Effects of increased pCO(2) and temperature on the North Atlantic spring bloom. I. The phytoplankton community and biogeochemical response

The North Atlantic spring bloom is one of the largest annual biological events in the ocean, and is characterized by dominance transitions from siliceous (diatoms) to calcareous (coccolithophores) algal groups. To study the effects of future global change on these phytoplankton and the biogeochemical cycles they mediate, a shipboard continuous culture experiment (Ecostat) was conducted in June 2005 during this transition period. Four treatments were examined: (1) 12 degrees C and 390 ppm CO2 (ambient control), (2) 12 degrees C and 690 ppm CO2 (high pCO(2)) (3) 16 degrees C and 390 ppm CO2 (high temperature), and (4) 16 degrees C and 690 ppm CO2 ('greenhouse'). Nutrient availability in all treatments was designed to reproduce the low silicate conditions typical of this late stage of the bloom. Both elevated pCO(2) and temperature resulted in changes in phytoplankton community structure. Increased temperature promoted whole community photosynthesis and particulate organic carbon (POC) production rates per unit chlorophyll a. Despite much higher coccolithophore abundance in the greenhouse treatment, particulate inorganic carbon production (calcification) was significantly decreased by the combination of increased pCO(2) and temperature. Our experiments suggest that future trends during the bloom could include greatly reduced export of calcium carbonate relative to POC, thus providing a potential negative feedback to atmospheric CO2 concentration. Other trends with potential climate feedback effects include decreased community biogenic silica to POC ratios at higher temperature. These shipboard experiments suggest the need to examine whether future pCO2 and temperature increases on longer decadal timescales will similarly alter the biological and biogeochemical dynamics of the North Atlantic spring bloom.

Biological impacts of enhanced alkalinity in Carcinus maenas

Further steps are needed to establish feasible alleviation strategies that are able to reduce the impacts of ocean acidification, whilst ensuring minimal biological side-effects in the process. Whilst there is a growing body of literature on the biological impacts of many other carbon dioxide reduction techniques, seemingly little is known about enhanced alkalinity. For this reason, we investigated the potential physiological impacts of using chemical sequestration as an alleviation strategy. In a controlled experiment, Carcinus maenas were acutely exposed to concentrations of Ca(OH)2 that would be required to reverse the decline in ocean surface pH and return it to pre-industrial levels. Acute exposure significantly affected all individuals' acid-base balance resulting in slight respiratory alkalosis and hyperkalemia, which was strongest in mature females. Although the trigger for both of these responses is currently unclear, this study has shown that alkalinity addition does alter acid-base balance in this comparatively robust crustacean species.

Reduced pH sea water disrupts chemo-responsive behaviour in an intertidal crustacean

Chemoreception is a key activity by which many aquatic animals perceive their environment, and therefore abiotic disruptions to this process could have serious impacts on the survival and fitness of individuals, and on species interactions. Hermit crabs are subject to cyclical reductions in the pH of the water in the intertidal rock pools that they inhabit. Such reductions may be further exacerbated by ongoing ocean acidification and/or leakage of carbon dioxide from geological storage sites and coastal upwelling events. Here we test the chemo-sensory responses of the hermit crab Pagurus bernhardus (Linnaeus) to a food odour under reduced pH conditions (pHNBS = 6.80). Acidifying the odour had no effect on its attractiveness indicating no permanent degradation of the cue; however, the pH of the sea water did affect the crabs' responses. Hermit crabs kept and tested in reduced pH sea water had lower antennular flicking rates (the ‘sniffing’ response in decapods); were less successful in locating the odour source, and showed an overall decline in locomotory activity compared to those in untreated sea water. Analysis of their haemolymph revealed a greater concentration of chloride ions ([Cl−]) in the reduced pH treatment group, suggesting iono-regulatory disruption; however, there was no correlation between [Cl−] and locomotory activity, suggesting a specific effect on chemoreception. This study shows that the chemo-responsiveness of a crustacean may be influenced by both naturally occurring pH fluctuations and future anthropogenically-induced changes in ocean pH.

Review of climate change impacts on marine fisheries in the UK and Ireland

1.Commercial fishing is an important socio-economic activity in coastal regions of the UK and Ireland. Ocean–atmospheric changes caused by greenhouse gas emissions are likely to affect future fish and shellfish production, and lead to increasing challenges in ensuring long-term sustainable fisheries management. 2.The paper reviews existing knowledge and understanding of the exposure of marine ecosystems to ocean-atmospheric changes, the consequences of these changes for marine fisheries in the UK and Ireland, and the adaptability of the UK and Irish fisheries sector. 3.Ocean warming is resulting in shifts in the distribution of exploited species and is affecting the productivity of fish stocks and underlying marine ecosystems. In addition, some studies suggest that ocean acidification may have large potential impacts on fisheries resources, in particular shell-forming invertebrates. 4.These changes may lead to loss of productivity, but also the opening of new fishing opportunities, depending on the interactions between climate impacts, fishing grounds and fleet types. They will also affect fishing regulations, the price of fish products and operating costs, which in turn will affect the economic performance of the UK and Irish fleets. 5.Key knowledge gaps exist in our understanding of the implications of climate and ocean chemistry changes for marine fisheries in the UK and Ireland, particularly on the social and economic responses of the fishing sectors to climate change. However, these gaps should not delay climate change mitigation and adaptation policy actions, particularly those measures that clearly have other ‘co-benefits’.

The carbonate system of the NW European shelf: sensitivity and model validation

The ocean plays an important role in regulating the climate, acting as a sink for carbon dioxide, perturbing the carbonate system and resulting in a slow decrease of seawater pH. Understanding the dynamics of the carbonate system in shelf sea regions is necessary to evaluate the impact of Ocean Acidification (OA) in these societally important ecosystems. Complex hydrodynamic and ecosystem coupled models provide a method of capturing the significant heterogeneity of these areas. However rigorous validation is essential to properly assess the reliability of such models. The coupled model POLCOMS–ERSEM has been implemented in the North Western European shelf with a new parameterization for alkalinity explicitly accounting for riverine inputs and the influence of biological processes. The model has been validated in a like with like comparison with North Sea data from the CANOBA dataset. The model shows good to reasonable agreement for the principal variables, physical (temperature and salinity), biogeochemical (nutrients) and carbonate system (dissolved inorganic carbon and total alkalinity), but simulation of the derived variables, pH and pCO2, are not yet fully satisfactory. This high uncertainty is attributed mostly to riverine forcing and primary production. This study suggests that the model is a useful tool to provide information on Ocean Acidification scenarios, but uncertainty on pH and pCO2 needs to be reduced, particularly when impacts of OA on ecosystem functions are included in the model systems.

Effect of elevated CO2 on the dynamics of particle-attached and free-living bacterioplankton communities in an Arctic fjord

In the frame of the European Project on Ocean Acidification (EPOCA), the response of an Arctic pelagic community (<3 mm) to a gradient of seawater pCO(2) was investigated. For this purpose 9 large-scale in situ mesocosms were deployed in Kongsfjorden, Svalbard (78 degrees 56.2' N, 11 degrees 53.6' E), in 2010. The present study investigates effects on the communities of particle-attached (PA; >3 mu m) and free-living (FL; <3 mu m > 0.2 mu m) bacteria by Automated Ribosomal Intergenic Spacer Analysis (ARISA) in 6 of the mesocosms, ranging from 185 to 1050 mu atm initial pCO(2), 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 (similar to 185-685 mu atm) by about 25 %, while they were more or less stable at high CO2 (similar to 820-1050 mu atm). 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.

Assessing the environmental consequences of CO2 leakage from geological CCS: Generating evidence to support environmental risk assessment

At the start of the industrial revolution (circa 1750) the atmospheric concentration of carbon dioxide (CO2) was around 280 ppm. Since that time the burning of fossil fuel, together with other industrial processes such as cement manufacture and changing land use, has increased this value to 400 ppm, for the first time in over 3 million years. With CO2 being a potent greenhouse gas, the consequence of this rise for global temperatures has been dramatic, and not only for air temperatures. Global Sea Surface Temperature (SST) has warmed by 0.4–0.8 °C during the last century, although regional differences are evident (IPCC, 2007). This rise in atmospheric CO2 levels and the resulting global warming to some extent has been ameliorated by the oceanic uptake of around one quarter of the anthropogenic CO2 emissions (Sabine et al., 2004). Initially this was thought to be having little or no impact on ocean chemistry due to the capacity of the ocean’s carbonate buffering system to neutralise the acidity caused when CO2 dissolves in seawater. However, this assumption was challenged by Caldeira and Wickett (2005) who used model predictions to show that the rate at which carbonate buffering can act was far too slow to moderate significant changes to oceanic chemistry over the next few centuries. Their model predicted that since pre-industrial times, ocean surface water pH had fallen by 0.1 pH unit, indicating a 30% increase in the concentration of H+ ions. Their model also showed that the pH of surface waters could fall by up to 0.4 units before 2100, driven by continued and unabated utilisation of fossil fuels. Alongside increasing levels of dissolved CO2 and H+ (reduced pH) an increase in bicarbonate ions together with a decrease in carbonate ions occurs. These chemical changes are now collectively recognised as “ocean acidification”. Concern now stems from the knowledge that concentrations of H+, CO2, bicarbonate and carbonate ions impact upon many important physiological processes vital to maintaining health and function in marine organisms. Additionally, species have evolved under conditions where the carbonate system has remained relatively stable for millions of years, rendering them with potentially reduced capacity to adapt to this rapid change. Evidence suggests that, whilst the impact of ocean acidification is complex, when considered alongside ocean warming the net effect on the health and productivity of the oceans will be detrimental.

Long-term responses of North Atlantic calcifying plankton to climate change

The global increase in atmospheric carbon dioxide concentration is potentially threatening marine biodiversity in two ways. First, carbon dioxide and other greenhouse gases accumulating in the atmosphere are causing global warming1. Second, carbon dioxide is altering sea water chemistry, making the ocean more acidic2. Although temperature has a cardinal influence on all biological processes from the molecular to the ecosystem level3, acidification might impair the process of calcification or exacerbate dissolution of calcifying organisms4. Here, we show however that North Atlantic calcifying plankton primarily responded to climate-induced changes in temperatures during the period 1960–2009, overriding the signal from the effects of ocean acidification. We provide evidence that foraminifers, coccolithophores, both pteropod and nonpteropod molluscs and echinoderms exhibited an abrupt shift circa 1996 at a time of a substantial increase in temperature5 and that some taxa exhibited a poleward movement in agreement with expected biogeographical changes under sea temperature warming6,7. Although acidification may become a serious threat to marine calcifying organisms, our results suggest that over the study period the primary driver of North Atlantic calcifying planktonwas oceanic temperature.

The importance of zooplankton in reducing levels of atmospheric CO sub(2) via the biological pump

Zooplankton play a key role in climate change through the transfer of large quantities of CO sub(2) to the deep ocean by a process known as the biological pump. Plankton composition is crucial as associated mineral material facilitates sinking of carbon rich debris and some taxa package faecal and detrital material. Ocean acidification may impact calcareous groups. Zooplankton have also been shown to be highly sensitive indicators of environmental change. Results will be presented to show that ocean temperature, circulation and planktonic ecosystems (using data from the Continuous Plankton Recorder, CPR survey) in the North Atlantic are changing rapidly in concert and that there is evidence to suggest that the changes are an ocean wide response to global warming with potential feedback effects. Given the importance of the oceans to the carbon cycle, even a minor change in the flux of carbon to the deep ocean would have a big impact increasing growth of atmospheric CO sub(2). We have virtually no understanding of the spatial and temporal variability in the efficiency of the biological pump for most of the world's ocean. Establishing new plankton monitoring programmes backed up by appropriate research to help understand processes is needed to address this gap in knowledge. There is little doubt within a global change context and the future of mankind that a potential acceleration in the growth of atmospheric carbon due to a reduction in the efficiency of the biological pump is a key issue for future research in zooplankton ecology.

Impacts of the Oceans on Climate Change

The oceans play a key role in climate regulation especially in part buffering (neutralising) the effects of increasing levels of greenhouse gases in the atmosphere and rising global temperatures. This chapter examines how the regulatory processes performed by the oceans alter as a response to climate change and assesses the extent to which positive feedbacks from the ocean may exacerbate climate change. There is clear evidence for rapid change in the oceans. As the main heat store for the world there has been an accelerating change in sea temperatures over the last few decades, which has contributed to rising sea‐level. The oceans are also the main store of carbon dioxide (CO2), and are estimated to have taken up ∼40% of anthropogenic-sourced CO2 from the atmosphere since the beginning of the industrial revolution. A proportion of the carbon uptake is exported via the four ocean ‘carbon pumps’ (Solubility, Biological, Continental Shelf and Carbonate Counter) to the deep ocean reservoir. Increases in sea temperature and changing planktonic systems and ocean currents may lead to a reduction in the uptake of CO2 by the ocean; some evidence suggests a suppression of parts of the marine carbon sink is already underway. While the oceans have buffered climate change through the uptake of CO2 produced by fossil fuel burning this has already had an impact on ocean chemistry through ocean acidification and will continue to do so. Feedbacks to climate change from acidification may result from expected impacts on marine organisms (especially corals and calcareous plankton), ecosystems and biogeochemical cycles. The polar regions of the world are showing the most rapid responses to climate change. As a result of a strong ice–ocean influence, small changes in temperature, salinity and ice cover may trigger large and sudden changes in regional climate with potential downstream feedbacks to the climate of the rest of the world. A warming Arctic Ocean may lead to further releases of the potent greenhouse gas methane from hydrates and permafrost. The Southern Ocean plays a critical role in driving, modifying and regulating global climate change via the carbon cycle and through its impact on adjacent Antarctica. The Antarctic Peninsula has shown some of the most rapid rises in atmospheric and oceanic temperature in the world, with an associated retreat of the majority of glaciers. Parts of the West Antarctic ice sheet are deflating rapidly, very likely due to a change in the flux of oceanic heat to the undersides of the floating ice shelves. The final section on modelling feedbacks from the ocean to climate change identifies limitations and priorities for model development and associated observations. Considering the importance of the oceans to climate change and our limited understanding of climate-related ocean processes, our ability to measure the changes that are taking place are conspicuously inadequate. The chapter highlights the need for a comprehensive, adequately funded and globally extensive ocean observing system to be implemented and sustained as a high priority. Unless feedbacks from the oceans to climate change are adequately included in climate change models, it is possible that the mitigation actions needed to stabilise CO2 and limit temperature rise over the next century will be underestimated.

1H NMR metabolomics reveals contrasting response by male and female mussels exposed to reduced seawater pH, increased temperature and a pathogen

Human activities are fundamentally altering the chemistry of the world's oceans. Ocean acidification (OA) is occurring against a background of warming and an increasing occurrence of disease outbreaks, posing a significant threat to marine organisms, communities, and ecosystems. In the current study, (1)H NMR spectroscopy was used to investigate the response of the blue mussel, Mytilus edulis, to a 90-day exposure to reduced seawater pH and increased temperature, followed by a subsequent pathogenic challenge. Analysis of the metabolome revealed significant differences between male and female organisms. Furthermore, males and females are shown to respond differently to environmental stress. While males were significantly affected by reduced seawater pH, increased temperature, and a bacterial challenge, it was only a reduction in seawater pH that impacted females. Despite impacting males and females differently, stressors seem to act via a generalized stress response impacting both energy metabolism and osmotic balance in both sexes. This study therefore has important implications for the interpretation of metabolomic data in mussels, as well as the impact of environmental stress in marine invertebrates in general.

Heterogeneity of impacts of high CO2 on the North Western European Shelf

The increase in atmospheric CO2 is a dual threat to the marine environment: from one side it drives climate change, leading to modifications in water temperature, circulation patterns and stratification intensity; on the other side it causes a decrease in marine pH (ocean acidification, or OA) due to the increase in dissolved CO2. Assessing the combined impact of climate change and OA on marine ecosystems is a challenging task. The response of the ecosystem to a single driver can be highly variable and remains still uncertain; additionally the interaction between these can be either synergistic or antagonistic. In this work we use the coupled oceanographic–ecosystem model POLCOMS-ERSEM driven by climate forcing to study the interaction between climate change and OA. We focus in particular on carbonate chemistry, primary and secondary production. The model has been run in three different configurations in order to assess separately the impacts of climate change on net primary production and of OA on the carbonate chemistry, which have been strongly supported by scientific literature, from the impact of biological feedbacks of OA on the ecosystem, whose uncertainty still has to be well constrained. The global mean of the projected decrease of pH at the end of the century is about 0.27 pH units, but the model shows significant interaction among the drivers and high variability in the temporal and spatial response. As a result of this high variability, critical tipping point can be locally and/or temporally reached: e.g. undersaturation with respect to aragonite is projected to occur in the deeper part of the central North Sea during summer. Impacts of climate change and of OA on primary and secondary production may have similar magnitude, compensating in some area and exacerbating in others.

Short-term responses of the cold water coral Lophelia pertusa to predicted rises in atmospheric CO2

Cold-water corals are associated with high local biodiversity, but despite their importance as ecosystem engineers, little is known about how these organisms will respond to projected ocean acidification. Since preindustrial times, average ocean pH has decreased from 8.2 to ~8.1, and predicted CO2 emissions will decrease by up to another 0.3 pH units by the end of the century. This decrease in pH may have a wide range of impacts upon marine life, and in particular upon calcifiers such as cold-water corals. Lophelia pertusa is the most widespread cold-water coral (CWC) species, frequently found in the North Atlantic. Here, we present the first short-term (21 days) data on the effects of increased CO2 (750 ppm) upon the metabolism of freshly collected L. pertusa from Mingulay Reef Complex, Scotland, for comparison with net calcification. Over 21 days, corals exposed to increased CO2 conditions had significantly lower respiration rates (11.4±1.39 SE, µmol O2 g−1 tissue dry weight h−1) than corals in control conditions (28.6±7.30 SE µmol O2 g−1 tissue dry weight h−1). There was no corresponding change in calcification rates between treatments, measured using the alkalinity anomaly technique and 14C uptake. The decrease in respiration rate and maintenance of calcification rate indicates an energetic imbalance, likely facilitated by utilisation of lipid reserves. These data from freshly collected L. pertusa from the Mingulay Reef Complex will help define the impact of ocean acidification upon the growth, physiology and structural integrity of this key reef framework forming species.

Fine-scale nutrient and carbonate system dynamics around cold-water coral reefs in the northeast Atlantic.

Ocean acidification has been suggested as a serious threat to the future existence of cold-water corals (CWC). However, there are few fine-scale temporal and spatial datasets of carbonate and nutrients conditions available for these reefs, which can provide a baseline definition of extant conditions. Here we provide observational data from four different sites in the northeast Atlantic that are known habitats for CWC. These habitats differ by depth and by the nature of the coral habitat. At depths where CWC are known to occur across these sites the dissolved inorganic carbon ranged from 2088 to 2186 μmol kg−1, alkalinity ranged from 2299 to 2346 μmol kg−1, and aragonite Ω ranged from 1.35 to 2.44. At two sites fine-scale hydrodynamics caused increased variability in the carbonate and nutrient conditions over daily time-scales. The observed high level of variability must be taken into account when assessing CWC sensitivities to future environmental change.

Response of an Arctic Sediment Nitrogen Cycling Community to Increased CO2

Ocean acidification influences sediment/water nitrogen fluxes, possibly by impacting on the microbial process of ammonia oxidation. To investigate this further, undisturbed sediment cores collected from Ny Alesund harbour (Svalbard) were incubated with seawater adjusted to CO2 concentrations of 380, 540, 760, 1,120 and 3,000 μatm. DNA and RNA were extracted from the sediment surface after 14 days' exposure and the abundance of bacterial and archaeal ammonia oxidising (amoA) genes and transcripts quantified using quantitative polymerase chain reaction. While there was no change to the abundance of bacterial amoA genes, an increase to 760 μatm pCO2 reduced the abundance of bacterial amoA transcripts by 65 %, and this was accompanied by a shift in the composition of the active community. In contrast, archaeal amoA gene and transcript abundance both doubled at 3,000 μatm, with an increase in species richness also apparent. This suggests that ammonia oxidising bacteria and archaea in marine sediments have different pH optima, and the impact of elevated CO2 on N cycling may be dependent on the relative abundances of these two major microbial groups. Further evidence of a shift in the balance of key N cycling groups was also evident: the abundance of nirS-type denitrifier transcripts decreased alongside bacterial amoA transcripts, indicating that NO3 − produced by bacterial nitrification fuelled denitrification. An increase in the abundance of Planctomycete-specific 16S rRNA, the vastmajority of which grouped with known anammox bacteria, was also apparent at 3,000 μatm pCO2. This could indicate a possible shift from coupled nitrification–denitrification to anammox activity at elevated CO2.