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Sediments are considered to be refractory materials with limited influences on dissolved iron (dFe) pool in the ocean. However, recent field observations and laboratory experiments suggest that iron released from resuspended sediment particles and transported from continental margins is prone to fertilize large areas of the world ocean. Here we conducted a dissolution experiment to quantify the amount of dFe released from two types of resuspended sediments (silicate and calcite-rich) to open ocean surface seawater under two temperatures (5 and 15 °C). We followed pH, dissolved oxygen (dO2), phosphate, silicate, dissolved Fe and Mn concentrations (dFe and dMn), and bacterial abundance over 250 days. Extremely low and undetectable phosphate concentrations (<50 pmol kg−1) were measured throughout the duration of the experiment, causing limited bacteria growth and stable pH and dissolved O 2 concentrations under all conditions. Silicate and dFe concentrations increased through time and high temperature (15 °C) induced more iron dissolution from the two sediments than low temperature (5 °C). Temperature had no effect on the dissolution of Mn. Our results further show that Fe and Mn are not released concurrently from the sediment source and that their distribution can be very different. Scavenging of Fe likely caused a decrease of dFe observed during the experiment, which was probably linked to the formation of Mn oxides. We also observed elevated dissolved Fe isotope ratios after dissolution, around +0.16 to +0.27‰. Isotopically heavy Fe was released from sediments to the dissolved pool during the dissolution but no difference in Fe isotope ratios was observed between the two temperature conditions. The Fe isotope fractionation can likely be attributed to ligand complexation and scavenging of Fe. These two mechanisms can be important factors not only in controlling the amount of Fe released from sediments but also in fractionating Fe i

Iron is an essential nutrient that regulates productivity in ~30% of the ocean. Compared with deep (>2000 meter) hydrothermal activity at mid-ocean ridges that provide iron to the ocean’s interior, shallow (<500 meter) hydrothermal fluids are likely to influence the surface’s ecosystem. However, their effect is unknown. In this work, we show that fluids emitted along the Tonga volcanic arc (South Pacific) have a substantial impact on iron concentrations in the photic layer through vertical diffusion. This enrichment stimulates biological activity, resulting in an extensive patch of chlorophyll (360,000 square kilometers). Diazotroph activity is two to eight times higher and carbon export fluxes are two to three times higher in iron-enriched waters than in adjacent unfertilized waters. Such findings reveal a previously undescribed mechanism of natural iron fertilization in the ocean that fuels regional hotspot sinks for atmospheric CO2.

A number of trace metals play essential roles in marine ecosystem structure and biological productivity. Until recently, it has been argued that phytoplankton access primarily dissolved iron, while particulate iron was considered a refractory material with little use biologically and limited interaction with the dissolved pool. In order to assess the transfer mechanisms between sediment-sourced particulate trace metals and the dissolved pool, we conducted a 14-month incubation that reacted resuspended sediments with natural seawater, both originating from the Kerguelen area (KEOPS cruises; Southern Ocean), in the dark, and at concentrations replicating natural conditions. Three types of sediments were investigated (named BioSi, BioSi + Ca, and Basalt), mostly composed of (i) biogenic silica (bSiO2), (ii) bSiO2 and calcite, and (iii) basaltic fragments, respectively. The release of dissolved silicon (dSi), iron (dFe) and manganese (dMn) was monitored regularly throughout the incubation, as well as living bacteria density and Fe organic ligands. Depending on the origin and composition of the sediment, unique dFe and dMn fluxes were observed, including a strong decoupling between dFe and dMn. The basaltic sediment released up to 1.09 ± 0.04 nmol L−1 of dFe and 0.28 ± 0.09 nmol L−1 of dMn, while the biogenic sediments released a higher 3.91 ± 0.04 nmol L−1 and 8.03 ± 0.42 nmol L−1 of dFe and dMn, respectively. Several factors influencing the release and removal of dFe and dMn were discernable at the temporal sampling resolution of the incubation, including the structural composition of the sediment, bacterial abundance, and the formation of manganese oxides. The regular sampling over short timescales and the extended sampling over one year proved to be critical to constrain the processes and exchanges that govern the contribution of the particulate to the dissolved pools. Overall, this incubation provides a strong basis for reassessing the role of resuspended sedimentar

The dissolved iron supply controls half of the oceans' primary productivity. Resupply by the remineralization of sinking particles, and subsequent vertical mixing, largely sustains this productivity. However, our understanding of the drivers of dissolved iron resupply, and their influence on its vertical distribution across the oceans, is still limited due to sparse observations. There is a lack of empirical evidence as to what controls the subsurface iron remineralization due to difficulties in studying mesopelagic biogeochemistry. Here we present estimates of particulate transformations to dissolved iron, concurrent oxygen consumption and iron-binding ligand replenishment based on in situ mesopelagic experiments. Dissolved iron regeneration efficiencies (that is, replenishment over oxygen consumption) were 10- to 100-fold higher in low-dust subantarctic waters relative to higher-dust Mediterranean sites. Regeneration efficiencies are heavily influenced by particle composition. Their make-up dictates ligand release, controls scavenging, modulates ballasting and may lead to the differential remineralization of biogenic versus lithogenic iron. At high-dust sites, these processes together increase the iron remineralization length scale. Modelling reveals that in oceanic regions near deserts, enhanced lithogenic fluxes deepen the ferricline, which alter the vertical patterns of dissolved iron replenishment, and set its redistribution at the global scale. Such wide-ranging regeneration efficiencies drive different vertical patterns in dissolved iron replenishment across oceanic provinces.

© 2019 Elsevier B.V. The dinoflagellate Alexandrium minutum produces toxic compounds, including paralytic shellfish toxins, but also some unknown extracellular toxins. Although copper (Cu) is an essential element, it can impair microalgal physiology and increase their toxic potency. This study investigated the effect of different concentrations of dissolved Cu (7 nM, 79 nM and 164 nM) on A. minutum allelochemical potency, here defined as negative effects of a protist on competing protists through the release of chemicals. This was studied in relation to its physiology. The effects of Cu were assessed on A. minutum growth, reactive oxygen species level, photosynthesis proxies, lipid metabolism, exudation of dissolved organic compounds, allelochemical potency and on the associate free bacterial community of A. minutum. Only the highest Cu exposure (164 nM) inhibited and delayed the growth of A. minutum, and only in this treatment did the allelochemical potency significantly increase, when the dissolved Cu concentration was still toxic. Within the first 7 days of the high Cu treatment, the physiology of A. minutum was severely impaired with decreased growth and photosynthesis, and increased stress responses and free bacterial density per algal cell. After 15 days, A. minutum partially recovered from Cu stress as highlighted by the growth rate, reactive oxygen species level and photosystem II yields. This recovery could be attributed to the apparent decrease in background dissolved Cu concentration to a non-toxic level, suggesting that the release of exudates may have partially decreased the bioavailable Cu fraction. Overall, A. minutum appeared quite tolerant to Cu, and this work suggests that the modifications in the physiology and in the exudates help the algae to cope with Cu exposure. Moreover, this study shows the complex interplay between abiotic and biotic factors that can influence the dynamic of A. minutum blooms. Modulation in allelochemical potency of A. min

© 2018 Elsevier Ltd Harmful microalgal blooms are a threat to aquatic organisms, ecosystems and human health. Toxic dinoflagellates of the genus Alexandrium are known to produce paralytic shellfish toxins and to release bioactive extracellular compounds (BECs) with potent cytotoxic, hemolytic, ichtyotoxic and allelopathic activity. Negative allelochemical interactions refer to the chemicals that are released by the genus Alexandrium and that induce adverse effects on the physiology of co-occurring protists and predators. Releasing BECs gives the donor a competitive advantage that may help to form dense toxic blooms of phytoplankton. However BECs released by Alexandrium minutum are uncharacterized and it is impossible to quantify them using classical chemical methods. Allelochemical interactions are usually quantified through population growth inhibition or lytic-activity based bioassays using a secondary target organism. However these bioassays require time (for growth or microalgal counts) and/or are based on lethal effects. The use of pulse amplitude modulation (PAM) fluorometry has been widely used to assess the impact of environmental stressors on phytoplankton but rarely for allelochemical interactions. Here we evaluated the use of PAM and propose a rapid chlorophyll fluorescence based bioassay to quantify allelochemical BECs released from Alexandrium minutum. We used the ubiquitous diatom Chaetoceros muelleri as a target species. The bioassay, based on sub-lethal effects, quantifies allelochemical activity from different samples (filtrates, extracts in seawater) within a short period of time (2 h). This rapid bioassay will help investigate the role of allelochemical interactions in Alexandrium bloom establishment. It will also further our understanding of the potential relationship between allelochemical activities and other cytotoxic activities from BECs. While this bioassay was developed for the species A. minutum, it may be applicable to other species produ

© 2018 Elsevier B.V. Allelochemical interactions are likely to be a contributing factor explaining the success of large blooms of the harmful marine dinoflagellate Alexandrium, however, the physiological mechanisms of allelochemical interactions remain poorly described. Here we investigated the sub-lethal effects (on an hourly scale) of a filtrate containing allelochemicals from Alexandrium minutum on the physiology of the common diatom Chaetoceros muelleri. The filtrate induced deleterious effects to the diatom physiology within only 30 min of exposure. Esterase activity and photosynthesis were drastically inhibited, with up to 34% of the population being metabolically inactive and up to 30% reduction in photosystem II quantum yield when exposed to the filtrate. In addition, intracellular reactive oxygen species increased by 26% in response to allelochemical exposure. C. muelleri pigment and lipid analyses indicated that the photosystem II was inhibited, with photoinhibition-like responses (activation of xanthophyll cycles, and changes in associated lipids) upregulated to mitigate the toxic effects of allelochemicals. Changes in the proportions of membrane lipid classes and increased membrane fatty acids saturation by 9% may be an attempt to maintain membrane integrity and associated enzyme activity, or could be the result of deleterious effects on membranes. An 8% decrease in cellular storage lipids (triglycerides) revealed a mobilization of energy suggesting an energetic cost for the diatom to counteract the allelochemical effects. We hypothesize that the rapid alteration of physiological functions such as photosynthesis and some enzymatic activities may result from direct damage on external membranes. Overall this study describes the sub-lethal mechanisms and provides useful biomarkers to understand the role of allelochemical interactions and associated ecological processes in structuring plankton communities.

The Labrador Current is an important conduit of freshwater from the Arctic to the interior North Atlantic subpolar gyre. Here, we investigate the spatial variability of the freshwater sources over the southern Labrador shelf and slope during May-June 2014. Using measurements of seawater properties such as temperature, salinity, nutrients and oxygen isotopic composition, we estimate the respective contributions of saline water of Atlantic and Pacific origins, of brines released during sea ice formation, and of freshwater from sea ice melt and meteoric water origins. On the southern Labrador shelf, we find a large brine signal and Pacific Water influence indicating a large contribution of water from the Canadian Arctic. The brine signal implies that more than 4 m of sea ice formed upstream, either in the Arctic or in Baffin Bay and the northern Labrador Sea. Over the mid-shelf and slope at 52°N, we find a stronger influence of slope water from the West Greenland Current with a smaller contribution of Pacific water and no brine signal. Thus, there is advection of water from the slope region to the mid-shelf between 55°N and 52°N. Very fresh water with high meteoric content is found close to the coast in June 2014. Observations from 1995 and 2008 suggest a higher fraction of brine and Pacific water on the shelf compared to that observed in 2014.

We will present the combined results of the French GEOTRACES GEOVIDE cruise in the North Atlantic Ocean and the 2015 German GEOTRACES cruise TransArc II in the central Arctic Ocean. Research vessel "Pourquoi pas?" sailed on May 15th from Lisbon to Greenland to arrive in Newfoundland on June 30th 2014, and icebreaker "Polarstern" sailed on August 17th from Tromsoe to explore the Nansen, the Amundsen and the Makarov basins, to arrive in Bremerhaven on October 15th 2015. Total mercury was sampled using ultra-trace clean rosettes and determined on board. In the Atlantic Ocean, surface waters of the Gulf Stream are cooled down as they travel north, and mix at the same time with waters exiting the Arctic Ocean via Fram Strait. These cool and dense surface waters dive to depth in the Greenland and Labrador seas. The North Atlantic Ocean predominantly receives Hg via atmospheric deposition from Europe and North America where industrial Hg emissions peaked in the 1970s. The Hg inputs to the Arctic Ocean are less well-constrained if not unknown. The current debate opposes a primary atmospheric with a river-dominated scenario. We find consistent surface depleted profiles in the North Atlantic Ocean, while we exclusively observe surface enrichments in the Arctic Ocean, at all sampling stations. We will make use of the combined data sets of both cruises to investigate how climate may impact Hg marine biogeochemical cycle, how anthropogenic Hg makes its way into the deep ocean and whether the temporal evolution of emissions is traceable in water masses of different ages. We will also put our new observations in context with recent numerical model evaluations.

Iron availability in the Southern Ocean controls phytoplankton growth, community composition and the uptake of atmospheric CO2 by the biological pump. The KEOPS-2 experiment took place around the Kerguelen plateau in the Indian sector of the Southern Ocean, a region naturally fertilised with iron at the scale of hundreds to thousands of square kilometres, producing a mosaic of spring blooms which showed distinct biological and biogeochemical responses to fertilisation. This paper presents biogeochemical iron budgets (incorporating vertical and lateral supply, internal cycling, and sinks) for three contrasting sites: an upstream high-nutrient low-chlorophyll reference, over the plateau, and in the o�shore plume east of Kerguelen Island. These budgets show that distinct regional environments driven by complex circulation and transport pathways are responsible for di�erences in the mode and strength of iron supply, with vertical supply dominant on the plateau and lateral supply dominant in the plume. Iron supply from “new” sources to surface waters of the plume was double that above the plateau and 20 times greater than at the reference site, whilst iron demand (measured by cellular uptake) in the plume was similar to the plateau but 40 times greater than the reference. “Recycled” iron supply by bacterial regeneration and zooplankton grazing was a relative minor component at all sites (< 8% of “new” supply), in contrast to earlier findings from other biogeochemical iron budgets in the Southern Ocean. Over the plateau, a particulate iron dissolution term of 2.5% was invoked to balance the budget; this approximately doubled the standing stock of dissolved iron in the mixed layer. The exchange of iron between dissolved, biogenic and lithogenic particulate pools was highly dynamic in time and space, resulting in a decoupling of iron supply and carbon export and, importantly, controlling the effi�ciency of fertilisation.

The KEOPS2 project aims to elucidate the role of natural Fe fertilisation on biogeochemical cycles and ecosystem functioning, including quantifying the sources and processes by which iron is delivered in the vicinity of the Kerguelen Archipelago, Southern Ocean. The KEOPS2 process study used an upstream high-nutrient, low-chlorophyll (HNLC), deep water (2500 m), reference station to compare with a shallow (500 m), strongly fertilised plateau station and continued the observations to a downstream, bathymetrically trapped recirculation of the Polar Front where eddies commonly form and persist for hundreds of kilometres into the Southern Ocean. Over the Kerguelen Plateau, mean particulate (1-53 mu m) Fe and Al concentrations (pFe = 13.4 nM, pAl = 25.2 nM) were more than 20-fold higher than at an offshore (lower-productivity) reference station (pFe = 0.53 nM, pAl = 0.83 nM). In comparison, over the plateau dissolved Fe levels were only elevated by a factor of similar to 2. Over the Kerguelen Plateau, ratios of pMn / pAl and pFe / pAl resemble basalt, likely originating from glacial/fluvial inputs into shallow coastal waters. In downstream, offshore deep-waters, higher pFe / pAl, and pMn / pAl ratios were observed, suggesting loss of lithogenic material accompanied by retention of pFe and pMn. Biological uptake of dissolved Fe and Mn and conversion into the biogenic particulate fraction or aggregation of particulate metals onto bioaggregates also increased these ratios further in surface waters as the bloom developed within the recirculation structure. While resuspension of shelf sediments is likely to be one of the important mechanisms of Fe fertilisation over the plateau, fluvial and glacial sources appear to be important to areas downstream of the island. Vertical profiles within an offshore recirculation feature associated with the Polar Front show pFe and pMn levels that were 6-fold and 3.5-fold lower, respectively, than over the plateau in surface waters, though st

The concentrations of greenhouse gases (GHGs), such as nitrous oxide (N2O) and methane (CH4), were measured in the Kerguelen Plateau region (KPR). The KPR is affected by an annual microalgal bloom caused by natural iron fertilization, and this may stimulate the microbes involved in GHG cycling. This study was carried out during the KEOPS 2 cruise during the austral spring of 2011. Oceanographic variables, including N2O and CH4, were sampled (from the surface to 500 m depth) in two transects along and across the KRP, the north south (TNS) transect (46 degrees-51 degrees S, similar to 72 degrees E) and the east west (TEW) transect (66 degrees -75 degrees E, similar to 48.3 degrees S), both associated with the presence of a plateau, polar front (PF) and other mesoscale features. The 'FEW presented N2O levels ranging from equilibrium (105 %) to slightly supersaturated (120%) with respect to the atmosphere, whereas CH4 levels fluctuated dramatically, being highly supersaturated (120-970%) in areas close to the coastal waters of the Kerguelen Islands and in the PF. The TNS showed a more homogenous distribution for both gases, with N2O and CH4 levels ranging from 88 to 171 % and 45 to 666% saturation, respectively. Surface CH4 peaked at southeastern stations of the KPR (A3 stations), where a phytoplankton bloom was observed. Both gases responded significantly, but in contrasting ways (CH4 accumulation and N2O de- pletion), to the patchy distribution of chlorophyll a. This seems to be associated to the supply of iron from various sources. Air sea fluxes for N2O (from -10.5 to 8.65, mean 1.25 +/- 4.04 mu mol m(-2) d(-1)) and for CH4 (from 0.32 to 38.1, mean 10.01 9.97 mmol(-2) d(-1)) indicated that the KPR is both a sink and a source for N2O, as well as a considerable and variable source of CH4. This appears to be associated with biological factors, as well as the transport of water masses enriched with Fe and CH4 from the coastal area of the Kerguelen Islands. These previou