鱼类对海洋升温与酸化的响应

自工业革命以来,空气中人为排放CO₂量增加,引起温室效应,导致地球表面温度升高和海水升温;同时,由于海-气界面气体交换,大气中CO₂部分溶解于海洋,引起海洋酸化。海洋升温加快鱼体内生化反应和代谢速率,并通过影响生长、觅食和繁殖等生命过程中能量供给,间接影响到鱼类种群分布、群落结构及生态系统的功能。而海水酸化会干扰海洋鱼类仔稚鱼的感觉和行为,增加其被捕食率,并削弱其野外生存能力,可能威胁自然种群补给量。综述了海洋升温、海洋酸化及其两者共同作用对海洋鱼类的影响,为预测鱼类响应全球海洋环境变化的响应趋势提供相关依据。     

Microenvironmental changes support evidence of photosynthesis and calcification inhibition in Halimeda under ocean acidification and warming

The effects of elevated CO₂ and temperature on photosynthesis and calcification of two important calcifying reef algae (Halimeda macroloba and Halimeda cylindracea) were investigated with O₂ microsensors and chlorophyll a fluorometry through a combination of two pCO₂ (400 and 1,200 μatm) and two temperature treatments (28 and 32 °C) equivalent to the present and predicted conditions during the 2100 austral summer. Combined exposure to pCO₂ and elevated temperature impaired calcification and photosynthesis in the two Halimeda species due to changes in the microenvironment around the algal segments and a reduction in physiological performance. There were no significant changes in controls over the 5-week experiment, but there was a 50–70 % decrease in photochemical efficiency (maximum quantum yield), a 70–80 % decrease in O₂ production and a threefold reduction in calcification rate in the elevated CO₂ and high temperature treatment. Calcification in these species is closely coupled with photosynthesis, such that a decrease in photosynthetic efficiency leads to a decrease in calcification. Although pH seems to be the main factor affecting Halimeda species, heat stress also has an impact on their photosystem II photochemical efficiency. There was a strong combined effect of elevated CO₂ and temperature in both species, where exposure to elevated CO₂ or temperature alone decreased photosynthesis and calcification, but exposure to both elevated CO₂ and temperature caused a greater decline in photosynthesis and calcification than in each stress individually. Our study shows that ocean acidification and ocean warming are drivers of calcification and photosynthesis inhibition in Halimeda. Predicted climate change scenarios for 2100 would therefore severely affect the fitness of Halimeda, which can result in a strongly reduced production of carbonate sediments on coral reefs under such changed climate conditions.     

Antagonistic effects of ocean acidification and warming on hunting sharks

Ocean warming and acidification alter the physiological performance and behaviour of many small‐bodied fishes, yet the potential interactive effects of these stressors on larger predators remains poorly understood. In particular, the combined effects of elevated temperature on metabolism and of elevated CO₂ on the behaviour of large predators may not only affect their foraging behaviour, but also the communities in which their prey live. We used a factorial design to assess how projected warming and acidification create synergies or antagonisms between physiological and behavioural processes, such as swimming activity and feeding behaviour through odour tracking and vision. Temperature increased swimming activity during feeding, independent of CO₂. Although temperature also increased motivational drive to locate and accept prey, elevated CO₂ negated chemical and visual behavioural responses that enable effective hunting. Fundamental to these effects was the negligible effect of high CO₂ in isolation, but its power to negate the positive effects of temperature when brought in conjunction. The reduced potential to locate prey due to the interactive effects of ocean acidification and warming, in combination with increases in energetic demand, suggests that energetic tradeoffs will be needed for sharks to sustain themselves at an individual and population level in a future ocean.     

Synergistic effects of ocean acidification and warming on overwintering pteropods in the Arctic

Ocean acidification and warming will be most pronounced in the Arctic Ocean. Aragonite shell‐bearing pteropods in the Arctic are expected to be among the first species to suffer from ocean acidification. Carbonate undersaturation in the Arctic will first occur in winter and because this period is also characterized by low food availability, the overwintering stages of polar pteropods may develop into a bottleneck in their life cycle. The impacts of ocean acidification and warming on growth, shell degradation (dissolution), and mortality of two thecosome pteropods, the polar Limacina helicina and the boreal L. retroversa, were studied for the first time during the Arctic winter in the Kongsfjord (Svalbard). The abundance of L. helicina and L. retroversa varied from 23.5 to 120 ind m^?2 and 12 to 38 ind m^?2, and the mean shell size ranged from 920 to 981 μm and 810 to 823 μm, respectively. Seawater was aragonite‐undersaturated at the overwintering depths of pteropods on two out of ten days of our observations. A 7‐day experiment [temperature levels: 2 and 7 °C, pCO₂ levels: 350, 650 (only for L. helicina) and 880 μatm] revealed a significant pCO₂ effect on shell degradation in both species, and synergistic effects between temperature and pCO₂ for L. helicina. A comparison of live and dead specimens kept under the same experimental conditions indicated that both species were capable of actively reducing the impacts of acidification on shell dissolution. A higher vulnerability to increasing pCO₂ and temperature during the winter season is indicated compared with a similar study from fall 2009. Considering the species winter phenology and the seasonal changes in carbonate chemistry in Arctic waters, negative climate change effects on Arctic thecosomes are likely to show up first during winter, possibly well before ocean acidification effects become detectable during the summer season.     

Differences in neurochemical profiles of two gadid species under ocean warming and acidification

Background

Exposure to future ocean acidification scenarios may alter the behaviour of marine teleosts through interference with neuroreceptor functioning. So far, most studies investigated effects of ocean acidification on the behaviour of fish, either isolated or in combination with environmental temperature. However, only few physiological studies on this issue were conducted despite the putative neurophysiological origin of the CO₂-induced behavioural changes. Here, we present the metabolic consequences of long-term exposure to projected ocean acidification (396–548 μatm PCO₂ under control and 915–1272 μatm under treatment conditions) and parallel warming in the brain of two related fish species, polar cod (Boreogadus saida, exposed to 0 °C, 3 °C, 6 °C and 8 °C) and Atlantic cod (Gadus morhua, exposed to 3 °C, 8 °C, 12 °C and 16 °C). It has been shown that B. saida is behaviourally vulnerable to future ocean acidification scenarios, while G. morhua demonstrates behavioural resilience.

Results

We found that temperature alters brain osmolyte, amino acid, choline and neurotransmitter concentrations in both species indicating thermal responses particularly in osmoregulation and membrane structure. In B. saida, changes in amino acid and osmolyte metabolism at the highest temperature tested were also affected by CO₂, possibly emphasizing energetic limitations. We did not observe changes in neurotransmitters, energy metabolites, membrane components or osmolytes that might serve as a compensatory mechanism against CO₂ induced behavioural impairments. In contrast to B. saida, such temperature limitation was not detected in G. morhua; however, at 8 °C, CO₂ induced an increase in the levels of metabolites of the glutamate/GABA-glutamine cycle potentially indicating greater GABAergic activity in G.morhua. Further, increased availability of energy-rich substrates was detected under these conditions.

Conclusions

Our results indicate a change of GABAergic metabolism in the nervous system of Gadus morhua close to the optimum of the temperature range. Since a former study showed that juvenile G. morhua might be slightly more behaviourally resilient to CO₂ at this respective temperature, we conclude that the observed change of GABAergic metabolism could be involved in counteracting OA induced behavioural changes. This may serve as a fitness advantage of this respective species compared to B. saida in a future warmer, more acidified polar ocean.

     

Incorporating benthic community changes into hydrochemical-based projections of coral reef calcium carbonate production under ocean acidification

Coral Reefs - The existence of coral reefs is dependent on the production and maintenance of calcium carbonate (CaCO3) framework that is produced through calcification. The net production of CaCO3...     

Effects of ocean warming and acidification on the energy budget of an excavating sponge

Recent research efforts have demonstrated increased bioerosion rates under experimentally elevated partial pressures of seawater carbon dioxide (pCO₂) with or without increased temperatures, which may lead to net erosion on coral reefs in the future. However, this conclusion clearly depends on the ability of the investigated bioeroding organisms to survive and grow in the warmer and more acidic future environments, which remains unexplored. The excavating sponge Cliona orientalis Thiele, 1900 is a widely distributed bioeroding organism and symbiotic with dinoflagellates of the genus Symbiodinium. Using C. orientalis, an energy budget model was developed to calculate amounts of carbon directed into metabolic maintenance and growth. This model was tested under a range of CO₂ emission scenarios (temperature + pCO₂) appropriate to an Austral early summer. Under a pre‐industrial scenario, present day (control) scenario, or B1 future scenario (associated with reducing the rate of CO₂ emissions over the next few decades), C. orientalis maintained a positive energy budget, where metabolic demand was likely satisfied by autotrophic carbon provided by Symbiodinium and heterotrophic carbon via filter‐feeding, suggesting sustainability. Under B1, C. orientalis likely benefited by a greater supply of photosynthetic products from its symbionts, which increased by up to 56% per unit area, and displayed an improved condition with up to 52% increased surplus carbon available for growth. Under an A1FI future scenario (associated with ‘business‐as‐usual’ CO₂ emissions) bleached C. orientalis experienced the highest metabolic demand, but carbon acquired was insufficient to maintain the sponge, as indicated by a negative energy budget. These metabolic considerations suggest that previous observations of increased bioerosion under A1FI by C. orientalis may not last through the height of future A1FI summers, and survival of individual sponges may be dependent on the energy reserves (biomass) they have accumulated through the rest of the year.     

Amelioration of ocean acidification and warming effects through physiological buffering of a macroalgae

Concurrent anthropogenic global climate change and ocean acidification are expected to have a negative impact on calcifying marine organisms. While knowledge of biological responses of organisms to oceanic stress has emerged from single‐species experiments, these do not capture ecologically relevant scenarios where the potential for multi‐organism physiological interactions is assessed. Marine algae provide an interesting case study, as their photosynthetic activity elevates pH in the surrounding microenvironment, potentially buffering more acidic conditions for associated epiphytes. We present findings that indicate increased tolerance of an important epiphytic foraminifera, Marginopora vertebralis, to the effects of increased temperature (±3°C) and pCO₂ (~1,000 µatm) when associated with its common algal host, Laurencia intricata. Specimens of M. vertebralis were incubated for 15 days in flow‐through aquaria simulating current and end‐of‐century temperature and pH conditions. Physiological measures of growth (change in wet weight), calcification (measured change in total alkalinity in closed bottles), photochemical efficiency (Fv/Fm), total chlorophyll, photosynthesis (oxygen flux), and respiration were determined. When incubated in isolation, M. vertebralis exhibited reduced growth in end‐of‐century projections of ocean acidification conditions, while calcification rates were lowest in the high‐temperature, low‐pH treatment. Interestingly, association with L. intricata ameliorated these stress effects with the growth and calcification rates of M. vertebralis being similar to those observed in ambient conditions. Total chlorophyll levels in M. vertebralis decreased when in association with L. intricata, while maximum photochemical efficiency increased in ambient conditions. Net production estimates remained similar between M. vertebralis in isolation and in association with L. intricata, although both production and respiration rates of M. vertebralis were significantly higher when associated with L. intricata. These results indicate that the association with L. intricata increases the resilience of M. vertebralis to climate change stress, providing one of the first examples of physiological buffering by a marine alga that can ameliorate the negative effects of changing ocean conditions.     

Paternal identity influences response of Acanthaster planci embryos to ocean acidification and warming

Coral Reefs - The crown-of-thorns sea star Acanthaster planci is a key predator of corals and has had a major influence on the decrease in coral cover across the Indo-Pacific. To understand how...     

Scaling up experimental ocean acidification and warming research: from individuals to the ecosystem

Understanding long‐term, ecosystem‐level impacts of climate change is challenging because experimental research frequently focuses on short‐term, individual‐level impacts in isolation. We address this shortcoming first through an interdisciplinary ensemble of novel experimental techniques to investigate the impacts of 14‐month exposure to ocean acidification and warming (OAW) on the physiology, activity, predatory behaviour and susceptibility to predation of an important marine gastropod (Nucella lapillus). We simultaneously estimated the potential impacts of these global drivers on N. lapillus population dynamics and dispersal parameters. We then used these data to parameterize a dynamic bioclimatic envelope model, to investigate the consequences of OAW on the distribution of the species in the wider NE Atlantic region by 2100. The model accounts also for changes in the distribution of resources, suitable habitat and environment simulated by finely resolved biogeochemical models, under three IPCC global emissions scenarios. The experiments showed that temperature had the greatest impact on individual‐level responses, while acidification had a similarly important role in the mediation of predatory behaviour and susceptibility to predators. Changes in Nucella predatory behaviour appeared to serve as a strategy to mitigate individual‐level impacts of acidification, but the development of this response may be limited in the presence of predators. The model projected significant large‐scale changes in the distribution of Nucella by the year 2100 that were exacerbated by rising greenhouse gas emissions. These changes were spatially heterogeneous, as the degree of impact of OAW on the combination of responses considered by the model varied depending on local‐environmental conditions and resource availability. Such changes in macro‐scale distributions cannot be predicted by investigating individual‐level impacts in isolation, or by considering climate stressors separately. Scaling up the results of experimental climate change research requires approaches that account for long‐term, multiscale responses to multiple stressors, in an ecosystem context.     

海洋酸化和全球变暖对贝类生理生态的影响研究进展

研究表明海洋酸化和全球变暖已严重威胁到海洋生态系统稳的定性及生物多样性。由于人类活动,大气中不断增加的CO2不仅造成全球气候异常,而且大量的CO2被海洋吸收,造成了海水中H+浓度增加,即海洋酸化(Ocean Acidification)。海洋酸化严重影响海洋生物的生存和繁衍,尤其是有壳类生物,如贝类,甲壳类,棘皮类等。主要影响方面包括生物的产卵受精,孵化,早期发育,钙化,酸碱调节,免疫功能,蛋白质合成,基因表达,摄食及能量代谢等一系列和生理相关的机能,进而对个体行为学,种群结构和海洋生态系统造成严重危害。目前,已有大量海洋酸化对海洋贝类的生理生态影响的报道,与此同时,全球变暖导致海洋温度升高伴随着海洋酸化同步发生。因此,为了更加准确地预测海洋生物应对全球气候变化的生理生态应答,越来越多的学者开始致力于研究温度和海洋酸化的复合胁迫对海洋生物交互影响作用。综述了近年来海洋酸化对贝类生理生态的影响,主要从个体早期发育、钙化、免疫、繁殖等方面做了系统的阐述,还对酸化和温度对贝类的复合环境胁迫效应也做了综合分析,以期为今后的海洋酸化研究提供基础理论。     

Ocean warming and acidification alter the behavioral response to flow of the sea urchin Paracentrotus lividus

Ocean warming (OW) and acidification (OA) are intensively investigated as they pose major threats to marine organism. However, little effort is dedicated to another collateral climate change stressor, the increased frequency, and intensity of storm events, here referred to as intensified hydrodynamics. A 2‐month experiment was performed to identify how OW and OA (temperature: 21°C; pH_T: 7.7, 7.4; control: 17°C‐pH_T7.9) affect the resistance to hydrodynamics in the sea urchin Paracentrotus lividus using an integrative approach that includes physiology, biomechanics, and behavior. Biomechanics was studied under both no‐flow condition at the tube foot (TF) scale and flow condition at the individual scale. For the former, TF disk adhesive properties (attachment strength, tenacity) and TF stem mechanical properties (breaking force, extensibility, tensile strength, stiffness, toughness) were evaluated. For the latter, resistance to flow was addressed as the flow velocity at which individuals detached. Under near‐ and far‐future OW and OA, individuals fully balanced their acid‐base status, but skeletal growth was halved. TF adhesive properties were not affected by treatments. Compared to the control, mechanical properties were in general improved under pH_T7.7 while in the extreme treatment (21°C‐pH_T7.4) breaking force was diminished. Three behavioral strategies were implemented by sea urchins and acted together to cope with flow: improving TF attachment, streamlining, and escaping. Behavioral responses varied according to treatment and flow velocity. For instance, individuals at 21°C‐pH_T7.4 increased the density of attached TF at slow flows or controlled TF detachment at fast flows to compensate for weakened TF mechanical properties. They also showed an absence of streamlining favoring an escaping behavior as they ventured in a riskier faster movement at slow flows. At faster flows, the effects of OW and OA were detrimental causing earlier dislodgment. These plastic behaviors reflect a potential scope for acclimation in the field, where this species already experiences diel temperature and pH fluctuations.     

The oceans are changing: impact of ocean warming and acidification on biofouling communities

Climate change (CC) is driving modification of the chemical and physical properties of estuaries and oceans with profound consequences for species and ecosystems. Numerous studies investigate CC effects from species to ecosystem levels, but little is known of the impacts on biofilm communities and on bioactive molecules such as cues, adhesives and enzymes. CC is induced by anthropogenic activity increasing greenhouse emissions leading to rises in air and water temperatures, ocean acidification, sea level rise and changes in ocean gyres and rainfall patterns. These environmental changes are resulting in alterations within marine communities and changes in species ranges and composition. This review provides insights and synthesis of knowledge about the effect of elevated temperature and ocean acidification on microfouling communities and bioactive molecules. The existing studies suggest that CC will impact production of bioactive compounds as well as the growth and composition of biofouling communities. Undoubtedly, with CC fouling management will became an even greater challenge.     

Impacts of ocean acidification on marine organisms: quantifying sensitivities and interaction with warming

Ocean acidification represents a threat to marine species worldwide, and forecasting the ecological impacts of acidification is a high priority for science, management, and policy. As research on the topic expands at an exponential rate, a comprehensive understanding of the variability in organisms' responses and corresponding levels of certainty is necessary to forecast the ecological effects. Here, we perform the most comprehensive meta‐analysis to date by synthesizing the results of 228 studies examining biological responses to ocean acidification. The results reveal decreased survival, calcification, growth, development and abundance in response to acidification when the broad range of marine organisms is pooled together. However, the magnitude of these responses varies among taxonomic groups, suggesting there is some predictable trait‐based variation in sensitivity, despite the investigation of approximately 100 new species in recent research. The results also reveal an enhanced sensitivity of mollusk larvae, but suggest that an enhanced sensitivity of early life history stages is not universal across all taxonomic groups. In addition, the variability in species' responses is enhanced when they are exposed to acidification in multi‐species assemblages, suggesting that it is important to consider indirect effects and exercise caution when forecasting abundance patterns from single‐species laboratory experiments. Furthermore, the results suggest that other factors, such as nutritional status or source population, could cause substantial variation in organisms' responses. Last, the results highlight a trend towards enhanced sensitivity to acidification when taxa are concurrently exposed to elevated seawater temperature.     

Vulnerability of global coral reef habitat suitability to ocean warming,acidification and eutrophication

Coral reefs are threatened by global and local stressors. Yet, reefs appear to respond differently to different environmental stressors. Using a global dataset of coral reef occurrence as a proxy for the long‐term adaptation of corals to environmental conditions in combination with global environmental data, we show here how global (warming: sea surface temperature; acidification: aragonite saturation state, Ω_arag) and local (eutrophication: nitrate concentration, and phosphate concentration) stressors influence coral reef habitat suitability. We analyse the relative distance of coral communities to their regional environmental optima. In addition, we calculate the expected change of coral reef habitat suitability across the tropics in relation to an increase of 0.1°C in temperature, an increase of 0.02 μmol/L in nitrate, an increase of 0.01 μmol/L in phosphate and a decrease of 0.04 in Ω_arag. Our findings reveal that only 6% of the reefs worldwide will be unaffected by local and global stressors and can thus act as temporary refugia. Local stressors, driven by nutrient increase, will affect 22% of the reefs worldwide, whereas global stressors will affect 11% of these reefs. The remaining 61% of the reefs will be simultaneously affected by local and global stressors. Appropriate wastewater treatments can mitigate local eutrophication and could increase areas of temporary refugia to 28%, allowing us to ‘buy time’, while international agreements are found to abate global stressors.