ALBERT

Description: Responses to ocean acidification in plankton communities were studied during a CO2-enrichment experiment in the Arctic Ocean, accomplished from June to July 2010 in Kongsfjorden, Svalbard (78°56, 2' N, 11°53, 6' E). Enclosed in 9 mesocosms (volume: 43.9–47.6 m3), plankton was exposed to CO2 concentrations, ranging from glacial to projected mid-next-century levels. Fertilization with inorganic nutrients at day 13 of the experiment supported the accumulation of phytoplankton biomass, as indicated by two periods of high Chl a concentration. This study tested for CO2 sensitivities in primary production (PP) of particulate organic carbon (PPPOC) and of dissolved organic carbon (PPDOC). Therefore, 14C-bottle incubations (24 h) of mesocosm samples were performed at 1 m depth receiving about 60% of incoming radiation. PP for all mesocosms averaged 8.06 ± 3.64 μmol C l−1 d−1 and was slightly higher than in the outside fjord system. Comparison between mesocosms revealed significantly higher PPPOC at elevated compared to low pCO2 after nutrient addition. PPDOC was significantly higher in CO2 enriched mesocosms before as well as after nutrient addition, suggesting that CO2 had a direct influence on DOC production. DOC concentrations inside the mesocosms increased before nutrient addition and more in high CO2 mesocosms. After addition of nutrients, however, further DOC accumulation was negligible and not significantly different between treatments, indicating rapid utilization of freshly produced DOC. Bacterial biomass production (BP) was coupled to PP in all treatments, indicating that 3.5 ± 1.9% of PP, or 21.6 ± 12.5% of PPDOC provided sufficient carbon for synthesis of bacterial biomass. The response of 14C-based PP rates to CO2 enrichment was at odds with O2-based net community production (NCP) rates that were also determined during this study, albeit at lower light level. We conclude that the enhanced release of labile DOC during autotrophic production at high CO2 exceedingly stimulated activities of heterotrophic microorganisms. As a consequence, increased PP induced less NCP, as suggested earlier for carbon limited microbial systems in the Arctic.

Description: Extracellular release (ER) by phytoplankton is the major source of fresh dissolved organic carbon (DOC) in marine ecosystems and accompanies primary production during all growth phases. Little is known, so far, on size and composition of released molecules, and to which extent ER occurs passively, by leakage, or actively, by exudation. Here, we report on ER by the widespread and bloom-forming coccolithophore Emiliania huxleyi grown under steady state conditions in phosphorus controlled chemostats (N : P = 29, growth rate of μ = 0.2 d−1). 14C incubations were accomplished to determine primary production (PP), comprised by particulate (PO14C) and dissolved organic carbon (DO14C), and the concentration and composition of particulate combined carbohydrates (pCCHO), and of high molecular weight (〉1 kDa, HMW) dissolved combined carbohydrates (dCCHO) as major components of ER. Information on size distribution of ER products was obtained by investigating distinct size classes (

Description: The effect of elevated seawater carbon dioxide (CO2) on the activity of a natural bacterioplankton community in an Arctic fjord system was investigated by a mesocosm perturbation study in the frame of the European Project on Ocean Acidification (EPOCA). A pCO2 range of 175–1085 μatm was set up in nine mesocosms deployed in the Kongsfjorden (Svalbard). The bacterioplankton communities responded to rising chlorophyll a concentrations after a lag phase of only a few days with increasing protein production and extracellular enzyme activity and revealed a close coupling of heterotrophic bacterial activity to phytoplankton productivity in this experiment. The natural extracellular enzyme assemblages showed increased activity in response to moderate acidification. A decrease in seawater pH of 0.5 units roughly doubled rates of β-glucosidase and leucine-aminopeptidase. Activities of extracellular enzymes in the mesocosms were directly related to both seawater pH and primary production. Also primary production and bacterial protein production in the mesocosms at different pCO2 were positively correlated. Therefore, it can be suggested that the efficient heterotrophic carbon utilization in this Arctic microbial food web had the potential to counteract increased phytoplankton production that was achieved under elevated pCO2 in this study. However, our results also show that the transfer of beneficial pCO2-related effects on the cellular bacterial metabolism to the scale of community activity and organic matter degradation can be mitigated by the top-down control of bacterial abundances in natural microbial communities.

Description: 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.

Description: Emiliania huxleyi (strain B 92/11) was exposed to different growth, CO2 and temperature conditions in phosphorous controlled chemostats, to investigate effects on organic carbon exudation, and partitioning between the pools of particulate organic carbon (POC) and dissolved organic carbon (DOC). 14C incubation measurements for primary production (PP) and for extracellular release (ER) were performed. Chemical analysis included amount and composition of high molecular weight dissolved combined carbohydrates (〉1 kDa, HMW-dCCHO), particulate combined carbohydrates (pCCHO) and the carbon content of transparent exopolymer particles (TEP-C). Applied CO2 and temperature conditions were 300, 550 and 900 μatm pCO2 at 14 °C, and additionally 900 μatm pCO2 at 18 °C simulating a greenhouse ocean scenario. A reduction in growth rate from μ =0.3 d−1 to μ =0.1 d−1 induced the most profound effect on the performance of E. huxleyi, relative to the effect of elevated CO2 and temperature. At μ =0.3 d−1, PP was significantly higher at elevated CO2 and temperature. DO14C production correlated to PO14C production in all cultures, resulting in similar percentages of extracellular release (DO14C/PP × 100; PER) of averaged 3.74 ± 0.94%. At μ =0.1 d−1, PO14C decreased significantly, while exudation of DO14C increased, thus leading to a stronger partitioning from the particulate to the dissolved pool. Maximum PER of 16.3 ± 2.3% were observed at μ =0.1 d−1 at greenhouse conditions. Concentrations of HMW-dCCHO and pCCHO were generally higher at μ =0.1 d−1 compared to μ =0.3 d−1. At μ =0.3 d−1, pCCHO concentration increased significantly along with elevated CO2 and temperature. Despite of high PER, the percentage of HMW-dCCHO was smallest at greenhouse conditions. However, highest TEP-formation was observed under greenhouse conditions, together with a pronounced increase in pCCHO concentration, suggesting a stronger partitioning of PP from DOC to POC by coagulation of exudates. Our results imply that greenhouse condition will enhance exudation processes in E. huxleyi and may affect organic carbon partitioning in the ocean due to an enhanced transfer of HMW-dCCHO to TEP by aggregation processes.

Description: With the accumulation of anthropogenic carbon dioxide (CO2), a proceeding decline in seawater pH has been induced that is referred to as ocean acidification. The ocean's capacity for CO2 storage is strongly affected by biological processes, whose feedback potential is difficult to evaluate. The main source of CO2 in the ocean is the decomposition and subsequent respiration of organic molecules by heterotrophic bacteria. However, very little is known about potential effects of ocean acidification on bacterial degradation activity. This study reveals that the degradation of polysaccharides, a major component of marine organic matter, by bacterial extracellular enzymes was significantly accelerated during experimental simulation of ocean acidification. Results were obtained from pH perturbation experiments, where rates of extracellular α- and β-glucosidase were measured and the loss of neutral and acidic sugars from phytoplankton-derived polysaccharides was determined. Our study suggests that a faster bacterial turnover of polysaccharides at lowered ocean pH has the potential to affect the cycling of organic carbon in the future ocean by weakening the biological carbon pump and increasing the respiratory production of CO2.

Description: 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.

Description: 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.

Description: 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 on DOM 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 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 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.

Description: Emiliania huxleyi (strain B 92/11) was exposed to different nutrient supply, CO2 and temperature conditions in phosphorus controlled chemostats to investigate effects on organic carbon exudation and partitioning between the pools of particulate organic carbon (POC) and dissolved organic carbon (DOC). 14C incubation measurements for primary production (PP) and extracellular release (ER) were performed. Chemical analysis included the amount and composition of high molecular weight (〉1 kDa) dissolved combined carbohydrates (HMW-dCCHO), particulate combined carbohydrates (pCCHO) and the carbon content of transparent exopolymer particles (TEP-C). Applied CO2 and temperature conditions were 300, 550 and 900 μatm pCO2 at 14 °C, and additionally 900 μatm pCO2 at 18 °C simulating a greenhouse ocean scenario. Enhanced nutrient stress by reducing the dilution rate (D) from D = 0.3 d−1 to D = 0.1 d−1 (D = μ) induced the strongest response in E. huxleyi. At μ = 0.3 d−1, PP was significantly higher at elevated CO2 and temperature and DO14C production correlated to PO14C production in all treatments, resulting in similar percentages of extracellular release (PER; (DO14C production/PP) × 100) averaging 3.74 ± 0.94%. At μ = 0.1 d−1, PO14C production decreased significantly, while exudation of DO14C increased. Thus, indicating a stronger partitioning from the particulate to the dissolved pool. Maximum PER of 16.3 ± 2.3% were observed at μ = 0.1 d−1 at elevated CO2 and temperature. While cell densities remained constant within each treatment and throughout the experiment, concentrations of HMW-dCCHO, pCCHO and TEP were generally higher under enhanced nutrient stress. At μ = 0.3 d−1, pCCHO concentration increased significantly with elevated CO2 and temperature. At μ = 0.1 d−1, the contribution (mol % C) of HMW-dCCHO to DOC was lower at elevated CO2 and temperature while pCCHO and TEP concentrations were higher. This was most pronounced under greenhouse conditions. Our findings suggest a stronger transformation of primary produced DOC into POC by coagulation of exudates under nutrient limitation. Our results further imply that elevated CO2 and temperature will increase exudation by E. huxleyi and may affect organic carbon partitioning in the ocean due to an enhanced transfer of HMW-dCCHO to TEP by aggregation processes.