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The eastern tropical North Atlantic (ETNA) is characterized by a highly productive coastal upwelling system and a moderate oxygen minimum zone with lowest open-ocean oxygen (O2/concentrations of approximately 40 μmol kg-1. The recent discovery of re-occurring mesoscale eddies with close to anoxic O2 concentrations (< 1 μmol kg-1/ located just below the mixed layer has challenged our understanding of O2 distribution and biogeochemical processes in this area. Here, we present the first microbial community study from a deoxygenated anticyclonic modewater eddy in the open waters of the ETNA. In the eddy, we observed significantly lower bacterial diversity compared to surrounding waters, along with a significant community shift. We detected enhanced primary productivity in the surface layer of the eddy indicated by elevated chlorophyll concentrations and carbon uptake rates of up to three times as high as in surrounding waters. Carbon uptake rates below the euphotic zone correlated to the presence of a specific high-light ecotype of Prochlorococcus, which is usually underrepresented in the ETNA. Our data indicate that high primary production in the eddy fuels export production and supports enhanced respiration in a specific microbial community at shallow depths, below the mixed-layer base. The transcription of the key functional marker gene for dentrification, nirS, further indicated a potential for nitrogen loss processes in O2-depleted core waters of the eddy. Dentrification is usually absent from the open ETNA waters. In light of future projected ocean deoxygenation, our results show that even distinct events of anoxia have the potential to alter microbial community structure with critical impacts on primary productivity and biogeochemical processes of oceanic water bodies. [ABSTRACT FROM AUTHOR]

Ocean deoxygenation due to climate change may alter redox-sensitive nutrient cycles in the marine environment. The productive eastern tropical North Atlantic (ETNA) upwelling region may be particularly affected when the relatively moderate oxygen minimum zone (OMZ) deoxygenates further and microbially-driven nitrogen (N) loss processes are promoted. Consequently, water masses with a low N : P ratio could reach the euphotic layer, possibly influencing primary production in those waters. Previous mesocosm studies in the oligotrophic Atlantic Ocean identified N availability as controlling of primary production, while a possible co-limitation of nitrate and phosphate (P) could not be ruled out. To better understand the impact of changing N : P ratios on primary production and on N2 fixation in the ETNA surface ocean, we conducted land-based mesocosm experiments with natural plankton communities and applied a broad range of N : P ratios (2.67-48). Silicate was supplied at 15 µmol L-1 in all mesocosms. We monitored nutrient drawdown, bloom formation, biomass build up and diazotrophic feedback in response to variable nutrient stoichiometry. Our results confirmed N to be limiting to primary production. We found that excess P was channeled through particulate organic matter (POP) into the dissolved organic matter (DOP) pool. In mesocosms with low P availability, DOP was utilized while N2 fixation increased, suggesting a link between those two processes. Interestingly this observation was most pronounced in mesocosms where inorganic N was still available, indicating that bioavailable N does not necessarily has to have a negative impact on N2 fixation. We observed a shift from a mixed cyanobacterial/proteobacterial dominated active diazotrophic community towards diazotrophic diatom symbionts of the Richelia-Rhizosolenia symbiosis. We hypothesize that a potential change in nutrient stoichiometry in the ETNA might lead to a general shift within the diazotrophic community, potentially modifying primary productivity. [ABSTRACT FROM AUTHOR]

In open-ocean regions, as is the Eastern Tropical North Atlantic (ETNA), pelagic production is the main source of dissolved organic matter (DOM) and is affected by dissolved inorganic nitrogen (DIN) and phosphorus (DIP) concentrations. Changes in pelagic production under nutrient amendments were shown to also modify DOM quantity and quality. However, little information is available about the effects of nutrient variability on chromophoric (CDOM) and fluorescent (FDOM) DOM dynamics. Here we present results from two mesocosm experiments ("Varied P" and "Varied N") conducted with a natural plankton community from the ETNA, where the effects of DIP and DIN supply onDOM optical properties were studied. CDOM accumulated proportionally to phytoplankton biomass during the experiments. Spectral slope (S/ decreased over time indicating accumulation of high molecular weight DOM. In Varied N, an additional CDOM portion, as a result of bacterial DOM reworking, was determined. It increased the CDOM fraction in DOC proportionally to the supplied DIN. The humic-like FDOM component (Comp.1) was produced by bacteria proportionally to DIN supply. The protein-like FDOM component (Comp.2) was released irrespectively to phytoplankton or bacterial biomass, but depended on DIP and DIN concentrations. Under high DIN supply, Comp.2 was removed by bacterial reworking, leading to an accumulation of humiclike Comp.1. No influence of nutrient availability on amino acid-like FDOM component in peptide form (Comp.3) was observed. Comp.3 potentially acted as an intermediate product during formation or degradation of Comp.2. Our findings suggest that changes in nutrient concentrations may lead to substantial responses in the quantity and quality of optically active DOM and, therefore, might bias results of the applied in situ optical techniques for an estimation of DOC concentrations in open-ocean regions. [ABSTRACT FROM AUTHOR]

Calanoid copepods and euphausiids are key components of marine zooplankton communities worldwide. Most euphausiids and several copepod species perform diel vertical migrations (DVMs) that contribute to the export of particulate and dissolved matter to midwater depths. In vast areas of the global ocean, and in particular in the eastern tropical Atlantic and Pacific, the daytime distribution depth of many migrating organisms corresponds to the core of the oxygen minimum zone (OMZ). At depth, the animals experience reduced temperature and oxygen partial pressure (pO2) and an increased carbon dioxide partial pressure (pCO2) compared to their near-surface nighttime habitat. Although it is well known that low oxygen levels can inhibit respiratory activity, the respiration response of tropical copepods and euphausiids to relevant pCO2, pO2 and temperature conditions remains poorly parameterized. Further, the regulation of ammonium excretion at OMZ conditions is generally not well understood. It was recently estimated that DVM-mediated ammonium supply considerably fuels bacterial anaerobic ammonium oxidation - a major loss process for fixed nitrogen in the ocean. These estimates were based on the implicit assumption that hypoxia or anoxia in combination with hypercapnia (elevated pCO2) does not result in a downregulation of ammonium excretion. Here we show that exposure to OMZ conditions can result in strong depression of respiration and ammonium excretion in calanoid copepods and euphausiids from the Eastern Tropical North Atlantic and the Eastern Tropical South Pacific. These physiological responses need to be taken into account when estimating DVM-mediated fluxes of carbon and nitrogen into OMZs. [ABSTRACT FROM AUTHOR]

Gel particles such as the polysaccharidic transparent exopolymer particles (TEP) and the proteinaceous Coomassie stainable particles (CSP) play an important role in marine biogeochemical and ecological processes like particle aggregation and export, or microbial nutrition and growth. So far, effects of nutrient availability or of changes in nutrient ratios on gel particle production and fate are not well understood. The tropical ocean includes large oxygen minimum zones, where nitrogen losses due to anaerobic microbial activity result in a lower supply of nitrate relative to phosphate to the euphotic zone. Here, we report of two series of mesocosm experiments that were conducted with natural plankton communities collected from the eastern tropical North Atlantic (ETNA) close to Cape Verde in October 2012. The experiments were performed to investigate how different phosphate (experiment 1, Varied P: 0.15-1.58 µmol L-1/ or nitrate (experiment 2, Varied N: 1.9-21.9 µmol L-1/ concentrations affect the abundance and size distribution of TEP and CSP. In the days until the bloom peak was reached, a positive correlation between gel particle abundance and Chl a concentration was determined, linking the release of dissolved gel precursors and the subsequent formation of gel particles to autotrophic production. After the bloom peak, gel particle abundance remained stable or even increased, implying a continued partitioning of dissolved into particulate organic matter after biomass production itself ceased. During both experiments, differences between TEP and CSP dynamics were observed; TEP were generally more abundant than CSP. Changes in size distribution indicated aggregation of TEP after the bloom, while newly formed CSP decomposed. Abundance of gel particles clearly increased with nitrate concentration during the second experiment, suggesting that changes in [DIN] : [DIP] ratios can affect gel particle formation with potential consequences for carbon and nitrogen cycling as well as food web dynamics in tropical ecosystems. [ABSTRACT FROM AUTHOR]

The Eastern Tropical North Atlantic (ETNA) is an open ocean region with little input of terrestrial dissolved organic matter (DOM), suggesting that pelagic production has to be the main source of DOM. Inorganic nitrogen (DIN) and phosphorus (DIP) concen- trations affect pelagic production, leading to DOM modifications. The quantitative and qualitative changes in DOM are often estimated by its optical properties. Colored DOM (CDOM) is often used to estimate dissolved organic carbon (DOC) concentrations by applied techniques, e.g. through remote sensing, whereas DOM properties, such as molecular weight, can be estimated from the slopes of the CDOM absorption spectra (S). Fluorescence properties of CDOM (FDOM) allow discriminating between different structural CDOM properties. The investigation of distribution and cycling of CDOM and FDOM was recognized to be important for understanding of physical and biogeochemical processes, influencing DOM. However, little information is available about effects of nutrient variability on CDOM and FDOM dynamics. Here we present results from two mesocosm experiments conducted with a natural plankton community of the ETNA, where effects of DIP ("Varied P") and DIN ("Varied N") supply on optical properties of DOM were studied. CDOM accumulated proportionally to phytoplankton biomass during the experiments. S decreased over time indicating accumulation of high molecular weight DOM. In Varied N, an additional CDOM portion, as a result of bacterial DOM reworking, was determined. It increased the CDOM fraction in DOC proportionally to the supplied DIN. The humic-like FDOM component (Comp.1) was derived by bacteria proportionally to DIN supply. The bound-to-protein amino acid-like FDOM component (Comp.2) was released irrespectively to phytoplankton biomass, but depending on DIP and DIN concentrations, as a part of an overflow mechanism. Under high DIN supply, Comp.2 was removed by bacterial reworking processes, leading to an accumulation of humic-like Comp.1. No influence of nutrient availability on amino acid-like FDOM component in peptide form (Comp.3) was observed. Comp.3 potentially acted as an intermediate product during formation or degradation Comp.2. Our findings suggest that changes in nutrient concentrations may lead to substantial responses in the quantity and "quality" of optically active DOM and, therefore, might bias results of the applied techniques for an estimation of DOC concentrations in open ocean regions. [ABSTRACT FROM AUTHOR]

The eastern tropical North Atlantic (ETNA) is characterized by a highly productive coastal upwelling system and a moderate oxygen minimum zone with lowest open ocean oxygen (O2) concentrations of around 40 µmol kg-1. Only recently, the discovery of re-occurring mesoscale eddies with sometimes close to anoxic O2 concentrations (< 1 µmol kg-1) and located just below the mixed layer challenged our understanding of O2 distribution and biogeochemical processes in this area. Here, we present the first metagenomic dataset from a deoxygenated anticyclonic modewater eddy in the open waters of the ETNA. In the eddy, we observed a significantly lower bacterial diversity compared to surrounding waters, along with a significant community shift. We detected enhanced primary productivity in the surface layer of the eddy indicated by elevated chlorophyll concentrations and increased carbon uptake rates up to three times as high as in surrounding waters. Carbon uptake below the euphotic zone correlated to the presence of a specific high-light ecotype of Prochlorococcus, which is usually underrepresented in the ETNA. Our combined data indicate that high primary production in the eddy fuels export production and the presence of a specific microbial community responsible for enhanced respiration at shallow depths, below the mixed layer base. Progressively decreasing O2 concentrations in the eddy were found to promote transcription of the key gene for denitrification, nirS, in the O2- depleted core waters. This process is usually absent from the open ETNA waters. In the light of future ocean deoxygenation our results show exemplarily that even distinct events of anoxia have the potential to alter microbial community structures and with that critically impact primary productivity and biogeochemical processes of oceanic water bodies. [ABSTRACT FROM AUTHOR]

It has been suggested that oxygen minimum zones (OMZ) will expand in the tropical oceans as a result of global change with potential consequences for marine element cycling, such as an increase in anaerobic nitrogen loss, resulting in a lower supply of nitrate relative to phosphate to the euphotic zone. So far, the effects of changes in nutrient ratios on organic matter recycling and export fluxes are not well understood. Here, were investigated how different phosphate (Varied P: 0.15-1.58 μmol L-1) or nitrate (Varied N: 1.9-21.9 μmol L-1) concentrations affect the abundance and size distribution of polysaccharidic transparent exopolymer particles (TEP), which are suggested to enhance particle aggregation and export fluxes, and on proteinaceous coomassie stainable particles (CSP), a supposedly good substrate for heterotrophic bacteria. Two series of mesocosm bloom experiments were conducted with natural plankton communities collected from the Eastern Tropical North Atlantic (ETNA) close to Cape Verde in October 2012. Until bloom peak, a positive correlation between gel particle abundance and Chl a concentration was determined, linking the release of dissolved gel precursors and the subsequent formation of gel particles to autotrophic production. After bloom peak, gel particle abundance remained stable or even increased, implying a continued partitioning of dissolved into particulate organic matter after biomass production itself ceased. During both experiments, differences between TEP and CSP dynamics were observed; TEP were generally more abundant than CSP. Changes in size distribution indicated aggregation of TEP during the bloom, while newly formed CSP decomposed. Abundance of gel particles clearly increased with nitrate concentration during the second experiment, suggesting that changes in [DIN]:[DIP] ratios can affect gel particle formation with potential consequences for carbon and nitrogen cycling as well as food web dynamics in tropical ecosystems. [ABSTRACT FROM AUTHOR]

Ocean deoxygenation due to climate change may alter redox-sensitive nutrient cycles in the marine environment. The productive eastern tropical North Atlantic (ETNA) up- welling region may be particularly affected when the relatively moderate oxygen minimum zone (OMZ) deoxygenates further and microbially-driven nitrogen (N) loss processes are promoted. Consequently, water masses with a low N : P ratio could reach the euphotic layer, possibly influencing primary production in those waters. Previous mesocosm studies in the oligotrophic Atlantic Ocean identified N availability as controlling of primary production, while a possible co-limitation of nitrate and phosphate (P) could not be ruled out. To better understand the impact of changing N: P ratios on primary production and on N2 fixation in the ETNA surface ocean, we conducted land-based mesocosm experiments with natural plankton communities and applied a broad range of N: P ratios (2.67-48). Silicate was supplied at 15 µmol L-1 in all mesocosms. We monitored nutrient drawdown, bloom formation, biomass build up and diazotrophic feedback in response to variable nutrient stoichiometry. Our results confirmed N to be limiting to primary production. We found that excess P was channeled through particulate organic matter (POP) into the dissolved organic matter (DOP) pool. In mesocosms with low P availability, DOP was utilized while N2 fixation increased, suggesting a link between those two processes. Interestingly this observation was most pronounced in mesocosms where inorganic N was still available, indicating that bioavailable N does not necessarily has to have a negative impact on N2 fixation. We observed a shift from a mixed cyanobacterial/proteobacterial dominated active diazotrophic community towards diazotrophic diatom symbionts of the Richelia-Rhizosolenia symbiosis. We hypothesize that a potential change in nutrient stoichiometry in the ETNA might lead to a general shift within the diazotrophic community, potentially modifying primary productivity. [ABSTRACT FROM AUTHOR]

The Eastern Tropical North Atlantic (ETNA) is an open ocean region with little input of terrestrial dissolved organic matter (DOM), suggesting that pelagic production has to be the main source of DOM. Inorganic nitrogen (DIN) and phosphorus (DIP) concentrations affect pelagic production, leading to DOM modifications. The quantitative and qualitative changes in DOM are often estimated by its optical properties. Colored DOM (CDOM) is often used to estimate dissolved organic carbon (DOC) concentrations by applied techniques, e.g. through remote sensing, whereas DOM properties, such as molecular weight, can be estimated from the slopes of the CDOM absorption spectra (S). Fluorescence properties of CDOM (FDOM) allow discriminating between different structural CDOM properties. The investigation of distribution and cycling of CDOM and FDOM was recognized to be important for understanding of physical and biogeochemical processes, influencing DOM. However, little information is available about effects of nutrient variability on CDOM and FDOM dynamics. Here we present results from two mesocosm experiments conducted with a natural plankton community of the ETNA, where effects of DIP (''Varied P") and DIN ("Varied N") supply on optical properties of DOM were studied. CDOM accumulated proportionally to phytoplankton biomass during the experiments. S decreased over time indicating accumulation of high molecular weight DOM. In Varied N, an additional CDOM portion, as a result of bacterial DOM reworking, was determined. It increased the CDOM fraction in DOC proportionally to the supplied DIN. The humic-like FDOM component (Comp.1) was derived by bacteria proportionally to DIN supply. The bound-to-protein amino acid-like FDOM component (Comp.2) was released irrespectively to phytoplankton biomass, but depending on DIP and DIN concentrations, as a part of an overflow mechanism. Under high DIN supply, Comp.2 was removed by bacterial reworking processes, leading to an accumulation of humic-like Comp.1. No influence of nutrient availability on amino acid-like FDOM component in peptide form (Comp.3) was observed. Comp.3 potentially acted as an intermediate product during formation or degradation Comp.2. Our findings suggest that changes in nutrient concentrations may lead to substantial responses in the quantity and "quality" of optically active DOM and, therefore, might bias results of the applied techniques for an estimation of DOC concentrations in open ocean regions. [ABSTRACT FROM AUTHOR]

It has been suggested that oxygen minimum zones (OMZ) will expand in the tropical oceans as a result of global change with potential consequences for marine element cycling, such as an increase in anaerobic nitrogen loss, resulting in a lower supply of nitrate relative to phosphate to the euphotic zone. So far, the effects of changes in nutrient ratios on organic matter recycling and export fluxes are not well understood. Here, were investigated how different phosphate (Varied P: 0.15-1.58 µmol L-1) or nitrate (Varied N: 1.9-21.9 µmol L-1) concentrations affect the abundance and size distribution of polysaccharidic transparent exopolymer particles (TEP), which are suggested to enhance particle aggregation and export fluxes, and on proteinaceous coomassie stainable particles (CSP), a supposedly good substrate for heterotrophic bacteria. Two series of mesocosm bloom experiments were conducted with natural plankton communities collected from the Eastern Tropical North Atlantic (ETNA) close to Cape Verde in October 2012. Until bloom peak, a positive correlation between gel particle abun dance and Chl a concentration was determined, linking the release of dissolved gel precursors and the subsequent formation of gel particles to autotrophic production. After bloom peak, gel particle abundance remained stable or even increased, implying a continued partitioning of dissolved into particulate organic matter after biomass production itself ceased. During both experiments, differences between TEP and CSP dynamics were observed; TEP were generally more abundant than CSP. Changes in size distribution indicated aggregation of TEP during the bloom, while newly formed CSP decomposed. Abundance of gel particles clearly increased with nitrate concentration during the second experiment, suggesting that changes in [DIN] : [DIP] ratios can a_ect gel particle formation with potential consequences for carbon and nitrogen cycling as well as food web dynamics in tropical ecosystems. [ABSTRACT FROM AUTHOR]

Recent modeling results suggest that oceanic oxygen levels will decrease significantly over the next decades to centuries in response to climate change and altered ocean circulation. Hence the future ocean may experience major shifts in nutrient cycling triggered by the expansion and intensification of tropical oxygen minimum zones (OMZs). There are numerous feedbacks between oxygen concentrations, nutrient cycling and biological productivity; however, existing knowledge is insufficient to understand physical, chemical and biological interactions in order to adequately assess past and potential future changes. We investigated the pelagic biogeochemistry of OMZs in the eastern tropical North Atlantic and eastern tropical South Pacific during a series of cruise expeditions and mesocosm studies. The following summarizes the current state of research on the influence of low environmental oxygen conditions on marine biota, viruses, organic matter formation and remineralization with a particular focus on the nitrogen cycle in OMZ regions. The impact of sulfidic events on water column biogeochemistry, originating from a specific microbial community capable of highly efficient carbon fixation, nitrogen turnover and N2O production is further discussed. Based on our findings, an important role of sinking particulate organic matter in controlling the nutrient stochiometry of the water column is suggested. These particles can enhance degradation processes in OMZ waters by acting as microniches, with sharp gradients enabling different processes to happen in close vicinity, thus altering the interpretation of oxic and anoxic environments. [ABSTRACT FROM AUTHOR]

Oxygen-deficient waters in the ocean, generally referred to as oxygen minimum zones (OMZ), are expected to expand as a consequence of global climate change. Poor oxygenation is promoting microbial loss of inorganic nitrogen (N) and increasing release of sediment-bound phosphate (P) into the water column. These intermediate water masses, nutrient-loaded but with an N deficit relative to the canonical N: P Redfield ratio of 16 : 1, are transported via coastal up-welling into the euphotic zone. To test the impact of nutrient supply and nutrient stoichiometry on production, partitioning and elemental composition of dissolved (DOC, DON, DOP) and particulate (POC, PON, POP) organic matter, three nutrient enrichment experiments were conducted with natural microbial communities in shipboard mesocosms, during research cruises in the tropical waters of the southeast Pacific and the northeast Atlantic. Maximum accumulation of POC and PON was observed under high N supply conditions, indicating that primary production was controlled by N availability. The stoichiometry of microbial biomass was unaffected by nutrient N: P supply during exponential growth under nutrient saturation, while it was highly variable under conditions of nutrient limitation and closely correlated to the N: P supply ratio, although PON: POP of accumulated biomass generally exceeded the supply ratio. Microbial N: P composition was constrained by a general lower limit of 5 : 1. Channelling of assimilated P into DOP appears to be the mechanism responsible for the consistent offset of cellular stoichiometry relative to inorganic nutrient supply and nutrient drawdown, as DOP build-up was observed to intensify under decreasing N: P supply. Low nutrient N: P conditions in coastal upwelling areas overlying O2-deficient waters seem to represent a net source for DOP, which may stimulate growth of diazotrophic phytoplankton. These results demonstrate that microbial nutrient assimilation and partitioning of organic matter between the particulate and the dissolved phase are controlled by the N: P ratio of upwelled nutrients, implying substantial consequences for nutrient cycling and organic matter pools in the course of decreasing nutrient N: P stoichiometry. [ABSTRACT FROM AUTHOR]

Oxygen-deficient waters in the ocean, generally referred to as oxygen minimum zones (OMZ), are expected to expand as a consequence of global climate change. Poor oxygenation is promoting microbial loss of inorganic nitrogen (N) and increasing release of sediment-bound phosphate (P) into the water column. These intermediate water masses, nutrient-loaded but with an N deficit relative to the canonical N : P Redfield ratio of 16 : 1, are transported via coastal upwelling into the euphotic zone. To test the impact of nutrient supply and nutrient stoichiometry on production, partitioning and elemental composition of phytoplankton-derived dissolved (DOC, DON, DOP) and particulate (POC, PON, POP) organic matter, three nutrient enrichment experiments were conducted with natural phytoplankton communities in shipboard mesocosms, during research cruises in the tropical waters of the South East Pacific and the North East Atlantic. Maximum accumulation of POC and PON was observed under high N supply conditions, indicating that primary production was controlled by N availability. The stoichiometry of photoautotrophic biomass was unaffected by nutrient N : P supply during exponential growth under nutrient saturation, while it was highly variable under conditions of nutrient limitation and closely correlated to the N : P supply ratio, although PON : POP of accumulated phytoplankton generally exceeded the supply ratio. Phytoplankton N : P composition was constrained by a general lower limit of 5 : 1. Channelling of assimilated P into DOP appears to be the mechanism responsible for the consistent offset of cellular stoichiometry relative to inorganic nutrient supply and nutrient drawdown, as DOP build-up was observed to intensify under decreasing N : P supply. Low nutrient N : P conditions in coastal upwelling areas overlying O2-deficient waters seem to represent a net source for DOP, which may stimulate growth of diazotrophic phytoplankton. These results demonstrate that microalgal nutrient assimilation and partitioning of phytoplankton-derived organic matter between the particulate and the dissolved phase are controlled by the N : P ratio of upwelled nutrients, implying substantial consequences for nutrient cycling and organic matter pools in the course of decreasing nutrient N : P stoichiometry. [ABSTRACT FROM AUTHOR]