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Pelagic calcifying protists such as coccolithophores and foraminifera represent an important microbial component of the marine carbon cycle. Although their calcitic shells are preserved in oceanic sediments over millennia, their resilience in the future decades is uncertain. We review current literature describing the response of calcifying protists to ocean acidification and temperature warming. We examine these key ecological and biogeochemical processes through the cellular perspective, exploring the physiological, metabolic, and molecular responses of calcifying protists. Ocean acidification is a chemical process that takes place in the seawater outside the cell, whereas protists calcify inside a modified cellular microenvironment. The function of these calcification compartments depends on cellular response to ocean acidification, such as maintaining pH homeostasis. The response of calcifying protists to ocean acidification and temperature warming is species-specific, with no unifying trends but rather a range of sensitivity levels. Coccolithophores and foraminifera display physiological sensitivity that may hamper their ecological success in comparison to non-calcifying species. Yet, certain species may be more adaptable, especially when comparing to highly vulnerable calcifying molluscs as pteropods. As the molecular machinery mediating cellular calcification is not fully resolved, as well as the functional role of the calcitic shell, our ability to predict the fate of calcifying microorganisms in a warmer, more acidic ocean is limited. We propose the urgent need to expand the study of these model systems by advancing cell biology approaches, to better understand the impact of climate change on microbial food webs in the ocean., (© The Author(s) 2025. Published by Oxford University Press on behalf of the International Society for Microbial Ecology.)

The intensity, frequencye and duration of seawater acidification in coastal seas have already surpassed projections for open oceans. Bivalve larvae are extremely sensitive to intensifying coastal seawater acidificaiton during their initial shell building, a critical period constraining recruitment success and population maintenance, but underlying mechanisms of larval shell formation sensitivity to acidification remain largely debated. Here, we performed an integrated analysis of the transcriptome and lipidome of trochophore of Ruditapes philippinarum to compare the core molecular responses involved in initial shell formation under ambient (pH 8.1), moderately (pH 7.7), and severely (pH 7.4) acidified conditions. Ocean acidification (OA) affected the ion transport efficiency by inhibiting gene expression of key ion transporters, thereby inhibiting initial shell formation, but the gene downregulation in the moderate exposure group was more significant. OA also induced major membrane lipid remodeling in larvae, which also significantly affected the ion transport efficiency. The TAG content of larvae which sustained the energy supply for active transport of calcification substrates and synthesis of organic matrix in the severe exposure group was significantly reduced. Overall, OA inhibited the formation of the initial larval shell, but different levels of OA had different inhibitory mechanisms on the initial larval shell formation, and the present study also further identified the role of lipids in initial shell formation, which can provide a theoretical basis for for a more accurate and comprehensive assessment of the impact of OA on bivalve calcification in an acidifying ocean., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2025 Elsevier Inc. All rights reserved.)

Greenhouse gas emissions from land-use change, fossil fuel, agriculture, transportation, and electricity sectors expose marine ecosystems to overlapping environmental stressors. Existing climate vulnerability assessment methods analyze the frequency of extreme conditions but often minimally consider how environmental data gaps hinder assessments. Here, we show an approach that assesses vulnerability and the uncertainty introduced by monitoring data gaps, using a case study of ocean acidification and deoxygenation in coastal California. We employ 5 million publicly available oceanographic observations and existing studies on species responses to low pH, low oxygen conditions to calculate vulnerability for six ecologically and economically valuable benthic invertebrate species: red sea urchin (Mesocentrotus franciscanus), purple sea urchin (Strongylocentrotus purpurpatus), warty sea cucumber (Apostichopus parvimensis), pink shrimp (Pandalus jordani), California spiny lobster (Panulirus interruptus), and Dungeness crab (Metacarncinus magister). Further, we evaluate the efficacy of current monitoring programs by examining how data gaps heighten associated uncertainty. We find that most organisms experience low oxygen (<35% saturation) conditions less frequently than low pH ( < 7.6) conditions. It is only deeper dwelling (>75 m depth) life stages such as Dungeness crab adults and pink shrimp embryos, juveniles, and adults that experience more frequent exposure to low oxygen conditions. Adult Dungeness crabs experience the strongest seasonal variation in exposure. Though these trends are intriguing, exposure remains low for most species despite well-documented pH and oxygen declines and strengthening upwelling in the central portions of the California Current. Seasonal biases of data collection and sparse observations near the benthos and at depths where organisms most frequently experience stressful conditions undermine exposure estimates. Herein we provide concrete examples of how pink shrimp and Dungeness crab fisheries may be impacted by our findings, and provide suggestions for incorporating oceanographic data into management plans. By limiting our scope to California waters and assessing the limitations presented by current monitoring coverage, this study aims to provide a granular, actionable framework that policymakers and managers can build from to prioritize targeted enhancements and sustained funding of oceanographic monitoring recommendations., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2025 Zulian et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

Local adaptation plays a critical role in an organism's ability to survive and reproduce in diverse environmental conditions, potentially improving an organism's response to stressful conditions such as ocean acidification or pollution. In this study, the effects of lower pH coupled with the presence of environmental contaminants were assessed on sea urchins (Paracentrotus lividus) collected outside and inside a volcanic CO 2 -vent system, where the mean ambient pH is 8.1 and 7.7, respectively. Both groups of sea urchins were spawned, and offspring were reared at pH 8.1 and 7.7, and in the presence or absence of a mixture of 100 μg/L of glyphosate and its main metabolite aminomethylphosphonic acid. Offspring performance metrics (development, abnormalities, and growth) were investigated under the different exposure conditions. The exposure to reduced pH affected the development and larval growth in echinoplutei obtained from adults of both sites, although to a different extent. Chemicals mixture had an additive effect in slowing embryo development. Results revealed that sea urchins living within the lower pH Vents area exhibited significantly higher egg quality, which likely enhanced embryonic development, reduced abnormalities, and increased larval size compared to their counterparts outside the Vents system, both in the presence and absence of contaminants. Findings suggest that sea urchins living within the CO 2 -Vents system developed adaptations to thrive under lower pH conditions. Elevated egg quality and improved offspring performance suggest organisms' resilience to environmental stressors associated with seawater acidification. Although insights gained from this study are preliminary, mostly due to the limited number of replicates in the egg biochemical analysis, they contribute to unveiling the adaptive capabilities of sea urchins in facing ongoing ocean acidification challenges., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2025 Elsevier Inc. All rights reserved.)

Ocean acidification occurs at a rate unprecedented for millions of years, forcing sessile organisms, such as oysters, to respond in the short term by relying on their phenotypic plasticity. Phenotypic plasticity has limits, tipping points, beyond which species will have to adapt or disappear. These limits could be related to the adaptation of species to different habitat variabilities. Here we expose juvenile pearl oysters, Pinctada margaritifera, to a broad range of pH and determine the response at the gross physiological, lipidome and transcriptome levels. Thus, we identify its high tolerance with low tipping points at pH 7.3-6.8 below which most physiological parameters are impacted. We then compare the transcriptomic reaction norms of the tropical subtidal P. margaritifera, with those of an intertidal temperate oyster, Crassostrea gigas, reusing data from a previous study. Despite showing similar tipping points with C. gigas, P. margaritifera exhibits strong mortalities and depletion of energy reserves below the tipping points, which is not the case for C. gigas. This divergence relies mainly on the induction of metabolic depression, an adaptation to intertidal habitats in C. gigas, but not in P. margaritifera. Our method makes it possible to detect divergences in phenotypic plasticity, probably linked to the species' specific life-history strategies related to different habitats, which will determine the survival of species to ongoing global changes. Such an approach is particularly relevant for studying the physiology of species in a world where physiological tipping points will be increasingly exceeded., (© 2025. Published by The Company of Biologists.)

Climate change as a result of increases in greenhouse gas emissions, such as CO 2 , is causing significant alteration in global environmental conditions, including ocean acidification (OA). Although the depletion of the ozone layer has reduced, the penetration of ultraviolet-B (UVB) radiation into the oceans still remains an environmental factor that may potentially enhance the effects of OA on biota. Improved understanding of the complex interactions between multiple stressors, such as UV-B radiation and increased CO 2 levels, is thus important for safeguarding ecosystems and developing effective conservation and management strategies. A 72 h experiment was carried out to investigate the combined effects of UVB irradiance (0.5 W m -2 ) and varying CO 2 levels (350, 500, 1000 ppm) on the diatom Skeletonema pseudocostatum. The study aimed to characterize the potential combined effects at different levels of biological organization, including ROS formation, lipid peroxidation (LPO), photosynthesis, pigments, oxidative phosphorylation (OXPHOS) and growth. The findings indicate that exposure to elevated CO 2 (500 ppm) alone resulted in increased total carotenoid content and growth of S. pseudocostatum, but did not significantly impact photosystem efficiency, oxidative stress, and OXPHOS. Sole UVB exposure induced oxidative stress, inhibited photosynthesis and OXPHOS processes, and suppressed growth in S. pseudocostatum. However, when co-exposed with CO 2 , synergistic impacts were observed for reactive oxygen species (ROS), lipid peroxidation (LPO), and growth, while carotenoids were reduced in an antagonistic manner. A putative impact pathway was proposed as an initial effort to characterize the combined effects of these stressors under proposed future marine OA scenarios involving elevated CO 2 ., Competing Interests: Declaration of Competing Interest, (Copyright © 2025 The Authors. Published by Elsevier Inc. All rights reserved.)

Rhodolith holobionts are formed by calcareous coralline algae (e.g., Corallinales) and associated microbiomes. The largest rhodolith bank in the South Atlantic is located in the Abrolhos Bank, in southwestern Brazil, covering an area of 22,000 km 2 . Rhodoliths serve as nurseries for marine life. However, ocean acidification threatens them with extinction. The acute effects of high pCO₂ levels on rhodolith metatranscriptomes remain unknown. This study investigates the transcriptomic profiles of rhodoliths exposed to short-term (96-h) high pCO₂ levels (up to 1638 ppm). Metatranscriptomes were generated for both dead and alive rhodoliths (15.48 million Illumina reads in total). Alive rhodoliths showed an enrichment of gene transcripts related to environmental stress responses and photosynthesis (Cyanobacteria). In contrast, the metatranscriptomes of dead rhodoliths were dominated by heterotrophic (Proteobacteria and Bacteroidetes) metabolism and virulence factors. The rhodolith holobiont metatranscriptomes respond rapidly to short-term acidification (within 1 h), suggesting that these holobionts may have some capacity to cope with acute acidification effects. However, the negative impacts of prolonged ocean acidification on rhodolith health cannot be overlooked. Rhodoliths exposed to low pH (7.5) for 96 h exhibited a completely altered transcriptomic profile compared to controls. This study highlights the plasticity of rhodolith transcriptomes in the face of ocean acidification and climate change., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2025 Elsevier B.V. All rights reserved.)

Since the industrial revolution, the direct impacts of elevated CO 2 concentrations, such as ocean acidification, and indirect impacts, such as extreme drought events, have synergistically influenced coastal ecosystems, including seagrass meadow. Consequently, investigating the individual and combined effects of ocean acidification and extreme drought-induced increased salinity on seagrasses is crucial for enhancing the management and monitoring of these ecosystems. This study used a two-factor crossover indoor simulation experiment to thoroughly examine the effects of seawater acidification at pH 7.7 and elevated salinity levels at 43‰ and 51‰ on the physiological responses and growth status of the dominant tropical seagrass species Thalassia hemprichii. The results indicated that seawater acidification at pH 7.7 significantly enhanced the growth rate and photosynthetic activity of T. hemprichii across all salinity levels. A salinity of 43‰ activated certain antioxidant enzymes without inducing severe osmotic stress in T. hemprichii and positively influenced leaf photosynthetic activity, with a 15.6% increase in growth rate compared to the CK group. The extreme salinity of 51‰ imposed osmotic stress, leading to increase in reactive oxygen species and decreased photosynthetic activity and a 52% decrease in growth rate compared to seagrasses in the CK group. Under future scenarios of ocean acidification and frequent extreme droughts, T. hemprichii inhabiting enclosed marine environments may exhibit greater adaptability and secure an ecologically competitive edge. Our findings underscore the importance of conserving declining meadows, forecasting the ecological trajectory of these ecosystems, and managing salinity in lagoons for the well-being of seagrass ecosystems., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2025 Elsevier Ltd. All rights reserved.)

Ocean acidification (OA) associated with climate change is expected to lower the ocean's pH by 0.5 units by 2100. Whilst associated effects such as coral bleaching and shell calcification are well documented, lesser-known impacts are the 'invisible' effects on animal sensory systems. Olfactory disruption impacts the behaviour towards chemical cues in many marine species, including crustaceans. We examine the effects of microplastic odour and additional stressors on the European green crab C. maenas . Using uridine diphosphate (UDP) and uridine triphosphate (UTP) as a sex pheromone bouquet, glutathione (GSH) as a food cue, and polyethylene (PE) as plastic odour, cues were mixed with carboxycellulose to create slow-release gels. Crabs were exposed to gels in seawater pH values of 8.2, 7.6, and 7.2. Crabs took longer to react to all odours in reduced pH conditions (pH 8.2 to pH 7.2, p = 0.0017). At a low pH, PE-exposed crabs exhibited attraction towards microplastic odour and changed behavioural responses by burying. The study confirms low pH as disruptive to olfaction and highlights that plastic derivatives can become more bioactive at reduced pH levels, potentially increasing the threat posed by microplastic pollution. Further research is required to determine the potential long-term impacts of the combined threat of microplastics and reduced pH in the environment.

Increased carbon dioxide (CO 2 ) in the ocean is changing seawater chemistry. Behavioural alterations in CO 2 exposed fish have been linked to changes in the central nervous system (CNS). However, we hypothesise that receptor cells in direct contact with the environment are more susceptible to changes in water chemistry than the CNS. Electrophysiology, histology, and transcriptomics were used to explore the effect of exposure to CO 2 acidified water on the olfactory epithelium (OE) of the Senegalese sole (Solea senegalensis). The upper and lower OE of this flatfish detect different odorants and are in contact with different environments. Acute exposure to acidified water decreased olfactory sensitivity more in the upper than in the lower OE. After chronic exposure to high CO 2 there were no histological changes in the upper OE; however, in the lower OE, there was a massive infiltration of melanomacrophage (MMC) and tissue disorganization. In addition, in the upper OE, differential expressed gene transcripts (DETs) were related to inflammation and innate immune processes whereas in the lower OE, DETs were related to the adaptative immune response. Differential regulation of genes related to neurogenesis and plasticity occurred in both epithelia. The effects of ocean acidification in sole OE depends on the nostril; however, the occurrence of an exacerbated immune response, OE remodelling and reduced sensitivity indicate that ocean acidification is likely to have significant and unpredictable consequences for behaviour., Competing Interests: Declaration of competing interest The authors declare no competing or financial interests., (Copyright © 2025 The Authors. Published by Elsevier Inc. All rights reserved.)

As the most significant carbon sink on Earth, the ocean has been inevitably absorbing excess atmospheric CO 2 , leading to high concentration of CO 2 that threatens marine life. Like emissions of greenhouse gases, this issue demands urgent attention. Hence, comprehensive investigation and comparison, including both non-electrochemical and electrochemical direct ocean capture (DOC) methods, are revealed in this review. The non-electrochemical approach utilizes specialized materials such as gas-permeable membranes (GPM), hollow fiber membrane contactors (HFMC), and ion exchange resins to extract CO 2 from seawater. In contrast, the electrochemical method employs chemical reactions to generate H + or OH - ions, which adjust the pH value of seawater to either release CO 2 gas or precipitate carbonate, thereby removing dissolved carbon. This article comprehensively overviews each method, including the latest research findings, underlying principles, employed equipment, and performance metrics. Finally, the achievements, current gaps, corresponding perspectives, and potential solutions in CO 2 capture from seawater are also proposed., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier Ltd. All rights reserved.)

Comprehensive understanding of environmental multiple stressors on calcification in marine calcifiers remains an important topic of study, especially under ocean global change associated with multiple stressors. We explore the impact of multiple stressor on pteropod calcification in the southern Salish Sea (Washington, U.S.), a coastal estuarine system that exhibits a high degree of spatial and temporal variability in multiple environmental parameters across sampling locations. We hypothesized that such variability is associated with differences in pteropod calcification. Shell thickness and shell density across pteropod life history stages was compared with high-resolution outputs from a realistic model of regional circulation and biogeochemistry to explore how the mean and variability of multiple stressors (aragonite saturation state (Ω ar ), temperature, food availability) influence calcification. We found that both the mean and variability in multiple stressors play a major role in calcification in pteropods, with a generalized linear model explaining more than 60% of the variance. We suggest two different modes of shell building: stable conditions of lower mean Ω ar trigger the loss of shell thickness and density. In the more variable habitats, i.e., where the variability occurs over diel and seasonal scales, shell thickness increases at higher Ω ar variability and greater food availability, which might partially compensate for the loss of shell density. This plastic response appears to be consistent across life stages and could represent a response mechanism that allows some compensatory calcification under less favourable conditions. However, compensation is very limited, as evident by lower shell growth resulting in shell sizes comparable to early life stages. These results substantially improve the understanding of the variability in multiple stressors on the calcification process under multiple stressors and provide a foundation for the development of two new proxies for calcification monitoring, and with implications for marine carbon dioxide removal strategies., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Volcanic emissions in shallow vents influence the biogeochemistry of the sedimentary compartment, creating marked abiotic gradients. We assessed the spatial dynamics of the sediment compartment, as for the composition and origin of organic matter and associated prokaryotic community, in a volcanic shallow CO 2 vent (Vulcano Island, Italy). Based on elemental (carbon, nitrogen content and their ratio) and isotopic composition (δ 13 C, δ 15 N and δ 34 S), the contribution of vent-derived organic matter (microbial mats) to sedimentary organic matter was high close to the vent, while the marine-derived end-members (seagrasses) contributed highly at increasing distance. Chemoautotrophic Campylobacterota and hyperthermophilic Achaea prevailed close to the vent, whilst phototrophic and chemoheterotrophic members dominated at increasing distance. Abiotic gradients generated by the volcanic CO 2 vent drive relevant changes in the composition, origin and nutritional quality of sedimentary organic matter, and influence the structure and complexity of associated prokaryotic communities, with expected relevant impact on the entire food-web., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier Ltd.. All rights reserved.)

The ongoing decline in seawater pH, driven by the absorption of excess atmospheric CO 2 , represents a major environmental issue. This reduction in pH can interact with metal pollution, resulting in complex effects on marine phytoplankton. In this study, we examined the combined impacts of seawater acidification and copper (Cu) exposure on the marine diatom Phaeodactylum tricornutum. Our data indicate that elevated pCO 2 had a minor effect on the growth and photochemistry and overall performance of P. tricornutum. However, seawater acidification significantly influenced cell size, surface roughness, and adhesion. Higher pCO 2 levels led to increased Cu accumulation in P. tricornutum under low ambient Cu concentrations, while significantly reducing Cu accumulation. The smaller cell size and reduced negative charge on the cell surface may explain the decreased Cu accumulation and toxicity. In response to metal stress, P. tricornutum upregulated Cu efflux to mitigate the increased Cu stress in acidified seawater. The expression of the metal transporter gene CTR1 and the reductase gene FRE1 were significantly downregulated, while ATPase5-1B was upregulated in cells exposed to elevated Cu concentrations at 1200 μatm pCO 2 . Our study provides useful insights into the interactions between metals and diatoms in an increasingly acidified ocean., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier Ltd. All rights reserved.)

This study examines the seasonal variations in carbonate system parameters in the East China Sea (ECS) off the Yangtze River estuary (YRE) and analyzes the contributions of anthropogenic CO₂ and eutrophication to acidification. Carbonate parameters data were collected during summer 2019 and combined winter 2011. During winter, acidification is primarily driven by rising atmospheric CO₂, with minimal impact from biological processes. In contrast, summer presents a different pattern: enhanced photosynthesis due to eutrophication in surface waters helps mitigate the acidification effects of atmospheric CO₂ increases, while in bottom waters, the combined pressures of atmospheric CO₂ and intensified aerobic respiration leads to more severe acidification. Notably, biological processes now contribute more to acidification than increasing atmospheric CO₂ in the bottom waters. Our projections indicate that the summer bottom waters will experience the most pronounced acidification, with average pH levels expected to decline from 8.04 to 7.82 and aragonite saturation state (Ω ar ) values decreasing from 2.24 to 1.38 between 2000 and 2100. Additionally, our study indicates that winter acidification trends are also concerning, with pH only slightly higher than in summer bottom waters. The buffering capacity and the DIC:TA ratio play significant roles in determining the rate of future pH and Ω ar declines. The strong buffering capacity in summer surface waters mitigates the pH decline, while the low DIC:TA ratio results in a rapid drop in Ω ar ., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2025 Elsevier Ltd. All rights reserved.)

In order to explore the impact of CO 2 -driven ocean acidification (OA) on gene expression of sea urchins, gametes of Hemicentrotus pulcherrimus were fertilized and developed to the four-armed larvae in either seawater at current pH levels (pH NBS  = 7.98) or in three laboratory-controlled OA conditions (ΔpH NBS  = -0.3, -0.4, -0.5 units) based on the projections of the Intergovernmental Panel on Climate Change (IPCC) for 2100. Four-armed larval specimens were collected, and comparative transcriptome analysis was then performed. The results showed that 58 differentially expressed genes (DEGs) were identified in OA-treated groups as compared to the control. Moreover, more transition and transversion SNPs were observed in OA-treated groups than those in the control indicating a potential occurrence of adaption to OA in H. pulcherrimus larvae. Six candidate DEGs shared among OA-treated groups were identified as potential biomarkers correlated with low pH tolerance, mainly enriched in nine pathways associated with Notch signaling, dorso-ventral axis formation, oxidative phosphorylation, lysine degradation, valine, leucine and isoleucine degradation, lysosome, cell adhesion molecules, glutathione metabolism and PPAR signaling pathway. These results will not only enrich our knowledge of the impacts of OA on sea urchin larvae from the aspect of gene expression, provide a better understanding on larval forms coping with OA, but also offer more clues and biomarkers for developing protection or management strategies for sea urchins under near-future OA conditions., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2025 Elsevier Ltd. All rights reserved.)

Ocean acidification can significantly affect the physiological performance of macroalgae. While copper (Cu) is an essential element for macroalgae and has been extensively studied, the interactive effects of ocean acidification and Cu on these organisms remain less understood. In this study, we measured the photosynthetic characteristics of Ulva lactuca exposed to varying Cu concentrations at two CO 2 levels (415 ppmv, low concentration; 1000 ppmv, high concentration). The results indicated that during chronic toxicity testing, the growth of juvenile U. lactuca significantly increased at Cu concentrations of 0.001 μM, 0.01 μM, and 0.1 μM regardless of low CO 2 concentrations or high CO 2 concentrations condition. In acute toxicity tests, elevated Cu concentrations negatively impacted the growth rate, yield, and photosynthetic rate of U. lactuca under low CO 2 concentrations. Conversely, high CO 2 concentrations enhanced the photosynthetic capacity of U. lactuca with increased Cu concentrations, while the growth rate significantly decreased at Cu concentration of 1.5 μM. Additionally, the activities of peroxidase (POD) and ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) increased, with an enhancement of malondialdehyde (MDA) content at 1.5 μM Cu under high CO 2 conditions. However, the structure of the chloroplast thylakoid was disrupted by elevated Cu concentrations. These findings suggest that low Cu concentrations promote the growth of U. lactuca, whereas high Cu concentrations inhibit algal growth, and ocean acidification may exacerbate the adverse effects of Cu on U. lactuca in acute toxicity tests., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2025 Elsevier B.V. All rights reserved.)

Anthropogenic carbon dioxide emissions have been increasing rapidly in recent years, driving pH and oxygen levels to record low concentrations in the oceans. Eastern boundary upwelling systems such as the California Current System (CCS) experience exacerbated ocean acidification and hypoxia (OAH) due to the physical and chemical properties of the transported deeper waters. Research efforts have significantly increased in recent years to investigate the deleterious effects of climate change on marine species, but have not focused on the impacts of simultaneous OAH stressor exposure. Additionally, few studies have explored the physiological impacts of these environmental stressors on the earliest life stages, which are more vulnerable and represent natural population bottlenecks in organismal life cycles. The physiological response of the ecologically and commercially important red sea urchin (Mesocentrotus franciscanus) was assessed by exposing larvae to a variety of OAH conditions, mimicking the range of ecologically relevant conditions encountered currently and in the near future along the CCS. Skeleton dissolution, larval development, and gene expression show a response with clearly delineated thresholds that were related to OAH severity. Skeletal dissolution and the induction of Acid-sensing Ion Channel 1A at pH 7.94/5.70 DO mg/L provide particularly sensitive markers of OAH, with dramatic shifts in larval morphology and gene expression detected at the pH/DO transition of 7.71/3.71-7.27/2.72 mg/L. Experimental simulations that describe physiological thresholds and establish molecular markers of OAH exposure will provide fishery management with the tools to predict patterns of larval recruitment and forecast population dynamics., (© 2025 John Wiley & Sons Ltd.)

Ocean acidification, a major consequence of climate change, poses significant threats to marine organisms, particularly when combined with other environmental stressors such as chemical pollution. This study investigated the physiological responses of Trochus niloticus to a 28-day exposure of ocean acidification and/or sulfamethoxazole, a commonly detected antibiotic in the South China Sea. Exposure to either acidification or sulfamethoxazole individually triggered adaptive responses through immune activation, antioxidant reactions, and metabolic adjustments. However, concurrent exposure resulted in significant adverse effects, including compromised immunity, oxidative damage, and disrupted energy budget. These findings provide new insights into how ocean acidification interacts with antibiotic pollution to synergistically impact marine gastropods, suggesting that multiple stressors may pose greater threats to T. niloticus populations than single stressors alone., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier Inc. All rights reserved.)

The increase of anthropogenic CO 2 in the Earth's atmosphere reduces the pH and raises the temperature of the oceans. The combination of both factors impacts the physiological responses and calcium carbonate structures of marine organisms. This study assessed the performance of the juvenile stage of the gastropod Acanthina monodon, after it was continuously exposed to treatments at two pCO 2 levels (400 and 1200 μatm) at two temperatures (10 °C and 15 °C) during the periods of embryonic encapsulated development and the early post-hatching juvenile stage. Juvenile performance was evaluated by quantifying shell growth, survival, foot adhesion strength, shell breakage resistance, and oxygen consumption rates. The results indicate that the combination of increased temperature and decreased pH increased juvenile shell growth, while only the decrease in pH had a negative effect on shell strength. However, juveniles were able to attach more strongly to substrates following exposure to the higher temperature level. Furthermore, the interaction of treatments had no effect on the metabolic rate or survival of juveniles, suggesting a possible compensatory effect of the juveniles to the more adverse conditions to which they were exposed., Competing Interests: Declaration of competing interest The authors declare that they have no conflicts of interest., (Copyright © 2024 Elsevier Ltd. All rights reserved.)

The West Coast rock lobster, Jasus lalandii , is a key ecological species and provides an important fishery resource in South Africa and Namibia. It is found along the west coast of southern Africa in the dynamic Benguela Current upwelling system. The low seawater pH of this system is expected to decline further due to ocean acidification and increase in upwelling in terms of frequency and severity. The lobster has therefore to respond to frequent and rapid changes in pH and other environmental impacts that are predicted to become more adverse in future. Although responses to such conditions are known for mature male and juvenile lobsters, there is a lack of information on mature females and later embryonic development. We addressed this by analysing the sensitivity of ovigerous ("berried") female WCRLs and their eggs/embryos to hypercapnia (high pCO 2 , low pH) and formulated the following research questions: (1) Can berried female WCRLs respond swiftly to large changes in pH? (2) What physiological mechanisms facilitate a potential response to a rapidly declining pH, i.e., acute hypercapnia? (3) Does a potential response persist during prolonged hypercapnia? (4) Are eggs/embryos impacted by hypercapnia? To investigate this, we exposed berried WCRLs to acute (pH 7.5) and chronic (up to 60 days at pH 7.5 and 7.8) hypercapnia. We applied extracellular acid-base analysis, microscopic examination of egg growth and development, and SEM of female exoskeleton structure and egg membranes. The results revealed that berried females efficiently respond to acute and chronic hypercapnia by means of increasing bicarbonate concentrations in the haemolymph. Moreover, embryo growth and development are not impacted by chronic hypercapnia, but growth shows geographical area-specific differences. We conclude that females and embryos of J. lalandii are as resilient to hypercapnia as previously shown for males and juveniles.

Global changes can profoundly affect the sex determination and reproductive output of marine organisms, disrupting the population structure and ecosystems. High CO 2 driven low pH in the context of ocean acidification (OA) has been shown to severely affect various calcifiers, but less is known about the extent to which low pH influences sex determination and reproduction of marine organisms, particularly mollusks. This study is the first to report a biased sex ratio over multiple generations toward females, driven by exposure to high CO 2 -induced low pH environments, using the ecologically and economically important Portuguese oyster ( Crassostrea angulata ) as a model. This phenomenon, which we term pH-mediated sex determination (PSD), has no consequences for fecundity, gonadal development, or reproductive function in the offspring. Moreover, PSD persisted into a second year of reproduction and was inherited across multiple generations. Transcriptomic analysis indicates PSD is associated with the activation of the Wnt signaling pathway in females and inhibition of spermiogenesis-related functions in males. This work expands our understanding of environmental sex determination and highlights the possible impact of global changes on reproduction and population dynamics of mollusks and other marine organisms.

The Gulf of Maine holds significant ecological and economic value for fisheries and communities in north-eastern North America. However, there is apprehension regarding its vulnerability to the effects of increasing atmospheric CO 2 . Substantial recent warming and the inflow of low alkalinity waters into the Gulf of Maine have raised concerns about the impact of ocean acidification on resident marine calcifiers (e.g. oysters, clams, mussels). With limited seawater pH records, the natural variability and drivers of pH in this region remain unclear. To address this, we present coastal water pH proxy records using boron isotope (δ 11 B) measurements in long-lived, annually banded, crustose coralline algae (1920-2018 CE). These records indicate seawater pH was low (~ 7.9) for much of the last century. Contrary to expectation, we also find that pH has increased (+ 0.2 pH units) over the past 40 years, despite concurrent rising atmospheric CO 2 . This increase is attributed to an increased input of high alkalinity waters derived from the Gulf Stream. This delayed onset of ocean acidification is cause for concern. Once ocean circulation-driven buffering effects reach their limit, seawater pH decline may occur swiftly. This would profoundly harm shellfisheries and the broader Gulf of Maine ecosystem., Competing Interests: Declarations. Competing interests: The authors declare no competing interests., (© 2025. The Author(s).)

To explore the dynamic molecular responses to CO 2 -driven ocean acidification (OA) during the early developmental stages of sea urchins, gametes of Strongylocentrotus intermedius were fertilized and developed to the four-armed larva stage in either natural seawater (as a control; pH NBS  = 7.99 ± 0.01) or acidified conditions (ΔpH NBS  = -0.3, -0.4, and - 0.5 units) according to the prediction for ocean pH by the end of this century. Specimens from five developmental stages (fertilization, cleavage, blastula, prism, and four-armed larva) were collected and comparative microRNA (miRNA) and mRNA transcriptome analyses were performed. The results showed that 1) a total of 22,224 differentially expressed genes (DEGs) and 51 differentially expressed miRNAs (DEMs) were identified in the OA-treated groups compared with the control group. 2) The numbers of both DEGs and DEMs were the largest at the blastula stage, indicating dramatic changes in gene expression. 3) Five "miR-1/DEG" modules were identified as potential biomarkers reflecting the response of sea urchins to OA during the early developmental period. 4) The PI3K/Akt signaling pathway was a key pathway involved in the response of S. intermedius to OA in its early developmental stages. This study deepens our understanding of the dynamic molecular regulatory mechanisms underlying sea urchin responses to CO 2 -driven OA., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationship that could have appeared to influence the work reported in this paper., (Copyright © 2024. Published by Elsevier Ltd.)

As a result of human activity, Earth's atmosphere and climate are changing at an unprecedented pace. Models based on short-term experiments predict major changes will occur in marine phytoplankton communities in the future ocean, but rarely consider how evolution or interactions with other microbes may influence these changes. Here, we experimentally evolved several phytoplankton in coculture with a heterotrophic bacterium, Alteromonas sp. EZ55, under either present-day or predicted future pCO2 conditions. Growth rates of phytoplankton generally increased over time under both conditions, but only Thalassiosira oceanica had evidence of a growth rate tradeoff in the ancestral environment after evolution at elevated pCO2. The growth defects observed in ancestral Prochlorococcus cultures at elevated pCO2 and in axenic culture were diminished after evolution, possibly due to regulatory mutations in antioxidant genes. Except for Prochlorococcus, mutational profiles suggested phytoplankton experienced primarily purifying selection, but most Alteromonas lineages showed evidence of directional selection, where evolution appeared to favor a metabolic switch between growth on small organic acids with cyanobacteria versus catabolism of more complex carbon substrates with eukaryotic phytoplankton. Evolved Alteromonas were also poorer "helpers" for Prochlorococcus, consistent with that interaction being a competitive Black Queen process rather than a true mutualism. This work provides new insights on how phytoplankton will respond to increased pCO2 and on the evolutionary mechanisms governing phytoplankton:bacteria interactions. It also clearly demonstrates that both evolution and interspecies interactions must be considered to predict future marine biogeochemistry., (© The Author(s) 2024. Published by Oxford University Press on behalf of the International Society for Microbial Ecology.)

Dinoflagellates, both armored and unarmored, with distinct cell wall difference, are being affected by elevated CO 2 -induced ocean acidification (OA). However, their specific responses to OA are not well understood. In this study, we investigated the physiological and molecular response of the armored species Prorocentrum obtusidens and the unarmored species Karenia mikimotoi to OA over a 28-day period. The results show that the two species responded differently to OA. Cell growth rate, particulate organic carbon (POC) content, and the activities of C 4 pathway enzymes decreased in P. obtusidens under future acidified ocean condition (pH 7.8, 1000 μatm pCO 2 ), but the activities of carbonic anhydrase (CA), ribulose-1,5-bisphosphate carboxylase/oxygenase (RubisCO), and superoxide dismutase (SOD) increased. Whereas cell growth rate, contents of Chl a and PON, and SOD activity altered insignificantly in K. mikimotoi, but contents of POC and total carbohydrate, and the activity of RubisCO increased while the activities of CA and C 4 pathway enzymes decreased. Transcriptomic analysis indicates that genes associated with antioxidative response, heat shock protein, proteasome, signal transduction, ribosome, and pH regulation were up-regulated in P. obtusidens but down-regulated in K. mikimotoi. Notably, the synthesis of soluble organic matter (i.e., spermidine and trehalose) was enhanced in K. mikimotoi, thereby regulating intracellular pH and improving stress resistance. This study highlights the divergent response of the armored and unarmored dinoflagellates to OA, with the unarmored dinoflagellate exhibiting a higher ability to withstand this stressor. Therefore, caution should be exercised when predicting the behavior and the eventual fate of dinoflagellates in the future acidified ocean., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier B.V. All rights reserved.)

Corals residing in habitats that experience high-frequency seawater pCO 2 variability may possess an enhanced capacity to cope with ocean acidification, yet we lack a clear understanding of the molecular toolkit enabling acclimatisation to environmental extremes or how life-long exposure to pCO 2 variability influences biomineralisation. Here, we examined the gene expression responses and micro-skeletal characteristics of Pocillopora damicornis originating from the reef flat and reef slope of Heron Island, southern Great Barrier Reef. The reef flat and reef slope had similar mean seawater pCO 2 , but the reef flat experienced twice the mean daily pCO 2 amplitude (range of 797 v. 399 μatm day -1 , respectively). A controlled mesocosm experiment was conducted over 8 weeks, exposing P. damicornis from the reef slope and reef flat to stable (218 ± 9) or variable (911 ± 31) diel pCO 2 fluctuations (μatm; mean ± SE). At the end of the exposure, P. damicornis originating from the reef flat demonstrated frontloading of 25% of the expressed genes regardless of treatment conditions, suggesting constitutive upregulation. This included higher expression of critical biomineralisation-related genes such as carbonic anhydrases, skeletal organic matrix proteins, and bicarbonate transporters. The observed frontloading corresponded with a 40% increase of the fastest deposited areas of the skeleton in reef flat corals grown under non-native, stable pCO 2 conditions compared to reef slope conspecifics, suggesting a compensatory response that stems from acclimatisation to environmental extremes and/or relief from stressful pCO 2 fluctuations. Under escalating ocean warming and acidification, corals acclimated to environmental variability warrant focused investigation and represent ideal candidates for active interventions to build reef resilience while societies adopt strict policies to limit climate change., (© 2024 The Author(s). Molecular Ecology published by John Wiley & Sons Ltd.)

Scenarios to stabilize global climate and meet international climate agreements require rapid reductions in human carbon dioxide (CO 2 ) emissions, often augmented by substantial carbon dioxide removal (CDR) from the atmosphere. While some ocean-based removal techniques show potential promise as part of a broader CDR and decarbonization portfolio, no marine approach is ready yet for deployment at scale because of gaps in both scientific and engineering knowledge. Marine CDR spans a wide range of biotic and abiotic methods, with both common and technique-specific limitations. Further targeted research is needed on CDR efficacy, permanence, and additionality as well as on robust validation methods-measurement, monitoring, reporting, and verification-that are essential to demonstrate the safe removal and long-term storage of CO 2 . Engineering studies are needed on constraints including scalability, costs, resource inputs, energy demands, and technical readiness. Research on possible co-benefits, ocean acidification effects, environmental and social impacts, and governance is also required.

When humans drive rapid environmental change, is it favourable to be a generalist or specialist? To address this question, we compare how specialist and generalist marine herbivores adjust their isotopic niches (used as proxy for trophic niche) in response to predicted resource alterations under the simulated effects of ocean warming and acidification (based on a 6-month mesocosm experiment). Here, we show that when exposed to multiple climate stressors, food resources homogenized towards dominance of turf algae and suspended organic matter, with generalists and specialists adjusting their trophic niches in opposing ways. Whilst the niche breath of most generalists narrowed under climate stressors, those of specialists generally broadened, causing increasing overlap between their niches. The magnitude of this change was such that some generalists turned into specialists, and vice versa. Under ocean acidification, there was a greater probability of generalists increasing and specialists maintaining their biomass, respectively, but under warming the biomass of both specialists and generalists had a greater probability of collapse. For specialists, this collapse occurred even though they had adequate thermal tolerance and the capacity to expand their trophic niche. Climate change constrains or liberates resources, but where they are homogenized, generalists and specialists are likely to converge their trophic niches so they can exploit transforming environments for their survival or adaptive advantage., Competing Interests: Declaration of competing interest The authors declare that they have no competing interests., (Copyright © 2024 The Authors. Published by Elsevier B.V. All rights reserved.)

Ocean acidification (OA) driven by human activities and climate change presents new challenges to marine ecosystems. At the same time, the risks posed by micro(nano)plastics (MNPs) and engineered nanoparticles (ENPs) to marine ecosystems are receiving increasing attention. Although previous studies have uncovered the environmental behavior and the toxic effects of MNPs and ENPs under OA, there is a lack of comprehensive literature reviews in this field. Therefore, this paper reviews how OA affects the environmental behavior of MNPs and ENPs, and summarizes the effects and the potential mechanisms of their co-exposure on marine organisms. The review indicates that OA changes the marine chemical environment, thereby altering the behavior of MNPs and ENPs. These changes affect their bioavailability and lead to co-exposure effects. This impacts marine organisms' energy metabolism, growth and development, antioxidant systems, reproduction and immunity. The potential mechanisms involved the regulation of signaling pathways, abnormalities in energy metabolism, energy allocation, oxidative stress, decreased enzyme activity, and disruptions in immune and reproductive functions. Finally, based on the limitations of existing research, actual environment and hot issues, we have outlined future research needs and identified key priorities and directions for further investigation. This review deepens our understanding of the potential effects of MNPs and ENPs on marine organisms under OA, while also aiming to promote further research and development in related fields., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier Inc. All rights reserved.)

Anadromous salmon populations are declining in the Pacific Northwest, with high mortality during the transition from fresh- to seawater as smolts, a stage particularly vulnerable to adverse environmental conditions. This study seeks to explore the impacts of warming and ocean acidification on the transition of life in freshwater to life at sea in Chinook salmon smolts. In a fully factorial experiment, we transitioned Chinook salmon from fresh- to seawater at current and future conditions of temperature (13 °C and 16 °C, respectively) and ocean acidification (400 and 1400 atm CO 2 ), including a fluctuating CO 2 treatment (between control and high CO 2 ) that may be more representative of natural environmental conditions associated with upwelling and tidal cycling. We hypothesized that constant elevated CO 2 levels would impair smoltification success immediately following seawater transfer, but that fluctuating conditions would be even more physiologically challenging. We predicted that elevated temperatures would exacerbate these effects. To test this, we measured plasma ion concentrations, gill Na + /K + -ATPase (NKA) isoform mRNA and protein expression, as well as condition indices in freshwater and following 1, 3, 6, and 18 days in seawater at the respective treatments. We confirmed the existence of gill freshwater and seawater isoforms of NKA (α1a and α1b, respectively) in Chinook salmon for the first time, and found an upregulation of both isoforms in the fluctuating CO 2 treatment but a reduction of the number of NKA α1b cells 3-days post seawater transfer at 13 °C. At 16 °C, NKA α1b was upregulated in high CO 2 levels, with an elevated hematocrit indicating fish were likely stressed. Taken together, plasma ions, gill NKA and condition indices revealed a complex response to interacting warming and acidification during the first few days in seawater, however there were no longer-term adverse physiological responses. Thus, Chinook salmon appear to be relatively resilient to near-future climate change., Competing Interests: Declaration of competing interest The authors declare no competing interests., (Copyright © 2024 The Authors. Published by Elsevier B.V. All rights reserved.)

This study examined the effects of solar light driven plastic degradation on carbon chemistry in the coastal waters of eastern Arabian Sea along the west coast of India. The research was conducted through experimental incubations exposed to natural sunlight at multiple locations between December 2023-February 2024. Photodegradation induced a significant pH decrease (up to 0.38 ± 0.02) between controls and plastic incubations ranging from 8.17 ± 0.01 to 7.54 ± 0.02 with the highest variation in the Mumbai coast ranging from 8.13 ± 0.01 to 7.75 ± 0.03. pH variations are primarily caused by the leaching of organic acids and CO 2 release during solar irradiated incubation. Plastic leaching due to natural light irradiation and subsequent changes in the water chemistry is of prime significance with dissolved organic carbon (DOC) leaching of 0.002-0.03% of plastic weight into the coastal waters. Our estimations suggest 15-75 metric tonnes (MT) of DOC release per year by plastic pollution in the eastern Arabian Sea coastal waters. Further, the fluorescent dissolved organic matter (FDOM) fragmentation, a part of DOC, may act as an organic source of synthetic contaminants and would promote heterotrophic microbial action in the coastal waters. Photodegradation of plastic and the interaction of natural DOC and plastic-derived DOC resulted in longer wavelengths FDOM, which may affect the penetration of photosynthetically active radiation in the water column, thereby impacting primary production. Finally, future research work focussing on the role of plastic pollution in coastal ocean acidification and vice-versa is essential and will be increasingly intense in the upcoming decades., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier Ltd. All rights reserved.)

Our study explores genomic signs of adaptation in A. lixula to different water pH conditions. To achieve this, we analysed the genomics variation of A. lixula individuals living across a natural pH gradient in Canary Islands, Spain. We use a 2b-RADseq protocol with 74 samples from sites with varying pH levels (from 7.3 to 7.9 during low tide) and included a control site. We identified 14,883 SNPs, with 432 identified as candidate SNPs under selection to pH variations through redundancy analysis. While all SNPs indicated genomic homogeneity, the 432 candidate SNPs under selection displayed genomic differences among sites and along the pH gradient. Out of these 432 loci, 17 were annotated using published A. lixula transcriptomes, involved in biological functions such as growth. Therefore, our findings suggest local adaptation in A. lixula populations to acidification in CO 2 vents, even over short distances of 75 m, underscoring their potential resistance to future Ocean Acidification., Competing Interests: Declaration of competing interest The author(s) declare no competing interests., (Copyright © 2024 The Author(s). Published by Elsevier Ltd.. All rights reserved.)

Submarine groundwater discharge is a pivotal factor in modifying the structure of phytoplankton communities in coastal waters. The objective of the study was to investigate how variations in nutrient concentrations and ratios influence the composition of phytoplankton communities along the coastal waters of Bay of Bengal. The experiment involved mixing groundwater with coastal water at 5 % and 10 % proportions. Phytoplankton growth was more pronounced in 10 % groundwater than those with 5 % and control samples. In control samples, Chl-a and other pigments, experienced decrease from 20 % to 80 %, except in Odisha-Paradeep and Visakhapatnam-Andhra Pradesh, where peridinin concentrations increased by 60 % to 65 % owing to low Si:N ratios below 0.2. A shift was observed from diatoms to dinoflagellates due to low Si: N ratios. The results reaffirm the hypothesis that variations in nutrient concentrations and ratios play a substantial role in shaping the composition of phytoplankton in the adjacent coastal waters., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the research reported in this paper., (Copyright © 2024 Elsevier Ltd. All rights reserved.)

The combined effects of the novel antifouling biocide tralopyril (TP) nitrile and ocean acidification (OA) on marine organisms are still not well understood, despite the increasing attention given to the toxic effects of emerging pollutants and OA on marine organisms in recent years. In this study, Crassostrea gigas (C. gigas) was exposed to TP, OA, and a combination of TP and OA for 21 days with a 14-day depuration. This study investigated the inter-tissue variability in energy metabolism responses and the impacts on gonadal development in C. gigas under both single and combined exposures to TP and OA. The results indicate that TP exposure and OA resulted in up-regulation of energy metabolism genes in the C. gigas, with tissues exhibiting enhanced aerobic metabolism. Furthermore, OA influences the sex determination of C. gigas, promoting the development of female individuals. Moreover, following depuration, C. gigas is able to restore normal energy metabolism and sexual development through the accumulation of suitable energy reserves. This study provides a valuable reference for the environmental and ecological risk assessment of TP, addressing the research gap in understanding the combined toxicity of TP and OA on aquatic organisms., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier Ltd. All rights reserved.)

Ocean acidification (OA) and other environmental factors can collectively affect marine organisms. Deltamethrin (DM), a type II pyrethroid insecticide, has been widely detected in coastal and estuarine areas, while little attention has been given to the combined effects of DM and OA. In this study, Haliotis discus hannai was exposed to three pH levels (8.1, 7.7 and 7.4) and three DM nominal concentrations (0 μg/L, 0.6 μg/L and 6 μg/L) for 14 and 28 days. The results indicated that experimental acidification and/or DM exposure led to impaired immune function and pathological damage. Additionally, acidified conditions and DM exposure induced oxidative stress, and gills are more sensitive than digestive glands. With increasing pCO 2 and DM nominal concentrations, superoxide dismutase (SOD) activity decreased, whereas catalase (CAT) and glutathione S-transferase (GST) activities increased in the gills. Moreover, the expression levels of Toll-like receptor (TLR) pathway-related genes were upregulated after exposure. Integrated biomarker response (IBR) analysis proved that acidified conditions and/or DM detrimentally affected the overall fitness of H. discus hannai, and co-exposure to experimental acidification and DM was the most stressful condition. This study emphasizes the necessity of incorporating OA in future pollutant environmental assessments to better elucidate the risks of environmental disturbance., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier B.V. All rights reserved.)

Ocean acidification is one of the major global environmental problems facing humankind today, and it has far-reaching impacts on marine organisms and the entire marine ecosystem. Zostera japonica, an important supporting species of intertidal seagrass beds, exhibits high photosynthetic productivity and plays an important role in the carbon cycle of nearshore waters. However, little is known about the characteristics, processes, and mechanisms of its response to ocean acidification. In this study, we conducted a 120-day acidification experiment in Z. japonica; here, plants underwent four leaf regeneration cycles to reveal the response mechanism of Z. japonica to ocean acidification (OA). We found that acidification significantly affected the seedling stage of Z. japonica, impacting leaf regeneration cycles by altering physiological and molecular responses. In one leaf regeneration cycle, the short-term exposure to CO 2 affected the seagrass parameters, such as the regulation of inorganic carbon uptake modes and the regulation of photosynthesis between the dark and light reactions, with the potential to affect the carbon sinks of the marine organisms. The long-term effects on the regulation of antioxidant enzymes and antioxidant metabolites, caused an improvement in the marine life adaptation to OA. In a comparison of the different leaf regeneration cycles, the response pattern of Z. japonica showed an offset of the acidification during the short cycles and an adaption to the acidification during the long cycles. This study revealed the response mechanism of Z. japonica to OA at different time scales and could provide a theoretical basis for accurately assessing the impact of OA on seagrass and the entire seagrass bed ecosystem., Competing Interests: Declaration of competing interest The authors declare no competing interest and no generative AI in scientific writing., (Copyright © 2024 Elsevier B.V. All rights reserved.)

Chitin-synthase (CHS) is found in most eukaryotes and has a complex evolutionary history. Research into CHS has mainly been in the context of biomineralization of mollusc shells an area of high interest due to the consequences of ocean acidification. Exploration of CHS at the genomic level in molluscs, the evolution of isoforms, their tissue distribution, and response to environmental challenges are largely unknown. Exploiting the extensive molecular resources for mollusc species it is revealed that bivalves possess the largest number of CHS genes (12-22) reported to date in eukaryotes. The evolutionary tree constructed at the class level of molluscs indicates four CHS Type II isoforms (A-D) probably existed in the most recent common ancestor, and Type II-A (Type II-A-1/Type II-A-2) and Type II-C (Type II-C-1/Type II-C-2) underwent further differentiation. Non-specific loss of CHS isoforms occurred at the class level, and in some Type II (B-D groups) isoforms the myosin head domain, which is associated with shell formation, was not preserved and highly species-specific tissue expression of CHS isoforms occurred. These observations strongly support the idea of CHS functional diversification with shell biomineralization being one of several important functions. Analysis of transcriptome data uncovered the species-specific potential of CHS isoforms in shell formation and a species-specific response to ocean acidification (OA). The impact of OA was not CHS isoform-dependent although in Mytilus, Type I-B and Type II-D gene expression was down-regulated in both M. galloprovincialis and M. coruscus. In summary, during CHS evolution the gene family expanded in bivalves generating a large diversity of isoforms with different structures and with a ubiquitous tissue distribution suggesting that chitin is involved in many biological functions. These findings provide insight into CHS evolution in molluscs and lay the foundation for research into their function and response to environmental changes., (Copyright © 2024 The Author(s). Published by Elsevier Inc. All rights reserved.)

Polar marine invertebrates serve as bellwethers for species vulnerabilities in the face of changing climate at high latitudes of the Earth. Ocean acidification, warming/heatwaves, freshening, sea ice retreat and productivity change are challenges for polar species. Adaptations to life in cold water with intensely seasonal productivity has shaped species traits at both poles. Polar species have life histories often characterised as K-strategist or K-selected (e.g. slow growth and development, larval hypometabolism) that make them sensitive to climate stress and altered seasonal productivity. Moderate warming results in faster development and can have positive effects on development, up to a limit. However, ocean acidification can retard development, impair skeletogenesis and result in smaller larvae. Given the fast pace of warming, data on the thermal tolerance of larvae from diverse species is urgently needed, as well as knowledge of adaptive responses to ocean acidification and changes to sea ice and productivity. Predicted productivity increase would benefit energy-limited reproduction and development, while sea ice loss negatively impacts species with reproduction that directly or indirectly depend on this habitat. It is critical to understand the interactive effects between warming, acidification and other stressors. Polar specialists cannot migrate, making them susceptible to competition and extinction from range-extending subpolar species. The borealisation and australisation of Arctic and Antarctic ecosystems, respectively, is underway as these regions become more hospitable for the larval and adult life-history stages of lower-latitude species. Differences in biogeography and pace of change point to different prospects for Arctic and Antarctic communities. In this Commentary, we hypothesise outcomes for polar species based on life history traits and sensitivity to climate change and suggest research avenues to test our predictions., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2024. Published by The Company of Biologists Ltd.)

The interplay of global climate change and anthropogenic activities has significantly affected the carbon cycle in coastal ocean environments. Consequently, further investigation into the carbonate system, carbon source and sink processes, and acidification mechanisms is essential. This study examined the surface carbonate system offshore of Qingdao, utilizing data from nine spring cruises in 2011-2019. The inter-annual variations in sea surface temperature revealed a gradual upward trend, whereas chlorophyll a (Chl-a) and apparent oxygen utilization (AOU) exhibited alternating trends of increase and decrease. Over the period from 2011 to 2019, pH levels ranged from 8.00 to 8.19, partial pressure of CO 2 (pCO 2 ) values varied between 187.8 and 364.1 μatm, and calcium carbonate saturation (Ω arag ) ranged from 3.22 to 4.79. From 2011 to 2017, both pH and Ω arag showed a decreasing trend, attributed to the increasing influence of respiration. In contrast, an upward trend was observed for both parameters during 2018 and 2019, indicating a decline in respiration's contribution. Biological processes, especially the increase in respiration caused by the nutrients and organic matter released through human activities, were pivotal in determining spatial variations in pCO 2 and Ω arag offshore, while the impact of mixing processes and temperature was comparatively negligible. Furthermore, the Ulva prolifera bloom event in July 2015 significantly affected the coastal waters of Qingdao's carbonate system when compared to the no-bloom period in March 2015. Overall, the study area functioned as a sink for atmospheric CO 2 in the spring. The fluctuations in air-sea CO 2 fluxes corresponded with changes in ΔpCO 2 and wind speeds, with surface seawater air-sea CO 2 fluxes ranging from -17.7 to 12.2 mmol m -2 d -1 between 2011 and 2019, yielding an average flux of -9.4 ± 5.3 mmol m -2 d -1 ., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier Ltd. All rights reserved.)

Molecular mechanisms driving species-specific environmental sensitivity in coccolithophores are unclear but crucial in understanding species selection and adaptation to environmental change. This study examined proteomic and physiological changes in three species under varying pH conditions. We showed that changing pH drives intracellular oxidative stress and changes membrane potential. Upregulation in antioxidant, DNA repair and cell cycle-related protein-groups indicated oxidative damage across high (pH 8.8) and low pH (pH 7.6) compared to control pH (pH 8.2), and correlated with reduced growth rates. Upregulation of mitochondrial proteins suggested higher metabolite demand for restoring cellular homeostasis under pH-induced stress. Photosynthetic rates generally correlated with CO 2 availability, driving higher net carbon fixation rates at low pH. The intracellular pH-buffering capacity of the coastal Chrysotila carterae and high metabolic adaptability in the bloom-forming Gephyrocapsa huxleyi will likely facilitate their adaptation to ocean acidification or artificial ocean alkalinisation. However, the pH sensitivity of the ancient open-ocean Coccolithus braarudii will possibly result in reduced growth and shrinking of its ecological niche., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Rising carbon dioxide emissions are provoking ocean warming and acidification 1,2 , altering plankton habitats and threatening calcifying organisms 3 , such as the planktonic foraminifera (PF). Whether the PF can cope with these unprecedented rates of environmental change, through lateral migrations and vertical displacements, is unresolved. Here we show, using data collected over the course of a century as FORCIS 4 global census counts, that the PF are displaying evident poleward migratory behaviours, increasing their diversity at mid- to high latitudes and, for some species, descending in the water column. Overall foraminiferal abundances have decreased by 24.2 ± 0.1% over the past eight decades. Beyond lateral migrations 5 , our study has uncovered intricate vertical migration patterns among foraminiferal species, presenting a nuanced understanding of their adaptive strategies. In the temperature and calcite saturation states projected for 2050 and 2100, low-latitude foraminiferal species will face physicochemical environments that surpass their current ecological tolerances. These species may replace higher-latitude species through poleward shifts, which would reduce low-latitude foraminiferal diversity. Our insights into the adaptation of foraminifera during the Anthropocene suggest that migration will not be enough to ensure survival. This underscores the urgent need for us to understand how the interplay of climate change, ocean acidification and other stressors will impact the survivability of large parts of the marine realm., Competing Interests: Competing interests: The authors declare no competing interests., (© 2024. The Author(s).)

Ocean acidification and warming are significant stressors impacting marine ecosystems, exerting profound effects on the physiological ecology of marine organisms. We investigated the impact of ocean acidification and warming on the immune system of mussels, focusing on the regulatory mechanisms of intrinsic and extrinsic apoptosis. The study explored the effects on the immune response ability of mussels (Mytilus coruscus) after 14 and 21 days under combined conditions of different temperatures (20 °C and 30 °C) and pH (8.1 and 7.7), as expected for the year 2100. The experimental results indicated that ocean acidification and warming have significant interactive effects on various immune parameters of M. coruscus. Specifically, ocean acidification and warming lead to an increase in ROS (Reactive Oxygen Species), apoptosis, TNF-α (Tumor Necrosis Factor-alpha), TGF-β (Transforming Growth Factor-beta), Caspase-8, and a decrease in IL-17 (Interleukin 17). These findings suggest that ocean acidification and warming trigger an immune inflammatory response in mussels. Regulating immune functions through apoptosis pathways may be a crucial coping mechanism in response to environmental variations, but its long-term impact on population health and sustainability remains uncertain. Our findings offer important insights into the complex interactions between bivalve immune responses and environmental stressors. This also underscores the need for further research into the adaptive capabilities of marine organisms facing the compounded challenges of ocean acidification and warming., (Copyright © 2025 Elsevier Ltd. All rights reserved.)

The responses of seaweed species to increased CO 2 and lowered pH (Ocean Acidification: OA) depend on their carbon concentrating mechanisms (CCMs) and inorganic carbon (Ci) preferences. However, few studies have described these mechanisms in the early life stages of seaweeds or assessed the effects of OA and its interactions with other environmental drivers on their functionality and photophysiology. Our study evaluated the effects of pH, light (PAR), temperature, and their interactions on the Ci uptake strategies and photophysiology in the early stages of Macrocystis pyrifera . Gametophytes were cultivated under varying pH (7.80 and 8.20), light (20 and 50 µmol photons m -2 s -1 ), and temperature (12 and 16 °C) conditions for 25 days. We assessed photophysiological responses and CCMs (in particular, the extracellular dehydration of HCO 3 - to CO 2 mediated by the enzyme carbonic anhydrase (CA) and direct HCO 3 - uptake via an anion exchange port). This study is the first to describe the Ci uptake strategies in gametophytes of M. pyrifera , demonstrating that their primary CCM is the extracellular conversion of HCO 3 - to CO 2 mediated by CA. Additionally, our results indicate that decreased pH can positively affect their photosynthetic efficiency and maximum quantum yield; however, this response is dependent on the light and temperature conditions.

The escalating impacts of climate change, particularly ocean acidification and warming, are pivotal stressors for marine ecosystems and have profound effects on biota in polar regions. This study investigated the immunological responses of the Antarctic fish Notothenia rossii to environmental stressors indicative of future ocean conditions under the Intergovernmental Panel on Climate Change Shared Socioeconomic Pathways 5-8.5 scenario for 2100. We exposed N. rossii to conditions simulating present-day conditions: control, elevated temperature, acidification, and both stressors combined over six days. Utilizing RNA-Seq for comprehensive gene expression analysis, we identified significant upregulation and downregulation of immune-related pathways, highlighting a complex interplay of genes involved in complement and coagulation cascades, the intestinal immune network for immunoglobulin A production, cytosolic DNA sensing, natural killer cell-mediated cytotoxicity, and Interleukin 17 signaling pathways. Our findings revealed a predominantly antagonistic gene expression response, suggesting an intricate balance between energy allocation for maintaining homeostasis and the capacity of the immune system to combat stressors. This reflects a potential adaptive mechanism to combined environmental stressors, underscoring the complexity of immune responses in N. rossii and suggesting both potential vulnerabilities and resilience in the face of climate change. This study provides critical insights into the immunological impacts of acidification and warming on Antarctic marine species, emphasizing the need for further research to unravel the mechanisms underlying these observed changes and inform conservation strategies for polar ecosystems in a changing global climate., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier Inc. All rights reserved.)

A multi-generational experiment (F1-F4) was conducted for a marine copepod Tigriopus japonicus to investigate its physiological and molecular responses to mercury (Hg) pollution and/or its combination with ocean acidification (OA) plus ocean warming (OW). The projected future scenario, i.e., OA plus OW (AW) significantly increased methylmercury accumulation in copepods by 1.14 times, despite insignificant change for total Hg bioaccumulation. Transcriptomic analysis indicated that copepods initiated several detoxification defense processes, including reactive oxygen species metabolic process, glutathione metabolism, and protein refolding, in response to increased Hg toxicity under combined exposure of AW and Hg; meanwhile, inhibited energy metabolism was observed in this case, linking to reduced number of nauplii/clutch but accelerated development in copepods probably due to an energetic trade-off. Increased Hg toxicity due to AW could also be ascribed to the impairment in immune defense (e.g., lysosome and vitamin metabolism) and reproduction-related processes (e.g., growth factor activity). Collectively, this study reveals the multi-generational response mechanism of copepods to Hg pollution under global change, emphasizing an exacerbated adverse effect of Hg, and it provides a scientific basis for an accurate understanding of the potential impact of Hg pollution on marine ecosystems., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier B.V. All rights reserved.)

Coral reefs are among the most sensitive ecosystems affected by ocean warming and acidification, and are predicted to collapse over the next few decades. Reefs are predicted to shift from net accreting calcifier-dominated systems with exceptionally high biodiversity to net eroding algal-dominated systems with dramatically reduced biodiversity. Here, we present a two-year experimental study examining the responses of entire mesocosm coral reef communities to warming (+2 °C), acidification (-0.2 pH units), and combined future ocean (+2 °C, -0.2 pH) treatments. Contrary to modeled projections, we show that under future ocean conditions, these communities shift structure and composition yet persist as novel calcifying ecosystems with high biodiversity. Our results suggest that if climate change is limited to Paris Climate Agreement targets, coral reefs could persist in an altered state rather than collapse., Competing Interests: Competing interests statement:The authors declare no competing interest.

In an era of unprecedented environmental changes, understanding the combined effects of multiple stressors on species' performance is urgent. The increasing UV-filter incorporation in daily-life products raises concerns about their potential impact on marine-coastal environments upon release. As stressors rarely act alone, global change-induced factors, such as ocean acidification (OA), can amplify ecological hazards promoted by contaminants in coastal realms. This study investigated the combined impacts of UV-filters 4-methylbenzylidene camphor (4-MBC) and benzophenone-3 (BP-3), at ecologically relevant concentrations (1 and 10 µg/L), under two target pH levels (8.2 and 7.7, reflecting a ∆pH of 0 and -0.3 relative to the average pH at the sampling site), on the biological performance and male reproductive health of the mussel Mytilus galloprovincialis. Using sperm and adult assays alongside a multi-biomarker approach, the study revealed that pH was the primary driver of the decline in mussel physiological and biochemical performances, further intensifying UV-filters' impacts. While sperm cells showed adaptive responses to low pH conditions alone, characterized by reduced lipid peroxidation (LPO) levels and superoxide anion overproduction, adult mussels experienced more pronounced effects, particularly under simultaneous exposure to low pH and UV-filters. Specifically, the adults exhibited distinct bioconcentration patterns under low pH, enhanced cellular metabolic activity and energy-demand compensatory processes, activation of biotransformation pathways, and regulation of antioxidant defenses. Given the ecological and socio-economic importance of M. galloprovincialis and its demonstrated vulnerability to these stressors, these findings highlight the need for further studies on potential transgenerational impacts and evolutionary implications for mussel populations., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier B.V. All rights reserved.)

Ocean acidification (OA) can affect marine bivalves at various levels of biological organization. Yet, little effort has been devoted to understanding how OA affects the reproductive events of marine bivalves during multiple cycles of maturation. Here, we tested sex-specific reproductive responses of Manila clams (Ruditapes philippinarum) to OA during gonadal rematuration. Under acidified conditions, both male and female clams exhibited delayed gonadal rematuration following spawning and impairments in gonadal tissues, which can be likely ascribed to lowered concentrations of hormones and vitellogenin. The findings indicate that marine bivalves experience significant declines in reproductive capacity as a result of OA during their reproductive cycles, with clear sex-specific differences. Consequently, it is essential to consider sex-specific reproduction responses of marine bivalves to OA when developing conservation strategies and forecasting population sustainability in a rapidly acidifying marine environment., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier Ltd. All rights reserved.)

The global ocean is rapidly changing, posing a substantial threat to the viability of marine populations due to the co-occurrence of multiple drivers, such as ocean warming (OW) and ocean acidification (OA). To persist, marine species must undergo some combination of acclimation and adaptation in response to these changes. Understanding such responses is essential to measure and project the magnitude and direction of current and future vulnerabilities in marine ecosystems. Echinoderms have been recognised as a model in studies of OW-OA effects on marine biota. However, despite their global diversity, vulnerability and ecological importance in most marine habitats, brittle stars (ophiuroids) are poorly studied. A long-term mesocosm experiment was conducted on adult mottled brittle star (Ophionereis fasciata) as a case study to investigate the physiological response and trade-offs of marine organisms to ocean acidification, ocean warming and the combined effect of these two drivers. Long-term exposure of O. fasciata to high temperature and low pH affected survival, respiration and regeneration rates, growth rate, calcification/dissolution and righting response. Higher temperatures increased stress and respiration, and decreased regeneration and growth rates as well as survival. Conversely, changes in pH had more subtle or no effect, affecting only respiration and calcification. Our results indicate that exposure to a combination of high temperature and low pH produces complex responses for respiration, righting response and calcification. We address the knowledge gap of the impact of a changing ocean on ophiuroids in the context of echinoderm studies, proposing this class as an ideal alternative echinoderm for future research., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2024. Published by The Company of Biologists Ltd.)