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.     

Iron and mixing affect biological carbon uptake in SOIREE and EisenEx,two Southern Ocean iron fertilisation experiments

This study explores the changes in the surface water fugacity of carbon dioxide (fCO₂) and biological carbon uptake in two Southern Ocean iron fertilisation experiments with different hydrographic regimes. The Southern Ocean Iron Release Experiment (SOIREE) experiment was carried out south of the Antarctic Polar Front (APF) at 61°S, 141°E in February 1999 in a stable hydrographic setting. The EisenEx experiment was conducted in a cyclonic eddy north of the APF at 48°S, 21°E in November 2000 and was characterised by a rapid succession of low to storm-force wind speeds and dynamic hydrographic conditions. The iron additions promoted algal blooms in both studies. They alleviated algal iron limitation during the 13-day SOIREE experiment and probably during the first 12 days of EisenEx. The fCO₂ in surface water decreased at a constant rate of 3.8 μatm day⁻¹ from 4 to 5 days onwards in SOIREE. The fCO₂ reduction was 35 μatm after 13 days. The evolution of surface water fCO₂ in the iron-enriched waters (or ‘patch’) displayed a saw tooth pattern in EisenEx, in response to algal carbon uptake in calm conditions and deep mixing and horizontal dispersion during storms. The maximum fCO₂ reduction was 18–20 μatm after 12 and 21 days with lower values in between. The iron-enriched waters in EisenEx absorbed four times more atmospheric CO₂ than in SOIREE between 5 and 12 days, as a result of stronger winds. The total biological uptake of inorganic carbon across the patch was 1389 ton C (±10%) in SOIREE and 1433 ton C (±27%) in EisenEx after 12 days (1 ton=10⁶ g). This similarity probably reflects the comparable size of the iron additions, as well as algal growth at a similar near-maximum growth rate in these regions. The findings imply that the different mixing regimes had less effect on the overall biological carbon uptake across the iron-enriched waters than suggested by the evolution of fCO₂ in surface water.     

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.     

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.     

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.     

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.     

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.     

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.     

Microzooplankton composition,biomass and grazing rates along the WOCE SR3 line between Tasmania and Antarctica

Microzooplankton species composition and grazing rates on phytoplankton were investigated along a transect between ∼46 and 67°S, and between 140 and 145°E. Experiments were conducted in summer between November 2nd and December 14th in 2001. The structure of the microbial food web changed considerably along the transect and was associated with marked differences in the physical and chemical environment encountered in the different water masses and frontal regions. On average microzooplankton grazing experiments indicated that 91%, 102%, and 157%, (see results) of the phytoplankton production would be grazed in the <200, <20 and <2 μm size fractions, respectively, indicating microzooplankton grazing was potentially constraining phytoplankton populations (<200 μm) along most of the transect. Small ciliates in general and especially oligotrich species declined in importance from the relatively warm, Southern Subtropical Front waters (6.8 μg C/L) to the colder waters of the southern branch of the Polar Front (S-PF), (∼0.5 μg C/L) before increasing again near the Antarctic landmass. Large changes in microzooplankton dominance were observed, with heterotrophic nanoflagellates (HNF), ciliates and larger dinoflagellates having significant biomass in different water masses. HNF were the dominant grazers when chlorophyll a was low in areas such as the Inter-Polar Frontal Zone (IPFZ), while in areas of elevated biomass such as the S-PF and Southern Antarctic Circumpolar Current (SACC), a mix of copepod nauplii and large heterotrophic and mixotrophic dinoflagellates tended to dominate the grazing community. In the S-PF and SACC water masses the tight coupling observed between the microzooplankton grazers and phytoplankton populations over most of the rest of the transect was relaxed. In these regions grazing was low on the >20 μm size fraction of chlorophyll a, which dominated the biomass, while smaller diatoms and nanoplankton in the <20 μm size fraction were still heavily grazed. The lack of grazing pressure on large phytoplankton contributes to this region's potential to export carbon with larger cells known to have higher sinking rates.     

Physical behaviors of the iron-fertilized patch in SEEDS II

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

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

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

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.     

Phytoplankton of the western Arctic in the spring and summer of 2002: Structure and seasonal changes

We analyzed the taxonomic structure and spatial variability of phytoplankton abundance and biomass in the Chukchi and Beaufort Seas during spring and summer seasons of the SBI program. Phytoplankton samples were collected during two surveys from May 10 to June 13 and from July 19 to August 21 of 2002. In May and June, ice cover exceeded 80% over most of the study area and there was no vertical stratification, indicating that the successional state of the phytoplankton corresponded to the end of the winter biological season. The phytoplankton abundance ranged from a few tens to a few thousands of cells per liter, while biomass varied from 0.1 to 3.0 mg C m⁻³. Small areas of high phytoplankton abundance (0.13–1.3×10⁶ cells L⁻¹) and biomass (22–536 mg C m⁻³), dominated by early spring diatoms Pauliella taeniata and Fragilariopsis oceanica in the surface waters, which indicated the beginning of the spring bloom, were observed only in the southeastern part of the Chukchi shelf and off Point Barrow. In July and August summer period, more than a half of the study area had <50% ice cover and the water column was stratified by temperature and salinity. Over the Chukchi shelf and continental slope of the Beaufort Sea, the phytoplankton abundance and biomass were an order of magnitude higher in July–August than in May–June. The taxonomic diversity of algae also increased due to the appearance of late-spring and summer diatoms, dinoflagellates, and coccolithophorids (Emiliania huxleyi). Interestingly, the seasonal differences between phytoplankton abundance and taxonomic composition in the spring and summer periods varied the least over the Chukchi Sea slope and in the deep-water area of the Arctic Ocean. High algae concentrations in summer were located in the lower layers of the euphotic zone, suggesting that the spring bloom on both the Chukchi shelf and in the western part of the Beaufort Sea occurred in late June/early July. In the spring and summer, the microalgal community was characterized by a high abundance of 4–10 μm flagellates, which exceeded the abundance of all other taxonomic groups. In both seasons studied, phytoplankton reached its maximum abundance within restricted areas in the southern part of the Chukchi Sea southwest of Point Hope, in the northern part of the Chukchi shelf between the 50- and 100-m isobaths, on the shelf northwest of Point Barrow, and over the continental slope in the Beaufort Sea. The pronounced spatial difference in the seasonal state was a characteristic feature of the phytoplankton community in the western Arctic.     

Population structure and life history of Hemimysis lamornae mediterranea (Malacostraca: Mysida) in the Ebro Delta (NW Mediterranean)

Hemimysis lamornae mediterranea Bacescu, 1936 has been recently reported in the Ebro Delta (Spain, NW Mediterranean). Little is known about the biology and ecology of this mysid and we provide the first information about its population biology. H. l. mediterranea were collected from Sant Carles de la Ràpita harbor from June 2010 to March 2012 at night. The H. l. mediterranea population was composed of two main individual size categories: larger-sized winter/spring individuals and smaller-sized spring/summer individuals. The overall sex ratio is highly skewed; mature females are 2.5 times more abundant than mature males. Reproductive activity was higher during late winter and spring but was almost continuous throughout the year. The intra-marsupial development and growth of juveniles has been studied in the laboratory. The mean duration of incubation period (intra-marsupial stages) in laboratory conditions was 11 days (20 °C) and the age at first maturity ranged from14 to 20 days. The growth rate was faster in early juveniles and declined with age, showing a maximum of 0.152 mm d^− 1. The laboratory results and demographic data suggest that H. l. mediterranea will produce several generations per year in the Ebro Delta. H. l. mediterranea was characterized by a combination of early maturation of individuals (short juvenile period), rapid growth, small adult size, a continuous reproduction all year round, iteroparous females, a relatively high fecundity and a high number of generations per year.     

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.     

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.     

Depth-related distribution and abundance of seastars (Echinodermata: Asteroidea) in the Porcupine Seabight and Porcupine Abyssal Plain,N.E. Atlantic

The depth-related distribution of seastar (Echinodermata: Asteroidea) species between 150 and 4950 m in the Porcupine Seabight and Porcupine Abyssal Plain is described. 47 species of asteroid were identified from ∼14,000 individuals collected. The bathymetric range of each species is recorded. What are considered quantitative data, from an acoustically monitored epibenthic sledge and supplementary data from otter trawls, are used to display the relative abundance of individuals within their bathymetric range. Asteroid species are found to have very narrow centres of distribution in which they are abundant, despite much wider total adult depth ranges. Centres of distribution may be skewed. This might result from competition for resources or be related to the occurrence of favourable habitats at particular depths. The bathymetric distributions of the juveniles of some species extend outside the adult depth ranges. There is a distinct pattern of zonation with two major regions of faunal change and six distinct zones. An upper slope zone ranges from 150 to ∼700 m depth, an upper bathyal zone between 700 and 1100 m, a mid-bathyal zone from 1100 to1700 m and a lower bathyal zone between 1700 and 2500 m. Below 2500 m the lower continental slope and continental rise have a characteristic asteroid fauna. The abyssal zone starts at about 2800 m. Regions of major faunal change are identified at the boundaries of both upper and mid-bathyal zones and at the transition of bathyal to abyssal fauna. Diversity is greatest at ∼1800 m, decreasing with depth to ∼2600 m before increasing again to high levels at ∼4700 m.     

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

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