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Micro-/nano-plastic particles (MNPs) are present in the ocean with potential detrimental impacts on marine ecosystems. Bivalves are often used as marine bioindicators and are ideal to evaluate the threat posed by various-sized MNPs. We exposed the mussel Mytilus coruscus to MNPs with different particle sizes (70 and 500 nm, 5, 10 and 100 μm) for 3, 72 h and 30 days. The antioxidant responses in digestive gland and the hemolymph were then evaluated. The time of exposure played a strong modulating role in the biological response. A 3-hour exposure had no significant impact on the digestive gland. After 72 h, an increase in oxidative stress was observed in the digestive gland, including increased hydrogen peroxide (H 2 O 2 ) level, catalase (CAT), glutathione peroxidase (GPx) activities and malondialdehyde (MDA) production. After a 30-day exposure, the oxidative stress decreased while lipid peroxidation increased. A 30-day exposure increased hemocyte mortality (HM) and reactive oxygen species (ROS) levels in the hemolymph, while phagocytosis (PA), lysosome content (LC), mitochondrial number (MN) and mitochondrial membrane potential (MMP) significantly decreased. Longer-term exposure to MNPs caused oxidative stress in the digestive gland as well as impaired viability and immunity of hemocytes. Particle size also influenced the response with smaller particles having more severe effects. A depuration for 7 days was enough to reverse the negative effects observed on the digestive gland and hemolymph. This study provides new insights on the effects of small-sized MNPs, especially nanoplastic particles (NPs), on aquatic organisms, and provides a solid theoretical knowledge background for future studies on toxic effects of MNPs., 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.)

The prevalence and frequency of hypoxia events have increased worldwide over the past decade as a consequence of global climate change and coastal biological oxygen depletions. On the other hand, anthropogenic emissions of CO 2 and consequent accumulation in the sea surface result in a perturbation of the seawater carbonate system, including a decrease in pH, known as ocean acidification. While the effect of decreases in pH and dissolved oxygen (DO) concentration is better understood, their combined effects are still poorly resolved. Here, we exposed adult mussels (Mytilus galloprovincialis) to two pHs (8.27 and 7.63) and DO concentrations (7.65 and 2.75 mg L -1 ) over 17 days in a full-factorial design. These levels correspond to extremes of the present natural variability and are relevant in the context of ocean acidification and hypoxia. No mortality was observed during the experiment. However, sublethal effects were observed for clearance and oxygen consumption rates, as well as total haemocytes count and haemocytes viability and gene expression in mussels exposed to the combination of low pH and low DO. Respiration and excretion rates were not significantly impacted by low pH and DO, alone or in combination. Overall, low pH alone led to a decrease in all tested physiological parameters while low DO alone led to a decline in clearance rate, haemocyte parameters and an increase in carbohydrate content. Both parameters led to up- or down-regulation of most of the selected genes. Not surprisingly, the combined effect of low pH and low DO could not be predicted by a simple arithmetic additive response at the effect level, highlighting more complex and non-linear effects., 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 © 2023 Elsevier Ltd. All rights reserved.)

This study investigated the combined effects of seawater acidification and hypoxia on the antioxidant response in gonads of the thick shell mussel Mytilus coruscus. Mussels were collected along the Shengsi Island, East China Sea, where oxygen and pH fluctuations frequently occur in summer. Mussels were exposed to three pH (8.1, 7.7, and 7.3) and two dissolved oxygen (DO) levels (6 and 2 mg L -1 ) for 21 days followed by a 10-day recovery period (pH 8.1 and DO 6 mg L -1 ). Gonad surface area (GSA) and activity of superoxide dismutase (SOD), catalase (CAT), glutathione peroxidase (GPX), glutathione (GSH), glutathione S-transferase (GST), and malondialdehyde (MDA) in gonad were measured at days 21 and 31. Complex and enzyme-specific responses were observed after the 21-day exposure period. Overall, PCA analysis revealed a stronger effect of pH than DO. Integrated biomarker response (IBR) analysis demonstrated that low pH and DO decreased mussel's antioxidant system and increased oxidative damage with potential consequences for gonad development. Mussels exposed to low pH and DO were only partly able to recover a normal enzymatic activity after 10-day recovery period. This suggests that mussels exposed to short-term pH and DO fluctuations event in the field may suffer lasting negative impacts., (© 2021. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Ocean acidification and hypoxia have become increasingly severe in coastal areas, and their co-occurrence poses emerging threats to coastal ecosystems. Here, we investigated the combined effects of ocean acidification and hypoxia on the reproductive capacity of the thick-shelled mussel Mytilus coruscus. Our results demonstrated low pH but not low oxygen induced decreased gonadosomatic index (GSI) in mussels. Male mussels had a lower level of sex steroids (estradiol, testosterone, and progesterone) when kept at low pH. Expression of genes related to reproduction were also impacted by low pH with a downregulation of genes involved in gonad development in males (β-catenin and Wnt-7b involved in males) and an upregulation of testosterone synthesis inhibition-related gene (Wnt-4) in females. Overall, our results suggest that ocean acidification has an impact on the gonadal development through an alternation of gene expression and level of steroids while hypoxia had no significant effect., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

In the present study, the combined effects of pH, dissolved oxygen (DO) and temperature levels on the antioxidant responses of the mussel Mytilus coruscus were evaluated. Mussels were exposed to two pH (8.1, 7.7-acidification), two DO (6 mg L -1 , 2 mg L -1 -hypoxia) and two temperature levels (20 °C, 30 °C-warming) for 30 days. SOD, CAT, MDA, GPx, GSH, GST, TAOC, AKP, ACP, GPT, AST levels were measured in the gills of mussels. All tested biochemical parameters were altered by these three environmental stressors. Values for all the test parameters except GSH first increased and then decreased at various experimental treatments during days 15 and 30 as a result of acidification, hypoxia and warming. GSH content always increased with decreased pH, decreased DO and increased temperature. PCA showed a positive correlation among all the measured biochemical indexes. IBR results showed that M. coruscus were adversely affected by reduced pH, low DO and elevated temperature., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

The effects of short-term (7 days) experimental ocean acidification (-0.4 pH units) and warming (+5 °C) on anti-predator defenses of two sympatric Mytilus species from China, M. coruscus and M. edulis, in the presence and absence of predator cues were investigated. Results suggested species-specific independent negative effects of acidification and warming on the number and weight of byssal threads, the force of thread attachment, and total thread plaque area. Similar negative effects were observed for clustering behaviour, with acidification and warming independently increasing the number of solitary individuals and decreasing the percentage of mussels in clusters. Acidification effects on byssus were strongly exacerbated when predators were present. Ultimately, this study suggests that short-term exposure to experimental warming and acidification can negatively impact anti-predator defense strategies in mussels with potential ramifications for predator-prey interactions and ecological functioning in systems where mussel beds play a key ecological role., (Copyright © 2019. Published by Elsevier Ltd.)

Biochemical responses of the mussel Mytilus coruscus exposed to different concentrations of titanium dioxide nanoparticles (nano-TiO 2 ) (0, 2.5, 10 mg L -1 ) and two pH levels (pH 8.1 and pH 7.3) for 14 days. Mussel responses were also investigated after a 7 days recovery period (pH 8.1 and no nanoparticle). Exposure to nano-TiO 2 led changes in antioxidant indexes (superoxide dismutase (SOD), catalase (CAT), glutathione peroxidase (GPx), glutathione (GSH)), biotransformation enzyme activity (GST) and malondialdehyde level (MDA) in gills and digestive glands. An increase in MDA level and a decrease in SOD and GSH activities were observed in gill of mussels exposed to 10 mg L -1 nano-TiO 2 . This effect was more severe in mussels kept at pH 7.3 as compared to pH 8.1. A different response was observed in the digestive gland as SOD, CAT and GSH levels increased in mussels exposed to nano-TiO 2 . These contrasting results in digestive glands and gills were only evident at high concentration of nano-TiO 2 and low pH. A 7 days recovery period was not sufficient to fully restore SOD, GPx, GST, GSH and MDA levels to levels before exposure to nano-TiO 2 and low pH. Overall, our results confirmed that seawater acidification modulates effects of nanoparticles in mussels, and that gills are more sensitive to these stressors as compared with digestive glands., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

Increased production of engineered nanoparticles (NPs) has raised extensive concerns about the potential toxic effects on marine organisms. Extensive evidences documented the impact of ocean acidification (OA) on the physiology and fitness of bivalves. In the present study, we investigated the biochemical responses of the mussel Mytilus coruscus exposed to both nano-ZnO and low pH relevant for ocean acidification conditions for 14 d followed by a 7-d recovery period. Most biochemical indexes (superoxide dismutase (SOD), catalase (CAT), glutathione peroxidase (GPx), acid phosphatase (ACP) and alkaline phosphatase (ALP)) measured in gills and hemocytes were increased when the mussels were subject to low pH or high concentration of nano-ZnO, suggesting oxidative stress responses. No significant interactions between the two stressors were observed for most measured parameters. After a 1 week recovery period, low pH and nano-ZnO had less marked impact for SOD, GPx, ACP and ALP in hemocytes as compared to the end of the 14 d exposure. However, no recovery was observed in gills. Overall, our results suggest that both low pH and nano-ZnO induce an anti-oxidative response in Mytilus coruscus with gills being more sensitive than hemocytes., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

Ocean acidification (OA) and hypoxic events are increasing worldwide problems, their interactive effects have not been well clarified, although their co-occurrence is prevalent. The East China Sea (the Yangtze River estuary area) suffers from not only coastal hypoxia but also pH fluctuation, representing an ideal study site to explore the combined effect of OA and hypoxia on marine bivalves. We experimentally evaluated the antioxidant response of the mussel Mytilus coruscus exposed to three pH levels (8.1, 7.7 and 7.3) at two dissolved oxygen (DO) levels (2.0mgL -1 and 6.0mgL -1 ) for 72h. Activities of superoxide dismutase, catalase, glutathione peroxidase, acid phosphatase, and alkaline phosphatase and levels of malondialdehyde were measured in gills and hemolymph. All enzymatic activities in hemolymph and gills followed a similar pattern throughout the experiment duration. Generally, low DO showed greater effects on enzyme activities than elevated CO 2 . Significant interactions between DO, pH and time were only observed at superoxide dismutase and catalase in both tissues. PCA revealed positive relationships between most enzyme activities in both gills and hemolymph with the exception of alkaline phosphatase activity and the level of malondialdehyde in the hemolymph. Overall, our results suggested that decreased pH and low DO induced similar antioxidant responses in the hard shelled mussel, and showed an additive effect on most enzyme activities. The evaluation of multiple environmental stressors, a more realistic scenario than single ones, is crucial to predict the effect of future global changes on coastal species and our results supply some insights on the potential combined effects of reduced pH and DO on marine bivalves., (Copyright © 2016 Elsevier Inc. All rights reserved.)

Hypoxia and ocean acidification are two consequences of anthropogenic activities. These global trends occur on top of natural variability. In environments such as estuarine areas, short-term acute pH and O2 fluctuations are occurring simultaneously. The present study tested the combined effects of short-term seawater acidification and hypoxia on the physiology and energy budget of the thick shell mussel Mytilus coruscus. Mussels were exposed for 72 h to six combined treatments with three pH levels (8.1, 7.7 and 7.3) and two dissolved oxygen (DO) levels (2 mg L(-1), 6 mg L(-1)). Clearance rate (CR), food absorption efficiency (AE), respiration rate (RR), ammonium excretion rate (ER), O:N ratio and scope for growth (SFG) were significantly reduced, and faecal organic dry weight ratio (E) was significantly increased at low DO. Low pH did not lead to a reduced SFG. Interactive effects of pH and DO were observed for CR, E and RR. Principal component analysis (PCA) revealed positive relationships among most physiological indicators, especially between SFG and CR under normal DO conditions. These results demonstrate that Mytilus coruscus was sensitive to short-term (72 h) exposure to decreased O2 especially if combined with decreased pH levels. In conclusion, the short-term oxygen and pH variation significantly induced physiological changes of mussels with some interactive effects., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

This study evaluated the combined effects of seawater pH decrease and temperature increase on the activity of antioxidant enzymes in the thick shell mussel Mytilus coruscus, an ecological and economic bivalve species widely distributed along the East China Sea. Mussels were exposed to three pH levels (8.1, 7.7 and 7.3) and two temperatures (25 °C and 30 °C) for 14 days. Activities of superoxide dismutase (SOD), catalase (CAT), glutathione peroxidase (GPX), glutathione (GSH), acid phosphatase (ACP), alkaline phosphatase (AKP) and glutamic-pyruvic transaminase (GPT) were measured in gills and digestive glands after 1, 3, 7 and 14 days of exposure. All enzymatic activities were significantly impacted by pH, temperature. Enzymatic activities at the high temperature were significantly higher than those at the low temperature, and the mussels exposed to pH 7.3 showed significantly higher activities than those under higher pH condition for all enzymes except ACP. There was no interaction between temperature and pH in two third of the measured activities suggesting similar mode of action for both drivers. Interaction was only consistently significant for GPX. PCA revealed positive relationships between the measured biochemical indicators in both gills and digestive glands. Overall, our results suggest that decreased pH and increased temperature induce a similar anti-oxidative response in the thick shell mussel., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

Most molluscs possess shells, constructed from a vast array of microstructures and architectures. The fully formed shell is composed of calcite or aragonite. These CaCO3 crystals form complex biocomposites with proteins, which although typically less than 5% of total shell mass, play significant roles in determining shell microstructure. Despite much research effort, large knowledge gaps remain in how molluscs construct and maintain their shells, and how they produce such a great diversity of forms. Here we synthesize results on how shell shape, microstructure, composition and organic content vary among, and within, species in response to numerous biotic and abiotic factors. At the local level, temperature, food supply and predation cues significantly affect shell morphology, whilst salinity has a much stronger influence across latitudes. Moreover, we emphasize how advances in genomic technologies [e.g. restriction site‐associated DNA sequencing (RAD‐Seq) and epigenetics] allow detailed examinations of whether morphological changes result from phenotypic plasticity or genetic adaptation, or a combination of these. RAD‐Seq has already identified single nucleotide polymorphisms associated with temperature and aquaculture practices, whilst epigenetic processes have been shown significantly to modify shell construction to local conditions in, for example, Antarctica and New Zealand. We also synthesize results on the costs of shell construction and explore how these affect energetic trade‐offs in animal metabolism. The cellular costs are still debated, with CaCO3 precipitation estimates ranging from 1–2 J/mg to 17–55 J/mg depending on experimental and environmental conditions. However, organic components are more expensive (~29 J/mg) and recent data indicate transmembrane calcium ion transporters can involve considerable costs. This review emphasizes the role that molecular analyses have played in demonstrating multiple evolutionary origins of biomineralization genes. Although these are characterized by lineage‐specific proteins and unique combinations of co‐opted genes, a small set of protein domains have been identified as a conserved biomineralization tool box. We further highlight the use of sequence data sets in providing candidate genes for in situ localization and protein function studies. The former has elucidated gene expression modularity in mantle tissue, improving understanding of the diversity of shell morphology synthesis. RNA interference (RNAi) and clustered regularly interspersed short palindromic repeats ‐ CRISPR‐associated protein 9 (CRISPR‐Cas9) experiments have provided proof of concept for use in the functional investigation of mollusc gene sequences, showing for example that Pif (aragonite‐binding) protein plays a significant role in structured nacre crystal growth and that the Lsdia1 gene sets shell chirality in Lymnaea stagnalis. Much research has focused on the impacts of ocean acidification on molluscs. Initial studies were predominantly pessimistic for future molluscan biodiversity. However, more sophisticated experiments incorporating selective breeding and multiple generations are identifying subtle effects and that variability within mollusc genomes has potential for adaption to future conditions. Furthermore, we highlight recent historical studies based on museum collections that demonstrate a greater resilience of molluscs to climate change compared with experimental data. The future of mollusc research lies not solely with ecological investigations into biodiversity, and this review synthesizes knowledge across disciplines to understand biomineralization. It spans research ranging from evolution and development, through predictions of biodiversity prospects and future‐proofing of aquaculture to identifying new biomimetic opportunities and societal benefits from recycling shell products. [ABSTRACT FROM AUTHOR]

With the release of large amounts of CO2, ocean acidification is intensifying and affecting aquatic organisms. In addition, salinity also plays an important role for marine organisms and fluctuates greatly in estuarine and coastal ecosystem, where ocean acidification frequently occurs. In present study, flow cytometry was used to investigate immune parameters of haemocytes in the thick shellmusselMytilus coruscus exposed to different salinities (15, 25, and 35‰) and two pH levels (7.3 and 8.1). A 7-day in vivo and a 5-h in vitro experiments were performed. In both experiments, low pH had significant effects on all tested immune parameters. When exposed to decreased pH, total haemocyte count (THC), phagocytosis (Pha), esterase (Est), and lysosomal content (Lyso) were significantly decreased, whereas haemocyte mortality (HM) and reactive oxygen species (ROS) were increased. High salinity had no significant effects on the immune parameters of haemocytes as compared with low salinity. However, an interaction between pH and salinity was observed in both experiments for most tested haemocyte parameters. This study showed that high salinity, low salinity and low pH have negative and interactive effects on haemocytes of mussels. As a consequence, it can be expected that the combined effect of low pH and changed salinity will have more severe effects on mussel health than predicted by single exposure. [ABSTRACT FROM AUTHOR]

Startle response behaviours are important in predator avoidance and escape for a wide array of animals. For many marine invertebrates, however, startle response behaviours are understudied, and the effects of global change stressors on these responses are unknown. We exposed two size classes of blue mussels (Mytilus edulis × trossulus) to different combinations of temperature (15 and 19 °C) and pH (8.2 and 7.5 pH T) for 3 months and subsequently measured individual time to open following a tactile predator cue (i.e. startle response time) over a series of four consecutive trials. Time to open was highly repeatable in the short term and decreased linearly across the four trials. Individuals from the larger size class had a shorter time to open than their smaller-sized counterparts. High temperature increased time to open compared to low temperature, while pH had no effect. These results suggest that bivalve time to open is repeatable, related to relative vulnerability to predation and affected by temperature. Given that increased closure times impact feeding and respiration, the effect of temperature on closure duration may play a role in the sensitivity to ocean warming in this species and contribute to ecosystem level effects. • We measured startle response (time to open) in mussels following a predator cue. • We tested effects of temperature, pH and size and measured repeatability. • Larger mussels opened faster; repeatable startle response; evidence of habituation. • High temperature increased time to open; no effect of pH. • Blue mussels are sensitive to temperature and vulnerable to climate change. [ABSTRACT FROM AUTHOR]