海洋酸化对鱼类感觉和行为影响的研究进展

海洋酸化是指大气增多的二氧化碳(CO2)溶解于海水而导致海水p H值降低的过程。海洋酸化将改变海水碳酸盐平衡体系,使依赖于原化学环境的多种海洋生物乃至生态系统面临巨大威胁。海洋酸化对钙质生物影响的研究最早引起大家关注,而海洋鱼类具有较完善的酸碱调节机制,大家普遍认为酸化对其影响不大。但在过去的5年中,不少实验证明海洋酸化会影响海洋鱼类仔稚鱼的感觉和行为,减弱其野外的生存能力及增加被捕食率,很可能将威胁自然种群补给量和影响全球的渔业资源量。本文从嗅觉、听觉、视觉及高级意识和相关行为角度,综述近几年海洋酸化对鱼类感觉和行为影响的研究进展,介绍了鱼类神经行为生物学的研究,为全面了解和预测海洋酸化的生态、经济和社会效应提供科学依据。     

Towards improved socio-economic assessments of ocean acidification’s impacts

Ocean acidification is increasingly recognized as a component of global change that could have a wide range of impacts on marine organisms, the ecosystems they live in, and the goods and services they provide humankind. Assessment of these potential socio-economic impacts requires integrated efforts between biologists, chemists, oceanographers, economists and social scientists. But because ocean acidification is a new research area, significant knowledge gaps are preventing economists from estimating its welfare impacts. For instance, economic data on the impact of ocean acidification on significant markets such as fisheries, aquaculture and tourism are very limited (if not non-existent), and non-market valuation studies on this topic are not yet available. Our paper summarizes the current understanding of future OA impacts and sets out what further information is required for economists to assess socio-economic impacts of ocean acidification. Our aim is to provide clear directions for multidisciplinary collaborative research.     

Predicted impact of ocean acidification on a marine invertebrate: elevated CO<Subscript>2</Subscript> alters response to thermal stress in sea urchin larvae

Ocean acidification (OA) and the biological consequences of altered seawater chemistry have emerged as a significant environmental threat to healthy marine ecosystems. Because a more acidic ocean interferes with fixation of calcium carbonate to form shells or calcified skeletons, future ocean chemistry may significantly alter the physiology of calcifying marine organisms. These alterations may manifest themselves directly in the calcification process, or have synergistic effects with other environmental factors such as elevated temperatures. New tools permit us to explore subtle changes in gene expression patterns in response to environmental conditions. We raised sea urchins (Strongylocentrotus franciscanus) under conditions simulating future atmospheric CO₂ levels of 540 and 970 ppm. When larvae raised under elevated CO₂ conditions were subjected to 1-h acute temperature stress, their ability to mount a physiological response (as measured by expression of the molecular chaperone hsp70) was reduced relative to those raised under ambient CO₂ conditions. These results represent the first use of gene expression assays to study the effects of OA on sea urchin development. They highlight the importance of looking at multiple environmental factors simultaneously as this approach may reveal previously unsuspected biological impacts of atmospheric changes.     

Future ocean acidification will be amplified by hypoxia in coastal habitats

Ocean acidification is elicited by anthropogenic carbon dioxide emissions and resulting oceanic uptake of excess CO₂ and might constitute an abiotic stressor powerful enough to alter marine ecosystem structures. For surface waters in gas-exchange equilibrium with the atmosphere, models suggest increases in CO₂ partial pressure (pCO₂) from current values of ca. 390 μatm to ca. 700–1,000 μatm by the end of the century. However, in typically unequilibrated coastal hypoxic regions, much higher pCO₂ values can be expected, as heterotrophic degradation of organic material is necessarily related to the production of CO₂ (i.e., dissolved inorganic carbon). Here, we provide data and estimates that, even under current conditions, maximum pCO₂ values of 1,700–3,200 μatm can easily be reached when all oxygen is consumed at salinities between 35 and 20, respectively. Due to the nonlinear nature of the carbonate system, the approximate doubling of seawater pCO₂ in surface waters due to ocean acidification will most strongly affect coastal hypoxic zones as pCO₂ during hypoxia will increase proportionally: we calculate maximum pCO₂ values of ca. 4,500 μatm at a salinity of 20 (T = 10 °C) and ca. 3,400 μatm at a salinity of 35 (T = 10 °C) when all oxygen is consumed. Upwelling processes can bring these CO₂-enriched waters in contact with shallow water ecosystems and may then affect species performance there as well. We conclude that (1) combined stressor experiments (pCO₂ and pO₂) are largely missing at the moment and that (2) coastal ocean acidification experimental designs need to be closely adjusted to carbonate system variability within the specific habitat. In general, the worldwide spread of coastal hypoxic zones also simultaneously is a spread of CO₂-enriched zones. The magnitude of expected changes in pCO₂ in these regions indicates that coastal systems may be more endangered by future global climate change than previously thought.     

Populations of the Sydney rock oyster, Saccostrea glomerata, vary in response to ocean acidification

Acidifying oceans are predicted to fundamentally alter marine ecosystems. Over the next century, acute studies suggest that the impacts of climate change on marine organisms and ecosystems may be catastrophic. To date, however, little is known about whether the response of marine organisms varies within a species and whether this provides a potential ??adaptive capacity??. Here, we show that selectively bred lines of the ecologically and economically important estuarine mollusc, the Sydney rock oyster Saccostrea glomerata, are more resilient to ocean acidification than the wild populations. When reared at elevated pCO₂, we found a 25% reduction in shell growth of the selectively bred population of the Sydney rock oyster, Saccostrea glomerata, compared to a 64% reduction in shell growth of wild populations. This study shows that there are significantly different sensitivities to ocean acidification even within the same species, providing preliminary evidence that selective breeding may be a solution for important aquaculture industries to overcome the future effects of ocean acidification.     

Metal contamination increases the sensitivity of larvae but not gametes to ocean acidification in the polychaete Pomatoceros lamarckii (Quatrefages)

Ocean acidification is not happening in isolation but against a background of chronic low-level pollution for most coastal marine environments. The reproductive and larval stages of marine invertebrates can be highly sensitive to the impacts of both environmental pollutants and ocean acidification, but very little is currently known regarding the potential impacts of combined contaminant and high CO₂ exposures on the health of marine organisms. Ocean acidification research to date has focused heavily on the responses of calcifying marine invertebrate larvae and algae, and as such the polychaetes as a group, despite their ecological importance, remain understudied. Here, we investigate the effects of elevated seawater CO₂ (pH range 8.1–7.4, plus an extreme pH of 7.2 in the sperm motility experiments), in combination with the environmental pollutant copper (0.002 μM), on the early life history stages of the intertidal polychaete Pomatoceros lamarckii from two populations. P. lamarckii sperm appear to be robust to elevated seawater CO₂. Whilst all three of the sperm motility end points measured showed a response to elevated CO₂, these responses were small and not linear. The percentage of motile sperm and sperm curvilinear velocity were significantly reduced in the lower pH treatments of 7.4 and 7.2, whereas sperm straight-line velocity (VSL) was mostly unaffected except for an increased VSL at pH 8.0. Fertilisation success was investigated using two populations from the South West (UK), one from Torquay and one from Plymouth Sound. Fertilisation success was slightly but significantly reduced at the 7.6 and 7.4 pH treatments for both populations (a 9.0 % reduction in fertilisation success from pH 8.1 to 7.4 for Torquay), but with a greater effect observed in the population from Plymouth Sound (a 13.33 % reduction in fertilisation success). No additional impact of 0.002 μM copper exposure on fertilisation success was found. Larval survival was found to be much more sensitive to elevated CO₂ than sperm function or fertilisation, and a significant interaction with copper exposure was observed. These results demonstrate the potential for polychaete larvae to be affected by predicted ocean acidification conditions and that chronic coastal pollutants, such as copper, have the potential to alter larval susceptibility to ocean acidification conditions.     

Impacts of ocean acidification on marine shelled molluscs

Over the next century, elevated quantities of atmospheric CO₂ are expected to penetrate into the oceans, causing a reduction in pH (?0.3/?0.4 pH unit in the surface ocean) and in the concentration of carbonate ions (so-called ocean acidification). Of growing concern are the impacts that this will have on marine and estuarine organisms and ecosystems. Marine shelled molluscs, which colonized a large latitudinal gradient and can be found from intertidal to deep-sea habitats, are economically and ecologically important species providing essential ecosystem services including habitat structure for benthic organisms, water purification and a food source for other organisms. The effects of ocean acidification on the growth and shell production by juvenile and adult shelled molluscs are variable among species and even within the same species, precluding the drawing of a general picture. This is, however, not the case for pteropods, with all species tested so far, being negatively impacted by ocean acidification. The blood of shelled molluscs may exhibit lower pH with consequences for several physiological processes (e.g. respiration, excretion, etc.) and, in some cases, increased mortality in the long term. While fertilization may remain unaffected by elevated pCO₂, embryonic and larval development will be highly sensitive with important reductions in size and decreased survival of larvae, increases in the number of abnormal larvae and an increase in the developmental time. There are big gaps in the current understanding of the biological consequences of an acidifying ocean on shelled molluscs. For instance, the natural variability of pH and the interactions of changes in the carbonate chemistry with changes in other environmental stressors such as increased temperature and changing salinity, the effects of species interactions, as well as the capacity of the organisms to acclimate and/or adapt to changing environmental conditions are poorly described.     

Enhancing synergies between action on ocean acidification and the post-2020 global biodiversity framework

Ocean acidification is a substantial emergent threat to marine biodiversity and the goods and services it provides. Although efforts to address ocean acidification have been taken under the Convention on Biological Diversity (CBD), a far greater potential to do so exists by finding synergies between biodiversity conservation efforts and ocean acidification action. The ongoing process to develop a post-2020 global biodiversity framework offers an opportunity to ensure that opportunities for addressing ocean acidification are capitalized on and not overlooked. I argue that to achieve this, the following are needed: a technical integration of ocean acidification across the targets to be included in the post-2020 framework and a reframing of the issue as a biodiversity problem so as to highlight the synergies between existing biodiversity work and action needed to address ocean acidification. Given that the post-2020 framework is intended to establish the global biodiversity agenda for the coming decades, integration of ocean acidification will set a precedent for the other biodiversity-related conventions and encourage greater uptake of the issue across the wider international community. My approach is of direct relevance to those participating in the negotiations, both from a CBD Party perspective and the perspective of those advocating for a strong outcome to protect marine biodiversity and marine socioecological systems. My discussion of framing is relevant to those working beyond the CBD within other biodiversity-related conventions in which goals to address ocean acidification are sorely lacking.     

Effects of near‐future ocean acidification,fishing, and marine protection on a temperate coastal ecosystem

Understanding ecosystem responses to global and local anthropogenic impacts is paramount to predicting future ecosystem states. We used an ecosystem modeling approach to investigate the independent and cumulative effects of fishing, marine protection, and ocean acidification on a coastal ecosystem. To quantify the effects of ocean acidification at the ecosystem level, we used information from the peer‐reviewed literature on the effects of ocean acidification. Using an Ecopath with Ecosim ecosystem model for the Wellington south coast, including the Taputeranga Marine Reserve (MR), New Zealand, we predicted ecosystem responses under 4 scenarios: ocean acidification + fishing; ocean acidification + MR (no fishing); no ocean acidification + fishing; no ocean acidification + MR for the year 2050. Fishing had a larger effect on trophic group biomasses and trophic structure than ocean acidification, whereas the effects of ocean acidification were only large in the absence of fishing. Mortality by fishing had large, negative effects on trophic group biomasses. These effects were similar regardless of the presence of ocean acidification. Ocean acidification was predicted to indirectly benefit certain species in the MR scenario. This was because lobster (Jasus edwardsii) only recovered to 58% of the MR biomass in the ocean acidification + MR scenario, a situation that benefited the trophic groups lobsters prey on. Most trophic groups responded antagonistically to the interactive effects of ocean acidification and marine protection (46%; reduced response); however, many groups responded synergistically (33%; amplified response). Conservation and fisheries management strategies need to account for the reduced recovery potential of some exploited species under ocean acidification, nonadditive interactions of multiple factors, and indirect responses of species to ocean acidification caused by declines in calcareous predators.     

Tough as a rock-boring urchin: adult Echinometra sp. EE from the Red Sea show high resistance to ocean acidification over long-term exposures

Ocean acidification, a process caused by the continuous rise of atmospheric CO₂ levels, is expected to have a profound impact on marine invertebrates. Findings of the numerous studies conducted in this field indicate high variability in species responses to future ocean conditions. This study aimed at understanding the effects of long-term exposure to elevated pCO₂ conditions on the performance of adult Echinometra sp. EE from the Gulf of Aqaba (Red Sea). During an 11-month incubation under high pCO₂ (1,433 μatm, pH_NBS 7.7) and control (435 μatm, pH_NBS 8.1) conditions, we examined the urchins’ somatic and gonadal growth, gametogenesis and skeletal microstructure. Somatic and gonadal growths were exhibited with no significant differences between the treatments. In addition, all urchins in the experiment completed a full reproductive cycle, typical of natural populations, with no detectable impact of increased pCO₂ on the timing, duration or progression of the cycle. Furthermore, scanning electron microscopy imaging of urchin tests and spines revealed no signs of the usual observed effects of acidosis, such as skeletal dissolution, widened stereom pores or non-smoothed structures. Our results, which yielded no significant impact of the high pCO₂ treatment on any of the examined processes in the urchins studied, suggest high resistance of adult Echinometra sp. EE to near future ocean acidification conditions. With respect to other findings in this area, the outcome of this study provides an example of the complicated and diverse responses of echinoids to the predicted environmental changes.     

The physiological and molecular responses of larvae from the reef-building coral Pocillopora damicornis exposed to near-future increases in temperature and pCO2

Given the threats of greenhouse gas emissions and a changing climate to marine ecosystems, there is an urgent need to better understand the response of not only adult corals, which are particularly sensitive to environmental changes, but also their larvae, whose mechanisms of acclimation to both temperature increases and ocean acidification are not well understood. Brooded larvae from the reef coral Pocillopora damicornis collected from Nanwan Bay, Southern Taiwan, were exposed to ambient or elevated temperature (25 or 29 °C) and pCO₂ (415 or 635 μatm) in a factorial experiment for 9 days, and a variety of physiological and molecular parameters were measured. Respiration and rubisco protein expression decreased in larvae exposed to elevated temperature, while those incubated at high pCO₂ were larger in size. Collectively, these findings highlight the complex metabolic and molecular responses of this life history stage and the need to integrate our understanding across multiple levels of biological organization. Our results also suggest that for this pocilloporid larval life stage, the impacts of elevated temperature are likely a greater threat under near-future predictions for climate change than ocean acidification.     

Spatio-temporal variability of polychaete colonization at volcanic CO2 vents indicates high tolerance to ocean acidification

Ocean acidification is predicted to have negative effects on marine biota, resulting in the loss of biodiversity and changes in marine ecosystem structure and function. However, some species and life stages may be capable of thriving in low pH conditions, either due to their natural ability to tolerate stressful low pH–high pCO₂ conditions and/or alteration of species interactions caused by changes in pH profiles, or due to evolutionary trade-offs. A better understanding of which species may be capable of tolerating ocean acidification can guide future research into the mechanisms for physiological and ecological resilience to future carbon dioxide (CO₂) conditions. We investigated the colonization of selected polychaete species along a pH gradient originating from shallow, coastal volcanic CO₂ vents (Ischia, Italy). Colonization was quantified by exposing artificial invertebrate collectors attached to the substratum for 30 days during different periods of the year (late spring, fall and late winter). Three species, Amphiglena mediterranea, Platynereis dumerilii and Syllis prolifera, were present and abundant along the gradient throughout the year. All three species were significantly more abundant in the most acidified areas, confirming their high tolerance and capacity to cope with very low pH. Abundances of all three species were compared to data previously collected via collectors suspended in the water column. More individuals were found in the collectors attached to the substratum, suggesting that abundances may have previously been underestimated. This is likely due to the close proximity of these collectors with the natural rocky substratum. All three species exhibited similar temporal variability, consistent with their life cycle and reproductive biology. Our results demonstrate high tolerance of the species for low and variable pH and corroborate their use as robust models to explore the capacity to cope with low pH–high pCO₂ conditions, both in the natural vent systems and in the laboratory.     

Can ocean acidification affect population dynamics of the barnacle Semibalanus balanoides at its southern range edge?

The global ocean and atmosphere are warming. There is increasing evidence suggesting that, in addition to other environmental factors, climate change is affecting species distributions and local population dynamics. Additionally, as a consequence of the growing levels of atmospheric carbon dioxide (CO2), the oceans are taking up increasing amounts of this CO2, causing ocean pH to decrease (ocean acidification). The relative impacts of ocean acidification on population dynamics have yet to be investigated, despite many studies indicating that there will be at least a sublethal impact on many marine organisms, particularly key calcifying organisms. Using empirical data, we forced a barnacle (Semibalanus balanoides) population model to investigate the relative influence of sea surface temperature (SST) and ocean acidification on a population nearing the southern limit of its geographic distribution. Hindcast models were compared to observational data from Cellar Beach (southwestern United Kingdom). Results indicate that a declining pH trend (-0.0017 unit/yr), indicative of ocean acidification over the past 50 years, does not cause an observable impact on the population abundance relative to changes caused by fluctuations in temperature. Below the critical temperature (here T(crit) = 13.1 degrees C), pH has a more significant affect on population dynamics at this southern range edge. However, above this value, SST has the overriding influence. At lower SST, a decrease in pH (according to the National Bureau of Standards, pHNBs) from 8.2 to 7.8 can significantly decrease the population abundance. The lethal impacts of ocean acidification observed in experiments on early life stages reduce cumulative survival by approximately 25%, which again will significantly alter the population level at this southern limit. Furthermore, forecast predictions from this model suggest that combined acidification and warming cause this local population to die out 10 years earlier than would occur if there was only global warming and no concomitant decrease in pH.     

Health and population-dependent effects of ocean acidification on the marine isopod Idotea balthica

Three populations of the grazing isopod Idotea balthica were exposed to high CO₂ treatment for a period of 20 days to investigate the effect of ocean acidification (OA) on animal health and immunocompetence. The results of the populations from more saline habitats were comparable and showed a 60–80 % decrease in immune response as a result of the high CO₂ treatment. Analysis of protein carbonyls showed no treatment effect, indicating that short-term OA does not increase oxidative protein damage. Meanwhile, the third tested population from the lower saline Baltic Sea had higher background protein carbonyl levels. Ocean acidification in addition to this resulted in 100 % mortality. The results of this study show that OA reduced immunocompetence of this marine isopod. In addition, populations and individuals in poor health are potentially at greater risk to succumb under OA.     

Optimum growth conditions as opposed to calcite saturation as a control on the calcification rate and shell-weight of marine foraminifera

Processes that impact negatively on CaCO₃ production and calcification rates, such as a lowering of the carbonate saturation state of the surface ocean in response to higher atmospheric CO₂ levels, can have disastrous consequences for marine ecosystems. This study, however, shows that on a global basis planktonic foraminiferal calcification rates (as inferred from shell-weight) are not related to calcite saturation state, as has been inferred from culture experiments and across a short latitudinal core-top transect. Rather, foraminiferal calcification is apparently a function of the same complex interplay of environmental parameters as assemblage abundance patterns. Highest foraminiferal shell-weights are observed within the optimum ecological niche of each planktonic species and no simple relationship with calcite saturation, temperature or surface nutrient levels exists. The implications of calcification-saturation relationships in calcifying marine ecosystems should therefore be evaluated in a broader context than has been the case in recent studies.Communicated by O. Kinne, Oldendorf/Luhe     

Ocean acidification induces budding in larval sea urchins

Ocean acidification (OA), the reduction of ocean pH due to hydration of atmospheric CO₂, is known to affect growth and survival of marine invertebrate larvae. Survival and transport of vulnerable planktonic larval stages play important roles in determining population dynamics and community structures in coastal ecosystems. Here, we show that larvae of the purple urchin, Strongylocentrotus purpuratus, underwent high-frequency budding (release of blastula-like particles) when exposed to elevated pCO₂ level (>700 μatm). Budding was observed in >50 % of the population and was synchronized over short periods of time (~24 h), suggesting this phenomenon may be previously overlooked. Although budding can be a mechanism through which larval echinoids asexually reproduce, here, the released buds did not develop into viable clones. OA-induced budding and the associated reduction in larval size suggest new hypotheses regarding physiological and ecological tradeoffs between short-term benefits (e.g. metabolic savings and predation escape) and long-term costs (e.g. tissue loss and delayed development) in the face of climate change.     

Ocean acidification reduces biomineralization-related gene expression in the sea urchin, Hemicentrotus pulcherrimus

Although recent studies have demonstrated that calcification in a wide range of marine organisms is profoundly affected by CO₂-induced ocean acidification, the mechanism of this phenomenon is still unclear. To clarify the effects of ocean acidification on the calcification process at the molecular level, we evaluated the expression of three biomineralization-related genes in the sea urchin Hemicentrotus pulcherrimus exposed under control, 1,000, and 2,000?ppm CO₂ from egg to pluteus larval stage. We found that the expression of the gene msp130, which is proposed to transport Ca²⁺ to the calcification site, is suppressed by increased CO₂ at pluteus larval stage. Meanwhile, expression of the spicule protein matrix genes SM30 and SM50 was apparently not affected. The results suggest that the combined effects of ocean acidification on the expression of skeletogenesis-related genes as well as the change in seawater carbonate chemistry affect the biomineralization ability of sea urchins.     

海洋酸化与重金属、有机污染物和人工纳米颗粒的联合毒性效应研究进展

由于大量人类活动的影响,大气CO_2浓度持续增加,其中约1/3被海洋吸收,导致表层海水pH值降低和碳酸盐平衡体系波动,即"海洋酸化"现象。污染物的海洋环境效应一直是全球环境科学领域研究的热点。在实际环境中,海洋酸化往往与污染物共同存在并作用于海洋生态系统,且海洋酸化极有可能改变污染物的海洋环境行为从而影响其毒性效应。但现有研究大多针对海洋酸化或者污染物单独作用下的毒性效应展开,对海洋酸化与污染物的联合毒性效应的研究不足、亟待加强。为此,综述了近年来海洋酸化与典型污染物(重金属、有机污染物)及新型污染物(人工纳米颗粒)的相关文献,重点阐述了海洋酸化对污染物环境行为的影响和海洋酸化与污染物对海洋生物的联合毒性效应,指出当前的研究不足,并对未来的研究方向进行了展望。     

Growth response of an early successional assemblage of coralline algae and benthic diatoms to ocean acidification

The sustained absorption of anthropogenically released atmospheric CO₂ by the oceans is modifying seawater carbonate chemistry, a process termed ocean acidification (OA). By the year 2100, the worst case scenario is a decline in the average oceanic surface seawater pH by 0.3 units to 7.75. The changing seawater carbonate chemistry is predicted to negatively affect many marine species, particularly calcifying organisms such as coralline algae, while species such as diatoms and fleshy seaweed are predicted to be little affected or may even benefit from OA. It has been hypothesized in previous work that the direct negative effects imposed on coralline algae, and the direct positive effects on fleshy seaweeds and diatoms under a future high CO₂ ocean could result in a reduced ability of corallines to compete with diatoms and fleshy seaweed for space in the future. In a 6-week laboratory experiment, we examined the effect of pH 7.60 (pH predicted to occur due to ocean acidification just beyond the year 2100) compared to pH 8.05 (present day) on the lateral growth rates of an early successional, cold-temperate species assemblage dominated by crustose coralline algae and benthic diatoms. Crustose coralline algae and benthic diatoms maintained positive growth rates in both pH treatments. The growth rates of coralline algae were three times lower at pH 7.60, and a non-significant decline in diatom growth meant that proportions of the two functional groups remained similar over the course of the experiment. Our results do not support our hypothesis that benthic diatoms will outcompete crustose coralline algae under future pH conditions. However, while crustose coralline algae were able to maintain their presence in this benthic rocky reef species assemblage, the reduced growth rates suggest that they will be less capable of recolonizing after disturbance events, which could result in reduced coralline cover under OA conditions.