Net Calcification Rates Research Articles

Hidden cost of pH variability in seagrass beds on marine calcifiers under ocean acidification

Coastal ecosystems experience large environmental variability leading to local adaptation. The key role of variability and adaptation in modulating the biological sensitivity to ocean acidification is increasingly acknowledged. Monitoring and understanding the ecological niche at the right spatio-temporal scale is key to understand the sensitivity of any organism and ecosystems. However, the role of the variability in relevant carbonate chemistry parameters as a driver is often overlooked. For example, the balance between photosynthesis and respiration over the day/night cycle is leading to high pH/pCO2 variability in seagrass beds. We hypothesized that (i) the calcifying larvae of the sea urchin Echinus esculentus exposed to seagrass-driven variability would have some physiological mechanisms to respond to such variability; and (ii) these mechanisms would reach their limit under ocean acidification. We compared the presence and absence of the seagrass Zostera marina in flow through mesocosms fed with seawater with 4 pHs. The carbonate chemistry was monitored and biological response of a sea urchin larvae was documented over 3 weeks. Growth and net calcification rates were measured twice a day to encompass diurnal variability. Our results show that larvae growth rate significantly decreased with decreasing average pHT in both absence and presence of seagrass. Moreover, sea urchin larvae showed a slower growth rate in presence of seagrass, only visible in the lowest pH conditions. In addition, larvae raised in presence of seagrass, maximized calcification during the day, and lower their calcification during the night. In contrast, no significant difference was observed between day and night for the net calcification rate in larvae raised in absence of seagrass. Our results demonstrate the limit of local adaptation to the present range of variability under ocean acidification conditions. It also demonstrates that photosynthetic ecosystems such as seagrass may not play a role of refuge against future ocean acidification.

Physiological differences among cryptic species of the Mediterranean crustose coralline alga Lithophyllum stictiforme (Corallinales, Rhodophyta)

ABSTRACT The crustose coralline alga Lithophyllum stictiforme is an important reef builder, forming coralligenous concretions in the Mediterranean Sea. This algal species complex is composed of eight cryptic species (or clades). The objective of our study was to compare light-dependent physiological processes within and between the two most abundant cryptic species (C1 and C4) of L. stictiforme from the Bay of Marseille (France). Physiological rates of respiration, photosynthesis and calcification were measured using incubation chambers under various irradiance levels and in the dark during the summer period. We compared the physiology of the C1 and C4 cryptic species of L. stictiforme among three algal groups: C1 (the most common in the Marseille area) from 28 m depth, C4 from the same site and depth, and C1 from a deeper site at 45 m depth. We found both interspecific (between C1 and C4) and intraspecific (within C1) physiological differences. Photosynthetic parameters showed acclimation to light (or depth) with higher values of initial slope of photosynthetic light response curve (α) and lower values of saturating irradiance (Ek) and compensating irradiance (Ec) for C1 from 45 m than for C1 and C4 from 28 m depth. At the shallower depth, significant physiological differences were observed between C1 and C4 with a higher value of maximum rate of gross photosynthesis (Pg max) in C4, suggesting better ability to grow under high irradiance. Light and dark calcification rates differed only between C4 from 28 m and C1 from 45 m depth, being intermediate in C1 from 28 m depth. On a 24 h basis, diel net calcification rates were significantly higher in specimens from the shallower waters than from the deeper waters. Physiological differences suggest physiological plasticity within the regionally dominant cryptic species C1 and species-specific physiological traits between the cryptic species C1 and C4.

Seasonal upwelling conditions promote growth and calcification in reef-building coralline algae.

Crustose coralline algae (CCA) are important components of reef ecology contributing to reef framework construction. However, little is known about how seasonal upwelling systems influence growth and calcification of tropical CCA. We assessed marginal and vertical growth and net calcification rates of two dominant but morphologically different reef-building CCA, Porolithon antillarum and Lithophyllum cf. kaiseri, in a shallow coral reef of the Colombian Caribbean during upwelling and non-upwelling seasons. Growth and calcification rates varied seasonally with higher values during the upwelling compared to the non-upwelling (rainy) season. Annual vertical growth showed rates of 4.48 ± 1.58 and 4.31 ± 2.17 mm · y-1 , net calcification using crust growth estimates of 0.75 ± 0.30 g and 0.68 ± 0.60 g CaCO3 · cm-2 · y-1 and net calcification using the buoyant weight method of 1.49 ± 0.57 and 0.52 ± 0.11 g CaCO3 · cm-2 · y-1 in P. antillarum and L. kaiseri, respectively. Seawater temperature was inversely related with growth and calcification; however, complex oceanographic interactions between temperature and resource availability (e.g., light, nutrients, and CO2 ) are proposed to modulate CCA vital rates. Although CCA calcification rates are comparable to hard corals, CCA vertical accretion is much lower, suggesting that the main contribution of CCA to reef construction is via cementation processes. These results provide baseline data on CCA in the region and generate useful information for monitoring the impacts of environmental changes on tropical upwelling environments.

Temporal variability in seawater Sr/Ca ratios within a coral atoll as an indicator of marine calcifier community diversity

Coral reefs are marine environments where calcium carbonate production and dissolution along with photosynthesis and respiration can have a large impact on the daily biogeochemical cycles of seawater dissolved inorganic carbon (DIC), total alkalinity (TA), oxygen (O2), and major nutrients. Using seawater samples from Tetiaroa Atoll, French Polynesia, we apply isotope dilution thermal ionization mass spectrometry (ID-TIMS) to show that these daily cycles of variability extend to the dissolved seawater Sr/Ca ratio (Sr/Casw). Hourly measurements over a 27+ hour period indicate that Sr/Casw covaries with DIC and TA and exhibits up to a 2% relative difference compared to the regional Pacific Ocean source water Sr/Casw value of 8.53 mmol/mol. We hypothesize that the primary mechanism of hourly variability in Sr/Casw is carbonate precipitation/dissolution by corals and other marine calcifiers on the Tetiaroa reef, while sedimentary redox processes may significantly contribute to an apparent decoupling between net calcification rates estimated from TA compared to dissolved calcium [Ca]sw concentrations. Using a simple Rayleigh distillation model and a two-member mixing equation, we estimate the net partition coefficient of Sr (KD-Sr) for each hourly sample as a mixture of coral (aragonite; KD-Sr ∼ 1.06) and crustose coralline algae (high Mg-calcite; KD-Sr ∼ 0.39) contributions to the measured variability of [Ca]sw and Sr/Casw. As the high Mg-calcites of crustose coralline algae are (1) integral components to the structure and stability of the reef framework and (2) more soluble than aragonite under more acidic conditions, we propose that continued analyses of Sr/Casw within coral reef environments could be a useful tool for monitoring the ecological distribution and contributions of marine calcifiers on reefs as ocean warming and acidification intensify in response to anthropogenic climate change. Additionally, seasonal, interannual, and/or centennial-scale offsets between the Sr/Casw of open ocean source seawater and ambient reef seawater may have significant implications for interpreting past changes in sea surface temperatures derived from the coral Sr/Ca paleothermometer.

Seasonal Upwelling Conditions Modulate the Calcification Response of a Tropical Scleractinian Coral

Natural processes such as upwelling of deeper-water masses change the physical-chemical conditions of the water column creating localized ocean acidification events that can have an impact on the natural communities. This study was performed in a coral reef system of an archetypical bay within the Tayrona National Natural Park (PNNT) (Colombia), and aimed to quantify net calcification rates of a foundational coral species within a temporal context (6 months) taking into account the dynamics of seasonal upwelling that influence the study area. Net calcification rates of coral fragments were obtained in situ by the alkalinity anomaly technique in short-term incubations (~2.5 h). We found a significant effect of the upwelling on net calcification rates (Gnet) (p < 0.05) with an 42% increase in CaCO3 accretion compared to non-upwelling season. We found an increase in total alkalinity (AT) and dissolved inorganic carbon (DIC) with decreased aragonite saturation (Ωara) for the upwelling months, indicating an influence of the Subtropical Under Water mass (SAW) in the PNNT coral community. Significant negative correlations between net calcification with temperature and Ωara, which indicates a positive response of M. auretenra with the upwelling conditions, thus, acting as “enhancer” of resilience for coral calcification.

Production and accumulation of reef framework by calcifying corals and macroalgae on a remote Indian Ocean cay

Abstract. Coral reefs face increasing pressures in response to unprecedented rates of environmental change at present. The coral reef physical framework is formed through the production of calcium carbonate (CaCO3) and maintained by marine organisms, primarily hermatypic corals, and calcifying algae. The northern part of Western Australia, known as the Kimberley, has largely escaped land-based anthropogenic impacts and this study provides important metabolic data on reef-building organisms from an undisturbed set of marine habitats. From the reef platform of Browse Island, located on the mid-shelf just inside the 200 m isobath off the Kimberley coast, specimens of the dominant coral (six species) and algal (five species) taxa were collected and incubated ex situ in light and dark shipboard experimental mesocosms for 4 h to measure rates of calcification and production patterns of oxygen. During experimental light and dark incubations, all algae were net autotrophic producing 6 to 111 mmolO2m-2d-1. In contrast, most corals were net consumers of O2 with average net fluxes ranging from −42 to 47 mmolO2m-2d-1. The net change in pH was generally negative for corals and calcifying algae (−0.01 to −0.08 h−1). Resulting net calcification rates (1.9 to 9.9 gCaCO3m-2d-1) for corals and calcifying algae (Halimeda and Galaxura) were all positive and were strongly correlated with net O2 production. In intertidal habitats around Browse Island, estimated relative contributions of coral and Halimeda to the reef production of CaCO3 were similar at around 600 to 840 gm-2yr-1. The low reef platform had very low coral cover of < 3 % which made a smaller contribution to calcification of ∼ 240 gCaCO3m-2yr-1. Calcification on the subtidal reef slope was predominantly from corals, producing ∼ 1540 gCaCO3m-2yr-1, twice that of Halimeda. These data provide the first measures of community metabolism from the offshore reef systems of the Kimberley. The relative contributions of the main reef builders, in these undisturbed areas, to net community metabolism and CaCO3 production is important to understand exclusively climate-driven negative effects on tropical reefs.

Environmental memory gained from exposure to extreme pCO2 variability promotes coral cellular acid-base homeostasis.

Ocean acidification is a growing threat to coral growth and the accretion of coral reef ecosystems. Corals inhabiting environments that already endure extreme diel pCO2 fluctuations, however, may represent acidification-resilient populations capable of persisting on future reefs. Here, we examined the impact of pCO2 variability on the reef-building coral Pocillopora damicornis originating from reefs with contrasting environmental histories (variable reef flat versus stable reef slope) following reciprocal exposure to stable (218 ± 9) or variable (911 ± 31) diel pCO2 amplitude (μtam) in aquaria over eight weeks. Endosymbiont density, photosynthesis and net calcification rates differed between origins but not treatment, whereas primary calcification (extension) was affected by both origin and acclimatization to novel pCO2 conditions. At the cellular level, corals from the variable reef flat exhibited less intracellular pH (pHi) acidosis and faster pHi recovery rates in response to experimental acidification stress (pH 7.40) than corals originating from the stable reef slope, suggesting environmental memory gained from lifelong exposure to pCO2 variability led to an improved ability to regulate acid–base homeostasis. These results highlight the role of cellular processes in maintaining acidification resilience and suggest that prior exposure to pCO2 variability may promote more acidification-resilient coral populations in a changing climate.

Impacts of ocean warming on a reef-building coralline alga Amphiroa cf. fragilissima under high irradiance

Coralline algae, an important calcifying group, play vital roles in the primary productivity, reef frameworks construction, and carbon store. In this study, we investigated the responses of an articulated coralline alga Amphiroa cf. fragilissima to ocean warming under various light intensities. The relative growth rate significantly decreased under light or heat stress. When A. cf. fragilissima was exposed to high light intensity (120 μmol photons m−2 s−1) at 32°C, the relative growth rate was lowest, which reduced by 87% compared with that of group A1 (60 μmol photons m−2 s−1, 26°C). Meanwhile, a higher level of algal bleaching occurred when light intensity was 120 μmol photons m−2 s−1. Similarly, Fv/Fm and Chl-a content were negatively affected by light and heat stress, but they were more affected by light. Furthermore, the mineralogical responses to temperature and light were investigated. The net calcification rate declined from 92.27 (60 μmol photons m−2 s−1, 26°C) to 10.92 μmol CaCO3 g−1 DW day−1 (120 μmol photons m−2 s−1, 32°C). High temperature significantly decreased Ca content in live algal pigmented layer, whereas there were no significant differences in Ca content in the skeleton layer, implying that the pigmented layer could protect skeleton layer from mineral changes under ocean warming. The results revealed that A. cf. fragilissima was impaired by high light or thermal stress from various aspects, including growth, survival, photosynthesis, reproduction, and calcification. This study contributes to understand the effects of warming and light on coralline algae and provides a theoretical basis to protect the richness and diversity of calcified macroalgae.

Will daytime community calcification reflect reef accretion on future, degraded coral reefs?

Abstract. Coral bleaching events continue to drive the degradation of coral reefs worldwide, causing a shift in the benthic community from coral- to algae-dominated ecosystems. Critically, this shift may decrease the capacity of degraded coral reef communities to maintain net positive accretion during warming-driven stress events (e.g., reef-wide coral bleaching). Here we measured rates of net ecosystem calcification (NEC) and net ecosystem production (NEP) on a degraded coral reef lagoon community (coral cover < 10 % and algae cover > 20 %) during a reef-wide bleaching event in February 2020 at Heron Island on the Great Barrier Reef. We found that during this bleaching event, rates of NEP and NEC across replicate transects remained positive and did not change in response to bleaching. Repeated benthic surveys over a period of 20 d indicated an increase in the percent area of bleached coral tissue, corroborated by relatively low Symbiodiniaceae densities (∼ 0.6 × 106 cm−2) and dark-adapted photosynthetic yields in photosystem II of corals (∼ 0.5) sampled along each transect over this period. Given that a clear decline in coral health was not reflected in the overall NEC estimates, it is possible that elevated temperatures in the water column that compromise coral health enhanced the thermodynamic favorability for calcification in other ahermatypic benthic calcifiers. These data suggest that positive NEC on degraded reefs may not equate to the net positive accretion of a complex, three-dimensional reef structure in a future, warmer ocean. Critically, our study highlights that if coral cover continues to decline as predicted, NEC may no longer be an appropriate proxy for reef growth as the proportion of the NEC signal owed to ahermatypic calcification increases and coral dominance on the reef decreases.

Assessment of the juvenile vulnerability of symbiont-bearing giant clams to ocean acidification

Ocean acidification (OA) severely affects marine bivalves, especially their calcification processes. However, very little is known about the fate of symbiont-bearing giant clams in the acidified oceans, which hinders our ability to develop strategies to protect this ecologically and economically important group in coral reef ecosystems. Here, we explored the integrated juvenile responses of fluted giant clam Tridacna squamosa (Lamarck, 1819) to acidified seawater at different levels of biological organization. Our results revealed that OA did not cause a significant reduction in survival and shell growth performance, indicating that T. squamosa juveniles are tolerated to moderate acidification. Yet, significantly reduced net calcification rate demonstrated the calcifying physiology sensitivity to OA, in line with significant declines in symbiont photosynthetic yield and zooxanthellae density which in turn lowered the amount of energy supply for energetically expensive calcification processes. Subsequent transcriptome sequencing and comparative analysis of differentially expressed genes revealed that the regulation of calcification processes, such as transport of calcification substrates, acid-base regulation, synthesis of organic matrix in the calcifying fluid, as well as metabolic depression were the major response to OA. Taken together, the integration of physiological and molecular responses can provide a comprehensive understanding of how the early life history stages of giant clams respond to OA and make an important leap forward in assessing their fate under future ocean conditions.

Two temperate corals are tolerant to low pH regardless of previous exposure to natural CO2 vents

AbstractOcean acidification is perceived to be a major threat for many calcifying organisms, including scleractinian corals. Here, we investigate (1) whether past exposure to low pH environments associated with CO2 vents could increase corals tolerance to low pH and (2) whether zooxanthellate corals are more tolerant to low pH than azooxanthellate corals. To test these hypotheses, two Mediterranean colonial corals Cladocora caespitosa (zooxanthellate) and Astroides calycularis (azooxanthellate) were collected from CO2 vents and reference sites and incubated in the laboratory under present day (pH on the total scale, pHT 8.07) and low pH conditions (pHT 7.70). Rates of net calcification, dark respiration, and photosynthesis were monitored during a 6‐month experiment. Monthly net calcification was assessed every 27–35 d using the buoyant weight technique, whereas light and dark net calcification was estimated using the alkalinity anomaly technique during 1‐h incubations. Neither species showed any change in net calcification rates, respiration, and photosynthesis regardless of their environmental history, pH treatment, and trophic strategy. Our results indicate that C. caespitosa and As. calycularis could tolerate future ocean acidification conditions for at least 6 months. These results will aid in predicting species' future responses to ocean acidification, and thus improve the management and conservation of Mediterranean corals.

Exposure duration modulates the response of Caribbean corals to global change stressors

AbstractGlobal change, including rising temperatures and acidification, threatens corals globally. Although bleaching events reveal fine‐scale patterns of resilience, traits enabling persistence under global change remain elusive. We conducted a 95‐d controlled‐laboratory experiment investigating how duration of exposure to warming (~28, 31°C), acidification (pCO2 ~ 343 [present day], ~663 [end of century], ~3109 [extreme] μatm), and their combination influences physiology of reef‐building corals (Siderastrea siderea, Pseudodiploria strigosa) from two reef zones on the Belize Mesoamerican Barrier Reef System. Every 30 d, net calcification rate, host protein and carbohydrate, chlorophyll a, and symbiont density were quantified for the same coral individual to characterize acclimation potential under global change. Coral physiologies of the two species were differentially affected by stressors and exposure duration was found to modulate these responses. Siderastrea siderea exhibited resistance to end of century pCO2 and temperature stress, but calcification was negatively affected by extreme pCO2. However, S. siderea calcification rates remained positive after 95 d of extreme pCO2 conditions, suggesting acclimation. In contrast, P. strigosa was more negatively influenced by elevated temperatures, which reduced most physiological parameters. An exception was nearshore P. strigosa, which maintained calcification rates under elevated temperature, suggesting local adaptation to the warmer environment of their natal reef zone. This work highlights how tracking coral physiology across various exposure durations can capture acclimatory responses to global change stressors.

Juvenile Eastern Oysters More Resilient to Extreme Ocean Acidification than Their Mud Crab Predators

AbstractOcean acidification is predicted to impair marine calcifiers' abilities to produce shells and skeletons. We conducted laboratory experiments investigating the impacts of CO2‐induced ocean acidification (pCO2 = 478–519, 734–835, 8,980–9,567; Ωcalcite = 7.3–5.7, 5.6–4.3, 0.6–0.7) on calcification rates of two estuarine calcifiers involved in a classic predator‐prey model system: adult Panopeus herbstii (Atlantic mud crab) and juvenile Crassostrea virginica (eastern oyster). Both oyster and crab calcification rates significantly decreased at the highest pCO2 level. Notably, however, oysters maintained positive net calcification rates in the highest pCO2 treatment that was undersaturated with respect to calcite, while mud crabs exhibited net dissolution (i.e., net loss of shell mass) in calcite‐undersaturated conditions. Secondary electron imaging of oyster shells revealed minor microstructural alterations in the moderate‐pCO2 treatment, and major microstructural and macrostructural changes (including shell dissolution, delamination of periostracum) in the high‐pCO2 treatment. These results underscore the threat that ocean acidification poses for marine organisms that produce calcium carbonate shells, illustrate the strong biological control that some marine calcifiers exert over their shell‐building process, and shows that ocean acidification differentially impacts the crab and oyster species involved in this classical predator‐prey model system.

Irradiance, photosynthesis and elevated pCO2 effects on net calcification in tropical reef macroalgae

Calcifying tropical macroalgae produce sediment, build three-dimensional habitats, and provide substrate for invertebrate larvae on reefs. Thus, lower calcification rates under declining pH and increasing ocean pCO2, or ocean acidification, is a concern. In the present study, calcification rates were examined experimentally under predicted end-of-the-century seawater pCO2 (1116 μatm) and pH (7.67) compared to ambient controls (pCO2 409 μatm; pH 8.04). Nine reef macroalgae with diverse calcification locations, calcium carbonate structure, photophysiology, and site-specific irradiance were examined under light and dark conditions. Species included five from a high light patch reef on the Florida Keys Reef Tract (FKRT) and four species from low light reef walls on Little Cayman Island (LCI). Experiments on FKRT and LCI species were conducted at 500 and 50 μmol photons m−2 s−1 in situ irradiance, respectively. Calcification rates independent of photosystem-II (PSII) were also investigated for FKRT species. The most consistent negative effect of elevated pCO2 on calcification rates in the tropical macroalgae examined occurred in the dark. Most species (89%) had net calcification rates of zero or net dissolution in the dark at low pH. Species from the FKRT that sustained positive net calcification rates in the light at low pH also maintained ~30% of their net calcification rates without PSII at ambient pH. However, calcification rates in the light independent of PSII were not sustained at low pH. Regardless of these low pH effects, most FKRT species daily net calcification rates, integrating light/dark rates over a 24h period, were not significantly different between low and ambient pH. This was due to a 10-fold lower dark, compared to light, calcification rate, and a strong correspondence between calcification and photosynthetic rates. Interestingly, low-light species sustained calcification rates on par with high-light species without high rates of photosynthesis. Low-light species' morphology and physiology that promote high calcification rates at ambient pH, may increase their vulnerability to low pH. Our data indicate that the negative effect of elevated pCO2 and low pH on tropical macroalgae at the organismal level is their impact on dark net calcification, probably enhanced dissolution. However, elevated pCO2 and low pH effects on macroalgae daily calcification rates are greatest in species with lower net calcification rates in the light. Thus, macroalgae able to maintain high calcification rates in the light (high and low irradiance) at low pH, and/or sustain strong biotic control with high [H+] in the bulk seawater, are expected to dominate under global change.

Ocean acidification effects on calcification and dissolution in tropical reef macroalgae

Net calcification rates for coral reef and other calcifiers have been shown to decline as ocean acidification (OA) occurs. However, the role of calcium carbonate dissolution in lowering net calcification rates is unclear. The objective of this study was to distinguish OA effects on calcification and dissolution rates in dominant calcifying macroalgae of the Florida Reef Tract, including two rhodophytes (Neogoniolithon strictum, Jania adhaerens) and two chlorophytes (Halimeda scabra, Udotea luna). Two experiments were conducted: (1) to assess the difference in gross (45Ca uptake) versus net (total alkalinity anomaly) calcification rates in the light/dark and (2) to determine dark dissolution (45CaCO3), using pH levels predicted for the year 2100 and ambient pH. At low pH in the light, all species maintained gross calcification rates and most sustained net calcification rates relative to controls. Net calcification rates in the dark were ~84% lower than in the light. In contrast to the light, all species had lower net calcification rates in the dark at low pH with chlorophytes exhibiting net dissolution. These data are supported by the relationship (R2 = 0.82) between increasing total alkalinity and loss of 45Ca from pre-labelled 45CaCO3 thalli at low pH in the dark. Dark dissolution of 45CaCO3-labelled thalli was ~18% higher in chlorophytes than rhodophytes at ambient pH, and ~ twofold higher at low pH. Only Udotea, which exhibited dissolution in the light, also had lower daily calcification rates integrated over 24 h. Thus, if tropical macroalgae can maintain high calcification rates in the light, lower net calcification rates in the dark from dissolution may not compromise daily calcification rates. However, if organismal dissolution in the dark is additive to sedimentary carbonate losses, reef dissolution may be amplified under OA and contribute to erosion of the Florida Reef Tract and other reefs that exhibit net dissolution.

Giant clams in shallow reefs: UV-resistance mechanisms of Tridacninae in the Red Sea

The photosymbiosis of tropical giant clams (subfamily Tridacninae) with unicellular algae (Symbiodiniaceae) restricts their distribution to the sunlit, shallow waters of the euphotic zone where organisms are additionally exposed to potentially damaging levels of solar UV radiation. Metabolic and physiological responses of Red Sea Tridacna maxima clams, including net calcification and primary production, as well as valvometry (i.e., shell gaping behavior) were assessed when exposed to simulated high radiation levels received at 3 and 5 m underwater. The two levels of radiation included exposure treatments to photosynthetically active radiation (PAR; 400–700 nm) alone and to both, PAR and ultraviolet-B radiation (UV-B; 280–315 nm). The valvometry data obtained using flexible magnetic sensors indicated that specimens under PAR + UV-B exposure significantly reduced the proportion of their exposed mantle area, a potential photo-protective mechanism which, however, reduces the overall amount of PAR received by the algal symbionts. Consequently, specimens under PAR + UV-B displayed a slight, although non-significant, reduction in primary production rates but no signs of additional oxidative stress, changes in symbiont densities, chlorophyll content, or levels of mycosporine-like amino acids. Net calcification rates of T. maxima were not affected by exposure to UV-B; however, calcification was positively correlated with incident PAR levels. UV-B exposure changes the valvometry, reducing the exposed mantle area which consequently diminishes the available PAR for the photosymbionts. Still, T. maxima maintains high rates of primary production and net calcification, even under high levels of UV-B. This provides experimental support for a recently described, effective UV-defensive mechanism in Tridacninae, in which the photonic cooperation of the associated algal symbionts and giant clam iridocytes is assumed to establish optimal conditions for the photosynthetic performance of the clams’ symbionts.

Increased irradiance availability mitigates the physiological performance of species of the calcifying green macroalga Halimeda in response to ocean acidification

Although negative responses of tropical calcifying organisms to ocean acidification have been widely reported, the modulating potential of irradiance combined with elevated pCO2 has not been well studied. In this study, the interactive effects of elevated pCO2 and irradiance availability on the physiology of calcifying macroalgae Halimeda cylindracea and Halimeda lacunalis were investigated using a fully factorial, 28-day aquaria coupling experiment. The results of the present study demonstrate that elevated pCO2 negatively influences growth, photosynthesis, calcification and other physiological processes of both Halimeda species. However, these negative effects could be mitigated to some extent by increased irradiance availability. Specific growth rate (SGR), net calcification rates (Gnet) and maximum quantum yield (Fv/Fm) decreased significantly by 6.84%–86.70%, 51.78%–62.29% and 2.37%–28.91% in elevated pCO2 treatments. However, SGR, Gnet and Fv/Fm increased by 3.39%–84.78%, 29.61%–40.68% and 1.68%–6.92% in high irradiance conditions, respectively. Chl-a in elevated pCO2 treatments was 7.75%–61.25% lower than ambient pCO2 conditions, while the carotenoid content increased by 12.12%–57.45% in low irradiance conditions from day 20–28. Malondialdehyde (MDA) content was higher in elevated pCO2 treatments. However, there was also a two- to four-fold increase in proline content in elevated pCO2 treatments. Tissue total organic carbon (TCorg) and nitrogen (TN) were positively correlated to CO2 enrichment. The results of the current study suggested that elevated pCO2 negatively influenced the physiological responses of Halimeda, while increased irradiance availability may enhance the metabolic performance in response to ocean acidification.

Physiological performance of three calcifying green macroalgae Halimeda species in response to altered seawater temperatures

The effects of seawater temperature on the physiological performance of three Halimeda species were studied for a period of 28 d. Five treatments were established for Halimeda cylindracea, Halimeda opuntia and Halimeda lacunalis, in triplicate aquaria representing a factorial temperature with 24°C, 28°C, 32°C, 34°C and 36°C, respectively. The average Fv/Fm of these species ranged from 0.732 to 0.756 between 24°C and 32°C but declined sharply between 34°C (0.457±0.035) and 36°C (0.122±0.014). Calcification was highest at 28°C, with net calcification rates (Gnet) of (20.082±2.482) mg/(g•d), (12.825±1.623) mg/(g•d) and (6.411±1.029) mg/(g•d) for H. cylindracea, H. opuntia and H. lacunalis, respectively. Between 24°C and 32°C, the specific growth rate (SGR) of H. lacunalis (0.079%–0.110% d–1) was lower than that of H. cylindracea (0.652%–1.644% d–1) and H. opuntia (0.360%–1.527% d–1). Three Halimeda species gradually bleached at 36°C during the study period. Malondialdehyde (MDA) and proline levels in tissues of the three Halimeda were higher in 34–36°C than those in 24–32°C. The results indicate that seawater temperature with range of 24–32°C could benefit the growth and calcification of these Halimeda species, however, extreme temperatures above 34°C have negative impacts. The measured physiological parameters also revealed that H. cylindracea and H. opuntia displayed broader temperature tolerance than H. lacunalis.

Blue Mussel (Genus Mytilus) Transcriptome Response to Simulated Climate Change in the Gulf of Maine

The biogeochemistry of the Gulf of Maine (GOM) is rapidly changing in response to the changing climate, including rising temperatures, acidification, and declining primary productivity. These impacts are projected to worsen over the next 100 y and will apply selective pressure on populations of marine calcifiers. This study investigates the transcriptome expression response to these changes in ecologically and economically important marine calcifiers, blue mussels. Wild mussels (Mytilus edulis and Mytilus trossulus) were sampled from sites spanning the GOM and exposed to two different biogeochemical water conditions: (1) present-day conditions in the GOM and (2) simulated future conditions, which included elevated temperature, increased acidity, and decreased food supply. Patterns of gene expression were measured using RNA sequencing from 24 mussel samples and contrasted between ambient and future conditions. The net calcification rate, a trait predicted to be under climate-induced stress, was measured for each individual over a 2-wk exposure period and used as a covariate along with gene expression patterns. Generalized linear models, with and without the calcification rate, were used to identify differentially expressed transcripts between ambient and future conditions. The comparison revealed transcripts that likely comprise a core stress response characterized by the induction of molecular chaperones, genes involved in aerobic metabolism, and indicators of cellular stress. Furthermore, the model contrasts revealed transcripts that may be associated with individual variation in calcification rate and suggest possible biological processes that may have downstream effects on calcification phenotypes, such as zinc-ion binding and protein degradation. Overall, these findings contribute to the understanding of blue mussel adaptive responses to imminent climate change and suggest metabolic pathways are resilient in variable environments.

Linking Internal Carbonate Chemistry Regulation and Calcification in Corals Growing at a Mediterranean CO2 Vent

Corals exert a strong biological control over their calcification processes, but there is a lack of knowledge on their capability of long-term acclimatization to ocean acidification. We used a dual geochemical proxy approach to estimate the calcifying fluid pH (pHcf) and carbonate chemistry of a Mediterranean coral (Balanophyllia europaea) naturally growing along a pH gradient (range: pHTS 8.07-7.74). The pHcf derived from skeletal boron isotopic composition (11B) was 0.3-0.6 units above seawater values and homogeneous along the gradient (mean  sem: Site 1 = 8.39  0.03, Site 2 = 8.34  0.03, Site 3 = 8.34  0.02). Also carbonate ion concentration derived from B/Ca was homogeneous (mean  sem (µmol kg-1): Site 1 = 579  34, Site 2 = 541  27, Site 3 = 568  30) regardless of seawater pH. Furthermore, gross calcification rate (GCR, mass of CaCO3 deposited on the skeletal unit area per unit of time), estimated by a “bio-inorganic model” (IpHRAC), was homogeneous with decreasing pH. The homogeneous GCR, internal pH and carbonate chemistry confirm that the features of the ‘building blocks’ – the fundamental structural components - produced by the biomineralization process were substantially unaffected by increased acidification. Furthermore, the pH up-regulation observed in this study could potentially explain the previous hypothesis that less ‘building blocks’ are produced with increasing acidification ultimately leading to increased skeletal porosity and to reduced net calcification rate obtained by including the total volume of the pore space. In fact, assuming that the available energy at the three sites is the same, this energy at the low pH sites could be partitioned among fewer calicoblastic cells that consume more energy given the larger difference between external and internal pH compared to the control, leading to the production of less building blocks (i.e., formation of pores inside the skeleton structure, determining increased porosity). However, we cannot exclude that also dissolution may play a role in increasing porosity. Thus, the ability of scleractinian corals to maintain elevated pHcf relative to ambient seawater might not always be sufficient to counteract declines in net calcification under ocean acidification scenarios.