Grazing by meso- and microzooplankton on phytoplankton in the upper reaches of the Schelde estuary (Belgium/The Netherlands)

In contrast with the marine reaches of estuaries, few studies have dealt with zooplankton grazing on phytoplankton in the upper estuarine reaches, where freshwater zooplankton species tend to dominate the zooplankton community. In spring and early summer 2003, grazing by micro- and mesozooplankton on phytoplankton was investigated at three sites in the upper Schelde estuary. Grazing by mesozooplankton was evaluated by monitoring growth of phytoplankton in 200 μm filtered water in the presence or absence of mesozooplankton. In different experiments, the grazing impact was tested of the calanoïd copepod Eurytemora affinis, the cyclopoid copepods Acanthocyclops robustus and Cyclops vicinus and the cladocera Chydorus sphaericus, Moina affinis and Daphnia magna/pulex. No significant grazing impact of mesozooplankton in any experiment was found despite the fact that mesozooplankton densities used in the experiments (20 or 40 ind. l⁻¹) were higher than densities in the field (0.1–6.9 ind. l⁻¹). Grazing by microzooplankton was evaluated by comparing growth of phytoplankton in 30 and 200 μm filtered water. Microzooplankton in the 30–200 μm size range included mainly rotifers of the genera Brachionus, Trichocerca and Synchaeta, which were present from 191 to 1777 ind. l⁻¹. Microzooplankton had a significant grazing impact in five out of six experiments. They had a community grazing rate of 0.41–1.83 day⁻¹ and grazed up to 84% of initial phytoplankton standing stock per day. Rotifer clearance rates estimated from microzooplankton community grazing rates and rotifer abundances varied from 8.3 to 41.7 μl ind.⁻¹ h⁻¹. CHEMTAX analysis of accessory pigment data revealed a similar phytoplankton community composition after incubation with and without microzooplankton, indicating non-selective feeding by rotifers on phytoplankton.     

Downward fluxes of settling particles in the deep Cretan Sea (NE Mediterranean)

During the CINCS project (Pelagic–benthic Coupling IN the oligotrophic Cretan Sea—NE Mediterranean), a single mooring with two sediment traps (at 200 and 1515m water depth) and two current meters was deployed in the southern Cretan Sea margin at a depth of 1550 m. A second mooring deployed at the 500 m station was lost, as a result of fishing activities. The duration of the study was 12 months (November 1994 to November 1995) with sampling intervals of 15 or 16 days. The traps were retrieved, serviced and the sedimented material was collected every 6 months. In total, 48 samples were collected (24 from each trap) throughout the study period and fluxes of total particulate mass, opal, organic matter, carbonates, and lithogenic component were measured. Natural radionuclides (²¹⁰Po and ²¹⁰Pb) were determined for all trap samples. Total mass flux and the fluxes of four major constituents increased with depth, the total mass flux reaching values of nearly 550 mg m⁻² d⁻¹ at 1515 m and 187 mg m⁻² d⁻¹ at 200 m depth, following the same patterns observed in other experiments (ECOMARGE, SEEP-I, SEEP-II). The mean annual mass fluxes were 209 and 49.8 mg m⁻² d⁻¹ at the near bottom and near surface trap respectively. This suggests that lateral transport of particulate matter is of importance in the area. Total mass fluxes at the two depths were characterized by different seasonal fluctuations, although a general decreasing trend was observed from the I (winter) to the II (summer) deployment at both depths. This was mainly a result of reductions in aluminosilicate inputs during the summer dry period. At 200 m depth carbonates were more important during winter, because of a large carbonate input consisting mainly of coccoliths of Emiliania huxleyi, while during the summer decreased fluxes of carbonates and aluminosilicates resulted in a reduction of the mass flux. In contrast, at 1515 m depth the lithogenic component was the dominant component during the winter deployment, indicating a terrigenous input. During the summer period the decrease in mass flux was strongly effected by the decrease in aluminosilicates. There was a diminution in the organic carbon content with a concomitant increase in total mass flux, which, together with the almost negligible increase in the annual ²¹⁰Pb activity with depth and the increase of ²¹⁰Po activity with depth could be interpreted as indicating a contribution of resuspended material to the input at 1515 m. The complex mesoscale circulation of the Cretan Sea, consisting of a cyclone (east)–anticyclone (west) system, controls particle transfer in the area. This hydrodynamic system seems to move water masses towards the southern Cretan Sea margin, and consequently carry materials from the open sea to the upper slope and shelf.     

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.     

Mesozooplankton in the eastern and western subarctic Pacific: community structure, seasonal life histories, and interannual variability

The zooplankton community of the subarctic Pacific is relatively simple, and contains a similar set of major species in all deep water areas of the subarctic Pacific. Their role in the food web varies considerably between coastal and offshore locations. In the oceanic gyres, microzooplankton and other mesozooplankton taxa replace phytoplankton as the primary food source for the dominant mesozooplankton species. Micronekton and larger zooplankton probably replace pelagic fish as major direct predators. Productivity and upper ocean biomass concentrations are intensely seasonal, in part because of seasonality of the physical environment and food supply, but also because of life history patterns involving seasonal vertical migrations (400–2000 m range) and winter dormancy. During the spring–summer season of upper ocean growth, small scale horizontal and vertical patchiness is intense. This can create local zones of high prey availability for predators such as planktivorous fish, birds, and marine mammals. On average, the cores of the subarctic gyres have lower biomass and productivity than the margins of the gyres. There is also some evidence that the Western Gyre is more productive than the Alaska Gyre, but more research is needed to confirm whether this east–west gradient is permanent. There is increasing evidence that the pattern of zooplankton productivity is changing over time, probably in response to interdecadal ocean climate variability. These changes include 2–3 fold shifts in total biomass, 30–60 day shifts in seasonal timing, and 10–25% changes in average body length.     

Downward export of carbon by diel migrant mesozooplankton in the central equatorial Pacific

Using data collected during cruises of the JGOFS equatorial Pacific Study in March/April and October of 1992 at the equator (140°W), we examine the downward transport of carbon by three size classes of die] migrant mesozooplankton (200–500 gm, 500–1000 μm and 1000–2000 gm). In addition to respiratory carbon flux, we consider the flux due to mortality of migrators below the euphotic zone. Diel migrant mesozooplankton biomass was estimated from the difference between nighttime and daytime biomass within the euphotic zone. Except for a four-day period early in the March/April cruise, mesozooplankton nighttime biomass was significantly larger than daytime biomass within the euphotic zone during both cruises. We estimate that the downward flux of carbon from the euphotic zone due to mesozooplankton die] vertical migrators was an average of 0.6 mmol Cm⁻² d⁻¹ and 1.1 mmol C m⁻² d⁻¹ during the March/April and October cruises, respectively. Addition of this flux to the gravitational particle sinking flux estimated from²³⁴Th measurements during the same period results in a 31 % increase in the carbon export flux from the euphotic zone in the equatorial Pacific during the March/April cruise and a 44% increase in the October cruise. The migratory flux is strongly dependent on whether feeding takes place below the euphoric zone, the length of time migrators spend in the deep waters, and the mortality rate of migrators.     

Photosynthate allocation in a temperate sea over an annual cycle: the relationship between protein synthesis and phytoplankton physiological state

The seasonal and vertical variations in the patterns of photosynthate allocation into biomolecules by natural phytoplankton assemblages were determined, together with their species composition, in a coastal station of the central Cantabrian Sea (southern Bay of Biscay). Chlorophyll-a concentration ranged from values below 20 mg m⁻² in winter to values above 80 mg m⁻² during spring and during an upwelling event in summer. Low primary production rates (<300 mgC m⁻² d^-1) were measured during winter and during summer stratification periods. The rate of C fixation during summer upwelling conditions exceeded 3500 mgC m⁻² d⁻¹. In terms of photosynthate partitioning, proteins were the dominant fraction, as they typically accounted for >30% of total photo-assimilated C, with polysaccharides and low molecular weight metabolites showing incorporation percentages around 10–30%. Relative C incorporation into lipids was generally <15%. Recurrent patterns of vertical variability in photosynthate partitioning were observed: the relative synthesis of proteins increased toward the bottom of the euphotic zone, whereas the relative C incorporation into polysaccharides and lipids tended to be higher near the surface. When primary production decreased, the synthesis of proteins was maintained more than that of other molecules. Throughout the year, the relative synthesis of proteins was inversely correlated with phytoplankton biomass, production and growth rate. The conservation of protein synthesis under growth-limiting conditions and the enhancement of lipid and polysaccharide synthesis when irradiance is high seem to constitute general patterns of photosynthate partitioning in marine phytoplankton. In our study, these patterns represented metabolic strategies of phytoplankton in response to changing environmental factors, rather than the effect of variations in the species composition of the community.     

Seasonal dynamics of inorganic nutrients and phytoplankton biomass in the NW Alboran Sea

The seasonal dynamics of inorganic nutrients and phytoplankton biomass (chlorophyll a), and its relation with hydrological features, was studied in the NW Alboran Sea during four cruises conducted in February, April, July and October 2002. In the upper layers, the seasonal pattern of nutrient concentrations and their molar ratios (N:Si:P) was greatly influenced by hydrological conditions. The higher nutrient concentrations were observed during the spring cruise (2.54 μM NO₃⁻, 0.21 μM PO₄³⁻ and 1.55 μM Si(OH)₄, on average), coinciding with the increase of salinity due to upwelling induced by westerlies. The lowest nutrient concentrations were observed during summer (<0.54 μM NO₃⁻, 0.13 μM PO₄³⁻ and 0.75 μM Si(OH)₄, on average), when the lower salinities were detected. Nutrient molar ratios (N:Si:P) followed the same seasonal pattern as nutrient distribution. During all the cruises, the ratio N:P in the top 20 m was lower than 16:1, indicating a NO₃⁻ deficiency relative to PO₄³⁻. The N:P ratio increased with depth, reaching values higher than 16:1 in the deeper layers (200–300 m). The N:Si ratio in the top 20 m was lower than 1:1, excepting during spring when N:Si ratios higher than 1:1 were observed in some stations due to the upwelling event. The N:Si ratio increased with depth, showing a maximum at 50–100 m (>1.5:1), which indicates a shift towards Si-deficiency in these layers. The Si:P ratio was much lower than 16:1 throughout the water column during the four cruises. In general, the spatial and seasonal variation of phytoplankton biomass showed a strong coupling with hydrological and chemical fields. The higher chlorophyll a concentrations at the depth of the chlorophyll maximum were found in April (2.57 mg m⁻³ on average), while the lowest phytoplankton biomass corresponded to the winter cruise (0.74 mg m⁻³ on average). The low nitrate concentrations together with the low N:P ratios found in the upper layers (top 20 m) during the winter, summer and autumn cruises suggest that N-limitation could occur in these layers during great part of the year. However, N-limitation during the spring cruise was temporally overcome by nutrient enrichment caused by an intense wind-driven upwelling event.     

Water fluxes through the Cretan Arc Straits, Eastern Mediterranean Sea: March 1994 to June 1995

Five research cruises were undertaken incorporating ADCP sections along the Cretan Arc Straits and CTD surveys covering the entire area of the Straits and the Cretan Sea. In addition, six moorings (with 15 current meters) were deployed within the Straits, which monitored flows in the surface (50 m), intermediate (250 m), and deep (50 m from the bottom) layers. The ADCP, CM, and CTD datasets enable the derivation of water transports through the Cretan Arc Straits to be assessed. Flow structure through the Cretan Arc Straits is not the typical flow regime with a surface inflow and deep outflow, instead there is a persistent deep outflow of Cretan Deep Water (CDW) (σ_θ>29.2) with an annual mean of ˜0.6 Sv, through the Antikithira and Kassos Straits at depths below 400 m and 500 m, respectively. CDW outflowing transports are higher (˜0.8 Sv) in April–June, and lower (˜0.3 Sv) in October–December. Within the upper water layer (0–˜400 m), the transport and the water exchanges through the Straits are controlled by local circulation features, which weaken substantially below 200 m. The Asia Minor Current (AMC) influences the Rhodes and the Karpathos Straits, resulting in a net inflow of water. In contrast, the Mirtoan/West Cretan Cyclone influences the Antikithira and Kithira Straits, where there is a net outflow. In the Kassos Strait, there is a complex interaction between the East Cretan Cyclone, the Ierapetra Anticyclone and the westward extension of the Rhodes Gyre, which results in a variable flow regime. There is a net inflow in autumn and early winter, and a switch to a net outflow in early spring and summer. The total inflow and outflow, throughout all of the Straits, ranged from ˜2 to ˜3.5 Sv, with higher values in autumn and early winter and lower in summer. The AMC carries ˜2 Sv of inflow through the Rhodes and Karpathos Straits, and this accounts for 60–80% of the total inflow. About 10–15% of the total outflow is of CDW, and a further 45–70% occurs through the upper 400 m of the Kithira and Antikithira Straits. The Kassos Strait exhibits a net inflow of ˜0.7 Sv in autumn and early winter, with a net outflow of ˜0.5 Sv in early spring and summer.     

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.     

Grazing impact of mesozooplankton in an upwelling region off northern California, 2000–2003

Mesozooplankton (>200 μm) grazing impact (% phytoplankton standing crop consumed d⁻¹) was determined by the gut fluorescence method during three springs (2000, 2001 and 2002) and two winters (2002 and 2003) in a coastal upwelling region off northern California. Wind events, in terms of both magnitude and duration, varied inter-annually and seasonally and included both upwelling-favorable and relaxation events. Grazing impact of mesozooplankton also varied inter-annually and seasonally, and was highest during June 2000 (mean=129% of standing crop d⁻¹), a prolonged period of wind “relaxation” and phytoplankton bloom. In contrast, mean grazing impact was lower during periods of stronger, more persistent winds, more active upwelling, greater cross-shelf transport, and lower chlorophyll concentration (25% and 38% in May–June 2001 and 2002, respectively). Wintertime conditions (January 2002 and 2003) were characterized by weakly upwelling or downwelling-favorable winds, low chlorophyll concentration, and lower mean mesozooplankton grazing impact (13% and 12%, respectively). The larger (>500 μm) size class contributed proportionally more to total mesozooplankton (>200 μm) grazing impact than the smaller (200–500 μm) size class during all sampling periods except spring 2002. These results suggest that mesozooplankton grazing impact is higher in spring than in winter, and that during the spring upwelling season, grazing is higher during periods of wind relaxation (weak upwelling) than during periods of stronger upwelling. Further, these results suggest an important role of mesozooplankton grazers on phytoplankton dynamics in the upwelling region off northern California.     

Influence of basin-scale and mesoscale physical processes on biological productivity in the Bay of Bengal during the summer monsoon

Physical forcing plays a major role in determining biological processes in the ocean across the full spectrum of spatial and temporal scales. Variability of biological production in the Bay of Bengal (BoB) based on basin-scale and mesoscale physical processes is presented using hydrographic data collected during the peak summer monsoon in July–August, 2003. Three different and spatially varying physical processes were identified in the upper 300 m: (I) anticyclonic warm gyre offshore in the southern Bay; (II) a cyclonic eddy in the northern Bay; and (III) an upwelling region adjacent to the southern coast. In the warm gyre (>28.8 °C), the low salinity (33.5) surface waters contained low concentrations of nutrients. These warm surface waters extended below the euphotic zone, which resulted in an oligotrophic environment with low surface chlorophyll a (0.12 mg m⁻³), low surface primary production (2.55 mg C m⁻³ day⁻¹) and low zooplankton biovolume (0.14 ml m⁻³). In the cyclonic eddy, the elevated isopycnals raised the nutricline upto the surface (NO₃–N > 8.2 μM, PO₄–P > 0.8 μM, SiO₄–Si > 3.5 μM). Despite the system being highly eutrophic, response in the biological activity was low. In the upwelling zone, although the nutrient concentrations were lower compared to the cyclonic eddy, the surface phytoplankton biomass and production were high (Chl a – 0.25 mg m⁻³, PP – 9.23 mg C m⁻³ day⁻¹), and mesozooplankton biovolume (1.12 ml m⁻³) was rich. Normally in oligotrophic, open ocean ecosystems, primary production is based on ‘regenerated’ nutrients, but during episodic events like eddies the ‘production’ switches over to ‘new production’. The switching over from ‘regenerated production’ to ‘new production’ in the open ocean (cyclonic eddy) and establishment of a new phytoplankton community will take longer than in the coastal system (upwelling). Despite the functioning of a cyclonic eddy and upwelling being divergent (transporting of nutrients from deeper waters to surface), the utilization of nutrients leading to enhanced biological production and its transfer to upper trophic levels in the upwelling region imply that the energy transfer from primary production to secondary production (mesozooplankton) is more efficient than in the cyclonic eddy of the open ocean. The results suggest that basin-scale and mesoscale processes influence the abundance and spatial heterogeneity of plankton populations across a wide spatial scale in the BoB. The multifaceted effects of these physical processes on primary productivity thus play a prominent role in structuring of zooplankton communities and could consecutively affect the recruitment of pelagic fisheries.     

Temporal variations in the concentration and settling flux of carbon and phytoplankton pigments in a deep fjordlike estuary

The weekly mass flux of C and phytoplankton pigments at five depths in the main basin of Puget Sound, a deep (200 m) fjordlike estuary, was sampled for a year with moored sequentially-sampling sediment traps. Flux measurements were compared with weekly samples of suspended pigments in the euphotic zone and bi-monthly samples of total suspended matter and particulate C throughout the water column at the mooring site.Seasonal changes in the total mass flux at all depths were small; instead, physical (river runoff, bottom resuspension) and biological (phytoplankton blooms) events caused occasional sharp increases on a weekly scale. The dry weight concentration of pigments in the trap samples mirrored the concentration of pigments in the euphotic zone suspended matter, increasing from 0·01% in winter to a maximum of 0·65% in late summer. Bloom-induced changes in the pigment concentration were observed almost simultaneously in the euphotic zone and in the traps to a depth of 160 m, indicating a rapid vertical transfer of surface-originating particles by organic aggregates. In contrast to the strong seasonal signal in the pigment concentration, C concentration varied by only a factor of three during the year.The seasonal trend of C/pigment ratios in the C flux arises from at least two sources: (1) a balance between terrestrial sources of C during the high-runoff winter season and in-situ primary production in spring and summer, and (2) cycling of C through the zooplankton population. Budget calculations suggest that the loss of primary-produced C and pigment from the euphotic zone by settling is 5% regardless of season. On an annual basis, this C flux (16 g m⁻²) is sufficient to support previously measured values of benthic aerobic respiration at the mooring site. To account for other C sinks such as burial, predation and chemical oxidation, however, terrestrial C sources and alternate transport pathways, such as vertical advection and sediment movement down the steep basin walls, are necessary.     

Nutrient dynamics and seasonal succession of phytoplankton assemblages in a Southern European Estuary: Ria de Aveiro, Portugal

The spatial and temporal dynamics of dissolved inorganic nitrogen, dissolved phosphate, dissolved silica and chlorophyll a were measured seasonally at eight stations in the Ria de Aveiro. Between December 2000 and September 2001, the seasonal succession of phytoplankton assemblages, inferred after the spatial and seasonal variation of silica and of chlorophyll a concentrations, showed that diatoms (μmol Si L⁻¹) dominated from late autumn until early spring, while chlorophytes (μg Chl a L⁻¹) bloomed during late spring and summer. The Si:N:P ratios and Si concentrations indicated no seasonal depletion in dissolved silica, as in other temperate systems, possibly because of abnormal precipitation and flood events prolonging the supply of dissolved Si to the system. The Si:N:P ratios suggested P limitation at the system level. Despite the relative proportions of available nutrients, the mean phosphorus concentration (5.3 μmol L⁻¹) was above the reported half-saturation constants for P uptake by phytoplankton. Thus, in Ria de Aveiro, the seasonal succession of phytoplankton assemblages may also be dependent on the grazing capacity of the pelagic community through top-down regulation.     

Influence of upwelling and river runoff interaction on phytoplankton assemblages in a Middle Galician Ria and Comparison with northern and southern rias (NW Iberian Peninsula)

The first oceanographic research (hydrography, nutrient salts, chlorophyll, primary production and phytoplankton assemblages) in a Middle Galician Ria was carried out in Corme-Laxe during 2001, just a year before the Prestige oil spill, being the only reference to evaluate eventual changes in the phytoplankton community. Due to the small size of this ria (6.5 km²), oceanographic processes were driven by the continental water supplied by Anllons River during the wet season (20–30 m³ s⁻¹ in winter), and the strong oceanic influence from the nearby shelf during the dry season. The annual cycle showed a spring bloom with high levels of chlorophyll (up to 14 μg Chl-a L⁻¹) and primary production (3 g C m⁻² d⁻¹) and a summer upwelling bloom (up to 8 μg Chl-a L⁻¹ and 10 g C m⁻² d⁻¹) where the proximity of the Galician upwelling core (<13.5 °C at sea surface) favors the input of upwelled seawater (up to 9 μM of nitrate and silicate) to the bottom ria layer, even during summer stratification events (primary production around 2 g C m⁻² d⁻¹). Thus, phytoplankton assemblages form a “continuum” from spring to autumn with a predominance of diatoms and overlapping species between consecutive periods; only in autumn dinoflagellates and flagellates characterized the phytoplankton community. In the Middle Rias as Corme-Laxe, the nutrient values, Chl-a, primary production and phytoplankton abundance for productive periods were higher than those reported for the Northern (Ria of A Coruña) and Southern Rias (Ria of Arousa) for year 2001; this suggests the importance of the hydrographic events occurring in the zone of maximum upwelling intensity of the Western Iberian Shelf, where a lack of annual cycles studies exists.     

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.     

Effects of tidal flushing on phytoplankton in a eutrophic tropical lagoon in Taiwan

The effects of tidal flushing on the abundance, productivity, and community structure of phytoplankton in response to eutrophication were examined every 2–3 months for more than 2 years in Tapong Bay, a tropical lagoon with only 1 tidal inlet connecting it to the sea in southern Taiwan. Water flushing time ranged from 4–12 d (8–25% d⁻¹) in the outer region subject to fast flushing to 8–24 d (4–12% d⁻¹) in the inner region subject to poor flushing. Chlorophyll a, cell number, and gross production (GP) rate of phytoplankton were significantly greater in the inner region than in the outer region. However, while GP rate was normalized by chlorophyll a (P^B) and was expressed as photosynthetic intensity, no significant differences were detected among the study sites. These parameters exhibited a unimodal seasonal pattern across all study sites, with greater values in summer and lower values in winter. No significant differences in species richness or Shannon–Wiener diversity were detected among the study sites. Nevertheless, diversity indices were significantly higher in winter and lower in summer. Classification of phytoplankton communities showed that the grouping patterns were primarily determined by sampling time. Ordination by non-metric multidimensional scaling (MDS) revealed a clear temporal continuum of changes in species composition across all study sites, suggesting that the communities were primarily structured by time, but that it was little affected by study site. Analyses of similarities (ANOSIM) showed that phytoplankton communities sampled in winter could be separated from those in summer, but others were barely separable at all. In summer, the most frequently observed species were the diatom Skeletonema costatum and the cyanobacterium Oscillatoria sp., and these shifted to the diatoms S. costatum and Cylindrotheca closterium in winter. Our results suggest that tidal flushing is an important factor for regulating responses of phytoplankton abundance and productivity to eutrophication in tropical lagoons, but the community structure was little affected due to horizontal mixing by the tidal circulation.     

Plankton dynamics and carbon flux in an area of upwelling off the coast of Morocco

A carbon flux study was carried out off the coast of Morocco, at 31°N, in a region characterized by the presence of a persistent cyclonic eddy. Two short-term (4 and 3 day) deployments of free-floating sediment traps were combined with water column sampling and rate process measurements as the ship followed the traps. For a period of 36 h between trap deployments, a hydrographic section was run along 31°30'N as part of a larger scale survey being carried out simultaneously on the R.V. A. von Humboldt. The first trap deployment was near the eastern margin of the eddy and the traps moved to the north and west in a frontal jet associated with its northern boundary. After the second deployment, which was at the recovery point of the first, the traps moved to the west and then to the southwest. Throughout the study, chlorophyll concentrations varied between 27 and 125 mg m⁻² (0–100 m), with highest concentrations in the upwelled water nearest the coast and in upwelled water generated within the cyclonic eddy. Particulate organic carbon (POC) and particulate organic nitrogen (PON) concentrations were relatively uniform (13.6±1.8 and 1.63±28 g m⁻² with phytoplankton carbon accounting for 16–85% of total POC. Bacterial carbon was 5% of total POC and mesozooplankton carbon concentrations were equivalent to 9% of total POC. Microzooplankton biomass was not assessed but POC:PON ratios in the water column were often high, suggesting there was sometimes a large detrital component in the POC. Primary production rates varied between 1.0 and 2.5 g C m⁻² day⁻¹. Bacterial consumption accounted for 50% of primary production. Metabolic rates suggested that copepods were ingesting more than 0.4 g C m⁻² day⁻¹. while filtration rates suggested that ingestion of phytoplankton carbon was only 0.2 g C m⁻²day⁻¹, even when phytoplankton constituted 85% of the POC. f-ratios (based on uptake rates for ¹⁵N-nitrate and ammonia) were between 0.1 and 0.4, and excretion by mesozooplankton could account for 40% of the daily ammonium uptake by phytoplankton. HPLC pigment analysis showed that when chlorophyll biomass was high, diatoms were dominant, whereas when it was low, small prymnesiophytes, chlorophytes and diatoms were all important. The composition of the fluoresecent pigments in material in the sediment traps indicated that intact phytoplankton and copepod faecal pellets were the main sources but the relative rates of sedimentation of pigment, POC and PON for the two trapping periods did not reflect differences that were observed in the overlying water column. This was likely to be the result of spatial heterogeneity and strong horizontal currents heterogeneity and strong horizontal currents within the euphotic zone. Thus, material collected at 100 m probably did not originate in the water column immediately overlying the traps and trapping efficiencies might also have been variable.     

Microzooplankton herbivory in the Ross Sea, Antarctica

Microplankton abundances and phytoplankton mortality rates were determined at six stations during four cruises spanning three seasons in the Ross Sea polynya, Antarctica (early spring, Oct.–Nov. 1996; mid-late summer, Jan.–Feb. 1997; fall, Apr. 1997; mid-late spring, Nov.–Dec. 1997). Rates of microzooplankton herbivory were measured using a modified dilution technique, as well as by examining the rate of disappearance of phytoplankton (chlorophyll) in samples incubated in the dark (i.e. grazing in the absence of phytoplankton growth). Strong seasonal cycles of phytoplankton and microzooplankton abundance were observed during the study. Microzooplankton abundance varied by more than three orders of magnitude during the four cruises, and was positively correlated with phytoplankton biomass over the entire data set. Nevertheless, microzooplankton grazing was insufficient to impact significantly phytoplankton standing stocks during most of the experiments performed in this perenially cold environment. Only thirteen out of a total of 51 experiments yielded phytoplankton mortality rates that were significantly different from zero. The highest mortality rate observed in this study (0.26 d⁻¹) was modest compared with maximal rates that have been observed in temperate and tropical ecosystems. Results from twenty experiments examining the rate of decrease of phytoplankton biomass during incubations in the dark agreed quite well with the results of the dilution experiments performed at the same time. The range of mortality rates for the dark incubations was −0.09–0.06 d⁻¹, and the average was essentially zero (−0.01 d⁻¹). That is, chlorophyll concentration was virtually unchanged in samples incubated in the dark for 3 d. A number of factors appeared to contribute to the very low rates of microbial herbivory observed, including low water temperature, and the size and taxonomic composition of the phytoplankton assemblage. Based on our results we conclude that the seasonal, massive phytoplankton blooms observed in the Ross Sea are due, in part, to low rates of removal by microbial herbivores.     

Heterotrophic bacterial production in the Cretan Sea (NE Mediterranean)

In March and September 1995, bacterial production was measured by the ³H-leucine method in the oligotrophic Cretan Sea (Aegean Sea, Eastern Mediterranean) in the framework of the CINCS/MTP program. Samples were obtained from four stations (a coastal, a continental shelf and 2 open-sea stations) for the construction of vertical profiles of bacterial abundance and production. Bacterial production ranged from 0.1 μg C m⁻³ h⁻¹ at 1500 m depth, to 82 μg C m⁻³ h⁻¹ in March at 50 m at the coastal station. Higher bacterial integrated production was observed in March at the coastal station (131 mg C m⁻² d⁻¹ for the 0–100 m layer). Bacterial production, integrated through the water-column, was similar in March and September for the open-sea stations (60–70 mg C m⁻² d⁻¹). Relative to production, bacterial concentrations varied little between stations and seasons ranging from 9×10⁵ ml⁻¹ to 3×10⁵ ml⁻¹. Relationships between bacterial biomass and bacterial production indicated seasonal differences, likely reflecting resource limitation of bacterial biomass in March (bloom situation), and predator limitation of bacterial biomass in September (post-bloom situation).     

Plankton distribution associated with frontal zones in the vicinity of the Pribilof Islands

We studied the effect of four types of fronts, the coastal front, the middle front, the shelf partition front and the shelf break front on the quantitative distribution and the composition of plankton communities in the Pribilof area of the eastern Bering Sea shelf in late spring and summer of 1993 and 1994. The coastal fronts near St. Paul and St. George Islands and the coastal domains encircled by the fronts featured specific taxonomic composition of planktonic algae, high abundance and production of phytoplankton, as well as large numbers of heterotrophic nanoplankton. The coastal fronts also were characterized by high values of total mesozooplankton biomass, high concentrations of Calanus marshallae, as well as relatively high abundances of Parasagitta setosa and Euphausiacea compared to surrounding shelf waters. We hypothesize that wind-induced erosion of a weak thermocline in the inner part of the coastal front as well as transfrontal water exchange in subthermocline layers result in nutrient enrichment of the euphotic layer in the coastal fronts and coastal domains in summer time. This leads to prolonged high primary production and high phytoplankton biomass. In this paper a new type of front—the shelf partition front located 45–55 km to the north-east off St. Paul Island—is described, which is assumed to be formed by the flux of oceanic domain waters onto the shelf. This front features a high abundance of phytoplankton and a high level of primary production compared to the adjacent middle shelf. Near the southwestern periphery of the front a mesozooplankton peak occurred, composed of C. marshallae, with biomass in the subthermocline layer, reaching values typical for the shelf break front and the highest for the area. High abundance of phyto- and zooplankton as well as heterotrophic nanoplankton and elevated primary production were most often observed in the area adjacent to the shelf break front at its oceanic side. The phyto- and mesozooplankton peaks here were formed by oceanic community species. The summer levels of phytoplankton numbers, biomass and primary production in the shelf break frontal area were similar to those reported for the outer and middle shelf during the spring bloom and the coastal domains and coastal fronts in summer. In the environment with a narrow shelf to the south of St. George Island, the mesozooplankton peak was observed at the inner side of the shelf break front as close as 20 km from the island shore and was comprised of a “mixed” community of shelf and oceanic species. The biomass in the peak reached the highest values for the Pribilof area at 2.5 g mean wet weight m⁻³ in the 0–100 m layer. Details of the taxonomic composition and the numbers and production of phytoplankton hint at the similarity of processes that affect the phytoplankton summer community in the coastal domains of the islands, at the coastal fronts, and at the oceanic side of the shelf break front. The middle front was the only one that had no effect on plankton composition or its quantitative characteristics in June and July. Location of a variety of frontal productive areas within 100 km of the Pribilof Islands creates favorable foraging habitat for higher trophic level organisms, including sea birds and marine mammals, populating the islands.