minor groups, while the fall of 1998 zooplankton was richer in salps and ostracods. Historical data from previous Baja California CalCOFI cruises indicated that zooplankton volumes measured during the IMECOCAL cruises were above the long-term mean for the period 1951–1984. This suggests a differential response of plankton to the El Niño of 1997–1998 compared to the El Niño of 1957–1959. Regional differences in zooplankton volumes were also found, with central Baja California having 41% higher biomass than northern Baja California. Volumes from both regions were larger than those recorded by CalCOFI off southern California during 1997–1998, but the situation was reversed in 1999. The higher biomasses in the 1997–1998 El Niño can be attributed to high abundance of salps, which showed an affinity with warm, saline water.     

Standing stocks and production rates of phytoplankton and planktonic copepods in the Inland Sea of Japan

Standing stocks and production rates of phytoplankton and planktonic copepods were investigated at 15 stations in the Inland Sea of Japan during four cruises in October–November 1979, January, April and June 1980. The overall mean of phytoplankton biomass was relatively constant during the study period, ranging from 2.3 mg chl.a m^–3 in April to 3.6 mg chl.a m^–3 in October–November. Primary production was low in January (mean: 90 mg C m^–2 d^–1), but higher than 375 mg C m^–2 d^–1 on the other occasions. Integrated annual primary production was 122 g C m^–2 yr^–1. In terms of carbon weight,Paracalanus parvus was the most important copepod species. The variation of the mean copepod biomass (range: 7.6 mg C m^–3 in April to 20.2 mg C m^–3 in June) was smaller than that of copepod production, which was estimated by the Ikeda-Motoda's physiological method. Copepod producion was low in cold seasons (0.6 and 0.9 mg C m^–3 d^–1 in January and April, respectively), and increased, following the elevation of primary production, to 4.9 mg C m^–3 d^–1 in June. Annual copepod production was 33.7 g C m^–2 yr^–1, of which herbivore (secondary) production was 26.4 g C m^–2 yr^–1 (21.7% of primary production). The ratios of pelagic planktivorous fish catch and total fish catch to the primary production were 0.82 and 1.8%, respectively, indicating very high efficiency in exploiting fishery resources in the Inland Sea of Japan.     

Multi-decadal synthesis of benthic–pelagic coupling in the western arctic: Role of cross-shelf advective processes

Using geographic information systems (GIS) software and geostatistical techniques, we utilized three decades of water-column chlorophyll a data to examine the relative importance of autochthonous versus allochthonous sources of reduced carbon to benthic communities that occur from the northern Bering to the eastern Beaufort Sea shelf. Spatial trend analyses revealed areas of high benthic biomass (>300 g m⁻²) and chlorophyll (>150 mg m⁻²) on both the southern and northern Chukchi shelf; both areas are known as depositional centers for reduced organic matter that originates on the Bering Sea shelf and is advected northward in Anadyr and Bering shelf water masses. We found a significant correlation between biomass and chlorophyll a in the Chukchi Sea, reflective of the strong benthic–pelagic coupling in a system that is utilized heavily by benthic-feeding marine mammals. In contrast, there was no significant correlation between biomass and chlorophyll in the Beaufort Sea, which by comparison, is considerably less productive (biomass and chlorophyll, <75 g m⁻² and <50 mg m⁻², respectively). One notable exception is an area of relatively high biomass (50–100 g m⁻²) and chlorophyll (80 mg m⁻²) near Barter Island in the eastern Beaufort Sea. Compared to other adjacent areas in the Beaufort Sea, the chlorophyll values in the vicinity of Barter Island were considerably higher and likely reflect a long-hypothesized upwelling in that area and close coupling between the benthos and autochthonous production. In the Bering Sea, a drop in benthic biomass in 1994 compared with previous measurements (1974–1993) may support earlier observations that document a decline in biomass that began between the 1980s and 1990s in the Chirikov Basin and south of St. Lawrence Island. The results of this study indicate that the benthos is an excellent long-term indicator of both local and physical advective processes. In addition, this work provides further evidence that secondary production on arctic shelves can be significantly augmented by reduced carbon advected from highly productive adjacent shelves.     

Microzooplankton and mesozooplankton in an upwelling filament off Galicia: modelling and sensitivity analysis of the linkages and their impact on the carbon dynamics

This paper reports estimates of trophic flows of carbon off the Galician coast from a 1D ecological model, which are compared with field data from a two week Lagrangian drift experiment. The model consists of 9 biological components: nitrate, ammonium, >5μm phytoplankton, <5μm phytoplankton, heterotrophic nanoflagellates/dinoflagellates (5–20 μm), heterotrophic dinoflagellates (>20 μm), ciliates, fast sinking detritus and slow sinking detritus. Calculations were made for the fluxes of carbon between biological components within the upper 45m of the water column. The temporal development of primary production during the simulation period of two weeks was in good agreement with field estimates, which varied between 248 and 436mgC.m⁻².d⁻¹. Heterotrophic nanoflagellates had the greatest impact on carbon flux, with a grazing rate of 168mgC.m⁻².d⁻¹. Herbivorous grazing by microzooplankton amounted to 215mgC.m⁻².d⁻¹, whereas grazing by copepods on phytoplankton was 35mgC.m⁻² d⁻¹. Copepods grazing on microzooplankton was minor (0.47mgC.m⁻².d⁻¹) and the export flux from the upper 45m was 302mgC.m⁻².d⁻¹. Sensitivity analyses, in which the grazing parameters (i.e the functional relationship between ingestion and food concentration) were changed, were carried out on the heterotrophic dinoflagellate, ciliate and heterotrophic nanoflagellates/dinoflagellate components of the model. These changes did not alter the temporal development of heterotrophic nanoflagellates/dinoflagellates biomass significantly, but ciliates and heterotrophic dinoflagellates were more sensitive to variations in the grazing parameters. The overall conclusion from this modelling study is that the coupling between small phytoplankton and heterotrophic nanoflagellates was the quantitatively most important process controlling carbon flow in this region.     

Variability of C02 distributions and sea-air fluxes in the central and eastern equatorial Pacific during the 199–1994 El Nin˜o

As part of the U.S. JGOFS Program and the NOAA Ocean-Atmosphere Carbon Exchange Study (DACES), measurements of C02 partial pressure were made in the atmosphere and in the surface waters of the central and eastern equatorial Pacific during the boreal spring and autumn of 1992, the spring of 1993, and the spring and autumn of 1994. Surface-water pC0₂ data indicate significant diurnal, seasonal, and interannual variations. The largest variations were associated with the 1991–1994 ENSO event, which reached maximum intensity in the spring of 1992. The lower values of surface-water ΔpC0₂ observed during the 1991–1994 ENSO period were the result of the combined effects of both remotely and locally forced physical processes. The warm pool, which reached a maximum eastward extent in January-February of 1992, began in September of 1991 as a series of westerly wind events lasting about 30 days. Each wind event initiated an eastward propagating Kelvin wave which caused a deepening of the thermocline. By the end of January 1992 the thermocline was at its maximum depth, so that the upwelled water was warm and C0₂-depleted. In April of the same year, the local winds were weaker than normal, and the upwelling was from shallow depths. These changes resulted in a lower-than-normal C02 flux to the atmosphere. The results show that for the one-year period from the fall of 1991 until the fall of 1992, approximately 0.3 GtC were released to the atmosphere; 0.6 GtC were released in 1993, and 0.7 GtC in 1994, in good agreement with the model results of Ciais et al. [Science,269,1098–1102;J. Geophys. Res.,100, 5051–5070]. The net reduction of the ocean-atmosphere C02 flux during the 1991–1994 El Nifio was on the order of 0.8 – 1.2 GtC. Thus, the total amount of C02 sequestered in the equatorial oceans during the prolonged 1991–1994 El Nin˜o period was about 25% higher than the severe El Nin˜o of 1982–1983.     

Contrasts in sedimentation flux below the southern California Current in late 1996 and during the El Niño event of 1997–1998

The vertical flux of particulate matter at 330 m depth in San Lázaro Basin off southern Baja California ranged from 63 to 587 mg m⁻² d⁻¹ between August and November 1996. Organic carbon contents were between 5.6 and 14.8%, yielding flux rates of 9–40 mgC m⁻² d⁻¹. In December 1997 and January 1998, at the height of the strong El Niño event, the respective fluxes (47–202 mg m⁻² d⁻¹ and 3–8 mgC m⁻² d⁻¹) were comparable. The February–June 1998 records, however, revealed sharply reduced mass (1–6 mg m⁻² d⁻¹) and organic carbon (0.2–0.8 mgC m⁻² d⁻¹) fluxes. The organics collected in 1996 were predominantly autochthonous (δ¹³C=−22‰; C/N=8). The variations in δ¹⁵N (8.3–11.0‰) suggest an alternation of new and regenerated production, possibly associated with fluctuations in the intensity of deep mixing during that autumn. The relatively high organic matter fluxes in December 1997 appear to be associated with regenerated production. The average composition from February to June 1998 (δ¹³C=−23.6‰; ¹⁵N=11.7‰; C/N=10.5) indicates degraded material of marine origin. The maximum δ¹⁵N value found (14‰) suggests that deeper, denitrified waters were brought to the surface and possibly advected laterally. Regime changes in the waters of the basin occur at 6–10 week intervals, evidenced by concurrent shifts in most of the measured parameters, including fecal pellet types and metal chemistry. The marine snow-dominated detritus collected showed a shift from a mixed diatom-rich-radiolarian-coccolith assemblage in late 1996 to a coccolith-dominated assemblage, including the contents of fecal pellets, during the 1997–1998 El-Niño period. T–S profiles, plankton analysis and chlorophyll contents of the upper water column indicated that the strong phytoplankton bloom, normally associated with seasonal upwelling along the Pacific coast of Baja, did not occur during the spring of 1998. The persistence of oligotrophic conditions during the 1997–1998 El Niño event favored the dominance of nanoplankton and reduced the vertical flux of particles.     

Seasonal Variations of Plankton Food Web Structure in the Coastal Water off Usujiri Southwestern Hokkaido,Japan

The planktonic food web structure in the subarctic coastal water off Usujiri south-western Hokkaido, Japan was investigated from June 1997 to June 1999, based on seasonal biomass data of pico- (<2 µm), nano- (2–10 µm), micro- (10–200 µm) and mesoplankton (>200 µm), and path analysis using the structural equation model (SEM). In spring, microphytoplankton predominated due to diatom bloom, while pico- and nanophytoplankton predominated in the other seasons, except November and December 1997. The seasonal change in size distribution of heterotrophic plankton was almost similar to that of phytoplankton, and mesozooplankton biomass was high in spring. The path analyses suggest that the main channel in the microbial food web could vary according to phytoplankton size composition, indicating not only the classical food chain (microphytoplankton - copepods) but also the indirect route (microphytoplankton - naked dinoflagellates - copepods).     

Abundance, Biomass, Production and Trophic Roles of Micro- and Net-Zooplankton in Ise Bay, Central Japan, in Winter

We investigated the geographical variations in abundance and biomass of the major taxonomic groups of micro- and net-zooplankton along a transect through Ise Bay, central Japan, and neighboring Pacific Ocean in February 1995. The results were used to estimate their secondary and tertiary production rates and assess their trophic roles in this eutrophic embayment in winter. Ise Bay nourished a much higher biomass of both micro- and net-zooplankton (mean: 3.79 and 13.9 mg C m^–3, respectively) than the offshore area (mean: 0.76 and 4.47 mg C m^–3, respectively). In the bay, tintinnid ciliates, naked ciliates and copepod nauplii accounted for, on average, 69, 18 and 13% of the microzooplankton biomass, respectively. Of net-zooplankton biomass, copepods (i.e. Acartia, Calanus, Centropages, Microsetella and Paracalanus) formed the majority (mean: 63%). Average secondary production rates of micro- and net-zooplankton in the bay were 1.19 and 1.87 mg C m^–3d^–1 (or 23.1 and 36.4 mg C m^–2d^–1), respectively, and average tertiary production rate of net-zooplankton was 0.75 mg C m^–3d^–1 (or 14.6 mg C m^–2d^–1). Available data approximated average phytoplankton primary production rate as 1000 mg C m^–2d^–1 during our study period. The transfer efficiency from primary production to zooplankton secondary production was 6.0%, and the efficiency from secondary production to tertiary production was 25%. The amount of food required to support the zooplankton secondary production corresponded to 18% of the phytoplankton primary production or only 1.7% of the phytoplankton biomass, demonstrating that the grazing impact of herbivorous zooplankton was minor in Ise Bay in winter.     

Carbon cycling in a high-arctic marine ecosystem – Young Sound, NE Greenland

Young Sound is a deep-sill fjord in NE Greenland (74°N). Sea ice usually begins to form in late September and gains a thickness of 1.5 m topped with 0–40 cm of snow before breaking up in mid-July the following year. Primary production starts in spring when sea ice algae begin to flourish at the ice–water interface. Most biomass accumulation occurs in the lower parts of the sea ice, but sea ice algae are observed throughout the sea ice matrix. However, sea ice algal primary production in the fjord is low and often contributes only a few percent of the annual phytoplankton production. Following the break-up of ice, the immediate increase in light penetration to the water column causes a steep increase in pelagic primary production. Usually, the bloom lasts until August–September when nutrients begin to limit production in surface waters and sea ice starts to form. The grazer community, dominated by copepods, soon takes advantage of the increased phytoplankton production, and on an annual basis their carbon demand (7–11 g C m⁻²) is similar to phytoplankton production (6–10 g C m⁻²). Furthermore, the carbon demand of pelagic bacteria amounts to 7–12 g C m⁻² yr⁻¹. Thus, the carbon demand of the heterotrophic plankton is approximately twice the estimated pelagic primary production, illustrating the importance of advected carbon from the Greenland Sea and from land in fuelling the ecosystem.In the shallow parts of the fjord (<40 m) benthic primary producers dominate primary production. As a minimum estimate, a total of 41 g C m⁻² yr⁻¹ is fixed by primary production, of which phytoplankton contributes 15%, sea ice algae <1%, benthic macrophytes 62% and benthic microphytes 22%. A high and diverse benthic infauna dominated by polychaetes and bivalves exists in these shallow-water sediments (<40 m), which are colonized by benthic primary producers and in direct contact with the pelagic phytoplankton bloom. The annual benthic mineralization is 32 g C m⁻² yr⁻¹ of which megafauna accounts for 17%. In deeper waters benthic mineralization is 40% lower than in shallow waters and megafauna, primarily brittle stars, accounts for 27% of the benthic mineralization. The carbon that escapes degradation is permanently accumulated in the sediment, and for the locality investigated a rate of 7 g C m⁻² yr⁻¹ was determined.A group of walruses (up to 50 adult males) feed in the area in shallow waters (<40 m) during the short, productive, ice-free period, and they have been shown to be able to consume <3% of the standing stock of bivalves (Hiatella arctica, Mya truncata and Serripes Groenlandicus), or half of the annual bivalve somatic production. Feeding at greater depths is negligible in comparison with their feeding in the bivalve-rich shallow waters.     

Geographical and seasonal variations in abundance,biomass and estimated production rates of microzooplankton in the Inland Sea of Japan

We measured abundance and biomass of 3 major groups of microzooplankton, i.e. tintinnids, naked ciliates and copepod nauplii, at 21 stations in the Inland Sea of Japan in October 1993, January, April and June 1994. The average abundance of the microzooplankton over the entire Inland Sea of Japan ranged from 2.39×10⁵ indiv. m^–3 in January to 4.00×10⁵ indiv. m^–3 in April. Ciliated protozoans, i.e. tintinnids plus naked ciliates, numerically dominated the microzooplankton. The average biomass of the microzooplankton was exceedingly high in October (8.62 mg C m^–3) compared to that in the other months (2.06, 2.79 and 2.68 mg C m^–3 in January, April and June, respectively). The ciliated protozoans also dominated in terms of biomass except in October, when copepod nauplii were more important. Estimated production rate of the microzooplankton was highest in October (average: 6.02 mg C m^–3d^–1) and followed in order by June, April and January (1.94, 1.14 and 0.54 mg C m^–3d^–1, respectively). Due to higher specific growth rate, the production rate by the ciliated protozoans far exceeded that by the copepod nauplii. The trophic importance of the microzooplankton in the pelagic ecosystem of the Inland Sea of Japan was assessed by estimating carbon flow through the microzooplankton community.     

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.     

Vertical flux of biogenic matter during a Lagrangian study off the NW Spanish continental margin

Lagrangian experiments with short-term, drifting sediment traps were conducted during a cruise on RRS Charles Darwin to the NW coast of Spain to study the vertical flux and composition of settling biogenic matter. The cruise was split into two legs corresponding to (i) a period of increased production following an upwelling event on the continental shelf (3–10 August 1998) and (ii) an evolution of a cold water filament originating from the upwelled water off the shelf (14–19 August). The export of particulate organic carbon (POC) from the upper layer (0–60m) on the shelf was 90–240mgC.m⁻².d⁻¹ and off the shelf was 60–180mgC.m⁻².d⁻¹. Off shelf the POC flux at 200m was 50–60mg.m⁻².d⁻¹. A modest sedimentation of diatoms (15–30mgC.m⁻².d⁻¹) after the upwelling was associated with increased vertical flux of chlorophyll a (1.8–2.1mg.m⁻².d⁻¹) and a decrease of the POC:PON molar ratio of the settled material from 9 to 6.4. Most of the pico-, nano-, and microplankton in the settled material were flagellates; diatoms were significant during the on shelf and dinoflagellates during the off shelf leg. Off shelf, the exponential attenuation of POC flux indicated a strong retention capacity of the plankton community between 40 and 75m. POC:PON ratio of the settled particulate matter decreased with depth and the relative portion of flagellates increased, suggesting a novel, flagellate and aggregate mediated particulate flux in these waters. Export of POC from the euphotic layer comprised 14–26% of the integrated primary production per day during the on shelf leg and 25–42% during the off shelf leg, which characterises the importance of sedimentation in the organic carbon budget of these waters.     

Seasonal horizontal and vertical variability in primary production and standing stocks of phytoplankton and zooplankton in the Cretan Sea and the Straits of the Cretan Arc (March 1994–January 1995)

Phytoplankton communities, production rates and chlorophyll levels, together with zooplankton communities and biomass, were studied in relation to the hydrological properties in the euphotic zone (upper 100 m) in the Cretan Sea and the Straits of the Cretan Arc. The data were collected during four seasonal cruises undertaken from March 1994 to January 1995.The area studied is characterised by low nutrient concentrations, low ¹⁴C fixation rates, and impoverished phytoplankton and zooplankton standing stocks. Seasonal fluctuations in phytoplankton densities, chlorophyll standing stock and phytoplankton production are significant; maxima occur in spring and winter and minima in summer and autumn. Zooplankton also shows a clear seasonal pattern, with highest abundances occurring in autumn–winter, and smallest populations in spring–summer. During summer and early autumn, the phytoplankton distribution is determined by the vertical structure of the water column.Concentrations of all nutrients are very low in the surface waters, but increase at the deep chlorophyll maximum (DCM) layer, which ranges in depth from about 75–100 m. Chlorophyll-a concentrations in the DCM vary from 0.22–0.49 mg m⁻³, whilst the surface values range from 0.03–0.06 mg m⁻³. Maxima of phytoplankton, in terms of cell populations, are also encountered at average depths of 50–75 m, and do not always coincide with chlorophyll maxima. Primary production peaks usually occur within the upper layers of the euphotic zone.There is a seasonal succession of phytoplankton and zooplankton species. Diatoms and ‘others’ (comprising mainly cryptophytes and rhodophytes) dominate in winter and spring and are replaced by dinoflagellates in summer and coccolithophores in autumn. Copepods always dominate the mesozooplankton assemblages, contributing approximately 70% of total mesozooplankton abundance, and chaetognaths are the second most abundant group.     

Mesozooplankton dynamics in nearshore waters of the Great Barrier Reef

Zooplankton dynamics (community composition, juvenile somatic growth rate, adult egg production, secondary production) were studied in coastal waters of the Great Barrier Reef. Two sectors were compared, one adjacent to a catchment of near-pristine land use patterns, the other to a more intensively farmed catchment. Sampling was conducted in the austral winter (August) and summer (January–March) of two succeeding years. Gradients in zooplankton community composition were weak, with only moderate effects of season and sector. Overall, 37% of zooplankton biomass was in the 73–150 μm size fraction, 26% in the 150–350 μm fraction, and 38% was >350 μm. There was no biomass difference and only small differences in community composition between samples taken during the day and at night; ostracods and large calanoid copepods were occasionally more common at night. Carbon-specific growth rates averaged 0.29 d⁻¹ for cyclopoid copepods and 0.35 d⁻¹ for calanoid copepods, with no difference between sectors. Calanoid copepod growth showed a significant relationship to chlorophyll concentration, but cyclopoid copepods did not. Copepod egg production was low (7.9 ± 5.9 eggs female⁻¹ d⁻¹) and apparently food-limited. Copepod secondary production was lower in August (mean = 2.6, range 1.4–4.0 mg C m⁻² d⁻¹) than in January–March (mean = 8.5, range 2.4–15.5 mg C m⁻² d⁻¹). Secondary production by mesozooplankton in the 73–100 μm size range averaged 0.9% of total phytoplankton production.     

Picophytoplankton in the equatorial Pacific: vertical distributions in the warm pool and in the high nutrient low chlorophyll conditions

Abundance distribution and cellular characteristics of picophytoplankton were studied in two distinct regions of the equatorial Pacific: the western warm pool (0°, 167°E), where oligotrophic conditions prevail, and the equatorial upwelling at 150°W characterized by high-nutrient low-chlorophyll (HNLC) conditions. The study was done in September–October 1994 during abnormally warm conditions. Populations of Prochlorococcus, orange fluorescing Synechococcus and picoeukaryotes were enumerated by flow cytometry. Pigment concentrations were studied by spectrofluorometry. In the warm pool, Prochlorococcus were clearly the dominant organisms in terms of cell abundance, estimated carbon biomass and measured pigment concentration. Integrated concentrations of Prochlorococcus, Synechococcus and picoeukaryotes were 1.5×10¹³, 1.3×10¹¹ and 1.5×10¹¹ cells m⁻², respectively. Integrated estimated carbon biomass of picophytoplankton was 1 g m⁻², and the respective contributions of each group to the biomass were 69, 3 and 28%. In the HNLC waters, Prochlorococcus cells were slightly less numerous than in the warm pool, whereas the other groups were several times more abundant (from 3 to 5 times). Abundance of Prochlorococcus, Synechococcus and picoeukaryotes were 1.2×10¹³, 6.2×10¹¹ and 5.1×10¹¹ cells m⁻², respectively. The integrated biomass was 1.9 g C m⁻². Prochlorococcus was again the dominant group in terms of abundance and biomass (chlorophyll, carbon); the respective contributions of each group to the carbon biomass were 58, 7 and 35%. In the warm pool the total chlorophyll biomass was 28 mg m⁻², 57% of which was divinyl chlorophyll a. In the HNLC waters, the total chlorophyll biomass was 38 mg m⁻², 44% of which was divinyl chlorophyll a. Estimates of Prochlorococcus, Synechococcus and picoeukaryotes cell size were made in both hydrological conditions.     

Production of <Emphasis Type="Italic">Acartia steueri</Emphasis> (Copepoda: Calanoida) in Ilkwang Bay,Southeastern Coast of Korea

Production of the marine calanoid copepod Acartia steueri was measured from 2 October 1991 to 8 October 1992 at a station in Ilkwang Bay, on the southeastern coast of Korea. Phytoplankton standing stock ranged over 1.0 to 9.3 mg chl.a m⁻³, and annual primary productivity (by the C-14 method) at three stations was estimated at 200 gC m⁻² yr⁻¹. Acartia steueri (nauplii + copepodids + adults) were present in the plankton throughout the year, with seasonal variation in abundance. Biomass of A. steueri, excluding the NI stage, was 0.01–4.55 mgC m⁻³ (mean: 0.68 mgC m⁻³) with peaks in November, February, May and July-early August, and relatively low biomass in September– January. Instantaneous growth rates of the nauplius stages were higher than the copepodid stages. Annual production of A. steueri was 25.1 mgC m⁻³ yr⁻¹ (or 166 mgC m⁻² yr⁻¹), showing peaks in November, May and July–August with a small peak in February, and low production in December–April and September–October. There were no significant relationships between the daily production rate of A. steueri and temperature or chlorophyll a concentration, indicating that unknown other factors might be related to the variation of the production rate.     

Variation in mesozooplankton community structure in the Japan/East Sea (1991–1999) with possible influence of the ENSO scale climatic variability

Seasonal and interannual change in mesozooplankton community structure in the offshore Tsushima Current area of the Japan/East Sea was studied in relation to climatic events and temporal variability of the upper water column environment from 1991 to 1999. We observed a clear seasonal succession in zooplankton community structure from a cold-water copepod-dominated community in winter and spring to a gelatinous, carnivorous and warm-water copepod-dominated community in summer and autumn. The mean abundance (inds. m⁻³) of the spring community was 3–4-fold higher than that of the other season. The spring community structure varied considerably between years: the community characterized by the summer–autumn type zooplankton assemblage appeared in 1991–1993 and 1998, while the community characterized by high abundance of cold-water copepods appeared in the mid 1990s. Time series profiles of water density and nutrients showed the thickness of the surface warm Tsushima Current and the cold subsurface water increased and decreased, respectively, limiting nutrient supply to the surface water in 1992 and 1998. These results suggest that a thick, warm surface layer might reduce the reproductive success and survival ratio of the cold-water copepods both directly and indirectly, by hindering their upward migration to the surface where food is available, and by limiting phytoplankton growth due to nutrient depletion, respectively. A Monsoon Index (MOI) showed weaker winter wind stress in 1992 and 1998, which might have attenuated formation of the cold subsurface layer in the northern Japan/East Sea and been responsible for surface warming of the study area. Since 1992 and 1998 were El Niño years, this study revealed that ENSO related climatic variability on an interannual time scale considerably influenced the lower trophic level ecosystem in the Japan/East Sea.     

Export flux of particles at the equator in the western and central Pacific ocean

Export of particles was studied at the equator during an El Nin˜o warm event (October 1994) as part of the French ORSTOM/FLUPAC program. Particulate mass, carbon (organic and inorganic) (C), nitrogen (N), and phosphorus (P) export fluxes were measured at the equator in the western and central Pacific during two 6–7 day-long time-series stations located in the warm pool (TS-I at 0°, 167°E) and in the equatorial HNLC situation (TS-II at 0°, 150°W), using drifting sediment traps deployed for 48 h at four depths (between, approximately, 100 and 300 m).The particulate organic carbon (POC) fluxes at the base of the euphotic zone (0.1 % light level), were approximately four times lower at TS-I than at TS-11 (4.1 vs. 17.0 mmol C m^-2 day^-1). Conversely, fluxes measured at 300 m were similar at both sites (3.6vs. 3.7 mmol C m⁻² day⁻¹ at TS-I and TS-11, respectively). This change in export fluxes was in good agreement with food-web dynamics in the euphotic zone characterized by an increase in plankton biomasses and metabolic rates and a shift towards larger size from TS-1 to TS-II. The POC flux profiles indicated high remineralization (up to 78%) of the exported particles at TS-II, between 100 and 200 m in the Equatorial Undercurrent. According to zooplankton ingestion estimates from 100 – 300 m, 60% of this POC loss could be accounted for by zooplankton grazing. At TS-I, no marked increase of flux with depth was observed, and we assume that loss of particles was compensated by in-situ particle production by zooplankton. Fluxes of particulate nitrogen and phosphorus followed the same general patterns as the POC fluxes. The elemental and pigment composition of the exported particles was not very different between the two stations. In particular, the POCYN flux molar ratio at the base of the euphotic zone was low, 6.9 and 6.2 at TS-1 and TS-II, respectively.For particulate inorganic carbon (mainly carbonate) flux, values at the base of the euphotic zone averaged 0.9 mmol C m-2 day-1 at TS-I and 2.3 mmol C m-2 day-1 at TS-11 (corresponding to a 2.6-fold increase) and showed low depth changes at both stations.POC export flux (including active flux associated with the interzonal migrants) at the 0.1 % light level depth represented only 8% of primary production (1°C uptake) measured at TS-1 and 19% at TS-II. For the time and space scales considered in the present study, new primary production, as measured by the 15N method, was in good agreement with the total export flux in the HNLC situation, thus leading to negligible dissolved organic carbon (DOC) or nitrogen (DON) losses from the photic zone. Conversely, export flux was found to be only 50% (C units) and 60% (N) of new production in the oligotrophic system, either because of an overestimation by the 15N method or of a significant export of DOC and DON.Comparison with other oceanic regions shows that export flux in the warm pool was within the same range as in the central gyres. On the other hand, comparison with EgPac data in the central Pacific suggests that there is no straightforward relation between the magnitude of the export and surface nitrate concentrations.     

Seasonal appearance ofProchlorococcus in Suruga Bay,Japan in 1992–1993

Seasonal appearance ofProchlorococcus was studied by flow cytometry in Suruga Bay, Japan in 1992–1993.Prochlorococcus cells were in high concentrations (>1×10⁴ cells ml^–1) from July to October 1992 and September 1993, when the water temperature was over 20°C. The 16S rRNA of the isolated cells showed 98.5% sequence homology with that ofP. marinus (Sargasso strain), indicating that they are the same species. The former has a high divinyl-chlorophyll (DV-Chl.)a/b ratio similar to the Mediterranean strain and different from the Sargasso strain. Maximum concentration ofProchlorococcus at the surface water was 2.5×10⁴ cells ml^–1 in August 1992 and their DV-Chl.a accounted for 4.0% of the total chlorophylla. A decrease in cell density to less than 5×10³ cells ml^–1 was observed from December to May with an exceptional rise in January 1993. WhileProchlorococcus showed a maximum concentration of 3.6×10⁴ cells ml^–1 at 10 m depth in September 1992, phycoerythrin (PE)-richSynechococcus spp. were dominant with their maximum concentration of 2.2×10⁵ cells ml^–1 in the same water body. On the other hand, phycocyanin (PC)-richSynechococcus spp. and the larger phytoplankters showed maximum concentrations in the surface waters in May and June. BothProchlorococcus and PE-richSynechococcus showed their lowest concentrations in April. A significant positive correlation was obtained between cell concentrations of the PE-richSynechococcus andProchlorococcus.