Effect of ocean acidification on the structure and fatty acid composition of a natural plankton community in the Baltic Sea

Increasing atmospheric carbon dioxide (CO2) is changing seawater chemistry towards reduced pH, which consequently affects various properties of marine organisms. Coastal and brackish water communities are expected to be less affected by ocean acidification (OA) as these communities are typically adapted to high fluctuations in CO2 and pH. Here we investigate the response of a coastal brackish water plankton community to increasing CO2 levels as projected for the coming decades and the end of this century in terms of community and biochemical fatty acid (FA) composition. A Baltic Sea plankton community was enclosed in a set of off-shore mesocosms and subjected to a CO2 gradient ranging from natural concentrations (~347 μatm pCO2) up to values projected for the year 2100 (~1333 μatm pCO2). We show that the phytoplankton community composition was resilient to CO2 and did not diverge between the treatments. Seston FA composition was influenced by community composition, which in turn was driven by silicate and phosphate limitation in the mesocosms, and showed no difference between the CO2 treatments. These results suggest that CO2 effects are dampened in coastal communities that already experience high natural fluctuations in pCO2. Although this coastal plankton community was tolerant to high pCO2 levels, hypoxia and CO2 uptake by the sea can aggravate acidification and may lead to pH changes outside the currently experienced range for coastal organisms

Effect of ocean acidification on the structure and fatty acid composition of a natural plankton community in the Baltic Sea

Increasing atmospheric carbon dioxide (CO2) is changing seawater chemistry towards reduced pH, which affects various properties of marine organisms. Coastal and brackish water communities are expected to be less affected by ocean acidification (OA) as these communities are typically adapted to high fluctuations in CO2 and pH. Here we investigate the response of a coastal brackish water plankton community to increasing CO2 levels as projected for the coming decades and the end of this century in terms of community and biochemical fatty acid (FA) composition. A Baltic Sea plankton community was enclosed in a set of offshore mesocosms and subjected to a CO2 gradient ranging from natural concentrations (similar to 347 mu atm fCO(2)) up to values projected for the year 2100 (similar to 1333 mu atm fCO(2)). We show that the phytoplankton community composition was resilient to CO2 and did not diverge between the treatments. Seston FA composition was influenced by community composition, which in turn was driven by silicate and phosphate limitation in the mesocosms and showed no difference between the CO2 treatments. These results suggest that CO2 effects are dampened in coastal communities that already experience high natural fluctuations in pCO(2). Although this coastal plankton community was tolerant of high pCO(2) levels, hypoxia and CO2 uptake by the sea can aggravate acidification and may lead to pH changes outside the currently experienced range for coastal organisms.Peer reviewe

No observed effect of ocean acidification on nitrogen biogeochemistry in a summer Baltic Sea plankton community

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

No observed effect of ocean acidification on nitrogen biogeochemistry in a summer Baltic Sea plankton community

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 neither inorganic nor organic N pool 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

Concentrations and Uptake of Dissolved Organic Phosphorus Compounds in the Baltic Sea

The dissolved organic phosphorus (DOP) pool in marine waters contains a variety of different compounds. Knowledge of the distribution and utilization of DOP by phyto- and bacterioplankton is limited, but critical to our understanding of the marine phosphorus cycle. In the Baltic Sea, detailed information about the composition of DOP and its turnover is lacking. This study reports the concentrations and uptake rates of DOP compounds, namely, adenosine triphosphate (dATP), deoxyribonucleic acid (dDNA), and phospholipids (dPL), in the Baltic Proper and in Finnish coastal waters in the summers of 2011 and 2012. Both areas differed in their dissolved inorganic phosphorus (DIP) concentrations (0.16 and 0.02–0.04 μM), in the C:P (123–178) and N:P (18–27) ratios, and in abundances of filamentous cyanobacteria and of autotrophic and heterotrophic picoplankton. The mean concentrations of dATP-P, dDNA-P, and dPL-P were 4.3–6.4, 0.05–0.12, and 1.9–6.8 nM, respectively, together contributing between 2.4 and 5.2% of the total DOP concentration. The concentrations of the compounds varied between and within the investigated regions and the distribution patterns of the individual components are not linked to each other. DIP was taken up at rates of 10.1–380.8 nM d-1. dATP-P and dDNA-P were consumed simultaneously with DIP at rates of 6.9–24.1 and 0.09–0.19 nM d-1, respectively, with the main proportion taken up by the size fraction <3 μm and with DIP to be the dominant source. Groups of hydrographical and biological parameters were identified in the multiple regression analysis to impact the concentrations and uptake rates. It points to the complexity of the regulation. Our results indicate that the investigated DOP compounds, particularly dATP-P, can make significant contributions to the P nutrition of microorganisms and their use seems to be not intertwined. Therefore, more detailed knowledge of all DOP components including variation of concentrations and the utilization is required to understand the roles of DOP in marine ecosystems

Temporal biomass dynamics of an Arctic plankton bloom in response to increasing levels of atmospheric carbon dioxide

Ocean acidification and carbonation, driven by anthropogenic emissions of carbon dioxide (CO2), have been shown to affect a variety of marine organisms and are likely to change ecosystem functioning. High latitudes, especially the Arctic, will be the first to encounter profound changes in carbonate chemistry speciation at a large scale, namely the under-saturation of surface waters with respect to aragonite, a calcium carbonate polymorph produced by several organisms in this region. During a CO2 perturbation study in Kongsfjorden on the west coast of Spitsbergen (Norway), in the framework of the EU-funded project EPOCA, the temporal dynamics of a plankton bloom was followed in nine mesocosms, manipulated for CO2 levels ranging initially from about 185 to 1420 μatm. Dissolved inorganic nutrients were added halfway through the experiment. Autotrophic biomass, as identified by chlorophyll a standing stocks (Chl a), peaked three times in all mesocosms. However, while absolute Chl a concentrations were similar in all mesocosms during the first phase of the experiment, higher autotrophic biomass was measured as high in comparison to low CO2 during the second phase, right after dissolved inorganic nutrient addition. This trend then reversed in the third phase. There were several statistically significant CO2 effects on a variety of parameters measured in certain phases, such as nutrient utilization, standing stocks of particulate organic matter, and phytoplankton species composition. Interestingly, CO2 effects developed slowly but steadily, becoming more and more statistically significant with time. The observed CO2-related shifts in nutrient flow into different phytoplankton groups (mainly dinoflagellates, prasinophytes and haptophytes) could have consequences for future organic matter flow to higher trophic levels and export production, with consequences for ecosystem productivity and atmospheric CO2.publishedVersio

Negligible effects of ocean acidification on Eurytemora affinis (Copepoda) offspring production

Ocean acidification is caused by increasing amounts of carbon dioxide dissolving in the oceans leading to lower seawater pH. We studied the effects of lowered pH on the calanoid copepod Eurytemora affinis during a mesocosm experiment conducted in a coastal area of the Baltic Sea. We measured copepod reproductive success as a function of pH, chlorophyll a concentration, diatom and dinoflagellate biomass, carbon to nitrogen (C : N) ratio of suspended particulate organic matter, as well as copepod fatty acid composition. The laboratory-based experiment was repeated four times during 4 consecutive weeks, with water and copepods sampled from pelagic mesocosms enriched with different CO2 concentrations. In addition, oxygen radical absorbance capacity (ORAC) of animals from the mesocosms was measured weekly to test whether the copepod's defence against oxidative stress was affected by pH. We found no effect of pH on offspring production. Phytoplankton biomass, as indicated by chlorophyll a concentration and dinoflagellate biomass, had a positive effect. The concentration of polyunsaturated fatty acids in the females was reflected in the eggs and had a positive effect on offspring production, whereas monounsaturated fatty acids of the females were reflected in their eggs but had no significant effect. ORAC was not affected by pH. From these experiments we conclude that E. affinis seems robust against direct exposure to ocean acidification on a physiological level, for the variables covered in the study. E. affinis may not have faced acute pH stress in the treatments as the species naturally face large pH fluctuations.Peer reviewe

Effect of ocean acidification and elevated fCO2 on trace gas production from the Baltic Sea summer phytoplankton community.

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

Ocean acidification decreases plankton respiration : evidence from a mesocosm experiment

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 (similar to 55 m(3)) were set up in the Baltic Sea with a gradient of CO2 levels initially ranging from ambient (similar to 240 mu atm), used as control, to high CO2 (up to similar to 1330 mu atm). 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 similar to 26% at t0 to similar to 8% at t31, probably driven by a shift towards smaller plankton (<4 mu m) 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.Peer reviewe

Effects of ocean acidification on pelagic carbon fluxes in a mesocosm experiment

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 (similar to 370 mu atm) to high (similar to 1200 mu atm), were set up in mesocosm bags (similar to 55m(3)). 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 Cm-2 at the start of the experiment, and the initial CO2 additions increased the DIC pool by similar to 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 similar to 100 mmol C m(-2) day(-1), from which 75-95% was respired, similar to 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 similar 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 similar to 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.Peer reviewe