Cellular iron contents of plankton during the Southern Ocean Iron Experiment (SOFeX)

Iron (Fe) availability limits phytoplankton biomass and production in large regions of the Southern Ocean and influences community composition and size structure, which may affect C export and other system-level functions. To improve our understanding of Fe partitioning within communities and the response of different components to fertilization, we assessed the cellular Fe contents of individual diatoms, autotrophic flagellates, and heterotrophic flagellates during the recent Southern Ocean Fe Experiment using synchrotron-based X-ray fluorescence (SXRF). Dual ⁵⁵Fe/¹⁴C radioisotope incubations were also conducted to estimate Fe:C ratios in size-fractionated plankton. Cellular Fe quotas determined by the two techniques were in close agreement when low amounts of ⁵⁵Fe (0.2 nM) were added, but ⁵⁵Fe additions of 2 nM resulted in 2–3-fold higher quotas. SXRF assessments of cellular Fe quotas (normalized to C) were generally in good agreement with prior bulk analyses of natural assemblages, but revealed compositional differences among protistan taxa not previously detected. Mean Fe:C ratios for diatoms, autotrophic flagellates, and heterotrophic flagellates from unfertilized waters were 6.0, 8.7, and 14.1 μmol mol C⁻¹, respectively. Smaller cells had higher Fe:C ratios than larger cells. Fertilization enhanced Fe quotas in all cell types, with mean Fe:C ratios increasing approximately 4-fold (from about 10 to about 40 μmol mol C⁻¹) after two Fe additions. This study provides some of the first measurements of Fe quotas in phytoplankton cells from natural communities and the first measurements of Fe quotas in natural protozoa.     

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.     

Phytoplankton and iron limitation of photosynthetic efficiency in the Southern Ocean during late summer

As part of two USJGOFS cruises, we investigated spatial variability in phytoplankton properties across the strong environmental gradient associated with the Antarctic Polar Frontal Zone during late austral summers of 1997 and 1998. Cell properties, including size and an index of pigment content as well as photosynthetic efficiency (as indicated by relative variable fluorescence), changed dramatically across this frontal region. A general trend toward reduced photosynthetic efficiency south of the Polar Front was correlated with low dissolved iron concentration and is consistent with physiological iron limitation in the phytoplankton. We detected no significant differences in photosynthetic efficiency among different size classes of the dominant pico- to nanophytoplankton, despite a systematic community level shift toward larger sized cells south of the Polar Front. In contrast to other cells, those classified as cryptophyte algae showed relatively high photosynthetic efficiency in low iron waters; however, this group was never found in high abundance. One group, all cells ⩽2 μm, showed an unexpected increase in intracellular pigment content (based on single cell chlorophyll fluorescence measurements) south of the Polar Front where dissolved iron concentration and the cells’ relative abundance were low. Overall, these results suggest that group- or size-specific differences in physiological status were not directly regulating community structure in the pico- to nanophytoplankton during the late summer season; other processes, such as differential grazing or sinking losses, must be important.     

Matching carbon pools and fluxes for the Southern Ocean Iron Release Experiment (SOIREE)

The Lagrangian Southern Ocean Iron Release Experiment (SOIREE) allowed study of a gradually evolving iron-mediated phytoplankton bloom in water labelled with the inert tracer sulfur hexafluoride, SF₆. This article describes a pelagic carbon budget for the mixed layer in SOIREE and assesses the extent to which closure of the budget is achieved. Net community production (NCP) converted 837 mmol m⁻² of inorganic carbon to organic carbon in 12.0 d after the first iron addition. A large fraction (41%) of NCP remained as particulate organic carbon in the mixed layer of the iron-enriched patch, while 23% was lost by horizontal dispersion and 0–29% was exported. The closure of the carbon budget is hampered by the lack of measurements of dissolved organic carbon (DOC), by a major uncertainty in carbon export, and by use of empirical conversion factors in estimates of carbon biomass and metabolic rates. Lagrangian carbon-budget studies may be improved by direct measurement of all major carbon parameters and conversion factors. Carbon cycling in the SOIREE bloom resembled that in ‘natural’ algal blooms in the open Southern Ocean in some respects, but not in all. Daily NCP in the SOIREE bloom (70 mmol m⁻² d⁻¹) was higher than in natural blooms, partly because other studies did not account for horizontal dispersion, were for longer periods or included less productive areas. The build-up of POC stock and carbon export as a fraction of NCP in SOIREE were in the lower range of observations elsewhere.     

Organic complexation of iron in the Southern Ocean

The chemical speciation of iron was determined in the Southern Ocean along a transect from 48 to 70°S at 20°E. Dissolved iron concentrations were low at 0.1–0.6 nM, with average concentrations of 0.25±0.13 nM. Organic iron complexing ligands were found to occur in excess of the dissolved iron concentration at 0.72±0.23 nM (equivalent to an excess of 0.5 nM), with a complex stability of log K_FeL′=22.1±0.5 (on the basis of Fe³⁺ and L′). Ligand concentrations were higher in the upper water column (top 200 m) suggesting in situ production by microorganisms, and less at the surface consistent with photochemical breakdown. Our data are consistent with the presence of stable organic iron-complexing ligands in deep global ocean waters at a background level of ∼0.7 nM. It has been suggested that this might help stabilise iron at levels of ∼0.7 nM in deep ocean waters. However, much lower iron concentrations in the waters of the Southern Ocean suggest that these ligands do not prevent the removal of iron (by scavenging or biological uptake) to well below the concentration of these ligands. Scavenging reactions are probably inhibited by such ligand competition, so it is likely that biological uptake is the chief cause for the further removal of iron to these low levels in waters that suffer from very low iron inputs.     

Mechanisms determining species dominance in a phytoplankton bloom induced by the iron fertilization experiment EisenEx in the Southern Ocean

The dynamics of phytoplankton species populations recorded during the 3-week, iron-fertilization experiment EisenEx carried out in spring in the Antarctic Polar Frontal Zone are presented and discussed as the difference between growth and mortality rates. Only two cosmopolitan diatom species, the centric Chaetoceros debilis and the pennate Pseudo-nitzschia lineola, increased population density exponentially throughout the experiment to 150- and 90-fold of initial values, respectively. Because C. debilis initial abundance was tenfold lower than that of P. lineola, the two contributed 1% and 21% to bloom biomass, respectively at the end of the experiment, high-lighting the role of seeding in bloom formation. The other significant species increased population size at a linear rate throughout the experiment or for a short spurt phase to 3- to 18-fold of initial values. Conservative estimates of mortality rates within diatom species populations were obtained by comparing net accumulation rates of full cells with those of empty and broken frustules. The ratios were consistent over time for the various species but varied widely between them. The species-specific variation can be explained by differences in both growth and mortality rates, the latter partly due to either selective grazing or avoidance by the large protozoo- and metazooplankton populations present. Selective predation by the abundant copepod populations on protistan grazers of diatoms (ciliates and heterotrophic dinoflagellates) apparently aided diatom biomass build-up. The response patterns of populations of the phytoplankton species present fall into six categories comprising disparate species, indicating that phylogeny is a poor predictor of ecology. The group that did not respond to fertilization was the most diverse and included both endemic and cosmopolitan as well as background and bloom-forming species. This lack of response to the advent of favorable growth conditions indicates that proximate factors during EisenEx triggered growth only in some species but had little effect on others. We attribute the differences in behavior to ultimate factors such as seasonal effects on life cycles and other internal constraints on growth rates. The implications for our understanding of the evolutionary ecology of phytoplankton and its impact on global biogeochemical cycles are pointed out.     

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.     

Dissolved iron in the Australian sector of the Southern Ocean (CLIVAR SR3 section): Meridional and seasonal trends

We report measurements of dissolved iron (dFe, <0.4 μm) in seawater collected from the upper 300 m of the water column along the CLIVAR SR3 section south of Tasmania in March 1998 (between 42°S and 54°S) and November–December 2001 (between 47°S and 66°S). Results from both cruises indicate a general north-to-south decrease in mixed-layer dFe concentrations, from values as high as 0.76 nM in the Subtropical Front to uniformly low concentrations (<0.1 nM) between the Polar Front and the Antarctic continental shelf. Samples collected from the seasonal sea-ice zone in November–December 2001 provide no evidence of significant dFe inputs from the melting pack ice, which may explain the absence of pronounced ice-edge algal blooms in this sector of the Southern Ocean, as implied by satellite ocean-color images. Our data also allow us to infer changes in the dFe concentration of surface waters during the growing season. South of the Polar Front, a comparison of near-surface with subsurface (150 m depth) dFe concentrations in November–December 2001 suggests a net seasonal biological uptake of at least 0.14–0.18 nM dFe, of which 0.05–0.12 nM is depleted early in the growing season (before mid December). A comparison of our spring 2001 and fall 1998 data indicates a barely discernible seasonal depletion of dFe (0.03 nM) within the Polar Frontal Zone. Further north, most of our iron profiles do not exhibit near-surface depletions, and mixed-layer dFe concentrations are sometimes higher in samples from fall 1998 compared to spring 2001; here, the near-surface dFe distributions appear to be dominated by time-varying inputs of aerosol iron or advection of iron-rich subtropical waters from the north.     

Methane Distribution in Antarctic Sound (Southern Ocean)

Oceanology - The paper presents the first data on the methane distribution in the water column of Antarctic Sound (Atlantic sector of the Southern Ocean) acquired during an integrated expedition of...     

The role of iron sources and transport for Southern Ocean productivity

Iron has been found to limit primary productivity in high nutrient, low chlorophyll regions of the oceans, including the Southern Ocean. Here we assess the relative magnitudes and geographical distributions of the sources of iron (sedimentary, atmospheric, icebergs and sea ice) to the Southern Ocean, and their impact on productivity. We present an iron cycling model, based on the assumptions of iron and light limitation of primary production, which is embedded in an eddy resolving ocean general circulation model. We find that the injection depth of the various iron inputs determines their availability for driving production because dissolved iron may be scavenged prior to it entering the illuminated mixed layer where it can drive primary production. The model suggests that production is predominantly regulated by sediment-derived iron sources rather than icebergs, sea ice or atmospheric dust. We note non-linear response in productivity to changes in the strength of one or more iron sources due to scavenging. Sea ice influences productivity by modifying the timing of iron supply to the euphotic zone. We also show that in the Scotia Sea the majority of productivity is driven by sediment-sourced iron from the Antarctic Peninsula, with additional local hotspots driven by island sources.     

Phytoplankton and ice-algal communities in the seasonal ice zone during January (Southern Ocean,Indian sector)

Journal of Oceanography - In the seasonal ice zone (SIZ), sea-ice algae have been hypothesized to influence phytoplankton species composition in seawater after melting from sea ice. However,...     

Vertical distributions of dissolved organic carbon and nitrogen in the Southern Ocean

Detailed vertical profiles of dissolved organic carbon (DOC), nitrogen (DON) and DOC/DON ratios in the Australian sector of the Southern Ocean (56°–65°S around 140°W transect) were obtained by a high-temperature catalytic oxidation method with a modified Shimadzu TOC-5000 unit. The simultaneous analyses of DOC and total dissolved nitrogen (TDN) gave high precision results (±0.7 μMC and ±0.3 μMN). In surface layers, DOC concentrations were lower (45–55 μM) than those generally obtained from other oceanic environments (60–90 μM). The surface concentrations of DON varied from 4 to 9 μM, but within the range generally reported for other ocean regions. Surface excesses of DOC and DON were calculated against nearly constant subsurface concentrations. A consistent contribution of low C/N ratio (2.7–5.0) was found in the mid-surface layer (30–75 m), suggesting more extensive degradation of carbon-enriched materials and/or enhanced supply of nitrogen-enriched ones.     

Iron and manganese in the wake of the Kerguelen Islands (Southern Ocean)

As part of the ANTARES 3/F-JGOFS cruise, the distributions of dissolved iron and manganese were measured in October 1995 in the north–east wake of the Kerguelen archipelago (48°40′–49°40′S, 68°70′–70°50′E), an area that shows high phytoplankton biomass (CZCS and SeaWiFS data) in the middle of the High Nutrient Low Chlorophyll (HNLC) Southern Ocean. The study area (about 25,000 km²) comprised a branch of the Polar Front with Antarctic surface water (AASW) intruding northward, shouldering the shelf break of the Kerguelen Plateau. The coastal zone was clearly affected by material of lithogenic origin (riverine discharges, soil leaching by rain waters, aeolian inputs), as well as by inputs from the sediments (effluxes from the sediment–water interface, resuspension from the sediments), its near surface waters showing considerable enrichment in dissolved iron (5.3–12.6 nM) and in dissolved manganese (2.9–8.6 nM). The offshore waters, although less enriched in trace-metals, were also affected by trace-metal inputs from coastal and continental shelf origin. Dissolved iron and manganese concentrations in these waters were 0.46–0.71 and 0.68–1.3 nM, i.e. far over typical antarctic open ocean surface water concentrations of 0.16 nM for iron [Martin, J.H., Gordon, R.M., Fitzwater, S.E., 1990. Iron in Antarctic waters. Nature, 345: 156–158.] and around 0.1 nM for manganese [Martin, J.H., Gordon, R.M., Fitzwater, S.E., 1990. Iron in Antarctic waters. Nature, 345: 156–158; Sedwick, P.N., Edwards, P.R., Mackey, D.J., Griffiths, F.B., Parslow, J.S., 1997. Iron and manganese in surface waters of the Australian subantarctic region. Deep-Sea Res., 44: 1239–1253.]. The dissolved iron enrichment in coastal waters of the Kerguelen Islands is much more important (about 10 times for dissolved iron) than for the Galapagos Islands, another oasis in the HNLC Equatorial oceanic system, where the concentration increase in dissolved iron in the surface waters around the islands is mostly driven by upwelling of the Equatorial Under Current (EUC) as it reaches the Galapagos Platform.     

Ocean current estimation using a Multi-Model Ensemble Kalman Filter during the Grand Lagrangian Deployment experiment (GLAD)

In the summer and fall of 2012, during the GLAD experiment in the Gulf of Mexico, the Consortium for Advanced Research on Transport of Hydrocarbon in the Environment (CARTHE) used several ocean models to assist the deployment of more than 300 surface drifters. The Navy Coastal Ocean Model (NCOM) at 1 km and 3 km resolutions, the US Navy operational NCOM at 3 km resolution (AMSEAS), and two versions of the Hybrid Coordinates Ocean Model (HYCOM) set at 4 km were running daily and delivering 72-h range forecasts. They all assimilated remote sensing and local profile data but they were not assimilating the drifter’s observations. This work presents a non-intrusive methodology named Multi-Model Ensemble Kalman Filter that allows assimilating the local drifter data into such a set of models, to produce improved ocean currents forecasts. The filter is to be used when several modeling systems or ensembles are available and/or observations are not entirely handled by the operational data assimilation process. It allows using generic in situ measurements over short time windows to improve the predictability of local ocean dynamics and associated high-resolution parameters of interest for which a forward model exists (e.g. oil spill plumes). Results can be used for operational applications or to derive enhanced background fields for other data assimilation systems, thus providing an expedite method to non-intrusively assimilate local observations of variables with complex operators. Results for the GLAD experiment show the method can improve water velocity predictions along the observed drifter trajectories, hence enhancing the skills of the models to predict individual trajectories.     

The influence of UV irradiation on the photoreduction of iron in the Southern Ocean

An iron enrichment experiment, EisenEx, was performed in the Atlantic sector of the Southern Ocean during the Antarctic spring of 2000. Deck incubations of open ocean water were performed to investigate the influence of ultraviolet B (UVB: 280–315 nm) and ultraviolet A (UVA: 315–400 nm) on the speciation of iron in seawater, using an addition of the radioisotopes ⁵⁹Fe(III) (1.25 nM) or ⁵⁵Fe(III) (0.5 nM). Seawater was sampled inside and outside the iron-enriched region. The radioisotopic Fe(II) concentration was monitored during daylight under three different light conditions: the full solar spectrum (total), total minus UVB, and total minus UVB+UVA. A distinct diel cycle was observed with a clear distinction between the three different light regimes. A clear linear relationship was found for the concentration of radioisotopic Fe(II) versus irradiance. UVB produced most of the Fe(II) followed by UVA and visible light (VIS: 400–700 nm), respectively. UVB produced 4.89 and 0.69 pM m² W⁻¹ radioisotopic Fe(II) followed by UVA with 0.33 and 0.10 pM m² W⁻¹ radioisotopic Fe(II) and VIS with 0.04 and 0.03 pM m² W⁻¹ radioisotopic Fe(II).     

Annually recurrent phytoplanktonic assemblages during summer in the seasonal ice zone west of the Antarctic Peninsula (Southern Ocean)

The distribution of phytoplankton composition, cell abundance and biomass from an area along the Western Antarctic Peninsula was studied during three summers, with the aim of understanding its dynamics over spatial and interannual scales. The studied area is characterized by seasonal sea-ice retreat and advance. Algae composition and concentration were found to be highly variable through the area as well as from year to year. Small unidentified phytoflagellates, diatoms and cryptophytes were the main phytoplankton groups, contributing the major proportion of total phytoplankton cell abundance and biomass concentration. Three annually recurrent phytoplankton assemblages were recognized in the area according to the algae composition and abundance: a diatom bloom associated with the sea-ice edge, an assemblage dominated by small unidentified phytoflagellates and cryptophytes, and a diatom-enriched assemblage in open waters. The distribution of these assemblages varied from year-to-year. During the summers preceded by early sea-ice retreat, the diatom bloom was spatially restricted and the other two assemblages occupied extended regions, whereas during the late sea-ice retreat year, the diatom bloom extended over a larger region and the other assemblages occupied smaller regions or were just absent. It was detected that these assemblages resemble different stages of the phytoplankton seasonal cycle, and that their distribution through the area can be related to a latitudinal and longitudinal gradient in the phytoplankton growth onset timing, associated with the progressive sea-ice retreat during spring. The local environmental conditions associated with each assemblage were also analyzed, but further study is needed for understanding the causes of the replacement of one assemblage by another through the area. On the other hand, the interannual variability in the distribution of the assemblages can be related to year-to-year differences in the timing of phytoplankton growth onset, associated with variations in the timing of the sea-ice retreat.     

Bacterioplankton responses to iron enrichment during the SAGE experiment

We studied the microbial food web in the upper 100 m of the water column in iron-limited sub-Antarctic HNLC waters south-east of New Zealand in the SAGE experiment in 2004, with focus on bacterioplankton. Samples were collected daily from inside and outside the iron enriched patch. Short term enrichment experiments were conducted on board in 4 L polycarbonate bottles with water outside the iron enriched patch to study single and combined effects of micronutrient additions on microbial food web. Low bacterial growth was recorded in the study area with community turnover times of 50 h or more during the study period. Measurements of bacterial standing stocks and production rates in the study show minor responses to the large scale iron enrichment, with increase in rates and stocks after the first enrichment and at the end of the study period after the third iron enrichment when solar radiation increased and wind mixing decreased. The average daily bacterial production rates were 31.5 and 33.7 mgCm⁻² d⁻¹ for the OUT and IN stations, respectively; thus overall there was not a significant difference between the control and the iron-enriched patch. In the bottle experiments bacterial thymidine incorporation showed responses to single iron and silicic acid enrichments and a major growth response to the combined iron and sucrose enrichments. Phytoplankton chlorophyll-a showed clear stimulation by single additions of iron and silicic acid and silicic acid enhanced the iron impact. Cobalt additions had no effect on bacteria growth and a negative effect on phytoplankton growth. Low bacterial in situ growth rates and the enrichment experiments suggest that bacteria are co-limited by iron and carbon, and that bacterial iron uptake is dependent on carbon supply by the food web. With the high iron quota (??mol Fe mol C⁻¹) bacteria may scavenge considerable amounts of the excess iron, and thus influence the relative importance of the microbial food web as a carbon sink.     

Responses of pico- and nanophytoplankton to artificial iron infusions observed during the second iron enrichment experiment in the western subarctic Pacific (SEEDS II)

How pico- and nanophytoplankton responded to artificial iron infusions was investigated using flow cytometry during SEEDS II, the second mesoscale in situ iron enrichment experiment in the western subarctic North Pacific. Two iron infusions on days 0 and 7 caused a remarkable increase in cellular chlorophyll fluorescence and cell size of all the four phytoplankton groups investigated: Synechococcus, cryptophytes, picoeucaryotes and nanoeucaryotes other than cryptophytes. After the second infusion, the abundance of three phytoplankton groups, excluding Synechococcus, also started to increase. After surface dissolved iron concentration decreased to <0.2 nM on day 11, chlorophyll fluorescence of all the four groups returned to a level observed before the iron infusions, suggesting that pico- and nanophytoplankton were physiologically stressed by iron deficiency. Cell concentrations of pico- and nanoeucaryotes decreased to the pre-infusion level by day 23, while that of cryptophytes remained high until day 25. Flow cytometric diagnosis showed that cryptophytes were physiologically limited during this period and effective iron uptake from suspended particles, as reported for freshwater cryptophytes, was not observed. Thus their prosperity may have been due to alleviation from grazing. Cell concentration of Synechococcus started to increase at a net specific growth rate of 0.13 d⁻¹ after day 12, reaching more than 6 times that of the pre-infusion level on day 24. This may have been due to the elevation of surface water temperature observed during the survey period, together with trophic cascading effects of increased copepod grazing.