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.     

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.     

Physical behaviors of the iron-fertilized patch in SEEDS II

Sulfur hexafluoride (SF₆) tracer release experiments were carried out to trace the iron-fertilized water mass during the iron-fertilization experiments in the western North Pacific of Subarctic Pacific Iron Experiment for Ecosystem Dynamics Study II (SEEDS II) in 2004. A solution of Fe and SF₆ tracer was released into the surface mixed layer over an 8×8 km area, and the fertilized patch was traced by onboard SF₆ analysis for 12 days during each experiment. A Lagrangian frame of reference was maintained by the use of a drogued GPS buoy released at the center of the patch to reduce the advection effect on observations. The patch moved along the contour of sea-surface height (SSH) of a clockwise mesoscale eddy for 4 days after release. Then strong easterly winds dragged the patch across the contour of SSH. The patch behavior was affected by both the mesoscale eddy and surface winds. Apparent horizontal diffusivities were determined by the change of the distribution of SF₆ concentrations. The averaged apparent horizontal diffusivity was about 49 m² s⁻¹ during SEEDS II. It was larger than the one in SEEDS. Mixed-layer depth (MLD) was 8.5–18 m during SEEDS, and 12–33 m during SEEDS II. The larger horizontal diffusivity and deeper MLD in SEEDS II were disadvantages to maintain a high iron concentration in the surface layer compared to SEEDS. Temporal change of the MLD corresponded to the temporal change of chlorophyll-a concentration. Temporal change in the surface MLD was also important for the response of phytoplankton by iron fertilization.     

Community structure and photosynthetic physiology of phytoplankton in the northwest subarctic Pacific during an in situ iron fertilization experiment (SEEDS-II)

Temporal changes in the abundance, community composition, and photosynthetic physiology of phytoplankton in surface waters were investigated during the second in situ iron (Fe) fertilization experiment in the NW subarctic Pacific (SEEDS-II). Surface chlorophyll a concentration was 0.75 mg m⁻³ on the day before the first Fe enrichment (i.e. Day 0), increased ca. 3-fold until Day 13 after two Fe additions, and thereafter declined with time. The photochemical quantum efficiency (F_v/F_m) and functional absorption cross-section (σPSII) of photosystem II for total phytoplankton in surface waters increased and decreased inside the Fe-enriched patch through Day 13, respectively. These results indicate that the photosynthetic physiological condition of the phytoplankton improved after the Fe infusions. However, the maximum F_v/F_m value of 0.43 and the maximum quantum yield of carbon fixation (φ_max) of 0.041 mol C (mol photon)⁻¹ during the development phase of the bloom were rather low, compared to their theoretical maximum of ca. 0.65 and 0.10 mol C (mol photon)⁻¹, respectively. Diatoms, which were mainly composed of oceanic species, did not bloom, and autotrophic nanoflagellates such as cryptophytes and prasinophytes became predominant in the phytoplankton community inside the Fe-enriched patch. In ferredoxin/flavodoxin assays for micro-sized (20–200 μm in cell length) diatoms, ferredoxin was not detected but flavodoxin expressions consistently occurred with similar levels both inside and outside the Fe-enriched patch, indicating that the large-sized diatoms were stressed by Fe bioavailability inside the Fe-enriched patch even after the Fe enrichments. Our data suggest that the absence of a Fe-induced large-sized diatom bloom could be partly due to their Fe stress throughout SEEDS-II.     

Behavior of particulate materials during iron fertilization experiments in the Western Subarctic Pacific (SEEDS and SEEDS II)

During two mesoscale iron-enrichment studies in the northwestern subarctic Pacific (SEEDS in 2001 summer and SEEDS II in 2004 summer), particulate materials from the iron-induced phytoplankton bloom in the upper water column were monitored to analyze the export processes beneath the upper mixed layer, mainly with drifting sediment traps. We could not observe the total downward export process of the high accumulation of particulate organic carbon from the mixed layer induced by the large diatom bloom of SEEDS [e.g., Tsuda, A., Takeda, S., Saito, H., Nishioka, J., Nojiri, Y., Kudo, I., Kiyosawa, H., Shiomoto, A., Imai, K., Ono, T., Shimamoto, A., Tsumune, D., Yoshimura, T., Aono, T., Hinuma, A., Kinugasa, M., Suzuki, K., Sohrin, Y., Noiri, Y., Tani, H., Deguchi, Y., Tsurushima, N., Ogawa, H., Fukami, K., Kuma, K., Saino, T., 2003. A mesoscale iron enrichment in the western subarctic Pacific induces large centric diatom bloom. Science 300, 958–961] because the 2-week observation period was too short to examine the decline phase of the bloom. In contrast, in SEEDS II, the particulate organic carbon and particulate organic nitrogen were accumulated 123 and 23 mmol m⁻², respectively, in the mixed layer until day-15 (days from iron-enrichment), and then ca. 90% were removed from the mixed layer by day-25. The sediment traps at 40 m depth between day-15 and day-25 accounted for at least more than 35% of these particles. There was no large variation in chemical composition in settling particles above 100 m depth throughout the experimental periods both in SEEDS and SEEDS II. The content of biogenic opal remained more than 50% of all settling particles during SEEDS, while the content of biogenic calcium carbonate was relatively high, with a low biogenic opal content of consistently less than 30% during SEEDS II. These results suggest that high standing stock of seed population of diatoms before the iron fertilization, indicated by low C/Si ratio of particulate matter, is an important factor to induce the large diatom bloom in SEEDS.     

Behaviors of dissolved and particulate Co,Ni, Cu,Zn, Cd and Pb during a mesoscale Fe-enrichment experiment (SEEDS II) in the western North Pacific

During mesoscale Fe enrichment (SEEDS II) in the western North Pacific ocean, we investigated dissolved and particulate Co, Ni, Cu, Zn, Cd and Pb in seawater from both field observation and shipboard bottle incubation of a natural phytoplankton assemblage with Fe addition. Before the Fe enrichment, strong correlations between dissolved trace metals (Ni, Zn and Cd) and PO₄³⁻, and between particulate trace metals (Ni, Zn and Cd) and chlorophyll-a were obtained, suggesting that biogeochemical cycles mainly control the distributions of Ni, Zn and Cd in the study area. Average concentrations of dissolved Co, Ni, Cu, Zn, Cd and Pb in the surface mixed layer (0–20 m) were 70 pM, 4.9, 2.1, 1.6, 0.48 nM and 52 pM, respectively, and those for the particulate species were 1.7 pM, 0.052, 0.094, 0.46, 0.037 nM and 5.2 pM, respectively. After Fe enrichment, chlorophyll-a increased 3 fold (up to 3 μg L⁻¹) during developing phases of the bloom (<12 days). Mesozooplankton biomass also increased. Particulate Co, Ni, Cu and Cd inside the patch hinted at an increase in the concentrations, but there were no analytically significant differences between concentrations inside and outside the patch. The bottle incubation with Fe addition (1 nM) showed an increase in chlorophyll-a (8.9 μg L⁻¹) and raised the particulate fraction up to 3–45% for all the metals, accompanying changes in Si/P, Zn/P and Cd/P. These results suggest that Fe addition lead to changes in biogeochemical cycling of trace metals. The comparison between the mesoscale Fe enrichment and the bottle incubation experiment suggests that although Fe was a limiting factor for the growth of phytoplankton, the enhanced biomass of mesozooplankton also limited the growth of phytoplankton and the transformation of trace metal speciation during the mesoscale Fe enrichment. Sediment trap data and the elemental ratios taken up by phytoplankton suggest that export loss was another reason that no detectable change in the concentrations of particulate trace metals was observed during the mesoscale Fe enrichment.     

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.     

Standing stocks,production, and respiration of phytoplankton and heterotrophic bacteria in the western Arctic Ocean

Standing stocks and production rates for phytoplankton and heterotrophic bacteria were examined during four expeditions in the western Arctic Ocean (Chukchi Sea and Canada Basin) in the spring and summer of 2002 and 2004. Rates of primary production (PP) and bacterial production (BP) were higher in the summer than in spring and in shelf waters than in the basin. Most surprisingly, PP was 3-fold higher in 2004 than in 2002; ice-corrected rates were 1581 and 458 mg C m⁻² d⁻¹, respectively, for the entire region. The difference between years was mainly due to low ice coverage in the summer of 2004. The spatial and temporal variation in PP led to comparable variation in BP. Although temperature explained as much variability in BP as did PP or phytoplankton biomass, there was no relationship between temperature and bacterial growth rates above about 0 °C. The average ratio of BP to PP was 0.06 and 0.79 when ice-corrected PP rates were greater than and less than 100 mg C m⁻² d⁻¹, respectively; the overall average was 0.34. Bacteria accounted for a highly variable fraction of total respiration, from 3% to over 60% with a mean of 25%. Likewise, the fraction of PP consumed by bacterial respiration, when calculated from growth efficiency (average of 6.9%) and BP estimates, varied greatly over time and space (7% to >500%). The apparent uncoupling between respiration and PP has several implications for carbon export and storage in the western Arctic Ocean.     

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.     

Kinetic study reveals weak Fe-binding ligand,which affects the solubility of Fe in the Scheldt estuary

The chemistry of dissolved Fe(III) was studied in the Scheldt estuary (The Netherlands). Two discrete size fractions of the dissolved bulk (< 0.2 μm and < 1 kDa) were considered at three salinities (S = 26, 10 and 0.3).Within the upper estuary, where fresh river water meets seawater, the dissolved Fe concentration decreases steeply with increasing salinity, for the fraction < 0.2 μm from 536 nM at S = 0.3 to 104 nM at S = 10 and for the fraction < 1 kDa from 102 nM to 36 nM Fe. Further downstream, in the middle and lower estuary, this decrease in the Fe concentration continues, but is far less pronounced. For all samples, the traditionally recognised dissolved strong organic Fe-binding ligand concentrations are lower than the dissolved Fe concentrations.Characteristics of dissolved Fe-binding ligands were determined by observing kinetic interactions with adsorptive cathodic stripping voltammetry. From these kinetic experiments we concluded that apart from the well-known strong Fe-binding organic ligands (L, logK′ = 19–22) also weak Fe-binding ligands (P) existed with an α value (binding potential = K′ · [P]) varying between 10^11.1 and 10^11.9. The presence of this relatively weak ligand explained the high concentrations of labile Fe present in both size fractions in the estuary. This weak ligand can retard or prevent a direct precipitation after an extra input of Fe.The dissociation rate constants of the weak ligand varied between 0.5 × 10^− 4 and 4.3 × 10^− 4 s^− 1. The rate constants of the strong organic ligand varied between k_d = 1.5 × 10^− 3–17 × 10^− 2 s^− 1 and k_f = 2.2 × 10⁸–2.7 × 10⁹ M^− 1 s^− 1. The dissociation rate constant of freshly amorphous Fe-hydroxide was found to be between 4.3 × 10^− 4 and 3.7 × 10^− 3 s^− 1, more labile or equal to the values found by Rose and Waite [Rose, A.L., Waite, T.D., 2003a. Kinetics of hydrolysis and precipitation of ferric iron in seawater. Environ. Sci. Technol., 37, 3897–3903.] for freshly precipitated Fe in seawater.Kinetic rate constants of Fe with the ligand TAC (2-(2-Thiazolylazo)-p-cresol) were also determined. The formation rate constant of Fe(TAC)₂ varied between 0.1 × 10⁸ and 3.6 × 10⁸ M^− 1 s^− 1, the dissociation rate constant between 0.2 × 10^− 5 and 17 × 10^− 5 s^− 1 for both S = 26 and S = 10. The conditional stability constant of Fe(TAC)₂ (β_Fe(TAC)2′) varied between 22 and 23.4 for S = 10 and S = 26 more or less equal to that known from the literature (logβ_Fe(TAC)2′ = 22.4; [Croot, P.L., Johansson, M., 2000. Determination of iron speciation by cathodic stripping voltammetry in seawater using the competing ligand 2-(2-Thiazolylazo)-p-cresol (TAC). Electroanalysis, 12, 565–576.]). However, at S = 0.3 the logβ_Fe(TAC)2′ was 25.3, three orders of magnitude higher. Apparently the application of TAC to samples of low salinity can only be done when the correct β_Fe(TAC)2′ is known.     

Primary productivity,differential size fraction and pigment composition responses in two Southern Ocean in situ iron enrichments

Two in situ iron-enrichment experiments were conducted in the Pacific sector of the Southern Ocean during summer 2002 (SOFeX). The “north patch,” established within the Subantarctic Zone (∼56°S), was characterized by high nitrate (∼21 mmol m⁻³) but low silicic acid (2 mmol m⁻³) concentrations. North patch iron enrichment increased chlorophyll (Chl) by 12-fold to 2.1 mg m⁻³ and primary productivity (PP_EU) by 8-fold to 188 mmol C m⁻² d⁻¹. Surprisingly, despite low silicic acid concentrations, diagnostic pigment and size-fraction composition changes indicated an assemblage shift from prymnesiophytes toward diatoms. The “south patch,” poleward of the Southern Boundary of the Antarctic Circumpolar Current (SBACC) (∼66°S), had high concentrations of nitrate (∼27 mmol m⁻³) and silicic acid (64 mmol m⁻³). South patch iron enrichment increased Chl by 9-fold to 3.8 mg m⁻³ and PP_EU 5-fold to 161 mmol C m⁻² d⁻¹ but, notably, did not alter the phytoplankton assemblage from the initial composition of ∼50% diatoms. South patch iron addition also reduced total particulate organic carbon:Chl from ∼300 to 100; enhanced the presence of novel non-photosynthetic, but fluorescent, compounds; and counteracted a decrease in photosynthetic performance as photoperiod decreased. These experiments show unambiguously that in the contemporary, high nitrate Southern Ocean increasing iron supply increases primary productivity, confirming the initial premise of the Martin Iron Hypothesis. However, despite a 5-fold increase in PP_EU under iron-replete conditions in late summer, the effect of iron on annual productivity in the Southern Ocean poleward of the SBACC is limited by seasonal ice coverage and the dark of polar winter.     

Environmental controls on phytoplankton community composition in the Thames plume,U.K.

The aim of this study was to investigate controls on the phytoplankton community composition and biogeochemistry of the estuarine plume zone of the River Thames, U.K. using an instrumented moored buoy for in situ measurements and preserved sample collection, and laboratory-based measurements from samples collected at the same site. Instrumentation on the moored buoy enabled high frequency measurements of a suite of environmental variables including in situ chlorophyll, water-column integrated irradiance, macronutrients throughout an annual cycle for 2001 e.g. nitrate and silicate, and phytoplankton biomass and species composition. The Thames plume region acts as a conduit for fluvial nutrients into the wider southern North Sea with typical winter concentrations of 45 μM nitrate, 17 μM silicate and 2 μM phosphate measured. The spring bloom resulted from water-column integrated irradiance increasing above 60 W h m^− 2 d^− 1 and was initially dominated by a diatom bloom mainly composed of Nitzschia sp. and Odontella sinesis. The spring bloom then switched after ∼ 30 days to become dominated by the flagellate Phaeocystis reaching a maximum chlorophyll concentration of 37.8 μg L^− 1. During the spring bloom there were high numbers of the heterotrophic dinoflagellates Gyrodinium spirale and Katodinium glaucum that potentially grazed the phytoplankton bloom. This diatom–flagellate switch was predicted to be due to a combination of further increasing water-column integrated irradiance > 100 W h m^− 2 d^− 1 and/or silicate reaching potentially limiting concentrations (< 1 μM). Post spring bloom, diatom dominance of the lower continuous summer phytoplankton biomass occurred despite the low silicate concentrations (Av. 0.7 μM from June–August). Summer diatom dominance, generally due to Guinardia delicatula, was expected to be as a result of microzooplankton grazing, dominated by the heterotrophic dinoflagellate Noctiluca scintillans, controlling 0.7–5.0 μm ‘flagellate’ fraction of the phytoplankton community with grazing rates up to 178% of ‘flagellate’ growth rate. The Thames plume region was therefore shown to be an active region of nutrient and phytoplankton processing and transport to the southern North Sea. The use of a combination of moorings and ship-based sampling was essential in understanding the factors influencing nutrient transport, phytoplankton biomass and species composition in this shelf sea plume region.     

Early spring bloom phytoplankton-nutrient dynamics at the Celtic Sea Shelf Edge

Phytoplankton production was measured at the shelf edge region of the Celtic Sea in April/May 1994 at the beginning of the spring bloom. Size fractionated ¹⁴C uptake experiments showed that phytoplankton >2 μm dominated the bloom although, in the period immediately before the increase in phytoplankton biomass, picophytoplankton (<2 μm) was responsible for up to 42% of the production; in these late winter conditions, chlorophyll concentrations were generally <0.7 μg l^-1 and primary production was ca. 70 mmol C m^-2 d^-1. As the spring bloom developed, phytoplankton production rates of 120 mmol C m^-2 d^-1 were measured. Chlorophyll concentration increased to >2 μg l^-1 as a result of growth of larger phytoplankton, including diatoms, with large numbers of Nitzschia, Thalassionema and Chaetoceros dominating the assemblage. Picophytoplankton production declined as the spring bloom progressed. Nutrient concentrations were not depleted during the sampling period, and NO^-₃ concentrations were >6 μmol l^-1. Nutrient assimilation rates were measured at the same time as primary production was estimated. Before the development of any substantial phytoplankton biomass, the uptake rates for ammonium and nitrate were very similar, with f-ratios ranging from 0.5 to 0.6. Assimilation of ammonium remained relatively constant after the onset of stratification and bloom development, but nitrate uptake increased by a factor of 2 or more, resulting in f-ratios >0.8. There was significant phosphate uptake in the dark, which was generally ca. 50% of the rate in the light. The C : N : P assimilation ratios changed as the bloom developed; in the pre-bloom situation, when small phytoplankton cells dominated the assemblage, the C : N assimilation ratio was variable, with some stations having ratios less than (ca 2.5), and some higher than (ca. 9), the Redfield ratio. The most actively growing assemblages had N : P ratios close to the Redfield ratio, but the C : N ratios were consistently lower. New production was found to be closely correlated with the size of the species making up the phytoplankton assemblage, and high f ratios were measured when larger phytoplankton dominated the assemblage.     

Meso- and microzooplankton responses to an in situ iron fertilization experiment (SEEDS II) in the northwest subarctic Pacific

A mesoscale iron fertilization experiment was carried out in the western subarctic Pacific during summer 2004. The iron-patch was traced for 26 days after the enrichment, and the abundance and behavior of meso- and microzooplankton was compared with those outside of the patch. The surface chlorophyll-a concentration in the patch was high between days 10 and 13 (2.5 mg m⁻³) and decreased to the initial level after day 20. Microzooplankton grazing rates, estimated by a dilution method, was mostly balanced with phytoplankton growth rates throughout the observed period. Dominant mesozooplankton species in the upper 200 m were copepods: dominated by Eucalanus bungii, Neocalanus plumchrus and Metridia pacifica. Species composition did not change in the patch over the observation period. The copepod biomass was 3–5 times higher than in Subarctic Pacific Iron Experiment for Ecosystem Dynamics Study (SEEDS), the previous iron-enrichment experiment in the same area, before the bloom, and exponentially increased both inside and outside the patch, which was mainly brought by the development of N. plumchrus. The development rates of N. plumchrus were not significantly different between inside and outside the patch. Estimated grazing rate suggest that the copepod grazing was main cause of the low accumulation of phytoplankton biomass, and dominance of grazing-resistant organisms such as large ciliates, large diatoms and diatoms with extremely long setae. “Arrested migration” for M. pacifica and upward shift of vertical distribution by E. bungii were observed during the bloom period, even if the accumulation of phytoplankton biomass was very low compared to other iron-enrichment experiments. These results indicate that the copepod grazing shaped the food-web structure of the lower trophic levels (biomass and species composition) in SEEDS II.     

High-frequency fluxes of labile compounds in the central Ligurian Sea,northwestern Mediterranean

Sinking particles were collected every 4 h with drifting sediment traps deployed at 200 m depth in May 1995 in a 1-D vertical system during the DYNAPROC observations in the northwestern Mediterranean sea. POC, proteins, glucosamine and lipid classes were used as indicators of the intensity and quality of the particle flux. The roles of day/night cycle and wind on the particle flux were examined. The transient regime of production from late spring bloom to pre-oligotrophy determined the flux intensity and quality. POC fluxes decreased from, on average, 34 to 11 mg m⁻² d⁻¹, representing 6–14% of the primary production under late spring bloom conditions to 1–2% under pre-oligotrophic conditions. Total protein and chloroplast lipid fluxes correlated with POC and reflected the input of algal biomass into the traps. As the season proceeded, changes in the biochemical composition of the exported material were observed. The C/N ratio rose from 7.8 to 12. Increases of serine (10–28% of total proteins), total lipids (7–9 to 14–28% of POC) and reserve lipids (1–5 to 5–22% of total lipids) were noticeable, whereas total protein content in POC decreased (20–27 to 18–7%). N-acetyl glucosamine, a tracer of fecal pellet flux, showed that zooplankton grazing was a major vector of downward export during the decaying bloom. Against this background pattern, episodic events specifically increased the flux, modifying the quality and the settling velocity of particles. Day/night signals in biotracers (POC, N-acetyl glucosamine, protein and chloroplast lipids) showed that zooplankton migrations were responsible for sedimentation of fresh material through fast sinking particles (V=170–180 m d⁻¹) at night. Periodic signatures of re-processed material (high lipolysis and bacterial biomass indices) suggested that other zooplankton fecal pellets or small aggregates, probably of lower settling velocities (V<170 m d⁻¹), contributed to the flux during calm periods. At the beginning of the experiment, during the development of a prymnesiophyte bloom in the upper layers, the sterol signal with no periodicity enabled us to estimate high particle settling velocities (⩾600 m d⁻¹) likely related to large aggregate formation. A wind event increased biotracer fluxes (POC, protein, chloroplast lipids). The rapid transmission of surface signals through extremely fast sinking particles could be a general feature of particle fluxes in marine areas unaffected by horizontal advection.     

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.     

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.     

Vertical distribution of zooplankton in Canary Island waters: implications for export flux

Zooplankton biomass, gut fluorescence and electron transfer system (ETS) activity were measured in vertical profiles (0–900 m) in two different size classes (<1 and >1 mm) in Canary Island waters. Both size fractions displayed a typical pattern of distribution with higher biomass, gut fluorescence and ETS in the shallower layers at night. By day, however, the vertical distribution varied between the size fractions, with higher biomass of the small fraction in the 0–200 m and a layer of large organisms at depth (∼500 m). For both size fractions, average ETS activity was higher by day than at night at depths between 200 and 600 m. Similarly, gut fluorescence was slightly higher by day below 200 m. The downward export of respiratory carbon was 1.92 and 4.29 mg C m⁻² d⁻¹ for samples obtained southwest of Gran Canaria Island and west of Tenerife Island respectively, being 2.68 mg C m⁻² d⁻¹ for the whole area. These values represented 16–45% (22–28% for the area) of the calculated passive particulate export production resulting from primary production. The estimated “gut flux” accounted for 0.35 (western zone) and 2.37 mg C m⁻² d⁻¹ (southwest of Gran Canaria), being 1.28 mg C m⁻² d⁻¹ for the whole area and represented between 3 and 25% (11–14% for the whole area) of the estimated passive particle export flux. These results agree with previous estimates and suggest that diel-migrant zooplankton can play an important role in the downward flux of carbon.     

Vertical flux of particulate Al,Fe, Pb,and Ba from the upper ocean estimated from 234Th/238U disequilibria

The vertical sinking flux of particulate Al, Fe, Pb, and Ba from the upper 250 m of the Labrador Sea has been estimated from measurements of ²³⁴Th/²³⁸U disequilibrium and the respective metal/²³⁴Th ratios in >53 μm size particles. ²³⁴Th-derived particulate metal fluxes include in situ scavenged metals, labile lithogenic metals, and metals derived from external input (e.g., atmospheric supply). In contrast to the POC/²³⁴Th ratio, particle size-fractionated (0.4–10 μm, 10–53 μm, and >53 μm) Al/²³⁴Th, Fe/²³⁴Th and Pb/²³⁴Th, and Ba/²³⁴Th ratios generally increase with depth and exhibit no systematic change with particle diameter. Sinking fluxes of particulate Al (2.47–22.3 μmol m⁻² d⁻¹), Fe (2.69–16.3 μmol m⁻² d⁻¹), Pb (2.85–70 nmol m⁻² d⁻¹), and Ba (0.13–2.1 μmol m⁻² d⁻¹) at 50 m (base of the euphotic zone) and 100 m (base of the mixed layer) are largely within the range of previous sediment trap results from other ocean basins. Estimates of the upper ocean residence time of Al (0.07–0.28 yr) and Pb (0.8–2.9 yr) are short compared to previously reported values. The settling rate of >53 μm particles calculated from the ²³⁴Th data ranges from 14 to 38 m d⁻¹.     

Biogeochemical cycling of N and Si during the mesoscale iron-enrichment experiment in the western subarctic Pacific (SEEDS-II)

Biogeochemical cycles of N and Si were examined in the surface mixed layer during the mesoscale iron-enrichment (IE) experiment in the high-nutrient low-chlorophyll (HNLC) western subarctic Pacific (SEEDS-II). Although the IEs increased nitrate uptake, silicic acid utilization was not stimulated. The nitrate drawdown in the iron-patch (IN-patch, 140.3 mmol m⁻² in the surface mixed layer, 0–30 m) was only 25% of the initial inventory, which was 1/3–2/5 of the previous IE experiments in the subarctic Pacific. This relatively weak response of nutrient drawdown to IEs was due to the high biomass of mesozooplankton (MZ) dominated by copepod Neocalanus plumchrus. Feeding of MZ (247.2 mmol m⁻² during Day 0–21 from the first IE) in the IN-patch was higher than the nitrate drawdown and prevented further development of the phytoplankton bloom. In the later period of the experiment (Day 14–21), the increase in the feeding activity and resultant decrease in phytoplankton biomass induced the accumulation of dissolved organic nitrogen (DON) and ammonium. Among total growth of MZ (81.6 mmol N m⁻²), 89% (72.8 mmol N m⁻²) was transported to the depth by the ontogenetic downward migration of N. plumchrus. Although silicic acid drawdown was not increased by the IEs, Si export flux increased by 2.7 times. The increase in Si export was also due to the increase in MZ, which egested faecal pellets with higher Si:N ratio and faster sinking speed than diatoms. The export efficiency (78% of new production) and total amount of export flux (143.8 mmol N m⁻², 1392 mmol C m⁻²) were highest records within the IE experiments despite weak responses of nutrient drawdown to the IE. During SEEDS-II, the high biomass of MZ reduced the phytoplankton response and nutrient drawdown to the IEs but via grazing and ontogenetic vertical migration accelerated the export flux as well as accumulations of dissolved forms of N. Results of the present and previous IE experiments indicate that the ecosystem and biogeochemical responses to IEs in the HNLC region are quite sensitive to the ecosystem components, especially for grazers of diatoms such as copepods and heterotrophic dinoflagellates. More attention needs to be paid to the ecosystem components and their biogeochemical functions as well as physical and chemical properties of the ecosystems in order to hindcast or forecast the impacts of changes in atmospheric iron deposition.