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      Mechanisms of dissolved and labile particulate iron supply to shelf waters and phytoplankton blooms off South Georgia, Southern Ocean

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          Abstract

          <p><strong>Abstract.</strong> 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<span class="thinspace"></span><span class="inline-formula">∕</span><span class="thinspace"></span>Mn and Fe<span class="thinspace"></span><span class="inline-formula">∕</span><span class="thinspace"></span>Al ratios for shelf sediments and suspended particles in the water column. Less than 1<span class="thinspace"></span>% 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 0.1–44<span class="thinspace"></span><span class="inline-formula">µ</span>mol<span class="thinspace"></span>DFe<span class="thinspace"></span><span class="inline-formula">m</span><span class="inline-formula"><sup>−2</sup></span><span class="thinspace"></span>d<span class="inline-formula"><sup>−1</sup></span>. However, we estimate that only <span class="inline-formula">0.41±0.26</span><span class="thinspace"></span><span class="inline-formula">µ</span>mol<span class="thinspace"></span>DFe<span class="thinspace"></span><span class="inline-formula">m</span><span class="inline-formula"><sup>−2</sup></span><span class="thinspace"></span>d<span class="inline-formula"><sup>−1</sup></span> 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 <span class="inline-formula">6.5±8.2</span><span class="thinspace"></span><span class="inline-formula">µ</span>mol<span class="thinspace"></span><span class="inline-formula">m</span><span class="inline-formula"><sup>−2</sup></span><span class="thinspace"></span>d<span class="inline-formula"><sup>−1</sup></span> of labile particulate Fe was supplied to the surface mixed layer via faecal pellets formed by Antarctic krill (<i>Euphausia superba</i>), with a further <span class="inline-formula">1.1±2.2</span><span class="thinspace"></span><span class="inline-formula">µ</span>mol<span class="thinspace"></span>DFe<span class="thinspace"></span><span class="inline-formula">m</span><span class="inline-formula"><sup>−2</sup></span><span class="thinspace"></span>d<span class="inline-formula"><sup>−1</sup></span> 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.</p> <p>The Fe requirement of the phytoplankton blooms <span class="inline-formula">∼ 1250</span><span class="thinspace"></span><span class="inline-formula">km</span> downstream of South Georgia was estimated as <span class="inline-formula">0.33±0.11</span><span class="thinspace"></span><span class="inline-formula">µ</span>mol<span class="thinspace"></span><span class="inline-formula">m</span><span class="inline-formula"><sup>−2</sup></span><span class="thinspace"></span>d<span class="inline-formula"><sup>−1</sup></span>, with the DFe supply by horizontal/vertical mixing, deep winter mixing, and aeolian dust estimated as <span class="inline-formula">∼0.12</span><span class="thinspace"></span><span class="inline-formula">µ</span>mol<span class="thinspace"></span><span class="inline-formula">m</span><span class="inline-formula"><sup>−2</sup></span><span class="thinspace"></span>d<span class="inline-formula"><sup>−1</sup></span>. 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.</p>

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          Architecture and material properties of diatom shells provide effective mechanical protection.

          Diatoms are the major contributors to phytoplankton blooms in lakes and in the sea and hence are central in aquatic ecosystems and the global carbon cycle. All free-living diatoms differ from other phytoplankton groups in having silicified cell walls in the form of two 'shells' (the frustule) of manifold shape and intricate architecture whose function and role, if any, in contributing to the evolutionary success of diatoms is under debate. We explored the defence potential of the frustules as armour against predators by measuring their strength. Real and virtual loading tests (using calibrated glass microneedles and finite element analysis) were performed on centric and pennate diatom cells. Here we show that the frustules are remarkably strong by virtue of their architecture and the material properties of the diatom silica. We conclude that diatom frustules have evolved as mechanical protection for the cells because exceptional force is required to break them. The evolutionary arms race between diatoms and their specialized predators will have had considerable influence in structuring pelagic food webs and biogeochemical cycles.
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            Surface-water iron supplies in the Southern Ocean sustained by deep winter mixing

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              Ferritin is used for iron storage in bloom-forming marine pennate diatoms.

              Primary productivity in 30-40% of the world's oceans is limited by availability of the micronutrient iron. Regions with chronically low iron concentrations are sporadically pulsed with new iron inputs by way of dust or lateral advection from continental margins. Addition of iron to surface waters in these areas induces massive phytoplankton blooms dominated primarily by pennate diatoms. Here we provide evidence that the bloom-forming pennate diatoms Pseudo-nitzschia and Fragilariopsis use the iron-concentrating protein, ferritin, to safely store iron. Ferritin has not been reported previously in any member of the Stramenopiles, a diverse eukaryotic lineage that includes unicellular algae, macroalgae and plant parasites. Phylogenetic analyses suggest that ferritin may have arisen in this small subset of diatoms through a lateral gene transfer. The crystal structure and functional assays of recombinant ferritin derived from Pseudo-nitzschia multiseries reveal a maxi-ferritin that exhibits ferroxidase activity and binds iron. The protein is predicted to be targeted to the chloroplast to control the distribution and storage of iron for proper functioning of the photosynthetic machinery. Abundance of Pseudo-nitzschia ferritin transcripts is regulated by iron nutritional status, and is closely tied to the loss and recovery of photosynthetic competence. Enhanced iron storage with ferritin allows the oceanic diatom Pseudo-nitzschia granii to undergo several more cell divisions in the absence of iron than the comparably sized, oceanic centric diatom Thalassiosira oceanica. Ferritin in pennate diatoms probably contributes to their success in chronically low-iron regions that receive intermittent iron inputs, and provides an explanation for the importance of these organisms in regulating oceanic CO(2) over geological timescales.
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                Author and article information

                Journal
                Biogeosciences
                Biogeosciences
                Copernicus GmbH
                1726-4189
                2018
                August 22 2018
                : 15
                : 16
                : 4973-4993
                Article
                10.5194/bg-15-4973-2018
                17fcd01e-62f6-4cb4-a612-eaf826fac709
                © 2018

                https://creativecommons.org/licenses/by/4.0/

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