Description: Ongoing acidification of the ocean through uptake of anthropogenic CO2 is known to affect marine biota and ecosystems with largely unknown consequences for marine food webs. Changes in food web structure have the potential to alter trophic transfer, partitioning, and biogeochemical cycling of elements in the ocean. Here we investigated the impact of realistic end-of-the-century CO2 concentrations on the development and partitioning of the carbon, nitrogen, phosphorus, and silica pools in a coastal pelagic ecosystem (Gullmar Fjord, Sweden). We covered the entire winter-to-summer plankton succession (100 days) in two sets of five pelagic mesocosms, with one set being CO2 enriched (~760 μatm pCO2) and the other one left at ambient CO2 concentrations. Elemental mass balances were calculated and we highlight important challenges and uncertainties we have faced in the closed mesocosm system. Our key observations under high CO2 were: (1) A significantly amplified transfer of carbon, nitrogen, and phosphorus from primary producers to higher trophic levels, during times of regenerated primary production. (2) A prolonged retention of all three elements in the pelagic food web that significantly reduced nitrogen and phosphorus sedimentation by about 11 and 9%, respectively. (3) A positive trend in carbon fixation (relative to nitrogen) that appeared in the particulate matter pool as well as the downward particle flux. This excess carbon counteracted a potential reduction in carbon sedimentation that could have been expected from patterns of nitrogen and phosphorus fluxes. Our findings highlight the potential for ocean acidification to alter partitioning and cycling of carbon and nutrients in the surface ocean but also show that impacts are temporarily variable and likely depending upon the structure of the plankton food web.

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: The potential effect of ocean acidification (OA) on seawater halocarbons in the Arctic was investigated during a~mesocosm experiment in Spitsbergen in June–July 2010. Over a period of 5 weeks, natural phytoplankton communities in nine ~50 m3 mesocosms were studied under a range of pCO2 treatments from ~185 μatm to ~1420 μatm. In general, the response of halocarbons to pCO2 was subtle, or undetectable. A large number of significant correlations with a range of biological parameters (chlorophyll a, microbial plankton community, phytoplankton pigments) were identified, indicating a biological control on the concentrations of halocarbons within the mesocosms. The temporal dynamics of iodomethane (CH3I) alluded to active turnover of this halocarbon in the mesocosms and strong significant correlations with biological parameters suggested a biological source. However, despite a pCO2 effect on various components of the plankton community, and a strong association between CH3I and biological parameters, no effect of pCO2 was seen in CH3I. Diiodomethane (CH2I2) displayed a number of strong relationships with biological parameters. Furthermore, the concentrations, the rate of net production and the sea-to-air flux of CH2I2 showed a significant positive response to pCO2. There was no clear effect of pCO2 on bromocarbon concentrations or dynamics. However, periods of significant net loss of bromoform (CHBr3) were found to be concentration-dependent, and closely correlated with total bacteria, suggesting a degree of biological consumption of this halocarbon in Arctic waters. Although the effects of OA on halocarbon concentrations were marginal, this study provides invaluable information on the production and cycling of halocarbons in a region of the world's oceans likely to experience rapid environmental change in the coming decades.

Description: 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.

Description: 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.

Description: 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 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 applying 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 the mentioned uncertainties. Water-column concentrations of particulate and dissolved organic and inorganic matter 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. However, losses elements from the budgets into a sum of insufficiently determined pools were detected, and are principally unavoidable in mesocosm investigation. The comparison of variability patterns of all single measured datasets revealed analytic precision to be the main issue in determination of budgets. Uncertainties in dissolved organic carbon (DOC), nitrogen (DON) and particulate organic phosphorus (POP) were much higher than the summed error in determination of the same elements in all other pools. With estimates provided for all other major elemental pools, mass balance calculations could be used to infer the temporal development of DOC, DON and POP pools. Future elevated pCO2 was found to enhance net autotrophic community carbon uptake in two of the three experimental phases but did not significantly affect particle elemental composition. Enhanced carbon consumption appears to result in accumulation of dissolved organic carbon under nutrient-recycling summer conditions. This carbon over-consumption effect becomes evident from mass balance calculations, but was too small to be resolved by direct measurements of dissolved organic matter. Faster nutrient uptake by comparatively small algae at high CO2 after nutrient addition resulted in reduced production rates under future ocean CO2 conditions at the end of the experiment. This CO2 mediated shift towards smaller phytoplankton and enhanced cycling of dissolved matter restricted the development of larger phytoplankton, thus pushing the system towards a retention type food chain with overall negative effects on export potential.

Description: 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 2.5 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.

Description: 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.

Description: 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.

Description: The increasing CO2 concentration in the atmosphere caused by burning fossil fuels leads to increasing pCO2 and decreasing pH in the world oceans. These changes may have severe consequences for marine biota, especially in cold-water ecosystems due to higher solubility of CO2. However, studies on the response of mesozooplankton communities to elevated pCO2 are yet lacking. In order to test whether abundance and taxonomic composition change with pCO2, we have sampled nine mesocosms, which were deployed in Kongsfjorden, an Arctic fjord at Svalbard, and were adjusted to eight CO2 concentrations, initially ranging from 185 μatm to 1420 μatm. Samples were taken weekly over a six-week period with an Apstein net (55 μm mesh size) in all mesocosms and the surrounding fjord. In addition, sediment trap samples, taken every second day in the mesocosms, were analyzed to account for losses due to vertical migration and mortality. The taxonomic analysis revealed that meroplanktonic larvae (cirripeds, polychaetes, bivalves, gastropod, and decapods) dominated in the mesocosms while copepods (Calanus spp., Oithona similis, Acartia longiremis and Microsetella norvegica) were found in lower abundances. In the fjord copepods prevailed for most of our study. With time, abundance and taxonomic composition developed similarly in all mesocosms; the pCO2 had no significant effect on the overall community structure. However, single taxa responded to elevated CO2 concentrations. The ratio of cirripedia nauplii to cypris larvae, the next developmental stage, in the sediment traps averaged over the entire experiment increased with pCO2 and this suggests that increased pCO2 may have delayed their development. Also, the number of bivalves, averaged over the experimental period, decreased significantly with increasing pCO2. The nature of the CO2 effect, either direct or indirect, remains open and needs to be addressed in future.