Phytoplankton dynamics associated with the monsoon in the Sulu Sea as revealed by pigment signature

Phytoplankton dynamics during the northeast monsoon was investigated in the Sulu Sea from algal pigment analysis. We visited the Sulu Sea in February 2000, a mid period of the northeast monsoon, and in November and December 2002, the beginning of the northeast monsoon. SeaWiFS images showed generally low concentrations of surface chlorophyll a (Chl a) during the southwest monsoon and higher concentrations with several peaks during the northeast monsoon. In the beginning of the northeast monsoon, subsurface chlorophyll maxima (SCM) occurred, where vertical variation in class-specific composition as estimated from pigment signatures was prominent. Prochlorococcus, cyanobacteria, prymnesiophytes and crysophytes were important groups above the SCM, and the contribution of cyanobacteria to Chl a became much lower at and below the SCM. Contributions of chlorophytes and prasinophytes to Chl a generally showed maxima near the SCM. This distribution was accompanied by vertical changes in the concentration of photoprotective pigments relative to photosynthetic accessory pigments. During the mid northeast monsoon, the upward supply of nutrients was probably enhanced at some stations due to vertical mixing, and as a consequence diatoms dominated in the upper 100 m water column of these stations, and other eukaryotic flagellates including prymnesiophytes, chrysophytes and cryptophytes were secondary major components of the community. The elevation of Chl a concentration and changes in phytoplankton community during the northeast monsoon likely influence the variation in biological production at higher trophic levels in the Sulu Sea.     

Growth stimulation and inhibition of natural phytoplankton communities by model organic ligands in the western subarctic Pacific

The influence of organic ligands on natural phytoplankton growth was investigated in high-nitrate low-chlorophyll (HNLC) waters and during a phytoplankton bloom induced by a mesoscale iron enrichment experiment (SEEDS II) in the western subarctic Pacific. The growth responses of the phytoplankton in the treatments with iron complexed with model ligand were compared with those with inorganic iron or a control. Desferrioxamine B and protoporphyrin IX were used as models for hydroxamate-type siderophore and tetrapyrrole-type cell breakdown ligand, respectively. In the HNLC water, iron associated with protoporphyrin IX especially stimulated smaller phytoplankton (<10 μm) growth, 1.5-fold more than did inorganic iron. Surprisingly, only the addition of protoporphyrin IX stimulated small phytoplankton growth, suggesting that these cell breakdown ligands might be more bioavailable for them. The protoporphyrin IX’s stimulatory effect on small phytoplankton was not observed during bloom decline phase. The growth of phytoplankton was inhibited in the treatment with desferrioxamine B-complexed iron, suggesting its low bioavailability for the natural phytoplankton community. Its inhibitory effects were particularly pronounced in pico-eukaryotic phytoplankton. During the iron-induced bloom, the phytoplankton’s iron-stress response gradually increased with the desferrioxamine B concentration, suggesting that the competition for iron complexation between natural ligands and desferrioxamine B affected phytoplankton growth. However, the pico-eukaryotes did seem better able to utilize the desferrioxamine B-complexed iron during the bloom-developing phase. These results indicate that the iron bioavailability for phytoplankton differs between bloom-developing and bloom-decline phases.     

Effects of nitrogen and iron enrichments on phytoplankton communities in the Northwestern Indian Ocean

Nutrient-enrichment bottle experiments in the northwestern Indian Ocean surface waters were conducted to investigate phytoplankton growth following enrichments with either NH₄⁺, NO₃⁻, Fe or Fe + NO₃⁻. Stimulation of phytoplankton growth could be achieved by the addition of either NH₄⁺ or NO₃⁻ under the ambient Fe concentrations, but the most significant increases in Chl a, POC, and cell densities were observed in the Fe + NO₃⁻-amended culture. Iron addition caused more rapid responses of phytoplankton growth in the Fe + NO₃⁻ treatment than those in the NO₃⁻ and NH₄⁻ treatment. However, the Fe-enrichment treatment revealed minimal growth of phytoplankton because of severe major nutrient deficiency and was similar to the control treatment. Increases in the cell density of diatoms and spherical phytoplankton cells (< 10 μm) were significant in the NH₄⁺-enriched samples, whereas NO₃⁻ enrichment alone had little effect on the diatoms. Simultaneous addition of Fe and NO₃⁻ stimulated maximal growth of phytoplankton, in particular in diatoms, coccolithophorids and Phaeocystis type colonies. However, the dominance of coccolithophorids and Phaeocystis type colonies in the Fe + NO₃⁻ treatment may be interpreted as resulting from Si-limitation. The high N/P ratio for phytoplankton nutrient uptake in the N-amended culture indicates the possibility of some P-limited growth. From these results, we conclude that in the northwestern Indian Ocean, Fe and major nutrients are co-limiting phytoplankton production during the northeast monsoon. Iron appeared to affect the ability of phytoplankton to respond quickly to transient nutrient inputs.     

Optical,biological, and chemical properties ofAoshio,hypoxic milky blue-green water,observed at the head of Tokyo Bay

Aoshio-hypoxic water which can be distinguished from surrounding coastal water by its milky blue-green color-is often observed at the head of Tokyo Bay when coastal upwelling of sulfidic bottom water occurs during summer and early autumn. Optical properties and the biological-chemical composition of suspended matter in theaoshio water were investigated using nearby “normal” coastal water as a reference. Suspended matter in theaoshio water contributed to the large beam-attenuation coefficient and also to the strong upward radiance with maximal intensity around 550 nm as a result of the scattering process. No significant difference in bacterial density was observed between samples ofaoshio water and reference coastal water. The low chlorophylls concentration inaoshio water samples indicates that biological pigments may not play a dominant role in the color of theaoshio events. The suspended particles inaoshio water samples had much higher contents of sulfur and manganese and a lower silicon content than those in reference coastal water samples. The colloidal elemental sulfur and manganese-rich particles identified in theaoshio water appear to be oxidation products of dissolved sulfide and manganese (II) in the anoxic water. On the basis of these findings, it may be concluded that there is a high probability that elemental sulfur and manganese-rich particles are responsible for the color and turbidity inaoshio.     

Physical behavior of the SEEDS iron-fertilized patch by sulphur hexafluoride tracer release

The first iron (Fe) – fertilization experiment in the western North Pacific was carried out using SF₆ to trace the Fe-fertilized water mass. A solution in 10,800 liters of seawater of 350 kg of Fe and 0.48 M of SF₆ tracer was released into the mixed layer over a 8 × 10 km area. On the first underway transects through the patch after the Fe release, we observed a significant increase of dissolved Fe (ave. 2.89 nM). The fertilized patch was traced for 14 days by on-board SF₆ analysis. A Lagrangian frame of reference was maintained by the use of a drogued GPS buoy released at the center of the patch. The patch moved westward at a rate of 6.8 km d⁻¹. Mixed layer depth increased from 8.5 to 15 m during the experiment. Horizontal diffusivity was determined by the change of SF₆ concentration in the patch. The horizontal diffusivity increased during the experiment. We evaluate here the fate of Fe in a Fe-fertilized patch using the dilution rate determined from sulphur hexafluoride (SF₆) concentration. Dissolved Fe concentrations subsequently decreased rapidly to 0.15 nM on Day 13. However, the dissolved Fe half-life of 43 h was relatively longer than in previous Fe-enrichment studies, and we observed a larger increase of the centric diatom standing stock and corresponding drawdown of macro-nutrients and carbon dioxide than in the previous studies. The most important reason for the larger response was the phytoplankton species in the western North Pacific. In addition, the smaller diffusivity and shallower mixed layer were effective to sustain the higher dissolved Fe concentration compared to previous experiments. This might be one reason for the larger response of diatoms in SEEDS.     

Effects of long-term sample preservation on flow cytometric analysis of natural populations of pico-and nanophytoplankton

The effects of long-term preservation on flow cytometric parameters of natural oceanic populations of pico-and nanophytoplankton have been examined. Populations collected at oligotrophic subtropical and subarctic locations in the North Pacific were fixed with glutaraldehyde and frozen in liquid nitrogen, according to Vaulot et al. (1989). During six months’ storage, chlorophyll red fluorescence declined in all the groups examined, while forward light scatter was enhanced in Synechococcus and Prochlorococcus, and weakened in nanoeucaryotes. Cell loss was not significant except for Synechococcus. Caution is required when analyzing flow cytometric data of samples stored for more than a month.     

Influence of non-photosynthetic pigments on light absorption and quantum yield of photosynthesis in the western equatorial Pacific and the Subarctic North Pacific

Regional variations in the contribution of non-photosynthetic pigments (ā _np^*) to the total light absorption of phytoplankton (ā _ph^*) and its influence on the maximum quantum yield of photosynthesis (φ _m) were investigated. In the western equatorial Pacific, the surface ā _np^* : ā _ph^* ratio was higher in the western warm pool than that in the upwelling region. This difference appears to be attributable to severe nitrate depletion and higher percentage of prokaryotes, which can accumulate very high concentrations of zeaxanthin in the western warm pool. In the subarctic North Pacific, the ā _np^* : ā _ph^* ratio was expected to be higher in the Alaskan Gyre where the thermocline is sharper and iron limitation may possibly be more severe than in the Western Subarctic Gyre. However, the ratio was actually higher in the Western Subarctic Gyre, contradictory to our expectations. This east-west variation appears to be attributable to changes in the taxonomic composition; cyanobacteria were more abundant in the Western Subarctic Gyre. The values of ā _np^* : ā _ph^* and its vertical variations were relatively small in the subarctic North Pacific compared to those in the western equatorial Pacific. These inter-regional variations appear to be attributable to the lower solar radiation intensity, smaller percentage of cyanobacteria, and relatively strong vertical mixing in the subarctic North Pacific. The spatial variations in ā _np^* : ā _ph^* significantly influence φ _m. In comparison with φ _m based on the total light absorption (φ _{m ph}), the values corrected for the contribution of non-photosynthetic pigments (φ _{m ps}) showed an increase in both the western equatorial Pacific and the subarctic North Pacific.     

Dissolved iron and zinc in Sagami Bay and the Izu-Ogasawara Trench

Using a clean seawater sampling system for trace metals onboard the R. V. Shinsei-Maru, newly launched in 2013, we investigated the vertical distributions of dissolved iron and zinc in Sagami Bay and the Izu-Ogasawara Trench. We applied appropriate clean sampling and filtering processes for trace metals, so that uncontaminated seawater samples were successfully collected. The distribution of zinc in the trench area was similar to that of silicate and the same as that previously reported in the subtropical North Pacific. There were spatial variations in the iron (Fe) distribution in the trench areas. We used previously reported information about biogeochemical cycling in the trench area, and found that Fe has a residence time of 29 years in the water column. The short residence time of Fe (29 years) corresponds to the vertical variations of dissolved Fe in the water column.     

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.     

Mesoscale decrease of surface phosphate and associated phytoplankton dynamics in the vicinity of the subtropical South Pacific islands

Surface distributions of nutrients and phytoplankton were investigated in the vicinity of the subtropical South Pacific islands by using a continuous underway system with a highly sensitive nutrient analysis. A total of 17 transects, whose lengths ranged between 42 and 271 km, were sampled for continuous nutrient measurements. The study area was characterized by an overall depletion of nitrate+nitrite (<15 nM), but phosphate varied from 7 to 192 nM. The transects were grouped into 4 patterns according to distribution of phosphate concentrations. In 7 transects, a mesoscale decrease in phosphate occurred, coinciding with an elevation of in vivo chlorophyll fluorescence, which was accompanied by an increase in phytoplankton abundance as revealed by microscopy, flow cytometry, and accessory pigments. This mirror–image relationship between the phosphate concentration and phytoplankton abundance was most apparent on both a 99-km transect east of Tonga, where the phosphate concentration ranged from 17 to 125 nM, and on a 98-km transect west of Fiji, where the phosphate concentration ranged from 23 to 136 nM. Both these transects contained distinct blooms of Trichodesmium in areas with the lowest concentrations of phosphate. In other 2 transects, fluctuations in phosphate concentrations showed no distinct relationship with chlorophyll fluorescence. Other patterns that emerged included consistently high concentrations, ranging from 109 to 192 nM, in 5 transects and consistently low phosphate concentrations, ranging from 7 to 50 nM, in 3 transects. Abundance of Trichodesmium, Prochlorococcus, Synechococcus, and all accessory pigments examined tended to be higher in the low phosphate transects than in the high phosphate ones. In particular, Trichodesmium occurred in low phosphate water (<25 nM). There was no significant relationship between phosphate concentrations and nanoplanktonic unicellular cyanobacteria. Our observations suggest that surface phosphate decreases are associated with phytoplankton utilization of phosphate, and that nitrogen supply from Trichodesmium may contributes to this utilization.