Investigating the Impact of Ocean Acidification on Anti-Stress Mechanisms in Sepia esculenta Larvae Based on Transcriptome Profiling

With the rapid development of oil,energy,power and other industries,CO_(2) emissions rise sharply,which will cause a large amount of CO_(2) in the air be absorbed by the ocean and lead to ocean acidification.The growth and development of organisms can be seriously affected by acidified seawater.Sepia esculenta is a mollusk with high nutritional and economic value and is widely cultured in offshore waters of China.Larvae are the early life forms of the organism and are more vulnerable to changes in the external environment.Too low pH will lead to some adverse reactions in larvae,which will affect metabolism,immune response and other life activities.In this study,we sequenced the transcriptome of S.esculenta subjected to acidified seawater stress and identified 1072differentially expressed genes(DEGs).The detected atypical expression of DEGs substantiates cellular malformation and translocation in S.esculenta under low pH stimulation.Simultaneously,this also substantiates the notable impact of ocean acidification on mollusks.These DEGs were used for functional enrichment analysis of GO and KEGG,and the top twenty items of the biological process classification in GO terms and 11 KEGG signaling pathways were significantly enriched.Finally,the constructed proteinprotein interaction network(PPI)was used to analyze protein-protein interactions,and 12 key DEGs and 3 hub genes were identified.The reliability of 12 genes was verified by quantitative RT-PCR.A comprehensive analysis of the KEGG signaling pathway and PPI revealed that ocean acidification leads to abnormalities in lipid metabolism in S.esculenta larvae,which can lead to cancer development and metastasis,accompanied by some degree of inflammation.The results of the study will help to further investigate the physiological processes of S.esculenta when stimulated by ocean acidification,and provide a reference to cope with the captive breeding of S.esculenta affected by acidification.

Effects of Ocean Acidification on Nitrogen Metabolism of Skeletonema costatum

Ocean acidification(OA),caused by the rising concentration of atmospheric CO_(2),leads to changes in the marine carbonate system.This,in turn,affects the physiological processes of phytoplankton.In response to increased pCO_(2) levels,marine microalgae modulate their physiological responses to meet their energy and metabolic requirements.Nitrogen metabolism is a critical metabolic pathway,directly affecting the growth and reproductive capacity of marine microorganisms.Understanding the molecular mechanisms that regulate nitrogen metabolism in microalgae under OA conditions is therefore crucial.This study aimed to investi-gate how OA affects the expression profiles of key genes in the nitrogen metabolic pathway of the marine diatom Skeletonema costatum.Our findings indicate that OA upregulates key genes involved in the nitrogen metabolic pathway,specifically those related to nitrate assimilation and glutamate metabolism.Moreover,pCO_(2) has been identified as the predominant factor affecting the expression of these genes,with a more significant impact than pH variations in S.costatum.This research not only advances our understanding of the adaptive mechanisms of S.costatum in response to OA but also provides essential data for predicting the ecological consequences of OA on marine diatoms.

Effects of ocean acidification on the metabolic rates of three species of bivalve from southern coast of China

Oceanic uptake of anthropogenic carbon dioxide results in a decrease in seawater pH, a process known as "ocean acidification". The pearl oyster Pinctada fucata, the noble scallop Chlamys nobilis, and the green-lipped mussel Perna viridis are species of economic and ecological importance along the southern coast of China. We evaluated the effects of seawater acidification on clearance, respiration, and excretion rates in these three species. The animals were reared in seawater at pH 8.1 (control), 7.7, or 7.4. The clearance rate was highest at pH 7.7 for P. fucata and at pH 8.1 for C. nobilis and P. viridis. The pH had little effecton the respiration rate of P. fucata and P. viridis. In contrast, the respiration rate was significantly lower atpH 7.4 in C. nobilis. The excretion rate was significantly lower at pH 7.4 than pH 8.1 for all species. Theresults indicate that the reduction in seawater pH likely affected the metabolic process (food intake, oxygenconsumption, and ammonia excretion) of these bivalves. Different species respond differently to seawateracidification. Further studies are needed to demonstrate the exact mechanisms for this effect and evaluateadaptability of these bivalves to future acidified oceans.

Nutrient Enrichment Regulates the Growth and Physiological Responses of Saccharina japonica to Ocean Acidification

Environmental changes,such as ocean acidification and eutrophication,have created threats to kelp mariculture.In this study,the growth,photosynthesis,respiration and nutrient composition of Saccharina japonica were evaluated at different levels of pCO2(400 and 800μL L−1)and nutrients(nutrient-enriched and non-enriched seawater).Elevated pCO2 decreased the relative growth rate(RGR),net photosynthetic rate and contents of tissue carbon and tissue nitrogen under non-enriched nutrient conditions,but it had no significant effect on these parameters under nutrient-enriched conditions.The dark respiration rate was positively affected by elevated pCO2 regardless of the nutrient conditions.However,the C:N was unaffected by elevated pCO2 at both nutrient levels.These results implied that ocean acidification could reduce the production and nutrient contents in the tissues of S.japonica,which was associated with nutrient conditions.

The Brown Algae Saccharina japonica and Sargassum horneri Exhibit Species-Specific Responses to Synergistic Stress of Ocean Acidification and Eutrophication

Ocean acidification and eutrophication are two important environmental stressors.They inevitably impact marine macroalgae,and hence the coastal ecosystem of China.Saccharina japonica,as the main culture species in China,is suffering the harmful golden tide caused by Sargassum horneri.However,it remains unclear whether the detrimental effects of S.horneri on S.japonica cultivation become more severe in future acidified and eutrophic scenario.In this study,we respectively investigated the effects of pCO_(2)(400μatm and 1000μatm)and nutrients(non-enriched and enriched seawater)on the growth,photosynthesis,respiration,chlorophyll contents,and tissue nitrogen of S.japonica and S.horneri.Results indicated that enrichment of nutrients contributed S.horneri to utilize HCO_(3)^(−).The carbon acquisition pathway shifted from HCO_(3)^(−)to CO_(2) in S.japonica,while S.horneri re-mained using HCO_(3)^(−)regulated by nutrient enrichment.S.horneri exhibited better photosynthetic traits than S.japonica,with a higher level of net photosynthetic rate and chlorophyll contents at elevated pCO_(2) and enriched nutrients.Tissue nitrogen also accumulated richly in the thalli of S.horneri under higher pCO_(2) and nutrients.Significant enhancement in growth was only detected in S.horneri under synergistic stress.Together,S.horneri showed competitive dominance in current study.These findings suggest that increasing risk of golden tide in acidified and eutrophic ocean can most likely result in great damage to S.japonica cultivation.

Simulation of global ocean acidification and chemical habitats of shallow-and cold-water coral reefs

Using the UVic Earth System Model, this study simulated the change of seawater chemistry and analyzed the chemical habitat surrounding shallow- and cold-water coral reefs from the year 1800 to 2300 employing RCP2.6, RCP4.5, RCP6.0, and RCP8.5 scenarios. The model results showed that the global ocean will continue to absorb atmospheric CO2. Global mean surface ocean temperature will rise 1.1-2.8 K at the end of the 21st century across RCP scenarios. Meanwhile, the global mean surface ocean pH will drop 0.14--0.42 and the ocean surface mean con- centration of carbonate will decrease 20%--51% across the RCP scenarios. The saturated state of sea water with respect to calcite carbonate minerals （t2） will decrease rapidly. During the pre-industrial period, 99% of the shallow-water coral reefs were surrounded by seawater with t2 〉 3.5 and 87% of the deep-sea coral reefs were surrounded by seawater with aragonite supersaturation. Within the 21st century, except for the high mitigation scenario of RCP2.6, almost none shallow-water coral reefs will be surrounded by seawater with g2 〉 3.5. Under the intensive emission scenario of RCP8.5, by the year 2100, the aragonite saturation horizon will rise to 308 m under the sea surface from 1138 m at the pre- industrial period, thus 73% of the cold-water coral reefs will be surrounded by seawater with aragonite undersaturation. By the year 2300, only 5% of the cold-water coral reefs will be surrounded by seawater with aragonite supersaturation.

The potential of ocean acidification on suppressing larval development in the Pacific oyster Crassostrea gigas and blood cockle Area inflata Reeve

We evaluated the effect of pH on larval development in larval Pacific oyster(Crassostrea gigas) and blood cockle(Arca inflata Reeve).The larvae were reared at pH 8.2(control),7.9,7.6,or 7.3beginning 30 min or 24 h post fertilization.Exposure to lower pH during early embryonic development inhibited larval shell formation in both species.Compared with the control,larvae took longer to reach the D-veliger stage when reared under pH 7.6 and 7.3.Exposure to lower pH immediately after fertilization resulted in significantly delayed shell formation in the Pacific oyster larvae at pH 7.3 and blood cockle larvae at pH 7.6 and 7.3.However,when exposure was delayed until 24 h post fertilization,shell formation was only inhibited in blood cockle larvae reared at pH 7.3.Thus,the early embryonic stages were more sensitive to acidified conditions.Our results suggest that ocean acidification will have an adverse effect on embryonic development in bivalves.Although the effects appear subtle,they may accumulate and lead to subsequent issues during later larval development.

An ocean acidification-simulated system and its application in coral physiological studies

Due to the elevated atmospheric carbon dioxide, ocean acidification(OA) has recently emerged as a research theme in marine biology due to an expected deleterious effect of altered seawater chemistry on calcification. A system simulating future OA scenario is crucial for OA-related studies. Here, we designed an OA-simulated system(OASys) with three solenoid-controlled CO_2 gas channels. The OASys can adjust the pH of the seawater by bubbling CO_2 gas into seawaters via feedback systems. The OASys is very simple in structure with an integrated design and is new-user friendly with the instruction. Moreover, the OASys can monitor and record real-time pH values and can maintain pH levels within 0.02 pH unit. In a 15-d experiment, the OASys was applied to simulate OA in which the expected target pH values were 8.00, 7.80 and 7.60 to study the calcifying response of Galaxea fascicularis. The results showed daily mean seawater pH values held at pH 8.00±0.01, 7.80±0.01 and 7.61±0.01 over15 d. Correspondingly, the coral calcification of G. fascicularis gradually decreased with reduced pH.

The ecological response of natural phytoplankton population and related metabolic rates to future ocean acidification

Ocean acidifi cation(OA)and global warming-induced water column stratification can signifi cantly alter phytoplankton-related biological activity in the marine ecosystem.Yet how these changes may play out in the tropical Indian Ocean remains unclear.This study investigated the ecological and metabolic responses of the different phytoplankton functional groups to elevated CO_(2) partial pressure and nitrate deficiency in two different environments of the eastern Indian Ocean(EIO).It is revealed that phytoplankton growth and metabolic rates are more sensitive to inorganic nutrients rather than CO_(2).The combined interactive effects of OA and N-limitation on phytoplankton populations are functional groupspecific.In particular,the abundance and calcification rate of calcifying coccolithophores are expected to be enhanced in the future EIO.The underlying mechanisms for this enhancement may be ascribed to coccolithophore’s lower carbon concentrating mechanisms(CCMs)efficiency and OA-induced[HCO^(-)_(3)]increase.In comparison,the abundance of non-calcifying microphytoplankton(e.g.,diatoms and dinoflagellates)and primary productivity would be inhibited under those conditions.Diff erent from previous laboratory experiments,interspecifi c competition for resources would be an important consideration in the natural phytoplankton populations.These combined factors would roughly determine calcifying coccolithophores as“winners”and non-calcifying microphytoplankton as“losers”in the future ocean scenario.Due to the large species-specific differences in phytoplankton sensitivity to OA,comprehensive investigations on oceanic phytoplankton communities are essential to precisely predict phytoplankton ecophysiological response to ocean acidification.

Increased light availability enhances tolerance against ocean acidification-related stress in the calcifying macroalga Halimedaopuntia

Although the adverse impacts of ocean acidification(OA)on marine calcifiers have been investigated extensively,the anti-stress capabilities regulated by increased light availability are unclear.Herein,the interactive effects of three light levels(30μmol photons/(m^(2)·s),150μmol photons/(m^(2)·s),and 240μmol photons/(m^(2)·s)combined with two pCO_(2)concentrations(400 ppmv and 1400 ppmv)on the physiological acclimation of the calcifying macroalga Halimeda opuntia were investigated using a pCO_(2)-light coupling experiment.The OA negatively influenced algal growth,calcification,photosynthesis,and other physiological performances in H.opuntia.The relative growth rate under elevated pCO_(2)conditions significantly declined by 13.14%−41.29%,whereas net calcification rates decreased by nearly three-fold under OA conditions.Notably,increased light availability enhanced stress resistance through the accumulation of soluble organic molecules,especially soluble carbohydrate,soluble protein,and free amino acids,and in combination with metabolic enzyme-driven activities,OA stress was alleviated.The carotenoid content under low light conditions increased markedly,and the rapid light curve of the relative electron transport rate was enhanced significantly by increasing light intensities,indicating that this new organization of the photosynthetic machinery in H.opuntia accommodated light variations and elevated pCO_(2)conditions.Thus,the enhanced metabolic performance of the calcifying macroalga H.opuntia mitigated OA-related stress.

Atmospheric Carbon Dioxide Reconstruction and Ocean Acidification Deduced from Carbon Isotope Variations across the Triassic–Jurassic Boundary in the Qiangtang Area, Tibetan Plateau

Objective The end-Triassic mass extinction was one of the five most profound Phanerozoic extinction events.This event was accompanied by a series of significant environmental changes,of which the most notable is the emergence of warm climate and the world-wide disappearance of carbonate platform.

Response of ocean acidification to atmospheric carbon dioxide removal

Artificial CO_(2)removal from the atmosphere(also referred to as negative CO_(2)emissions)has been proposed as a potential means to counteract anthropogenic climate change.Here we use an Earth system model to examine the response of ocean acidification to idealized atmospheric CO_(2)removal scenarios.In our simulations,atmospheric CO_(2)is assumed to increase at a rate of 1%per year to four times its pre-industrial value and then decreases to the pre-industrial level at a rate of 0.5%,1%,2%per year,respectively.Our results show that the annual mean state of surface ocean carbonate chemistry fields including hydrogen ion concentration([H^(+)]),pH and aragonite saturation state respond quickly to removal of atmospheric CO_(2).However,the change of seasonal cycle in carbonate chemistry lags behind the decline in atmospheric CO_(2).When CO_(2)returns to the pre-industrial level,over some parts of the ocean,relative to the pre-industrial state,the seasonal amplitude of carbonate chemistry fields is substantially larger.Simulation results also show that changes in deep ocean carbonate chemistry substantially lag behind atmospheric CO_(2)change.When CO_(2)returns to its pre-industrial value,the whole-ocean acidity measured by[H^(+)]is 15%-18%larger than the pre-industrial level,depending on the rate of CO_(2)decrease.Our study demonstrates that even if atmospheric CO_(2)can be lowered in the future as a result of net negative CO_(2)emissions,the recovery of some aspects of ocean acidification would take decades to centuries,which would have important implications for the resilience of marine ecosystems.

Impacts of ocean acidification under multiple stressors on typical organisms and ecological processes

The oceans are taking up over one million tons of fossil CO_(2) per hour,resulting in increased/ρCO_(2) and declining pH,leading to ocean acidification(OA).At the same time,accumulation of CO_(2) and other greenhouse gases is causing ocean warming,which enhances stratification with thinned upper mixed layers,exposing planktonic organisms to increasing levels of daytime integrated UV radiation.Ocean warming also reduces dissolved oxygen in seawater,resulting in ocean deoxygenation.All these ocean global changes are impacting marine ecosystems and effects are well documented for each individual driver(pH,oxygen,temperature,UV).However,combined effects are still poorly understood,strongly limiting our ability to project impacts at regional or local levels.Different regions are often exposed(and often adapted)to contrastingly different physical and chemical environmental conditions and organisms,and ecosystems from different parts of the world will be exposed to unique combinations of stressors in the future.Understanding the modulating role of adaptation,species niche and stressors’interaction is key.This review,being a non-exhaustively explored one.aims to provide an overview on understandings of ecophysiological effects of OA and its combination with covarying drivers,mainly warming,deoxygenation and solar UV radiation.We propose a testable hypothetical model as well as future research perspectives.

Biocalcification crisis in the continental shelf under ocean acidification

Ocean acidification(OA)is a persistent challenge for humans and is predicted to have deleterious effects on marine organisms,especially marine calcifiers such as coral and foraminifera.Benthic foraminifera is an important component of sediment in the continental shelf,while little is known about the impact of ocean acidification on benthic foraminifera both at the community and individual level and associated calcium carbonate deposition.We conducted eight months continued culture experiment under the scenario of 400,800,1200 and 1600 ppm pCO_(2)gradients on living benthic foraminifera from four stations in the continental shelf of the West Pacific Ocean.Statistic results showed OA had a negative effect on the abundance of benthic foraminifera.In contrast,the diversity increased roughly under OA conditions implying OA might stimulate the emergence of rare species and promote community diversity to some extent.In addition,we confirmed that the offshore area wasn’t the refuge for benthic foraminifera while the nearshore one had more resistance to moderate acidification.Calcareous species Protelphidium tuberculatum was the dominant species occupying on average 75%in all treatments and its shell diameter,weight and thickness showed a decrease,indicating the decrease of calcification of benthic foraminifera.A relationship between the weight of P.tuberculatum and pCO_(2)(R^(2)=0.96)was established.Based on the present work,calcareous benthic foraminifera deposited 8.57104 t calcium carbonate per year and this might reduce by nearly half and 90%under 800 and 1200 ppm scenarios,which indicates a biocalcification crisis under ongoing OA.This work shows an analogy for palaeoceanic OA and also provides new insights into the sediment of calcium carbonate in the future.

Light history modulates growth and photosynthetic responses of a diatom to ocean acidification and UV radiation

To examine the synergetic effects of ocean acidification(OA)and light intensity on the photosynthetic performance of marine diatoms,the marine centric diatom Thalassiosira weissflogii was cultured under ambient low CO_(2)(LC,390μatm)and elevated high CO_(2)(HC,1000μatm)levels under low-light(LL,60μmol m^(-2)s^(-1))or high-light(HL,220μmol m^(-2)s^(-1))conditions for over 20 generations.HL stimulated the growth rate by 128 and 99%but decreased cell size by 9 and 7%under LC and HC conditions,respectively.However,HC did not change the growth rate under LL but decreased it by 9%under HL.LL combined with HC decreased both maximum quantum yield(FV/FM)and effective quantum yield(ΦPSII),measured under either low or high actinic light.When exposed to UV radiation(UVR),LL-grown cells were more prone to UVA exposure,with higher UVA and UVR inducing inhibition ofΦPSII compared with HL-grown cells.Light use efficiency(α)and maximum relative electron transport rate(rETRmax)were inhibited more in the HC-grown cells when UVR(UVA and UVB)was present,particularly under LL.Our results indicate that the growth light history influences the cell growth and photosynthetic responses to OA and UVR.

Simulated effects of interactions between ocean acidification,marine organism calcification, and organic carbon export on ocean carbon and oxygen cycles

Ocean acidification caused by oceanic uptake of anthropogenic carbon dioxide(CO_2) tends to suppress the calcification of some marine organisms. This reduced calcification then enhances surface ocean alkalinity and increases oceanic CO_2 uptake, a process that is termed calcification feedback. On the other hand, decreased calcification also reduces the export flux of calcium carbonate(Ca CO_3), potentially reducing Ca CO_3-bound organic carbon export flux and CO_2 uptake, a process that is termed ballast feedback. In this study, we incorporate a range of different parameterizations of the links between organic carbon export, calcification, and ocean acidification into an Earth system model, in order to quantify the long-term effects on oceanic CO_2 uptake that result from calcification and ballast feedbacks. We utilize an intensive CO_2 emission scenario to drive the model in which an estimated fossil fuel resource of 5000 Pg C is burnt out over the course of just a few centuries. Simulated results show that, in the absence of both calcification and ballast feedbacks, by year 3500, accumulated oceanic CO_2 uptake is2041 Pg C. Inclusion of calcification feedback alone increases the simulated uptake by 629 Pg C(31%), while the inclusion of both calcification and ballast feedbacks increase simulated uptake by 449–498 Pg C(22–24%), depending on the parameter values used in the ballast feedback scheme. These results indicate that ballast effect counteracts calcification effect in oceanic CO_2 uptake. Ballast effect causes more organic carbon to accumulate and decompose in the upper ocean, which in turn leads to decreased oxygen concentration in the upper ocean and increased oxygen at depths. By year 2600, the inclusion of ballast effect would decrease oxygen concentration by 11% at depth of ca. 200 m in tropics. Our study highlights the potentially critical effects of interactions between ocean acidification, marine organism calcification, and Ca CO3-bound organic carbon export on the ocean carbon and oxygen cycles.

Effects of solar radiation modification on the ocean carbon cycle:An earth system modeling study

Solar radiation modification(SRM,also termed as geoengineering)has been proposed as a potential option to counteract anthropogenic warming.The underlying idea of SRM is to reduce the amount of sunlight reaching the atmosphere and surface,thus offsetting some amount of global warming.Here,the authors use an Earth system model to investigate the impact of SRM on the global carbon cycle and ocean biogeochemistry.The authors simulate the temporal evolution of global climate and the carbon cycle from the pre-industrial period to the end of this century under three scenarios:the RCP4.5 CO_(2) emission pathway,the RCP8.5 CO_(2) emission pathway,and the RCP8.5 CO_(2) emission pathway with the implementation of SRM to maintain the global mean surface temperature at the level of RCP4.5.The simulations show that SRM,by altering global climate,also affects the global carbon cycle.Compared to the RCP8.5 simulation without SRM,by the year 2100,SRM reduces atmospheric CO_(2) by 65 ppm mainly as a result of increased CO_(2) uptake by the terrestrial biosphere.However,SRM-induced change in atmospheric CO_(2) and climate has a small effect in mitigating ocean acidification.By the year 2100,relative to RCP8.5,SRM causes a decrease in surface ocean hydrogen ion concentration([H^(+)])by 6% and attenuates the seasonal amplitude of[H^(+)]by about 10%.The simulations also show that SRM has a small effect on globally integrated ocean net primary productivity relative to the high-CO_(2) simulation without SRM.This study contributes to a comprehensive assessment of the effects of SRM on both the physical climate and the global carbon cycle.

Advances in Chinese and international biogeochemistry research in the western Arctic Ocean: a review

Over the past decades, the Arctic Ocean has experienced rapid warming under climate change, which has dramatically altered its physical and biogeochemical properties. Reduction in the sea-ice cover is one of the most important driving forces of biogeochemical changes in the Arctic Ocean. Between 1999 and 2016, seven Chinese National Arctic Research Expeditions have taken place in the Bering and Chukchi seas, allowing assessment of the biogeochemical response of the western Arctic Ocean to global warming. Herein, we summarize advances in Chinese and international marine biogeochemistry research in the western Arctic Ocean, reviewing results from the Chinese expeditions and highlighting future trends of biogeochemistry in the Pacific Arctic region. The findings reported in this paper contribute towards a better understanding of water masses, greenhouse gases, nutrients, ocean acidification, and organic carbon export and burial processes in this region.

Processes Controlling the Carbonate Chemistry of Surface Seawater Along the 150°E Transect in the Northwest Pacific Ocean

The problem of ocean acidification caused by the increase of atmospheric carbon dioxide concentration is becoming increasingly prominent.Field observation in the northwest Pacific Ocean was carried out along the 150°E transect in November 2019.The distribution characteristics and influencing factors of the surface seawater carbonate chemistry,including dissolved inorganic carbon(DIC),total alkalinity(TA),pH,partial pressure of carbon dioxide(pCO_(2))and aragonite saturation state(Ω_(arag))were investigated.DIC and TA ranged from 1915 to 2014μmol kg^(−1)and 2243 to 2291μmol kg^(−1),respectively;DIC in general decreased with decreasing latitude,but TA had no clear latitudinal gradient.pCO_(2)values increased with the decrease of latitude and were all below the atmospheric pCO_(2)level,ranging from 332 to 387μatm.pH on the total hydrogen ion concentration scale(pH_(T))decreased with the decrease of latitude in the range of 8.044–8.110,whileΩ_(arag) increased with the decrease of latitude in the range of 2.61–3.88,suggesting that the spatial distributions of pH_(T) andΩ_(arag) were out of phase.Compared with the present,the predicted values of pH_(T) and Ω_(arag) by the end of this century would decrease remarkedly;larger declines were found in the higher pH_(T) and Ω_(arag) regions,resulting in the differences along the meridional gradient becoming smaller for bothpH_(T) and Ω_(arag).

Impact of seawater acidification on shell property of the Manila clam Ruditapes philippinarum grown within and without sediment

Although the impact of ocean acidification on marine bivalves has been previously investigated under mainly controlled laboratory conditions,it is still unclear whether the impact of acidification on sediment-burrowing species differs between those within or without sediment.In order to fill this gap in our knowledge,we compared shell properties of the infaunal Manila clam(Ruditapes philippinarum)exposed to three pH concentrations(7.4,7.7,and 8.0),within and without sediments.In the first experiment(140 d),clams were exposed to seawater in an acidification system without sediment.A decrease in shell weight corresponding to the increase in dissolution rate was observed in the group ofpH 7.4,at which shell color disappeared or whitened.SEM observations confirmed the changes of the external shell surface.In the second experiment(170 d),sediment was placed at the bottom of each exposure chamber.The effects were found obvious in shell dissolution rate and shell color in the shell specimens exposed to overlying seawater but not found in the shell specimens exposed to sediment.Although the experimental period was longer in the second experiment,shell specimens in the first experiment were more seriously damaged than those in the second experiment under acidic seawater conditions.Our results,in relation to the defense function of the shell,show that marine bivalves in burrowing behavior are more adaptable to seawater acidification than those who do not burrow into sediment.