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Marine calcareous sediments provide a fundamental basis for paleoceanographic 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. The results demonstrate the complex interaction between environmental forcing and physiological control on the strontium partitioning in coccolithophore calcite. 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 results give an extended insight into the driving factors that lead to variations in the coccolith Mg/ Ca ratio and can be used for Sr / Ca and Mg/Ca paleoproxy calibration. [ABSTRACT FROM AUTHOR]

Natural variability in seawater pH and associated carbonate chemistry parameters is in part driven by biological activities such as photosynthesis and respiration. The amplitude of these variations is expected to increase with increasing seawater carbon dioxide (CO) concentrations in the future, because of simultaneously decreasing buffer capacity. Here, we address this experimentally during a diurnal cycle in a mesocosm CO perturbation study. We show that for about the same amount of dissolved inorganic carbon (DIC) utilized in net community production diel variability in proton (H) and CO concentrations was almost three times higher at CO levels of about 675 ± 65 in comparison with levels of 310 ± 30 μatm. With a simple model, adequately simulating our measurements, we visualize carbonate chemistry variability expected for different oceanic regions with relatively low or high net community production. Since enhanced diurnal variability in CO and proton concentration may require stronger cellular regulation in phytoplankton to maintain respective gradients, the ability to adjust may differ between communities adapted to low in comparison with high natural variability. [ABSTRACT FROM AUTHOR]

Net community production (NCP) and carbon to nutrient uptake ratios were studied during a large-scale mesocosm experiment on ocean acidification in Kongsfjorden, western Svalbard, during June--July 2010. Nutrient depleted fjord water with natural plankton assemblages, enclosed in nine mesocosms of ~50m³ in volume, was exposed to pCO2 levels ranging initially from 185 to 1420 μatm. NCP estimations are the cumulative change in dissolved inorganic carbon concentrations after accounting for gas exchange and total alkalinity variations. Stoichiometric coupling between inorganic carbon and nutrient net uptake is shown as a ratio of NCP to a cumulative change in inorganic nutrients. Phytoplankton growth was stimulated by nutrient addition half way through the experiment and three distinct peaks in chlorophyll a concentration were observed during the experiment. Accordingly, the experiment was divided in three phases. Cumulative NCP was similar in all mesocosms over the duration of the experiment. However, in phases I and II, NCP was higher and in phase III lower at elevated pCO2. Due to relatively low inorganic nutrient concentration in phase I, C :N and C : P uptake ratios were calculated only for the period after nutrient addition (phase II and phase III). For the total post-nutrient period (phase II+phase III) ratios were close to Redfield, however they were lower in of NCP, C :N and C : P uptake ratios in different phases reflects the effect of increasing CO2 on phytoplankton community composition and succession. The phytoplankton community was composed predominantly of haptophytes in phase I, prasinophytes, dinoflagellates, and cryptophytes in phase II, and haptophytes, prasinophytes, dinoflagellates and chlorophytes in phase III (Schulz et al., 2013). Increasing ambient inorganic carbon concentrations have also been shown to promote primary production and carbon assimilation. For this study, it is clear that the pelagic ecosystem response to increasing CO2 is more complex than that represented in previous work, e.g. Bellerby et al. (2008). Carbon and nutrient uptake representation in models should, where possible, be more focused on individual plankton functional types as applying a single stoichiometry to a biogeochemical model with regard to the effect of increasing pCO2 may not always be optimal. The phase variability in NCP and stoichiometry may be better understood if CO2 sensitivities of the plankton's functional type biogeochemical uptake kinetics and trophic interactions are better constrained. [ABSTRACT FROM AUTHOR]

The volume of water enclosed inside flexible-wall mesocosm bags is hard to estimate using geometrical calculations and can be strongly variable among bags of the same dimensions. Here we present a method for precise water volume determination in mesocosms using salinity as a tracer. Knowledge of the precise volume of water enclosed allows establishment of exactly planned treatment concentrations and calculation of elemental budgets. [ABSTRACT FROM AUTHOR]

One of the great challenges in ocean change research is to understand and forecast the effects of environmental changes on pelagic communities and the associated impacts on biogeochemical cycling. Mesocosms, experimental enclosures designed to approximate natural conditions, and in which environmental factors can be manipulated and closely monitored, provide a powerful tool to close the gap between small-scale laboratory experiments and observational and correlative approaches applied in field surveys. Existing pelagic mesocosm systems are stationary and/or restricted to well-protected waters. To allow mesocosm experimentation in a range of hydrographic conditions and in areas considered most sensitive to ocean change, we developed a mobile sea-going mesocosm facility, the Kiel Off-Shore Mesocosms for Future Ocean Simulations (KOSMOS). The KOSMOS platform, which can be transported and deployed by mid-sized research vessels, is designed for operation in moored and free-floating mode under low to moderate wave conditions (up to 2.5 m wave heights). It encloses a water column 2 m in diameter and 15 to 25 m deep (∼50-75 m3 in volume) without disrupting the vertical structure or disturbing the enclosed plankton community. Several new developments in mesocosm design and operation were implemented to (i) minimize differences in starting conditions between mesocosms, (ii) allow for extended experimental duration, (iii) precisely determine the mesocosm volume, (iv) determine air-sea gas exchange, and (v) perform mass balance calculations. After multiple test runs in the Baltic Sea, which resulted in continuous improvement of the design and handling, the KOSMOS platform successfully completed its first full-scale experiment in the high Arctic off Svalbard (78°56.2' N, 11°53.6' E) in June/July 2010. The study, which was conducted in the framework of the European Project on Ocean Acidification (EPOCA), focused on the effects of ocean acidification on a natural plankton community and its impacts on biogeochemical cycling and air-sea exchange of climate-relevant gases. This manuscript describes the mesocosm hardware, its deployment and handling, CO2 manipulation, sampling and cleaning, including some further modifications conducted based on the experiences gained during this study. [ABSTRACT FROM AUTHOR]

Impacts of ocean acidification (OA) on marine biota have been observed in a wide range of marine systems. We used a mesocosm approach to study the response of a high Arctic coastal microzooplankton community during the post-bloom period in Kongsfjorden (Svalbard) to direct and indirect effects of high pCO2/low pH. We found almost no direct effects of OA on microzooplankton composition and diversity. Both the relative shares of ciliates and heterotrophic dinoflagellates as well as the taxonomic composition of microzooplankton remained unaffected by changes in pCO2/pH. Although the different pCO2 treatments affected food availability and phytoplankton composition, no indirect effects (e.g. on the total carrying capacity and phenology of microzooplankton) could be observed. Our data point to a high tolerance of this Arctic microzooplankton community to changes in pCO2/pH. Future studies on the impact of OA on plankton communities should include microzooplankton in order to test whether the observed low sensitivity to OA is typical for coastal communities where changes in seawater pH occur frequently. [ABSTRACT FROM AUTHOR]

The effect of CO2 on carbon fluxes (production, consumption, and export) in Arctic plankton communities was investigated during the 2010 EPOCA (European project on Ocean Acidification) mesocosm study off Ny Ålesund, Svalbard. 13C labelled bicarbonate was added to nine mesocosms with a range in pCO2 (185 to 1420 μatm) to follow the transfer of carbon from dissolved inorganic carbon (DIC) into phytoplankton, bacterial and zooplankton consumers, and export. 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. The plankton community structure was characteristic for a post-bloom situation and retention food web and showed high bacterial production (∼ 31% of primary production), high abundance of mixotrophic phytoplankton, low mesozooplankton grazing (☤ 6% of primary production) and low export (∼ 7% of primary production). Zooplankton grazing and export of detritus were sensitive to CO2: grazing decreased and export increased with increasing pCO2. Nutrient addition halfway through the experiment increased the export, but not the production rates. Although mixotrophs showed initially higher production rates with increasing CO2, the overall production of POC (particulate organic carbon) after nutrient addition decreased with increasing CO2. Interestingly, and contrary to the low nutrient situation, much more material settled down in the sediment traps at low CO2. The observed CO2 related effects potentially alter future organic carbon flows and export, with possible consequences for the efficiency of the biological pump. [ABSTRACT FROM AUTHOR]

Mesocosms as large experimental units provide the opportunity to perform elemental mass balance calculations, e.g. to derive net biological turnover rates. However, the system is in most cases not closed at the water surface and gases exchange with the atmosphere. Previous attempts to budget carbon pools in mesocosms relied on educated guesses concerning the exchange of CO2 with the atmosphere. Here, we present a simple method for precise determination of air-sea gas exchange in mesocosms using N2O as a deliberate tracer. Beside the application for carbon budgeting, transfer velocities can be used to calculate exchange rates of any gas of known concentration, e.g. to calculate aquatic production rates of climate relevant trace gases. Using an arctic KOSMOS (Kiel Off Shore Mesocosms for future Ocean Simulation) experiment as an exemplary dataset, it is shown that the presented method improves accuracy of carbon budget estimates substantially. Methodology of manipulation, measurement, data processing and conversion to CO2 fluxes are explained. A theoretical discussion of prerequisites for precise gas exchange measurements provides a guideline for the applicability of the method under various experimental conditions. [ABSTRACT FROM AUTHOR]

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: 43.9-47.6m³), 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 on average sufficient carbon for synthesis of bacterial biomass. During the later course of the bloom, the response of 14C-based PP rates to CO2 enrichment differed from net community production (NCP) rates that were also determined during this mesocosm campaign. 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. [ABSTRACT FROM AUTHOR]

The Arctic Ocean ecosystem is particularly vulnerable to ocean acidification (OA) related alterations due to the relatively high CO2 solubility and low carbonate saturation states of its cold surface waters. Thus far, however, there is only little known about the consequences of OA on the base of the food web. In a mesocosm CO2-enrichment experiment (overall CO2 levels ranged from ~180 to 1100 µatm) in Kongsfjorden off Svalbard, we studied the consequences of OA on a natural pelagic microbial community. OA distinctly affected the composition and growth of the Arctic phytoplankton community, i.e. the picoeukaryotic photoau- totrophs and to a lesser extent the nanophytoplankton thrived. A shift towards the smallest phytoplankton as a result of OA will have direct consequences for the structure and functioning of the pelagic food web and thus for the biogeochemical cycles. Besides being grazed, the dominant pico- and nanophytoplankton groups were found prone to viral lysis, thereby shunting the carbon accumulation in living organisms into the dissolved pools of organic carbon and subsequently affecting the efficiency of the biological pump in these Arctic waters. [ABSTRACT FROM AUTHOR]

The effect of ocean acidification on the fatty acid composition of a natural plankton community in the Arctic was studied in a large-scale mesocosm experiment, carried out in Kongsfjorden (Svalbard, Norway) at 79°N. Ninemesocosms of ~50 m³ each were exposed to 8 different pCO2 levels (from natural background conditions to ~1420 µatm), yielding pH values (on the total scale) from ~8.3 to 7.5. Inorganic nutrients were added on day 13. The phytoplank- ton development during this 30-day experiment passed three distinct phases: (1) prior to the addition of inorganic nutrients, (2) first bloom after nutrient addition, and (3) second bloom after nutrient addition. The fatty acid composition of the natural plankton community was analysed and showed, in general, high percentages of polyunsaturated fatty acids (PUFAs): 44–60% of total fatty acids. Positive correlations with pCO2 were found for most PUFAs during phases 2 and/or 3, with the exception of 20:5n3 (eicosapentaenoic acid, EPA), an important diatom marker. These correlations are probably linked to changes in taxonomic composition in response to pCO2. While diatoms (together with prasinophytes and hap- tophytes) increased during phase 3 mainly in the low and intermediate pCO2 treatments, dinoflagellates were favoured by high CO2 concentrations during the same time period. This is reflected in the development of group-specific fatty acid trophic markers. No indications were found for a generally detrimental effect of ocean acidification on the planktonic food quality in terms of essential fatty acids. [ABSTRACT FROM AUTHOR]

The impact of ocean acidification and carbonation on microbial community structure was assessed during a large-scale in situ costal pelagic mesocosm study, included as part of the EPOCA 2010 Arctic campaign. The mesocosm experiment included ambient conditions (fjord) and nine mesocosms with pCO2 levels ranging from ~145 to ~1420 µatm. Samples for the present study were collected at ten time points (t-1, t1, t5, t7, t12, t14, t18, t22, t26 to t28) in seven treatments (ambient fjord (~145), 2 x ~185, ~270, ~685, ~820, ~1050 µatm) and were analysed for "small" and "large" size fraction microbial community composition using 16S rRNA (ribosomal ribonucleic acid) amplicon sequencing. This high-throughput sequencing analysis produced ~20 000 000 16S rRNA V4 reads, which comprised 7000 OTUs. The main variables structuring these communities were sample origins (fjord or mesocosms) and the community size fraction (small or large size fraction). The community was significantly different between the unenclosed fjord water and enclosed mesocosms (both control and elevated CO2 treatments) after nutrients were added to the mesocosms, suggesting that the addition of nutrients is the primary driver of the change in mesocosm community structure. The relative importance of each structuring variable depended greatly on the time at which the community was sampled in relation to the phytoplankton bloom. The sampling strategy of separating the small and large size fraction was the second most important factor for community structure. When the small and large size fraction bacteria were analysed separately at different time points, the only taxon pCO2 was found to significantly affect were the Gamma proteobacteria after nutrient addition. Finally, pCO2 treatment was found to be significantly correlated (non-linear) with 15 rare taxa, most of which increased in abundance with higher CO2. [ABSTRACT FROM AUTHOR]

The effect of ocean acidification on the balance between gross community production (GCP) and community respiration (CR) (i.e., net community production, NCP) of plankton communities was investigated in summer 2010 in Kongsfjorden, west of Svalbard. Surface water, which was characterized by low concentrations of dissolved inorganic nutrients and chlorophyll α (a proxy of phytoplankton biomass), was enclosed in nine mesocosms and subjected to eight pCO2 levels (two replicated controls and seven enhanced pCO2 treatments) for one month. Nutrients were added to all mesocosms on day 13 of the experiment, and thereafter increase of chlorophyll α was provoked in all mesocosms. No clear trend in response to increasing pCO2 was found in the daily values of NCP, CR, and GCP. For further analysis, these parameters were cumulated for the following three periods: phase 1 - end of CO2 manipulation until nutrient addition (t4 to t13); phase 2 - nutrient addition until the second chlorophyll α minimum (t14 to t21); phase 3 - the second chlorophyll α minimum until the end of this study (t22 to t28). A significant response was detected as a decrease of NCP with increasing pCO2 during phase 3. CR was relatively stable throughout the experiment in all mesocosms. As a result, the cumulative GCP significantly decreased with increasing pCO2 during phase 3. After the nutrient addition, the ratios of cumulative NCP to cumulative consumption of NO3 and PO4 showed a significant decrease during phase 3 with increasing pCO2. The results suggest that elevated pCO2 influenced cumulative NCP and stoichio-metric C and nutrient coupling of the plankton community in a high-latitude fjord only for a limited period. However provided that there were some differences or weak correlations between NCP data based on different methods in the same experiment, this conclusion should be taken with caution. [ABSTRACT FROM AUTHOR]

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 activity of natural extracellular enzyme assemblages increased in response to acidification. Rates of β-glucosidase and leucine-aminopeptidase increased along the gradient of mesocosm pCO2. A decrease in seawater pH of 0.5 units almost doubled rates of both enzymes. Heterotrophic bacterial activity was closely coupled to phytoplankton productivity in this experiment. The bacterio-plankton community responded to rising chlorophyll a concentrations after a lag phase of only a few days with increasing protein production and extracellular enzyme activity. Time-integrated primary production and bacterial protein production were positively correlated, strongly suggesting that higher amounts of phytoplankton-derived organic matter were assimilated by heterotrophic bacteria at increased primary production. Primary production increased under high pCO2 in this study, and it can be suggested that the efficient heterotrophic carbon utilisation had the potential to counter-act the enhanced autotrophic CO2 fixation. However, our results also show that beneficial pCO2-related effects on bacterial activity can be mitigated by the top-down control of bacterial abundances in natural microbial communities. [ABSTRACT FROM AUTHOR]

In the frame of the European Project on Ocean Acidification (EPOCA), the response of an Arctic pelagic community (<3mm) to a gradient of seawater 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. [ABSTRACT FROM AUTHOR]

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 α), peaked three times in all mesocosms. However, while absolute Chl α 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. [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]

The impact of ocean acidification and carbonation on microbial community structure was assessed during a large-scale in situ costal pelagic mesocosm study, included as part of the EPOCA 2010 Arctic campaign. The mesocosm experiment included ambient conditions (fjord) and nine mesocosms, with pCO2 range from ~145 to ~1420 µatm. Samples collected at nine time points (t-1, t1, t5, t7, t12, t14, t22, t26 to t28) in seven treatments (ambient fjord (~145), 2x ~185, ~270, ~685, ~820, ~1050 µatm) were analysed for "free-living" and "particle associated" microbial community composition using 16S rRNA amplicon sequencing. This high-throughput sequencing analysis produced ~20 000 000 16S rRNA V4 reads, which comprised 7000 OTUs. The main variables structuring these communities were, sample origin (fjord or mesocosms) and the filter size fraction (free-living or particle associated). The community was significantly different between the fjord and both the control and elevated CO2 mesocosms (which were not significant different) after nutrients were added to the mesocosms; suggesting that the addition of nutrients is the primary driver of the change in mesocosm community structure. The relative importance of each structuring variable depended greatly on the time at which the community was sampled in relation to the phytoplankton bloom. The size fraction was the second most important factor for community structure; separating free-living from particle-associated bacteria. When free-living and particleassociated bacteria were analysed separately at different time points, the only taxon pCO2 was found to significantly affect were the Gammaproteobacteria after nutrient addition. Finally, pCO2 treatment was found to be significantly correlated (non-linear) with 15 rare taxa, most of which increased in abundance with higher CO2. [ABSTRACT FROM AUTHOR]

Impacts of ocean acidification (OA) on marine biota have been observed in a wide range of marine systems. We used a mesocosm approach to study the response of a high Arctic coastal protozooplankton (PZP in the following) community during the post-bloom period in the Kongsfjorden (Svalbard) to direct and indirect effects of high pCO2/low pH. We found almost no direct effects of OA on PZP composition and diversity. Both, the relative shares of ciliates and heterotrophic dinoflagellates as well as the taxonomic composition of protozoans remained unaffected by changes in pCO2/pH. The different pCO2 treatments did not have any effect on food availability and phytoplankton composition and thus no indirect effects e.g. on the total carrying capacity and phenology of PZP could be observed. Our data points at a high tolerance of this Arctic PZP community to changes in pCO2/pH. Future studies on the impact of OA on plankton communities should include PZP in order to test whether the observed low sensitivity of protozoans to OA is typical for coastal communities where changes in seawater pH occur frequently. [ABSTRACT FROM AUTHOR]

One of the great challenges in ocean change research is to understand and forecast the effects of environmental changes on pelagic communities and the associated impacts on biogeochemical cycling. Mesocosms, experimental enclosures designed to approximate natural conditions, and in which environmental factors can be manipulated and closely monitored, provide a powerful tool to close the gap between single species laboratory experiments and observational and correlative approaches applied in field surveys. Existing pelagic mesocosm systems are stationary and/or restricted to well-protected waters. To allow mesocosm experimentation in a range of hydrographic conditions and in areas considered most sensitive to ocean change, we developed a mobile, sea-going mesocosm facility, the Kiel Off-Shore Mesocosms for Future Ocean Simulations (KOSMOS). The KOSMOS platform, which can be transported and deployed by mid-sized research vessels, is designed for operation in moored and free- floating mode under low to moderate wave conditions (up to 2.5m wave heights). It encloses a water column 2m in diameter and 15 to 25m deep (~50-75m³ in volume) without disrupting the vertical structure or disturbing the enclosed plankton community. Several new developments in mesocosm design and operation were implemented to (i) minimize differences in starting conditions between mesocosms, (ii) allow for extended experimental duration, (iii) precisely determine the mesocosm volume, (iv) determine air-sea gas exchange, and (v) perform mass balance calculations. After multiple test runs in the Baltic Sea, which resulted in continuous improvement of the design and handling, the KOSMOS platform successfully completed its first full-scale experiment in the high Arctic off Svalbard (78° 56.2′ N, 11° 53.6′ E) in June/July 2010. The study, which was conducted in the framework of the European Project on Ocean Acidification (EPOCA), focused on the effects of ocean acidification on a natural plankton community and its impacts on biogeochemical cycling and air/sea exchange of climate relevant gases. This manuscript describes the mesocosm hardware, its deployment and handling, CO2 manipulation, sampling and cleaning, including some further modifications conducted based on the experiences gained during this study. [ABSTRACT FROM AUTHOR]

The volume of water enclosed inside flexible-wall mesocosm bags is hard to estimate using geometrical calculations and can be strongly variable among bags of the same dimensions. Here we present a method for precise water volume determination in mesocosms using salinity as a tracer. Knowledge of the precise volume of water enclosed allows establishment of exactly planed treatment concentrations and calculation of elemental budgets. [ABSTRACT FROM AUTHOR]

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 2010, 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 at 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 diatoms, 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. [ABSTRACT FROM AUTHOR]

The Arctic Ocean ecosystem is particular vulnerable for ocean acidification (OA) related alterations due to the relatively high CO2 solubility and low carbonate saturation states of its cold surface waters. Thus far, however, there is only little known about the consequences of OA on the base of the food web. In a mesocosm CO2-enrichment experiment (overall CO2 levels ranged from ~180 to 1100 µatm) in the Kongsfjord off Svalbard, we studied the consequences of OA on a natural pelagic microbial community. The most prominent finding of our study is the profound effect of OA on the composition and growth of the Arctic phytoplankton community, i.e. the picoeukaryotic photoautotrophs and to a lesser extent the nanophytoplankton prospered. A shift towards the smallest phytoplankton as a result of OA will have direct consequences for the structure and functioning of the pelagic food web and thus for the biogeochemical cycles. Furthermore, the dominant pico- and nanophytoplankton groups were found prone to viral lysis, thereby shunting the carbon accumulation in living organisms into the dissolved pools of organic carbon and subsequently affecting the efficiency of the biological pump in these Arctic waters. [ABSTRACT FROM AUTHOR]

Mesocosms as large experimental vessels principally provide the opportunity of performing elemental budget calculations e.g. to derive net biological turnover rates. However, the system is in most cases not closed at the water surface and gases can exchange with the atmosphere. Previous attempts to budget carbon pools in mesocosms relied on educated guesses concerning the exchange of CO2 with the atmosphere. Nevertheless, net primary production rates derived from these budget calculations were, despite large uncertainties in air/sea gas exchange, often more reasonable than cumulative extrapolations of bioassays. While bioassays have limitations representing the full spectrum of trophic levels and abiotic conditions inside the mesocosms, calculating dissolved inorganic carbon uptake inside the mesocosms has the potential to deliver net community production rates representative of the enclosed system. Here, we present a simple method for precise determination of air/sea gas exchange velocities in mesocosms using N2O as a deliberate tracer. Beside the application for carbon budgeting, exchange velocities can be used to calculate exchange rates of any gas of known concentration, e.g. to calculate aquatic production rates of climate relevant trace gases. Using an arctic (Kiel Off Shore Mesocosms for future Ocean Simulation) mesocosm experiment as an exemplary dataset, it is shown that application of the presented method largely improves accuracy of carbon budget estimates. Methodology of manipulation, measurement, data processing and conversion to CO2 fluxes are explained. A theoretical discussion of prerequisites for precise gas exchange measurements provides a guideline for the applicability of the method under various experimental conditions. [ABSTRACT FROM AUTHOR]

Increasing concentrations of atmospheric carbon dioxide are projected to lead to an increase in sea surface temperatures, potentially impacting marine ecosystems and biogeochemical cycling. Here we conducted an indoor mesocosm experiment with a natural plankton community taken from the Baltic Sea in summer. We induced a plankton bloom via nutrient addition and followed the dynamics of the different carbon and nitrogen pools for a period of one month at temperatures ranging from 9.5 °C to 17.5 °C, representing a range of ±4 °C relative to ambient temperature. The uptake of dissolved inorganic carbon (DIC) and the net build-up of both particulate (POC) and dissolved organic carbon (DOC) were all enhanced at higher temperatures and almost doubled over a temperature gradient of 8 °C. Furthermore, elemental ratios of carbon and nitrogen (C : N) in both particulate and dissolved organic matter increased in response to higher temperatures, both reaching very high C : N ratios of >30 at +4 °C. Altogether, these observations suggest a pronounced increase in excess carbon fixation in response to elevated temperatures. Most of these findings are contrary to results from similar experiments conducted with plankton populations sampled in spring, revealing large uncertainties in our knowledge of temperature sensitivities of key processes in marine carbon cycling. Since a major difference to previous mesocosm experiments was the dominant phytoplankton species, we hypothesize that species composition might play an important role in the response of biogeochemical cycling to increasing temperatures. [ABSTRACT FROM AUTHOR]

Recent studies on the impacts of ocean acidification on pelagic communities have identified changes in carbon to nutrient dynamics with related shifts in elemental stoichiometry. In principle, mesocosm experiments provide the opportunity of determining the temporal dynamics of all relevant carbon and nutrient pools and, thus, calculating elemental budgets. In practice, attempts to budget mesocosm enclosures are often hampered by uncertainties in some of the measured pools and fluxes, in particular due to uncertainties in constraining air/sea gas exchange, particle sinking, and wall growth. In an Arctic mesocosm study on ocean acidification using KOSMOS (Kiel Off-Shore Mesocosms for future Ocean Simulation) all relevant element pools and fluxes of carbon, nitrogen and phosphorus were measured, using an improved experimental design intended to narrow down some of the mentioned uncertainties. Water column concentrations of particulate and dissolved organic and inorganic constituents were determined daily. New approaches for quantitative estimates of material sinking to the bottom of the mesocosms and gas exchange in 48 h temporal resolution, as well as estimates of wall growth were developed to close the gaps in element budgets. Future elevated pCO2 was found to enhance net autotrophic community carbon uptake in 2 of the 3 experimental phases but did not significantly affect particle elemental composition. Enhanced carbon consumption appears to result in accumulation of dissolved organic compounds under nutrient recycling summer conditions. This carbon over-consumption effect becomes evident from budget calculations, but was too small to be resolved by direct measurements of dissolved organics. The out-competing of large diatoms by comparatively small algae in nutrient uptake caused reduced production rates under future ocean CO2 conditions in the end of the experiment. This CO2 induced shift away from diatoms towards smaller phytoplankton and enhanced cycling of dissolved organics was pushing the system towards a retention type food chain with overall negative effects on export potential. [ABSTRACT FROM AUTHOR]

Net community production (NCP) and ratios of carbon to nutrient consumption were studied during a large-scale mesocosm experiment on ocean acidification in Kongsfjorden, West Spitsbergen, during June-July 2010. Nutrient-deplete fjord water with natural phyto- and bacteriaplankton assemblages, enclosed in nine mesocosms of ∼50m³ volume, was exposed to pCO2 levels ranging from 185 to 1420 µatm on initial state. Mean values of pCO2 levels during experiment ranged from 175 to 1085 µatm in different mesocosms. Phytoplankton growth was stimulated by nutrient addition. In this study NCP is estimated as a cumulative change in dissolved inorganic carbon concentrations. Stoichiometric couping between inorganic carbon and nutrient is shown as a ratio of a cumulative NCP to a cumulative change in inorganic nutrients. Three peaks of chlorophyll a concentration occurred during the experiment. Accordingly the experiment was divided in three phases. Overall cumulative NCP was similar in all mesocosms by the final day of experiment. However, NCP varied among phases, showing variable response to CO2 perturbation. Carbon to nitrogen (C:N) and carbon to phosphorus (C: P) uptake ratios were estimated only for the period after nutrient addition (post-nutrient period). For the total post-nutrient period ratios were close to Redfield proportions, however varied from it in different phases. The response of C:N and C: P uptake ratios to CO2 perturbation was different for three phases of the experiment, reflecting variable NCP and dependence on changing microbial community. Through the variable NCP, C:N and C: P uptake ratios for 31 days of the experiment we show a flexibility of biogeochemical response establishing a strong microbial loop in Kongsfjorden under different CO2 scenarios. [ABSTRACT FROM AUTHOR]

The effect of ocean acidification on the balance between gross community production (GCP) and community respiration (CR) (i.e. net community production, NCP) of plankton communities was investigated in summer 2010 in Kongsfjorden, West of Svalbard. Surface water, which was characterized by low concentrations of dissolved inorganic nutrients and chlorophyll, was enclosed in 9 mesocosms and subjected to 8 pCO2 levels (2 replicated controls and 7 enhanced pCO2 treatments) for one month. Nutrients were added to all mesocosms on day 13 of the experiment, and thereafter increase of chlorophyll (index of phytoplankton biomass) was provoked in all mesocosms. No clear trend in response to increasing pCO2 was found in the daily values of NCP, CR, and GCP. For further analysis, these parameters were cumulated for the following three periods: phase 1: end of CO2 manipulation until nutrient addition (t4 to t13); phase 2: nutrient addition until the second chlorophyll minimum (t14 to t21); phase 3: the second chlorophyll minimum until the end of this study (t22 to t28). Significant response was detected as a decrease of NCP with increasing pCO2 during phase 3. CR was relatively stable throughout the experiment in all mesocosms. As a result, the cumulative GCP significantly decreased with increasing pCO2 during phase 3. After the nutrient addition, the ratios of cumulative NCP to cumulative consumption of NO3 and PO4 showed significant decrease during phase 3 with increasing pCO2. The results suggest that elevated pCO2 influenced cumulative NCP and stoichiometric C and nutrient coupling of the plankton community in a high latitude fjord only for a limited period. However provided that there were some differences or weak correlations between NCP data based on different methods in the same experiment, this conclusion should be taken with caution. [ABSTRACT FROM AUTHOR]

The increase in atmospheric carbon dioxide (CO2) results in acidification of the oceans, expected to lead to the fastest drop in ocean pH in the last 300 million years, if anthropogenic emissions are continued at present rate. Due to higher solubility of gases in cold waters and increased exposure to the atmosphere by decreasing ice cover, the Arctic Ocean will be among the areas most strongly affected by ocean acidification. Yet, the response of the plankton community of high latitudes to ocean acidification has not been studied so far. This work is part of the Arctic campaign of the European Project on Ocean Acidification (EPOCA) in 2010, employing 9 in situ mesocosms of about 45 000 l each to simulate ocean acidification in Kongsfjorden, Svalbard (78° 56.20 N 11 ° 53.60 E). In the present study, we investigated effects of elevated CO2 on the composition and richness of particle attached (PA; >3 µm) and free living (FL; < 3 µm > 0.2 µm) bacterial communities by Automated Ribosomal Intergenic Spacer Analysis (ARISA) in 6 of the mesocosms and the surrounding fjord, ranging from 185 to 1050 initial µatm pCO2. ARISA was able to resolve about 20-30 bacterial band-classes per sample and allowed for a detailed investigation of the explicit richness. 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 (phase 3 of the experiment), number of ARISA-band classes in the PA-community were reduced at low and medium CO2 (∼180-600 µatm) by about 25%, while it was more or less stable at high CO2 (∼650-800 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-treatments, suggesting a positive effect of community richness on this function and on carbon cycling by bacteria. [ABSTRACT FROM AUTHOR]

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 effcient 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. [ABSTRACT FROM AUTHOR]

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, 20 N, 11° 53, 60 E). Enclosed in 9 mesocosms (volume: 43.9-47.6m³ ), 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, 14 C-bottle incubations (24 h) of mesocosm samples were performed at 1m depth receiving about 60% of incoming radiation. PP for all mesocosms averaged 8.06±3.64 µmolCl-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 suffcient 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. [ABSTRACT FROM AUTHOR]

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 (HCO Due to image rights restrictions, multiple line equation(s) cannot be graphically displayed.), carbonate ion (CO Due to image rights restrictions, multiple line equation(s) cannot be graphically displayed.), 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 [HCO Due to image rights restrictions, multiple line equation(s) cannot be graphically displayed.] and [H+] although influence of [CO Due to image rights restrictions, multiple line equation(s) cannot be graphically displayed.] 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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

The effect of ocean acidification on the fatty acid composition of a natural plankton community in the Arctic was studied in a large-scale mesocosm experiment, carried out in Kongsfjorden (Svalbard, Norway) at 79° N. Nine mesocosms of ∼50 cbm each were exposed to different pCO2 levels (from natural background conditions to ∼1420 µatm), yielding pH values (on the total scale) from ∼8.3 to 7.5. Inorganic nutrients were added on day 13. The phytoplankton development during this 30 days experiment passed three distinct phases: (1) prior to the addition of inorganic nutrients, (2) first bloom after nutrient addition, and (3) second bloom after nutrient addition. The fatty acid composition of the natural plankton community was analysed and showed, in general, high percentages of polyunsaturated fatty acids (PUFAs): 44-60% of total fatty acids. Positive correlations with pCO2 were found for most PUFAs during phases 2 and/or 3, with the exception of 20 : 5n3 (eicosapentaenoic acid, EPA), an important diatom marker. There are strong indications for these correlations being mediated indirectly through taxonomic changes and the natural development of the communities in the mesocosms exposed to different pCO2 levels. While diatoms increased during phase 3 mainly in the low and intermediate pCO2 treatments, dinoflagellates were favoured by high CO2 concentrations during the same time period. This is reflected in the development of group-specific fatty acid trophic markers. No indications were found for a generally detrimental effect of ocean acidification on the planktonic food quality in terms of essential fatty acids. The significant positive correlations between most PUFAs and pCO2 reflected treatment-dependent differences in the community composition between the mesocosms rather than a direct positive effect of pCO2 on specific fatty acids. [ABSTRACT FROM AUTHOR]

Most scleractinian corals and many other cnidarians host intracellular photosynthetic dinoflagellate symbionts ("zooxanthellae"). The zooxanthellae contribute to host metabolism and skeletogenesis to such an extent that this symbiosis is well recognised for its contribution in creating the coral reef ecosystem. The stable functioning of cnidarian symbioses is however dependent upon the host's ability to maintain demographic control of its algal partner. In this review, I explain how the modern envelope of seawater conditions found within many coral reef ecosystems (characterised by elevated temperatures, rising pCO2, and enriched nutrient levels) are antagonistic toward the dominant host processes that restrict excessive symbiont proliferation. Moreover, I outline a new hypothesis and initial evidence base, which support the suggestion that the additional "excess" zooxanthellae fraction permitted by seawater pCO2 levels beyond 260 ppmv significantly increases the propensity for symbiosis breakdown ("bleaching") in response to temperature and irradiance extremes. The relevance of this biological threshold is discussed in terms of historical reef extinction events, glacial-interglacial climate cycles and the modern decline of coral reef ecosystems. [ABSTRACT FROM AUTHOR]