Response of bacterioplankton community structure to an artificial gradient of pCO2 in the Arctic Ocean (update)

In order to test the influences of ocean acidification on the ocean pelagic ecosystem, so far the largest CO₂ manipulation mesocosm study (European Project on Ocean Acidification, EPOCA) was performed in Kings Bay (Kongsfjorden), Spitsbergen. During a 30 day incubation, bacterial diversity was investigated using DNA fingerprinting and clone library analysis of bacterioplankton samples. Terminal restriction fragment length polymorphism (T-RFLP) analysis of the PCR amplicons of the 16S rRNA genes revealed that general bacterial diversity, taxonomic richness and community structure were influenced by the variation of productivity during the time of incubation, but not the degree of ocean acidification. A BIOENV analysis suggested a complex control of bacterial community structure by various biological and chemical environmental parameters. The maximum apparent diversity of bacterioplankton (i.e., the number of T-RFs) in high and low pCO₂ treatments differed significantly. A negative relationship between the relative abundance of Bacteroidetes and pCO₂ levels was observed for samples at the end of the experiment by the combination of T-RFLP and clone library analysis. Our study suggests that ocean acidification affects the development of bacterial assemblages and potentially impacts the ecological function of the bacterioplankton in the marine ecosystem.

Results of laboratory and field experiments of the direct effect of increasing CO2 on net primary production of macroalgal species in brackish-water ecosystems

Studies on the effects of increasing acidification on marine communities have been previously mostly carried out in truly marine areas whereas brackish-water ecosystems such as the Baltic Sea have been less studied. The current study analyses how acidification induced by elevated atmospheric carbon dioxide affects the photosynthetic net production of different macroalgal species in the brackish Baltic Sea. Research methods include sets of laboratory and field experiments carried out in shallow coastal brackish waters. The aim of the laboratory experiments was to develop the necessary techniques and experience for the mesocosm experiments. Laboratory experiments were carried out using specimens of the red alga Furcellaria lumbricalis collected from Kakumäe Bay. The mesocosm experiments were conducted in Kõiguste Bay during the field season of 2011. Separate mesocosms were operated in each set with different CO₂ concentrations and a control treatment in natural conditions. Field experiments were carried out with three species representing three different morphological and ecological groups: Ulva intestinalis, a fast-growing green alga; Fucus vesiculosus, a perennial brown alga with a slow metabolism; and Furcellaria lumbricalis, a perennial red alga. Photosynthetic activity was used as the response variable. In the laboratory decreasing pH increased the net primary production of F. lumbricalis with the lowest net primary production values measured at pH 8.0 and the highest at pH 6.5. Results of the field experiments indicated that increased CO₂ levels in seawater favoured photosynthetic activity of the macroalgae U. intestinalis and F. lumbricalis, but F. vesiculosus showed no response to elevated CO₂. Elevated CO₂ levels are suggested to favour the production of fast-growing filamentous species, which thus may indirectly enhance the effect of eutrophication in the shallow coastal brackish waters.

Effect of CO2 enrichment on bacterial metabolism in an Arctic fjord (update)

The anthropogenic increase of carbon dioxide (CO₂) alters the seawater carbonate chemistry, with a decline of pH and an increase in the partial pressure of CO₂ (pCO₂). Although bacteria play a major role in carbon cycling, little is known about the impact of rising pCO₂ on bacterial carbon metabolism, especially for natural bacterial communities. In this study, we investigated the effect of rising pCO₂ on bacterial production (BP), bacterial respiration (BR) and bacterial carbon metabolism during a mesocosm experiment performed in Kongsfjorden (Svalbard) in 2010. Nine mesocosms with pCO₂ levels ranging from ca. 180 to 1400 μatm were deployed in the fjord and monitored for 30 days. Generally BP gradually decreased in all mesocosms in an initial phase, showed a large (3.6-fold average) but temporary increase on day 10, and increased slightly after inorganic nutrient addition. Over the wide range of pCO₂ investigated, the patterns in BP and growth rate of bulk and free-living communities were generally similar over time. However, BP of the bulk community significantly decreased with increasing pCO₂ after nutrient addition (day 14). In addition, increasing pCO₂ enhanced the leucine to thymidine (Leu : TdR) ratio at the end of experiment, suggesting that pCO₂ may alter the growth balance of bacteria. Stepwise multiple regression analysis suggests that multiple factors, including pCO₂, explained the changes of BP, growth rate and Leu : TdR ratio at the end of the experiment. In contrast to BP, no clear trend and effect of changes of pCO₂ was observed for BR, bacterial carbon demand and bacterial growth efficiency. Overall, the results suggest that changes in pCO₂ potentially influence bacterial production, growth rate and growth balance rather than the conversion of dissolved organic matter into CO₂.

Implications of elevated CO2 on pelagic carbon fluxes in an Arctic mesocosm study – an elemental mass balance approach (update)

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.

Consumers mediate the effects of experimental ocean acidification and warming on primary producers

It is well known that ocean acidification can have profound impacts on marine organisms. However, we know little about the direct and indirect effects of ocean acidification and also how these effects interact with other features of environmental change such as warming and declining consumer pressure. In this study, we tested whether the presence of consumers (invertebrate mesograzers) influenced the interactive effects of ocean acidification and warming on benthic microalgae in a seagrass community mesocosm experiment. Net effects of acidification and warming on benthic microalgal biomass and production, as assessed by analysis of variance, were relatively weak regardless of grazer presence. However, partitioning these net effects into direct and indirect effects using structural equation modeling revealed several strong relationships. In the absence of grazers, benthic microalgae were negatively and indirectly affected by sediment-associated microalgal grazers and macroalgal shading, but directly and positively affected by acidification and warming. Combining indirect and direct effects yielded no or weak net effects. In the presence of grazers, almost all direct and indirect climate effects were nonsignificant. Our analyses highlight that (i) indirect effects of climate change may be at least as strong as direct effects, (ii) grazers are crucial in mediating these effects, and (iii) effects of ocean acidification may be apparent only through indirect effects and in combination with other variables (e.g., warming). These findings highlight the importance of experimental designs and statistical analyses that allow us to separate and quantify the direct and indirect effects of multiple climate variables on natural communities.

Response of halocarbons to ocean acidification in the Arctic (update)

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.

Contrasting responses of DMS and DMSP to ocean acidification in Arctic waters (update)

Increasing atmospheric CO₂ is decreasing ocean pH most rapidly in colder regions such as the Arctic. As a component of the EPOCA (European Project on Ocean Acidification) pelagic mesocosm experiment off Spitzbergen in 2010, we examined the consequences of decreased pH and increased pCO₂ on the concentrations of dimethylsulphide (DMS). DMS is an important reactant and contributor to aerosol formation and growth in the Arctic troposphere. In the nine mesocosms with initial pH_T 8.3 to 7.5, equivalent to pCO₂ of 180 to 1420 μatm, highly significant but inverse responses to acidity (hydrogen ion concentration [H⁺]) occurred following nutrient addition. Compared to ambient [H⁺], average concentrations of DMS during the mid-phase of the 30 d experiment, when the influence of altered acidity was unambiguous, were reduced by approximately 60% at the highest [H⁺] and by 35% at [H⁺] equivalent to 750 μatm pCO₂, as projected for 2100. In contrast, concentrations of dimethylsulphoniopropionate (DMSP), the precursor of DMS, were elevated by approximately 50% at the highest [H⁺] and by 30% at [H⁺] corresponding to 750 μatm pCO₂. Measurements of the specific rate of synthesis of DMSP by phytoplankton indicate increased production at high [H⁺], in parallel to rates of inorganic carbon fixation. The elevated DMSP production at high [H⁺] was largely a consequence of increased dinoflagellate biomass and in particular, the increased abundance of the species Heterocapsa rotundata. We discuss both phytoplankton and bacterial processes that may explain the reduced ratios of DMS:DMSPt (total dimethylsulphoniopropionate) at higher [H⁺]. The experimental design of eight treatment levels provides comparatively robust empirical relationships of DMS and DMSP concentration, DMSP production and dinoflagellate biomass versus [H⁺] in Arctic waters.

Technical Note: the determination of enclosed water volume in large flexible-wall mesocosms “KOSMOS” (update)

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.

Technical Note: A mobile sea-going mesocosm system – new opportunities for ocean change research (update)

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.

Red coral extinction risk enhanced by ocean acidification

The red coral Corallium rubrum is a habitat-forming species with a prominent and structural role in mesophotic habitats, which sustains biodiversity hotspots. This precious coral is threatened by both over-exploitation and temperature driven mass mortality events. We report here that biocalcification, growth rates and polyps’ (feeding) activity of Corallium rubrum are significantly reduced at pCO₂ scenarios predicted for the end of this century (0.2 pH decrease). Since C. rubrum is a long-living species (>200 years), our results suggest that ocean acidification predicted for 2100 will significantly increases the risk of extinction of present populations. Given the functional role of these corals in the mesophotic zone, we predict that ocean acidification might have cascading effects on the functioning of these habitats worldwide.