Solar UV irradiances modulate effects of ocean acidification on the Coccolithophorid Emiliania huxleyi

Emiliania huxleyi, the most abundant coccolithophorid in the oceans, is naturally exposed to solar UV radiation (UVR, 280-400 nm) in addition to photosynthetically active radiation (PAR). We investigated the physiological responses of E. huxleyi to the present day and elevated CO2 (390 vs 1000 µatm; with pH(NBS) 8.20 vs 7.86) under indoor constant PAR and fluctuating solar radiation with or without UVR. Enrichment of CO2 stimulated the production rate of particulate organic carbon (POC) under constant PAR, but led to unchanged POC production under incident fluctuating solar radiation. The production rates of particulate inorganic carbon (PIC) as well as PIC/POC ratios were reduced under the elevated CO2, ocean acidification (OA) condition, regardless of PAR levels, and the presence of UVR. However, moderate levels of UVR increased PIC production rates and PIC/POC ratios. OA treatment interacted with UVR to influence the alga's physiological performance, leading to reduced specific growth rate in the presence of UVA (315-400 nm) and decreased quantum yield, along with enhanced nonphotochemical quenching, with addition of UVB (280-315 nm). The results clearly indicate that UV radiation needs to be invoked as a key stressor when considering the impacts of ocean acidification on E. huxleyi.

Effects of elevated CO2 on fish behaviour undiminished by transgenerational acclimation

Behaviour and sensory performance of marine fishes are impaired at CO2 levels projected to occur in the ocean in the next 50-100 years, and there is limited potential for within-generation acclimation to elevated CO2. However, whether fish behaviour can acclimate or adapt to elevated CO2 over multiple generations remains unanswered. We tested for transgenerational acclimation of reef fish olfactory preferences and behavioural lateralization at moderate (656 µatm) and high (912 µatm) end-of-century CO2 projections. Juvenile spiny damselfish, Acanthochromis polyacanthus, from control parents (446 µatm) exhibited an innate avoidance to chemical alarm cue (CAC) when reared in control conditions. In contrast, juveniles lost their innate avoidance of CAC and even became strongly attracted to CAC when reared at elevated CO2 levels. Juveniles from parents maintained at mid-CO2 and high-CO2 levels also lost their innate avoidance of CAC when reared in elevated CO2, demonstrating no capacity for transgenerational acclimation of olfactory responses. Behavioural lateralization was also disrupted for juveniles reared under elevated CO2, regardless of parental conditioning. Our results show minimal potential for transgenerational acclimation in this fish, suggesting that genetic adaptation will be necessary to overcome the effects of ocean acidification on behaviour.

Effects of ocean acidification on the photosynthetic performance, carbonic anhydrase activity and growth of the giant kelp Macrocystis pyrifera

Under ocean acidification (OA), the 200 % increase in CO2(aq) and the reduction of pH by 0.3-0.4 units are predicted to affect the carbon physiology and growth of macroalgae. Here we examined how the physiology of the giant kelp Macrocystis pyrifera is affected by elevated pCO2/low pH. Growth and photosynthetic rates, external and internal carbonic anhydrase (CA) activity, HCO3 (-) versus CO2 use were determined over a 7-day incubation at ambient pCO2 400 µatm/pH 8.00 and a future OA treatment of pCO2 1200 µatm/pH 7.59. Neither the photosynthetic nor growth rates were changed by elevated CO2 supply in the OA treatment. These results were explained by the greater use of HCO3 (-) compared to CO2 as an inorganic carbon (Ci) source to support photosynthesis. Macrocystis is a mixed HCO3 (-) and CO2 user that exhibits two effective mechanisms for HCO3 (-) utilization; as predicted for species that possess carbon-concentrating mechanisms (CCMs), photosynthesis was not substantially affected by elevated pCO2. The internal CA activity was also unaffected by OA, and it remained high and active throughout the experiment; this suggests that HCO3 (-) uptake via an anion exchange protein was not affected by OA. Our results suggest that photosynthetic Ci uptake and growth of Macrocystis will not be affected by elevated pCO2/low pH predicted for the future, but the combined effects with other environmental factors like temperature and nutrient availability could change the physiological response of Macrocystis to OA. Therefore, further studies will be important to elucidate how this species might respond to the global environmental change predicted for the ocean.

Annual response of two Mediterranean azooxanthellate temperate corals to low-pH and high-temperature conditions

Ocean acidification (OA) and warming related to the anthropogenic increase in atmospheric CO2 have been shown to have detrimental effects on several marine organisms, especially those with calcium carbonate structures such as corals. In this study, we evaluate the response of two Mediterranean shallow-water azooxanthellate corals to the projected pH and seawater temperature (ST) scenarios for the end of this century. The colonial coral Astroides calycularis and the solitary Leptopsammia pruvoti were grown in aquaria over a year under two fixed pH conditions, control (8.05 pHT units) and low (7.72 pHT units), and simulating two annual ST cycles, natural and high (+3 °C). The organic matter (OM), lipid and protein content of the tissue and the skeletal microdensity of A. calycularis were not affected by the stress conditions (low pH, high ST), but the species exhibited a mean 25 % decrease in calcification rate at high-ST conditions at the end of the warm period and a mean 10 % increase in skeletal porosity under the acidified treatment after a full year cycle. Conversely, an absence of effects on calcification and skeletal microdensity of L. pruvoti exposed to low-pH and high-ST treatments contrasted with a significant decrease in the OM, lipid and protein content of the tissue at high-ST conditions and a 13 % mean increase in the skeletal porosity under low-pH conditions following a full year of exposure. This species-specific response suggests that different internal self-regulation strategies for energy reallocation may allow certain shallow-water azooxanthellate corals to cope more successfully than others with global environmental changes.

Effects of Ocean Acidification and Warming on Sperm Activity and Early Life Stages of the Mediterranean Mussel (Mytilus galloprovincialis)

Larval stages are among those most vulnerable to ocean acidification (OA). Projected atmospheric CO2 levels for the end of this century may lead to negative impacts on communities dominated by calcifying taxa with planktonic life stages. We exposed Mediterranean mussel (Mytilus galloprovincialis) sperm and early life stages to pHT levels of 8.0 (current pH) and 7.6 (2100 level) by manipulating pCO2 level (380 and 1000 ppm). Sperm activity was examined at ambient temperatures (16-17 °C) using individual males as replicates. We also assessed the effects of temperature (ambient and = 20 °C) and pH on larval size, survival, respiration and calcification of late trochophore/early D-veliger stages using a cross-factorial design. Increased pCO2 had a negative effect on the percentage of motile sperm (mean response ratio R= 71%) and sperm swimming speed (R= 74%), possibly indicating reduced fertilization capacity of sperm in low concentrations. Increased temperature had a more prominent effect on larval stages than pCO2, reducing performance (RSize = 90% and RSurvival = 70%) and increasing energy demand (RRespiration = 429%). We observed no significant interactions between pCO2 and temperature. Our results suggest that increasing temperature might have a larger impact on very early larval stages of M. galloprovincialis than OA at levels predicted for the end of the century.

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.

Effects of ultraviolet radiation and CO2 increase on winter phytoplankton assemblages in a temperate coastal lagoon

Increases in ultraviolet radiation (UVR) and CO2 affect phytoplankton growth and mortality in a variety of different ways. However, in situ responses of natural phytoplankton communities to climate change, as well as its effects on phytoplankton annual cycles, are still largely unknown. Although temperature and UVR have been increasing in temperate latitudes during winter, this season is still particularly neglected in climate change studies, being considered a non-active season regarding phytoplankton growth and production. Additionally, coastal lagoons are highly productive ecosystems and very vulnerable to climate change. This study aims, therefore, to evaluate the short-term effects of increased UVR and CO2 on the composition and growth of winter phytoplankton assemblages in a temperate coastal lagoon. During winter 2012, microcosm experiments were used to evaluate the isolated and combined effects of UVR and CO2, under ambient and high CO2 treatments, exposed to ambient UV levels and photosynthetically active radiation (PAR), or to PAR only. Phytoplankton composition, abundance, biomass and photosynthetic parameters were evaluated during the experiments. Significant changes were observed in the growth of specific phytoplankton groups, leading to changes in community composition. The cyanobacterium Synechococcus was dominant at the beginning of the experiment, but it was negatively affected by UVR and CO2. Diatoms clearly benefited from high CO2 and UVR, particularly Thalassiosira. Despite the changes observed in specific phytoplankton groups, growth and production of the whole phytoplankton community did not show significant responses to UVR and/or CO2.

Seawater carbonate chemistry and combined mechanistic effects of CO2 and light on the N2-fixing cyanobacterium Trichodesmium IMS101, 2010

The marine diazotrophic cyanobacterium Trichodesmium responds to elevated atmospheric CO2 partial pressure (pCO2) with higher N2 fixation and growth rates. To unveil the underlying mechanisms, we examined the combined influence of pCO2(150 and 900 µatm) and light (50 and 200 µmol photons m-2 s-1) on TrichodesmiumIMS101. We expand on a complementary study that demonstrated that while elevated pCO2 enhanced N2 fixation and growth, oxygen evolution and carbon fixation increased mainly as a response to high light. Here, we investigated changes in the photosynthetic fluorescence parameters of photosystem II, in ratios of the photosynthetic units (photosystem I:photosystem II), and in the pool sizes of key proteins involved in the fixation of carbon and nitrogen as well as their subsequent assimilation. We show that the combined elevation in pCO2 and light controlled the operation of the CO2-concentrating mechanism and enhanced protein activity without increasing their pool size. Moreover, elevated pCO2 and high light decreased the amounts of several key proteins (NifH, PsbA, and PsaC), while amounts of AtpB and RbcL did not significantly change. Reduced investment in protein biosynthesis, without notably changing photosynthetic fluxes, could free up energy that can be reallocated to increase N2 fixation and growth at elevated pCO2 and light. We suggest that changes in the redox state of the photosynthetic electron transportchain and posttranslational regulation of key proteins mediate the high flexibility in resources and energy allocation in Trichodesmium. This strategy should enableTrichodesmium to flourish in future surface oceans characterized by elevated pCO2, higher temperatures, and high light.

The effects of thermal and high-CO2 stresses on the metabolism and surrounding microenvironment of the coral Galaxea fascicularis

The effects of elevated temperature and high pCO2 on the metabolism of Galaxea fascicularis were studied with oxygen and pH microsensors. Photosynthesis and respiration rates were evaluated from the oxygen fluxes from and to the coral polyps. High-temperature alone lowered both photosynthetic and respiration rates. High pCO2 alone did not significantly affect either photosynthesis or respiration rates. Under a combination of high-temperature and high-CO2, the photosynthetic rate increased to values close to those of the controls. The same pH in the diffusion boundary layer was observed under light in both (400 and 750 ppm) CO2 treatments, but decreased significantly in the dark as a result of increased CO2. The ATP contents decreased with increasing temperature. The effects of temperature on the metabolism of corals were stronger than the effects of increased CO2. The effects of acidification were minimal without combined temperature stress. However, acidification combined with higher temperature may affect coral metabolism due to the amplification of diel variations in the microenvironment surrounding the coral and the decrease in ATP contents.

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

Mixed effects of elevated pCO2 on fertilisation, larval and juvenile development and adult responses in the mobile subtidal scallop Mimachlamys asperrima (Lamarck, 1819)

Ocean acidification is predicted to have severe consequences for calcifying marine organisms especially molluscs. Recent studies, however, have found that molluscs in marine environments with naturally elevated or fluctuating CO2 or with an active, high metabolic rate lifestyle may have a capacity to acclimate and be resilient to exposures of elevated environmental pCO2. The aim of this study was to determine the effects of near future concentrations of elevated pCO2 on the larval and adult stages of the mobile doughboy scallop, Mimachlamys asperrima from a subtidal and stable physio-chemical environment. It was found that fertilisation and the shell length of early larval stages of M. asperrima decreased as pCO2 increased, however, there were less pronounced effects of elevated pCO2 on the shell length of later larval stages, with high pCO2 enhancing growth in some instances. Byssal attachment and condition index of adult M. asperrima decreased with elevated pCO2, while in contrast there was no effect on standard metabolic rate or pHe. The responses of larval and adult M. asperrima to elevated pCO2 measured in this study were more moderate than responses previously reported for intertidal oysters and mussels. Even this more moderate set of responses are still likely to reduce the abundance of M. asperrima and potentially other scallop species in the world's oceans at predicted future pCO2 levels.

Ocean acidification outweighs nutrient effects in structuring seagrass epiphyte communities

1. Developing a framework for assessing interactions between multiple anthropogenic stressors remains an important goal in environmental research. In coastal ecosystems, the relative effects of aspects of global climate change (e.g. CO2 concentrations) and localized stressors (e.g. eutrophication), in combination, have received limited attention. 2. Using a long-term (11 month) field experiment, we examine how epiphyte assemblages in a tropical seagrass meadow respond to factorial manipulations of dissolved carbon dioxide (CO2(aq)) and nutrient enrichment. In situ CO2(aq) manipulations were conducted using clear, open-top chambers, which replicated carbonate parameter forecasts for the year 2100. Nutrient enrichment consisted of monthly additions of slow-release fertilizer, nitrogen (N) and phosphorus (P), to the sediments at rates equivalent to theoretical maximum rates of anthropogenic loading within the region (1.54 g N/m**2/d and 0.24 g P m**2/d). 3. Epiphyte community structure was assessed on a seasonal basis and revealed declines in the abundance of coralline algae, along with increases in filamentous algae under elevated CO2(aq). Surprisingly, nutrient enrichment had no effect on epiphyte community structure or overall epiphyte loading. Interactions between CO2(aq) and nutrient enrichment were not detected. Furthermore, CO2(aq)-mediated responses in the epiphyte community displayed strong seasonality, suggesting that climate change studies in variable environments should be conducted over extended time-scales. 4. Synthesis. The observed responses indicate that for certain locations, global stressors such as ocean acidification may take precedence over local eutrophication in altering the community structure of seagrass epiphyte assemblages. Given that nutrient-driven algal overgrowth is commonly cited as a widespread cause of seagrass decline, our findings highlight that alternate climate change forces may exert proximate control over epiphyte community structure.

Study on the effects of near-future ocean acidification on marine yeasts: a microcosm approach

Marine yeasts play an important role in biodegradation and nutrient cycling and are often associated with marine flora and fauna. They show maximum growth at pH levels lower than present-day seawater pH. Thus, contrary to many other marine organisms, they may actually profit from ocean acidification. Hence, we conducted a microcosm study, incubating natural seawater from the North Sea at present-day pH (8.10) and two near-future pH levels (7.81 and 7.67). Yeasts were isolated from the initial seawater sample and after 2 and 4 weeks of incubation. Isolates were classified by matrix-assisted laser desorption/ionization time-of-flight mass spectrometry (MALDI-TOF MS) and representative isolates were identified by partial sequencing of the large subunit rRNA gene. From the initial seawater sample, we predominantly isolated a yeast-like filamentous fungus related to Aureobasidium pullulans, Cryptococcus sp., Candida sake, and various cold-adapted yeasts. After incubation, we found more different yeast species at near-future pH levels than at present-day pH. Yeasts reacting to low pH were related to Leucosporidium scottii, Rhodotorula mucilaginosa, Cryptococcus sp., and Debaryomyces hansenii. Our results suggest that these yeasts will benefit from seawater pH reductions and give a first indication that the importance of yeasts will increase in a more acidic ocean.