Population dynamics of phytoplankton, heterotrophic bacteria, and viruses during the spring bloom in the western subarctic Pacific

We characterized the community composition of phytoplankton in the western subarctic Pacific from the pre-bloom to the decline phase of the spring bloom with special reference to decreases in the silicic acid concentration in surface waters as an index for diatom bloom development. Furthermore, responses of heterotrophic bacteria and viruses to the spring bloom were also concomitantly investigated. Under pre-bloom conditions when nutrients were abundant but the surface mixed layer depth was relatively deep, chlorophyll (Chl) a concentrations were consistently low and green algae (chlorophytes and prasinophytes), cryptophytes, and diatoms were predominant in the phytoplankton assemblages as estimated by algal pigment signatures. Together with the shallowing of the mixed layer depth and the decrease in silicic acid concentration, diatoms bloomed remarkably in the Oyashio region, though the magnitude of the bloom in the Kuroshio-Oyashio transition (hereafter Transition) region was relatively small. A total of 77 diatom species were identified, with the bloom-forming diatoms mainly consisting of Thalassiosira, Chaetoceros, and Fragilariopsis species. It has become evident that the carotenoid fucoxanthin can serve as a strong indicator of the diatom carbon biomass during the spring diatom bloom. Differences in the species richness of diatoms among stations generally enabled us to separate the Oyashio bloom stations from the Transition and the Oyashio pre-bloom stations. Relatively high values of the Shannon-Wiener index for the diatom species were also maintained during the Oyashio bloom, indicating that a wide variety of species then shared dominance. In the decline phase of the Oyashio bloom when surface nutrient concentrations decreased, senescent diatom cells increased, as inferred from the levels of chlorophyllide a. Although the cell density of heterotrophic bacteria changed little with the development of the diatom bloom, viral abundance increased toward the end of the bloom, suggesting an increased likelihood of mortality among organisms including diatoms resulting from viral infection. This is the first report on the microbial trophodynamics, including viruses, during the spring diatom bloom in the western subarctic Pacific.     

Enrichment of alkanes within a phytoplankton bloom during an in situ iron enrichment experiment in the western subarctic Pacific

During the Subarctic Pacific Iron Experiment for Ecosystem Dynamics Study ΙΙ (SEEDS-II), we monitored variations in the concentrations of non-methane hydrocarbons (NMHCs), CH₃Cl, N₂O, and CH₄ within a phytoplankton bloom. Stable isotopic compositions were also determined to evaluate the sources of the variations. Although there was little variation in either the concentrations or the stable isotopic compositions of alkenes, CH₃Cl, N₂O, and CH₄ during the 23-day observation period, alkane concentrations increased substantially as the phytoplankton bloomed. The column-integrated quantities of alkanes increased to 3 times pre-bloom levels for C₂H₆, 5 times for C₃H₈, and 20 times for n-C₄H₁₀. The δ¹³C values of both C₂H₆ and C₃H₈ remained almost constant while concentrations increased, whereas that of n-C₄H₁₀ increased by about 12‰. To evaluate the sources of the alkanes produced during the bloom, we compared their δ¹³C values with those of alkanes produced in axenic phytoplankton cultures in our laboratory. We concluded that during the SEEDS-ΙΙ experiment the major portions of C₂H₆ and C₃H₈ were produced during the autolysis of diatoms cells, whereas n-C₄H₁₀ was produced during autolysis of other phytoplankton cells such as cryptophytes and dinoflagellates.     

Responses of diatoms to iron-enrichment (SEEDS) in the western subarctic Pacific, temporal and spatial comparisons

Phytoplankton species composition was analyzed inside and outside of the iron-enriched patch during the SEEDS experiment. Before the iron-enrichment, the phytoplankton community consisted of similar proportions of pico-, nano- and micro-sized phytoplankton, and the micro-phytoplankton was dominated by the pennate diatom Pseudo-nitzschia turgidula. Although all the diatoms, except the nano-sized Fragilariopsis sp., increased during the two weeks of the observation period, the flora in the patch dramatically changed with the increase of phytoplankton biomass to a centric diatom-dominated community. Neritic diatoms, especially Chaetoceros debilis, showed higher growth rates than other diatoms, without any delay in the initiation of growth after the enrichment, and accounted for 90% of the micro-phytoplankton after day 9. In contrast, the oceanic diatoms showed distinct delays in the initiation of growth. We conclude that the responses of the diatoms to the manipulation of iron concentration were different by species, and the fast and intensive response of the phytoplankton to iron-enrichment resulted from the presence of a small amount of neritic diatoms at the study site. The important factors that determine the dominant species in the bloom are the potential growth rates under an iron-replete condition and the growth lag. Abundant species in the patch are widely distributed in the North Pacific and their relative contributions in the Oyashio area and at Stn KNOT are high from spring to summer. However, a characteristic difference of species composition between the SEEDS bloom and natural blooms was the lack of Thalassiosira and Coscinodiscus species in the patch, which usually account for a major part of the phytoplankton community under blooming conditions in the western North Pacific.     

Seasonal and interannual variability in phytoplankton and nutrient dynamics along Line P in the NE subarctic Pacific

We analysed mixed-layer seasonal and interannual variability in phytoplankton biomass and macronutrient (NO₃ and Si(OH)₄) concentrations from three decades of observations, and nitrogen uptake rates from the 1990s along Line P in the NE subarctic Pacific. Chlorophyll a concentrations near 0.35 mg m⁻³ were observed year-round along Line P except at the nearshore station (P4) where chlorophyll a concentrations in spring were on average 2.4 times the winter values. In contrast, the temporal variability in carbon-to-chlorophyll ratios at the two main end members of Line P (P4 and OSP) was high. Large seasonal and interannual variability in NO₃ and Si(OH)₄ concentration were observed along Line P. Highest upper mixed-layer (top 15 m) nutrient concentrations occurred on the continental shelf in late summer and early fall due to seasonal coastal upwelling. Beyond the shelf, maximum nutrient concentrations increased gradually offshore, and were highest in late winter and early spring due to mixing by winter storms. Interannual variations in upper mixed-layer nutrient concentrations beyond the shelf (>128°W) were correlated with E-W winds and the PDO since 1988 but were not correlated with either climate index between 1973 and 1981. Despite differences in nutrient concentration, nutrient utilization (ΔNO₃ and ΔSi(OH)₄) during the growing season were about 7.5 μM at all offshore stations. Variations in ΔNO₃ were correlated with those of ΔSi(OH)₄. The annual cycle of absolute NO₃ uptake (ρNO₃) and NH₄ uptake (ρNH₄) rates by phytoplankton in the upper mixed-layer showed a weak increasing trend from winter to spring/summer for the period 1992-1997. Rates were more variable at the nearshore station (P4). Rates of ρNO₃ were low along the entire line despite abundant NO₃ and low iron (Fe), at the offshore portion of Line P and sufficient Fe at the nearshore station (P4). As a result, new production contributed on average to only 32 ± 15% of the total nitrogen (N) uptake along Line P. NO₃ utilization in the NE subarctic Pacific is probably controlled by a combination of environmental variables, including Fe, light and ambient NH₄ levels. Elevated ambient NH₄ concentrations seem to decrease the rates of new production (and f-ratios) in surface waters of the oceanic subarctic NE Pacific. Contrary to expectation, phytoplankton biomass, nutrient utilization (ΔNO₃ and ΔSi(OH)₄), and nitrogen uptake (ρNO₃ + ρNH₄) varied relatively little along Line P, despite significant differences in the factors controlling phytoplankton composition assemblages and production. Future studies would benefit from including other variables, especially light limitation, to improve our understanding of the seasonal and interannual variability in phytoplankton biomass and nutrients in this region.     

An in situ iron-enrichment experiment in the western subarctic Pacific (SEEDS): Introduction and summary

To test the iron hypothesis in the subarctic Pacific Ocean, an in situ iron-enrichment experiment (SEEDS) was performed in the western subarctic gyre in July–August 2001. About 350 kg of iron (as acidic iron sulfate) and 0.48 mol of the inert chemical tracer sulfur hexafluoride were introduced into a 10-m deep surface mixed layer over an 80 km² area. This single iron infusion raised dissolved iron levels to 2.9 nM initially. Dissolved iron concentrations rapidly decreased after the infusion, but levels remained close to 0.15 nM even at the end of the 14-day experimental period. During SEEDS there were iron-mediated increases in chlorophyll a concentrations (up to 20 μg l⁻¹), primary production rates, biomass and photosynthetic energy conversion efficiency relative to waters outside the iron-enriched patch. The rapid and very high accumulation of phytoplankton biomass in response to the iron addition appeared to be partly attributable to shallow mixed-layer depth and moderate water temperature in the western subarctic Pacific. However, the main reason was a floristic shift to fast-growing centric diatom Chaetoceros debilis, unlike the previous iron-enrichment experiments in the equatorial Pacific and the Southern Ocean, in both of which iron stimulated the growth of pennate diatoms. The iron-mediated blooming of diatoms resulted in a marked consumption of macronutrients and drawdown of pCO₂. Biological and physiological measurements indicate that phytoplankton growth in the patch became both light- and iron-limited, making phytoplankton biomass relatively constant after day 9. The increase in microzooplankton grazing rate after day 9 also influenced the net growth rate of phytoplankton. There was no significant increase in the export flux of carbon to depth during the 14-day occupation of the experimental site. The export flux between day 4 and day 13 was estimated to be only 13% of the integrated primary production in the iron-enriched patch. The major part of the carbon fixed by the diatom bloom remained in the surface mixed layer as biogenic particulate matter. Our findings support the hypothesis that iron limits phytoplankton growth and biomass in a ‘bottom up’ manner in this area, but the fate of algal carbon remains unknown.     

Mesozooplankton responses to iron-fertilization in the western subarctic Pacific (SEEDS2001)

A mesoscale iron-fertilization experiment was carried out in the western subarctic Pacific during summer 2001. The iron-patch was traced for 14 days after the fertilization, and the abundance and behavior of mesozooplankton were compared with those outside of the patch. The phytoplankton biomass in the patch rapidly increased to over 15 times the initial level by the later half of the observation period, and was composed of large-sized (>10 mm), centric diatoms. Dominant zooplankton species in the upper 200-m depth were large copepods: Neocalanus plumchrus, Neocalanus cristatus, Eucalanus bungii and Metridia pacifica. Mesozoplankton biomass as well as species composition did not change significantly in the patch over the observation period. Furthermore, no changes of vertical distribution or diel vertical migration were observed for any species or stages of mesozooplankton throughout the observation period. However, the abundance of the first copepodite stages of N. plumchrus and E. bungii increased several fold in the patch after the diatom bloom formation compared to the densities outside the patch. The increases of both species are considered to be due to lowered mortality during the egg and nauplius stages. Spawning of N. plumchrus takes place at depth using lipid storage, while spawning of E. bungii takes place in the surface layer supported by grazing. These facts suggest that the relative importance of nauplii in the diets of the large copepods was decreased in the patch by the diatom bloom. Gut-pigment contents of dominant copepods in the patch increased 4–18 times, and the maximum values were observed during the bloom peak. However, the grazing impact on phytoplankton was low throughout the experiment, especially during the bloom period (<6% of the primary production).     

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.     

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.     

Single particle analysis of oceanic suspended matter during the SEEDS II iron fertilization experiment

Suspended particles collected from surface seawater during the SEEDS II (Subarctic Iron Experiment for Ecosystem and Dynamics Study II) experiment were analyzed individually using an electron probe X-ray micro analyzer and characterized by size and elemental composition. Their numbers, relative abundances, and relative particle volume all showed clear differences between samples collected inside vs. outside the phytoplankton bloom that developed following the addition of iron. Throughout the study, Si-rich, Ca-rich and Organic particles were dominant and their number increased inside the fertilized patch; these particles accounted for 21%, 13% and 58% of the particles examined, respectively. Si-rich or Ca-rich particles commonly consisted of fragments of diatom frustules and coccolithophorids. There was consistently greater percentage of Ca-rich particles and lower percentage of Si-rich particles inside the patch than outside of it in number, but both types of these particles apparently occupied a larger volume inside the patch than outside of it. Organic particles, that showed having peaks in smaller diameter particles, increased apparently inside the patch with time after iron fertilization. The Organic particles had a more diverse mixture of both bio-related and crustal trace elements than the other types of particles. These results suggest that the increase in suspended particles following the iron enrichment was due to a combination of detrital material and live phytoplankton.     

Evidence for the grazing hypothesis: Grazing reduces phytoplankton responses of the HNLC ecosystem to iron enrichment in the western subarctic pacific (SEEDS II)

A mesoscale iron-enrichment study (SEEDS II) was carried out in the western subarctic Pacific in the summer of 2004. The iron patch was traced for 26 days, which included observations of the development and the decline of the bloom by mapping with sulfur hexafluoride. The experiment was conducted at almost the same location and the same season as SEEDS (previous iron-enrichment experiment). However, the results were very different between SEEDS and SEEDS II. A high accumulation of phytoplankton biomass (∼18 mg chl m⁻³) was characteristic of SEEDS. In contrast, in SEEDS II, the surface chlorophyll-a accumulation was lower, 0.8 to 2.48 mg m⁻³, with no prominent diatom bloom. Photosynthetic competence in terms of F _v/F _m for the total phytoplankton community in the surface waters increased after the iron enrichments and returned to the ambient level by day 20. These results suggest that the photosynthetic physiology of the phytoplankton assemblage was improved by the iron enrichments and returned to an iron-stressed condition during the declining phase of the bloom. Pico-phytoplankton (<2 μm) became dominant in the chlorophyll-a size distribution after the bloom. We observed a nitrate drawdown of 3.8 μM in the patch (day 21), but there was no difference in silicic acid concentration between inside and outside the patch. Mesozooplankton (copepod) biomass was three to five times higher during the bloom-development phase in SEEDS II than in SEEDS. The copepod biomass increased exponentially. The grazing rate estimation indicates that the copepod grazing prevented the formation of an extensive diatom bloom, which was observed in SEEDS, and led to the change to a pico-phytoplankton dominated community towards the end of the experiment.     

Effect of iron enrichment on the dynamics of transparent exopolymer particles in the western subarctic Pacific

Dynamics of transparent exopolymer particles (TEP) was studied during the first in situ iron-enrichment experiment conducted in the western subarctic Pacific in July–August 2001, with the goal of evaluating the contribution of TEP to vertical flux as a result of increased primary production following iron enrichment in open ocean ecosystems. Subsequent to the enhancement of phytoplankton production, we observed increase in TEP concentration in the surface layer and sedimentation of organic matter beneath it. Vertical profiles of TEP, chlorophyll a (Chl a) and particulate organic carbon (POC) were obtained from six depths between 5 and 70 m, from a station each located inside and outside the enriched patch. TEP and total mass flux were estimated from the floating sediment traps deployed at 200 m depth. Chl a and TEP concentrations outside the patch varied from 0.2 to 1.9 μg L⁻¹ and 40–60 μg XG equiv. L⁻¹, respectively. Inside the patch, Chl a increased drastically from day 7 reaching the peak of 19.2 μg L⁻¹ on day 13, which coincided with the TEP peak of 189 μg XG equiv. L⁻¹. TEP flux in the sediment trap increased from 41 to 88 mg XG equiv. m⁻² d⁻¹, with 8–14% contribution of TEP to total mass flux. This forms the basic data set on ambient concentrations of TEP in the western subarctic Pacific, and evaluation of the effect of iron enrichment on TEP.     

Temporal changes in community composition of heterotrophic bacteria during in situ iron enrichment in the western subarctic Pacific (SEEDS-II)

Little is known about the effects of iron enrichment in high-nitrate low-chlorophyll (HNLC) waters on the community composition of heterotrophic bacteria, which are crucial to nutrient recycling and microbial food webs. Using denaturing gradient gel electrophoresis (DGGE) of 16S rDNA fragments, we investigated the heterotrophic eubacterial community composition in surface waters during an in situ iron-enrichment experiment (SEEDS-II) in the western subarctic Pacific in the summer of 2004. DGGE fingerprints representing the community composition of eubacteria differed inside and outside the iron-enriched patch. Sequencing of DGGE bands revealed that at least five phylotypes of α-proteobacteria including Roseobacter, Cytophaga-Flavobacteria-Bacteroides (CFB), γ-proteobacteria, and Actinobacteria occurred in almost all samples from the iron-enriched patch. Diatoms did not bloom during SEEDS-II, but the eubacterial composition in the iron-enriched patch was similar to that in diatom blooms observed previously. Although dissolved organic carbon (DOC) accumulation was not detected in surface waters during SEEDS-II, growth of the Roseobacter clade might have been particularly stimulated after iron additions. Two identified phylotypes of CFB were closely related to the genus Saprospira, whose algicidal activity might degrade the phytoplankton assemblages increased by iron enrichment. These results suggest that the responses of heterotrophic bacteria to iron enrichment could differ among phylotypes during SEEDS-II.     

The response of phytoplankton to iron enrichment in Sub-Antarctic HNLCLSi waters: Results from the SAGE experiment

Areas of high nutrients and low chlorophyll a comprise nearly a third of the world’s oceans, including the equatorial Pacific, the Southern Ocean and the Sub-Arctic Pacific. The SOLAS Sea-Air Gas Exchange (SAGE) experiment was conducted in late summer, 2004, off the east coast of the South Island of New Zealand. The objective was to assess the response of phytoplankton in waters with low iron and silicic acid concentrations to iron enrichment. We monitored the quantum yield of photochemistry (F_v/F_m) with pulse amplitude modulated fluorometry, chlorophyll a, primary productivity, and taxonomic composition. Measurements of F_v/F_m indicated that the phytoplankton within the amended patch were relieved from iron stress (F_v/F_m approached 0.65). Although there was no significant difference between IN and OUT stations at points during the experiment, the eventual enhancement in chlorophyll a and primary productivity was twofold by the end of the 15-day patch occupation. However, no change in particulate carbon or nitrogen pools was detected. Enhancement in primary productivity and chlorophyll a were approximately equal for all phytoplankton size classes, resulting in a stable phytoplankton size distribution. Initial seed stocks of diatoms were extremely low, <1% of the assemblage based on HPLC pigment analysis, and did not respond to iron enrichment. The most dominant groups before and after iron enrichment were type 8 haptophytes and prasinophytes that were associated with ∼75% of chlorophyll a. Twofold enhancement of biomass estimated by flow cytometry was detected only in eukaryotic picoplankton, likely prasinophytes, type 8 haptophytes and/or pelagophytes. These results suggest that factors other than iron, such as silicic acid, light or physical disturbance limited the phytoplankton assemblage during the SAGE experiment. Furthermore, these results suggest that additional iron supply to the Sub-Antarctic under similar seasonal conditions and seed stock will most likely favor phytoplankton <2 ??m. This implies that any iron-mediated gain of fixed carbon will most likely be remineralized in shallow water rather than sink and be sequestered in the deep ocean.     

Nutrient and Plankton Dynamics in the NE and NW Gyres of the Subarctic Pacific Ocean

The western subarctic gyre (WSG) and the eastern Alaska Grye (AG) on each side of the subarctic North Pacific, have many similarities. In both gyres, macronutrients are generally high and chl is low, and hence both gyres are High Nitrate, Low Chlorophyll (HNLC) regions. Despite the general similarities between these two gyres, there are many important differences. The time series station established at Stn KNOT on the southwest edge of the WSG and two in situ mesoscale iron enrichment experiments at each of the gyres has provided more information on iron concentrations, the dual role of iron and silicate limitation and seasonal cycles in the gyres. There is more seasonality in many parameters at Stn KNOT than at Stn P. There is an increase in Chl and primary productivity at Stn KNOT in May followed by increased iron limitation in summer. Low DIC:NO₃ ratios and high Si:NO₃ ratios in the WSG, indicate lower calcification and higher diatom production than at Stn P. The sources of iron for these areas are still not clear, but horizontal transport of iron rich coastal water and vertical transport could be important sources at certain times of the year in addition to dust input. Satellite images show that chl-rich coastal waters occasionally extend to the vicinity of Stn KNOT and therefore Stn KNOT may not always represent conditions in the main part of the WSG. This review focuses mainly on a comparison of Stn KNOT and Stn P, two time series stations on the edge of two very large gyres. At present, we have a limited understanding of the temporal and spatial variability within each of these large gyres. This revised version was published online in July 2006 with corrections to the Cover Date.     

Flow cytometry and pigment analyses as tools to investigate the toxicity of herbicides to natural phytoplankton communities

Characterisation of natural phytoplanktonic communities is currently being advanced through flow cytometry and high resolution pigment analyses. To date, toxicological methods to assess impacts of herbicides on natural phytoplankton populations are lacking. Here, we report the novel use of these techniques in combination to study changes in phytoplankton populations exposed to 2-methylthio-4-tertiary-butylamino-6-cyclopropylamino-s-triazine (Irgarol 1051), a herbicide used in antifouling paints. Flow cytometry results revealed that following a 72-h exposure to approximately 100 ngL(-1), eukaryote abundance was less than half that in the controls. High performance liquid chromatographic analyses of pigments demonstrated that 19'-hexanoyloxyfucoxanthin was selectively lost relative to the control. This carotenoid is specific to the prymnesiophytes which are key constituents of phytoplanktonic communities within temperate marine waters. Values of EC(50) (72 h) as low as 70 ngL(-1) were calculated from the selective reduction in this compound. Concentrations substantially exceeding this level have been reported in UK and other European coastal waters.     

Community Structure and Dynamics of Phytoplankton in the Western Subarctic Pacific Ocean: A Synthesis

The phytoplankton community in the western subarctic Pacific (WSP) is composed mostly of pico- and nanophytoplankton. Chlorophyll a (Chl a) in the <2 μm size fraction accounted for more than half of the total Chl a in all seasons, with higher contributions of up to 75% of the total Chl a in summer and fall. The exception is the western boundary along the Kamchatka Peninsula and Kuril Islands and the Oyashio region where diatoms make up the majority of total Chl a during the spring bloom. Among the picophytoplankton, picoeukaryotes and Synechococcus are approximately equally abundant, but the former is more important in term of carbon biomass. Despite the lack of a clear seasonal variation in Chl a concentration, primary productivity showed a large seasonal variation, and was lowest in winter and highest in spring. Seasonal succession in the phytoplankton community is also evident with the abundance of diatoms peaking in May, followed by picoeukaryotes and Synechococcus in summer. The growth of phytoplankton (especially >10 μm cell size) in the western subarctic Pacific is often limited by iron bioavailability, and microzooplankton grazing keeps the standing stock of pico- and nano-phytoplankton low. Compared to the other HNLC regions (the eastern equatorial Pacific, the Southern Ocean, and the eastern subarctic Pacific), iron limitation in the Western Subarctic Gyre (WSG) may be less severe probably due to higher iron concentrations. The Oyashio region has similar physical condition, macronutrient supply and phytoplankton species compositions to the WSG, but much higher phytoplankton biomass and primary productivity. The difference between the Oyashio region and the WSG is also believed to be the results of difference in iron bioavailability in both regions. This revised version was published online in July 2006 with corrections to the Cover Date.     

Modelling the response of the planktonic food web to iron fertilization and warming in the NE subarctic Pacific

A one-dimensional ecosystem model with two explicit size classes of phytoplankton was developed for the NE subarctic Pacific to investigate variations in the export of organic particles to the ocean interior due to potential changes in the environment. Specifically, the responses of the planktonic ecosystem to permanent removal of iron limitation and to warming (of 2 and 5 °C) were explored. The ecosystem model consists of five components (small and large phytoplankton, microzooplankton, detritus and nitrogen), and includes grazing by mesozooplankton that varies in time according to long-term observations at Ocean Station Papa (OSP). The model addresses the role of iron limitation on phytoplankton growth and includes temperature dependence of physiological rates. The ecosystem model was forced with annual wind and solar heating from OSP. The model best reproduced the low chlorophyll high nitrate conditions of the NE subarctic Pacific when both small and large phytoplankton were limited by iron such that their maximum specific growth rate was reduced by 10 and 70%, respectively. Sensitivity analysis showed that model results depended on the value of the iron limitation parameter of large phytoplankton (L_Fe-L) and the grazing parameters of micro- and mesozooplankton. To explore the effect of iron limitation, simulations were carried out varying the iron limitation parameters while maintaining the nitrogen flux at the base of the model constant and the grazing pressure by mesozooplankton unchanged. In the warming case, simulations were carried out increasing ocean temperatures by 2° and 5 °C applied only to the ecological components, the flux of nitrate at the base of the model was increased to obtain a steady annual cycle, and grazing by mesozooplankton remained constant. When compared with the standard case, model simulations indicated that both permanent removal of iron limitation and warming cause changes in food web structure and the carbon cycle. The response was more dramatic in the iron-replete case where the phytoplankton community structure in spring changed from one dominated by pico- and nanoplankton to one dominated by large phytoplankton, and primary production increased until it consumed all the external nutrient (N) supply to the upper layer. However, reducing iron deficiency actually led to lower annual primary production due to a decrease in the regeneration of nitrogen in the euphotic zone. These changes in food web structure influenced the magnitude, composition and seasonal cycle of sinking particles.     

Responses of DMS in the seawater and atmosphere to iron enrichment in the subarctic western North Pacific (SEEDS-II)

Simultaneous measurements of dimethylsulfide (DMS) in the seawater and atmosphere were conducted during SEEDS-II to investigate the responses of DMS to iron (Fe) fertilization in the subarctic North Pacific. No significant increases in the seawater DMS (DMSw) concentration were observed inside the fertilized patch compared to those outside the patch, while particulate dimethylsulfoniopropionate (DMSPp) concentration inside the patch increased 2-fold compared to those outside the patch in the phytoplankton bloom of major DMSP producers such as prasinophytes, cryptophytes, diatoms and prymnesiophytes. In the decline phase of the bloom, maximum DMSw was observed both inside the patch (ca. 6.2 nM) and outside the patch (ca. 9.3 nM). In this period, increases in mesozooplankton and decreases in the DMSP producers (prymnesiophytes and diatoms) were observed both sides of the patch, but larger inside the patch than outside the patch. Large decreases in the DMSPp inside the patch, which was probably related to the large increases in mesozooplankton inside the patch, did not result in increases in the DMSw concentration. Considering biological and nonbiological parameters, we discussed these results, although they could not be completely explained. Unfortunately, the impact of Fe fertilization on the atmospheric DMS (DMSa) concentration was not detected due to no significant changes in DMSw. However, it is noted that DMSa concentrations were dependent on the sea–air DMS flux in the air from higher latitudes and/or the Eurasian continent, though the DMS flux was a minor role to the budget of DMSw. Therefore if DMSw were significantly changed by Fe fertilization, DMSa might be affected through changes in the sea-air flux in this condition.     

Impact of environmental factors on in situ determination of iron in seawater by flow injection analysis

A sensitive method for iron determination in seawater has been adapted on a submersible chemical analyser for in situ measurements. The technique is based on flow injection analysis (FIA) coupled with spectrophotometric detection. When direct injection of seawater was used, the detection limit was 1.6 nM, and the precision 7%, for a triplicate injection of a 4 nM standard. At low iron concentrations, on line preconcentration using a column filled with 8-hydroxyquinoline (8HQ) resin was used. The detection limit was 0.15 nM (time of preconcentration = 240 s), and the precision 6%, for a triplicate determination of a 1 nM standard, allowing the determination of Fe in most of the oceanic regimes, except the most depleted surface waters. The effect of temperature, pressure, salinity, copper, manganese, and iron speciation on the response of the analyser was investigated. The slope of the calibration curves followed a linear relation as a function of pressure (C_p = 2.8 × 10^{− 5}P + 3.4 × 10^{− 2} s nmol^{− 1}, R² = 0.997, for Θ = 13 °C) and an exponential relation as a function of temperature (C_Θ = 0.009e^0.103Θ, R² = 0.832, for P = 3 bar). No statistical difference at 95% confidence level was observed for samples of different salinities (S = 0, 20, 35). Only very high concentration of copper (1000 × [Fe]) produced a detectable interference. The chemical analyser was deployed in the coastal environment of the Bay of Brest to investigate the effect of iron speciation on the response of the analyser. Direct injection was used and seawater samples were acidified on line for 80 s. Dissolved iron (DFe, filtered seawater (0.4 μm), acidified and stored at pH 1.8) corresponded to 29 ± 4% of Fe_a (unfiltered seawater, acidified in line at pH 1.8 for 80 s). Most of Fe_a (71 ± 4%) was probably a fraction of total dissolvable iron (TDFe, unfiltered seawater, acidified and stored at pH 1.8).     

Silicate to Nitrate Ratio of the Upper Sub-Arctic Pacific and the Bering Sea Basin in Summer: Its Implication for Phytoplankton Dynamics

Consumption of silicate and nitrate (Si:N molar ratio) in the upper layer of the pelagic subarctic Pacific in summer was evaluated by a regression analysis of silicate vs. nitrate concentrations at the upper 100 m depth. Based on data of three cruises, the pelagic subarctic Pacific can be classified into two groups. First group is characterized by roughly 1:1 consumption of silicate and nitrate, and occupies rather larger area of subarcfic Pacific, i.e., the Gulf of Alaska and the Western Subarctic gyre (averaged slope of Si:N linear regression: 1.21, n = 10 and 1.45, n = 9, respectively). Second group is the regions of the Bering Sea basin and the Oyashio region, and showed higher silicate consumption compared to that of nitrate (averaged slope of Si:N linear regression: 2.14, n = 9 and 2.36, n = 3, respectively). The Si:N difference observed is possibly attributed to relative contribution of diatoms production among the phytoplankton assemblages in the regions, i.e., dominance of diatoms production in the regions of the second group. Higher accumulation of ammonium at the bottom of euphotic layer in the summer Bering Sea basin would also contribute to increase consumption ratio of Si:N amounts.