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The shallower oxygen-poor water masses of the ocean confine a majority of the microbial communities that can produce up to 90 % of oceanic N2. This effective N2-yielding section encloses a suspended small-particle layer, inferred from particle backscattering (bbp) measurements. It is thus hypothesized that this layer (hereafter, the bbp-layer) is linked to microbial communities involved in N2 yielding such as nitrate-reducing SAR11 as well as sulfur-oxidizing, anammox, and denitrifying bacteria – a hypothesis yet to be evaluated. Here, data collected by three BGC-Argo floats deployed in the Black Sea are used to investigate the origin of this bbp-layer. To this end, we evaluate how the key drivers of N2-yielding bacteria dynamics impact the vertical distribution of bbp and the thickness of the bbp-layer. In conjunction with published data on N2 excess, our results suggest that the bbp-layer is at least partially composed of the bacteria driving N2 yielding for three main reasons: (1) strong correlations are recorded between bbp and nitrate; (2) the top location of the bbp-layer is driven by the ventilation of oxygen-rich subsurface waters, while its thickness is modulated by the amount of nitrate available to produce N2; and (3) the maxima of both bbp and N2 excess coincide at the same isopycnals where bacteria involved in N2 yielding coexist. We thus advance that bbp and O2 can be exploited as a combined proxy to delineate the N2-yielding section of the Black Sea. This proxy can potentially contribute to refining delineation of the effective N2-yielding section of oxygen-deficient zones via data from the growing BGC-Argo float network.

Upper suboxic water masses confine a majority of the microbial communities that can produce up to 90 % of oceanic N2. This effective N2-yielding section encloses a suspended small-particle layer, inferred from particle backscattering (bbp) measurements. It is thus hypothesized that this layer (hereafter, the bbp-layer) is linked to N2-yielding microbial communities such as anammox and denitrifying bacteria – a hypothesis yet to be evaluated. Here, data collected by three BGC-Argo floats deployed in the Black Sea are used to investigate the origin of this bbp-layer. To this end, we evaluate how key drivers of anammox-denitrifying bacteria dynamics impact on the vertical distribution of bbp and the thickness of the bbp-layer. In conjunction with published data on N2 excess, our results suggest that the bbp-layer is at least partially composed of anammox-denitrifying bacteria for three main reasons: (1) strong correlations are recorded between bbp and nitrate; (2) the top location of the bbp-layer is driven by the ventilation of oxygen-rich subsurface waters, while its thickness is modulated by the amount of nitrate available to produce N2; (3) the maxima of both bbp and N2 excess coincide at the same isopycnals where denitrifying-anammox bacteria coexist. We thus advance that bbp and O2 can be exploited as a combined proxy to delineate the N2-yielding section of the Black Sea. This proxy can potentially contribute to refining delineation of the effective N2-yielding section of oxygen-deficient zones via data from the growing BGC-Argo float network.

The extracellular concentration of H2O2 in surface aquatic environments is controlled by a balance between photochemical production and the microbial synthesis of catalase and peroxidase enzymes to remove H2O2 from solution. In any kind of incubation experiment, the formation rates and equilibrium concentrations of reactive oxygen species (ROSs) such as H2O2 may be sensitive to both the experiment design, particularly to the regulation of incident light, and the abundance of different microbial groups, as both cellular H2O2 production and catalase–peroxidase enzyme production rates differ between species. Whilst there are extensive measurements of photochemical H2O2 formation rates and the distribution of H2O2 in the marine environment, it is poorly constrained how different microbial groups affect extracellular H2O2 concentrations, how comparable extracellular H2O2 concentrations within large-scale incubation experiments are to those observed in the surface-mixed layer, and to what extent a mismatch with environmentally relevant concentrations of ROS in incubations could influence biological processes differently to what would be observed in nature. Here we show that both experiment design and bacterial abundance consistently exert control on extracellular H2O2 concentrations across a range of incubation experiments in diverse marine environments. During four large-scale (>1000 L) mesocosm experiments (in Gran Canaria, the Mediterranean, Patagonia and Svalbard) most experimental factors appeared to exert only minor, or no, direct effect on H2O2 concentrations. For example, in three of four experiments where pH was manipulated to 0.4–0.5 below ambient pH, no significant change was evident in extracellular H2O2 concentrations relative to controls. An influence was sometimes inferred from zooplankton density, but not consistently between different incubation experiments, and no change in H2O2 was evident in controlled experiments using different densities of the copepod Calanus finmarchicus grazing on the diatom Skeletonema costatum (<1 % change in [H2O2] comparing copepod densities from 1 to 10 L−1). Instead, the changes in H2O2 concentration contrasting high- and low-zooplankton incubations appeared to arise from the resulting changes in bacterial activity. The correlation between bacterial abundance and extracellular H2O2 was stronger in some incubations than others (R2 range 0.09 to 0.55), yet high bacterial densities were consistently associated with low H2O2. Nonetheless, the main control on H2O2 concentrations during incubation experiments relative to those in ambient, unenclosed waters was the regulation of incident light. In an open (lidless) mesocosm experiment in Gran Canaria, H2O2 was persistently elevated (2–6-fold) above ambient concentrations; whereas using closed high-density polyethylene mesocosms in Crete, Svalbard and Patagonia H2O2 within incubations was always reduced (median 10 %–90 %) relative to ambient waters.

The speciation of dissolved iron (DFe) in the ocean is widely assumed to consist almost exclusively of Fe(III)-ligand complexes. Yet in most aqueous environments a poorly defined fraction of DFe also exists as Fe(II), the speciation of which is uncertain. Here we deploy flow injection analysis to measure in situ Fe(II) concentrations during a series of mesocosm/microcosm/multistressor experiments in coastal environments in addition to the decay rate of this Fe(II) when moved into the dark. During five mesocosm/microcosm/multistressor experiments in Svalbard and Patagonia, where dissolved (0.2 µm) Fe and Fe(II) were quantified simultaneously, Fe(II) constituted 24 %–65 % of DFe, suggesting that Fe(II) was a large fraction of the DFe pool. When this Fe(II) was allowed to decay in the dark, the vast majority of measured oxidation rate constants were less than calculated constants derived from ambient temperature, salinity, pH, and dissolved O2. The oxidation rates of Fe(II) spikes added to Atlantic seawater more closely matched calculated rate constants. The difference between observed and theoretical decay rates in Svalbard and Patagonia was most pronounced at Fe(II) concentrations <2 nM, suggesting that the effect may have arisen from organic Fe(II) ligands. This apparent enhancement of Fe(II) stability under post-bloom conditions and the existence of such a high fraction of DFe as Fe(II) challenge the assumption that DFe speciation in coastal seawater is dominated by ligand bound-Fe(III) species.

Bottom trawling in the deep sea is one of the main drivers of sediment resuspension, eroding the seafloor and altering the content and composition of sedimentary organic matter (OM). The physical and biogeochemical impacts of bottom trawling were studied on the continental slope of the Gulf of Castellammare, Sicily (southwestern Mediterranean), through the analysis of two triplicate sediment cores collected at trawled and untrawled sites (∼550 m water depth) during the summer of 2016. Geochemical and sedimentological parameters (excess 210Pb, excess 234Th, 137Cs, dry bulk density, and grain size), elemental (organic carbon and nitrogen) and biochemical composition of sedimentary OM (proteins, carbohydrates, lipids), as well as its freshness (phytopigments) and degradation rates were determined in both coring locations. The untrawled site had a sedimentation rate of 0.15 cm yr−1 and presented a 6 cm thick surface mixed layer that contained siltier sediment with low excess 210Pb concentrations, possibly resulting from the resuspension, posterior advection, and eventual deposition of coarser and older sediment from adjacent trawling grounds. In contrast, the trawled site was eroded and presented compacted century-old sediment highly depleted in OM components, which were between 20 % and 60 % lower than those in the untrawled site. However, the upper 2 cm of the trawled site consisted of recently accumulated sediments enriched in excess 234Th, excess 210Pb, and phytopigments, while OM contents were similar to those from the untrawled core. This fresh sediment supported protein turnover rates of 0.025 d−1, which doubled those quantified in surface sediments of the untrawled site. The enhancement of remineralization rates in surface sediment of the trawled site was associated with the arrival of fresh particles on a chronically trawled deep-sea region that is generally deprived of OM. We conclude that the detrimental effects of bottom trawling can be temporarily and partially abated by the arrival of fresh and nutritionally rich OM, which stimulate the response of benthic communities. However, these ephemeral deposits are likely to be swiftly eroded due to the high trawling frequency over fishing grounds, highlighting the importance of establishing science-based management strategies to mitigate the impacts of bottom trawling.

The North Atlantic Ocean is a major sink region for atmospheric CO2 and contributes to the storage of anthropogenic carbon (Cant). While there is general agreement that the intensity of the meridional overturning circulation (MOC) modulates uptake, transport and storage of Cant in the North Atlantic Subpolar Ocean, processes controlling their recent variability and evolution over the 21st century remain uncertain. This study investigates the relationship between transport, air–sea flux and storage rate of Cant in the North Atlantic Subpolar Ocean over the past 53 years. Its relies on the combined analysis of a multiannual in situ data set and outputs from a global biogeochemical ocean general circulation model (NEMO–PISCES) at 1∕2∘ spatial resolution forced by an atmospheric reanalysis. Despite an underestimation of Cant transport and an overestimation of anthropogenic air–sea CO2 flux in the model, the interannual variability of the regional Cant storage rate and its driving processes were well simulated by the model. Analysis of the multi-decadal simulation revealed that the MOC intensity variability was the major driver of the Cant transport variability at 25 and 36∘ N, but not at OVIDE. At the subpolar OVIDE section, the interannual variability of Cant transport was controlled by the accumulation of Cant in the MOC upper limb. At multi-decadal timescales, long-term changes in the North Atlantic storage rate of Cant were driven by the increase in air–sea fluxes of anthropogenic CO2. North Atlantic Central Water played a key role for storing Cant in the upper layer of the subtropical region and for supplying Cant to Intermediate Water and North Atlantic Deep Water. The transfer of Cant from surface to deep waters occurred mainly north of the OVIDE section. Most of the Cant transferred to the deep ocean was stored in the subpolar region, while the remainder was exported to the subtropical gyre within the lower MOC.

A significant decrease of dissolved iron (DFe) concentration has been observed after dust addition into mesocosms during the DUst experiment in a low Nutrient low chlorophyll Ecosystem (DUNE), carried out in the summer of 2008. Due to low biological productivity at the experiment site, biological consumption of iron can not explain the magnitude of DFe decrease. To understand processes regulating the observed DFe variation, we simulated the experiment using a one-dimensional model of the Fe biogeochemical cycle, coupled with a simple ecosystem model. Different size classes of particles and particle aggregation are taken into account to describe the particle dynamics. DFe concentration is regulated in the model by dissolution from dust particles and adsorption onto particle surfaces, biological uptake, and photochemical mobilisation of particulate iron. The model reproduces the observed DFe decrease after dust addition well. This is essentially explained by particle adsorption and particle aggregation that produces a high export within the first 24 h. The estimated particle adsorption rates range between the measured adsorption rates of soluble iron and those of colloidal iron, indicating both processes controlling the DFe removal during the experiment. A dissolution timescale of 3 days is used in the model, instead of an instantaneous dissolution, underlining the importance of dissolution kinetics on the short-term impact of dust deposition on seawater DFe. Sensitivity studies reveal that initial DFe concentration before dust addition was crucial for the net impact of dust addition on DFe during the DUNE experiment. Based on the balance between abiotic sinks and sources of DFe, a critical DFe concentration has been defined, above which dust deposition acts as a net sink of DFe, rather than a source. Taking into account the role of excess iron binding ligands and biotic processes, the critical DFe concentration might be applied to explain the short-term variability of DFe after natural dust deposition in various different ocean regions.

The boron isotopic composition (δ11B) of marine carbonates (e.g. corals) is increasingly utilised as a proxy for paleo-pH, with the strong correlation between δ11B of marine calcifiers and seawater pH now well documented. However, the potential roles of other environmental parameters that may also influence both the boron isotopic composition and boron concentration into coral aragonite are poorly known. To overcome this, the tropical scleractinian coral Acropora sp. was cultured under 3 different temperatures (22, 25 and 28 °C) and two light conditions (200 and 400 μmol photon m−2 s−1). The δ11B indicates an increase in internal pH that is dependent on the light conditions. Changes in light intensities from 200 to 400 μmol photon m−2 s−1 seem to indicate an apparent decrease in pH at the site of calcification, contrary to what is expected in most models of light-enhanced calcification. Thus, variations in light conditions chosen to mimic average annual variations of the natural environments where Acropora sp. colonies can be found could bias pH reconstructions by about 0.05 units. For both light conditions, a significant impact of temperature on δ11B can be observed between 22 and 25 °C, corresponding to an increase of about 0.02 pH-units, while no further δ11B increase can be observed from 25 to 28 °C. This non-linear temperature effect complicates the determination of a correction factor. B / Ca ratios decrease with increasing light, consistent with the decrease in pH at the site of calcification under enhanced light intensities. When all the other parameters are constant, boron concentrations in Acropora sp. increase with increasing temperatures and increasing carbonate ion concentrations. These observations contradict previous studies where B / Ca in corals was found to vary inversely with temperature, suggesting that the controlling factors driving boron concentrations have not yet been adequately identified and might be influenced by other environmental variables and/or species-specific responses.

The Eastern Boundary Upwelling Systems (EBUS) contribute to one fifth of the global catches in the ocean. Often associated with Oxygen Minimum Zones (OMZs), EBUS represent key regions for the oceanic nitrogen (N) cycle. Important bioavailable N loss due to denitrification and anammox processes as well as greenhouse gas emissions (e.g, N2O) occur also in these EBUS. However, their dynamics are currently crudely represented in global models. In the climate change context, improving our capability to properly represent these areas is crucial due to anticipated changes in the winds, productivity, and oxygen content. We developed a biogeochemical model (BioEBUS) taking into account the main processes linked with EBUS and associated OMZs. We implemented this model in a 3-D realistic coupled physical/biogeochemical configuration in the Namibian upwelling system (northern Benguela) using the high-resolution hydrodynamic ROMS model. We present here a validation using in situ and satellite data as well as diagnostic metrics and sensitivity analyses of key parameters and N2O parameterizations. The impact of parameter values on the OMZ off Namibia, on N loss, and on N2O concentrations and emissions is detailed. The model realistically reproduces the vertical distribution and seasonal cycle of observed oxygen, nitrate, and chlorophyll a concentrations, and the rates of microbial processes (e.g, NH4+ and NO2− oxidation, NO3− reduction, and anammox) as well. Based on our sensitivity analyses, biogeochemical parameter values associated with organic matter decomposition, vertical sinking, and nitrification play a key role for the low-oxygen water content, N loss, and N2O concentrations in the OMZ. Moreover, the explicit parameterization of both steps of nitrification, ammonium oxidation to nitrate with nitrite as an explicit intermediate, is necessary to improve the representation of microbial activity linked with the OMZ. The simulated minimum oxygen concentrations are driven by the poleward meridional advection of oxygen-depleted waters offshore of a 300 m isobath and by the biogeochemical activity inshore of this isobath, highlighting a spatial shift of dominant processes maintaining the minimum oxygen concentrations off Namibia. In the OMZ off Namibia, the magnitude of N2O outgassing and of N loss is comparable. Anammox contributes to about 20% of total N loss, an estimate lower than currently assumed (up to 50%) for the global ocean.

Biological communities populating the Mediterranean Sea, which is situated at the northern boundary of the subtropics, are often claimed to be particularly affected by global warming. This is indicated, for instance, by the introduction of (sub)tropical species of fish or invertebrates that can displace local species. This raises the question of whether microbial communities are similarly affected, especially in the Levantine basin where sea surface temperatures have significantly risen over the last 25 years (0.50 ± 0.11 °C in average per decade, P Prochlorococcus and Synechococcus, was examined during two cruises through both eastern and western Mediterranean Sea basins held in September 1999 (PROSOPE cruise) and in June–July 2008 (BOUM cruise). Diversity was studied using dot blot hybridization with clade-specific 16S rRNA oligonucleotide probes and/or clone libraries of the 16S-23S ribosomal DNA Internal Transcribed Spacer (ITS) region, with a focus on the abundance of clades that may constitute bioindicators of warm waters. During both cruises, the dominant Prochlorococcus clade in the upper mixed layer at all stations was HLI, a clade typical of temperate waters, whereas the HLII clade, the dominant group in (sub)tropical waters, was only present at very low concentrations. The Synechococcus community was dominated by clades I, III and IV in the northwestern waters of the Gulf of Lions and by clade III and groups genetically related to clades WPC1 and VI in the rest of the Mediterranean Sea. In contrast, only a few sequences of clade II, a group typical of warm waters, were observed. These data indicate that local cyanobacterial populations have not yet been displaced by their (sub)tropical counterparts.