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Krill and salps are important for carbon flux in the Southern Ocean, but the extent of their contribution and the consequences of shifts in dominance from krill to salps remain unclear. We present a direct comparison of the contribution of krill and salp faecal pellets (FP) to vertical carbon flux at the Antarctic Peninsula using a combination of sediment traps, FP production, carbon content, microbial degradation, and krill and salp abundances. Salps produce 4-fold more FP carbon than krill, but the FP from both species contribute equally to the carbon flux at 300 m, accounting for 75% of total carbon. Krill FP are exported to 72% to 300 m, while 80% of salp FP are retained in the mixed layer due to fragmentation. Thus, declining krill abundances could lead to decreased carbon flux, indicating that the Antarctic Peninsula could become a less efficient carbon sink for anthropogenic CO 2 in future.
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Although zooplankton are the primary energy pathway from phytoplankton to fish, we understand little about how climate change will modify zooplankton communities and their role in marine ecosystems. Using a trait-based marine ecosystem model resolving key zooplankton groups, we assess climate change impacts on zooplankton community composition and implications for marine food webs globally. We find that future oceans favour food webs increasingly dominated by carnivorous (chaetognaths, jellyfish and carnivorous copepods) and gelatinous filter-feeding zooplankton (larvaceans and salps). By providing a direct energetic pathway from small phytoplankton to fish, the rise of gelatinous filter-feeders largely offsets the increase in trophic steps between primary producers and fish from declining phytoplankton production and increasing carnivorous zooplankton. However, our results indicate that future fish communities face not only reduced carrying capacity from falling primary production, but also lower quality diets as environmental conditions increasingly favour gelatinous zooplankton.
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The impact of marine animals on the iron (Fe) cycle has mostly been considered in terms of their role in supplying dissolved Fe to phytoplankton at the ocean surface. However, little attention has been paid to how the transformation of ingested food into fecal matter by animals alters the relative Fe-richness of particles, which could have consequences for Fe cycling in the water column and for the food quality of suspended and sinking particles. Here, we compile observations to show that the Fe to carbon (C) ratio (Fe:C) of fecal pellets of various marine animals is consistently enriched compared to their food, often by more than an order of magnitude. We explain this consistent enrichment by the low assimilation rates that have been measured for Fe in animals, together with the respiratory conversion of dietary organic C to excreted dissolved inorganic C. Furthermore, we calculate that this enrichment should cause animal fecal matter to constitute a major fraction of the global sinking flux of biogenic Fe, a component of the marine iron cycle that has been previously unappreciated. We also estimate that this fecal iron pump provides an important source of Fe to marine animals via coprophagy, particularly in the mesopelagic, given that fecal matter Fe:C can be many-fold higher than the Fe:C of local phy-toplankton. Our results imply that the fecal iron pump is important both for global Fe cycling and for the iron nutrition of pelagic and mesopelagic communities.
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Fe is a critical nutrient to the marine biological pump, which is the process that exports photosynthetically fixed carbon in the upper ocean to the deep ocean. Fe limitation controls...
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Phytoplankton growth in large parts of the world ocean is limited by low availability of dissolved iron (dFe), restricting oceanic uptake of atmospheric CO2. The bioavailability of dFe in seawater is however difficult to appraise since it is bound by a variety of poorly characterized organic ligands. Here, we propose a new approach for evaluating seawater dFe bioavailability based on its uptake rate constant by Fe-limited cultured phytoplankton. We utilized seven phytoplankton species of diverse classes, sizes, and provenances to probe for dFe bioavailability in 12 seawater samples from several ocean basins and depths. All tested phytoplankton acquired organically bound Fe in any given sample at similar rates (after normalizing to cellular surface area), confirming that multiple, Fe-limited phytoplankton species can be used to probe dFe bioavailability in seawater. These phytoplankton-based uptake rate constants allowed us to compare water types, and obtain a grand average estimate of seawater dFe bioavailability. Among water types, dFe bioavailability varied by approximately four-fold, and did not clearly correlate with Fe concentrations or any of the measured Fe speciation parameters. Compared with well-studied Fe complexes, seawater dFe is more available than model siderophore Fe, but less available than inorganic Fe. Exposure of seawater to sunlight, however, significantly enhanced dFe bioavailability. The rate constants established in this work, not only facilitate comparison between water types, but also allow calculation of Fe uptake rates by phytoplankton in the ocean based on measured dFe concentrations. The approach established and verified in this study, opens a new way for determining dFe bioavailability in samples across the ocean, and enables modeling of in situ Fe uptake rates by phytoplankton using dFe concentrations from GEOTRACES datasets.
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Iron (Fe) is a paradox in the modern ocean – it is central to many life-critical enzymes but is scarce across most surface waters. The high cellular demand and low bioavailability of Fe likely puts selective pressure on marine microorganisms. Previous observations suggest that heterotrophic bacteria are outcompeted by small diatoms for Fe supply in the subantarctic zone of Southern Ocean, thereby challenging the idea of heterotrophic bacteria being more competitive than phytoplankton in the access to this trace metal. To test this hypothesis, incubation experiments were carried out at the Southern Ocean Time Series site (March–April 2016). We investigated (a) whether dissolved organic carbon (DOC), dissolved Fe, or both limit the growth of heterotrophic bacteria and, (b) if the presence of potential competitors has consequences on the bacterial Fe acquisition. We observed a pronounced increase in both bulk and cell-specific bacterial production in response to single (+C) and combined (+Fe+C) additions, but no changes in these rates when only Fe was added (+Fe). Moreover, we found that +Fe+C additions promoted increases in cell-specific bacterial Fe uptake rates, and these increases were particularly pronounced (by 13-fold) when phytoplankton were excluded from the incubations. These results suggest that auto- and heterotrophs could compete for Fe when DOC limitation of bacterial growth is alleviated. Such interactions between primary producers and nutrient-recyclers are unexpected drivers for the duration and magnitude of phytoplankton blooms in the Southern Ocean.
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Iron is an essential micronutrient that limits primary production in up to 40% of the surface ocean and influences carbon dioxide uptake and climate change. Dissolved iron is mostly associated with loosely characterised organic molecules, called ligands, which define key aspects of the iron cycle such as its residence time, distribution and bioavailability to plankton. Models based on in situ ligand distributions and the behaviour of purified compounds include long-lived ligands in the deep ocean, bioreactive ligands in the surface ocean and photochemical processes as important components of the iron cycle. Herein, we further characterise biologically refractory ligands in dissolved organic matter (DOM) from the deep ocean and labile ligands in DOM from the surface ocean, and their photochemical and biological reactivities. Experimental results indicated that photodegradation of upwelled refractory iron-binding ligands can fuel iron remineralisation and its association with labile organic ligands, thus enhancing iron bioavailability in surface waters. These observations better elucidate the roles of biologically refractory and labile molecules and global overturning circulation in the ocean iron cycle, with implications for the initiation and sustainment of biological activity in iron-limited regions and the residence time of iron in the ocean.
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New estimates of pCO2 from profiling floats deployed by the Southern Ocean Carbon and Climate Observations and Modeling (SOCCOM) project have demonstrated the importance of wintertime outgassing south of the Polar Front, challenging the accepted magnitude of Southern Ocean carbon uptake (Gray et al., 2018, https://doi:10.1029/2018GL078013). Here, we put 3.5 years of SOCCOM observations into broader context with the global surface carbon dioxide database (Surface Ocean CO2 Atlas, SOCAT) by using the two interpolation methods currently used to assess the ocean models in the Global Carbon Budget (Le Quéré et al., 2018, https://doi:10.5194/essd-10-2141-2018) to create a ship-only, a float-weighted, and a combined estimate of Southern Ocean carbon fluxes (<35°S). In our ship-only estimate, we calculate a mean uptake of -1.14 ± 0.19 Pg C/yr for 2015-2017, consistent with prior studies. The float-weighted estimate yields a significantly lower Southern Ocean uptake of -0.35 ± 0.19 Pg C/yr. Subsampling of high-resolution ocean biogeochemical process models indicates that some of the differences between float and ship-only estimates of the Southern Ocean carbon flux can be explained by spatial and temporal sampling differences. The combined ship and float estimate minimizes the root-mean-square pCO2 difference between the mapped product and both data sets, giving a new Southern Ocean uptake of -0.75 ± 0.22 Pg C/yr, though with uncertainties that overlap the ship-only estimate. An atmospheric inversion reveals that a shift of this magnitude in the contemporary Southern Ocean carbon flux must be compensated for by ocean or land sinks within the Southern Hemisphere.
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Antarctic krill (Euphausia superba) are swarming, oceanic crustaceans, up to two inches long, and best known as prey for whales and penguins – but they have another important role. With their large size, high biomass and daily vertical migrations they transport and transform essential nutrients, stimulate primary productivity and influence the carbon sink. Antarctic krill are also fished by the Southern Ocean’s largest fishery. Yet how krill fishing impacts nutrient fertilisation and the carbon sink in the Southern Ocean is poorly understood. Our synthesis shows fishery management should consider the influential biogeochemical role of both adult and larval Antarctic krill.
Article
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In many areas of the world’s ocean such as the Southern Ocean (SO), primary production is low despite an abundance of macronutrients. In these high nutrient low chlorophyll (HNLC) regions the trace metal (TM) iron (Fe) limits phytoplankton biomass and subsequently the biological carbon pump. Besides Fe, the TMs zinc (Zn), cobalt (Co), and the vitamin cobalamin (B12) have also been shown to limit biomass and/or influence plankton species composition. While the impacts of Fe limitation and, to a lesser degree of Zn and Co, on the cellular physiology of Antarctic phytoplankton have been investigated, studies focusing simultaneously on several TMs and vitamins are still lacking. This study measured the impacts of Fe, Zn, Co, and B12 limitation on the Antarctic diatom Chaetoceros simplex and Fe and Zn limitation on the Antarctic cryptophyte Geminigera cryophila. Both species responded to all limitation scenarios by reducing their growth and particulate organic carbon (POC) production rates. For both algae limitation by Fe and Zn resulted in a reduction of light harvesting pigments, a significant reduction in the photosynthetic yield (Fv/Fm) and increase in the C:N ratio. Most interestingly, with a few exceptions, limitation by one TM also resulted in a significant decrease of the cellular quotas of other TMs measured. These observations suggest that one consequence of limitation by one TM may be a secondary and perhaps more fatal limitation by another.
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High-latitude ecosystems are among the fastest warming on the planet¹. Polar species may be sensitive to warming and ice loss, but data are scarce and evidence is conflicting2–4. Here, we show that, within their main population centre in the southwest Atlantic sector, the distribution of Euphausia superba (hereafter, ‘krill’) has contracted southward over the past 90 years. Near their northern limit, numerical densities have declined sharply and the population has become more concentrated towards the Antarctic shelves. A concomitant increase in mean body length reflects reduced recruitment of juvenile krill. We found evidence for environmental controls on recruitment, including a reduced density of juveniles following positive anomalies of the Southern Annular Mode. Such anomalies are associated with warm, windy and cloudy weather and reduced sea ice, all of which may hinder egg production and the survival of larval krill⁵. However, the total post-larval density has declined less steeply than the density of recruits, suggesting that survival rates of older krill have increased. The changing distribution is already perturbing the krill-centred food web⁶ and may affect biogeochemical cycling7,8. Rapid climate change, with associated nonlinear adjustments in the roles of keystone species, poses challenges for the management of valuable polar ecosystems³.
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The island of South Georgia is situated in the iron (Fe)-depleted Antarctic Circumpolar Current of the Southern Ocean. Iron emanating from its shelf system fuels large phytoplankton blooms downstream of the island, but the actual supply mechanisms are unclear. To address this, we present an inventory of Fe, manganese (Mn), and aluminium (Al) in shelf sediments, pore waters, and the water column in the vicinity of South Georgia, alongside data on zooplankton-mediated Fe cycling processes, and provide estimates of the relative dissolved Fe (DFe) fluxes from these sources. Seafloor sediments, modified by authigenic Fe precipitation, were the main particulate Fe source to shelf bottom waters as indicated by the similar Fe∕Mn and Fe∕Al ratios for shelf sediments and suspended particles in the water column. Less than 1% of the total particulate Fe pool was leachable surface-adsorbed (labile) Fe and therefore potentially available to organisms. Pore waters formed the primary DFe source to shelf bottom waters, supplying 0.1–44µmolDFem⁻²d⁻¹. However, we estimate that only 0.41±0.26µmolDFem⁻²d⁻¹ was transferred to the surface mixed layer by vertical diffusive and advective mixing. Other trace metal sources to surface waters included glacial flour released by melting glaciers and via zooplankton egestion and excretion processes. On average 6.5±8.2µmolm⁻²d⁻¹ of labile particulate Fe was supplied to the surface mixed layer via faecal pellets formed by Antarctic krill (Euphausia superba), with a further 1.1±2.2µmolDFem⁻²d⁻¹ released directly by the krill. The faecal pellets released by krill included seafloor-derived lithogenic and authigenic material and settled algal debris, in addition to freshly ingested suspended phytoplankton cells. The Fe requirement of the phytoplankton blooms ∼ 1250km downstream of South Georgia was estimated as 0.33±0.11µmolm⁻²d⁻¹, with the DFe supply by horizontal/vertical mixing, deep winter mixing, and aeolian dust estimated as ∼ 0.12µmolm⁻²d⁻¹. We hypothesize that a substantial contribution of DFe was provided through recycling of biogenically stored Fe following luxury Fe uptake by phytoplankton on the Fe-rich shelf. This process would allow Fe to be retained in the surface mixed layer of waters downstream of South Georgia through continuous recycling and biological uptake, supplying the large downstream phytoplankton blooms.
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The availability of iron controls primary productivity in large areas of the Southern Ocean. Iron is largely supplied via atmospheric dust deposition, melting ice, the weathering of shelf sediments, upwelling, sediment resuspension, mixing (deep water, biogenic, and vertical mixing) and hydrothermal vents with varying degrees of temporal and spatial importance. However, large areas of the Southern Ocean are remote from these sources, leading to regions of low primary productivity. Recent studies suggest that recycling of iron by animals in the surface layer could enhance primary productivity in the Southern Ocean. The aim of this review is to provide a quantitative and qualitative assessment of the current literature on pelagic iron recycling by marine animals in the Southern Ocean and highlight the next steps forward in quantifying the retention and recycling of iron by higher trophic levels in the Southern Ocean. Phytoplankton utilize the iron in seawater to meet their metabolic demand. Through grazing, pelagic herbivores transfer the iron in phytoplankton cells into their body tissues and organs. Herbivores can recycle iron through inefficient feeding behavior that release iron into the water before ingestion, and through the release of fecal pellets. The iron stored within herbivores is transferred to higher trophic levels when they are consumed. When predators consume iron beyond their metabolic demand it is either excreted or defecated. Waste products from pelagic vertebrates can thus contain high concentrations of iron which may be in a form that is available to phytoplankton. Bioavailability of fecal iron for phytoplankton growth is influenced by a combination of the size of the fecal particle, presence of organic ligands, the oxidation state of the iron, as well as biological (e.g., remineralization, coprochaly, coprorhexy, and coprophagy) and physical (e.g., dissolution, fragmentation) processes that lead to the degradation and release of fecal iron. The flux of dissolved iron from pelagic recycling is comparable to other sources in the region such as atmospheric dust, vertical diffusivity, vertical flux, lateral flux and upwelling, but lower than sea ice, icebergs, sediment resuspension, and deep winter mixing. The temporal and seasonal importance of these various factors requires further examination.
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The projected rise in anthropogenic CO2 and associated ocean acidification (OA) will change trace metal solubility and speciation, potentially altering Southern Ocean (SO) phytoplankton productivity and species composition. As iron (Fe) sources are important determinants of Fe bioavailability, we assessed the effect of Fe-laden dust versus inorganic Fe (FeCl3) enrichment under ambient and high pCO² levels (390 and 900 μatm) in a naturally Fe-limited SO phytoplankton community. Despite similar Fe chemical speciation and net particulate organic carbon (POC) production rates, CO²-dependent species shifts were controlled by Fe sources. Final phytoplankton communities of both control and dust treatments were dominated by the same species, with an OA-dependent shift from the diatom Pseudo-nitzschia prolongatoides towards the prymnesiophyte Phaeocystis antarctica. Addition of FeCl3 resulted in high abundances of Nitzschia lecointei and Chaetoceros neogracilis under ambient and high pCO2, respectively. These findings reveal that both the characterization of the phytoplankton community at the species level and the use of natural Fe sources are essential for a realistic projection of the biological carbon pump in the Felimited pelagic SO under OA. As dust deposition represents a more realistic scenario for the Felimited pelagic SO under OA, unaffected net POC production and dominance of P. antarctica can potentially weaken the export of carbon and silica in the future.
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Planktonic grazers such as salps may have a dominant role in iron (Fe) cycling in surface waters of the Southern Ocean (SO). Salps have high ingestion rates and egest large, fast sinking fecal pellets (FPs) that potentially contribute to the vertical flux of carbon. In this study, we determined the impact of FPs from Salpa thompsoni, the most abundant salp in the SO, on Fe biogeochemistry. During the Polarstern expedition ANT-XXVII/3, salps were sampled from a large diatom bloom area in the Atlantic sector of the SO. Extensive work on carbon export and salp FPs export at the sampling location had shown that salps were a minor component of zooplankton and were responsible for only a 0.2% consumption of the daily primary production. Furthermore, at 100 m, export efficiency of salp FPs was ∼2–3 fold higher than that of the bulk of sinking particulate organic carbon (POC). After collection, salps were maintained in 200 µm screened seawater and their FPs were collected for further experiments. To investigate whether the FPs release Fe and/or Fe-binding ligands into the filtered seawater (FSW) under different experimental conditions, they were either incubated in the dark or under full sunlight at in situ temperatures for 24 h, or placed into the dark after a freeze/thaw treatment. We observed that none of the treatments caused release of dissolved Fe (dFe) or strong Fe ligands from the salp FPs. However, humic-substance like (HS-like) compounds, weak Fe ligands, were released at a rate of 8.2 ± 4.7 µg HS-like FP −1 d −1. Although the Fe content per salp FP was high at 0.33 −1 ± 0.02 nmol dFe FP , the small contribution of salps to the zooplankton pool resulted in an estimated dFe export flux of 11.3 nmol Fe m −2 d −1 at 300 m. Since salp FPs showed an export efficiency at 100 m well above that shown by the bulk of sinking POC, our results suggest that in those areas of the SO where salps play a major role in the grazing of primary production, they could be actively contributing to the depletion of the dFe pool in surface water.
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The microbial degradation of marine particles is an important process in the remineralization of nutrients including iron. As part of the GEOTRACES process study (FeCycle II), we conducted incubation experiments with marine particles obtained from 30 and 100 m depth at two stations during austral spring in the subtropical waters east of the North Island of New Zealand. The particles were collected using in-situ pumps, and comprised mainly of suspended and slow sinking populations along with associated attached heterotrophic bacteria. In treatments with live bacteria, increasing concentrations of Fe binding ligands were observed with an average stability constant of logKFeL,Fe3+ = 21.11±0.37 for station 1 and 20.89±0.25 for station 2. The ligand release rates varied between 2.54 and 11.8 pmol L-1 d-1 (calculated for ambient seawater particle concentration) and were similar to those found in two Southern Ocean subsurface studies from ~110 m depths in subpolar and polar waters. Dissolved iron (DFe) was released at a rate between 0.33 and 2.09 pmol Fe L-1 d-1 with a column integrated (30 -100 m) flux of 107 and 58 nmol Fe m-2 day-1 at station 1 and 2, respectively. Given a mixed layer DFe inventory of ~48 µmol m-2 and ~4 µmol m-2 at the time of sampling for station 1 and 2, this will therefore result in a DFe residence time of 1.2 and 0.18 years, assuming particle remineralization was the only source of iron in the mixed layer. The DFe release rates calculated were comparable to those found in the previously mentioned study of Southern Ocean water masses. Fe-binding ligand producing bacteria (CAS positive) abundance was found to increase throughout the duration of the experiment of 7 to 8 days. For the first time ferrioxamine type siderophores, including the well-known ferrioxamine B and G, have been quantified using chemical assays and LC-ESI-MS. Our subtropical study corroborates prior reports from the Southern Ocean of particle remineralization being an important source of DFe and ligands, and adds unprecedented detail by revealing that siderophores are probably an important component of the ligands released into subsurface waters during particle remineralisation.
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Summary Iron is an essential nutrient for phytoplankton, but low concentrations limit primary production and associated atmospheric carbon drawdown in large parts of the world’s oceans [ 1, 2 ]. Lithogenic particles deriving from aeolian dust deposition, glacial runoff, or river discharges can form an important source if the attached iron becomes dissolved and therefore bioavailable [ 3–5 ]. Acidic digestion by zooplankton is a potential mechanism for iron mobilization [ 6 ], but evidence is lacking. Here we show that Antarctic krill sampled near glacial outlets at the island of South Georgia (Southern Ocean) ingest large amounts of lithogenic particles and contain 3-fold higher iron concentrations in their muscle than specimens from offshore, which confirms mineral dissolution in their guts. About 90% of the lithogenic and biogenic iron ingested by krill is passed into their fecal pellets, which contain ∼5-fold higher proportions of labile (reactive) iron than intact diatoms. The mobilized iron can be released in dissolved form directly from krill or via multiple pathways involving microbes, other zooplankton, and krill predators. This can deliver substantial amounts of bioavailable iron and contribute to the fertilization of coastal waters and the ocean beyond. In line with our findings, phytoplankton blooms downstream of South Georgia are more intensive and longer lasting during years with high krill abundance on-shelf. Thus, krill crop phytoplankton but boost new production via their nutrient supply. Understanding and quantifying iron mobilization by zooplankton is essential to predict ocean productivity in a warming climate where lithogenic iron inputs from deserts, glaciers, and rivers are increasing [ 7–10 ].
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This study aimed to create the first model of biological iron (Fe) cycling in the Southern Ocean food web. Two biomass mass-balanced Ecopath models were built to represent pre- and post-whaling ecosystem states (1900 and 2008). Functional group biomasses (tonnes wet weight km ⁻² ) were converted to biogenic Fe pools (kg Fe km ⁻² ) using published Fe content ranges. In both models, biogenic Fe pools and consumption in the pelagic Southern Ocean were highest for plankton and small nektonic groups. The production of plankton biomass, particularly unicellular groups, accounted for the highest annual Fe demand. Microzooplankton contributed most to biological Fe recycling, followed by carnivorous zooplankton and krill. Biological Fe recycling matched previous estimates, and, under most conditions, could entirely meet the Fe demand of bacterioplankton and phytoplankton. Iron recycling by large baleen whales was reduced 10-fold by whaling between 1900 and 2008. However, even under the 1900 scenario, the contribution of whales to biological Fe recycling was negligible compared with that of planktonic consumers. These models are a first step in examining oceanic-scale biological Fe cycling, highlighting gaps in our present knowledge and key questions for future research on the role of marine food webs in the cycling of trace elements in the sea. This article is part of the themed issue ‘Biological and climatic impacts of ocean trace element chemistry’.
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It has univocally been shown that iron (Fe) is the primary limiting nutrient for phytoplankton metabolism in High Nutrient Low Chlorophyll (HNLC) oceans, yet, the question of how this trace metal affects heterotrophic microbial activity is far less understood. We investigated the role of Fe for bacterial heterotrophic production and growth at three contrasting sites in the naturally Fe-fertilized region east of Kerguelen Islands and at one site in HNLC waters during the KEOPS2 (Kerguelen Ocean and Plateau Compared Study 2) cruise in spring 2011. We performed dark incubations of natural microbial communities amended either with iron (Fe, as FeCl3), or carbon (C, as trace-metal clean glucose), or a combination of both, and followed bacterial abundance and heterotrophic production for up to 7 days. Our results show that single and combined additions of Fe and C stimulated bulk and cell-specific bacterial production at all sites, while bacterial growth was enhanced only in two out of four occasions. The extent of stimulation of bulk bacterial heterotrophic production by single Fe or C additions increased with increasing in situ bacterial Fe uptake rates in the surface mixed layer. Our results provide evidence that both Fe and C are present at limiting concentrations for bacterial heterotrophic activity, in HNLC and fertilized regions, in spring. The observation that the extent of stimulation by both elements was related to Fe-uptake rates highlights the tight interaction between the C- and Fe-cycles through bacterial heterotrophic metabolism in the Southern Ocean.
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Photophysiological processes as well as uptake characteristics of iron and inorganic carbon were studied in inshore phytoplankton assemblages of the Western Antarctic Peninsula (WAP) and offshore assemblages of the Drake Passage. Chlorophyll a concentrations and primary productivity decreased from in- to offshore waters. The inverse relationship between low maximum quantum yields of photochemistry in PSII (Fv/Fm) and large sizes of functional absorption cross sections (σPSII) in offshore communities indicated iron-limitation. Congruently, the negative correlation between Fv/Fm values and iron uptake rates across our sampling locations suggest an overall better iron uptake capacity in iron-limited pelagic phytoplankton communities. Highest iron uptake capacities could be related to relative abundances of the haptophyte Phaeocystis antarctica. As chlorophyll a-specific concentrations of humic-like substances were similarly high in offshore and inshore stations, we suggest humic-like substances may play an important role in iron chemistry in both coastal and pelagic phytoplankton assemblages. Regarding inorganic carbon uptake kinetics, the measured maximum short-term uptake rates (Vmax(CO2)) and apparent half-saturation constants (K1/2(CO2)) did not differ between offshore and inshore phytoplankton. Moreover, Vmax(CO2) and K1/2(CO2) did not exhibit any CO2-dependent trend over the natural pCO2 range from 237 to 507 µatm. K1/2(CO2) strongly varied among the sampled phytoplankton communities, ranging between 3.5 and 35.3 µmol L−1 CO2. While in many of the sampled phytoplankton communities, the operation of carbon-concentrating mechanisms (CCMs) was indicated by low K1/2(CO2) values relative to ambient CO2 concentrations, some coastal sites exhibited higher values, suggesting down-regulated CCMs. Overall, our results demonstrate a complex interplay between photophysiological processes, iron and carbon uptake of phytoplankton communities of the WAP and the Drake Passage.
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The micronutrient iron (Fe) is rapidly cycled in surface waters, and regenerated Fe supports much of the phytoplankton growth in open ocean waters. Meso- and microzooplankton grazing are both important mechanisms to regenerate Fe, but the chemical conditions in the respective digestive systems are different and might affect the bioavailability of Fe. We conducted radiotracer grazing experiments with the copepod Acartia tonsa or the heterotrophic dinoflagellate Oxyrrhis marina feeding on the diatom Thalassiosira pseudonana or the coccolithophore Emiliania huxleyi. Uptake of regenerated Fe-55 by a separate T. pseudonana culture was compared to the uptake of inorganic Fe. Iron regenerated by A. tonsa was taken up 4- to 7-fold faster than inorganic iron. In contrast, no difference was detected between the uptake rate of inorganic Fe and Fe regenerated by O. marina. Ingestion of different prey by A. tonsa revealed that Fe released during diatom digestion was taken up 1.8-fold faster than Fe released from coccolithophore digestion. Digestive systems and the chemical makeup of the ingested prey are crucial in determining the bioavailability of regenerated Fe. Differences in pH and oxygen saturation between digestive vacuoles and guts may affect the speciation of regenerated Fe, and the release of ferrous Fe might contribute to the bioavailability of regenerated Fe. The ecological structure determines the importance of regenerated Fe for a particular ecosystem.
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Phytoplankton are often limited by iron in aquatic environments. Here we examine Fe bioavailability to phytoplankton by analyzing iron uptake from various Fe substrates by several species of phytoplankton grown under conditions of Fe limitation and comparing the measured uptake rate constants (Fe uptake rate/ substrate concentration). When unchelated iron, Fe′, buffered by an excess of the chelating agent EDTA is used as the Fe substrate, the uptake rate constants of all the eukaryotic phytoplankton species are tightly correlated and proportional to their respective surface areas (S.A.). The same is true when FeDFB is the substrate, but the corresponding uptake constants are one thousand times smaller than for Fe′. The uptake rate constants for the other substrates we examined fall mostly between the values for Fe′ and FeDFB for the same S.A. These two model substrates thus empirically define a bioavailability envelope with Fe′ at the upper and FeDFB at the lower limit of iron bioavailability. This envelope provides a convenient framework to compare the relative bioavailabilities of various Fe substrates to eukaryotic phytoplankton and the Fe uptake abilities of different phytoplankton species. Compared with eukaryotic species, cyanobacteria have similar uptake constants for Fe′ but lower ones for FeDFB. The unique relationship between the uptake rate constants and the S.A. of phytoplankton species suggests that the uptake rate constant of Fe-limited phytoplankton has reached a universal upper limit and provides insight into the underlying uptake mechanism.
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The “biological pump” is the process by which photosynthetically-produced organic matter in the ocean descends from the surface layer to depth by a combination of sinking particles, advection or vertical mixing of dissolved organic matter, and transport by animals. Particulate organic matter that is exported downward from the euphotic zone is composed of combinations of fecal pellets from zooplankton and fish, organic aggregates known as “marine snow” and phytodetritus from sinking phytoplankton. Previous reviews by Turner and Ferrante (1979) and Turner (2002) focused on publications that appeared through late 2001. Since that time, studies of the biological pump have continued, and there have been > 300 papers on vertical export flux using sediment traps, large-volume filtration systems and other techniques from throughout the global ocean. This review will focus primarily on recent studies that have appeared since 2001. Major topics covered in this review are 1) an overview of the biological pump, and its efficiency and variability, and the role of dissolved organic carbon in the biological pump; 2) zooplankton fecal pellets, including the contribution of zooplankton fecal pellets to export flux, epipelagic retention of zooplankton fecal pellets due to zooplankton activities, zooplankton vertical migration and fecal pellet repackaging, microbial ecology of fecal pellets, sinking velocities of fecal pellets and aggregates, ballasting of sinking particles by mineral contents, phytoplankton cysts, intact cells and harmful algae toxins in fecal pellets, importance of fecal pellets from various types of zooplankton, and the role of zooplankton fecal pellets in picoplankton export; 3) marine snow, including the origins, abundance, and distributions of marine snow, particles and organisms associated with marine snow, consumption and fragmentation of marine snow by animals, pathogens associated with marine snow; 4) phytodetritus, including pulsed export of phytodetritus, phytodetritus from Phaeocystis spp., picoplankton in phytodetritus, the summer export pulse (SEP) of phytodetritus in the subtropical North Pacific, benthic community responses to phytodetritus; 5) other components of the biological pump, including fish fecal pellets and fish-mediated export, sinking carcasses of animals and macrophytes, feces from marine mammals, transparent exopolymer particles (TEP); 6) the biological pump and climate, including origins of the biological pump, the biological pump and glacial/interglacial cycles, the biological pump and contemporary climate variations, and the biological pump and anthropogenic climate change. The review concludes with potential future modifications in the biological pump due to climate change.
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Advances in iron biogeochemistry have transformed our understanding of the oceanic iron cycle over the past three decades: multiple sources of iron to the ocean were discovered, including dust, coastal and shallow sediments, sea ice and hydrothermal fluids. This new iron is rapidly recycled in the upper ocean by a range of organisms; up to 50% of the total soluble iron pool is turned over weekly in this way in some ocean regions. For example, bacteria dissolve particulate iron and at the same time release compounds - iron-binding ligands - that complex with iron and therefore help to keep it in solution. Sinking particles, on the other hand, also scavenge iron from solution. The balance between these supply and removal processes determines the concentration of dissolved iron in the ocean. Whether this balance, and many other facets of the biogeochemical cycle, will change as the climate warms remains to be seen.
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A majority of eukaryotic phytoplankton species require an exogenous source of vitamin B12 for growth and recent field studies in some coastal and polar regions indicate that the addition of vitamin B12 alone, or with another limiting nutrient can influence the accumulation of phytoplankton biomass. We quantified the concentrations and uptake rates of vitamin B12, characterized phytoplankton community composition, and examined the ability of vitamin B12 to alter the growth and composition of phytoplankton communities in the Gulf of Alaska. Picoplankton (0.2-2 μm) were responsible for the majority of vitamin B12 uptake in both coastal and high-nutrient low-chlorophyll (HNLC) regions and B12 concentrations and uptake rates were higher in HNLC regions compared to coastal regions with higher iron (Fe) concentrations. During vitamin amendment experiments, B12 alone or in conjunction with other limiting nutrients (N or Fe) significantly enhanced algal biomass and increased the growth rates of multiple groups of larger (> 2 μm) phytoplankton. This included ecologically significant, B12 auxotrophs such as Gymnodinium sp. and Alexandrium sp. (in the costal experiment) as well as Chaetoceros sp. and Gymnodinium sp. (in the HNLC experiment). The ability of vitamin B12 to shape algal community composition in coastal and HNLC areas of the Gulf of Alaska, even in cases where it does not limit total phytoplankton production, suggests that it may influence carbon export in this and other polar ecosystems.
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Krill (Euphausia superba) provide a direct link between primary producers and higher trophic levels in the Antarctic marine food web. The pelagic tunicate Salpa thompsoni can also be important during spring and summer through the formation of extensive and dense blooms. Although salps are not a major dietary item for Antarctic vertebrate predators,, their blooms can affect adult krill reproduction and survival of krill larvae. Here we provide data from 1995 and 1996 that support hypothesized relationships between krill, salps and region-wide sea-ice conditions,. We have assessed salp consumption as a proportion of net primary production, and found correlations between herbivore densities and integrated chlorophyll-a that indicate that there is a degree of competition between krill and salps. Our analysis of the relationship between annual sea-ice cover and a longer time series of air temperature measurements, indicates a decreased frequency of winters with extensive sea-ice development over the last five decades. Our data suggest that decreased krill availability may affect the levels of their vertebrate predators. Regional warming and reduced krill abundance therefore affect the marine food web and krill resource management.
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In large parts of the Southern Ocean, primary production is limited due to shortage of iron (Fe). We measured vertical Fe profiles in the western Weddell Sea, Weddell-Scotia Confluence, and Antarctic Circumpolar Current (ACC), showing that Fe is derived from benthic Fe diffusion and sediment resuspension in areas characterized by high turbulence due to rugged bottom topography. Our data together with literature data reveal an exponential decrease of dissolved Fe (DFe) concentrations with increasing distance from the continental shelves of the Antarctic Peninsula and the western Weddell Sea. This decrease can be observed 3500 km eastward of the Antarctic Peninsula area, downstream the ACC. We estimated DFe summer fluxes into the upper mixed layer of the Atlantic sector of the Southern Ocean and found that horizontal advection dominates DFe supply, representing 54 ± 15% of the total flux, with significant vertical advection second most important at 29 ± 13%. Horizontal and vertical diffusion are weak with 1 ± 2% and 1 ± 1%, respectively. The atmospheric contribution is insignificant close to the Antarctic continent but increases to 15 ± 10% in the remotest waters (>1500 km offshore) of the ACC. Translating Southern Ocean carbon fixation by primary producers into biogenic Fe fixation shows a twofold excess of new DFe input close to the Antarctic continent and a one-third shortage in the open ocean. Fe recycling, with an estimated “fe” ratio of 0.59, is the likely pathway to balance new DFe supply and Fe fixation.
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Salp fecal pellets are rich in organic matter and have been shown to sink at very high velocities. In recent years, salp abundances have been increasing in the Southern Ocean where they seem to be replacing krill as the dominant grazers on phytoplankton. As salps can form large swarms with high pellet production rates, it has been suggested that they will become increasingly important for the vertical export of particulate organic matter in the Southern Ocean. However, detailed studies combining both investigations of pellet production rates, turnover, and export are still needed in order to determine whether salp pellets are important for export (‘sinkers’) or recycling (‘floaters’) of organic matter. Our results suggest that pellets are produced at high rates in the upper few hundred meters of the water column. Although we observed high sinking velocities and low microbial degradation rates of the produced salp pellets, only about one third of the produced pellets were captured in sediment traps placed at 100 m and about ~13% of the produced pellets were exported to sediment traps placed at 300 m. The high retention of these fast-settling pellets seems to be caused by break-up and loosening of the pellets, possibly by zooplankton and salps themselves. We measured 3-fold lower size-specific sinking velocities in loosened and fragmented compared to freshly produced intact pellets-. This enhanced the residence times (>1 day) of both small and large pellets in the upper water column. We postulate that the fragile nature of salp pellets make them more important for recycling of organic matter in the upper mesopelagic layer rather than as a conduit for export of particulate organic matter to the seafloor.
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Iron limits phytoplankton growth in large areas of the Southern Ocean. A new study shows that Antarctic krill play a crucial role in the recycling of iron in the iron-limited waters.
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Antarctic krill may play a significant role in the Southern Ocean iron cycle. However, understanding the control on iron budgets by Antarctic krill is hampered by the large range in the reported iron concentration of krill. The aim of this study was to investigate the causes of the large range of iron concentrations in krill reported in the literature (6-190 mg kg-1). Antarctic krill samples were collected from three research voyages to Pyrdz Bay, Antarctica, and analysed individually. Iron concentrations were measured using sector field inductively coupled plasma mass spectrometry in whole krill specimens and in the isolated stomach, digestive gland, muscle, body (whole krill excluding stomach and digestive gland), exoskeleton and faecal pellets. Iron concentrations in stomach (6-98 mg/kg), digestive gland (14-82 mg kg-1), and faecal pellet (683-1039 mg kg-1) were higher compared to muscle (4-7 mg kg-1), exoskeleton (6-15 mg kg-1), and body (4-18 mg kg-1) indicating that krill may ingest more iron than they require for physiological processes. Iron concentrations in whole krill from March 2012 (10±3 mg kg-1) were significantly lower compared to February 2003 (19±7 mg kg-1) and February 2015 (18±12 mg kg-1). Overall, the iron concentrations in krill from this study were consistently at the lower end of the published range. We propose that the large range in reported whole iron concentrations of krill can be accounted for by a combination of seasonal and regional differences in sampling, reflecting differences in the quantity and quality of their diet. © 2016 Association for the Sciences of Limnology and Oceanography.
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Field-collected living specimens of the benthic foraminifera Haynesina germanica were maintained in the laboratory and fed a naturally occurring motile benthic diatom assemblage dominated by Pleurosigma angulatum. The extracellular removal of diatom contents was inferred for P. angulatum in controlled experiments. A characteristic pattern of fracturing of the diatom frustule was observed that was directly attributed to foraminiferal feeding/sequestration mechanisms. These feeding/sequestration mechanisms have a potentially important bearing on our understanding of foraminiferal aperture morphology, foraminiferal evolution and the preservation of diatoms in marine sediments. Recognition of this characteristic breakage pattern of diatom frustules may provide insight into the natural importance of foraminifera in grazing diatom biofilms. (c) 2005 Elsevier B.V All rights reserved.
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In the Southern Ocean, phytoplankton growth is largely limited by the lack of iron, affecting the biogeochemical cycling not only of iron itself but also of other elements, including nutrients and carbon. It is now recognized that iron limitation affects carbon cycling globally and thus plays a role in Earth's climate regulation. The bioavailable fraction of iron is the fraction that can effectively interact with phytoplankton to support their iron-dependent metabolic reactions and growth. As such, it is the bioavailable iron pool that shapes phytoplankton communities in most of the Southern Ocean. Despite numerous studies, parameters controlling iron bioavailability to phytoplankton are still poorly understood, probably due to an extremely complex and dynamic interplay between iron chemistry and biology in surface waters. Iron bioavailability depends on chemical and physical speciation and the different uptake strategies of the phyto- and bacterio-plankton communities. In the Southern Ocean, 99% of the dissolved iron is complexed by organic ligands, which likely controls its bioavailability. Furthermore, microorganisms also exert feedback on iron chemistry, for instance, by releasing organic iron-binding ligands through production, cell lysis, or degradation of fecal pellets, as well as by reducing iron at the cell surface. Regeneration of iron, through grazing as well as bacterial and viral activities, is another pathway that supplies iron to phytoplankton communities. Field investigations of iron speciation in the Southern Ocean are discussed in conjunction with laboratory assessments of iron speciation and bioavailability using natural assemblages and strains isolated from the Southern Ocean. Methods to measure iron bioavailability and recent developments in mathematical models are also presented.
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Growth limitation of marine algae due to lack of iron occurs in up to 40 % of the global ocean. Despite important advances on the impact of organic compounds on iron biogeochemistry, their roles in controlling iron availability to prokaryotic and eukaryotic phytoplankton remain unclear. Whether algal and bacterial exopolymeric substances (EPS) include organic ligands which may help iron-limited phytoplankton growth remains an unknown. If so, then EPS could relieve phytoplankton iron limitation with implications for the biological carbon pump and hence the regulation of atmospheric CO2. Here we compared the biological impact of algal, bacterial and in situ EPS with model compounds, a siderophore and two saccharides on biological parameters including, iron bioavailability, phytoplankton growth, photo-physiology and community structure. Laboratory and field experiments demonstrated that EPS produced by marine microorganisms are efficient in sustaining biological iron uptake as well as algal growth, and can affect natural phytoplankton community structure. Our data suggest that natural phytoplankton growth enhancement in the presence of EPS was not solely due to highly bioavailable iron forms, but also because EPS contains other micronutrients. Stronger ligands were detected following iron-siderophore enrichments (log KFe’L = 12.0) and weaker ligands were measured in the presence of EPS (log KFe’L = 10.4-11.0). The trend of the conditional stability constants of organic ligands did not seem to be affected as a result of biological activity and photo-chemistry during our four days incubations. The shift in the phytoplankton community observed during our field experiments was not uniformly observed between different sites rendering it difficult to extrapolate which functional group(s) would benefit the most from iron bound to EPS.
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Biological vectors are important for redistribution of nutrients in many ecological systems. While availability of iron (Fe) to phyto - plankton limits pelagic productivity in the Southern Ocean, biomagnification within marine food webs can lead to high concentrations of Fe in the diet of seabirds and marine mammals. We investigated patterns in concentrations of the micronutrients Fe, Co, Zn and Mn, and the toxins Cd and As, in the guano of oceanic, coastal and predatory seabirds and in faeces of 2 species of marine mammals that congregate to breed in the sub-Antarctic Auckland Islands. We found that much of the variability in concentrations of Fe, Co, Zn and Mn among species could be explained by foraging behaviour and by trophic position. We observed concentrations of Fe to be 8 orders of magnitude higher in the guano of predators and coastal foragers than in the sub-Antarctic mixed layer. High concentrations of As and Cd were associated with organic matter sources from macroalgae. Analyses of the molar ratio Fe:Al indicated that Fe within food webs supporting seabirds has likely been extensively recycled from its lithogenic source. Patterns in Fe:N among species indicated that Fe is concentrated 2 to 4 orders of magnitude in the guano of seabirds compared to limiting conditions for phytoplankton growth in sub-Antarctic waters. These data highlight the potential role of seabirds and marine mammals in the redistribution of micronutrients in the Southern Ocean and their likely role as key nutrient vectors in the ecosystem, particularly around the sub-Antarctic islands during the breeding season.
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In order to establish the potential role of Antarctic krill (Euphausia superba) in the recycling of bioactive elements, we have quantified the release of iron, phosphate, and ammonia by these organisms along the Antarctic Peninsula sector of the Southern Ocean. The experimental results suggested that the presence of krill has a significant impact on ambient iron concentrations, as large amounts of this trace element were released by the krill (22-689 nmol Fe g Dry Weight-1 h-1, equivalent to 0.2 to 4.3 nmol Fe L-1 d-1). Half of this iron release occurred within the first hour of the experiment, and differences in iron and phosphate release rates (3.1 to 14.0 mumol PO4 3- g DW-1 h-1) seemed to reflect differences in food availability. These results identify krill as a major node in iron cycling in the Southern Ocean, potentially influencing iron residence time in the upper water column of this region.
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Available data on the spatial distribution and feeding ecophysiology of Antarctic krill, Euphausia superba, and the tunicate, Salpa thompsoni, in the Southern Ocean are summarized in this study. Antarctic krill and salps generally display pronounced spatial segregation at all spatial scales. This appears to be the result of a clear biotopical separation of these key species in the Antarctic pelagic food web. Krill and salps are found in different water masses or water mass modifications, which are separated by primary or secondary frontal features. On the small-scale (
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The solubilities of iron(III) hydroxides in seawater were determined in Gulf Stream seawater as a function of pH (2 to 9), temperature (5 to 50 °C) and salinity (0 to 36). Our results at S=36 and 25 °C near a pH of 8 are in agreement with the measurements of Byrne and Kester [Mar. Chem. 4 (1976a) 255] and Kuma et al. [Limnol. Oceanogr. 41 (1996) 396] (0.2 to 0.3 nM). The solubilities at 5 °C are considerably higher than at 25 °C and decrease with a decrease in salinity. Near a pH of 8, the solubilities as a function of temperature (T/K) and ionic strength [I=19.922S/(1000−1.005S)] can be estimated from
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The atmospheric transport of continental weathering products is responsible for much of the mineral material and Fe entering the open ocean and is probably the dominant source of nutrient Fe in the photic zone. In regions where other nutrients are present in high concentrations, the flux of Fe from the atmosphere may be a limiting factor in primary productivity. Due to the larger source regions for dust north of the equator, â¼8 times more atmospheric Fe is deposited in the northern hemisphere than in the southern hemisphere. The mineral aerosol and Fe transport and deposition are highly variable due to the episodic nature of dust generation and its transport and deposition processes. Between 10 and 50% of the total atmospheric Fe entering the world ocean appears to dissolve rapidly when the mineral matter enters the ocean. Much of the atmospheric Fe is present as Fe(II), apparently produced as a result of photochemical reduction reactions taking place during atmospheric transport. This readily soluble Fe(II) should be available immediately for use as a nutrient by phytoplankton. Atmospheric transport from the continents is estimated to supply â¼3 times as much dissolved Fe to the oceans as that delivered via rivers.
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Concentrations of 11 trace elements-scandium, lanthanum, cerium, europium, thorium, rubidium, cesium, strontium, iron, zinc, and cobalt-have been detcrmincd by activation analysis in two tunicates. Their concentration factors were calculated and are discussed in relation to radioccology and to seawater pollution with radionuclides. The treatment of samples and the radiochemical procedure are described.
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The role of heterotrophic bacteria in iron recycling, the influence of complexation on iron remineralization, and iron mobilization rates from lithogenic vs. biogenic particulate iron (PFe) were examined using field experiments and modeling simulations. During summer, we measured the mobilization rate of algal iron by heterotrophic bacteria in the mixed layer at a polar and a subpolar site south of Australia, and conducted shipboard incubations to track the release of dissolved iron (DFe) and iron-binding ligands from subsurface settling particles sampled from 120-m depth. Bacteria mobilized > 25% PFe d−1 in surface waters relative to mobilization at depth (< 2% d−1). Our incubations provide the first evidence of the concurrent release of weak iron-binding ligands and DFe from sinking particles. Simulated profiles of PFe remineralization, based on proxies, point to greater dissolution from biogenic PFe than from lithogenic PFe. Together our findings point to different biogeochemical functions for lithogenic vs. biogenic PFe: biogenic PFe is probably the main source of both DFe and ligands, whereas lithogenic PFe may contribute most to DFe scavenging and ballasting of biogenic PFe. The relative proportions of lithogenic vs. biogenic PFe flux vary regionally and set the contribution of scavenging and ballasting vs. dissolution and ligand release, and hence the fate of iron in the water column.
Article
Enrichment experiments were performed in the Ross Sea to test the hypothesis that iron deficiency is responsible for the phytoplankton's failure to use up the luxuriant major nutrient supplies found in these and all other offshore Antarctic ocean waters . Nitrate uptake rates in the controls without added trace elements ranged from 0.58 to 1.22 μmol kg-1 d-1; the addition of 1 to 5 nmol of unchelated Fe per liter resulted in rates that were 2 to 10 times higher (2.54 to 6.00 μmol NO3 kg-1 d-1). Rates in bottles with 2 nmol Mn added were identical to those in the controls (0.57 to 1.04 μmol NO3 kg-1 d-1). Total decreases in NO3 were balanced by increases in particulate organic N. These results suggest that Fe deficiency is the primary reason that the present-day southern ocean biological pump is shut off. In contrast, iron was 50 times more abundant during the last glacial maximum; greater Fe availability may have stimulated the biological pump and contributed to the ice age drawdown of atmospheric C02. These results also imply that large-scale southern ocean Fe fertilization is feasible, at least in terms of the total amounts of Fe required; i.e., 100,000 to 500,000 tons yr-1.