Causes and consequences of acidification in the Baltic Sea: implications for monitoring and management.

Increasing atmospheric CO2 drives ocean acidification globally. In coastal seas, acidification trends can however be either counteracted or enhanced by other processes. Ecosystem effects of acidification are so far small in the Baltic Sea, but changes should be anticipated unless CO2 emissions are curbed. Possible future acidification trends in the Baltic Sea, conditional on CO2 emissions, climate change, and changes in productivity, can be assessed by means of model simulations. There are uncertainties regarding potential consequences for marine organisms, partly because of difficulties to assign critical thresholds, but also because of knowledge gaps regarding species' capacity to adapt. Increased temporal and spatial monitoring of inorganic carbon system parameters would allow a better understanding of current acidification trends and also improve the capacity to predict possible future changes. An additional benefit is that such measurements also provide quantitative estimates of productivity. The technology required for precise measurements of the inorganic carbon system is readily available today. Regularly updated status evaluations of acidification, and the inorganic carbon system in general, would support management when assessing climate change effects, eutrophication or characteristics of the pelagic habitats. This would, however, have to be based on a spatially and temporally sufficient monitoring program.

Effects of different oxygen regimes on ecological performance and bioenergetics of a coastal marine bioturbator, the soft shell clam Mya arenaria.

Benthic species are exposed to oxygen fluctuations that can affect their performance and survival. Physiological effects and ecological consequences of fluctuating oxygen are not well understood in marine bioturbators such as the soft-shell clam Mya arenaria. We explored the effects of different oxygen regimes (21 days of exposure to constant hypoxia (~4.1 kPa PO2), cyclic hypoxia (~2.1-~10.4 kPa PO2) or normoxia (~21 kPa PO2)) on energy metabolism, oxidative stress and ecological behaviors (bioirrigation and bioturbation) of M. arenaria. Constant hypoxia and post-hypoxic recovery in cyclic hypoxia led to oxidative injury of proteins and lipids, respectively. Clams acclimated to constant hypoxia maintained aerobic capacity similar to the normoxic clams. In contrast, clams acclimated to cyclic hypoxia suppressed aerobic metabolism and activated anaerobiosis during hypoxia, and strongly upregulated aerobic metabolism during recovery. Constant hypoxia led to decreased lipid content, whereas in cyclic hypoxia proteins and glycogen accumulated during recovery and were broken down during the hypoxic phase. Digging of clams was impaired by constant and cyclic hypoxia, and bioirrigation was also suppressed under constant hypoxia. Overall, cyclic hypoxia appears less stressful for M. arenaria due to the metabolic flexibility that ensures recovery during reoxygenation and mitigates the negative effects of hypoxia, whereas constant hypoxia leads to depletion of energy reserves and impairs ecological functions of M. arenaria potentially leading to negative ecological consequences in benthic ecosystems.

Effects of variable oxygen regimes on mitochondrial bioenergetics and reactive oxygen species production in a marine bivalve, Mya arenaria.

Estuarine and coastal benthic organisms often experience fluctuations in oxygen levels that can negatively impact their mitochondrial function and aerobic metabolism. To study these impacts, we exposed a common sediment-dwelling bivalve, the soft-shell clam Mya arenaria, for 21 days to chronic hypoxia (PO2  âˆ¼4.1 kPa), cyclic hypoxia (PO2  âˆ¼12.7-1.9 kPa, mean 5.7 kPa) or normoxia (PO2  âˆ¼21.1 kPa). pH was manipulated to mimic the covariation in CO2/pH and oxygen levels in coastal hypoxic zones. Mitochondrial respiration, including proton leak, the capacity for oxidative phosphorylation (OXPHOS), the maximum activity of the electron transport system (ETS), reactive oxygen species (ROS) production, and activity and oxygen affinity of cytochrome c oxidase (CCO) were assessed. Acclimation to constant hypoxia did not affect the studied mitochondrial traits except for a modest decrease in the OXPHOS coupling efficiency. Cyclic hypoxia had no effect on OXPHOS or ETS capacity, but increased proton leak and lowered mitochondrial OXPHOS coupling efficiency. Furthermore, mitochondria of clams acclimated to cyclic hypoxia had higher rates of ROS generation compared with the clams acclimated to normoxia or chronic hypoxia. CCO activity was upregulated under cyclic hypoxia, but oxygen affinity of CCO did not change. These findings indicate that long-term cyclic hypoxia has a stronger impact on the mitochondria of M. arenaria than chronic hypoxia and might lead to impaired ATP synthesis, higher costs of mitochondrial maintenance and oxidative stress. These changes might negatively affect populations of M. arenaria in the coastal Baltic Sea under increasing hypoxia pressure.

Methane-carbon flow into the benthic food web at cold seeps--a case study from the Costa Rica subduction zone.

Cold seep ecosystems can support enormous biomasses of free-living and symbiotic chemoautotrophic organisms that get their energy from the oxidation of methane or sulfide. Most of this biomass derives from animals that are associated with bacterial symbionts, which are able to metabolize the chemical resources provided by the seeping fluids. Often these systems also harbor dense accumulations of non-symbiotic megafauna, which can be relevant in exporting chemosynthetically fixed carbon from seeps to the surrounding deep sea. Here we investigated the carbon sources of lithodid crabs (Paralomis sp.) feeding on thiotrophic bacterial mats at an active mud volcano at the Costa Rica subduction zone. To evaluate the dietary carbon source of the crabs, we compared the microbial community in stomach contents with surface sediments covered by microbial mats. The stomach content analyses revealed a dominance of epsilonproteobacterial 16S rRNA gene sequences related to the free-living and epibiotic sulfur oxidiser Sulfurovum sp. We also found Sulfurovum sp. as well as members of the genera Arcobacter and Sulfurimonas in mat-covered surface sediments where Epsilonproteobacteria were highly abundant constituting 10% of total cells. Furthermore, we detected substantial amounts of bacterial fatty acids such as i-C15∶0 and C17∶1ω6c with stable carbon isotope compositions as low as -53‰ in the stomach and muscle tissue. These results indicate that the white microbial mats at Mound 12 are comprised of Epsilonproteobacteria and that microbial mat-derived carbon provides an important contribution to the crab's nutrition. In addition, our lipid analyses also suggest that the crabs feed on other (13)C-depleted organic matter sources, possibly symbiotic megafauna as well as on photosynthetic carbon sources such as sedimentary detritus.

Metabolically active microbial communities in marine sediment under high-CO(2) and low-pH extremes.

Sediment-hosting hydrothermal systems in the Okinawa Trough maintain a large amount of liquid, supercritical and hydrate phases of CO(2) in the seabed. The emission of CO(2) may critically impact the geochemical, geophysical and ecological characteristics of the deep-sea sedimentary environment. So far it remains unclear whether microbial communities that have been detected in such high-CO(2) and low-pH habitats are metabolically active, and if so, what the biogeochemical and ecological consequences for the environment are. In this study, RNA-based molecular approaches and radioactive tracer-based respiration rate assays were combined to study the density, diversity and metabolic activity of microbial communities in CO(2)-seep sediment at the Yonaguni Knoll IV hydrothermal field of the southern Okinawa Trough. In general, the number of microbes decreased sharply with increasing sediment depth and CO(2) concentration. Phylogenetic analyses of community structure using reverse-transcribed 16S ribosomal RNA showed that the active microbial community became less diverse with increasing sediment depth and CO(2) concentration, indicating that microbial activity and community structure are sensitive to CO(2) venting. Analyses of RNA-based pyrosequences and catalyzed reporter deposition-fluorescence in situ hybridization data revealed that members of the SEEP-SRB2 group within the Deltaproteobacteria and anaerobic methanotrophic archaea (ANME-2a and -2c) were confined to the top seafloor, and active archaea were not detected in deeper sediments (13-30 cm in depth) characterized by high CO(2). Measurement of the potential sulfate reduction rate at pH conditions of 3-9 with and without methane in the headspace indicated that acidophilic sulfate reduction possibly occurs in the presence of methane, even at very low pH of 3. These results suggest that some members of the anaerobic methanotrophs and sulfate reducers can adapt to the CO(2)-seep sedimentary environment; however, CO(2) and pH in the deep-sea sediment were found to severely impact the activity and structure of the microbial community.

Comment on "A persistent oxygen anomaly reveals the fate of spilled methane in the deep Gulf of Mexico".

Kessler et al. (Reports, 21 January 2011, p. 312) reported that methane released from the 2010 Deepwater Horizon blowout, approximately 40% of the total hydrocarbon discharge, was consumed quantitatively by methanotrophic bacteria in Gulf of Mexico deep waters over a 4-month period. We find the evidence explicitly linking observed oxygen anomalies to methane consumption ambiguous and extension of these observations to hydrate-derived methane climate forcing premature.

Experimental investigation of the rising behavior of CO2 droplets in seawater under hydrate-forming conditions.

In a laboratory-based test series, seven experiments along a simulated Pacific hydrotherm at 152 degrees W, 40 degrees N were carried out to measure the rise velocities of liquefied CO2 droplets under (clathrate) hydrate forming conditions. The impact of a hydrate skin on the rising behavior was investigated by comparing the results with those from outside the field of hydrate stability at matching buoyancy. A thermostatted high-pressure tank was used to establish conditions along the natural oceanic hydrotherm. Under P-/T-conditions allowing hydrate formation, the majority of the droplets quickly developed a skin of CO2 hydrate upon contact with seawater. Rise rates of these droplets support the parametrization by Chen et al. (Tellus 2003, 55B, 723-730), which is based on empirical equations developed to match momentum of hydrate covered, deformed droplets. Our data do not support other parametrizations recently suggested in the literature. In the experiments from 5.7 MPa, 4.8 oC to 11.9 MPa, 2.8 degrees C positive and negative deviations from predicted rise rates occurred, which we propose were caused by lacking hydrate formation and reflect intact droplet surface mobility and droplet shape oscillations, respectively. This interpretation is supported by rise rates measured at P-/T-conditions outside the hydrate stability field atthe same liquid CO2-seawater density difference (delta rho) matching the rise rates of the deviating data within the stability field. The results also show that droplets without a hydrate skin ascend up to 50% faster than equally buoyant droplets with a hydrate skin. This feature has a significant impact on the vertical pattern of dissolution of liquid CO2 released into the ocean. The experiments and data presented considerably reduce the uncertainty of the parametrization of CO2 droplet rise velocity, which in the past emerged partly from their scarcity and contradictions in constraints of earlier experiments.

Experimental determination of the fate of rising CO2 droplets in seawater.

Direct oceanic disposal of fossil fuel CO2 is being considered as a possible means to moderate the growth rate of CO2 in the atmosphere. We have measured the rise rate and dissolution rate of freely released CO2 droplets in the open ocean to provide fundamental data for carbon sequestration options. A small amount of liquid CO2 was released at 800 m, at 4.4 degrees C, and the rising droplet stream was imaged with a HDTV camera carried on a remotely operated vehicle. The initial rise rate for 0.9-cm diameter droplets was 10 cm/s at 800 m, and the dissolution rate was 3.0 micromol cm(-2) s(-1). While visual contact was maintained for 1 h and over a 400 m ascent, 90% of the mass loss occurred within 30 min over a 200 m ascent above the release point. Images of droplets crossing the liquid-gas-phase boundary showed formation of a gas head, pinching off of a liquid tail, and rapid gas bubble separation and dissolution.