Calcium carbonate formation in coastal macroalgal ecosystems via multiple pathways
Highlights
Focused on a novel inorganic carbon sequestration pathway of non-calcifying macroalgae.
Clarified algae-bacteria synergistic driving mechanism for calcium carbonate precipitation.
Systematically sorted out potential mineralizing microbial taxa within the phycosphere and their core metabolic pathways.
Abstract
Macroalgae are dominant primary producers that drive carbon sequestration in coastal ecosystems. Macroalgal carbon sequestration primarily refers to the long-term storage of macroalgae-derived organic carbon in the ocean. However, calcium carbonate (CaCO3) formation is frequently observed in non-calcifying macroalgal environments, suggesting the existence of an overlooked inorganic carbon process in macroalgal ecosystems. Here, we introduce multiple pathways that may drive CaCO3 formation in macroalgal ecosystems. These include the effects of macroalgal photosynthesis and carbon-concentrating mechanisms on the seawater carbonate system, the role of phycosphere interfacial properties in facilitating CaCO3 nucleation, and the macroalgae-bacteria synergy that promotes CaCO3 formation. We identified several current knowledge gaps—the unclear carbon sequestration or source effect of CaCO3 formation in macroalgal ecosystems and the stability of CaCO3 minerals in macroalgal ecosystems—that require further investigation. This review advances the understanding of macroalgal carbon cycling beyond organic pathways and emphasizes the importance of a comprehensive assessment of macroalgal carbon sequestration, including that of inorganic carbon.
table isotopes are fundamental proxies for deciphering past environmental conditions and carbon cycle perturbations. However, applying these geochemical tools with a forward-looking perspective can also help predict the resilience of marine biota in future acidified oceans. This study investigates carbon uptake and biomineralization pathways under reduced pH in two ecologically keystone calcifying species: the bivalve Mytilus galloprovincialis and the gastropod Phorcus sp. To unravel these mechanisms, we adopted a dual approach, comparing isotopic trajectories (δ13C) under strictly controlled laboratory mesocosms with observations from natural CO2 vent systems in Ischia, Italy. Our results reveal a striking discrepancy between settings. Under laboratory conditions, both species exhibited a significant depletion in shell δ13C as pH decreased, directly tracking seawater carbonate chemistry. Conversely, specimens from the volcanic vents displayed a paradoxical δ13C enrichment at low-pH sites (pH ~7.4). These divergent trajectories indicate that in complex natural environments, the geochemical signal of ocean acidification can be overridden by metabolic overprints and trophically driven vital processes. Consequently, while laboratory experiments are essential to isolate kinetic fractionation, natural analogues remain crucial to capture the biological resilience and complex ecological feedback of future oceans. Future research combining these geochemical trends with direct, in vivo physiological assessments will be essential to precisely constrain the underlying metabolic kinetics and refine predictive metabolic models.
Low salinity linked to warming may intensify coastal pH reduction.
Low salinity reduced righting and emergence in the limpet Patelloida pygmaea.
Low salinity reduces Mg/Ca ratios in newly formed shell layers.
Low pH increases shell thickness and Mg/Ca ratios despite dissolution.
CHS2 upregulation suggests compensatory resistance to low pH.
Abstract
Rising anthropogenic carbon dioxide emissions have driven ongoing ocean warming and associated climate changes. In the Yellow Sea, this warming is associated with enhanced monsoonal rainfall, which increases freshwater inputs and lowers coastal salinity. Increased freshwater input can weaken seawater buffering capacity, thereby leading to lower pH conditions in coastal environments. Here, we examined the effects of low pH and low salinity on the intertidal limpet Patelloida pygmaea. Adult limpets were exposed for 31 days to four experimental artificial seawater conditions combining two pH levels (8.0 and 7.5) and two salinity levels (30 and 21 psu). Survival and condition factor were not influenced by pH or salinity. However, low salinity reduced righting and emergence behavior. In addition, the Mg/Ca ratio in the M + 1 layer was lower at 21 psu than at 30 psu. Low pH resulted in a thicker M + 2 layer with higher Mg/Ca ratios despite shell dissolution, potentially helping to maintain shell integrity. While there was no change in heat shock protein (HSP70) expression, these shell modifications were accompanied by an upregulation of chitin synthase (CHS2) genes under low pH. These findings suggest that P. pygmaea is negatively influenced by low pH and low salinity, but also demonstrate compensatory mechanisms that enhance resistance to low pH.
Marine fish survival is threatened by ocean acidification, but the hormonal mechanisms for pH compensation are not well understood, limiting mechanistic understanding of stress responses in marine fish. We examined isotocin signaling in marine medaka (Oryzias melastigma) exposed to year-2100 ocean acidification conditions (Pco2 ∼0.14 kPa, pH 7.6). Our analysis demonstrated that isotocin receptor b (ITRb) was selectively upregulated at 6 h postexposure in adult gills, though it showed only a nonsignificant trend at 5 days postfertilization (dpf) embryos, whereas adenylyl cyclase 5 (ADCY5) showed hypercapnia responsiveness primarily at hatching. Using immunofluorescence and confocal microscopy, we found that both ITRb and ADCY5 proteins localize to the basolateral membrane of Na+-K+-ATPase-positive ionocytes, partially separated from apical H+-secretion machinery. Knockdown experiments showed that ITRb-ADCY5 coupling is crucial for pH compensation, with individual knockdown moderately reducing H+ secretion and combined knockdown causing severe impairment (>70% reduction) and decreasing transcription of acid-secretion genes (nhe3, ca2, and rhcgb) by 44%–60%. Paradoxically, double knockdown triggered a twofold cAMP increase that failed to restore function, whereas wild-type embryos maintained stable cAMP levels across pH conditions, consistent with the hypothesis that ITRb-ADCY5 coupling may organize cAMP production within specific basolateral microdomains, though direct subcellular imaging would be required to validate this compartmentalization model. The developmental asynchrony between ITRb (5 dpf) and ADCY5 (hatching) responses indicates life-stage-specific vulnerabilities. Our findings reveal that basolateral ITRb-ADCY5 coupling represents a critical control point for pH compensation capacity.
NEW & NOTEWORTHY Isotocin signaling, through ITRb-ADCY5 coupling, enables marine fish to compensate for ocean acidification via compartmentalized rather than global cAMP production. Paradoxically, disrupting this pathway doubles cAMP levels yet abolishes H+ secretion, revealing that signal localization determines function. The developmental asynchrony between ITRb and ADCY5 responses suggests that pH-stress vulnerabilities are specific to each life stage and require further investigation.
The present study experimentally investigates how temperature and acidification affect the polar lipid fatty acid (FA) composition of European sea bass (Dicentrarchus labrax) juveniles. Fish were reared for 92 days and fed ad libitum under four conditions: a control (CT; with natural fluctuating temperature and pH); a warming (WT, +4°C above control, and control pH), an acidification (AT, control temperature and −0.4 pH units below the control), and a combined warming and acidification treatment (WAT, +4°C and −0.4 pH units relative to the control). Results showed that warm treatments (WT and WAT) increased final weight and length similarly, whether or not combined with acidification, and that acidification alone (AT) had no effect on these variables. Polar lipid FA profiles of muscle and brain were only mildly impacted by temperature and not by acidification. Overall, the results suggest that European sea bass, under unrestricted quantity of food is able to maintain a very stable polar lipid FA composition. However, complementary studies with individuals under different feeding regimes or from wild populations subject to natural dietary variability are needed.
Marine diatoms are key components of the planetary ocean, playing crucial roles in trophic networks: fixing carbon, producing fatty acids (FA) that cannot be synthesized de novo by higher trophic organisms. To better comprehend how the combined action of climatic change influences the dietary value and the impacts on the upper trophic levels, we used a 2 × 2 factorial experiment to investigate how the FA profiles of Skeletonema marinoi changed in response to the individual and combined effects of warming (7, 19°C) and acidification (400, 1 000 ppm CO2). Three S. marinoi strains were exposed for ~ 40 weeks to ambient conditions, warming, acidification and their combination and analyzed for FA profiles, focusing on polyunsaturated (PUFA), omega-3 and omega-6 FA. We found that omega-3 FA increased under warming exposure, while acidification alone led to lower omega-3/omega-6 ratios. In contrast, the ratios increased under warming, alone or coupled with acidification, in all three strains, indicating better food quality for higher trophic levels. Our results suggest the long-term exposure to both drivers will help the marine diatoms to acclimatize to their combined effects, allowing them to buffer the changes brought by warming and ocean acidification.
We innovatively define two biophysical parameters (pH change rate and acceleration) of Zostera japonica photosynthesis.
pH change acceleration is sensitive to the regulation of carbon utilization pathways during Z. japonica photosynthesis.
A progressive pH–salinity-driven regulation mode of carbon utilization pathways in Z. japonica photosynthesis is summarized.
The impact of ocean acidification and alkalization on seagrass is characterized from a new biophysical standpoint.
Abstract
Seagrass carbon concentration mechanisms are modulated by seawater pH and salinity, yet their progressive regulation in photosynthetic inorganic carbon utilization pathways remain poorly characterized. This study novelly mathematically characterized first-order (pH change rate) and second-order (pH change acceleration) derivatives from pH-drift experiments in the intertidal seagrass Zostera japonica along China’s coastline. The pH change-based method effectively highlights the dissolved inorganic carbon (DIC) utilization and biomass accumulation of seagrass, while pH change acceleration is sensitive to the progressive switch of DIC utilization pathways during Z. japonica photosynthesis. As pH increases, six significant regulations emerge that have ecophysiological significance: a. primary regulation via reaching the CO2 compensation point; b. extracellular carbonic anhydrase (exCA) activity dropping to negligible levels; c. diminished efficiency of proton pump-mediated extracellular acidification in supplying CO2; d. regulation via reaching the bicarbonate saturation point; e. regulation via reaching the bicarbonate compensation point; and f. regulation via enhanced respiratory CO2 into seawater temporarily stimulating photosynthesis as a feedback. We summarize a progressive and universal pH–salinity-driven regulation mode reflecting different combinations of DIC utilization pathways and their respective intensities. Specifically, seawater pH modulates the mode’s fluctuating thresholds, while salinity governs the amplitude. Unexpectedly, elevated salinity serves as an effective stimulant for Z. japonica to maintain strong DIC utilization intensity in high pH (> 9.1) environments. Ocean acidification could increase pH change acceleration by 166%, enhancing carbon fixation, whereas artificial ocean alkalinity enhancement (Ca(OH)2 supplementation and olivine-seawater weathering) could reduce it by 165% and 105%, respectively, risking mortality of Z. japonica. Additionally, the salinity-dependent braking point (where acceleration drops to zero) serves as a critical threshold for seagrass photosynthesis and a new factor for transplantation-based restoration.
Mixotrophic protists capable of both photosynthesis and phagotrophy are key members of marine plankton communities. Yet, little is known about their responses to the combination of ocean acidification and warming.
A marine mixotrophic chrysophyte, Ochromonas CCMP2951, was subjected to two levels of pCO2 (300 and 800 ppm, resulting in pH of 8.2 and 7.8) and temperature (21°C and 26°C) in a factorial design.
Enhanced growth rates were observed in both the high CO2 and high temperature treatments, while cell size significantly decreased with temperature. Strongly decreased cellular phosphorus content led to increased N : P and C : P ratios of Ochromonas with temperature. Furthermore, warming increased grazing rates, while elevated CO2 reduced the Chl content but increased photosynthetic carbon acquisition, albeit only at low temperature. The combination of warming and elevated CO2 had antagonistic effects on the balance between autotrophic and heterotrophic carbon acquisition, keeping the net role of this mixotroph in the marine carbon cycle stable.
Altogether, both the direct stimulation of growth and the indirect effects of altered stoichiometry may favor mixotrophs under future ocean conditions. However, their contribution to future carbon cycling in complex natural communities will need further study.
Graphical Abstract
Grazing rates of Ochromonas CCMP2951 expressed as carbon obtained per mean cell volume (a) and Caut/het-photosynthetically fixed carbon divided by carbon obtained through grazing (b) in the four experimental treatments.
Low pH conditions at CO₂ vents increase THg bioavailability and fish exposure.
Functional traits explain THg patterns better than species identity.
Under low pH, less-mobile benthic fish accumulate more THg than highly-mobile fish.
Trophic position drives stronger THg biomagnification at low pH sites.
Abstract
Ocean acidification is not an isolated climate-related threat to marine organisms, as it may act in combination with other stressors such as trace element contamination. Here, we took advantage of naturally acidified conditions and mercury spill-out from the shallow CO2 vent of Vulcano Island (Italy) to test whether fish functional traits explain contaminant dynamics better than species identity under ocean acidification scenarios. Specifically, we investigated if trophic group and mobility influence total mercury (THg) bioaccumulation and trophic transfer in fish inhabiting Cymodocea nodosa seagrass meadows under low and ambient pH conditions. Low-mobility benthic fish, mainly represented by invertebrate feeders and small piscivores, exhibited higher THg concentration at the vent site than at reference sites, while highly mobile zooplanktivorous fish showed no significant differences, highlighting the importance of fish mobility and local trophic benthic pathways in shaping contaminant exposure and bioaccumulation. Moreover, the relationship between trophic position and THg concentration in fish was stronger in the low pH site, indicating enhanced trophic transfer and biomagnification when Hg availability is higher. In contrast, weak or absent biomagnification in the reference sites suggests that Hg transfer depends more on contaminant availability and food web structure than trophic position alone. Overall, these findings indicate that functional traits better predict Hg bioaccumulation patterns than species identity, supporting trait-based approach to assess contaminant dynamics in future ocean acidification scenarios.
Anthropogenic CO2 emissions are altering marine ecosystems through two interconnected processes: climate change and ocean acidification, both of which are particularly affecting the Southern Ocean. However, the molecular responses of cold-water corals to these stressors remain poorly understood. In this study, we investigated the transcriptomic response of the Antarctic coral Malacobelemnon daytoni exposed to ocean acidification (pH ~7.7; LpH) and elevated temperatures (+2°C and +4°C above ambient; T1 and T2, respectively) under controlled laboratory conditions. Using high-throughput RNA sequencing (RNA-Seq), we compared gene expression profiles across six treatments, including a control (pH ~8.1, CpH; 0–1°C; CT), acidification, warming, and their combinations. Principal component analysis revealed treatment-specific clustering and greater transcriptomic dispersion under LpH conditions. Differential expression analysis identified between 475 and 767 differentially expressed genes (DEGs), with the strongest transcriptional responses observed under combined stress conditions (LpH + T1 and LpH + T2). Combined exposure to ocean acidification and warming elicited the largest gene expression response, suggesting enhanced, potentially non-additive effects of multiple stressors. Functional enrichment analyses revealed differential regulation of genes associated with protein folding, signal transduction, energy metabolism, oxidative stress, and regulation of cell death. Our findings demonstrate that Malacobelemnon daytoni exhibits complex, treatment-dependent gene expression responses to environmental stressors. These results highlight the capacity of this Antarctic octocoral to mount molecular responses to projected future ocean conditions while also emphasizing its potential vulnerability to the interacting effects of ocean warming and acidification.
Climate change models predict future ocean-wide decreases in pH and increases in temperature, posing a serious threat to calcifying marine invertebrates. In Hawaiʻi, local seawater temperature and pH are predicted to change even more rapidly. The Hawaiian oyster Dendostrea sandvichensis, an endemic species, remains understudied despite its ecological roles in reef- building, biofiltration, and as a food source. While previous studies have revealed the alarming impacts of ocean acidification and warming on bivalves, little is known about how the planktonic larvae of D. sandvichensis will respond to projected climate conditions.
To investigate these effects, the larvae of D. Sandvichensis were reared under present-day conditions in Pearl Harbor, HI (pH 8.1, 26.5 °C) and projected future conditions (pH 7.7, 30.0°C) for one week. Shell growth, density, and degradation were then measured using micro-CT, confocal, and scanning electron microscopy. Additionally, two larval husbandry methods were evaluated, with a static-flow system paired with UV water treatment yielding approximately 11- fold higher survival than an open-flow system. Elevated temperature reduced larval survival (~67%) compared to ambient conditions. Reduced pH significantly decreased shell length (~11.8μm), while both stressors reduced shell density in Experiment 3. Shell density decreased by ~12% under elevated temperature, ~18% under reduced pH, and ~43% under the combined stressors, and was accompanied by increased shell degradation and abnormalities.
These findings indicate that elevated temperature and reduced pH impact larval oysters through multiple pathways, including reduced survival and compromised shell integrity. This study provides new insights into the vulnerability of an endemic Hawaiian species and essential data for predicting its resilience under future climate change scenarios.
Ocean acidification (OA) refers to the increase in the partial pressure of CO2 and decrease in the pH of seawater resulting from the absorption of atmospheric CO2 by the ocean. This study investigated OA impacts across multiple generations (F0-F3) of the copepod Acartia tonsa which were exposed to four pH conditions (8.1, 7.8, 7.6 and 7.1). Key reproductive and developmental traits were evaluated, including egg production, hatchability, development time, fecal pellet production, and sex ratio. Results showed that pH 7.1 significantly reduced egg production, hatchability and fecal pellet production, while prolonging the N–C time; N-A time remained unaffected by pH across all groups. For sex ratio, a downward trend was observed with increasing generations and decreasing pH: in F3, female proportion in pH 7.8, 7.6 and 7.1 groups was significantly lower than in the control group, and pH 7.1 and 7.6 groups had lower F3 female proportion than in the F1 generation. A. tonsa showed some tolerance under short-term acidification conditions at pH 7.1, but its physiological functions were further reduced after long-term exposure, suggesting a limited adaptive capacity. Ecologically, A. tonsa is a dominant coastal zooplankton species that links phytoplankton to higher trophic levels (e.g., fish larvae) and mediates energy flow and biogeochemical cycling. These results highlight the risks to key zooplankton populations and associated ecosystem functioning. Future studies should focus on the long-term effects of acidification on A. tonsa and combine multi-species experiments with field surveys to assess the ecological risks of OA.
Climate change explains ∼40% of biological stress variance in Mediterranean mussels.
Ocean acidification drives 67% of metal bioavailability changes via pH-metal speciation.
64% of climate effects on biomarkers operate indirectly through metal bioaccumulation.
Multi-method convergence (LMG, GAM, SEM, Bayesian Networks) confirms stressor ranking.
Only SSP1-2.6 keeps biological stress below the chronic-stress threshold through 2050.
Abstract
Marine coastal ecosystems face concurrent pressure from climate change and anthropogenic contamination, yet their relative contributions to biological stress remain poorly quantified. Here we present a decade-long (2014–2023) biomonitoring study on Mytilus galloprovincialis in the Ligurian Sea (NW Mediterranean), integrating quarterly biomarker measurements, heavy metal bioaccumulation data (12 metals), and high-resolution oceanographic records at a control site (Gorgona Island Marine Protected Area) and an offshore regasification terminal.
Biological stress variance was partitioned using five complementary analytical frameworks — Lindeman–Merenda–Gold (LMG) variance decomposition, Hierarchical and Generalised Additive Models (HGAM/GAM), Structural Equation Modelling (SEM), and Bayesian Networks — applied to four biomarkers: DNA damage, lysosomal membrane stability, gill tissue integrity, and immune response. A campaign-specific T0 baseline normalisation isolated environmental signals from initial population variability.
Climate change emerged as the dominant driver, consistently explaining ∼40% of variance across all methods, significantly exceeding metal bioaccumulation (∼30%), terminal influence (∼13%), and seasonal effects (∼2%). Ocean acidification was the primary climate mechanism, influencing 67% of analysed metals. Causal mediation analysis revealed that 64% of the climate effect operates indirectly through enhanced metal bioaccumulation (Climate→Metals→Biomarkers), while 36% acts directly. Climate and biological stress indices co-varied strongly (ρ = 0.78, p < 0.001), with marine heatwaves coinciding with peak biomarker responses.
Under IPCC Shared Socioeconomic Pathway (SSP) scenarios, the Biological Stress Index is projected to cross chronic-stress thresholds by 2035–2040 under the high-emission scenario (SSP5-8.5) and the intermediate-emission scenario (SSP2-4.5), with only the low-emission scenario (SSP1-2.6) maintaining stress below critical levels through 2050.
These findings challenge pollution-centric monitoring paradigms and demonstrate that CO2 mitigation now constitutes the highest-leverage intervention for marine invertebrate health in the Mediterranean.
Graphical abstract
This graphical abstract illustrates the main findings of a decade-long field study (2014–2023) on the effects of climate change and metal contamination on Mytilus galloprovincialis in the Ligurian Sea (NW Mediterranean). Three panels summarise the causal chain from environmental drivers to biological outcomes. The first panel depicts the key oceanographic trends recorded at the study site: ocean warming (+0.41°C/decade), acidification (−0.020 pH units/decade), deoxygenation, and a doubling of marine heatwave frequency after 2018. The second panel shows how pH decline enhances the bioavailability of 67% of the metals analysed, driving a predominantly indirect pathway (64%) from climate stressors to biological stress, mediated by metal bioaccumulation, as revealed by structural equation modelling and Bayesian network analysis. The final panel presents the four biomarkers used to compute the Biological Stress Index (BSI), its strong temporal correlation with the Climate Change Index (ρ = 0.78), and BSI projections to 2050 under three IPCC emission scenarios, showing that only SSP1-2.6 keeps BSI below the chronic-stress threshold throughout the projection period.
Absorption of CO2 by global oceans is decreasing pH resulting in ocean acidification (OA). Impacts on shellfish have been documented in ecologically and commercially important species. We examined the influence of diet and OA between two populations of red abalone (Haliotis rufescens) a species of aquaculture importance and declining wild populations. Populations experience different exposure histories: strong upwelling (Van Damme, California [VD]) historically exposed to low-pH conditions and weak-intermittent upwelling (Santa Barbara, California [SB]). Abalone were cultured under control-pH or OA-conditions and fed crustose coralline algae (CCA) or diatoms used in aquaculture. We tested treatment effects of population, settlement diet, and OA-exposure on survival as influenced by larval-energy stores. Survival in both populations was enhanced by CCA when cultured under both treatment conditions; however, by later stages, this effect remained only for SB. SB had reduced post-settlement survival when cultured under OA-conditions, whereas post-settlement survival of VD was not. Diet affected the relationship between larval-energy and post-settlement survival; a positive relationship when fed diatoms and a negative relationship with CCA. The relationship between larval energy and post-settlement survival was stronger in VD. CCA enhanced juvenile growth in SB cultured abalone at both three-months and one-year post-settlement. Settlement diets can reduce the impacts of OA on early-life stages of abalone, but population differences driven by underlying energetics affect the consistency of this outcome. These findings illuminate the impacts from OA, suggesting populations may be at risk, and inform strategies for developing and sustaining shellfish aquaculture in the face of changing ocean conditions.
To investigate the effects of ocean acidification (OA) on macroalgae, we conducted in situ surveys along a natural CO2 vent gradient in Shikine Island, Japan, together with complementary laboratory culture experiments. The in-situ surveys revealed that near the CO2 vent (where pH dropped by 0.37), macroalgal diversity and species-richness were less than half those at the reference sites under ambient pH conditions. Nevertheless, the rates of CO2 assimilation of several common macroalgae increased from the reference site to the areas near the CO2 vent. This enhancement coincided with decreased photosynthetic CO2 affinity, reflecting that the acidified area down-regulated CO2-concentrating mechanisms in the algae. Measured photosystem II activity revealed that macroalgae at reference sites had lower electron transport rate and light utilization efficiency. The laboratory culture experiments, in which the dominant species (Gelidium elegans and Dictyopteris undulata), were cross-exposed to ambient and elevated CO2 conditions, further demonstrated that the stress near the CO2 vent significantly exacerbated photoinhibition under high light stress. Our results demonstrate that reduced pH and high sunlight act synergistically to impair macroalgal photosynthesis through exacerbated photoinhibition. This effect was more pronounced in red algae (e.g., G. elegans) than in brown algae (e.g., D. undulata). These different physiological responses provide a mechanistic explanation for an observed community shift, from red algal dominance in ambient pCO2 areas to brown algal dominance near the vent. Our findings imply that future OA, when combined with high-light stress, may selectively disadvantage high-light-sensitive species, thereby altering macroalgal community structure in coastal waters.
Natural CO2 vents reveal transcriptomic responses to chronic ocean acidification.
Acute low-pH exposure triggers rapid but limited plastic responses in Arbacia lixula.
Vent-origin Arbacia lixula exhibit extensive metabolic reprogramming and antioxidant activation.
Low pH supresses biomineralization genes and up-regulates collagen and extracellular matrix pathways.
Persistence under ocean acidification is associated with energetic trade-offs and skeletal homeostasis.
Abstract
Ocean acidification is reshaping coastal ecosystems as a consequence of anthropogenic CO2 emissions. Natural CO2 vent systems provide valuable analogues for investigating organismal responses to long-term acidified conditions under ecologically realistic scenarios. Here, we examined genome-wide transcriptomic responses of the sea urchin Arbacia lixula, an ecosystem-relevant grazer inhabiting a natural CO2 vent system in La Palma (Canary Islands, Spain). Using RNA sequencing of 24 adults (n = 8 per treatment), we compared: (i) acute experimental exposure of ambient-origin individuals to low pH, (ii) chronic exposure by comparing ambient and vent-origin populations in their native pH conditions, and (iii) a genotype-of-origin comparison under shared low pH. Acute exposure triggered a limited transcriptional response (116 differentially expressed genes, DEG), characterized by activation of ion transport, redox regulation, and NAD-associated metabolism. In contrast, chronically exposed vent-origin urchins showed a tenfold increase in transcriptional changes (1053 DEG), reflecting metabolic reprogramming involving lipid, carbohydrate and amino acid pathways, and strengthened antioxidant capacity. Chronic low-pH exposure was also associated with suppression of biomineralization and developmental genes, alongside strong upregulation of collagen and extracellular matrix–associated genes that may help maintain skeletal performance under reduced carbonate availability. Genotype-of-origin effects (131 DEGs) revealed constitutive differences in metabolic, redox, extracellular matrix, and biomineralization pathways in vent populations. Together, these findings indicate that persistence under natural acidification involves both rapid plastic responses and sustained physiological reorganization, providing mechanistic insight into how calcifying species maintain functional performance under ongoing ocean acidification.
Cold-water corals (CWCs) are key ecosystem engineers that create complex three-dimensional habitats much like tropical reefs, but in deep, cold seas. However, like other reef-building systems, they are increasingly threatened by climate change and ocean acidification. CWC communities in the Mediterranean Sea may be especially vulnerable because these waters absorb more atmospheric CO2 than the global ocean, making it a mesocosm that mirrors broader global trends affecting marine life. Since calcification is energetically costly and likely becomes even more demanding as pH and carbonate ion availability decline, understanding how the decrease in aragonite saturation state (Ωarag) affects biomineralization is essential for predicting the future of these corals.
Results
Here, we investigated skeletal structural and compositional changes of the scleractinian CWC Desmophyllum dianthus along an Ωarag gradient in the Mediterranean Sea using specimens collected between 400 and 1200 m depth. Our findings indicate that skeletal porosity increases at the macro-scale with decreasing Ωarag, while micro- and nano-scale structural and compositional features remained unaffected.
Conclusions
The persistence of micro- and nano-scale skeletal features across an 800 m depth gradient suggests that D. dianthus maintains tight biological control over mineralization at these scales, even as Ωarag declines. This control does not extend to the macro-scale, where increasing porosity alters the skeleton’s overall architecture under lower Ωarag. D. dianthus thus appears to preserve the fundamental “building blocks” of its skeleton while changing its larger-scale structure, a decoupling that may make macro-scale porosity an early marker of acidification stress in CWCs.
Acidification in coastal habitats is increasing in duration and amplitude under the continued influence of ocean acidification and contributing coastal processes. The impacts of low pH conditions on calcifying organisms, especially echinoderms, is well established, with the early developmental stages being especially vulnerable. This is the first study to assess the impact of locally relevant coastal acidification scenarios on the early development of the Cape urchin Parechinus angulosus. Our findings suggest that the early larval stages of this species are unlikely to survive when exposed to low pH conditions, specifically during the onset of skeletogenesis. In our laboratory experiments, larvae that were exposed to the low pH treatment (pH 7.32) showed significantly reduced growth (GLMM, Time × Treatment interaction: β = −0.361 ± 0.019, z = −19.06, p < 0.001) and developmental regression compared with those from the control treatment (pH 7.95). Substantially slower growth rates were observed in the low pH treatment (length = 72.3 hpf0.18) compared with in the control treatment (length = 24.24 hpf0.54). There was also evidence of abnormal and delayed development and potential dissolution of skeletal structures under the low pH condition. However, fertilisation success and larval survival did not differ significantly between the experimental treatments, suggesting that developmental impacts of low pH over short durations, even though substantial, may be sublethal. The developmental impacts are likely to impair the transition of larvae to the adult stages, which may ultimately affect populations of this ecologically important species under future coastal acidification scenarios.
Ocean acidification, driven by rising atmospheric CO₂, threatens the ability of corals to build their skeletons by reducing their capacity to maintain an elevated pH at the calcification site (pHcf), a process essential for calcium carbonate precipitation. Boron isotopes have commonly been used to show that the response of pHcf to ocean acidification is highly species-specific. However, the physiological mechanisms underlying this variability remain poorly understood. Recently, lithium (Li) isotopes have been used to trace the activity of ionic transport involved in cellular pH regulation and calcification (e.g. H+, Na+ and Ca2+), and may therefore help resolve these mechanisms. Here, we investigate multiple coral species from Tutum Bay (Papua New Guinea), a natural CO₂ seep system creating pH gradients (mean pHT = 7.66 at seeps vs. 8.01 at control sites) analogous to future ocean acidification scenarios. Our results show a relationship between seawater pH, calcifying fluid chemistry, and lithium isotopic composition. Corals exposed to low seawater pH exhibit significantly altered δ⁷Li values relative to colonies from the control site, with some species becoming enriched in ⁷Li (up to 2‰) as pHcf declines. This isotopic shift is consistent with reduced efficiency of Na⁺/H⁺ exchangers (NHEs), active transporters that preferentially incorporate the lighter ⁶Li isotope under optimal conditions but may become less effective under elevated proton concentrations. By linking Li isotopes to calcifying-fluid chemistry, these results provide geochemical evidence that ocean acidification may disrupt ionic regulation in corals and that Li isotopes can help to resolve biogeochemical controls of carbonate-systems.
Marine ecosystems are undergoing rapid transformation under climate change, yet the responses of many marine invertebrates remain vastly understudied. In particular, for many benthic gastropods there is a striking imbalance between their traditional appreciation by shell collectors—and, consequently, their consistent representation in Natural History Collections—and the limited attention they receive in ecological and conservation studies. Focusing on the northeastern Atlantic and the Mediterranean, the cowries Luria lurida, Naria spurca, Zonaria pyrum and the frog-shell Talisman scrobilator are emblematic examples of this knowledge gap, despite being frequently mentioned as species of conservation concern. Using long-term occurrence records spanning more than a century, we modelled past and present distributions of these species and explored their potential responses to future climate scenarios through a multi-temporal Species Distribution Modelling framework. Our results show that intermediate climatic conditions—both in time (2050–2060 vs. 2090–2100) and scenario intensity (moderate SSP2-4.5 versus high-emission SSP5-8.5)—may represent a critical transition phase, leading to habitat contractions without compensatory gains in newly emerging suitable areas. The Mediterranean Sea is expected to increasingly function as a cul-de-sac, with the dominant circulation patterns strongly limiting outward movements towards cooler regions for species relying on planktic larvae for dispersal. Furthermore, incorporating larval sensitivity to reduced pH suggests that large areas of the Atlantic Ocean may actually result unsuitable for larval persistence, substantially reducing the habitat effectively available for completion of the full life cycle; this highlights the need to account for connectivity, life-history constraints and juvenile-stage sensitivity when assessing climate-driven range shifts in shelled organisms with planktic larvae.
