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Gravitational sinking of photosynthetically fixed particulate organic carbon (POC) constitutes a key component of the biological carbon pump. The fraction of POC leaving the surface ocean depends on POC sinking velocity (SV) and remineralization rate (Cremin), both of which depend on plankton community structure. However, the key drivers in plankton communities controlling SV and Cremin are poorly constrained. In fall 2014, we conducted a 6 weeks mesocosm experiment in the subtropical NE Atlantic Ocean to study the influence of plankton community structure on SV and Cremin. Oligotrophic conditions prevailed for the first 3 weeks, until nutrient‐rich deep water injected into all mesocosms stimulated diatom blooms. SV declined steadily over the course of the experiment due to decreasing CaCO3 ballast and – according to an optical proxy proposed herein – due to increasing aggregate porosity mostly during an aggregation event after the diatom bloom. Furthermore, SV was positively correlated with the contribution of picophytoplankton to the total phytoplankton biomass. Cremin was highest during a Synechococcus bloom under oligotrophic conditions and in some mesocosms during the diatom bloom after the deep‐water addition while it was particularly low during harmful algal blooms. The temporal changes were considerably larger in Cremin (max. 15‐fold) than in SV (max. 3‐fold). Accordingly, estimated POC transfer efficiency to 1000 m was mainly dependent on how the plankton community structure affected Cremin. Our approach revealed key players and interactions in the plankton food web influencing POC export efficiency thereby improving our mechanistic understanding of the biological carbon pump. Key points Sinking velocity was higher during oligotrophy than during blooms which is linked to ballast, porosity, and phytoplankton size structure Remineralization was highly variable but tended to be higher during Synechococcus or diatom blooms and lower during harmful algal blooms Plankton community structure had a considerably larger influence on particle remineralization rate than on sinking velocity

Marine species and ecosystems are exposed to a wide range of environmental change – both detrimental (threats) and beneficial – due to human activities. Some of the changes are global, whereas others are regional or local. It is important to distinguish the scale of each threat as the solutions will differ. For example, the mitigation of a global problem requires a global response, which is more difficult to achieve than addressing a local problem with a local response. These wide-ranging changes are often referred to drivers or stressors. The term multiple drivers refers to the concurrent alteration of multiple environmental properties, that are each biologically-influential, by anthropogenic pressures including climate change. These multiple environmental properties are commonly referred to as drivers or stressors, and include temperature, carbon dioxide, pH, oxygen, salinity, density, irradiance and nutrients, eutrophication, UV exposure, and point source pollutants (Figure 1). The multiple drivers framework represents a complex matrix of changing ocean properties, that will vary from locale to locale, and may also alter with season. Published Refereed Current 14.1.1 Nutrients Best Practice

Marine calcareous sediments provide a fundamental basis for palaeoceanographic studies aiming to reconstruct past oceanic conditions and understand key biogeochemical element cycles. Calcifying unicellular phytoplankton (coccolithophores) are a major contributor to both carbon and calcium cycling by photosynthesis and the production of calcite (coccoliths) in the euphotic zone, and the subsequent long-term deposition and burial into marine sediments. Here we present data from controlled laboratory experiments on four coccolithophore species and elucidate the relation between the divalent cation (Sr, Mg and Ca) partitioning in coccoliths and cellular physiology (growth, calcification and photosynthesis). Coccolithophores were cultured under different seawater temperature and carbonate chemistry conditions. The partition coefficient of strontium (DSr) was positively correlated with both carbon dioxide (pCO2) and temperature but displayed no coherent relation to particulate organic and inorganic carbon production rates. Furthermore, DSr correlated positively with cellular growth rates when driven by temperature but no correlation was present when changes in growth rates were pCO2-induced. Our results demonstrate the complex interaction between environmental forcing and physiological control on the strontium partitioning in coccolithophore calcite and challenge interpretations of the coccolith Sr / Ca ratio from high-pCO2 environments (e.g. Palaeocene–Eocene thermal maximum). The partition coefficient of magnesium (DMg) displayed species-specific differences and elevated values under nutrient limitation. No conclusive correlation between coccolith DMg and temperature was observed but pCO2 induced a rising trend in coccolith DMg. Interestingly, the best correlation was found between coccolith DMg and chlorophyll a production, suggesting that chlorophyll a and calcite associated Mg originate from the same intracellular pool. These and previous findings indicate that Mg is transported into the cell and to the site of calcification via different pathways than Ca and Sr. Consequently, the coccolith Mg / Ca ratio should be decoupled from the seawater Mg / Ca ratio. This study gives an extended insight into the driving factors influencing the coccolith Mg / Ca ratio and should be considered for future palaeoproxy calibrations.

The surface ocean absorbs large quantities of the CO2 emitted to the atmosphere from human activities. As this CO2 dissolves in seawater, it reacts to form carbonic acid. While this phenomenon, called ocean acidification, has been found to adversely affect many calcifying organisms, some photosynthetic organisms appear to benefit from increasing [CO2]. Among these is the cyanobacterium Trichodesmium, a predominant diazotroph (nitrogen-fixing) in large parts of the oligotrophic oceans, which responded with increased carbon and nitrogen fixation at elevated pCO2. With the mechanism underlying this CO2 stimulation still unknown, the question arises whether this is a common response of diazotrophic cyanobacteria. In this study we therefore investigate the physiological response of Nodularia spumigena, a heterocystous bloom-forming diazotroph of the Baltic Sea, to CO2-induced changes in seawater carbonate chemistry. N. spumigena reacted to seawater acidification/carbonation with reduced cell division rates and nitrogen fixation rates, accompanied by significant changes in carbon and phosphorus quota and elemental composition of the formed biomass. Possible explanations for the contrasting physiological responses of Nodularia compared to Trichodesmium may be found in the different ecological strategies of non-heterocystous (Trichodesmium) and heterocystous (Nodularia) cyanobacteria.

Availability of phosphate for phytoplankton and bacteria and of glucose for bacteria at different pCO2 levels were studied in a mesocosm experiment (PeECE III). Using nutrient-depleted SW Norwegian fjord waters, three different levels of pCO2 (350 μatm: 1×CO2; 700 μatm: 2×CO2; 1050 μatm: 3×CO2) were set up, and nitrate and phosphate were added at the start of the experiment in order to induce a phytoplankton bloom. Despite similar responses of total particulate P concentration and phosphate turnover time at the three different pCO2 levels, the size distribution of particulate P and 33PO4 uptake suggested that phosphate transferred to the >10 μm fraction was greater in the 3×CO2 mesocosm during the first 6–10 days when phosphate concentration was high. During the period of phosphate depletion (after Day 12), specific phosphate affinity and specific alkaline phosphatase activity (APA) suggested a P-deficiency (i.e. suboptimal phosphate supply) rather than a P-limitation for the phytoplankton and bacterial community at the three different pCO2 levels. Specific phosphate affinity and specific APA tended to be higher in the 3×CO2 than in the 2×CO2 and 1×CO2 mesocosms during the phosphate depletion period, although no statistical differences were found. Glucose turnover time was correlated significantly and negatively with bacterial abundance and production but not with the bulk DOC concentration. This suggests that even though constituting a small fraction of the bulk DOC, glucose was an important component of labile DOC for bacteria. Specific glucose affinity of bacteria behaved similarly at the three different pCO2 levels with measured specific glucose affinities being consistently much lower than the theoretical maximum predicted from the diffusion-limited model. This suggests that bacterial growth was not severely limited by the glucose availability. Hence, it seems that the lower availability of inorganic nutrients after the phytoplankton bloom reduced the bacterial capacity to consume labile DOC in the upper mixed layer of the stratified mesocosms.

International audience; A dynamic model has been developed to represent biogeochemical variables and processes observed during a bloom of Emiliania huxleyi coccolithophore. This bloom was induced in a mesocosm experiment during which the ecosystem development was followed over a period of 23-days through changes in various biogeochemical parameters such as inorganic nutrients (nitrate, ammonium and phosphate), total alkalinity (TA), dissolved inorganic carbon (DIC), partial pressure of CO2 (pCO2), dissolved oxygen (O2), photosynthetic pigments, particulate organic carbon (POC), dissolved organic carbon (DOC), Transparent Exopolymer Particles (TEP), primary production, and calcification. This dynamic model is based on unbalanced algal growth and balanced bacterial growth. In order to adequately reproduce the observations, the model includes an explicit description of phosphorus cycling, calcification, TEP production and an enhanced mortality due to viral lysis. The model represented carbon, nitrogen and phosphorus fluxes observed in the mesocosms. Modelled profiles of algal biomass and final concentrations of DIC and nutrients are in agreement with the experimental observations.

The effect of CO2 on carbon fluxes in Arctic plankton communities was investigated during the 2010 EPOCA mesocosm study in Ny Ålesund, Svalbard. Nine mesocosms were set up with initial pCO2 levels ranging from 185 to 1420 μatm for 5 weeks. 13C labelled bicarbonate was added at the start of the experiment to follow the transfer of carbon from dissolved inorganic carbon (DIC) into phytoplankton, bacteria, total particulate organic carbon (POC), zooplankton, and settling particles. Polar lipid derived fatty acids (PLFA) were used to trace carbon dynamics of phytoplankton and bacteria and allowed distinction of two groups of phytoplankton: phyto I (autotrophs) and phyto II (mixotrophs). Nutrients were added on day 13. A nutrient-phytoplankton-zooplankton-detritus model amended with 13C dynamics was constructed and fitted to the data to quantify uptake rates and carbon fluxes in the plankton community during the phase prior to nutrient addition (phase 1, days 0–12). During the first 12 days, a phytoplankton bloom developed that was characterized by high growth rates (0.87 days−1) for phyto I and lower growth rates (0.18 days−1) for phyto II. A large part of the carbon fixed by phytoplankton (~31%) was transferred to bacteria, while mesozooplankton grazed only ~6% of the production. After 6 days, the bloom collapsed and part of the organic matter subsequently settled into the sediment traps. The sedimentation losses of detritus in phase 1 were low (0.008 days−1) and overall export was only ~7% of production. Zooplankton grazing and detritus sinking losses prior to nutrient addition were sensitive to CO2: grazing decreased with increasing CO2, while sinking increased. Phytoplankton production increased again after nutrient addition on day 13. Although phyto II showed initially higher growth rates with increasing CO2 (days 14–22), the overall production of POC after nutrient addition (phase 2, days 14–29) decreased with increasing CO2. Significant sedimentation occurred towards the end of the experiment (after day 24) and much more material settled down in the sediment traps at low CO2.

The ocean's potential to export carbon to depth partly depends on the fraction of primary production (PP) sinking out of the euphotic zone (i.e., the e-ratio). Measurements of PP and export flux are often performed simultaneously in the field, although there is a temporal delay between those parameters. Thus, resulting e-ratio estimates often incorrectly assume an instantaneous downward export of PP to export flux. Evaluating results from four mesocosm studies, we find that peaks in organic matter sedimentation lag chlorophyll a peaks by 2 to 15 days. We discuss the implications of these time lags (TLs) for current e-ratio estimates and evaluate potential controls of TL. Our analysis reveals a strong correlation between TL and the duration of chlorophyll a buildup, indicating a dependency of TL on plankton food web dynamics. This study is one step further toward time-corrected e-ratio estimates

Ohne das Meer gäbe es kein Leben auf unserem Planeten. Es regelt weitgehend das Klima, gibt uns Nahrung und liefert Energie. Darüber hinaus ist es ein wichtiger Verkehrsweg, ein Erholungsraum und ein Quell ästhetischen Vergnügens. Aber das Meer steht unter Stress, denn das alte Prinzip von der „Freiheit der Meere“ hat zu Überfischung, Artenverlust und einer immensen Verschmutzung der Ozeane geführt. Deshalb muss der Umgang mit dem Meer auf nachhaltige und gerechte Grundlagen gestellt werden. Der Meeresatlas 2017 liefert dazu die Daten, Fakten und Zusammenhänge. Er zeigt in zahlreichen Beiträgen und über 50 Grafiken, in welch schlechtem Zustand sich die Weltmeere befinden, warum das so ist und was man tun muss, um die Situation der Ozeane zu verbessern.

Ocean acidification resulting from the uptake of anthropogenic carbon dioxide (CO2) by the ocean is considered a major threat to marine ecosystems. Here we examined the effects of ocean acidification on microbial community dynamics in the eastern Baltic Sea during the summer of 2012 when inorganic nitrogen and phosphorus were strongly depleted. Large-volume in situ mesocosms were employed to mimic present, future and far future CO2) scenarios. All six groups of phytoplankton enumerated by flow cytometry (

The oceans absorb about a quarter of the annually produced anthropogenic atmospheric carbon dioxide (CO2), resulting in a decrease in surface water pH, a process termed ocean acidification (OA). Surprisingly little is known about how OA affects the physiology of heterotrophic bacteria or the coupling of heterotrophic bacteria to phytoplankton when nutrients are limited. Previous experiments were, for the most part, undertaken during productive phases or following nutrient additions designed to stimulate algal blooms. Therefore, we performed an in situ large-volume mesocosm ( ∼ 55m3) experiment in the Baltic Sea by simulating different fugacities of CO2 (fCO2) extending from present to future conditions. The study was conducted in July–August after the nominal spring bloom, in order to maintain low-nutrient conditions throughout the experiment. This resulted in phytoplankton communities dominated by small-sized functional groups (picophytoplankton). There was no consistent fCO2-induced effect on bacterial protein production (BPP), cell-specific BPP (csBPP) or biovolumes (BVs) of either free-living (FL) or particle-associated (PA) heterotrophic bacteria, when considered as individual components (univariate analyses). Permutational Multivariate Analysis of Variance (PERMANOVA) revealed a significant effect of the fCO2 treatment on entire assemblages of dissolved and particulate nutrients, metabolic parameters and the bacteria–phytoplankton community. However, distance-based linear modelling only identified fCO2 as a factor explaining the variability observed amongst the microbial community composition, but not for explaining variability within the metabolic parameters. This suggests that fCO2 impacts on microbial metabolic parameters occurred indirectly through varying physicochemical parameters and microbial species composition. Cluster analyses examining the co-occurrence of different functional groups of bacteria and phytoplankton further revealed a separation of the four fCO2-treated mesocosms from both control mesocosms, indicating that complex trophic interactions might be altered in a future acidified ocean. Possible consequences for nutrient cycling and carbon export are still largely unknown, in particular in a nutrient-limited ocean.

In the frame of the European Project on Ocean Acidification (EPOCA), the response of an Arctic pelagic community (pCO2 was investigated. For this purpose 9 large-scale in situ mesocosms were deployed in Kongsfjorden, Svalbard (78°56.2´ N, 11°53.6´ E), in 2010. The present study investigates effects on the communities of particle-attached (PA; >3 μm) and free-living (FL; < 3 μm > 0.2 μm) bacteria by Automated Ribosomal Intergenic Spacer Analysis (ARISA) in 6 of the mesocosms, ranging from 185 to 1050 μatm initial pCO2, and the surrounding fjord. ARISA was able to resolve, on average, 27 bacterial band classes per sample and allowed for a detailed investigation of the explicit richness and diversity. Both, the PA and the FL bacterioplankton community exhibited a strong temporal development, which was driven mainly by temperature and phytoplankton development. In response to the breakdown of a picophytoplankton bloom, numbers of ARISA band classes in the PA community were reduced at low and medium CO2 (~ 185–685 μatm) by about 25%, while they were more or less stable at high CO2 (~ 820–1050 μatm). We hypothesise that enhanced viral lysis and enhanced availability of organic substrates at high CO2 resulted in a more diverse PA bacterial community in the post-bloom phase. Despite lower cell numbers and extracellular enzyme activities in the post-bloom phase, bacterial protein production was enhanced in high CO2 mesocosms, suggesting a positive effect of community richness on this function and on carbon cycling by bacteria.

The Baltic Sea is a unique environment as the largest body of brackish water in the world. Acidification of the surface oceans due to absorption of anthropogenic CO2 emissions is an additional stressor facing the pelagic community of the already challenging Baltic Sea. To investigate its impact on trace gas biogeochemistry, a large-scale mesocosm experiment was performed off Tvärminne Research Station, Finland, in summer 2012. During the second half of the experiment, dimethylsulfide (DMS) concentrations in the highest-fCO2 mesocosms (1075–1333 µatm) were 34 % lower than at ambient CO2 (350 µatm). However, the net production (as measured by concentration change) of seven halocarbons analysed was not significantly affected by even the highest CO2 levels after 5 weeks' exposure. Methyl iodide (CH3I) and diiodomethane (CH2I2) showed 15 and 57 % increases in mean mesocosm concentration (3.8 ± 0.6 increasing to 4.3 ± 0.4 pmol L−1 and 87.4 ± 14.9 increasing to 134.4 ± 24.1 pmol L−1 respectively) during Phase II of the experiment, which were unrelated to CO2 and corresponded to 30 % lower Chl a concentrations compared to Phase I. No other iodocarbons increased or showed a peak, with mean chloroiodomethane (CH2ClI) concentrations measured at 5.3 (±0.9) pmol L−1 and iodoethane (C2H5I) at 0.5 (±0.1) pmol L−1. Of the concentrations of bromoform (CHBr3; mean 88.1 ± 13.2 pmol L−1), dibromomethane (CH2Br2; mean 5.3 ± 0.8 pmol L−1), and dibromochloromethane (CHBr2Cl, mean 3.0 ± 0.5 pmol L−1), only CH2Br2 showed a decrease of 17 % between Phases I and II, with CHBr3 and CHBr2Cl showing similar mean concentrations in both phases. Outside the mesocosms, an upwelling event was responsible for bringing colder, high-CO2, low-pH water to the surface starting on day t16 of the experiment; this variable CO2 system with frequent upwelling events implies that the community of the Baltic Sea is acclimated to regular significant declines in pH caused by up to 800 µatm fCO2. After this upwelling, DMS concentrations declined, but halocarbon concentrations remained similar or increased compared to measurements prior to the change in conditions. Based on our findings, with future acidification of Baltic Sea waters, biogenic halocarbon emissions are likely to remain at similar values to today; however, emissions of biogenic sulfur could significantly decrease in this region.

The coccolithophore Emiliania huxleyi is a marine phytoplankton species capable of forming small calcium carbonate scales (coccoliths) which cover the organic part of the cell. Calcification rates of E. huxleyi are known to be sensitive to changes in seawater carbonate chemistry. It has, however, not yet been clearly determined how these changes are reflected in size and weight of individual coccoliths and which specific parameter(s) of the carbonate system drive morphological modifications. Here, we compare data on coccolith size, weight, and malformation from a set of five experiments with a large diversity of carbonate chemistry conditions. This diversity allows distinguishing the influence of individual carbonate chemistry parameters such as carbon dioxide (CO2), bicarbonate (HCO3−), carbonate ion (CO32−), and protons (H+) on the measured parameters. Measurements of fine-scale morphological structures reveal an increase of coccolith malformation with decreasing pH suggesting that H+ is the major factor causing malformations. Coccolith distal shield area varies from about 5 to 11 μm2. Changes in size seem to be mainly induced by varying [HCO3−] and [H+] although influence of [CO32−] cannot be entirely ruled out. Changes in coccolith weight were proportional to changes in size. Increasing CaCO3 production rates are reflected in an increase in coccolith weight and an increase of the number of coccoliths formed per unit time. The combined investigation of morphological features and coccolith production rates presented in this study may help to interpret data derived from sediment cores, where coccolith morphology is used to reconstruct calcification rates in the water column.

Ocean acidification and associated shifts in carbonate chemistry speciation induced by increasing levels of atmospheric carbon dioxide (CO2) have the potential to impact marine biota in various ways. The process of biogenic calcification, for instance, is usually shown to be negatively affected. In coccolithophores, an important group of pelagic calcifiers, changes in cellular calcification rates in response to changing ocean carbonate chemistry appear to differ among species. By applying a wider CO2 range we show that a species previously reported insensitive to seawater acidification, Coccolithus braarudii, responds both in terms of calcification and photosynthesis, although at higher levels of CO2. Thus, observed differences between species seem to be related to individual sensitivities while the underlying mechanisms could be the same. On this basis we develop a conceptual model of coccolithophorid calcification and photosynthesis in response to CO2-induced changes in seawater carbonate chemistry speciation.

International audience; No abstract available.

International audience; The potential impact of seawater acidification on the concentrations of dimethylsulfide (DMS) and dimethylsulfoniopropionate (DMSP), and the activity of the enzyme DMSP-lyase was investigated during a pelagic ecosystem CO2 enrichment experiment (PeECE III) in spring 2005. Natural phytoplankton blooms were studied for 24 days under present, double and triple partial pressures of CO2 (pCO2; pH=8.3, 8.0, 7.8) in triplicate 25 m3 enclosures. The results indicate similar DMSP concentrations and DMSP-lyase activity (DLA) patterns for all treatments. Hence, DMSP and DLA do not seem to have been affected by the CO2 treatment. In contrast, DMS concentrations showed small but statistically significant differences in the temporal development of the low versus the high CO2 treatments. The low pCO2 enclosures had higher DMS concentrations during the first 10 days, after which the levels decreased earlier and more rapidly than in the other treatments. Integrated over the whole study period, DMS concentrations were not significantly different from those of the double and triple pCO2 treatments. Pigment and flow-cytometric data indicate that phytoplanktonic populations were generally similar between the treatments, suggesting a certain resilience of the marine ecosystem under study to the induced pH changes, which is reflected in DMSP and DLA. However, there were significant differences in bacterial community structure and the abundance of one group of viruses infecting nanoeukaryotic algae. The amount of DMS accumulated per total DMSP or chlorophyll-a differed significantly between the present and future scenarios, suggesting that the pathways for DMS production or bacterial DMS consumption were affected by seawater pH. A comparison with previous work (PeECE II) suggests that DMS concentrations do not respond consistently to pelagic ecosystem CO2 enrichment experiments.