Seasonal dynamics of primary and new production in the northern South China Sea: The significance of river discharge and nutrient advection

Biochemical and productivity measurements and nutrient enrichment experiments were conducted on three cruises in summer and two cruises in winter on the shelf and the basin of the northern South China Sea (SCS) between 2001 and 2004. Phytoplankton production, in terms of depth-integrated new production (INP) or depth-integrated primary production (IPP), was higher in winter than in summer and on the shelf than in the basin. In winter, with deepening of the mixed layer, nitrate from the shallow nitracline that characterized the SCS waters was made available in the surface and supported the highest production of the year. Averaged INP measured in winter (0.25 g C m⁻² d⁻¹) was about twice the summer average (0.12 g C m⁻² d⁻¹) and was 0.19 g C m⁻² d⁻¹ on the shelf compared with 0.15 g C m⁻² d⁻¹ in the basin. In winter, average INP on the shelf was higher than the basin (0.34 versus 0.21 g C m⁻² d⁻¹); whereas in summer, averaged INP on the shelf (0.13 g C m⁻² d⁻¹) and the basin (0.11 g C m⁻² d⁻¹) were similar. While averaged IPP measured in the basin was higher in winter than in summer (0.53 versus 0.35 g C m⁻² d⁻¹), IPP on the shelf showed little temporal variation (0.82 in winter versus 0.84 g C m⁻² d⁻¹ in summer). Considerable spatial and inter-annual variation in production was measured in the shelf waters during summer, which could be linked to discharge volume and plume flow direction of the Zhujiang River. While the shelf waters in summer were mostly nitrogen starved or nitrogen and phosphorus co-limited, excessive river runoff may cause the nutritive state to shift to phosphorus deficiency. Waters with low surface salinities and high fluorescence from riverine mixing could be found extending from the Zhujiang mouth to as far as offshore southern Taiwan after a typhoon passed the northern SCS and brought heavy rainfall. Overall, both nutrient advection in winter and river discharge from the China coast in summer made new nitrogen available and shaped the dynamics of phytoplankton production in these oligotrophic waters.     

Effects of mesoscale phytoplankton variability on the copepods Neocalanus flemingeri and N. plumchrus in the coastal Gulf of Alaska

The copepods Neocalanus flemingeri and N. plumchrus are major components of the mesozooplankton on the shelf of the Gulf of Alaska, where they feed, grow and develop during April–June, the period encompassing the spring phytoplankton bloom. Satellite imagery indicates high mesoscale variability in phytoplankton concentration during this time. Because copepod ingestion is related to food concentration, we hypothesized that phytoplankton ingestion by N. flemingeri and N. plumchrus would vary in response to mesoscale variability of phytoplankton. We proposed that copepods on the inner shelf, where the phytoplankton bloom is most pronounced, would be larger and have more lipid stores than animals collected from the outer shelf, where phytoplankton concentrations are typically low. Shipboard feeding experiments with both copepods were done in spring of 2001 and 2003 using natural water as food medium. Chlorophyll concentration ranged widely, between 0.32 and 11.44 μg l⁻¹ and ingestion rates varied accordingly, between 6.0 and 627.0 ng chl cop⁻¹ d⁻¹. At chlorophyll concentrations<0.50 μg l⁻¹, ingestion is always low, <40 ng cop⁻¹ d⁻¹. Intermediate ingestion rates were observed at chlorophyll concentrations between 0.5 and 1.5 μg l⁻¹, and maximum rates at chlorophyll concentrations>1.5 μg l⁻¹. Application of these feeding rates to the phytoplankton distribution on the shelf allowed locations and time periods of low, intermediate and high daily feeding to be calculated for 2001 and 2003. A detailed cross-shelf survey of body size and lipid store in these copepods, however, indicated they were indistinguishable regardless of collection site. Although the daily ingestion of phytoplankton by N. flemingeri and N. plumchrus varied widely because of mesoscale variability in phytoplankton, these daily differences did not result in differences in final body size or lipid storage of these copepods. These copepods efficiently dealt with small and mesoscale variations in their food environment such that mesoscale structure in phytoplankton did not affect their final body size.     

Physical drivers of phytoplankton production in the southern Benguela upwelling system

Investigations of primary production (PP) were undertaken in the southern Benguela ecosystem during two research surveys in October 2006 and May 2007. Significant differences in environmental conditions, as well as biomass and PP, were observed between October and May. During October, integrated biomass and PP were significantly higher, ranging from 20.43 to 355.01 mg m⁻², and 0.71 to 6.98 g C m⁻² d⁻¹, respectively, than in May, where the range was 47.92–141.79 mg m⁻², and 0.70–3.35 g C m⁻² d⁻¹, respectively. Distribution patterns indicated low biomass and PP in newly upwelled water along the coast, higher biomass and PP in the mid-shelf region, while lower values were observed at and beyond the shelf edge. Latitudinal variations showed consistently higher biomass and PP in the St. Helena Bay region compared to biomass and PP south of Cape Town. During both surveys, phytoplankton communities were comprised primarily of diatoms and small flagellates, with no significant differences. Phytoplankton adaptation to environmental variability was characterised by increased P_m^B and E_k under elevated temperatures and irradiance, while no clear relationships were evident for α^B. Generalised Additive Models (GAMs) showed that photosynthetic parameters were all significant predictors of photosynthesis rates (P_z), with P_m^B being the most important, accounting for 36.97% of the deviance in P_z. However, biomass levels and environmental conditions exerted a much greater influence on P_z, with irradiance explaining the largest proportion (68.24%) of the deviance. Multiple predictor GAMs revealed that 96.26% of the deviance in P_z could be explained by a model which included nitrate, chlorophyll a, and irradiance.     

Exposure of pink salmon embryos to dissolved polynuclear aromatic hydrocarbons delays development,prolonging vulnerability to mechanical damage

Exposure to dissolved polynuclear aromatic hydrocarbons (PAHs) from crude oil delays pink salmon (Oncorhynchus gorbuscha) embryo development, thus prolonging their susceptibility to mechanical damage (shock). Exposure also caused mortality, edema, and anemia consistent with previous studies. Hatching and yolk consumption were delayed, indicating the rate of embryonic development was slowed by PAH exposure. The net result was that exposed embryos were more susceptible to shock than normal, unexposed embryos. Susceptibility to shock was protracted by 4–6 d for more than a month in embryos exposed to exponentially declining, dissolved PAH concentrations in water passed through oiled rock; the initial total PAH concentration was 22.4 μg L⁻¹ and the geometric mean concentration was 4.5 μg L⁻¹ over the first 20 d. Protracted susceptibility to shock caused by exposure to PAHs dissolved from oil could potentially increase the reported incidence of mortality in oiled stream systems, such as those in Prince William Sound after the Exxon Valdez oil spill, if observers fail to discriminate between direct mortality and shock-induced mortality.     

Top-down control of spring surface phytoplankton blooms by microzooplankton in the Central Yellow Sea,China

Phytoplankton growth and microzooplankton grazing were studied during the 2007 spring bloom in Central Yellow Sea. The surveyed stations were divided to pre-bloom phase (Chl a concentration less than 2 μg L⁻¹), and bloom phase (Chl a concentration greater than 2 μg L⁻¹). Shipboard dilution incubation experiments were carried out at 19 stations to determine the phytoplankton specific growth rates and the specific grazing rates of microzooplankton on phytoplankton. Diatoms dominated in the phytoplankton community in surface waters at most stations. For microzooplankton, Myrionecta rubra and tintinnids were dominant, and heterotrophic dinoflagellate was also important in the community. Phytoplankton-specific growth rates, with an average of 0.60±0.19 d⁻¹, were higher at pre-bloom stations (average 0.62±0.17 d⁻¹), and lower at the bloom stations (average 0.59±0.21 d⁻¹), but the difference of growth rates between bloom and pre-bloom stations was not statistically significant (t test, p=0.77). The phytoplankton mortality rate by microzooplankton grazing averaged 0.41±0.23 d⁻¹ at pre-bloom stations, and 0.58±0.31 d⁻¹ during the blooms. In contrast to the growth rates, the statistic difference of grazing rates between bloom and pre-bloom stations was significant (after removal of outliers, t test, p=0.04), indicating the importance of the top-down control in the phytoplankton bloom processes. Average potential grazing efficiency on primary productivity was 66% at pre-bloom stations and 98% at bloom stations, respectively. Based on our results, the biomass maximum phase (bloom phase) was not the maximum growth rate phase. Both phytoplankton specific growth rate and net growth rate were higher in the pre-bloom phase than during the bloom phase. Microzooplankton grazing mortality rate was positively correlated with phytoplankton growth rate during both phases, but growth and grazing were highly coupled during the booming phase. There was no correlation between phytoplankton growth rate and cell size during the blooms, but they were positive correlated during the pre-bloom phase. Our results indicate that microzooplankton grazing is an important process controlling the growth of phytoplankton in spring bloom period in the Central Yellow Sea, particularly in the “blooming” phase.     

High primary productivity and f-ratio in summer in the Ulleung basin of the East/Japan Sea

To better understand the cause of high summer primary productivity in the Ulleung Basin located in the southwest part of the East/Japan Sea, the spatial dynamics of primary, new, and regenerated productivities (PP, NP, and RP) were examined along the path of the Tsushima Warm Current system in summer 2008. We compared hydrographic and chemical parameters in the Ulleung Basin with those of the Kuroshio Current in the Western Pacific Ocean and the East China Sea. In summer, integrated primary productivity (IPP, 0.37–0.96 g C m⁻² d⁻¹) and integrated new productivity (INP, 26–221 mg N m⁻² d⁻¹) within the euphotic zone in the Ulleung Basin were higher than those in the East China Sea and the Western Pacific Ocean (0.17–0.28 g C m⁻² d⁻¹, 2−5 mg N m⁻² d⁻¹, respectively). In contrast, there was no pronounced spatial variation in integrated regenerated productivity (IRP, 43–824 mg N m⁻² d⁻¹). Strong positive correlations between IPP and INP (also the f-ratio), and between nitrate uptake rate in the mixed layer and nitrate upward flux through the top of pycnocline in summer in the Ulleung Basin imply that the high IPP was mainly supported by supply of nitrate from the underlying water in the euphotic zone. Shallowing of the pycnocline depth as the current enters the East/Japan Sea facilitates nitrate supply from the nutrient-replete cold water immediately below the pycnocline through nitrate upward flux. A subsurface maximum in PP at or above the pycnocline and a high f-ratio further support the importance of this source of nitrate for maintaining the high summer PP in the Ulleung Basin. In comparison, the high PP layer was observed at the surface in the following fall and spring in the Ulleung Basin. Our results demonstrate the importance of hydrographic features in enhancing PP in this oligotrophic Tsushima Warm Current system.     

Influence of sex and age on PCBs accumulation in the commercial fish Chelon labrosus

Thicklip grey mullet, Chelon labrosus, is an important commercial fish species and has been studied worldwide. However, no recent studies have been made regarding polychlorinated biphenyls (PCBs) in wild C. labrosus. Due to that, the concentration of 13 PCBs congeners was measured in muscles and livers, of males and females, of C. labrosus of different ages, allowing the estimation of PCB bioaccumulation throughout the species lifespan, in the Mondego estuary, a southern European temperate estuary. Male muscle sample concentrations ranged from 32 to 96 ng g^− 1 (lipid wt.) and in females from 32 to 62 ng g^− 1 (lipid wt.). In male liver sample concentrations ranged from 106 to 158 ng g^− 1 (lipid wt.), while female concentrations ranged from 88 to 129 ng g^− 1 (lipid wt.). The most abundant congeners presenting higher percentages in all samples were CB 138, 153 and 180. No significant differences were found between the concentrations in both sexes, but muscle and liver PCB concentrations in males tended to increase with age whereas in females concentrations remained stable throughout the species lifespan. Significant differences were found between concentrations in muscle and liver.     

Microzooplankton grazing impact in the Western Arctic Ocean

Microzooplankton grazing impact on phytoplankton was assessed using the Landry–Hassett dilution technique in the Western Arctic Ocean during spring and summer 2002 and 2004. Forty experiments were completed in a region encompassing productive shelf regions of the Chukchi Sea, mesotrophic slope regions of the Beaufort Sea off the North Slope of Alaska, and oligotrophic deep-water sites in the Canada Basin. A variety of conditions were encountered, from heavy sea-ice cover during both spring cruises, moderate sea-ice cover during summer of 2002, and light to no sea ice during summer of 2004, with a concomitant range of trophic conditions, from low chlorophyll-a (Chl-a; <0.5 μg L⁻¹) during heavy ice cover in spring and in the open basin, to late spring and summer shelf and slope open-water diatom blooms with Chl-a >5 μg L⁻¹. The microzooplankton community was dominated by large naked ciliates and heterotrophic gymnodinoid dinoflagellates. Significant, but low, rates of microzooplankton herbivory were found in half of the experiments. The maximum grazing rate was 0.16 d⁻¹ and average grazing rate, including experiments with no significant grazing, was 0.04±0.06 d⁻¹. Phytoplankton intrinsic growth rates varied from the highest values of about 0.4 d⁻¹ to the lowest values of zero to slightly negative growth, on average 0.16±0.15 d⁻¹. Light limitation in spring and post-bloom senescence during summer were likely explanations of observed low phytoplankton growth rates. Microzooplankton grazing consumed 0–120% (average 22±26%) of phytoplankton daily growth. Grazing and growth rates found in this study were low compared to rates reported in another Arctic system, the Barents Sea, and in major geographic regions of the world ocean.     

Time-series observation of POC fluxes estimated from 234Th in the northwestern North Pacific

Time-series measurements of ²³⁴Th activities and particulate organic carbon (POC) concentrations were made at time-series stations (K1, K2, K3, and KNOT) in the northwestern North Pacific from October 2002 to August 2004. Seasonal changes in POC export fluxes from the surface layer (∼100 m) were estimated using ²³⁴Th as a tracer. POC fluxes varied seasonally from approximately 0 to 180 mg C m⁻² d⁻¹ and were higher in spring–summer than in autumn–winter. The export ratio (e-ratio) ranged from 6% to 55% and was also higher in spring–summer. Annual POC fluxes were estimated to be 31 g C m⁻² y⁻¹ in the subarctic region (station K2) and 23 g C m⁻² y⁻¹ in the region between the subarctic and subtropical gyres (station K3). POC fluxes and e-ratios in the northwestern North Pacific were much higher than those in most other oceans. The annual POC flux corresponded to 69% of annual new production estimated from the seasonal difference of the nutrient in the Western Subarctic Gyre (45 g C m⁻² y⁻¹). These results indicate that much of the organic carbon assimilated in the surface layer of the northwestern North Pacific is transferred to the deep ocean in particulate form. Our conclusions support previous reports that diatoms play an important role in the biological pump.     

Biogenic fluxes in the Cariaco Basin: a combined study of sinking particulates and underlying sediments

The fluxes of total mass, organic carbon (OC), biogenic opal, calcite (CaCO₃) and long-chain C₃₇ alkenones (ΣAlk₃₇) were measured at three water depths (275, 455 and 930 m) in the Cariaco Basin (Venezuela) over three separate annual upwelling cycles (1996–1999) as part of the CARIACO sediment trap time-series. The strength and timing of both the primary and secondary upwelling events in the Cariaco Basin varied significantly during the study period, directly affecting the rates of primary productivity (PP) and the vertical transport of biogenic materials. OC fluxes showed a weak positive correlation (r²=0.3) with PP rates throughout the 3 years of the study. The fluxes of opal, CaCO₃ and ΣAlk₃₇ were strongly correlated (0.6<r²<0.8) with those of OC. The major exception was the lower than expected ΣAlk₃₇ fluxes measured during periods of strong upwelling. All sediment trap fluxes were significantly attenuated with depth, consistent with marked losses during vertical transport. Annually, strong upwelling conditions, such as those observed during 1996–1997, led to elevated opal fluxes (e.g., 35 g m⁻² yr⁻¹ at 275 m) and diminished ΣAlk₃₇ fluxes (e.g., 5 mg m⁻² yr⁻¹ at 275 m). The opposite trends were evident during the year of weakest upwelling (1998–1999), indicating that diatom and haptophyte productivity in the Cariaco Basin are inversely correlated depending on upwelling conditions.The analyses of the Cariaco Basin sediments collected via a gravity core showed that the rates of OC and opal burial (10–12 g m⁻² yr⁻¹) over the past 5500 years were generally similar to the average annual water column fluxes measured in the deeper traps (10–14 g m⁻² yr⁻¹) over the 1996–1999 study period. CaCO₃ burial fluxes (30–40 g m⁻² yr⁻¹), on the other hand, were considerably higher than the fluxes measured in the deep traps (∼10 g m⁻² yr⁻¹) but comparable to those obtained from the shallowest trap (i.e. 38 g m⁻² yr⁻¹ at 275 m). In contrast, the burial rates of ΣAlk₃₇ (0.4–1 mg m⁻² yr⁻¹) in Cariaco sediments were significantly lower than the water column fluxes measured at all depths (4–6 mg m⁻² yr⁻¹), indicating the large attenuation in the flux of these compounds at the sediment–water interface. The major trend throughout the core was the general decrease in all biogenic fluxes with depth, most likely due to post-depositional in situ degradation. The major exception was the relatively low opal fluxes (∼5 g m⁻² yr⁻¹) and elevated ΣAlk₃₇ fluxes (∼2 mg m⁻² yr⁻¹) measured in the sedimentary interval corresponding to 1600–2000 yr BP. Such compositions are consistent with a period of low diatom and high haptophyte productivity, which based on the trends observed from the sediment traps, is indicative of low upwelling conditions relative to the modern day.     

Basin-scale variability of phytoplankton biomass,production and growth in the Atlantic Ocean

The latitudinal distributions of phytoplankton biomass, composition and production in the Atlantic Ocean were determined along a 10,000-km transect from 50°N to 50°S in October 1995, May 1996 and October 1996. Highest levels of euphotic layer-integrated chlorophyll a (Chl a) concentration (75–125 mg Chl m⁻²) were found in North Atlantic temperate waters and in the upwelling region off NW Africa, whereas typical Chl a concentrations in oligotrophic waters ranged from 20 to 40 mg Chl m⁻². The estimated concentration of surface phytoplankton carbon (C) biomass was 5–15 mg C m⁻² in the oligotrophic regions and increased over 40 mg C m⁻² in richer areas. The deep chlorophyll maximum did not seem to constitute a biomass or productivity maximum, but resulted mainly from an increase in the Chl a to C ratio and represented a relatively small contribution to total integrated productivity. Primary production rates varied from 50 mg C m⁻² d⁻¹ at the central gyres to 500–1000 mg C m⁻² d⁻¹ in upwelling and higher latitude regions, where faster growth rates (μ) of phytoplankton (>0.5 d⁻¹) were also measured. In oligotrophic waters, microalgal growth was consistently slow [surface μ averaged 0.21±0.02 d⁻¹ (mean±SE)], representing <20% of maximum expected growth. These results argue against the view that the subtropical gyres are characterized by high phytoplankton turnover rates. The latitudinal variations in μ were inversely correlated to the changes in the depth of the nitracline and positively correlated to those of the integrated nitrate concentration, supporting the case for the role of nutrients in controlling the large-scale distribution of phytoplankton growth rates. We observed a large degree of temporal variability in the phytoplankton dynamics in the oligotrophic regions: productivity and growth rates varied in excess of 8-fold, whereas microalgal biomass remained relatively constant. The observed spatial and temporal variability in the biomass specific rate of photosynthesis is at least three times larger than currently assumed in most satellite-based models of global productivity.     

Fluxes and exchange of suspended sediment in tidal inlets draining a degraded mangrove forest in Kenya

This study focuses on sediment exchange in the degraded Mwache mangrove forest wetland located in southern Kenya. It involved measurement of total and particulate organic suspended sediment concentrations (TSSC and POSC), tidal water elevation and current velocities. Results showed that in the heavily degraded backwater zone mangrove forest, the ebb and flood tide total sediment fluxes were of same order of magnitude, however, flood tide sediment fluxes were slightly higher than the ebb ones. In the moderately degraded frontwater zone mangrove forest, the flood tide sediment fluxes were more than 50% higher than the ebb tide fluxes. The peak net sedimentation in the highly degraded backwater zone was 4 g m⁻² tide⁻¹ but that in the moderately degraded frontwater zone was 63 g m⁻² tide⁻¹. In the frontwater zone of the mangrove forest, the peak instantaneous ebb tide sediment flux was 3206 kg tide⁻¹ equivalent to 35.6 g m⁻² tide⁻¹ and the flood one 8574 kg tide⁻¹ (95 g m⁻² tide⁻¹). The peak instantaneous flood and ebb tide particulate organic sediment (POS) fluxes in the frontwater zone mangrove forest were 1316 kg tide⁻¹ (15 g m⁻² tide⁻¹) and 587 kg tide⁻¹ (6.5 g m⁻² tide⁻¹), respectively. The peak ebb and flood tide sediment fluxes in the backwater mangrove forest were 3206 kg tide⁻¹ (36 g m⁻² tide⁻¹) and 3305 kg tide⁻¹ (36.7 g m⁻² tide⁻¹), respectively. In case of POS fluxes in the backwater zone mangrove forest, the peak flood period POS flux was 969 kg tide⁻¹ (10.7 g m⁻² tide⁻¹) while the ebb period one was 484 kg tide⁻¹ (5.4 g m⁻² tide⁻¹). In both highly degraded backwater and moderately degraded frontwater zone of the mangrove forest, there is net import of sediments. However, the net import is relatively lower in the backwater zone forest where the trapping efficiency is 27%. In the moderately degraded frontwater zone of the mangrove forest, the sediment trapping efficiency is 65%. The net sediment import occurs mainly in periods of high river discharge in both neap and spring tides, but occurs only in spring tides during dry season. The net accretion rates in the backwater and frontwater zone mangrove forests are 0.25 and 3.5 cm year⁻¹, respectively.     

Particulate organic carbon export fluxes and size-fractionated POC/234Th ratios in the Ligurian,Tyrrhenian and Aegean Seas

Measurements of ²³⁴Th/²³⁸U disequilibria and particle size-fractionated (1, 10, 20, 53, 70, 100 μm) organic C and ²³⁴Th were made to constrain estimates of the export flux of particulate organic C (POC) from the surface waters of the Ligurian, Tyrrhenian and Aegean Seas in March–June 2004. POC exported from the surface waters (75–100 m depth) averaged 9.2 mmol m⁻² d⁻¹ in the Ligurian and Tyrrhenian Seas (2.3±0.5–14.9±3.0 mmol m⁻² d⁻¹) and 0.9 mmol m⁻² d⁻¹ in the Aegean Sea. These results are comparable to previous measurements of ²³⁴Th-derived and sediment-trap POC fluxes from the upper 200 m in the Mediterranean Sea. Depth variations in the POC/²³⁴Th ratio suggest two possible controls. First, decreasing POC/²³⁴Th ratios with depth were attributed to preferential remineralization of organic C. Second, the occurrence of maxima or minima in the POC/²³⁴Th ratio near the DCM suggests influence by phytoplankton dynamics. To assess the accuracy of these data, the empirical ²³⁴Th-method was evaluated by quantifying the extent to which the ²³⁴Th-based estimate of POC flux, P_POC, deviates from the true flux, F_POC, defined as the p-ratio (p-ratio=P_POC/F_POC=S_Th/S_POC, where S=particle sinking rate). Estimates of the p-ratio made using Stokes’ Law and the particle size distributions of organic C and ²³⁴Th yield values ranging from 0.93–1.45. The proximity of the p-ratio to unity implies that differences in the sinking rates of POC- and ²³⁴Th-carrying particles did not bias ²³⁴Th-normalized POC fluxes by more than a factor of two.     

Latitudinal distribution of microbial plankton abundance,production, and respiration in the Equatorial Atlantic in autumn 2000

Phytoplankton and bacterial abundance, size-fractionated phytoplankton chlorophyll-a (Chl-a) and production together with bacterial production, microbial oxygen production and respiration rates were measured along a transect that crossed the Equatorial Atlantic Ocean (10°N–10°S) in September 2000, as part of the Atlantic Meridional Transect 11 (AMT 11) cruise. From 2°N to 5°S, the equatorial divergence resulted in a shallowing of the pycnocline and the presence of relatively high nitrate (>1 μM) concentrations in surface waters. In contrast, a typical tropical structure (TTS) was found near the ends of the transect. Photic zone integrated ¹⁴C primary production ranged from ∼200 mg C m⁻² d⁻¹ in the TTS region to ∼1300 mg C m⁻² d⁻¹ in the equatorial divergence area. In spite of the relatively high primary production rates measured in the equatorial upwelling region, only a moderate rise in phytoplankton biomass was observed as compared to nearby nutrient-depleted areas (22 vs. 18 mg Chl-a m⁻², respectively). Picophytoplankton were the main contributors (>60%) to both Chl-a biomass and primary production throughout the region. The equatorial upwelling did not alter the phytoplankton size structure typically found in the tropical open ocean, which suggests a strong top-down control of primary producers by zooplankton. However, the impact of nutrient supply on net microbial community metabolism, integrated over the euphotic layer, was evidenced by an average net microbial community production within the equatorial divergence (1130 mg C m⁻² d⁻¹) three-fold larger than net production measured in the TTS region (370 mg C m⁻² d⁻¹). The entire region under study showed net autotrophic community metabolism, since respiration accounted on average for 51% of gross primary production integrated over the euphotic layer.     

Microbial biomass and viral infections of heterotrophic prokaryotes in the sub-surface layer of the central Arctic Ocean

Seawater samples were collected for microbial analyses between 55 and 235 m depth across the Arctic Ocean during the SCICEX 97 expedition (03 September–02 October 1997) using a nuclear submarine as a research platform. Abundances of prokaryotes (range 0.043–0.47×10⁹ dm⁻³) and viruses (range 0.68–11×10⁹ dm⁻³) were correlated (r=0.66, n=150) with an average virus:prokaryote ratio of 26 (range 5–70). Biomass of prokaryotes integrated from 55 to 235 m ranged from 0.27 to 0.85 g C m⁻² exceeding that of phytoplankton (0.005–0.2 g C m⁻²) or viruses (0.02–0.05 g C m⁻²) over the same depth range by an order of magnitude on average. Using transmission electron microscopy (TEM), we estimated that 0.5% of the prokaryote community on average (range 0–1.4%) was visibly infected with viruses, which suggests that very little of prokaryotic secondary production was lost due to viral lysis. Intracellular viruses ranged from 5 to >200/cell, with an average apparent burst size of 45±38 (mean±s.d.; n=45). TEM also revealed the presence of putative metal-precipitating bacteria in 8 of 13 samples, which averaged 0.3% of the total prokaryote community (range 0–1%). If these prokaryotes are accessible to protistan grazers, the Fe and Mn associated with their capsules might be an important source of trace metals to the planktonic food web. After combining our abundance and mortality data with data from the literature, we conclude that the biomass of prokaryoplankton exceeds that of phytoplankton when averaged over the upper 250 m of the central Arctic Ocean and that the fate of this biomass is poorly understood.     

Fate of biogenic carbon in the upper 200 m of the central Greenland Sea

Sedimentation of particulate carbon from the upper 200–300 m in the central Greenland Sea from August 1993 to June 1995 was less than 2 g C m⁻² yr⁻¹. Daily rates of sedimentation of particulate organic carbon reached highest values of about 18 mg m⁻² d⁻¹ in fall 1994. For total particulate material, maximum rates of sedimentation of about 250 mg m⁻² d⁻¹ were recorded in spring and fall 1994. For chlorophyll equivalent, highest rates of sedimentation of about 140 μg m⁻² d⁻¹ were recorded in spring 1994. As reported in related investigations, the transient accumulation of DOC in surface waters during summer, as well as respiration and mortality of deep overwintering zooplankton stocks, appeared to dominate the fate of photosynthetically fixed organic carbon. The above processes may account for roughly 43 g C m⁻² in the upper 200 m of the central Greenland Sea. For comparison, the seasonal deficit in dissolved inorganic carbon was reported to be about 23 g C m⁻² in the upper 20 m of surface water, and estimates for new annual production were reported to be about 57 g C m⁻². In our investigation, the biological carbon pump was not unusually effective in transporting carbon out of the productive surface layer.     

Nutrient and phytoplankton dynamics off the west coast of Vancouver Island during the 1997/98 ENSO event

Six research cruises were conducted off the west coast of Vancouver Island between April and October of 1997 and 1998 as part of the Canadian GLOBEC project to compare nutrient and phytoplankton dynamics between ENSO (1997) and non-ENSO (1998) years. Limited sampling also was conducted during three cruises in 1999. During the 1997 ENSO period, there was a shallow thermocline (∼10 m) that resulted in a shallower mixed layer, lower salinity and density, and stronger summer stratification. In general on the shelf, the 1997 growing season was characterized by higher nitrate (7.5 μM) and silicic acid (17 μM) concentrations, lower total chlorophyll (∼76 mg m⁻²), lower phytoplankton carbon biomass (0.2 mg C L⁻¹), and lower diatom abundance and biomass than in 1998. Phytoplankton assemblages were dominated by nanoplankton in 1997 and by diatoms in 1998. These results suggest that the 1997 ENSO was responsible for a reduction in the growth and biomass of larger phytoplankton cells. In mid-1998, the hydrographic characteristics off the west coast of Vancouver Island changed suddenly. The 1997 poleward transport of warm water reversed to an equatorward transport of coastal water in July 1998, which was accompanied by normal summer upwelling. During 1998, a large diatom bloom (mainly dominated by Chaetoceros debilis, Leptocylindrus danicus and to a lesser extent by Skeletomema and Pseudo-nitzschia sp.) was observed in July over the continental shelf. This large bloom resulted in chlorophyll concentrations of up to 400 mg m⁻², primary productivity of up to 11 g C m⁻² d⁻¹, and near undetectable dissolved nitrogen concentrations at some of the shelf stations in 1998. In contrast, during 1997, the sub-tropical waters that were advected over the slope, resulted in low chlorophyll a and primary productivity (generally <1 g C m⁻² d⁻¹). Therefore, there was a sharp contrast between the very high primary productivity on the shelf in July 1998, due to normal nutrient replenishment from summer upwelling and outflow from the Strait of Juan de Fuca, and the lower primary productivity during the 1997 ENSO year. During 1998, non-ENSO conditions resulted in phytoplankton biomass that was twice as high on the shelf as that measured in regions beyond the continental shelf of the west coast of Vancouver Island.     

Phytoplankton dynamics in the NE subarctic Pacific

Ocean Station Papa (OSP, 50°N 145°W) in the NE subarctic Pacific is characterised as high nitrate low chlorophyll (HNLC). However, little is known about the spatial extent of these HNLC waters or the phytoplankton dynamics on the basin scale. Algal biomass, production and size-structure data are presented from winter, spring and summer between 1992 and 1997 for five stations ranging from coastal to open-ocean conditions. The inshore stations (P04–P16) are characterised by the classical seasonal cycle of spring and late summer blooms (production >3 g C m⁻² d⁻¹), diatoms are not Fe-stressed, and growth rate is probably controlled by macronutrient supply. The fate of the phytoplankton is likely sedimentation by diatom-dominated spring blooms, with a pelagic recycling system predominating at other times. The offshore stations (P20/OSP) display low seasonality in biomass and production (OSP, mean winter production 0.3 g C m⁻² d⁻¹, mean spring/summer production 0.85 g C m⁻² d⁻¹), and are dominated by small algal cells. Low Fe availability prevents the occurrence of diatom blooms observed inshore. The main fate of phytoplankton is probably recycling through the microbial food web, with relatively low sedimentation compared to inshore. However, the supply of macro- and micro-nutrients to the coastal and open ocean, respectively, may vary between years. Variability in macro-nutrient supply to the coastal ocean may result in decreased winter reserve nitrate, summer nitrate limitation, subsequent floristic shifts towards small cells, and reduced primary production. Offshore, higher diatom abundances are occasionally observed, perhaps indicating episodic Fe supply. The two distinct oceanic regimes have different phytoplankton dynamics resulting in different seasonality, community structure and fate of algal carbon. These differences will strongly influence the biogeochemical signatures of the coastal and open-oceanic NE subarctic Pacific.