Combined effects of CO2 and light on large and small isolates of the unicellular N2-fixing cyanobacterium Crocosphaera watsonii from the western tropical Atlantic Ocean

We examined the combined effects of light and pCO2 on growth, CO2-fixation and N2-fixation rates by strains of the unicellular marine N2-fixing cyanobacterium Crocosphaera watsonii with small (WH0401) and large (WH0402) cells that were isolated from the western tropical Atlantic Ocean. In low-pCO2-acclimated cultures (190 ppm) of WH0401, growth, CO2-fixation and N2-fixation rates were significantly lower than those in cultures acclimated to higher (present-day 385 ppm, or future 750 ppm) pCO2 treatments. Growth rates were not significantly different, however, in low-pCO2-acclimated cultures of WH0402 in comparison with higher pCO2 treatments. Unlike previous reports for C. watsonii (strain WH8501), N2-fixation rates did not increase further in cultures of WH0401 or WH0402 when acclimated to 750 ppm relative to those maintained at present-day pCO2. Both light and pCO2 had a significant negative effect on gross : net N2-fixation rates in WH0402 and trends were similar in WH0401, implying that retention of fixed N was enhanced under elevated light and pCO2. These data, along with previously reported results, suggest that C. watsonii may have wide-ranging, strain-specific responses to changing light and pCO2, emphasizing the need for examining the effects of global change on a range of isolates within this biogeochemically important genus. In general, however, our data suggest that cellular N retention and CO2-fixation rates of C. watsonii may be positively affected by elevated light and pCO2 within the next 100 years, potentially increasing trophic transfer efficiency of C and N and thereby facilitating uptake of atmospheric carbon by the marine biota.

Reduced calcification and lack of acclimatization by coral colonies growing in areas of persistent natural acidification

As the surface ocean equilibrates with rising atmospheric CO2, the pH of surface seawater is decreasing with potentially negative impacts on coral calcification. A critical question is whether corals will be able to adapt or acclimate to these changes in seawater chemistry. We use high precision CT scanning of skeletal cores of Porites astreoides, an important Caribbean reef-building coral, to show that calcification rates decrease significantly along a natural gradient in pH and aragonite saturation (Omega arag). This decrease is accompanied by an increase in skeletal erosion and predation by boring organisms. The degree of sensitivity to reduced ?arag measured on our field corals is consistent with that exhibited by the same species in laboratory CO2 manipulation experiments. We conclude that the Porites corals at our field site were not able to acclimatize enough to prevent the impacts of local ocean acidification on their skeletal growth and development, despite spending their entire lifespan in low pH, low Omega arag seawater.

Impact of CO2-driven acidification on the development of the sea cucumber Apostichopus japonicus (Selenka) (Echinodermata: Holothuroidea)

We evaluated the impact of ocean acidification on the early development of sea cucumber Apostichopus japonicus. The effect of pH-levels (pH 8.04, 7.85, 7.70 and 7.42) were tested on post-fertilization success, developmental (stage duration) and growth rates. Post-fertilization success decreased linearly with pH leading to a 6% decrease at pH 7.42 as compared to pH 8.1. The impact of pH on developmental time was stage-dependent: (1) stage duration increased linearly with decreasing pH in early-auricularia stage; (2) decreased linearly with decreasing pH in the mid-auricularia stage; but (3) pH decline had no effect on the late-auricularia stage. At the end of the experiment, the size of doliolaria larvae linearly increased with decreasing pH. In conclusion, a 0.62 unit decrease in pH had relatively small effects on A. japonicus early life-history compared to other echinoderms, leading to a maximum of 6% decrease in post-fertilization success and subtle effects on growth and development.

Rapid acclimation of juvenile corals to CO2-mediated acidification by upregulation of heat shock protein and Bcl-2 genes

Corals play a key role in ocean ecosystems and carbonate balance, but their molecular response to ocean acidification remains unclear. The only previous whole-transcriptome study documented extensive disruption of gene expression, particularly of genes encoding skeletal organic matrix proteins, in juvenile corals (Acropora millepora) after short-term (3 d) exposure to elevated pCO2. In this study, whole-transcriptome analysis was used to compare the effects of such 'acute' (3 d) exposure to elevated pCO2 with a longer ('prolonged'; 9 d) period of exposure beginning immediately post-fertilization. Far fewer genes were differentially expressed under the 9-d treatment, and although the transcriptome data implied wholesale disruption of metabolism and calcification genes in the acute treatment experiment, expression of most genes was at control levels after prolonged treatment. There was little overlap between the genes responding to the acute and prolonged treatments, but heat shock proteins (HSPs) and heat shock factors (HSFs) were over-represented amongst the genes responding to both treatments. Amongst these was an HSP70 gene previously shown to be involved in acclimation to thermal stress in a field population of another acroporid coral. The most obvious feature of the molecular response in the 9-d treatment experiment was the upregulation of five distinct Bcl-2 family members, the majority predicted to be anti-apoptotic. This suggests that an important component of the longer term response to elevated CO2 is suppression of apoptosis. It therefore appears that juvenile A. millepora have the capacity to rapidly acclimate to elevated pCO2, a process mediated by upregulation of specific HSPs and a suite of Bcl-2 family members.

Differing responses of three Southern Ocean Emiliania huxleyi ecotypes to changing seawater carbonate chemistry

The invasion of anthropogenic carbon dioxide into the surface ocean is altering seawater carbonate speciation, a process commonly called ocean acidification. The high latitude waters of the Southern Ocean are one of the primary and most severely affected regions. Coccolithophores are an important phytoplankton group, responsible for the majority of pelagic calcium carbonate production in the world's oceans, with a distribution that ranges from tropical to polar waters. Emiliania huxleyi is numerically the most abundant coccolithophore species and appears in several different ecotypes. We tested the effects of ocean acidification on 3 carefully selected E. huxleyi ecotypes isolated from the Southern Ocean. Their responses were measured in terms of growth, photosynthesis, calcification, cellular geometry, and stoichiometry. The 3 ecotypes exhibited differing sensitivities in regards to seawater carbonate chemistry when cultured at the same temperature (14°C) and continuous light (110 µmol photons/m2/s). Under future ocean acidification scenarios, particulate inorganic to organic carbon ratios (PIC:POC) decreased by 38-44, 47-51 and 71-98% in morphotype A 'over-calcified' (A o/c), A and B/C, respectively. All ecotypes reduced their rate of calcification, but the cold-water adapted ecotype (morphotype B/C) was by far the most sensitive, and almost ceased calcification at partial pressure of carbon dioxide ( pCO2) levels above 1000 µatm. We recommend that future surveys for E. huxleyi cells in the Southern Ocean should include the capability of recognising 'naked cells' by molecular and microscopic tools. The distinct differences in the physiological responses of these 3 dominant Southern Ocean coccolithophore ecotypes are likely to have consequences for future coccolithophore community structures and thereby the Southern Ocean carbon cycle.

The effects of water acidification, temperature and salinity on the regenerative capacity of the polychaete Diopatra neapolitana

Changes in seawater pH, temperature and salinity are expected to occur in the near future, which can be a threat to aquatic systems, mainly for marine coastal areas, and their inhabiting species. Hence, the present study proposes to evaluate the effects of temperature shifts, pH decrease and salinity changes in the tissue's regenerative capacity of the polychaete Diopatra neapolitana. This study evidenced that D. neapolitana individuals exposed to lower pH exhibited a significantly lower capacity to regenerate their body, while with the increase of temperature individuals showed a higher capacity to regenerate their tissues. Furthermore, the present work demonstrated that individuals exposed to salinities 28 and 35 did not present significant differences between them, while salinities 21 and 42 negatively influenced the regenerative capacity of D. neapolitana. At the end of regeneration, comparing all conditions, high salinity (42) seemed to have a greater impact on the regenerative capacity of individuals than the other factors, since under this condition individuals took longer to completely regenerate. Overall, this study demonstrated that variations in abiotic factors can strongly affect D. neapolitana's performance.

Photophysiological responses of Southern Ocean phytoplankton to changes in CO2 concentrations: Short-term versus acclimation effects

The present study examines how different pCO2 acclimations affect the CO2- and light-dependence of photophysiological processes and O2 fluxes in four Southern Ocean (SO) key phytoplankton species. We grew Chaetoceros debilis (Cleve), Pseudo-nitzschia subcurvata (Hasle), Fragilariopsis kerguelensis (O'Meara) and Phaeocystis antarctica (Karsten) under low (160 µatm) and high (1000 ?atm) pCO2. The CO2- and light-dependence of fluorescence parameters of photosystem II (PSII) were determined by means of a fluorescence induction relaxation system (FIRe). In all tested species, nonphotochemical quenching (NPQ) is the primary photoprotection strategy in response to short-term exposure to high light or low CO2 concentrations. In C. debilis and P. subcurvata, PSII connectivity (p) and functional absorption cross-sections of PSII in ambient light (sigma PSII') also contributed to photoprotection while changes in re-oxidation times of Qa acceptor (tQa) were more significant in F. kerguelensis. The latter was also the only species being responsive to high acclimation pCO2, as these cells had enhanced relative electron transport rates (rETRs) and sigma PSII' while tQa and p were reduced under short-term exposure to high irradiance. Low CO2-acclimated cells of F. kerguelensis and all pCO2 acclimations of C. debilis and P. subcurvata showed dynamic photoinhibition with increasing irradiance. To test for the role and presence of the Mehler reaction in C. debilis and P. subcurvata, the light-dependence of O2 fluxes was estimated using membrane inlet mass spectrometry (MIMS). Our results show that the Mehler reaction is absent in both species under the tested conditions. We also observed that dark respiration was strongly reduced under high pCO2 in C. debilis while it remained unaltered in P. subcurvata. Our study revealed species-specific differences in the photophysiological responses to pCO2, both on the acclimation as well as the short-term level.

European sea bass, Dicentrarchus labrax, in a changing ocean

Ocean acidification, caused by rising concentrations of carbon dioxide (CO2), is widely considered to be a major global threat to marine ecosystems. To investigate the potential effects of ocean acidification on the early life stages of a commercially important fish species, European sea bass (Dicentrarchus labrax), 12 000 larvae were incubated from hatch through metamorphosis under a matrix of two temperatures (17 and 19 °C) and two seawater pCO2 levels (ambient and 1,000 µatm) and sampled regularly for 42 days. Calculated daily mortality was significantly affected by both temperature and pCO2, with both increased temperature and elevated pCO2 associated with lower daily mortality and a significant interaction between these two factors. There was no significant pCO2 effect noted on larval morphology during this period but larvae raised at 19 °C possessed significantly larger eyes and lower carbon:nitrogen ratios at the end of the study compared to those raised under 17 °C. Similarly, when the incubation was continued to post-metamorphic (juvenile) animals (day 67-69), fish raised under a combination of 19 °C and 1000 µatm pCO2 were significantly heavier. However, juvenile D. labrax raised under this combination of 19 °C and 1000 µatm pCO2 also exhibited lower aerobic scopes than those incubated at 19 °C and ambient pCO2. Most studies investigating the effects of near-future oceanic conditions on the early life stages of marine fish have used incubations of relatively short durations and suggested that these animals are resilient to ocean acidification. Whilst the increased survival and growth observed in this study supports this view, we conclude that more work is required to investigate whether the differences in juvenile physiology observed in this study manifest as negative impacts in adult fish.

Seawater carbonate chemistry and protein and chlorophyll a content per nubbin of Acropora muricata during observations of La Saline fringing reef, La Reunion Island, western Indian Ocean, 2011

The effect of decreasing aragonite saturation state (Omega Arag) of seawater (elevated pCO2) on calcification rates of Acropora muricata was studied using nubbins prepared from parent colonies located at two sites of La Saline reef (La Réunion Island, western Indian Ocean): a back-reef site (BR) affected by nutrient-enriched groundwater discharge (mainly nitrate), and a reef flat site (RF) with low terrigenous inputs. Protein and chlorophyll a content of the nubbins, as well as zooxanthellae abundance, were lower at RF than BR. Nubbins were incubated at ~27°C over 2 h under sunlight, in filtered seawater manipulated to get differing initial pCO2 (1,440-340 µatm), Omega Arag (1.4-4.0), and dissolved inorganic carbon (DIC) concentrations (2,100-1,850 µmol/kg). Increasing DIC concentrations at constant total alkalinity (AT) resulted in a decrease in Omega Arag and an increase in pCO2. AT at the beginning of the incubations was kept at a natural level of 2,193 ± 6 µmol/kg (mean ± SD). Net photosynthesis (NP) and calcification were calculated from changes in pH and AT during the incubations. Calcification decrease in response to doubling pCO2 relative to preindustrial level was 22% for RF nubbins. When normalized to surface area of the nubbins, (1) NP and calcification were higher at BR than RF, (2) NP increased in high pCO2 treatments at BR compared to low pCO2 treatments, and (3) calcification was not related to Omega Arag at BR. When normalized to NP, calcification was linearly related to Omega Arag at both sites, and the slopes of the relationships were not significantly different. The increase in NP at BR in the high pCO2 treatments may have increased calcification and thus masked the negative effect of low Omega Arag on calcification. Removing the effect of NP variations at BR showed that calcification declined in a similar manner with decreased Omega Arag (increased pCO2) whatever the nutrient loading.

Mesocosm experiment on warming and acidification effects in 2012: Zooplankton data

Concerns about increasing atmospheric CO2 concentrations and global warming have initiated studies on the consequences of multiple-stressor interactions on marine organisms and ecosystems. We present a fully-crossed factorial mesocosm study and assess how warming and acidification affect the abundance, body size, and fatty acid composition of copepods as a measure of nutritional quality. The experimental set-up allowed us to determine whether the effects of warming and acidification act additively, synergistically, or antagonistically on the abundance, body size, and fatty acid content of copepods, a major group of lower level consumers in marine food webs. Copepodite (developmental stages 1-5) and nauplii abundance were antagonistically affected by warming and acidification. Higher temperature decreased copepodite and nauplii abundance, while acidification partially compensated for the temperature effect. The abundance of adult copepods was negatively affected by warming. The prosome length of copepods was significantly reduced by warming, and the interaction of warming and CO2 antagonistically affected prosome length. Fatty acid composition was also significantly affected by warming. The content of saturated fatty acids increased, and the ratios of the polyunsaturated essential fatty acids docosahexaenoic- (DHA) and arachidonic acid (ARA) to total fatty acid content increased with higher temperatures. Additionally, here was a significant additive interaction effect of both parameters on arachidonic acid. Our results indicate that in a future ocean scenario, acidification might partially counteract some observed effects of increased temperature on zooplankton, while adding to others. These may be results of a fertilizing effect on phytoplankton as a copepod food source. In summary, copepod populations will be more strongly affected by warming rather than by acidifying oceans, but ocean acidification effects can modify some temperature impacts