Decreasing Seawater pH Research Articles

A decrease in pH, increase in temperature, and pollution exposure elicit distinct stress responses in a scleractinian coral (Desmophyllum pertusum)

Scleractinian corals create habitat that persists on geological timescales and supports some of the most diverse ecosystems on Earth. Throughout their depth range, these ecosystems are threatened by anthropogenic disturbances, including global ocean change and hydrocarbon extraction. While numerous studies have focused on how shallow-water corals respond to stressors, there is a paucity of research on deep-sea corals. Here, we analyze the gene expression patterns from a reef-forming deep-sea coral, Desmophyllum pertusum (previously Lophelia pertusa), exposed to oil and dispersant mixtures under current and projected future temperature and pH conditions. The overall gene expression patterns varied among coral colonies (genets), but dispersant exposure induced the strongest response. A Weighted-Gene Correlation Network Analysis (WGCNA) identified networks of co-expressed genes in response to different stressors. Gene ontology (GO) enrichment analysis revealed that D. pertusum exhibited the coral cellular stress response (CSR) during exposure to oil, dispersant, and a decrease in pH. Dispersant exposure elicited a response that included up-regulation of apoptosis, immune system, wound healing, and stress-related pathways. Coral nubbins exposed to oil exhibited signs of resource reallocation and a reduction in growth to maintain cellular homeostasis. The decrease in seawater pH resulted in a less severe stress response than dispersant exposure. These results support the idea that there is an underlying environmental stress response (ESR) shared by scleractinians, but that this response varies depending on the type and intensity of the stress with which they are challenged.

CO2‐induced seawater acidification impairs the stinging cells of a jellyfish

AbstractCO2‐induced seawater acidification has been shown to modify predator–prey interactions in many marine taxa. Scyphozoans play an important role in the trophic dynamics of marine ecosystems during their blooms in coastal waters; however, the impacts of seawater acidification on the predation behavior of these animals are poorly understood. Here, we aimed to examine the impact of a decrease in seawater pH on the feeding behavior and growth of ephyrae (juvenile medusae) of the scyphozoan Aurelia coerulea. Combining bulk and single‐cell RNA sequencing approaches, we assessed transcriptomic changes of ephyrae under a laboratory‐based pH 7.6 condition. We found that the feeding rates and growth of ephyrae were significantly inhibited by a decrease in seawater pH. Furthermore, transcriptome analysis showed that a decline in pH significantly reduced the expression of genes related to toxins and nematocyst structure in ephyrae. These findings were further confirmed by single‐cell transcriptomic analyses and revealed that low pH impaired the toxin activity and energy metabolism of stinging cells. The pH recovery experiment indicated that moving ephyrae from seawater with pH 7.6 into seawater with pH 8.1 greatly restored their feeding, growth, and toxin‐related and nematocyst structure–related gene expression. However, exposure to pH 7.6 for 23 d could not recover the decrease in the feeding and growth of ephyrae. Together, these findings indicate that CO2‐induced acidification compromised the stinging cells of A. coerulea ephyrae, with concomitant negative consequences on predation and growth that are likely to alter predator–prey interactions, with consequent effects on community structure and ecosystem.

High nutrient availability modulates photosynthetic performance and biochemical components of the economically important marine macroalga Kappaphycus alvarezii (Rhodophyta) in response to ocean acidification

Increased atmospheric CO2 concentrations not only change the components of inorganic carbon system in seawater, resulting in ocean acidification, but also lead to decreased seawater pH, resulting in ocean acidification. Consequently, increased inorganic carbon concentrations in seawater provide a sufficient carbon source for macroalgal photosynthesis and growth. Increased domestic sewage and industrial wastewater discharge into coastal areas has led to nutrient accumulation in coastal seawaters. Combined with elevated pCO2 (1200 ppmv), increased nutrient availability always stimulates the growth of non-calcifying macroalgae, such as red economical macroalga Gracilariopsis lemaneiformis. Here, we evaluated the interactive effects of nutrients with elevated pCO2 on the economically important marine macroalga Kappaphycus alvarezii (Rhodophyta) in a factorial 21-day coupling experiment. The effects of increased nutrient availability on photosynthesis and photosynthetic pigments of K. alvarezii were greater than those of pCO2 concentration. The highest Fv/Fm values (0.660 ± 0.019 and 0.666 ± 0.030, respectively) were obtained at 2 μmol L−1 of NO3–N at two pCO2 levels. Under the elevated pCO2 condition, the Chl-a content was lowest (0.007 ± 0.004 mg g−1) at 2 μmol L−1 of NO3–N and highest (0.024 ± 0.002 mg g−1) at 50 μmol L−1 of NO3–N. The phycocyanin content was highest (0.052 ± 0.012 mg g−1) at 150 μmol L−1 of NO3–N under elevated pCO2 condition. The malondialdehyde content declined from 32.025 ± 4.558 nmol g−1 to 26.660 ± 3.124 nmol g−1 with the increased nutrients at under low pCO2. To modulate suitable adjustments, soluble biochemical components such as soluble carbohydrate, soluble protein, free amino acids, and proline were abundantly secreted and were likely to protect the integrity of cellular structures under elevated nutrient availability. Our findings can serve as a reference for cultivation and bioremediation methods under future environmental conditions.

Spatial symmetry and contrasting controls of surface pH and aragonite saturation state in the western North Pacific

Oceanic uptake of anthropogenic CO2 causes a decrease in seawater pH and aragonite saturation state (Ωarag), a process known as ocean acidification (OA). The western North Pacific is a hotspot for anthropogenic CO2 sinks; however, the spatiotemporal variability of pH and Ωarag and their controlling mechanisms remain unexplored. In this study, we provide high-frequency and high-precision underway measurements of sea surface pCO2 and pH to investigate the distribution and drivers of OA metrics across different hydrochemical gradients in the western North Pacific in late spring 2018, a season with the highest primary production in the year. Our results show that the surface pH reached near air-sea equilibrium in the subtropical zone but gradually increased northward across the Kuroshio Recirculation (KR) zone and peaked in the Kuroshio Extension (KE) zone. We found that sea surface temperature played the most prominent role in regulating pH, which was also counteracted by the effects of air–sea gas exchange and vertical mixing. In contrast, the distribution of Ωarag largely mirrored the pH and was governed by air–sea gas exchange and vertical mixing, the effects of which on Ωarag were enhanced by temperature. Biological activity thrived in the KE zone to increase both pH and Ωarag, which further reinforced the latitudinal pattern of pH, but weakened that of Ωarag. These findings are based on direct in situ measurements of pH and improve our understanding of the spatiotemporal variability of OA metrics in the western North Pacific region.

The Effects of Short-Term Exposure to pH Reduction on the Behavioral and Physiological Parameters of Juvenile Black Rockfish (Sebastes schlegelii).

Coastal areas are subject to greater pH fluctuation and more rapid pH decline as a result of both natural and anthropogenic influences in contrast to open ocean environments. Such variations in pH have the potential to pose a threat to the survival and physiological function of offshore fishes. With the aim of evaluating the impact of short-term pH reduction on the behavioral performance and physiological response of costal fish, the black rockfish (Sebastes schlegelii), one of the principal stock-enhanced species, was examined. In the present study, juveniles of the black rockfish with a mean body length of 6.9 ± 0.3 cm and weight of 8.5 ± 0.5 g were exposed to a series of pHs, 7.0, 7.2, 7.4, 7.6, 7.8, and normal seawater (pH 8.0) for 96 h. At the predetermined time points post-exposure (i.e., 0, 12, 24, 48, and 96 h), fish movement behavior was recorded and the specimens were sampled to assess their physiological responses. The results indicate that the lowered pH environment (pH 7.0-7.8) elicited a significant increase in highly mobile behavior, a decrease in immobile behavior, and a significant rise in the metabolic levels of the black rockfish juveniles. Specifically, carbohydrate metabolism was significantly elevated in the pH 7.2 and 7.4 treatments, while lipid metabolism was significantly increased in the pH 7.0, 7.4, and 7.8 treatments. The results of the present study indicate that short-term reductions in pH could ramp up boldness and boost energy expenditure in the black rockfish juveniles, leading to an increased metabolic cost. Additionally, the present investigation revealed that the black rockfish juveniles were capable of adapting to a short-term pH reduction. The findings may provide insight into the underlying physiological mechanisms that govern fish responses to potential decreases in seawater pH in the future.

Corals adapted to extreme and fluctuating seawater pH increase calcification rates and have unique symbiont communities.

Ocean acidification (OA) is a severe threat to coral reefs mainly by reducing their calcification rate. Identifying the resilience factors of corals to decreasing seawater pH is of paramount importance to predict the survivability of coral reefs in the future. This study compared corals adapted to variable pHT (i.e., 7.23-8.06) from the semi-enclosed lagoon of Bouraké, New Caledonia, to corals adapted to more stable seawater pHT (i.e., 7.90-8.18). In a 100-day aquarium experiment, we examined the physiological response and genetic diversity of Symbiodiniaceae from three coral species (Acropora tenuis, Montipora digitata, and Porites sp.) from both sites under three stable pHNBS conditions (8.11, 7.76, 7.54) and one fluctuating pHNBS regime (between 7.56 and 8.07). Bouraké corals consistently exhibited higher growth rates than corals from the stable pH environment. Interestingly, A. tenuis from Bouraké showed the highest growth rate under the 7.76 pHNBS condition, whereas for M. digitata, and Porites sp. from Bouraké, growth was highest under the fluctuating regime and the 8.11 pHNBS conditions, respectively. While OA generally decreased coral calcification by ca. 16%, Bouraké corals showed higher growth rates than corals from the stable pH environment (21% increase for A. tenuis to 93% for M. digitata, with all pH conditions pooled). This superior performance coincided with divergent symbiont communities that were more homogenous for Bouraké corals. Corals adapted to variable pH conditions appear to have a better capacity to calcify under reduced pH compared to corals native to more stable pH condition. This response was not gained by corals from the more stable environment exposed to variable pH during the 100-day experiment, suggesting that long-term exposure to pH fluctuations and/or differences in symbiont communities benefit calcification under OA.

Ocean acidification causes fundamental changes in the cellular metabolism of the Arctic copepod Calanus glacialis as detected by metabolomic analysis

Using a targeted metabolomic approach we investigated the effects of low seawater pH on energy metabolism in two late copepodite stages (CIV and CV) of the keystone Arctic copepod species Calanus glacialis. Exposure to decreasing seawater pH (from 8.0 to 7.0) caused increased ATP, ADP and NAD+ and decreased AMP concentrations in stage CIV, and increased ATP and phospho-L-arginine and decreased AMP concentrations in stage CV. Metabolic pathway enrichment analysis showed enrichment of the TCA cycle and a range of amino acid metabolic pathways in both stages. Concentrations of lactate, malate, fumarate and alpha-ketoglutarate (all involved in the TCA cycle) increased in stage CIV, whereas only alpha-ketoglutarate increased in stage CV. Based on the pattern of concentration changes in glucose, pyruvate, TCA cycle metabolites, and free amino acids, we hypothesise that ocean acidification will lead to a shift in energy production from carbohydrate metabolism in the glycolysis toward amino acid metabolism in the TCA cycle and oxidative phosphorylation in stage CIV. In stage CV, concentrations of most of the analysed free fatty acids increased, suggesting in particular that ocean acidification increases the metabolism of stored wax esters in this stage. Moreover, aminoacyl-tRNA biosynthesis was enriched in both stages indicating increased enzyme production to handle low pH stress.

Transcriptome analysis of hepatopancreas in penaeus monodon under acute low pH stress

The decrease of seawater pH can affect the metabolism, acid-base balance, immune response and immunoprotease activity of aquatic animals, leading to aquatic animal stress, impairing the immune system of aquatic animals and weakening disease resistance, etc. In this study, we performed high-throughput sequencing analysis of the hepatopancreas transcriptome library of low pH stress penaeus monodon, and after sequencing quality control, a total of 43488612–56271828 Clean Reads were obtained, and GO annotation and KEGG pathway enrichment analysis were performed on the obtained Clean Reads, and a total of 395 DEGs were identified. we mined 10 differentially expressed and found that they were significantly enriched in the Metabolic pathways (ko01100), Biosynthesis of secondary metabolites (ko01110), Nitrogen metabolism (ko00910) pathways, such as PIGA, DGAT1, DGAT2, UBE2E on Metabolic pathways; UGT, GLT1, TIM genes on Biosynthesis of secondary metabolites; CA, CA2, CA4 genes on Nitrogen metabolism, are involved in lipid metabolism, induction of oxidative stress and inflammation in the muscular body of spot prawns. These genes play an important role in lipid metabolism, induction of oxidative stress and inflammatory response in the muscle of the shrimp. In summary, these genes provide valuable reference information for future breeding of low pH-tolerant shrimp.

Abiotic plastic leaching contributes to ocean acidification

Ocean acidification and plastic pollution are considered as potential planetary boundary threats for which crossing certain thresholds could be very harmful for the world's societies and ecosystems well-being. Surface oceans have acidified around 0.1 units since the Industrial Revolution, and the amount of plastic reaching the ocean in 2018 was quantified to 13 million metric tonnes. Currently, both ocean threats are worsening with time. Plastic leaching is known to alter the biogeochemistry of the ocean through the release of dissolved organic matter. However, its impact in the inorganic chemistry of the seawater is less studied. Here we show, from laboratory experiments, that abiotic plastic degradation induces a decrease in seawater pH, particularly if the plastic is already aged, as that found in the ocean. The pH decrease is enhanced by solar radiation, and it is probably induced from a combination of the release of organic acids and the production of CO2. It is also related to the amount of leached dissolved organic carbon, with higher acidification as leaching increases. In coastal areas, where plastic debris accumulates in large quantities, plastic leaching could lead to a seawater pH decrease up to 0.5 units. This is comparable to the projected decrease induced in surface oceans by the end of the twenty-first century for the most pessimistic anthropogenic emissions scenarios.

Ocean acidification impacts sperm swimming performance and pHi in the New Zealand sea urchin Evechinus chloroticus.

In sea urchins, spermatozoa are stored in the gonads in hypercapnic conditions (pH<7.0). During spawning, sperm are diluted in seawater of pH>8.0, and there is an alkalinization of the sperm's internal pH (pHi) through the release of CO2 and H+. Previous research has shown that when pHi is above 7.2-7.3, the dynein ATPase flagellar motors are activated, and the sperm become motile. It has been hypothesized that ocean acidification (OA), which decreases the pH of seawater, may have a narcotic effect on sea urchin sperm by impairing the ability to regulate pHi, resulting in decreased motility and swimming speed. Here, we used data collected from the same individuals to test the relationship between pHi and sperm motility/performance in the New Zealand sea urchin Evechinus chloroticus under near-future (2100) and far-future (2150) atmospheric PCO2conditions (RCP 8.5: pH 7.77, 7.51). Decreasing seawater pH significantly negatively impacted the proportion of motile sperm, and four of the six computer-assisted sperm analysis (CASA) sperm performance measures. In control conditions, sperm had an activated pHi of 7.52. Evechinus chloroticus sperm could not defend pHi in future OA conditions; there was a stepped decrease in the pHi at pH 7.77, with no significant difference in mean pHi between pH 7.77 and 7.51. Paired measurements in the same males showed a positive relationship between pHi and sperm motility, but with a significant difference in the response between males. Differences in motility and sperm performance in OA conditions may impact fertilization success in a future ocean.

PH trends and seasonal cycle in the coastal Balearic Sea reconstructed through machine learning

The decreasing seawater pH trend associated with increasing atmospheric carbon dioxide levels is an issue of concern due to possible negative consequences for marine organisms, especially calcifiers. Globally, coastal areas represent important transitional land-ocean zones with complex interactions between biological, physical and chemical processes. Here, we evaluated the pH variability at two sites in the coastal area of the Balearic Sea (Western Mediterranean). High resolution pH data along with temperature, salinity, and also dissolved oxygen were obtained with autonomous sensors from 2018 to 2021 in order to determine the temporal pH variability and the principal drivers involved. By using environmental datasets of temperature, salinity and dissolved oxygen, Recurrent Neural Networks were trained to predict pH and fill data gaps. Longer environmental time series (2012–2021) were used to obtain the pH trend using reconstructed data. The best predictions show a rate of -,0.0020pm 0.00054 pH units year^{-1}, which is in good agreement with other observations of pH rates in coastal areas. The methodology presented here opens the possibility to obtain pH trends when only limited pH observations are available, if other variables are accessible. Potentially, this could be a way to reliably fill the unavoidable gaps present in time series data provided by sensors.

Caribbean king crab larvae and juveniles show tolerance to ocean acidification and ocean warming

Coastal habitats are experiencing decreases in seawater pH and increases in temperature due to anthropogenic climate change. The Caribbean king crab, Maguimithrax spinosissimus, plays a vital role on Western Atlantic reefs by grazing macroalgae that competes for space with coral recruits. Therefore, identifying its tolerance to anthropogenic stressors is critically needed if this species is to be considered as a potential restoration management strategy in coral reef environments. We examined the effects of temperature (control: 28 °C and elevated: 31 °C) and pH (control: 8.0 and reduced pH: 7.7) on the king crab’s larval and early juvenile survival, molt-stage duration, and morphology in a fully crossed laboratory experiment. Survival to the megalopal stage was reduced (13.5% lower) in the combined reduced pH and elevated temperature treatment relative to the control. First-stage (J1) juveniles delayed molting by 1.5 days in the reduced pH treatment, while second-stage (J2) crabs molted 3 days earlier when exposed to elevated temperature. Juvenile morphology did not differ among treatments. These results suggests that juvenile king crabs are tolerant to changes associated with climate change. Given the important role of the king crab as a grazer of macroalgae, its tolerance to climate stressors suggests that it could benefit restoration efforts aimed at making coral reefs more resilient to increasingly warm and acidic oceans into the future.

Genome-wide association study reveals genetic variations associated with ocean acidification resilience in Yesso scallop Patinopecten yessoensis.

Ocean acidification (OA), which refers to a gradual decrease in seawater pH due to the absorption of atmospheric carbon dioxide, profoundly affects the growth, development and survival of bivalves. Relatively limited studies have assessed the resilience of bivalve to OA. In the present study, Patinopecten yessoensis, an economically and ecologically significant species, were exposed to low pH (pH=7.5) for 4 weeks. Forty-seven scallops that died in the first week were considered pH-sensitive population, and 20 that were alive at the end of the experiment were considered pH-tolerant population. A genome-wide association study was conducted to identify the genomic loci associated the resilience of P. yessoensis to OA. Twenty-one single nucleotide polymorphisms were significantly associated with resilience, which were distributed in 11 linkage groups. Within the linkage disequilibrium block region (± 300kb) surrounding the 21 SNPs, 193 candidate genes were successfully identified. Particularly, five associated SNPs were directly located on five genes, including SP24, CFDH, 5HTR3, HSDL1 and ZFP346. The GO enrichment and KEGG pathway analyses showed that the molecular response of P. yessoensis to OA mainly involved neural signal transmission, energy metabolism and redox reaction. Candidate genes were expressed during larval development and in adult tissues. Furthermore, the expression of 30 candidate genes changed significantly under low pH stress in the mantle. Our results reveal certain SNPs and candidate genes that could elucidate the different responses of P. yessoensis to OA. The genetic variations indicated molecular resilience in P. yessoensis populations, which may enable adaptation to future acidification stress.

Ocean Acidification and Mollusc Settlement in Posidonia oceanica Meadows: Does the Seagrass Buffer Lower pH Effects at CO2 Vents?

Ocean acidification has been broadly recognised to have effects on the structure and functioning of marine benthic communities. The selection of tolerant or vulnerable species can also occur during settlement phases, especially for calcifying organisms which are more vulnerable to low pH–high pCO2 conditions. Here, we use three natural CO2 vents (Castello Aragonese north and south sides, and Vullatura, Ischia, Italy) to assess the effect of a decrease of seawater pH on the settlement of Mollusca in Posidonia oceanica meadows, and to test the possible buffering effect provided by the seagrass. Artificial collectors were installed and collected after 33 days, during April–May 2019, in three different microhabitats within the meadow (canopy, bottom/rhizome level, and dead matte without plant cover), following a pH decreasing gradient from an extremely low pH zone (pH &lt; 7.4), to ambient pH conditions (pH = 8.10). A total of 4659 specimens of Mollusca, belonging to 57 different taxa, were collected. The number of taxa was lower in low and extremely low pH conditions. Reduced mollusc assemblages were reported at the acidified stations, where few taxa accounted for a high number of individuals. Multivariate analyses revealed significant differences in mollusc assemblages among pH conditions, microhabitat, and the interaction of these two factors. Acanthocardia echinata, Alvania lineata, Alvania sp. juv, Eatonina fulgida, Hiatella arctica, Mytilys galloprovincialis, Musculus subpictus, Phorcus sp. juv, and Rissoa variabilis were the species mostly found in low and extremely low pH stations, and were all relatively robust to acidified conditions. Samples placed on the dead matte under acidified conditions at the Vullatura vent showed lower diversity and abundances if compared to canopy and bottom/rhizome samples, suggesting a possible buffering role of the Posidonia on mollusc settlement. Our study provides new evidence of shifts in marine benthic communities due to ocean acidification and evidence of how P. oceanica meadows could mitigate its effects on associated biota in light of future climate change.

Effects of ocean acidification on the growth, photosynthetic performance, and domoic acid production of the diatom Pseudo-nitzschia australis from the California Current System

Pseudo-nitzschia australis (Frenguelli), a toxigenic pennate diatom capable of producing the neurotoxin domoic acid (DA), was examined in unialgal laboratory cultures to quantify its physiological response to ocean acidification (OA) - the decline in pH resulting from increasing partial pressure of CO2 (pCO2) in the oceans. Toxic blooms of P. australis are common in the coastal waters of eastern boundary upwelling systems (EBUS), including those of the California Current System (CCS) off the west coast of the United States where increased pCO2 and decreased seawater pH are well-known. This study determined the production of dissolved (dDA) and particulate DA (pDA), the rates of growth and nutrient (nitrate, silicate and phosphate) utilization, cellular elemental ratios of carbon and nitrogen, and the photosynthetic response to declining pH during the exponential and stationary growth phases of a strain of P. australis isolated during a massive toxic bloom that persisted for months along much of the U.S. west coast during 2015. Our controlled lab studies showed that DA production significantly increased as pCO2 increased, and total DA (pDA + dDA) normalized to cell density was 2.7 fold greater at pH 7.8 compared to pH 8.1 (control) during nutrient-limited stationary growth. However, exponential growth rates did not increase with declining pH, but remained constant until pH of 7.8 was reached, and then specific growth rates declined by ca. 30%. The toxin results demonstrate that despite minimal effects of OA observed during the nutrient-replete exponential growth phase, the enhancement of DA production, notably the 3-fold increase in particulate DA per cell, with declining pH from 8.1 to 7.8 during the nutrient-depleted stationary phase, supports the hypothesis that increasing pCO2 will result in greater toxic risk to coastal ecosystems from elevated ambient concentrations of particulate DA. The ecological consequences of decreasing silicate uptake rates and increasing cellular carbon quotas with declining pH may potentially ameliorate some negative impacts of OA on Pseudo-nitzschia growth in natural systems.

Establishing an Ocean Acidification Monitoring System for the Tropical Waters of Indonesia Facing Regional Climate Variability

The emission of greenhouse gases, including high CO2 and other materials, initiates global warming and climate change. Atmospheric CO2 that affects the carbonate system of seawater causes ocean acidification (OA). OA affects marine organisms directly, as well as humans economically and ecologically. Considering the high impact of OA and following the United Nations' Sustainable Development Goals, systematic research and monitoring of OA is necessary in Indonesia, whose seas play an important role in this emerging phenomenon. This review discusses the urgency of OA monitoring systems and suggests carbonate system monitoring, as well as carbon biogeochemistry. OA significantly affects marine production and alters ecosystem services, and it is likely to have an impact on habitats shifting from calcified to non-calcified and reducing benthic complexity. Its effect on calcifying organisms can also be found, i.e., coral calcification and/or dissolution of CaCO3 of calcifying organisms. Acidity (pH), as well as the carbonate system variables of seawater, fluctuate, especially with variations in space and time. Coastal ecosystems that are directly affected by terrestrial input will have carbonate system variables that fluctuate more. The annual rate of decreasing seawater pH, especially over an open and large spatial scale, may indicate OA. Therefore, a monitoring system must be implemented to obtain systematic and comprehensive information on OA. Here, we also introduce a biogeochemical monitoring initiative for OA in Lombok with the established protocols. Improvement of many aspects, including analysis instruments, analysis methods, sample treatment, and sampling frequency will provide new insight into further research and monitoring of OA.

Controls on boron isotopes in a cold-water coral and the cost of resilience to ocean acidification

Coral skeletal growth is sensitive to environmental change and may be adversely impacted by an acidifying ocean. However, physiological processes can also buffer biomineralization from external conditions, providing apparent resilience to acidification in some species. These same physiological processes affect skeletal composition and can impact paleoenvironmental proxies. Understanding the mechanisms of coral calcification is thus crucial for predicting the vulnerability of different corals to ocean acidification and for accurately interpreting coral-based climate records. Here, using boron isotope (δ11B) measurements on cultured cold-water corals, we explain fundamental features of coral calcification and its sensitivity to environmental change. Boron isotopes are one of the most widely used proxies for past seawater pH, and we observe the expected sensitivity between δ11B and pH. Surprisingly, we also discover that coral δ11B is independently sensitive to seawater dissolved inorganic carbon (DIC). We can explain this new DIC effect if we introduce boric acid diffusion across cell membranes as a new flux within a geochemical model of biomineralization. This model independently predicts the sensitivity of the δ11B-pH proxy, without being trained to these data, even though calcifying fluid pH (pHCF) is constant. Boric acid diffusion can resolve why δ11B is a useful proxy across a range of calcifiers, including foraminifera, even when calcifying fluid pH differs from seawater. Our modeling shows that δ11B cannot be interpreted unequivocally as a direct tracer of pHCF. Constant pHCF implies similar calcification rates as seawater pH decreases, which can explain the resilience of some corals to ocean acidification. However, we show that this resilience has a hidden energetic cost such that calcification becomes less efficient in an acidifying ocean.

Responses of coral gastrovascular cavity pH during light and dark incubations to reduced seawater pH suggest species-specific responses to the effects of ocean acidification on calcification

Coral polyps have a fluid-filled internal compartment, the gastrovascular cavity (GVC). Respiration and photosynthesis cause large daily excursions in GVC oxygen concentration (O2) and pH, but few studies have examined how this correlates with calcification rates. We hypothesized that GVC chemistry can mediate and ameliorate the effects of decreasing seawater pH (pHSW) on coral calcification. Microelectrodes were used to monitor O2 and pH within the GVC of Montastraea cavernosa and Duncanopsammia axifuga (pH only) in both the light and the dark, and three pHSW levels (8.2, 7.9, and 7.6). At pHSW 8.2, GVC O2 ranged from ca. 0 to over 400% saturation in the dark and light, respectively, with transitions from low to high (and vice versa) within minutes of turning the light on or off. For all three pHSW treatments and both species, pHGVC was always significantly above and below pHSW in the light and dark, respectively. For M. cavernosa in the light, pHGVC reached levels of pH 8.4–8.7 with no difference among pHSW treatments tested; in the dark, pHGVC dropped below pHSW and even below pH 7.0 in some trials at pHSW 7.6. For D. axifuga in both the light and the dark, pHGVC decreased linearly as pHSW decreased. Calcification rates were measured in the light concurrent with measurements of GVC O2 and pHGVC. For both species, calcification rates were similar at pHSW 8.2 and 7.9 but were significantly lower at pHSW 7.6. Thus, for both species, calcification was protected from seawater acidification by intrinsic coral physiology at pHSW 7.9 but not 7.6. Calcification was not correlated with pHGVC for M. cavernosa but was for D. axifuga. These results highlight the diverse responses of corals to changes in pHSW, their varying abilities to control pHGVC, and consequently their susceptibility to ocean acidification.

The role of gastropod shell composition and microstructure in resisting dissolution caused by ocean acidification

Organisms, such as molluscs, that produce their hard parts from calcium carbonate are expected to show increased difficulties growing and maintaining their skeletons under ocean acidification (OA). Any loss of shell integrity increases vulnerability, as shells provide protection against predation, desiccation, and disease. Not all species show the same responses to OA, which may be due to the composition and microstructural arrangement of their shells. We explore the role of shell composition and microstructure in resisting dissolution caused by decreases in seawater pH using a combination of microCT scans, XRD analysis, and SEM imaging. Two gastropods with different shell compositions and microstructure, Tegula funebralis and Nucella ostrina, were exposed to simulated ocean acidification conditions for six months. Both species showed signs of dissolution on the exterior of their shells, but changes in density were significantly more pronounced in T. funebralis. XRD analysis indicated that the exterior layer of both shell types was made of calcite. T. funebralis may be more prone to dissolution because their outer fibrous calcite layer has more crystal edges and faces exposed, potentially increasing the surface area on which dissolution can occur. These results support a previous study where T. funebralis showed significant decreases in both shell growth and strength, but N. ostrina only showed slight reductions in shell strength, and unaffected growth. We suggest that microstructural arrangement of shell layers in molluscs, more so than their composition alone, is critical for determining the vulnerability of mollusc shells to OA.

Short-Term Response of Cytosolic to Inorganic Carbon Increase in Posidonia oceanica Leaf Cells.

The concentration of CO2 in the atmosphere has increased over the past 200 years and is expected to continue rising in the next 50 years at a rate of 3 ppm·year−1. This increase has led to a decrease in seawater pH that has changed inorganic carbon chemical speciation, increasing the dissolved . Posidonia oceanica is a marine angiosperm that uses as an inorganic carbon source for photosynthesis. An important side effect of the direct uptake of is the diminution of cytosolic Cl− (Cl−c) in mesophyll leaf cells due to the efflux through anion channels and, probably, to intracellular compartmentalization. Since anion channels are also permeable to we hypothesize that high , or even CO2, would also promote a decrease of cytosolic (). In this work we have used - and Cl−-selective microelectrodes for the continuous monitoring of the cytosolic concentration of both anions in P. oceanica leaf cells. Under light conditions, mesophyll leaf cells showed a of 5.7 ± 0.2 mM, which rose up to 7.2 ± 0.6 mM after 30 min in the dark. The enrichment of natural seawater (NSW) with 3 mM NaHCO3 caused both a decrease of 1 ± 0.04 mM and a decrease of 3.5 ± 0.1 mM. The saturation of NSW with 1000 ppm CO2 also produced a diminution of the , but lower (0.4 ± 0.07 mM). These results indicate that the rise of dissolved inorganic carbon ( or CO2) in NSW would have an effect on the cytosolic anion homeostasis mechanisms in P. oceanica leaf cells. In the presence of 0.1 mM ethoxyzolamide, the plasma membrane-permeable carbonic anhydrase inhibitor, the CO2-induced cytosolic diminution was much lower (0.1 ± 0.08 mM), pointing to as the inorganic carbon species that causes the cytosolic leak. The incubation of P. oceanica leaf pieces in 3 mM -enriched NSW triggered a short-term external net concentration increase consistent with the leak. As a consequence, the cytosolic diminution induced in high inorganic carbon could result in both the decrease of metabolic N flux and the concomitant biomass N impoverishment in P. oceanica and, probably, in other aquatic plants.