Iron-induced alterations of bacterial DMSP metabolism in the western subarctic Pacific during SEEDS-II

The effect of added iron on bacterial cycling of the climate-active gas dimethylsulfide (DMS) and its precursor dimethylsulfoniopropionate (DMSP) was tested during the second Subarctic Pacific Iron Experiment for Ecosystem Dynamics Study (SEEDS II) from 19 July to 21 August 2004 aboard the R/V Hakuho-Maru. The study area in the northwest Pacific Ocean (48°N 165°E) was enriched with Fe and the conservative tracer, SF₆, allowing the fertilized patch to be tracked. Microbial DMSP cycling rates were determined in the surface mixed layer (5 m) during incubations using the ³⁵S-DMSP technique. The addition of iron resulted in a 4-fold increase in concentrations of chlorophyll a (chl a) within the surface mixed layer (5 m depth), and the length of the sampling period allowed the observation of both bloom and post-bloom conditions. Inside the fertilized patch, the alleviation of resource limitation gave rise to the concurrent increase in bacterial abundance and production. Changes in the phytoplankton community within the Fe-enriched patch translated into a sustained decrease in chl a-normalized particulate DMSP (DMSP_p) concentrations, suggesting a preferential stimulation of the growth of DMSP_p-poor phytoplankton species. Despite short-lived peaks of DMSP_p within the Fe-enriched area, concentrations of DMSP_p generally remained stable during the entire sampling period inside and outside the fertilized patch. During the Fe-induced bloom, microbial DMSP-sulfur (DMSP-S) assimilation efficiency increased 2.6-fold inside the Fe-enriched area, which indicated that as bacterial production increased, a greater proportion of DMSP-S was assimilated and possibly diverted away from the bacterial cleavage pathway (i.e. production of DMS). Our results suggest that iron-induced stimulation of weak DMSP_p-producers and DMSP-assimilating bacteria may diminish the potential production of DMS and thus limit its flux towards the atmosphere over the subarctic Pacific Ocean.     

Spatial variations of dimethylsulfide and dimethylsulfoniopropionate in the surface microlayer and in the subsurface waters of the South China Sea during springtime

Spatial variations in dimethylsulfide (DMS) and dimethylsulfoniopropionate (DMSP) were surveyed in the surface microlayer and in the subsurface waters of the low productivity South China Sea in May 2005. Overall, average subsurface water concentrations of DMS and DMSP of dissolved (DMSPd) and particulate (DMSPp) fractions were 1.74 (1.00-2.50), 3.92 (2.21-6.54) and 6.06 (3.40-8.68) nM, respectively. No enrichment in DMS and DMSPp was observed in the microlayer. In contrast, the microlayer showed a DMSPd enrichment, with an average enrichment factor (EF, defined as the ratio of the microlayer concentration to subsurface water concentration) of 1.40. In the study area, none of the sulfur components were correlated with chlorophyll a. An important finding in this study was that DMS, DMSP and chlorophyll a concentrations in the surface microlayer were respectively correlated with those in the subsurface water, suggesting a close linkage between these two water bodies. The ratios of DMS:Chl-a and DMSPp:Chl-a showed a gradually increasing trend from North to South. This might be due to changes in the proportion of DMSP producers in the phytoplankton community with the increased surface seawater temperature. A clear diurnal variation in the DMS and DMSP concentrations was observed at an anchor station with the highest concentrations appearing during the day and the lowest concentrations during the night. The higher DMS and DMSP concentrations during daytime might be attributed to the light-induced increase in both algal synthesis and exudation of DMSP and biological production of DMS. The mean flux of DMS from the investigated area to the atmosphere was estimated to be 2.06 micromo lm(-2)d(-1). This low DMS emission flux, together with the low DMS surface concentrations was attributed to the low productivity in this sea.     

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.     

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.     

Changes in iron concentrations and bio-availability during an open-ocean mesoscale iron enrichment in the western subarctic Pacific,SEEDS II

A patch of water in the western subarctic gyre (low iron concentration, <0.02 nM) was fertilized twice with 322 and 159 kg of iron to induce a phytoplankton bloom. In order to understand the changes in iron distribution and bio-availability throughout the evolution and termination phase of the iron-induced bloom, iron concentrations were measured at stations inside and outside of the iron-fertilized patch, and shipboard culture experiments using iron and desferrioxamine B (DFB) inoculation to regulate iron availability were conducted 5 times with water collected from the center of the iron-fertilized patch on D2, D7, D11, D17 and D23.After the iron fertilization, we observed a significant increase in dissolved iron (1.38 nM at 5 m depth) at the center of the patch (D1). Dissolved iron concentrations subsequently decreased to an ambient level (~0.08 nM) on D16–D17, despite the second iron fertilization made on D6. During the 4-day incubations of the shipboard culture experiments, excess DFB-inoculated treatment inhibited the phytoplankton growth compared to the controls for D2, D7 and D11 patch water. This indicated that available iron existed in the iron-fertilized patch at least until D11. Moreover, iron-inoculated treatments induced growth of large-sized phytoplankton with an accompanying silicate decrease for D7, D11 and D17 patch water, but not for D23 patch water. These results indicated that large diatoms, which can respond to additional iron inoculation, existed in the iron-fertilized patch in evolution and early termination phase of the iron-induced bloom (at least until D17); however, there was no significant amount of large diatoms, which could rapidly respond to iron, in late termination phase (D23) of the iron-induced phytoplankton bloom.     

Dimethylsulphide and dimethylsulphoniopropionate in Antarctic sea ice and their release during sea ice melting

This study presents concentrations of dimethylsulphide (DMS) and its precursor compound dimethylsulphoniopropionate (DMSP) in a variety of sea ice and seawater habitats in the Antarctic Sea Ice Zone (ASIZ) during spring and summer. Sixty-two sea ice cores of pack and fast ice were collected from twenty-seven sites across an area of the eastern ASIZ (64°E to 110°E; and the Antarctic coastline north to 62°S). Concentrations of DMS in 81 sections of sea ice ranged from < 0.3 to 75 nM, with an average of 12 nM. DMSP in 60 whole sea ice cores ranged from 25 to 796 nM and showed a negative relationship with ice thickness (y = 125x^− 0.8). Extremely high DMSP concentrations were found in 2 cores of rafted sea ice (2910 and 1110 nM). The relationship of DMSP with ice thickness (excluding rafted ice) suggests that the release of large amounts of DMSP during sea ice melting may occur in discrete areas defined by ice thickness distribution, and may produce ‘hot spots’ of elevated seawater DMS concentration of the order of 100 nM. During early summer across a 500 km transect through melting pack ice, elevated DMS concentrations (range 21–37 nM, mean 31 nM, n = 15) were found in surface seawater. This band of elevated DMS concentration appeared to have been associated with the release of sea ice DMS and DMSP rather than in situ production by an ice edge algal bloom, as chlorophyll a concentrations were relatively low (0.09–0.42 μg l^− 1). During fast ice melting in the area of Davis station, Prydz Bay, sea ice DMSP was released mostly as extracellular DMSP, since intracellular DMSP was negligible in both hyposaline brine (5 ppt) and in a melt water lens (4–5 ppt), while extracellular DMSP concentrations were as high as 149 and 54 nM, respectively in these habitats. DMS in a melt water lens was relatively high at 11 nM. During the ice-free summer in the coastal Davis area, DMS concentrations in surface seawater were highest immediately following breakout of the fast ice cover in late December (range 5–14 nM), and then remained at relatively low concentrations through to late February (< 0.3–6 nM). These measurements support the view that the melting of Antarctic sea ice produces elevated seawater DMS due to release of sea ice DMS and DMSP.     

Occurrence and turnover of DMSP and DMS in deep waters of the Ross Sea,Antarctica

High concentrations of the phytoplankton metabolite dimethylsulfoniopropionate (DMSP) and its degradation product dimethylsulfide (DMS) are associated with blooms of Phaeocystis antarctica in the Ross Sea, Antarctica. Episodic and rapid vertical export of Phaeocystis biomass to deep water has been reported for the Ross Sea, therefore we examined the distribution and microbial consumption rates of DMSP and DMS throughout the sub-euphotic water column. Total DMSP (dissolved+particulate; DMSPt) was present at 0.5–22 nM at depths between 70 and 690 m during both the early bloom (November) and the late bloom (January). Sub-euphotic peaks of DMSP were sometimes associated with mid-water temperature maxima, and elevated DMSP below 70 m was found mainly in water masses characterized as Modified Circumpolar Deep Water or Antarctic Shelf Water. Overall, 50–94% of the integrated water-column DMSPt was found below the euphotic zone. At one station during the early bloom, local maxima of DMSPt (14 nM) and DMS (20 nM) were observed between 113 and 240 m and these maxima corresponded with high chlorophyll a concentrations, P. antarctica cell numbers, and Fv/Fm (the quantum yield of photosystem II). During the late bloom, a sub-euphotic maximum of DMSPt (15.8 nM) at 250 m cooccurred with peaks of chlorophyll a concentration, DMSP lyase activity, bacterial production and dissolved DMSP consumption rates. DMSP turnover contributed ～12% of the bacterial carbon demand between 200 and 400 m. DMS concentrations peaked at 286 m but the maximum concentration (0.42 nM) was far lower than observed during the early bloom, probably because of relatively rapid biological consumption of DMS (1–3 turnovers per day) which, in turn, contributed to elevated dissolved dimethylsulfoxide (DMSO) concentrations. Relatively stable DMSPt distributions at some sites suggest that rapid sinking of Phaeocystis biomass is probably not the major mechanism responsible for mesopelagic DMSP accumulations. Rather, subduction of near-surface water masses, lateral advective transport or trapping of slowly sinking P. antarctica biomass in intermediate water masses are more likely mechanisms. We found that a culture of P. antarctica maintained cellular integrity during 34 days of darkness, therefore the presence of intact cells (and DMSP) at depth can be explained even under a slow sinking/advection scenario. Whatever the mechanism, the large pools of DMSP and DMS below the euphotic zone suggest that export exerts a control on potential DMS emission from the surface waters of the Ross Sea.     

Seasonal variations of dimethylsulfide (DMS) and dimethylsulfoniopropionate (DMSP) in the sea-surface microlayer and subsurface water of Jiaozhou Bay and its adjacent area

The distributions of DMS and its precursor dimethylsulfoniopropionate, in both dissolved (DMSP_d) and particulate fractions (DMSP_p) were determined in the seasurface microlayer and corresponding subsurface water of the Jiaozhou Bay, China and its adjacent area in May and August 2006. The concentrations of all these components showed a clear seasonal variation, with higher concentrations occurring in summer. This can be mainly attributed to the higher phytoplankton biomass observed in summer. Simultaneously, the enrichment extents of DMSP_d and DMSP_p in the microlayer also exhibited seasonal changes, with higher values in spring and lower ones in summer. Higher water temperature and stronger radiant intensity in summer can enhance their solubility and photochemical reaction in the microlayer water, reducing their enrichment factors (the ratio of concentration in the microlayer to that in the corresponding subsurface water). A statistically significant relationship was found between the microlayer and subsurface water concentrations of DMS, DMSP and chlorophyll a, demonstrating that the biogenic materials in the microlayer come primarily from the underlying water. Moreover, our data show that the concentrations of DMSP_p and DMS were significantly correlated with the levels of chlorophyll a, indicating that phytoplankton biomass might play an important role in controlling the distributions of biogenic sulfurs in the study area. The ratios of DMS/chlorophyll a and DMSP_p/chlorophyll a varied little from spring to summer, suggesting that there was no obvious change in the proportion of DMSP producers in the phytoplankton community. The mean sea-to-air flux of DMS from the study area was estimated to be 5.70 μmol/(m²·d), which highlights the effects of human impacts on DMS emission.     

Profiles of DMSP,algal pigments,nutrients and salinity in pack ice from eastern Antarctica

Concentrations of dimethylsulphoniopropionate (DMSP) were measured in seven pack ice cores from three sites in eastern Antarctica to determine their relation to algal pigments, nutrients (nitrate, silicate and phosphate) and bulk salinity. The algal groups haptophytes, dinoflagellates and diatoms were identified in surface, interior and bottom assemblages in the pack ice cores using the photosynthetic marker pigments 19′-hexanoyloxyfucoxanthin (HEX), peridinin (PER) and fucoxanthin (FUC), respectively. DMSP concentrations were significantly correlated (P<0.01, Pearson) with chlorophyll-a (r=0.58), HEX (r=0.75), PER (r=0.79) and FUC (r=0.63) concentrations. The pool of DMSP within the pack ice (mean 107 nM) was contributed mainly by interior and bottom algal assemblages (mean 94 and 268 nM, respectively), whilst the surface algal assemblages were minor contributors (mean 18 nM). DMSP production and/or accumulation appears to differ between surface, interior and bottom pack ice algal assemblages due to differences in biomass, class composition, and possibly the unique environmental conditions experienced by each assemblage. In pack ice, diatoms appear to be important producers of DMSP, due to their dominance of algal assemblages.     

Particle flux characterisation and sedimentation patterns of protistan plankton during the iron fertilisation experiment LOHAFEX in the Southern Ocean

The taxonomic composition and types of particles comprising the downward particle flux were examined during the mesoscale artificial iron fertilisation experiment LOHAFEX. The experiment was conducted in low-silicate waters of the Atlantic Sector of the Southern Ocean during austral summer (January–March 2009), and induced a bloom dominated by small flagellates. Downward particle flux was low throughout the experiment, and not enhanced by addition of iron; neutrally buoyant sediment traps contained mostly faecal pellets and faecal material apparently reprocessed by mesozooplankton. TEP fluxes were low, ≤5 mg GX eq. m⁻² d⁻¹, and a few phytodetrital aggregates were found in the sediment traps. Only a few per cent of the POC flux was found in the traps consisting of intact protist plankton, although remains of taxa with hard body parts (diatoms, tintinnids, thecate dinoflagellates and foraminifera) were numerous, far more so than intact specimens of these taxa. Nevertheless, many small flagellates and coccoid cells, belonging to the pico- and nanoplankton, were found in the traps, and these small, soft-bodied cells probably contributed the majority of downward POC flux via mesozooplankton grazing and faecal pellet export. TEP likely played an important role by aggregating these small cells, and making them more readily available to mesozooplankton grazers.     

Significance of vertical flux as a sink for surface water DMSP and as a source for the sediment surface in coastal zones of northern Europe

In April 1997 and 1998 the significance of sedimentation as a sink for epipelagic dimethylsulphoniopropionate (DMSP) production and as a source for marine sediments was reassessed using a newly designed sediment trap. The behaviour of the traps in immersion was monitored continuously and the collection efficiency was evaluated with ²³⁴Th measurements. Net DMS(P) fluxes were corrected for some physical and biological losses during the whole sedimentation process providing reliable estimates of gross DMSP fluxes. It is shown that daily losses by sedimentation account for between 0.1% and 16% of seawater particulate DMSP (DMSPp) standing stocks, and between 3% and 75% of daily DMSPp production. In the Malangen fjord we observed temporal increases of DMSP production and standing stocks which resulted also in increases of DMSP vertical fluxes and DMS(P) concentrations at the sediment surface. This result illustrates how tight the coupling can be between pelagos and benthos, and confirms that DMS(P) concentration in the sediment was a reliable diagnostic indicator of vertical export from overlying waters in Malangen fjord. In Ullsfjord, however, DMS(P) concentrations in the sediment were poorly indicators of Phaeocystis pouchetii export during the early stage of growth of a bloom. The high load of DMS(P) in Balsfjord's sediments could neither be attributed to local vertical sedimentation nor to short-term lateral advection of fresh DMSP-containing phytoplanktonic material, and provides indication that this tracer sometimes also can be misleading. The highest loads of DMS(P) in sediments and the fastest rates of sedimentation occurred in the Southern Bight of the North Sea.     

Production and air–sea flux of halomethanes in the western subarctic Pacific in relation to phytoplankton pigment concentrations during the iron fertilization experiment (SEEDS II)

Iron could play a key role in controlling phytoplankton biomass and productivity in high-nutrient, low-chlorophyll regions. As a part of the iron fertilization experiment carried out in the western subarctic Pacific from July to August 2004 (Subarctic Pacific iron Experiment for Ecosystem Dynamics Study II—SEEDS II), we analysed the concentrations of trace gases in the seawater for 12 d following iron fertilization. The mean concentrations of chlorophyll a in the mixed layer (5–30 m depth) increased from 0.94 to 2.81 μg L^–1 for 8 d in the iron patch. The mean concentrations of methyl bromide (CH₃Br; 5–30 m depth) increased from 6.4 to 13.4 pmol L^–1 for 11 d; the in-patch concentration increased relative to the out-patch concentration. A linear correlation was observed between the concentrations of 19′-hexanoyloxyfucoxanthin, which is a biomarker of several prymnesiophytes, and CH₃Br in the seawater. After fertilization, the air–sea flux of CH₃Br inside the patch changed from influx to efflux from the ocean. There was no clear evidence for the increase in saturation anomaly of methyl chloride (CH₃Cl) due to iron fertilization. Furthermore, CH₃Cl fluxes did not show a tendency to increase after fertilization of the patch. In contrast to CH₃Br, no change was observed in the concentrations of bromoform (in-patch day 11 and out-patch day 11: 1.7 and 1.7 pmol L^–1), dibromomethane (2.1 and 2.2 pmol L^–1), and dibromochloromethane (1.0 and 1.2 pmol L^–1, respectively). The concentration of isoprene, which is known to have a relationship with chlorophyll a, did not change in this study. The responses of trace gases during SEEDS II differed from the previous findings (in situ iron enrichment experiment—EisenEx, Southern Ocean iron experiment—SOFeX, and Subarctic Ecosystem Response to Iron Enrichment Study—SERIES). Thus, in order to estimate the concomitant effect of iron fertilization on the climate, it is important to assess the induction of biological activity and the distributions/air–sea fluxes of trace gases by iron addition.     

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.     

Mesozooplankton grazing manipulations during in vitro iron enrichment studies in the NE subarctic Pacific

IronEx I demonstrated a rapid and marked response by grazers to Fe-induced increases in phytoplankton stocks, which was thought to be due, in part, to arrested vertical migration by mesozooplankton. These observations prompted an investigation of the relative roles of Fe enrichment and grazing pressure in controlling the magnitude of phytoplankton stocks in the NE subarctic Pacific. The grazing impact of increased mesozooplankton abundance in response to a localised Fe-induced enhancement of algal biomass was simulated by performing in vitro (6 d) grazer perturbation experiments in May 1994 and September 1995 at Ocean Station Papa (OSP), when pelagic mesozooplankton stocks are usually at their annual maximum and submaximal, respectively. Manipulations were designed to increase mesozooplankton stocks in 25L carboys after various lag-times corresponding to grazing pressure greater or equal to that in situ, and to monitor changes in chlorophyll a levels as a proxy for grazing pressure. At the onset of the experiments, in vitro mesozooplankton abundances were comparable to those in situ. Despite the addition of mesozooplankton to selected Fe-enriched carboys in May after 24, 48 and 72 h, corresponding to ca. two-fold increases in their abundances, chlorophyll a increased to ca. 2 μg l⁻¹ in all treatments. In September, chlorophyll a levels increased five-fold to 2 μg l⁻¹ after 4 days – but little thereafter – in the presence of up to ten-fold higher animal abundances (added at t=0) than observed in situ. Thus, Fe-induced increases in diatom growth rates were sufficiently high to escape both initial and additional grazing pressure. If and when Fe is supplied to this region, it is unlikely that mesozooplankton can respond and graze down the resulting elevated algal abundance. Theoretical calculations, based on algal growth and grazing rate data from May in this study, suggested that a greater than five-fold increase in mesozooplankton abundance, after a 48-h lag, is required to exert sufficient grazing pressure to prevent Fe-mediated increases in algal biomass. These findings are discussed in relation to the scale dependency of such events, and the pelagic ecology of other High Nitrate Low Chlorophyll regions.     

春秋季南黄海二甲基硫(DMS)与二甲巯基丙酸(DMSP)的浓度分布研究

Temporal distributions of dimethylsulfide(DMS) and dimethylsulfoniopropionate(DMSP) were studied in the southern Yellow Sea(SYS) during April and September 2010. The mean concentrations(range) of DMS, dissolved and particulate DMSP(DMSPd and DMSPp) in the surface waters in spring are 1.69(0.48–4.92), 3.18(0.68–6.75)and 15.81(2.82–52.33) nmol/L, respectively, and those in autumn are 2.80(1.33–5.10), 5.45(2.19–11.30) and 30.63(6.24–137.87) nmol/L. On the whole, the distributions of DMS and DMSP in spring are completely different from those in autumn. In the central part of the SYS, the concentrations of DMS and DMSP in spring are obviously higher than those in autumn, but the opposite situation is found on the south of 34°N, which can be attributed to the differences in nutrients and phytoplankton biomass and composition between spring and autumn. Besides,the seasonal variations of water column stability and the Changjiang diluted water also have significant impact on the distributions of DMS and DMSP in spring and autumn on the south of 34°N. DMS and DMSPp concentrations coincide well with chlorophyll a(Chl a) levels in the spring cruise, suggesting that phytoplankton biomass may play an important role in controlling the distributions of DMS and DMSPp in the study area. Annual DMS emission rates range from 0.015 to 0.033 Tg/a(calculated by S), respectively, using the equations of Liss and Merlivat(1986) and Wanninkhof(1992). This result implies a significant relative contribution of the SYS to the global oceanic DMS fluxes.     

Response of microzooplankton (protists and small copepods) to an iron-induced phytoplankton bloom in the Southern Ocean (EisenEx)

The dynamics, composition and grazing impact of microzooplankton were studied during the in situ iron fertilisation experiment EisenEx in the Antarctic Polar Frontal Zone in austral spring (November 2000). During the 21 day experiment, protozooplankton and small metazooplankton were sampled from the mixed layer inside and outside the patch using Niskin bottles. Aplastidic dinoflagellates increased threefold in abundance and biomass in the first 10 days of the experiment, but decreased thereafter to values twofold higher than pre-fertilisation values. The decline after day 10 is attributed to increasing grazing pressure by copepods. They also constrained ciliate abundances and biomass which were higher inside the fertilised patch than outside but highly variable. Copepod nauplii abundance remained stable whereas biomass doubled. Numbers of copepodites and adults of small copepod species (<1.5 mm) increased threefold inside the patch, but doubled in surrounding waters. Grazing rates estimated using the dilution method suggest that microzooplankton grazing constrained pico- and nanoplankton populations, but species capable of feeding on large diatoms (dinoflagellates and small copepods including possibly nauplii) were selectively predated by the metazoan community. Thus, iron fertilisation of a developing spring phytoplankton assemblage resulted in a trophic cascade which favoured dominance of the bloom by large diatoms.     

Impacts of the wintertime mesozooplankton community to downward carbon flux in the subarctic and subtropical Pacific Oceans

We compared wintertime depth distributions of the mesozooplankton community and dominant copepods between the subtropical (S1) and subarctic (K2) Pacific Oceans to evaluate the relative importance of actively transported carbon by vertical migrants to sinking particulate organic carbon flux. Primary production was higher and the ratio of sinking particulate organic carbon flux to primary production was lower at S1 compared with those at K2. The mesozooplankton community was lower in abundance and biomass at S1 compared to K2. Copepods were the dominant group among both mesozooplankton abundance and biomass throughout the water column down to 1000 m at both sites. The depth distribution showed that diel vertical migration was obvious for the mesozooplankton abundance and biomass at S1 but was not apparent for the abundance at K2, because the dominant component was diurnally migrating species at S1 and overwintering species residing at mesopelagic depths at K2. The major components of diel migrants were copepods and euphausiids at S1 and only euphausiids at K2. Respiratory flux by the diurnally migrating mesozooplankton was estimated to be 2 mgC m⁻² day⁻¹ at S1 and 7 mgC m⁻² day⁻¹ at K2. The respiratory flux was equivalent to 131% of sedimentary fecal pellet flux at S1 and 136% of that at K2. Because pathways of downward carbon flux are facilitated by the mesozooplankton community, the actively transported carbon (respiration of dissolved inorganic carbon, excretion of dissolved organic carbon and egestion of fecal pellets at depth) might be larger during winter than the flux of sinking fecal pellets.     

Spatial distributions of dimethylsulfide in the South China Sea

The concentration of dimethylsulfide (DMS) and supporting parameters were determined in surface seawater and vertical profiles at 26 stations in the South China Sea. The concentrations of DMS in surface seawater ranged from 61 to 148 ng S/l, with a mean of 82 ng S/l. High concentrations of DMS were found in the productive regions. The vertical profiles of DMS were characterized by a maximum at depths typically between 20 and 75 m. The concentrations of DMS were correlated with the levels of chlorophyll a both in the surface seawater and in the vertical distribution. The concentrations of DMS were higher than expected for this chlorophyll-poor tropical sea, as indicated by markedly high DMS (ng S/l)/chlorophyll a (μg/l) ratios ranging from 315 to 3524 with a mean of 1768 for all the surface seawater samples. DMS concentration was significantly correlated with seawater temperature and dissolved oxygen, but it showed an inverse relationship to nutrients (including nitrate, phosphate and silicate). On the basis of sea surface concentrations of DMS and gas exchange calculations, the mean flux of DMS from the South China Sea to the atmosphere was estimated to be 5.5 μmol m⁻² d⁻¹.     

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.     

Response of the larger protozooplankton to an iron-induced phytoplankton bloom in the Polar Frontal Zone of the Southern Ocean (EisenEx)

The responses of larger (>50 μm in diameter) protozooplankton groups to a phytoplankton bloom induced by in situ iron fertilization (EisenEx) in the Polar Frontal Zone (PFZ) of the Southern Ocean in austral spring are presented. During the 21 days of the experiment, samples were collected from seven discrete depths in the upper 150 m inside and outside the fertilized patch for the enumeration of acantharia, foraminifera, radiolaria, heliozoa, tintinnid ciliates and aplastidic thecate dinoflagellates. Inside the patch, acantharian numbers increased twofold, but only negligibly in surrounding waters. This finding is of major interest, since acantharia are suggested to be involved in the formation of barite (BaSO₄), a palaeoindicator of both ancient and modern high-productivity regimes. Foraminifera increased significantly in abundance inside and outside the fertilized patch. However, the marked increase of juveniles after a full-moon event suggests a lunar periodicity in the reproduction cycle of some foraminiferan species rather than a reproductive response to enhanced food availability. In contrast, adult radiolaria showed no clear trend during the experiment, but juveniles increased threefold, indicating elevated reproduction. Aplastidic thecate dinoflagellates almost doubled in numbers and biomass but also increased outside the patch. Tintinnid numbers decreased twofold, although biomass remained constant because of a shift in the size spectrum. Empty tintinnid loricae, however, increased by a factor of two, indicating that grazing pressure on this group mainly by copepods, intensified during EisenEx. The results show that iron-fertilization experiments can shed light on the biology and the role of these larger protists in pelagic ecosystem, which will improve their use as proxies in paleoceanography.