Description: Every year, the oceans absorb about 30% of anthropogenic carbon dioxide (CO2) leading to a re-equilibration of the marine carbonate system and decreasing seawater pH. Today, there is increasing awareness that these changes–summarized by the term ocean acidification (OA)–could differentially affect the competitive ability of marine organisms, thereby provoking a restructuring of marine ecosystems and biogeochemical element cycles. In winter 2013, we deployed ten pelagic mesocosms in the Gullmar Fjord at the Swedish west coast in order to study the effect of OA on plankton ecology and biogeochemistry under close to natural conditions. Five of the ten mesocosms were left unperturbed and served as controls (~380 μatm pCO2), whereas the others were enriched with CO2-saturated water to simulate realistic end-of-the-century carbonate chemistry conditions (~760 μatm pCO2). We ran the experiment for 113 days which allowed us to study the influence of high CO2 on an entire winter-to-summer plankton succession and to investigate the potential of some plankton organisms for evolutionary adaptation to OA in their natural environment. This paper is the first in a PLOS collection and provides a detailed overview on the experimental design, important events, and the key complexities of such a “long-term mesocosm” approach. Furthermore, we analyzed whether simulated end-of-the-century carbonate chemistry conditions could lead to a significant restructuring of the plankton community in the course of the succession. At the level of detail analyzed in this overview paper we found that CO2-induced differences in plankton community composition were non-detectable during most of the succession except for a period where a phytoplankton bloom was fueled by remineralized nutrients. These results indicate: (1) Long-term studies with pelagic ecosystems are necessary to uncover OA-sensitive stages of succession. (2) Plankton communities fueled by regenerated nutrients may be more responsive to changing carbonate chemistry than those having access to high inorganic nutrient concentrations and may deserve particular attention in future studies.

Description: Sediment traps are the most common tool to investigate vertical particle flux in the marine realm. However, the spatial and temporal decoupling between particle formation in the surface ocean and particle collection in sediment traps at depth often handicaps reconciliation of production and sedimentation even within the euphotic zone. Pelagic mesocosms are restricted to the surface ocean, but have the advantage of being closed systems and are therefore ideally suited to studying how processes in natural plankton communities influence particle formation and settling in the ocean's surface. We therefore developed a protocol for efficient sample recovery and processing of quantitatively collected pelagic mesocosm sediment trap samples for biogeochemical analysis. Sedimented material was recovered by pumping it under gentle vacuum through a silicon tube to the sea surface. The particulate matter of these samples was subsequently separated from bulk seawater by passive settling, centrifugation or flocculation with ferric chloride, and we discuss the advantages and efficiencies of each approach. After concentration, samples were freeze-dried and ground with an easy to adapt procedure using standard lab equipment. Grain size of the finely ground samples ranged from fine to coarse silt (2–63 µm), which guarantees homogeneity for representative subsampling, a widespread problem in sediment trap research. Subsamples of the ground material were perfectly suitable for a variety of biogeochemical measurements, and even at very low particle fluxes we were able to get a detailed insight into various parameters characterizing the sinking particles. The methods and recommendations described here are a key improvement for sediment trap applications in mesocosms, as they facilitate the processing of large amounts of samples and allow for high-quality biogeochemical flux data.

Description: Anthropogenic carbon dioxide (CO2) emissions are reducing the pH in the world's oceans. The plankton community is a key component driving biogeochemical fluxes, and the effect of increased CO2 on plankton is critical for understanding the ramifications of ocean acidification on global carbon fluxes. We determined the plankton community composition and measured primary production, respiration rates and carbon export (defined here as carbon sinking out of a shallow, coastal area) during an ocean acidification experiment. Mesocosms ( ∼  55 m3) were set up in the Baltic Sea with a gradient of CO2 levels initially ranging from ambient ( ∼  240 µatm), used as control, to high CO2 (up to  ∼  1330 µatm). The phytoplankton community was dominated by dinoflagellates, diatoms, cyanobacteria and chlorophytes, and the zooplankton community by protozoans, heterotrophic dinoflagellates and cladocerans. The plankton community composition was relatively homogenous between treatments. Community respiration rates were lower at high CO2 levels. The carbon-normalized respiration was approximately 40 % lower in the high-CO2 environment compared with the controls during the latter phase of the experiment. We did not, however, detect any effect of increased CO2 on primary production. This could be due to measurement uncertainty, as the measured total particular carbon (TPC) and combined results presented in this special issue suggest that the reduced respiration rate translated into higher net carbon fixation. The percent carbon derived from microscopy counts (both phyto- and zooplankton), of the measured total particular carbon (TPC), decreased from  ∼  26 % at t0 to  ∼  8 % at t31, probably driven by a shift towards smaller plankton (〈 4 µm) not enumerated by microscopy. Our results suggest that reduced respiration leads to increased net carbon fixation at high CO2. However, the increased primary production did not translate into increased carbon export, and consequently did not work as a negative feedback mechanism for increasing atmospheric CO2 concentration.

Description: About a quarter of anthropogenic CO2 emissions are currently taken up by the oceans, decreasing seawater pH. We performed a mesocosm experiment in the Baltic Sea in order to investigate the consequences of increasing CO2 levels on pelagic carbon fluxes. A gradient of different CO2 scenarios, ranging from ambient ( ∼  370 µatm) to high ( ∼  1200 µatm), were set up in mesocosm bags ( ∼  55 m3). We determined standing stocks and temporal changes of total particulate carbon (TPC), dissolved organic carbon (DOC), dissolved inorganic carbon (DIC), and particulate organic carbon (POC) of specific plankton groups. We also measured carbon flux via CO2 exchange with the atmosphere and sedimentation (export), and biological rate measurements of primary production, bacterial production, and total respiration. The experiment lasted for 44 days and was divided into three different phases (I: t0–t16; II: t17–t30; III: t31–t43). Pools of TPC, DOC, and DIC were approximately 420, 7200, and 25 200 mmol C m−2 at the start of the experiment, and the initial CO2 additions increased the DIC pool by  ∼  7 % in the highest CO2 treatment. Overall, there was a decrease in TPC and increase of DOC over the course of the experiment. The decrease in TPC was lower, and increase in DOC higher, in treatments with added CO2. During phase I the estimated gross primary production (GPP) was  ∼  100 mmol C m−2 day−1, from which 75–95 % was respired,  ∼  1 % ended up in the TPC (including export), and 5–25 % was added to the DOC pool. During phase II, the respiration loss increased to  ∼  100 % of GPP at the ambient CO2 concentration, whereas respiration was lower (85–95 % of GPP) in the highest CO2 treatment. Bacterial production was  ∼  30 % lower, on average, at the highest CO2 concentration than in the controls during phases II and III. This resulted in a higher accumulation of DOC and lower reduction in the TPC pool in the elevated CO2 treatments at the end of phase II extending throughout phase III. The “extra” organic carbon at high CO2 remained fixed in an increasing biomass of small-sized plankton and in the DOC pool, and did not transfer into large, sinking aggregates. Our results revealed a clear effect of increasing CO2 on the carbon budget and mineralization, in particular under nutrient limited conditions. Lower carbon loss processes (respiration and bacterial remineralization) at elevated CO2 levels resulted in higher TPC and DOC pools than ambient CO2 concentration. These results highlight the importance of addressing not only net changes in carbon standing stocks but also carbon fluxes and budgets to better disentangle the effects of ocean acidification.

Description: Nitrogen fixation by filamentous cyanobacteria supplies significant amounts of new nitrogen (N) to the Baltic Sea. This balances N loss processes such as denitrification and anammox and forms an important N source supporting primary and secondary production in N-limited post-spring bloom plankton communities. Laboratory studies suggest that filamentous diazotrophic cyanobacteria growth and N2-fixation rates are sensitive to ocean acidification with potential implications for new N supply to the Baltic Sea. In this study, our aim was to assess the effect of ocean acidification on diazotroph growth and activity as well as the contribution of diazotrophically-fixed N to N supply in a natural plankton assemblage. We enclosed a natural plankton community in a summer season in the Baltic Sea near the entrance to the Gulf of Finland in six large-scale mesocosms (volume ~ 55 m3) and manipulated fCO2 over a range relevant for projected ocean acidification by the end of this century (average treatment fCO2: 365–1231 μatm). The direct response of diazotroph growth and activity was followed in the mesocosms over a 47 day study period during N-limited growth in the summer plankton community. Diazotrophic filamentous cyanobacteria abundance throughout the study period and N2-fixation rates (determined only until day 21 due to subsequent use of contaminated commercial 15N-N2 gas stocks) remained low. Thus estimated new N inputs from diazotrophy were too low to relieve N limitation and stimulate a summer phytoplankton bloom. Instead regeneration of organic N sources likely sustained growth in the plankton community. We could not detect significant CO2-related differences in inorganic or organic N pools sizes, or particulate matter N : P stoichiometry. Additionally, no significant effect of elevated CO2 on diazotroph activity was observed. Therefore, ocean acidification had no observable impact on N cycling or biogeochemistry in this N-limited, post-spring bloom plankton assemblage in the Baltic Sea.

Description: Studies investigating the effect of increasing CO2 levels on the phosphorus cycle in natural waters are lacking although phosphorus often controls phytoplankton development in aquatic systems. The aim of our study was to analyze effects of elevated CO2 levels on phosphorus pool sizes and uptake. Therefore, we conducted a CO2-manipulation mesocosm experiment in the Storfjärden (western Gulf of Finland, Baltic Sea) in summer 2012. We compared the phosphorus dynamics in different mesocosm treatments but also studied them outside the mesocosms in the surrounding fjord water. In the mesocosms as well as in surface waters of Storfjärden, dissolved organic phosphorus (DOP) concentrations of 0.26 ± 0.03 and 0.23 ± 0.04 μmol L−1, respectively, formed the main fraction of the total P-pool (TP), whereas phosphate (PO4) constituted the lowest fraction with mean concentration of 0.15 ± 0.02 μmol L−1 and 0.17 ± 0.07 μmol L−1 in the mesocosms and in the fjord, respectively. Uptake of PO4 ranged between 0.6 and 3.9 nmol L−1 h−1 of which ~ 86 % (mesocosms) and ~ 72 % (fjord) were realized by the size fraction 〈 3 μm. Adenosine triphosphate (ATP) uptake revealed that additional P was supplied from organic compounds accounting for 25–27 % of P provided by PO4 only. CO2 additions did not cause significant changes in phosphorus (P) pool sizes, DOP composition, and uptake of PO4 and ATP when the whole study period was taken into account. About 18 % of PO4 was transformed into POP, whereby the major proportion (~ 82 %) was converted into DOP suggesting that the conversion of PO4 to DOP is the main pathway of the PO4 turnover. We observed that significant relationships (e.g., between POP and Chl a) in the untreated mesocosms vanished under increased fCO2 conditions. Consequently, it can be hypothesized that the relationship between POP formation and phytoplankton growth changed under elevated CO2 conditions. Significant short-term effects were observed for PO4 and particulate organic phosphorus (POP) pool sizes in CO2 treatments 〉 1000 μatm during periods when phytoplankton started to grow.

Description: Anthropogenic carbon dioxide (CO2) emissions are reducing the pH in the world's oceans. The plankton community is a key component driving biogeochemical fluxes, and the effect of increased CO2 on plankton is critical for understanding the ramifications of ocean acidification on global carbon fluxes. We determined the plankton community composition and measured primary production, respiration rates and carbon export (defined here as carbon sinking out of a shallow, coastal area) during an ocean acidification experiment. Mesocosms (~ 55 m3) were set up in the Baltic Sea with a gradient of CO2 levels initially ranging from ambient (~ 240 µatm), used as control, to high CO2 (up to ~ 1330 µatm). The phytoplankton community was dominated by dinoflagellates, diatoms, cyanobacteria and chlorophytes, and the zooplankton community by protozoans, heterotrophic dinoflagellates and cladocerans. The plankton community composition was relatively homogenous between treatments. Community respiration rates were lower at high CO2 levels. The carbon-normalized respiration was approximately 40 % lower in the high CO2 environment compared with the controls during the latter phase of the experiment. We did not, however, detect any effect of increased CO2 on primary production. This could be due to measurement uncertainty, as the measured total particular carbon (TPC) and combined results presented in this special issue suggest that the reduced respiration rate translated into higher net carbon fixation. The percent carbon derived from microscopy counts (both phyto- and zooplankton), of the measured total particular carbon (TPC) decreased from ~ 26 % at t0 to ~ 8 % at t31, probably driven by a shift towards smaller plankton (〈 4 µm) not enumerated by microscopy. Our results suggest that reduced respiration lead to increased net carbon fixation at high CO2. However, the increased primary production did not translate into increased carbon export, and did consequently not work as a negative feedback mechanism for increasing atmospheric CO2 concentration.

Description: Coccolithophores -single-celled calcifying phytoplankton- are an important group of marine primary producers and the dominant builders of calcium carbonate globally. Coccolithophores form extensive blooms and increase the density and sinking speed of organic matter via calcium carbonate ballasting. Thereby, they play a key role in the marine carbon cycle. Coccolithophore physiological responses to experimental ocean acidification have ranged from moderate stimulation to substantial decline in growth and calcification rates, combined with enhanced malformation of their calcite platelets. Here we report on a mesocosm experiment conducted in a Norwegian fjord in which we exposed a natural plankton community to a wide range of CO2-induced ocean acidification, to test whether these physiological responses affect the ecological success of coccolithophore populations. Under high-CO2 treatments, Emiliania huxleyi, the most abundant and productive coccolithophore species, declined in population size during the pre-bloom period and lost the ability to form blooms. As a result, particle sinking velocities declined by up to 30% and sedimented organic matter was reduced by up to 25% relative to controls. There were also strong reductions in seawater concentrations of the climate-active compound dimethylsulfide in CO2-enriched mesocosms. We conclude that ocean acidification can lower calcifying phytoplankton productivity, potentially creating a positive feedback to the climate system.

Description: Every year, the oceans absorb about 30% of anthropogenic carbon dioxide (CO2) leading to a re-equilibration of the marine carbonate system and decreasing seawater pH. Today, there is increasing awareness that these changes–summarized by the term ocean acidification (OA)–could differentially affect the competitive ability of marine organisms, thereby provoking a restructuring of marine ecosystems and biogeochemical element cycles. In winter 2013, we deployed ten pelagic mesocosms in the Gullmar Fjord at the Swedish west coast in order to study the effect of OA on plankton ecology and biogeochemistry under close to natural conditions. Five of the ten mesocosms were left unperturbed and served as controls (~380 μatm pCO2), whereas the others were enriched with CO2-saturated water to simulate realistic end-of-the-century carbonate chemistry conditions (~760 μatm pCO2). We ran the experiment for 113 days which allowed us to study the influence of high CO2 on an entire winter-to-summer plankton succession and to investigate the potential of some plankton organisms for evolutionary adaptation to OA in their natural environment. This paper is the first in a PLOS collection and provides a detailed overview on the experimental design, important events, and the key complexities of such a “long-term mesocosm” approach. Furthermore, we analyzed whether simulated end-of-the-century carbonate chemistry conditions could lead to a significant restructuring of the plankton community in the course of the succession. At the level of detail analyzed in this overview paper we found that CO2-induced differences in plankton community composition were non-detectable during most of the succession except for a period where a phytoplankton bloom was fueled by remineralized nutrients. These results indicate: (1) Long-term studies with pelagic ecosystems are necessary to uncover OA-sensitive stages of succession. (2) Plankton communities fueled by regenerated nutrients may be more responsive to changing carbonate chemistry than those having access to high inorganic nutrient concentrations and may deserve particular attention in future studies.