Impact of a cyclonic eddy on phytoplankton community structure and photosynthetic competency in the subtropical North Pacific Ocean

A synoptic spatial examination of the eddy Haulani (17–20 November 2000) revealed a structure typical of Hawaiian cyclonic eddies with divergent surface flow forcing the upward displacement of deep waters. Hydrographic surveys revealed that surface water in the eddy center was ca. 3.5°C cooler, 0.5 saltier, and 1.4 kg m⁻³ denser than surface waters outside the eddy. Vertically integrated concentrations of nitrate+nitrite, phosphate and silicate were enhanced over out-eddy values by about 2-fold, and nitrate+nitrite concentrations were ca. 8× greater within the euphotic zone inside the eddy than outside. Si:N ratios were lower within the upper mixed layer of the eddy, indicating an enhanced Si uptake relative to nitrate+nitrite. Chlorophyll a concentrations were higher within the eddy compared to control stations outside, when integrated over the upper 150 m, but were not significantly different when integrated over the depth of the euphotic zone. Photosynthetic competency, assessed using fast repetition-rate fluorometry, varied with the doming of the isopycnals and the supply of macro-nutrients to the euphotic zone. The physical and chemical environment of the eddy selected for the accumulation of larger phytoplankton species. Photosynthetic bacteria (Prochlorococcus and Synechococcus) and small (<3 μm diameter) photosynthetic eukaryotes were 3.6-fold more numerically abundant outside the eddy as compared to inside. Large photosynthetic eukaryotes (>3 μm diameter) were more abundant inside the eddy than outside. Diatoms of the genera Rhizosolenia and Hemiaulus outside the eddy contained diazotrophic endosymbiontic cyanobacteria, but these endosymbionts were absent from the cells of these species inside the eddy. The increase in cell numbers of large photosynthetic eukaryotes with hard silica or calcite cell walls is likely to have a profound impact on the proportion of the organic carbon production that is exported to deep water by sinking of senescent cells and cells grazed by herbivorous zooplankton and repackaged as large fecal pellets.     

Organic complexation of iron in the Southern Ocean

The chemical speciation of iron was determined in the Southern Ocean along a transect from 48 to 70°S at 20°E. Dissolved iron concentrations were low at 0.1–0.6 nM, with average concentrations of 0.25±0.13 nM. Organic iron complexing ligands were found to occur in excess of the dissolved iron concentration at 0.72±0.23 nM (equivalent to an excess of 0.5 nM), with a complex stability of log K_FeL′=22.1±0.5 (on the basis of Fe³⁺ and L′). Ligand concentrations were higher in the upper water column (top 200 m) suggesting in situ production by microorganisms, and less at the surface consistent with photochemical breakdown. Our data are consistent with the presence of stable organic iron-complexing ligands in deep global ocean waters at a background level of ∼0.7 nM. It has been suggested that this might help stabilise iron at levels of ∼0.7 nM in deep ocean waters. However, much lower iron concentrations in the waters of the Southern Ocean suggest that these ligands do not prevent the removal of iron (by scavenging or biological uptake) to well below the concentration of these ligands. Scavenging reactions are probably inhibited by such ligand competition, so it is likely that biological uptake is the chief cause for the further removal of iron to these low levels in waters that suffer from very low iron inputs.     

Phytoplankton growth and microzooplankton grazing in the Sea of Okhotsk during late summer of 2006

The Sea of Okhotsk is one of the most productive marine basins in the world ocean and plays an important role in transport of organic carbon and iron to the western subarctic Pacific. We report the first measurements of phytoplankton growth and microzooplankton grazing rates in the Sea of Okhotsk, in late summer of 2006. The study area can be divided into two areas: nutrient-sufficient waters on the continental shelf along the east coast of Sakhalin Island and in the vicinity of Bussol Strait, and surface nutrient-depleted waters beyond the shelf break and in the vicinity of Sakhalin Bay. Phytoplankton growth rate in the studied area was strongly affected by nutrient availability, with high phytoplankton growth rate (0.55±0.14 d^?1) in the nutrient-replete region and severely depressed growth (0.03±0.05 d^?1) in the nutrient-depleted region. On the other hand, microzooplankton grazing rates in both the nutrient-replete and nutrient-depleted regions were approximately the same (0.26±0.20 d^?1 vs. 0.27±0.24 d^?1). Consequently, microzooplankton grazing consumed <50% of the phytoplankton growth in nutrient-rich waters but >3 times the phytoplankton growth in nutrient-depleted waters. Phytoplankton physiological condition as measured by the maximum photochemical quantum efficiency (F_v/F_m) of algal photosystem II (PS II) showed a general trend in agreement with the in situ growth rate of phytoplankton. In contrast to the phytoplankton community, picophytoplankton, especially the cyanobacteria Synechococcus, showed no nutrient effect on their growth, and the growth and mortality rates were well balanced, suggesting that they have a low nutrient requirement and their biomass was controlled principally by microzooplankton grazing.     

Seamounts and organic matter—Is there an effect? The case of Sedlo and Seine seamounts,Part 2. Composition of suspended particulate organic matter

The suspended particulate organic matter (sPOM) around two isolated NE Atlantic seamounts, Seine (33°46′N 14°21′W; summit at ∼170 m) and Sedlo (40°19′N 26°40′W; summit at ∼780 m), was studied over a period of 2 years during four 2–4-week oceanographic surveys. Elemental (C and N), chlorophyll a and lipid biomarker concentrations and N stable isotopic values were variable close to the surface (40–90 m), although some chlorophyll a enrichment above the summits was discerned sporadically. Results from near-surface waters showed a generally “fresh”, mainly phytoplankton signature in sPOM with some seasonality, which was more pronounced around Sedlo. sPOM concentrations and composition changed with depth, apparently controlled by seasonality and proximity to the seamounts. A few metres above the Seine summit, the suspended particulate organic carbon (sPOC) concentrations and labile polyunsaturated fatty acids (% of lipids) were higher than elsewhere at similar depths (∼200 m) in summer 2004. In the same season at Sedlo, polyunsaturated fatty acids were also relatively more abundant (up to 43% of total lipids) around the topographic feature throughout the water column, indicating supply of more labile sPOM, perhaps by advection, downwelling or passive sinking of locally produced phytoplankton and/or in situ production. The high-quality sPOM that seems to be present around the seamounts could provide an important food source to the biological community.     

Diatom biomass and productivity in oceanic and plume-influenced waters of the western tropical Atlantic ocean

Whereas diatoms (class Bacillariophyceae) often dominate phytoplankton taxa in the Amazon estuary and shelf, their contribution to phytoplankton dynamics and impacts on regional biogeochemistry are poorly understood further offshore in the western tropical Atlantic Ocean (WTAO). Thus, relative contribution of diatoms to phytoplankton biomass and primary production rates and associated environmental conditions were quantified during three month-long cruises in January–February 2001, July–August 2001, and April–May 2003. The upper water column was sampled at 6 light depths (100%, 50%, 25%, 10%, 1% and 0.1% of surface irradiance) at 64 stations between 3° and 14°N latitude and 41° and 58°W longitude. Each station was categorized as ‘oceanic’ or ‘plumewater’, based on principal component analysis of eight physical, chemical and biological variables. All stations were within the North Brazil Current, and plumewater stations were characterized by shallower mixed layers with lower surface salinities and higher dissolved silicon (dSi) concentrations than oceanic stations. The major finding was a much greater role of diatoms in phytoplankton biomass and productivity at plumewater stations relative to oceanic stations. Mean depth-integrated bSi concentrations at the plumewater and oceanic stations were 14.2 and 3.7 mmol m⁻², respectively. Mean depth-integrated SiP rates at the plumewater and oceanic stations were 0.17 and 0.02 mmol m⁻² h⁻¹, respectively. Based on ratios of SiP and PP rates, and typical Si:C ratios, diatoms contributed on average 29% of primary productivity at plumewater stations and only 3% of primary productivity at oceanic stations. In contrast, phytoplankton biomass (as chlorophyll a concentrations) and primary production (PP) rates (as ¹⁴C uptake rates) integrated over the euphotic zone were not significantly different at plumewater and oceanic stations. Chlorophyll a concentrations ranged from 8.5 to 42.4 mg m⁻² and 4.0 to 38.0 mg m⁻² and PP rates ranged from 2.2 to 11.2 mmol m⁻² h⁻² and 1.8 to 10.8 mmol m⁻² h⁻² at plumewater and oceanic stations, respectively. A conservative estimate of annual integrated SiP in offshore waters of Amazon plume between April and August is 0.59 Tmol Si, based on mean SiP rates in plumewaters and satellite-derived estimates of the area of the Amazon plume. In conclusion, river plumewaters dramatically alter the silicon dynamics of the WTAO, forming extensive diatom-dominated phytoplankton blooms that may contribute significantly to the global Si budget as well as contributing to energy and matter flow off of the continental shelf.     

Phytoplankton community structure in the Arabian Sea during and after the SW monsoon, 1994

Diatoms, dinoflagellates, coccolithophores, nanoflagellates, picophytoplankton and procaryote algae (Synechococcus spp. and prochlorophytes) were quantified by microscopy and flow cytometry, and their biomass determined, at 12 stations along a 1600 km transect across the Arabian Sea at the end of the SW monsoon in September, and during the inter-monsoon period of November/December 1994. The transect spanned contrasting oceanic conditions that varied from seasonally eutrophic, upwelling waters through mesotrophic, downwelling waters to permanently oligotrophic, stratified waters. The overall diversity of diatoms, dinoflagellates and coccolithophores along the transect was not significantly different between the SW monsoon and inter-monsoon. However, diatoms showed greatest diversity during the SW monsoon and coccolithophores were most diverse during the inter-monsoon. Integrated phytoplankton standing stocks during the SW monsoon ranged from 3 to 9 g C m^-2 in the upwelling eutrophic waters, from 3 to 5 g C m^-2 in downwelling waters, and from 1 to 2 g C m^-2 in oligotrophic waters. Similar phytoplankton standing stocks were found in oligotrophic waters during the inter-monsoon, but were ca. 40% lower compared to the SW monsoon in the more physically dynamic waters. Phytoplankton abundance and biomass was dominated by procaryote taxa. Synechococcus spp. were abundant (often >10⁸ cells l^-1) during both the SW monsoon and inter-monsoon, where the nitrate concentration was ⩾0.1 μ mol l^-1, and often dominated the phytoplankton standing stocks. Prochlorophytes were restricted to oligotrophic stratified waters during the SW monsoon period but were found at all stations along the transect during the inter-monsoon, dominating the phytoplankton standing stocks (>40%) in the oligotrophic region during this period. Of the nano- and micro-phytoplankton, only diatoms contributed significantly to phytoplankton standing stocks, and then only in near-shore upwelling waters during the SW monsoon. There were significant changes in the temporal composition of the phytoplankton community. In nearshore waters a mixed community of diatoms and Synechococcus spp. dominated during the SW monsoon. This gave way to a community dominated by Synechococcus spp. in the intermonsoon. In the downwelling zone, a Synechococcus spp. dominated community was replaced by a mixed procaryote community of Synechococcus spp. and prochlorophytes. In the oligotrophic stratified waters, the mix of procaryote algae was replaced by one dominated by prochlorophytes alone.     

Tropical instability wave interactions within the Galápagos Archipelago

The effects of tropical instability waves (TIW) within the eastern equatorial Pacific during the boreal fall of 2005 were observed in multiple data sets. The TIW cause oscillations of the sea surface temperature (SST), meridional currents (V), and 20 °C isotherm (thermocline). A particularly strong 3-wave packet of ～15-day period TIW passed through the Galápagos Archipelago in Sep and Oct 2005 and their effects were recorded by moored near-surface sensors. Repeat Argo profiles in the archipelago showed that the large temperature (>5 °C) oscillations that occurred were associated with a vertical adjustment within the water column. Numerical simulations report strong oscillations and upwelling magnitudes of ～5.0 m d^?1 near the Tropical Atmosphere Ocean (TAO) buoy at 0°, 95°W and in the Archipelago at 92°W and 90°W. A significant biological response to the TIW passage was observed within the archipelago. Chlorophyll a measured by the Sea-viewing Wide Field-of-view Sensor (SeaWiFS) increased by >30% above 1998–2007 mean concentrations within the central archipelago. The increases coincide with coldest temperatures and the much larger increases within the archipelago as compared to those of 95°W indicate that TIW induced upwelling over the island platform itself brought more iron-enriched upwelling waters into the euphotic zone.     

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.     

Suspended particle organic composition and cycling in surface and midwaters of the equatorial Pacific Ocean

In this study we relate spatial and temporal variation in the organic composition of suspended particles to current conceptual models of open-ocean particle cycling. Suspended particles in surface (0–200 m) and midwaters (200–1000 m) of the equatorial Pacific Ocean were collected during the 1992 US JGOFS Equatorial Pacific (EqPac) program. Samples collected during El Niño (Survey I) and normal conditions (Survey II) were analyzed for pigment, amino acid, fatty acid, and neutral lipid concentrations and compositions. Principal Components Analysis (PCA) and other statistical methods were used to assess changes in particulate organic composition between Surveys I and II, over 24° of latitude, from 15 to 850 m depth, and to compare our compositional data with previously published data from EqPac sinking particles. These analyses indicated that surface suspended particles (0–200 m) were similar in composition to surface ocean phytoplankton and were less degraded than particles sinking out of the euphotic zone (105 m). The organic composition of suspended particles in surface waters varied with latitudinal and El-Niño-induced changes in phytoplankton assemblages. Midwater suspended particles (200–1000 m) contained labile phytodetrital material derived from particles exiting the euphotic zone (105 m). However, labile organic constituents of midwater suspended particles were increasingly degraded by microbes or consumed by midwater metazoans with depth. The increase in degradation state observed for midwater suspended particles may also have been caused by dilution of deeper (450–850 m) suspended particle pools with more refractory material originating from fast-sinking particles, e.g., fecal pellets. However, the mechanism controlling midwater particle degradation state varied with flux regime; dilution of midwater suspended particles dominated only in the higher flux regime found at equatorial latitudes (5°N–5°S) during Survey II (normal conditions). In summary, it is apparent that organic matter alteration in midwaters, and not cycling within the euphotic zone, has the larger effect on organic composition of suspended particles in the deep equatorial Pacific Ocean.     

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.     

Dissolved organic carbon and apparent oxygen utilization in the Atlantic Ocean

Dissolved organic carbon (DOC) distributions along two Atlantic Meridional Transects conducted in 2005 in the region between 47°N and 34°S showed clear latitudinal patterns. The DOC concentrations in the epipelagic zone (0–100 m) were the highest (70–90 µM) in tropical and subtropical waters with stable mixed layers, and lowest (50–55 µM) at the poleward extremities of the transects due to deep convective mixing supplying low DOC waters to the surface. A decrease in DOC occurred with depth, and lowest DOC concentrations (41–45 µM) in the 100–300 m depth range were observed in the equatorial region due to upwelling of low DOC waters. A strong relationship between DOC and AOU was observed in the σ–t 26–26.5 isopycnal layer which underlies the euphotic zone and outcrops at the poleward extremities of the North and South Atlantic Subtropical Gyres (NASG and SASG) in the region ventilating the thermocline waters. Our observations reveal significant north–south variability in the DOC–AOU relationship. The gradient of the relationship suggests that 52% of the AOU in the σ–t 26–26.5 density range was driven by DOC degradation in the NASG and 36% in the SASG, with the remainder due to the remineralisation of sinking particulate material. We assess possible causes for the greater contribution of DOC remineralisation in the NASG compared to the SASG.     

Atmospheric iron deposition and sea-surface dissolved iron concentrations in the eastern Atlantic Ocean

Atmospheric iron and underway sea-surface dissolved (<0.2 μm) iron (DFe) concentrations were investigated along a north–south transect in the eastern Atlantic Ocean (27°N/16°W–19°S/5°E). Fe concentrations in aerosols and dry deposition fluxes of soluble Fe were at least two orders of magnitude higher in the Saharan dust plume than at the equator or at the extreme south of the transect. A weaker source of atmospheric Fe was also observed in the South Atlantic, possibly originating in southern Africa via the north-easterly outflow of the Angolan plume. Estimations of total atmospheric deposition fluxes (dry plus wet) of soluble Fe suggested that wet deposition dominated in the intertropical convergence zone, due to the very high amount of precipitation and to the fact that a substantial part of Fe was delivered in dissolved form. On the other hand, dry deposition dominated in the other regions of the transect (73–97%), where rainfall rates were much lower. Underway sea-surface DFe concentrations ranged 0.02–1.1 nM. Such low values (0.02 nM) are reported for the first time in the Atlantic Ocean and may be (co)-limiting for primary production. A significant correlation (Spearman's rho=0.862, p<0.01) was observed between mean DFe concentrations and total atmospheric deposition fluxes, confirming the importance of atmospheric deposition on the iron cycle in the Atlantic. Residence time of DFe in the surface waters relative to atmospheric inputs were estimated in the northern part of our study area (17±8 to 28±16 d). These values confirmed the rapid removal of Fe from the surface waters, possibly by colloidal aggregation.     

Nutrient gradients in the western North Atlantic Ocean: Relationship to microbial community structure and comparison to patterns in the Pacific Ocean

Studies of nitrogen and phosphorus dynamics in the oligotrophic surface waters of the western North Atlantic Ocean have been constrained because ambient concentrations are typically at or below the detection limits of standard colorometric methods, except during periods of deep vertical mixing. Here we report the application of high-sensitivity analytical methods—determinations of nitrate plus nitrite (N+N) by chemiluminescence and soluble reactive phosphorus (SRP) by the magnesium induced co-precipitation (MAGIC) protocol—to surface waters along a transect from the Sargasso Sea at 26°N through the Gulf Stream at 37°N, including sampling at the JGOFS Bermuda Atlantic Time-series Study (BATS) station. The results were compared with data from the BATS program, and the HOT station in the Pacific Ocean, permitting cross-ecosystem comparisons. Microbial populations were analyzed along the transect, and an attempt was made to interpret their distributions in the context of the measured nutrient concentrations.Surface concentrations of N+N and SRP during the March 1998 transect separated into 3 distinct regions, with the boundaries corresponding roughly to the locations of the BATS station (∼31°N) and the Gulf Stream (∼37°N). Although N+N and SRP co-varied, the [N+N] : [SRP] molar ratios increased systematically from ∼1 to 10 in the southern segment, remained relatively constant at ∼40–50 between 31°N and 37°N, then decreased again systematically to ratios <10 north of the Gulf Stream. Dissolved organic N (DON) and P (DOP) dominated (⩾90%) the total dissolved N (TDN) and P (TDP) pools except in the northern portion of the transect. The [DON] : [DOP] molar ratios were relatively invariant (∼30–60) across the entire transect.Heterotrophic prokaryotes (operationally defined as “bacteria”), Prochlorococcus, Synechococcus, ultra- and nanophytoplankton, cryptophytes, and coccolithophores were enumerated by flow cytometry. The abundance of bacteria was well correlated with the concentration of SRP, and that of the ultra- and nanophytoplankton was well correlated with the concentration of N+N. The only group whose concentration was correlated with temperature was Prochlorococcus, and its abundance was unrelated to the concentrations of nutrients measured at the surface.We combined our transect results with time-series measurements from the BATS site and data from select depth profiles, and contrasted these North Atlantic data sets with time-series of N and P nutrient measurements from a station in the North Pacific subtropical gyre near Hawaii [Hawaii Ocean Time-series (HOT) site]. Two prominent differences are readily observed from this comparison. The [N+N] : [SRP] molar ratios are much less than 16 : 1 during stratified periods in surface waters at the BATS site, as is the case at the HOT site year round. However, following deep winter mixing, this ratio is much higher than 16 : 1 at BATS. Also, SRP concentrations in the upper 100 m at BATS fall in the range 1–10 nM during stratified periods, which is at least one order of magnitude lower than at the HOT site. That two ecosystems with comparable rates of primary and export production would differ so dramatically in their nutrient dynamics is intriguing, and highlights the need for detailed cross ecosystem comparisons.     

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

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

Thermocline ventilation and oxygen utilization rates in the subtropical North Pacific based on CFC distributions during WOCE

Thermocline ventilation rates for the subtropical North Pacific are determined using a 1-dimensional (meridional) along-isopycnal advective–diffusive model tuned to chlorofluorocarbon (CFC) concentrations measured along 152°W in 1991 during WOCE P16. Mean southward advection rates in the subtropics range from 1.03 to 0.56 cm s^-1 between σ_θ=25.5 and 26.6. Model-derived ventilation times for the subtropical gyre increase from about 10 to 27 years for that isopycnal range. Oxygen utilization rates (OURs) determined using the advective-diffusive model decrease with depth from 6.6 to 3.2 μmol kg^-1 yr^-1 between σ_θ=25.5 and 26.6. Extrapolation of the OUR versus depth trend to the base of the euphotic zone with the 1/Z power function of Martin et al. (1987) and integration from 500 to 100 m depth implies a carbon export rate from the overlying euphotic zone of 2.2±0.5 moles C m^-2 yr^-1 at 30°N, 152°W. Analysis of the WOCE radiocarbon and salinity distributions indicates that zonal and cross-isopycnal transport terms would have to be considered in modeling these tracers in the subtropical North Pacific.     

Monsoonal influence on the distribution of phytoplankton pigments in the Arabian Sea

Variations in the distribution of chemotaxonomic pigments were monitored in the Arabian Sea and the Gulf of Oman at the end of the SW monsoon in September 1994 and during the inter-monsoon period in November/December 1994 to determine the seasonal changes in phytoplankton composition. The Gulf of Oman was characterized by sub-surface chlorophyll maxima at 20-40 m during both seasons, and low levels of divinyl chlorophyll a indicated that prochlorophytes did not contribute significantly to the total chlorophyll a. Prymnesiophytes (19′-hexanoyloxyfucoxanthin), diatoms (fucoxanthin) and chlorophyll b containing organisms accounted for most of the phytoplankton biomass in September, while prymnesiophytes dominated in November/December. In the Arabian Sea in September, high total chlorophyll a concentrations up to 1742 ng l^-1 were measured in the coastal upwelling region and a progressive decline was monitored along the 1670 km offshore transect to oligotrophic waters at 8°N. Divinyl chlorophyll a was not detected along this transect except at the two most southerly stations where prochlorophytes were estimated to contribute 25–30% to the total chlorophyll a. Inshore, the dominance of fucoxanthin and/or hexanoyloxyfucoxanthin indicated that diatoms and prymnesiophytes generally dominated the patchy phytoplankton community, with zeaxanthin-containing Synechococcus also being important, especially in surface waters. At the southern oligotrophic localities, Synechococcus and prochlorophytes dominated the upper 40 m and prymnesiophytes were the most prominent at the deep chlorophyll maximum. During the inter-monsoon season, total chlorophyll a concentrations were generally half those measured in September and highest levels were found on the shelf (1170 ng l^-1). Divinyl chlorophyll a was detected at all stations along the Arabian Sea transect, and we estimated that prochlorophytes contributed between 3 and 28% to the total chlorophyll a, while at the two oligotrophic stations this proportion increased to 51–52%. While procaryotes were more important in November/December than September, eucaryotes still accounted for >50% of the total chlorophyll a. Pigment/total chlorophyll a ratios indicated that 19′-hexanoyloxyfucoxanthin-containing prymnesiophytes were the dominant group, although procaryotes accounted for 65% at the two southerly oligotrophic stations.     

Responses of pico- and nanophytoplankton to artificial iron infusions observed during the second iron enrichment experiment in the western subarctic Pacific (SEEDS II)

How pico- and nanophytoplankton responded to artificial iron infusions was investigated using flow cytometry during SEEDS II, the second mesoscale in situ iron enrichment experiment in the western subarctic North Pacific. Two iron infusions on days 0 and 7 caused a remarkable increase in cellular chlorophyll fluorescence and cell size of all the four phytoplankton groups investigated: Synechococcus, cryptophytes, picoeucaryotes and nanoeucaryotes other than cryptophytes. After the second infusion, the abundance of three phytoplankton groups, excluding Synechococcus, also started to increase. After surface dissolved iron concentration decreased to <0.2 nM on day 11, chlorophyll fluorescence of all the four groups returned to a level observed before the iron infusions, suggesting that pico- and nanophytoplankton were physiologically stressed by iron deficiency. Cell concentrations of pico- and nanoeucaryotes decreased to the pre-infusion level by day 23, while that of cryptophytes remained high until day 25. Flow cytometric diagnosis showed that cryptophytes were physiologically limited during this period and effective iron uptake from suspended particles, as reported for freshwater cryptophytes, was not observed. Thus their prosperity may have been due to alleviation from grazing. Cell concentration of Synechococcus started to increase at a net specific growth rate of 0.13 d⁻¹ after day 12, reaching more than 6 times that of the pre-infusion level on day 24. This may have been due to the elevation of surface water temperature observed during the survey period, together with trophic cascading effects of increased copepod grazing.     

Variability in ultraplankton at the Porcupine Abyssal Plain study site

Observations of the ultraplankton (<5 μm) are presented from a 4 day mesoscale survey centred on the Porcupine Abyssal Plain (PAP) study site (49°00’N 16°30’W), in July 2006. The organisms enumerated include two groups of phytoplankton, Synechococcus cyanobacteria, heterotrophic bacteria, large viruses, and two size classes of heterotrophic protist. The dataset comprises over 400 samples from the mixed layer taken over a 100 × 100 km² area at a spatial resolution of typically 2-3 km.For phytoplankton and heterotrophic bacteria there is a clear bimodal structure to the histograms of abundance indicative of two distinct communities in the region. Using the strong bimodality of one of the phytoplankton groups’ histogram as a basis, the dataset is split into two subsets, with roughly 200 points in each, corresponding to the two histogram peaks. Doing so provides evidence that Synechococcus and viruses may also have a bimodal structure. Correlations between all pairings of these five organisms (both phytoplankton groups, Synechococcus, heterotrophic bacteria and viruses) are positive and quite high (r>0.7). The two communities can therefore be characterised as high and low abundance. Although there is a coincidence of low abundances with high temperatures in the southwest corner of the region, where there was known to be an eddy present, the spatial distributions of these organisms over the whole region is poorly predicted by temperature (or salinity or density). Furthermore, the spatial distributions of heterotrophic protists are found to differ strongly from those of the other organisms, having a unimodal structure and no obvious large scale structure. The more random structure of the heterotrophs’ spatial distribution compared to their prey is consistent with previous results from the continental shelf, but is demonstrated for the open ocean here for the first time.Spatial variability is a large potential source of error in point samples, such as those comprising time series or transect cruises, unless a sufficient number of samples are taken. This large dataset is further used to provide guidance on the number of samples that would be required to estimate the mean abundance for the organisms accurately in this spatially variable region. Even if the bimodal structure was known initially, many of the organisms would require 10 or more samples to estimate the mean with 25% accuracy.     

Speciation of Fe in the Eastern North Atlantic Ocean

In the Eastern North Atlantic Ocean iron (Fe) speciation was investigated in three size fractions: the dissolvable from unfiltered samples, the dissolved fraction (<0.2 μm) and the fraction smaller than 1000 kDa (<1000 kDa). Fe concentrations were measured by flow injection analysis and the organic Fe complexation by voltammetry. In the research area the water column consisted of North Atlantic Central Water (NACW), below which Mediterranean Overflow Water (MOW) was found with the core between 800 and 1000 m depth. Below 2000 m depth the North Atlantic Deep Water (NADW) proper was recognised. Dissolved Fe and Fe in the <1000 kDa fraction showed a nutrient like profile, depleted at the surface, increasing until 500–1000 m depth below which the concentration remained constant. Fe in unfiltered samples clearly showed the MOW with high concentrations (4 nM) compared to the overlying NACW and the underlying NADW, with 0.9 nM and 2 nM Fe, respectively. By using excess ligand (Excess L) concentrations as parameter we show a potential to bind Fe. The surface mixed layer had the highest excess ligand concentrations in all size fractions due to phytoplankton uptake and possible ligand production. The ratio of Excess L over Fe proved to be a complementary tool in revealing the relative saturation state of the ligands with Fe. In the whole water column, the organic ligands in the larger colloidal fraction (between 0.2 μm and 1000 kDa) were saturated with Fe, whereas those in the smallest fraction (<1000 kDa) were not saturated with Fe, confirming that this fraction was the most reactive one and regulates dissolution and colloid aggregation and scavenging processes. This regulation was remarkably stable with depth since the alpha factor (product of Excess L and K′), expressing the reactivity of the ligands, did not vary and was 10¹³. Whereas, in the NACW and the MOW, the ligands in the particulate (>0.2 μm) fraction were unsaturated with Fe with respect to the dissolved fraction, thus these waters had a scavenging potential.     

Dissolved iron(II) in the Pacific Ocean: Measurements from the PO2 and P16N CLIVAR/CO2 repeat hydrography expeditions

The redox speciation of dissolved iron in seawater was evaluated at 121 locations in the Pacific Ocean at depths of 15-1000 m, using the method of luminol chemiluminescence. The results indicate that reduced iron, Fe(II), is ubiquitous in surface seawater with a relatively consistent pattern of occurrence. Surface maxima were present in most profiles, with median concentrations of 25-30 pM representing 12-14% of the total dissolved iron. Concentrations decreased monotonically with depth to<12 pM within the upper euphotic zone. This pattern was observed during both day and nighttime sampling events, which suggests that non-photochemical production mechanisms can produce photochemical-like signatures. Further, if theoretical rates of Fe(II) oxidation are applicable to the open ocean, then the employed sampling methods precluded assessment of photochemically-produced Fe(II), regardless of ambient light conditions. For this and other reasons, the concentrations reported here for the upper water column likely represent lower limits of labile iron concentration, and suggest that dissolved iron may be more available for uptake than previously believed. Deeper in the water column, Fe(II) was also frequently detected, though it constituted a small fraction of the total dissolved iron. Possible source mechanisms at these depths include thermal (dark) reduction of Fe(III) organic complexes or remineralization of sinking biogenic particles containing Fe(II). In the northern Philippine Sea between the Japanese coast and the Izu-Bonin volcanic arc system, Fe(II) concentrations were found to be atypically high, possibly because of high atmospheric dust deposition near the surface and transport of sediment-derived iron at depth.