Seaweed fails to prevent ocean acidification impact on foraminifera along a shallow‐water CO2 gradient

Ocean acidification causes biodiversity loss, alters ecosystems, and may impact food security, as shells of small organisms dissolve easily in corrosive waters. There is a suggestion that photosynthetic organisms could mitigate ocean acidification on a local scale, through seagrass protection or seaweed cultivation, as net ecosystem organic production raises the saturation state of calcium carbonate making seawater less corrosive. Here, we used a natural gradient in calcium carbonate saturation, caused by shallow‐water CO₂ seeps in the Mediterranean Sea, to assess whether seaweed that is resistant to acidification (Padina pavonica) could prevent adverse effects of acidification on epiphytic foraminifera. We found a reduction in the number of species of foraminifera as calcium carbonate saturation state fell and that the assemblage shifted from one dominated by calcareous species at reference sites (pH ~8.19) to one dominated by agglutinated foraminifera at elevated levels of CO₂ (pH ~7.71). It is expected that ocean acidification will result in changes in foraminiferal assemblage composition and agglutinated forms may become more prevalent. Although Padina did not prevent adverse effects of ocean acidification, high biomass stands of seagrass or seaweed farms might be more successful in protecting epiphytic foraminifera.     

Ocean acidification and the loss of phenolic substances in marine plants

Rising atmospheric CO(2) often triggers the production of plant phenolics, including many that serve as herbivore deterrents, digestion reducers, antimicrobials, or ultraviolet sunscreens. Such responses are predicted by popular models of plant defense, especially resource availability models which link carbon availability to phenolic biosynthesis. CO(2) availability is also increasing in the oceans, where anthropogenic emissions cause ocean acidification, decreasing seawater pH and shifting the carbonate system towards further CO(2) enrichment. Such conditions tend to increase seagrass productivity but may also increase rates of grazing on these marine plants. Here we show that high CO(2) / low pH conditions of OA decrease, rather than increase, concentrations of phenolic protective substances in seagrasses and eurysaline marine plants. We observed a loss of simple and polymeric phenolics in the seagrass Cymodocea nodosa near a volcanic CO(2) vent on the Island of Vulcano, Italy, where pH values decreased from 8.1 to 7.3 and pCO(2) concentrations increased ten-fold. We observed similar responses in two estuarine species, Ruppia maritima and Potamogeton perfoliatus, in in situ Free-Ocean-Carbon-Enrichment experiments conducted in tributaries of the Chesapeake Bay, USA. These responses are strikingly different than those exhibited by terrestrial plants. The loss of phenolic substances may explain the higher-than-usual rates of grazing observed near undersea CO(2) vents and suggests that ocean acidification may alter coastal carbon fluxes by affecting rates of decomposition, grazing, and disease. Our observations temper recent predictions that seagrasses would necessarily be "winners" in a high CO(2) world.     

Calcification, growth and mortality of juvenile clams Ruditapes decussatus under increased pCO2 and reduced pH: Variable responses to ocean acidification at local scales?

We investigated the effects of ocean acidification on juvenile clams Ruditapes decussatus (average shell length 10.24 mm) in a controlled CO₂ perturbation experiment. The carbonate chemistry of seawater was manipulated by diffusing pure CO₂, to attain two reduced pH levels (by −0.4 and −0.7 pH units), which were compared to unmanipulated seawater. After 75 days we found no differences among pH treatments in terms of net calcification, size or weight of the clams. The naturally elevated total alkalinity of local seawater probably contributed to buffer the effects of increased pCO₂ and reduced pH. Marine organisms may, therefore, show diverse responses to ocean acidification at local scales, particularly in coastal, estuarine and transitional waters, where the physical-chemical characteristics of seawater are most variable. Mortality was significantly reduced in the acidified treatments. This trend was probably related to the occurrence of spontaneous spawning events in the control and intermediate acidification treatments. Spawning, which was unexpected due to the small size of the clams, was not observed for the pH −0.7 treatment, suggesting that the increased survival under acidified conditions may have been associated with a delay in the reproductive cycle of the clams. Future research about the impacts of ocean acidification on marine biodiversity should be extended to other types of biological and ecological processes, apart from biological calcification.     

海洋酸化效应对海水鱼类的综合影响评述

人类活动引起的大气CO2浓度的升高,除了使全球温度升高外,导致的另一个严重生态问题——海洋酸化(Ocean acidification,OA),受到社会各界包括科研界的高度重视,该领域的大部分研究结果都是在近十年才发表出来的,目前还有很多需要解决的问题。海洋酸化的研究涉及到很多学科的交叉包括化学、古生物学、生态学、生物地球化学等等。在生物学领域,海洋酸化主要围绕敏感物种,例如由碳酸钙形成贝壳或外骨骼的贝类,珊瑚礁群体等。鱼类作为海洋脊椎动物的代表生物类群,自身具有一定的酸碱平衡调节能力,但相关海洋酸化方向的研究并不是很多。尽管人们对于海洋酸化对鱼类的影响了解甚少,这并不说明海洋酸化对鱼类没有作用或者效应小,在相关研究逐步展开的同时,发现鱼类同样受到海洋酸化的危害,几乎涉及到鱼类整个生活史和几乎大部分生理过程,尤其是早期生活史的高度敏感。因此就目前国内外对此领域研究结果做综述,以期待业界同行能够对海水鱼类这个大的类群引起重视。     

Effects of CO(2) enrichment on photosynthesis, growth, and biochemical composition of seagrass Thalassia hemprichii (Ehrenb.) Aschers

The effects of CO2 enrichment on various ecophysiological parameters of tropical seagrass Thalassia hemprichii(Ehrenb.)Aschers were tested.T.hemprichii,collected from a seagrass bed in Xincun Bay,Hainan island of Southern China,was cultured at 4 CO2(aq)concentrations in flow-through seawater aquaria bubbled with CO2.CO2 enrichment considerably enhanced the relative maximum electron transport rate(RETRmax)and minimum saturating irradiance(Ek)of T.hemprichii.Leaf growth rate of CO2enriched plants was significantly higher than that in unenriched treatment.Nonstructural carbohydrates(NSC)of T.hemprichii,especially in belowground tissues,increased strongly with elevated CO2(aq),suggesting a translocation of photosynthate from aboveground to belowground tissues.Carbon content in belowground tissues showed a similar response with NSC,while in aboveground tissues,carbon content was not affected by CO2 treatments.In contrast,with increasing CO2(aq),nitrogen content in aboveground tissues markedly decreased,but nitrogen content in belowground was nearly constant.Carbon: nitrogen ratio in both tissues were obviously enhanced by increasing CO2(aq).Thus,these results indicate that T.hemprichii may respond positively to CO2-induced acidification of the coastal ocean.Moreover,the CO2-stimulated improvement of photosynthesis and NSC content may partially offset negative effects of severe environmental disturbance such as underwater light reduction.     

Effects of CO2 Enrichment on Photosynthesis, Growth, and Biochemical Composition of Seagrass Thalassia hemprichii (Ehrenb.) Aschers

The effects of CO2 enrichment on various ecophysiological parameters of tropical seagrass Thalassia hemprichii(Ehrenb.)Aschers were tested.T.hemprichii,collected from a seagrass bed in Xincun Bay,Hainan island of Southern China,was cultured at 4 CO2(aq)concentrations in flow-through seawater aquaria bubbled with CO2.CO2 enrichment considerably enhanced the relative maximum electron transport rate(RETRmax)and minimum saturating irradiance(Ek)of T.hemprichii.Leaf growth rate of CO2enriched plants was significantly higher than that in unenriched treatment.Nonstructural carbohydrates(NSC)of T.hemprichii,especially in belowground tissues,increased strongly with elevated CO2(aq),suggesting a translocation of photosynthate from aboveground to belowground tissues.Carbon content in belowground tissues showed a similar response with NSC,while in aboveground tissues,carbon content was not affected by CO2 treatments.In contrast,with increasing CO2(aq),nitrogen content in aboveground tissues markedly decreased,but nitrogen content in belowground was nearly constant.Carbon: nitrogen ratio in both tissues were obviously enhanced by increasing CO2(aq).Thus,these results indicate that T.hemprichii may respond positively to CO2-induced acidification of the coastal ocean.Moreover,the CO2-stimulated improvement of photosynthesis and NSC content may partially offset negative effects of severe environmental disturbance such as underwater light reduction.     

Slow‐flow habitats as refugia for coastal calcifiers from ocean acidification

The pH of the oceans’ surface water is dropping, termed ocean acidification (OA), and the 0.4 unit reduction in pH by 2100 is projected to negatively impact benthic coastal organisms that produce calcium carbonate “skeletons.” Research has focussed on identifying species that are susceptible to OA, but there is an urgent need to discover refuge habitats that will afford protection to vulnerable species. The susceptibility of calcium carbonate skeletons to dissolution by OA depends on the pH at their surface, and this is controlled by the interaction between seawater velocity and organismal metabolism. This perspective considers how seawater velocity modifies the responses of calcifying organisms (seaweed, shellfish, and tropical corals) to OA through its action on controlling diffusion boundary layer thickness and thereby the pH and calcium carbonate saturation state (Ω) at the organisms’ surface. Evidence is presented to support the idea that slow‐flow habitats, such as wave‐sheltered bays or the within canopies of seaweed/seagrass beds, might provide inexpensive refugia from OA for vulnerable coastal calcifiers.     

海洋酸化对珊瑚礁生态系统的影响研究进展

目前,大气CO2浓度的升高已导致海水pH值比工业革命前下降了约0.1,海水碳酸盐平衡体系随之变化,进而影响珊瑚礁生态系统的健康。近年来的研究表明海洋酸化导致造礁石珊瑚幼体补充和群落恢复更加困难,造礁石珊瑚和其它造礁生物(Reef-building organisms)钙化率降低甚至溶解,乃至影响珊瑚礁鱼类的生命活动。虽然海洋酸化对造礁石珊瑚光合作用的影响不显著,但珊瑚-虫黄藻共生体系会受到一定影响。建议选择典型海区进行长期系统监测,结合室内与原位模拟试验,从个体、种群、群落到系统不同层面,运用生理学和分子生物学技术,结合生态学研究手段,综合研究珊瑚的相应响应,以期深入认识海洋酸化对珊瑚礁生态系统健康(例如珊瑚白化)的影响及其效应。     

Impact of Ocean Warming and Ocean Acidification on Larval Development and Calcification in the Sea Urchin Tripneustes gratilla

Background

As the oceans simultaneously warm, acidify and increase in P _CO2, prospects for marine biota are of concern. Calcifying species may find it difficult to produce their skeleton because ocean acidification decreases calcium carbonate saturation and accompanying hypercapnia suppresses metabolism. However, this may be buffered by enhanced growth and metabolism due to warming.

Methodology/Principal Findings

We examined the interactive effects of near-future ocean warming and increased acidification/P _CO2 on larval development in the tropical sea urchin Tripneustes gratilla. Larvae were reared in multifactorial experiments in flow-through conditions in all combinations of three temperature and three pH/P _CO2 treatments. Experiments were placed in the setting of projected near future conditions for SE Australia, a global change hot spot. Increased acidity/P _CO2 and decreased carbonate mineral saturation significantly reduced larval growth resulting in decreased skeletal length. Increased temperature (+3°C) stimulated growth, producing significantly bigger larvae across all pH/P _CO2 treatments up to a thermal threshold (+6°C). Increased acidity (-0.3-0.5 pH units) and hypercapnia significantly reduced larval calcification. A +3°C warming diminished the negative effects of acidification and hypercapnia on larval growth.

Conclusions and Significance

This study of the effects of ocean warming and CO₂ driven acidification on development and calcification of marine invertebrate larvae reared in experimental conditions from the outset of development (fertilization) shows the positive and negative effects of these stressors. In simultaneous exposure to stressors the dwarfing effects of acidification were dominant. Reduction in size of sea urchin larvae in a high P _CO2 ocean would likely impair their performance with negative consequent effects for benthic adult populations.     

The role of coccolithophore calcification in bioengineering their environment

Coccolithophorids are enigmatic plankton that produce calcium carbonate coccoliths, which over geological time have buried atmospheric CO₂ into limestone, changing both the atmosphere and geology of the Earth. However, the role of coccoliths for the proliferation of these organisms remains unclear; suggestions include roles in anti-predation, enhanced photosynthesis and sun-screening. Here we test the hypothesis that calcification stabilizes the pH of the seawater proximate to the organisms, providing a level of acidification countering the detrimental basification that occurs during net photosynthesis. Such bioengineering provides a more stable pH environment for growth and fits the empirical evidence for changes in rates of calcification under different environmental conditions. Under this scenario, simulations suggest that the optimal production ratio of inorganic to organic particulate C (PIC : POC_prod) will be lower (by approx. 20%) with ocean acidification and that overproduction of coccoliths in a future acidified ocean, where pH buffering is weaker, presents a risk to calcifying cells.     

Detrimental effects of ocean acidification on the economically important Mediterranean red coral (Corallium rubrum)

The mean predicted decrease of 0.3–0.4 pH units in the global surface ocean by the end of the century has prompted urgent research to assess the potential effects of ocean acidification on the marine environment, with strong emphasis on calcifying organisms. Among them, the Mediterranean red coral (Corallium rubrum) is expected to be particularly susceptible to acidification effects, due to the elevated solubility of its Mg‐calcite skeleton. This, together with the large overexploitation of this species, depicts a bleak future for this organism over the next decades. In this study, we evaluated the effects of low pH on the species from aquaria experiments. Several colonies of C. rubrum were long‐term maintained for 314 days in aquaria at two different pH levels (8.10 and 7.81, pH_T). Calcification rate, spicule morphology, major biochemical constituents (protein, carbohydrates and lipids) and fatty acids composition were measured periodically. Exposure to lower pH conditions caused a significant decrease in the skeletal growth rate in comparison with the control treatment. Similarly, the spicule morphology clearly differed between both treatments at the end of the experiment, with aberrant shapes being observed only under the acidified conditions. On the other hand, while total organic matter was significantly higher under low pH conditions, no significant differences were detected between treatments regarding total carbohydrate, lipid, protein and fatty acid composition. However, the lower variability found among samples maintained in acidified conditions relative to controls, suggests a possible effect of pH decrease on the metabolism of the colonies. Our results show, for the first time, evidence of detrimental ocean acidification effects on this valuable and endangered coral species.     

Effects of variations in carbonate chemistry on the calcification rates of Madracis auretenra (= Madracis mirabilis sensu Wells, 1973): bicarbonate concentrations best predict calcification rates

Physiological data and models of coral calcification indicate that corals utilize a combination of seawater bicarbonate and (mainly) respiratory CO₂ for calcification, not seawater carbonate. However, a number of investigators are attributing observed negative effects of experimental seawater acidification by CO₂ or hydrochloric acid additions to a reduction in seawater carbonate ion concentration and thus aragonite saturation state. Thus, there is a discrepancy between the physiological and geochemical views of coral biomineralization. Furthermore, not all calcifying organisms respond negatively to decreased pH or saturation state. Together, these discrepancies suggest that other physiological mechanisms, such as a direct effect of reduced pH on calcium or bicarbonate ion transport and/or variable ability to regulate internal pH, are responsible for the variability in reported experimental effects of acidification on calcification. To distinguish the effects of pH, carbonate concentration and bicarbonate concentration on coral calcification, incubations were performed with the coral Madracis auretenra (= Madracis mirabilis sensu Wells, 1973) in modified seawater chemistries. Carbonate parameters were manipulated to isolate the effects of each parameter more effectively than in previous studies, with a total of six different chemistries. Among treatment differences were highly significant. The corals responded strongly to variation in bicarbonate concentration, but not consistently to carbonate concentration, aragonite saturation state or pH. Corals calcified at normal or elevated rates under low pH (7.6–7.8) when the seawater bicarbonate concentrations were above 1800 μm . Conversely, corals incubated at normal pH had low calcification rates if the bicarbonate concentration was lowered. These results demonstrate that coral responses to ocean acidification are more diverse than currently thought, and question the reliability of using carbonate concentration or aragonite saturation state as the sole predictor of the effects of ocean acidification on coral calcification.     

Sea urchin Arbacia dufresnei (Blainville 1825) larvae response to ocean acidification

Increased atmospheric CO₂ emissions are inducing changes in seawater carbon chemistry, lowering its pH, decreasing carbonate ion availability and reducing calcium carbonate saturation state. This phenomenon, known as ocean acidification, is happening at a faster rate in cold regions, i.e., polar and sub-polar waters. The larval development of Arbacia dufresnei from a sub-Antarctic population was studied at high (8.0), medium (7.7) and low (7.4) pH waters. The results show that the offspring from sub-Antarctic populations of A. dufresnei are susceptible to a development delay at low pH, with no significant increase in abnormal forms. Larvae were isometric between pH treatments. Even at calcium carbonate (CaCO₃) saturation states (of both calcite and aragonite, used as proxies of the magnesium calcite) <1, skeleton deposition occurred. Polar and sub-polar sea urchin larvae can show a certain degree of resilience to acidification, also emphasizing A. dufresnei potential to poleward migrate and further colonize southern regions.     

海洋酸化生态学研究进展

工业革命以来,人类排放的大量二氧化碳引起温室效应的同时,也被海洋吸收使得全球海洋出现了严重的酸化。海洋酸化及伴随的海水碳酸盐化学体系的变化对海洋生物产生深远的影响。以海洋酸化对钙化作用和光合作用的影响为重点,总结了近年来关于海洋酸化的研究,介绍了海洋中不同生态系统对海洋酸化的响应。一方面,海水中CO23-浓度和碳酸钙饱和度的降低对海洋钙化生物造成严重损害,生活在高纬的冷水珊瑚和翼足目等文石生产者是最早的受害者;贝类和棘皮动物在钙化早期对海洋酸化尤其敏感,其幼体存活率受到海洋酸化的严重制约。另一方面,CO2浓度的增加能促进海洋植物的光合作用和生长,增加初级生产力,改变浮游植物的群落组成。此外,海洋酸化可以促进固氮和脱氮作用同时削弱硝化作用,改变溶氧浓度分布和金属的生物可利用性,从而对海洋生物产生间接影响。海洋酸化对海洋生态系统的影响机制复杂,影响程度深远。为了能准确的评估海洋酸化的生态学效应,需要更全面深入的研究。     

Similar controls on calcification under ocean acidification across unrelated coral reef taxa

Ocean acidification (OA) is a major threat to marine ecosystems, particularly coral reefs which are heavily reliant on calcareous species. OA decreases seawater pH and calcium carbonate saturation state (Ω), and increases the concentration of dissolved inorganic carbon (DIC). Intense scientific effort has attempted to determine the mechanisms via which ocean acidification (OA) influences calcification, led by early hypotheses that calcium carbonate saturation state (Ω) is the main driver. We grew corals and coralline algae for 8–21 weeks, under treatments where the seawater parameters Ω, pH, and DIC were manipulated to examine their differential effects on calcification rates and calcifying fluid chemistry (Ω_cf, pH_cf, and DIC_cf). Here, using long duration experiments, we provide geochemical evidence that differing physiological controls on carbonate chemistry at the site of calcification, rather than seawater Ω, are the main determinants of calcification. We found that changes in seawater pH and DIC rather than Ω had the greatest effects on calcification and calcifying fluid chemistry, though the effects of seawater carbonate chemistry were limited. Our results demonstrate the capacity of organisms from taxa with vastly different calcification mechanisms to regulate their internal chemistry under extreme chemical conditions. These findings provide an explanation for the resistance of some species to OA, while also demonstrating how changes in seawater DIC and pH under OA influence calcification of key coral reef taxa.     

Ocean acidification and temperature increase impact mussel shell shape and thickness: problematic for protection?

Ocean acidification threatens organisms that produce calcium carbonate shells by potentially generating an under‐saturated carbonate environment. Resultant reduced calcification and growth, and subsequent dissolution of exoskeletons, would raise concerns over the ability of the shell to provide protection for the marine organism under ocean acidification and increased temperatures. We examined the impact of combined ocean acidification and temperature increase on shell formation of the economically important edible mussel Mytilus edulis. Shell growth and thickness along with a shell thickness index and shape analysis were determined. The ability of M. edulis to produce a functional protective shell after 9 months of experimental culture under ocean acidification and increasing temperatures (380, 550, 750, 1000 μatm pCO ₂, and 750, 1000 μatm pCO ₂ + 2°C) was assessed. Mussel shells grown under ocean acidification conditions displayed significant reductions in shell aragonite thickness, shell thickness index, and changes to shell shape (750, 1000 μatm pCO ₂) compared to those shells grown under ambient conditions (380 μatm pCO ₂). Ocean acidification resulted in rounder, flatter mussel shells with thinner aragonite layers likely to be more vulnerable to fracture under changing environments and predation. The changes in shape presented here could present a compensatory mechanism to enhance protection against predators and changing environments under ocean acidification when mussels are unable to grow thicker shells. Here, we present the first assessment of mussel shell shape to determine implications for functional protection under ocean acidification.     

Persistent natural acidification drives major distribution shifts in marine benthic ecosystems

Ocean acidification is receiving increasing attention because of its potential to affect marine ecosystems. Rare CO₂ vents offer a unique opportunity to investigate the response of benthic ecosystems to acidification. However, the benthic habitats investigated so far are mainly found at very shallow water (less than or equal to 5 m depth) and therefore are not representative of the broad range of continental shelf habitats. Here, we show that a decrease from pH 8.1 to 7.9 observed in a CO₂ vent system at 40 m depth leads to a dramatic shift in highly diverse and structurally complex habitats. Forests of the kelp Laminaria rodriguezii usually found at larger depths (greater than 65 m) replace the otherwise dominant habitats (i.e. coralligenous outcrops and rhodolith beds), which are mainly characterized by calcifying organisms. Only the aragonite-calcifying algae are able to survive in acidified waters, while high-magnesium-calcite organisms are almost completely absent. Although a long-term survey of the venting area would be necessary to fully understand the effects of the variability of pH and other carbonate parameters over the structure and functioning of the investigated mesophotic habitats, our results suggest that in addition of significant changes at species level, moderate ocean acidification may entail major shifts in the distribution and dominance of key benthic ecosystems at regional scale, which could have broad ecological and socio-economic implications.     

Effects of seawater acidification on Diopatra neapolitana (Polychaete,Onuphidae): Biochemical and regenerative capacity responses

Changes in atmospheric CO₂ concentrations and, consequently, ocean acidification, together with changes in seawater chemistry, are likely to have a large impact on marine ecosystems. Much of the current research concerning the consequences of ocean acidification has been limited to organisms dependent on the availability of carbonate ions in seawater, especially bivalves. Nevertheless, many marine organisms do not rely on calcium carbonate structures, such as most of the polychaete species that might also be affected by seawater acidification. However the effects of ocean acidification on these organisms are almost unknown. Thus, in the present study, the effects of pH decrease were studied in the polychaete Diopatra neapolitana, using tissue regenerative capacity and biochemical alterations as biomarkers. The results obtained revealed that individuals exposed for 28 days to low pH levels (7.5, 7.3, 7.1) exhibited lower capacity to regenerate their body, in comparison to control organisms (pH 7.8). This study also evidenced that seawater acidification induced oxidative stress in D. neapolitana, with individuals under lower pH levels showing higher LPO, lower GSH/GSSG values and higher enzymatic activity (CAT, SOD and GSTs). Additionally, organisms exposed to low pH presented significantly lower glycogen and protein content. Thus, the present work revealed the effects of pH on the polychaete species D. neapolitana, both at physiological and biochemical levels. Furthermore, our results validate the use of D. neapolitana as a test organism in laboratory-based bioassays, and also as an adequate bioindicator species to pH decrease in the environment. Finally, the regenerative capacity appears to be a promising endpoint to evaluate the effects of pH on D. neapolitana.     

Changes in coral microbial communities in response to a natural pH gradient

Surface seawater pH is currently 0.1 units lower than pre-industrial values and is projected to decrease by up to 0.4 units by the end of the century. This acidification has the potential to cause significant perturbations to the physiology of ocean organisms, particularly those such as corals that build their skeletons/shells from calcium carbonate. Reduced ocean pH could also have an impact on the coral microbial community, and thus may affect coral physiology and health. Most of the studies to date have examined the impact of ocean acidification on corals and/or associated microbiota under controlled laboratory conditions. Here we report the first study that examines the changes in coral microbial communities in response to a natural pH gradient (mean pH_T 7.3–8.1) caused by volcanic CO₂ vents off Ischia, Gulf of Naples, Italy. Two Mediterranean coral species, Balanophyllia europaea and Cladocora caespitosa, were examined. The microbial community diversity and the physiological parameters of the endosymbiotic dinoflagellates (Symbiodinium spp.) were monitored. We found that pH did not have a significant impact on the composition of associated microbial communities in both coral species. In contrast to some earlier studies, we found that corals present at the lower pH sites exhibited only minor physiological changes and no microbial pathogens were detected. Together, these results provide new insights into the impact of ocean acidification on the coral holobiont.     

TESTING THE EFFECTS OF OCEAN ACIDIFICATION ON ALGAL METABOLISM: CONSIDERATIONS FOR EXPERIMENTAL DESIGNS1

Ocean acidification describes changes in the carbonate chemistry of the ocean due to the increased absorption of anthropogenically released CO₂. Experiments to elucidate the biological effects of ocean acidification on algae are not straightforward because when pH is altered, the carbon speciation in seawater is altered, which has implications for photosynthesis and, for calcifying algae, calcification. Furthermore, photosynthesis, respiration, and calcification will themselves alter the pH of the seawater medium. In this review, algal physiologists and seawater carbonate chemists combine their knowledge to provide the fundamental information on carbon physiology and seawater carbonate chemistry required to comprehend the complexities of how ocean acidification might affect algae metabolism. A wide range in responses of algae to ocean acidification has been observed, which may be explained by differences in algal physiology, timescales of the responses measured, study duration, and the method employed to alter pH. Two methods have been widely used in a range of experimental systems: CO₂ bubbling and HCl/NaOH additions. These methods affect the speciation of carbonate ions in the culture medium differently; we discuss how this could influence the biological responses of algae and suggest a third method based on HCl/NaHCO₃ additions. We then discuss eight key points that should be considered prior to setting up experiments, including which method of manipulating pH to choose, monitoring during experiments, techniques for adding acidified seawater, biological side effects, and other environmental factors. Finally, we consider incubation timescales and prior conditioning of algae in terms of regulation, acclimation, and adaptation to ocean acidification.