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.     

Dissolved inorganic carbon,alkalinity, nutrient and oxygen seasonal and interannual variations at the Antarctic Ocean JGOFS-KERFIX site

JGOFS-KERFIX (KERguelen point FIXe) time-series station, located south of the polar front in the Indian sector of the Antarctic Ocean, was occupied monthly between January 1990 and March 1995. Annual cycles of dissolved inorganic carbon (DIC), total alkalinity (TALK), oxygen (O₂) and nutrients (nitrate, silicate, phosphate and ammonia) in the upper ocean are presented for this site. From seasonal drawdown of nutrients and DIC, we estimate a spring–summer net community production of 3.2±0.5 mol m⁻² and C/N/P ratios of 100/16/1. The Si/N ratio varies between 1.8 and 3, suggesting low iron concentrations. The spring–summer biogenic silicon export derived from silicate drawdown is 1.18 mol m⁻², consistent with model estimates of silicate export at this site. Seasonal and interannual variations of oxygen, nitrate and DIC due to physical and biological processes are quantified using a simple month-to-month budget formulation. From these budgets, an annual net community production of 5.7±3.3 mol m⁻² yr⁻¹ is estimated, about twice the averaged spring–summer production, indicating that, at KERFIX, there is a positive net community production throughout the year. Air–sea CO₂ fluxes show that KERFIX is a strong CO₂ sink for the atmosphere of 2.4–5.1 mol m⁻² yr⁻¹ in 1993, depending on the gas exchange formulation used. A 2.1–3.3 mol m⁻² yr⁻¹ outgassing of O₂ is observed at KERFIX except in 1993 and 1994 where a decreasing trend of temperature induces an increase of O₂ solubility.     

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.     

Interannual variability of Si and N cycles at the time-series station KERFIX between 1990 and 1995 – a 1-D modelling study

Interannual variability of nutrients and plankton cycles were studied at the time-series station KERFIX (50°40′S, 68°25′E) using a 1-D coupled physical-biogeochemical model that is descended from that of Pondaven et al. (1998). At KERFIX, a high half saturation constant for silicic acid uptake (K_Si) and a high Si/N uptake ratio are required to reproduce the Si and N cycles. Although very high in comparison with most data from temperate systems, these values are consistent with K_Si and Si/N uptake ratios measured in the Indian sector of the Southern Ocean. Past and recent finding on the role of light and iron limitation on nutrient consumption ratios might explain these “unusual” silicon uptake kinetic parameters. Comparison of model results with observations show that the model correctly reproduces the observed interannual variability of nutrients and plankton cycles at KERFIX between 1992 and 1995. Characteristic features of this region are a spring phytoplankton bloom of 1.0–1.5 mg Chlorophyll a m⁻³ and a net excess of silicic acid utilisation over that of nitrate. This high silicic acid utilisation leads to low Si concentrations in late summer and subsequent Si limitation of diatom growth. The interannual variability of production of silicon and nitrogen predicted by the model is 1.93±0.04 mol Si m⁻² yr⁻¹ and 1.35±0.07 mol N m⁻² yr⁻¹ (±SD). In parallel, the predicted export is 1.12±0.04 mol Si m⁻² yr⁻¹ and 0.06±0.01 mol N m⁻² yr⁻¹. It is shown that diatoms may contribute significantly to export if diatom sinking is taken into account. An interannual variability of the predicted Si and N cycles is detected. This variability is associated with changes in the mixed layer properties, which have been documented to be linked to the Pacific El Niño Southern Oscillation or displacement of the Polar Front.     

Picophytoplankton dynamics and production in the Arabian Sea during the 1995 Southwest Monsoon

Phytoplankton community structure is expected to shift to larger cells (e.g., diatoms) with monsoonal forcing in the Arabian Sea, but recent studies suggest that small primary producers remain active and important, even in areas strongly influenced by coastal upwelling. To better understand the role of smaller phytoplankton in such systems, we investigated growth and grazing rates of picophytoplankton populations and their contributions to phytoplankton community biomass and primary productivity during the 1995 Southwest Monsoon (August–September). Environmental conditions at six study stations varied broadly from open-ocean oligotrophic to coastal eutrophic, with mixed-layer nitrate and chlorophyll concentrations ranging from 0.01 to 11.5 μM NO₃ and 0.16 to 1.5 μg Chl a. Picophytoplankton comprised up to 92% of phytoplankton carbon at the oceanic stations, 35% in the diatom-dominated coastal zone, and 26% in a declining Phaeocystis bloom. Concurrent in situ dilution and ¹⁴C-uptake experiments gave comparable ranges of community growth rates (0.53–1.05 d⁻¹ and 0.44–1.17 d⁻¹, to the 1% light level), but uncertainties in C:Chl a confounded agreement at individual stations. Microzooplankton grazing utilized 81% of community phytoplankton growth at the oligotrophic stations and 54% at high-nutrient coastal stations. Prochlorococcus (PRO) was present at two oligotrophic stations, where its maximum growth approached 1.4 d⁻¹ (two doublings per day) and depth-integrated growth varied from 0.2 to 0.8 d⁻¹. Synechococcus (SYN) growth ranged from 0.5 to 1.1 d⁻¹ at offshore stations and 0.6 to 0.7 d⁻¹ at coastal sites. Except for the most oligotrophic stations, growth rates of picoeukaryotic algae (PEUK) exceeded PRO and SYN, reaching 1.3 d⁻¹ offshore and decreasing to 0.8 d⁻¹ at the most coastal station. Microzooplankton grazing impact averaged 90, 70, and 86% of growth for PRO, SYN, and PEUK, respectively. Picoplankton as a group accounted for 64% of estimated gross carbon production for all stations, and 50% at high-nutrient, upwelling stations. Prokaryotes (PRO and SYN) contributed disproportionately to production relative to biomass at the most oligotrophic station, while PEUK were more important at the coastal stations. Even during intense monsoonal forcing in the Arabian Sea, picoeukaryotic algae appear to account for a large portion of primary production in the coastal upwelling regions, supporting an active community of protistan grazers and a high rate of carbon cycling in these areas.     

Microzooplankton grazing and selective feeding during bloom periods in the Tolo Harbour area as revealed by HPLC pigment analysis

Dilution experiments were conducted to investigate microzooplankton grazing impact on phytoplankton of different taxonomic groups and size fractions (< 5, 5–20, 20–200 μm) during spring and summer bloom periods at two different sites (inner Tolo Harbour and Tolo Channel) in the Tolo Harbour area, the northeastern coastal area of Hong Kong. Experiments combined with HPLC pigment analysis in three phytoplankton size fractions measured pigment and size specific phytoplankton growth rates and microzooplankton grazing rates. Pigment-specific phytoplankton growth rates ranged between 0.08 and 3.53 d^− 1, while specific grazing rates of microzooplankton ranged between 0.07 and 2.82 d^− 1. Highest specific rates of phytoplankton growth and microzooplankton grazing were both measured in fucoxanthin in 5–20 μm size fraction in inner Tolo Harbour in summer, which coincided with the occurrence of diatom bloom. Results showed significant correlations between phytoplankton growth and microzooplankton grazing rates. Microzooplankton placed high grazing pressure on phytoplankton community. High microzooplankton grazing impact on alloxanthin (2.63–5.13) suggested strong selection toward cryptophytes. Our results provided no evidence for size selective grazing on phytoplankton by microzooplankton.     

Factors controlling silicon and nitrogen biogeochemical cycles in high nutrient,low chlorophyll systems (the Southern Ocean and the North Pacific): Comparison with a mesotrophic system (the North Atlantic)

A 1-D coupled physical-biogeochemical model is used to study the seasonal cycles of silicon and nitrogen in two High Nutrient Low Chlorophyll (HNLC) systems, the Antarctic Circumpolar Current (ACC) and the North Pacific Ocean, and a mesotrophic system, the North Atlantic Ocean. The biological model consists of nine compartments (diatoms, nano-flagellates, microzooplankton, mesozooplankton, two types of detritus, nitrate, ammonium and silicic acid) forced by irradiance, temperature, mixing and deep nitrate and silicic acid concentrations. At all sites, nanophytoplankton standing crop variations are low, in spite of variations in primary production, because of a “top–down” control by microzooplankton. Although nanophytoplankton sustain more than 60% of the annual primary production in these areas, their contribution to the export production does not exceed 1% of the total. The differences in the seasonal plankton cycle among these regions come mainly from differences in the dynamics of large phytoplankton (here diatoms). In the ACC, the chlorophyll maximum remains <1.5 mg m⁻³, as an unfavourable light/mixing regime and a likely trace-metal limitation keep diatoms from blooming. In the northeast Pacific, trace-metal limitation seems to keep diatoms from blooming throughout the year. In both these systems, light or iron limitations induce high Si/N uptake ratios. Incidentally these high Si/N uptake ratios lead to a net excess of silicic acid utilization over nitrate, and to a subsequent silicic acid limitation during the summertime. In the North Atlantic, under favourable light/mixing regime and nutrient-replete conditions at the onset of the growing period, diatoms outburst and sustain a bloom >3.5 mg Chl-a m⁻³. Thereafter, mesozooplankton grazing pressure and silicic acid limitation induce the collapse of the chlorophyll maximum and the persistence of lower chlorophyll concentrations in summer. Although the ACC and the North Pacific show HNLC features, they support a high biogenic silica production (1.9 and 1.07 mol Si m⁻² yr⁻¹) and export flux (0.79 and 0.61 mol Si m⁻² yr⁻¹), compared to the North Atlantic (production: 0.23 mol Si m⁻² yr⁻¹, export: 0.12 mol Si m⁻² yr⁻¹). The differences in Si production and export between the HNLC systems and the mesotrophic North Atlantic come from both higher Si concentrations and Si/N uptake ratios in the HNLC areas compared to the North Atlantic. Also, the low dissolution rate of biogenic silica compared to nitrogen degradation rate, and the inhibition of nitrate uptake by ammonium, reinforce the net excess of silicic acid utilization over nitrate. As a result, the model also illustrates the efficiency of the silica pump for the three sites: about 50% of the biogenic silica synthesized in the euphotic layer is exported out of the first 100 m, while only 4–11% of the particulate organic nitrogen escapes recycling in the surface layer.     

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.     

Phytoplankton of the western Arctic in the spring and summer of 2002: Structure and seasonal changes

We analyzed the taxonomic structure and spatial variability of phytoplankton abundance and biomass in the Chukchi and Beaufort Seas during spring and summer seasons of the SBI program. Phytoplankton samples were collected during two surveys from May 10 to June 13 and from July 19 to August 21 of 2002. In May and June, ice cover exceeded 80% over most of the study area and there was no vertical stratification, indicating that the successional state of the phytoplankton corresponded to the end of the winter biological season. The phytoplankton abundance ranged from a few tens to a few thousands of cells per liter, while biomass varied from 0.1 to 3.0 mg C m⁻³. Small areas of high phytoplankton abundance (0.13–1.3×10⁶ cells L⁻¹) and biomass (22–536 mg C m⁻³), dominated by early spring diatoms Pauliella taeniata and Fragilariopsis oceanica in the surface waters, which indicated the beginning of the spring bloom, were observed only in the southeastern part of the Chukchi shelf and off Point Barrow. In July and August summer period, more than a half of the study area had <50% ice cover and the water column was stratified by temperature and salinity. Over the Chukchi shelf and continental slope of the Beaufort Sea, the phytoplankton abundance and biomass were an order of magnitude higher in July–August than in May–June. The taxonomic diversity of algae also increased due to the appearance of late-spring and summer diatoms, dinoflagellates, and coccolithophorids (Emiliania huxleyi). Interestingly, the seasonal differences between phytoplankton abundance and taxonomic composition in the spring and summer periods varied the least over the Chukchi Sea slope and in the deep-water area of the Arctic Ocean. High algae concentrations in summer were located in the lower layers of the euphotic zone, suggesting that the spring bloom on both the Chukchi shelf and in the western part of the Beaufort Sea occurred in late June/early July. In the spring and summer, the microalgal community was characterized by a high abundance of 4–10 μm flagellates, which exceeded the abundance of all other taxonomic groups. In both seasons studied, phytoplankton reached its maximum abundance within restricted areas in the southern part of the Chukchi Sea southwest of Point Hope, in the northern part of the Chukchi shelf between the 50- and 100-m isobaths, on the shelf northwest of Point Barrow, and over the continental slope in the Beaufort Sea. The pronounced spatial difference in the seasonal state was a characteristic feature of the phytoplankton community in the western Arctic.     

Seasonal and interannual variability in particle fluxes of carbon,nitrogen and silicon from time series of sediment traps at Ocean Station P, 1982–1993: relationship to changes in subarctic primary productivity

An extended time series of particle fluxes at 3800 m was recorded using automated sediment traps moored at Ocean Station Papa (OSP, 50°N, 145°W) in the northeast Pacific Ocean for more than a decade (1982–1993). Time-series observations at 200 and 1000 m, and short-term measurements using surface-tethered free-drifting sediment traps also were made intermittently. We present data for fluxes of total mass (dry weight), particulate organic carbon (POC), particulate organic nitrogen (PON), biogenic Si (BSi), and particulate inorganic carbon (PIC) in calcium carbonate. Mean monthly fluxes at 3800 m showed distinct seasonality with an annual minimum during winter months (December–March), and maximum during summer and fall (April–November). Fluxes of total mass, POC, PIC and BSi showed 4-, 10-, 7- and 5-fold increases between extreme months, respectively. Mean monthly fluxes of PIC often showed two plateaus, one in May–August dominated by <63 μm particles and one in October–November, which was mainly >63 μm particles. Dominant components of the mass flux throughout the year were CaCO₃ and opal in equal amounts. The mean annual fluxes at 3800 m were 32±9 g dry weight g m⁻² yr⁻¹, 1.1±0.5 g POC m⁻² yr⁻¹, 0.15±0.07 g PON m⁻² yr⁻¹, 5.9±2.0 g BSi m⁻² yr⁻¹ and 1.7±0.6 g PIC m⁻² yr⁻¹. These biogenic fluxes clearly decreased with depth, and increased during “warm” years (1983 and 1987) of the El Niño, Southern Oscillation cycle (ENSO). Enhancement of annual mass flux rates to 3800 m was 49% in 1983 and 36% in 1987 above the decadal average, and was especially rich in biogenic Si. Biological events allowed estimates of sinking rates of detritus that range from 175 to 300 m d⁻¹, and demonstrate that, during periods of high productivity, particles sink quickly to deep ocean with less loss of organic components. Average POC flux into the deep ocean approximated the “canonical” 1% of the surface primary production.     

Oxygen dynamics in the North Atlantic subtropical gyre

Dissolved oxygen (DO) in the ocean is a tracer for most ocean biogeochemical processes including net community production and remineralization of organic matter which in turn constrains the biological carbon pump. Knowledge of oxygen dynamics in the North Atlantic Ocean is mainly derived from observations at the Bermuda Atlantic Time-series Study (BATS) site located in the western subtropical gyre which may skew our view of the biogeochemistry of the subtropical North Atlantic. This study presents and compares a 15 yr record of DO observations from ESTOC (European Station for Time-Series in the Ocean, Canary Islands) in the eastern subtropical North Atlantic with the 20 yr record at BATS. Our estimate for net community production of oxygen was 2.3±0.4 mol O₂ m⁻² yr⁻¹ and of oxygen consumption was −2.3±0.5 mol O₂ m⁻² yr⁻¹ at ESTOC, and 4 mol O₂ m⁻² yr⁻¹ and −4.4±1 mol m⁻² yr⁻¹ at BATS, respectively. These values were determined by analyzing the time-series using the Discrete Wavelet Transform (DWT) method. These flux values agree with similar estimates from in-situ observational studies but are higher than those from modeling studies. The difference in net oxygen production rates supports previous observations of a lower carbon export in the eastern compared to the western subtropical Atlantic. The inter-annual analysis showed clear annual cycles at BATS whereas longer cycles of nearly 4 years were apparent at ESTOC. The DWT analysis showed trends in DO anomalies dominated by long-term perturbations at a basin scale for the consumption zones at both sites, whereas yearly cycles dominated the production zone at BATS. The long-term perturbations found are likely associated with ventilation of the main thermocline, affecting the consumption and production zones at ESTOC.     

Structure of potential upwelling areas in the Philippines

The three-dimensional structure of two potential mesoscale upwelling areas that are located in the external waters of the Philippine archipelago (i.e. northwest of Luzon and east of Mindanao) were constructed by analysis of historical data. A unique characteristic of both upwelling sites is that they can be identified by their anomalously cold subsurface temperatures rather than sea surface temperature distributions. As such, they cannot be observed in sea surface temperature fields derived by satellite imagery. The data used in the analysis were obtained from the National Oceanographic Data Center hydrographic database. Objective analysis was performed to produce monthly temperature fields at several standard depths within the upper 500 m of the region 0–30°N and 100–140°E with a horizontal grid resolution of 0.5°. The extent and timing of these upwelling areas are described. A review of existing hypothesis on the mechanisms for their evolution and seasonal modulation are presented. The change in heat content during the upwelling season is greater than 300 W m⁻² in both areas. Based on the excursion of isotherms, vertical velocities of 83 cm day⁻¹ and 26 cm day⁻¹ were obtained for upwelling northwest of Luzon and east of Mindanao, respectively.     

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.     

Modelling phytoplankton deposition to Chesapeake Bay sediments during winter–spring: interannual variability in relation to river flow

The often-rapid deposition of phytoplankton to sediments at the end of the spring phytoplankton bloom is an important component of benthic–pelagic coupling in temperate and high latitude estuaries and other aquatic systems. However, quantifying the flux is difficult, particularly in spatially heterogeneous environments. Surficial sediment chlorophyll-a, which can be measured quickly at many locations, has been used effectively by previous studies as an indicator of phytoplankton deposition to estuarine sediments. In this study, surficial sediment chlorophyll-a was quantified in late spring at 20–50 locations throughout Chesapeake Bay for 8 years (1993–2000). A model was developed to estimate chlorophyll-a deposition to sediments using these measurements, while accounting for chlorophyll-a degradation during the time between deposition and sampling. Carbon flux was derived from these estimates via C:chl-a = 75.Bay-wide, the accumulation of chlorophyll-a on sediments by late spring averaged 171 mg m⁻², from which the chlorophyll-a and carbon sinking fluxes, respectively, were estimated to be 353 mg m⁻² and 26.5 gC m⁻². These deposition estimates were ∼50% of estimates based on a sediment trap study in the mid-Bay. During 1993–2000, the highest average chlorophyll-a flux was in the mid-Bay (248 mg m⁻²), while the lowest was in the lower Bay (191 mg m⁻²). Winter–spring average river flow was positively correlated with phytoplankton biomass in the lower Bay water column, while phytoplankton biomass in that same region of the Bay was correlated with increased chlorophyll-a deposition to sediments. Responses in other regions of the Bay were less clear and suggested that the concept that nutrient enrichment in high flow years leads to greater phytoplankton deposition to sediments may be an oversimplification. A comparison of the carbon flux associated with the deposition of the spring bloom with annual benthic carbon budgets indicated that the spring bloom did not contribute a disproportionately large fraction of annual carbon inputs to Chesapeake Bay sediments. Regional patterns in chlorophyll-a deposition did not correspond with the strong regional patterns that have been found for plankton net community metabolism during spring.     

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.     

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.     

Carbon dioxide in surface seawater of the eastern North Pacific Ocean (Line P), 1973–2005

Partial pressure of CO₂ (pCO₂) in surface seawater has been measured in the northeastern Pacific Ocean at Station P and along Line P since 1973. These data have been divided into ‘oceanic’ and ‘coastal/transition’ zones, and the seasonal and interannual variability and the long-term trends for each zone have been examined. The oceanic zone shows little seasonality in surface seawater pCO₂, with undersaturation throughout the year. A strong, biologically-driven seasonal cycle is offset by variation in temperature-dependent solubility of CO₂. The coastal/transition zone shows a decline in pCO₂ from winter–spring through summer and fall that is likely the result of seasonal stratification and convection rather than coastal upwelling. Interannual variability all along Line P is correlated with the multivariate ENSO index (MEI), with lower seawater pCO₂ associated with El Niño conditions. Correlations with the Pacific Decadal Oscillation Index are similar but weaker, in part because there are few data prior to the 1976 regime shift. The long-term trend in seawater pCO₂ in the oceanic zone is +1.36±0.16 μatm year^?1, indistinguishable from the atmospheric growth rate, and varies little among the seasons. In the coastal/transition zone a slow increase in the pCO₂ of surface seawater relative to that of the atmosphere has led to increasing undersaturation, particularly in spring. Aliasing of the seasonal and interannual variability due to sampling frequency may explain part of the observed trend in the coastal/transition zone, but real changes in physical or biological processes are also possible and require more detailed study.     

Winter and spring phytoplankton composition and production in a shallow eutrophic Baltic lagoon

Taxonomic composition and productivity of winter and spring phytoplankton in a eutrophic estuary have been investigated in order to elucidate the carbon flux under conditions of limitation by physical factors – light and temperature. In spite of the important differences in nutrients, solar radiation and water temperature between winter and spring season, mean concentrations of particulate organic carbon were equal to 13.2 and 13.0 mgC l⁻¹, respectively. Chlorophyll a averaged at 79 μgChl l⁻¹ in winter, that is 69% of spring. Although community respiration accounted for only 6–26% of light saturated photosynthesis, integrated net primary production of the 1.2 m deep water column was negative until April. High attenuation of the water body (K_o = 2.9 m⁻¹) lead to a negative carbon balance (net heterotrophy) below 35 cm for all sampling dates. Thus, the high winter POC and phytoplankton values can only originate from summer or autumn primary production. This assumption was supported by a carbon loss rate of just 3% of total organic carbon per day for the whole water column. The composition of phytoplankton was very constant through both seasons: 39% Chlorophyceae, 33% Cyanobacteria and 25% Bacillariophyceae. As expected, phytoplankton was low light acclimated, having high α values (slope of light limited photosynthesis), but moderate maximum photosynthesis rates at saturating irradiances, which were heavily affected by temperature. Calculation of net carbon flux yet showed net heterotrophy of the Bodden waters in winter and early spring were caused by external physical limitation (low surface irradiance and low temperature) in combination with a high light attenuation of the water body.     

Sea-ice algae: Major contributors to primary production and algal biomass in the Chukchi and Beaufort Seas during May/June 2002

Sea-ice and water samples were collected at 14 stations on the shelves and slope regions of the Chukchi and Beaufort Seas during the spring 2002 expedition as part of the Shelf–Basin Interaction Studies. Algal pigment content, particulate organic carbon and nitrogen, and primary productivity were estimated for both habitats based on ice cores, brine collection and water samples from 5-m depth. The pigment content (0.2–304.3 mg pigments m⁻²) and primary productivity (0.1–23.0 mg C m⁻³ h⁻¹) of the sea-ice algae significantly exceeded water-column parameters (0.2 and 1.0 mg pigments m⁻³; <0.1–0.4 mg C m⁻³ h⁻¹), making sea ice the habitat with the highest food availability for herbivores in early spring in the Chukchi and Beaufort Seas. Stable isotope signatures for ice and water samples did not differ significantly for δ¹⁵N, but for δ¹³C (ice: −25.1‰ to −14.2‰; water: −26.1‰ to −22.4‰). The analysis of nutrient concentrations and the pulse-amplitude-modulated fluorescence signal of ice algae and phytoplankton indicate that nutrients were the prime limiting factor for sea-ice algal productivity. The estimated spring primary production of about 1–2 g C m⁻² of sea-ice algae on the shelves requires the use of substantial nutrient reservoirs from the water column.     

The influence of phytoplankton assemblage composition on biogeochemical characteristics and cycles in the southern Ross Sea,Antarctica

To test the hypothesis that phytoplankton assemblages dominated by different taxa have distinct biogeochemical characteristics and cycles, the temporal and spatial variations in phytoplankton biomass and composition were studied within the Ross Sea polynya, where diatoms and the haptophyte Phaeocystis antarctica are thought to have spatially distinct distributions. Two cruises were completed, with the first conducted in spring, 1994, and the second in late spring–early summer, 1995/1996. Ice concentrations decreased substantially from spring to summer. Mixed layer depths for the region decreased markedly in early spring and were relatively invariant thereafter; the strength of the stratification varied both in time and space. Mixed layers were greater in spring in assemblages dominated by diatoms (as determined by HPLC pigment concentrations) than those dominated by Phaeocystis antarctica, whereas in summer no difference was observed. Nutrient concentrations were initially high and near winter values, but decreased throughout November and December. Nitrate : phosphate removal ratios varied widely, with ratios exceeding 20 in spring but decreasing below 14 in summer. N : P removal ratios at stations dominated by diatoms were less than the Redfield ratio in both spring and summer, and at those stations dominated by P. antarctica the N : P removal ratio was ca. 19 in both seasons. Chlorophyll and particulate matter concentrations increased as nutrients decreased. Spatial and temporal variations of phytoplankton pigments occurred, with 19′-hexanoylfucoxanthin, a pigment of P. antarctica, exceeding 3.9 μg l⁻¹ during spring in the south-central polynya, and fucoxanthin, an accessory pigment of diatoms, found in concentrations >1 μg l⁻¹ in the western Ross Sea. The distributions were not mutually exclusive, and concentrations of both pigments were greatest in spring. The early growth of P. antarctica appears to be related to earlier stratification and disappearance of ice from the south-central Ross Sea. Ratios of FUCO/CHL were relatively invariant, but substantial changes in the HEX/CHL and POC/CHL ratios were observed through time. A one-dimensional nitrogen budget for the spring–early summer period suggests that much of the surface production was partitioned into particles, most (53%) of which remained in the upper 200 m. The rest was partitioned into dissolved organic matter (14%), remineralized as ammonium (19%), or sank from the surface layer as particles (13%). The region may serve as a useful analog to other polar systems, and an understanding of the processes controlling assemblage composition, production, and biomass accumulation may provide insights into biogeochemical cycles of other Antarctic environments.