Coral calcification responds to seawater acidification: a working hypothesis towards a physiological mechanism

The decrease in the saturation state of seawater, Ω, following seawater acidification, is believed to be the main factor leading to a decrease in the calcification of marine organisms. To provide a physiological explanation for this phenomenon, the effect of seawater acidification was studied on the calcification and photosynthesis of the scleractinian tropical coral Stylophora pistillata. Coral nubbins were incubated for 8 days at three different pH (7.6, 8.0, and 8.2). To differentiate between the effects of the various components of the carbonate chemistry (pH, CO₃²⁻, HCO₃⁻, CO₂, Ω), tanks were also maintained under similar pH, but with 2-mM HCO₃⁻ added to the seawater. The addition of 2-mM bicarbonate significantly increased the photosynthesis in S. pistillata, suggesting carbon-limited conditions. Conversely, photosynthesis was insensitive to changes in pH and pCO₂. Seawater acidification decreased coral calcification by ca. 0.1-mg CaCO₃ g⁻¹ d⁻¹ for a decrease of 0.1 pH units. This correlation suggested that seawater acidification affected coral calcification by decreasing the availability of the CO₃²⁻ substrate for calcification. However, the decrease in coral calcification could also be attributed either to a decrease in extra- or intracellular pH or to a change in the buffering capacity of the medium, impairing supply of CO₃²⁻ from HCO₃⁻.     

The Physiological Response of Two Green Calcifying Algae from the Great Barrier Reef towards High Dissolved Inorganic and Organic Carbon (DIC and DOC) Availability

Increasing dissolved inorganic carbon (DIC) concentrations associated with ocean acidification can affect marine calcifiers, but local factors, such as high dissolved organic carbon (DOC) concentrations through sewage and algal blooms, may interact with this global factor. For calcifying green algae of the genus Halimeda, a key tropical carbonate producer that often occurs in coral reefs, no studies on these interactions have been reported. These data are however urgently needed to understand future carbonate production. Thus, we investigated the independent and combined effects of DIC (pCO₂ 402 μatm/ pH_tot 8.0 and 996 μatm/ pH_tot 7.7) and DOC (added as glucose in 0 and 294 μmol L^-1) on growth, calcification and photosynthesis of H. macroloba and H. opuntia from the Great Barrier Reef in an incubation experiment over 16 days. High DIC concentrations significantly reduced dark calcification of H. opuntia by 130 % and led to net dissolution, but did not affect H. macroloba. High DOC concentrations significantly reduced daily oxygen production of H. opuntia and H. macroloba by 78 % and 43 %, respectively, and significantly reduced dark calcification of H. opuntia by 70%. Combined high DIC and DOC did not show any interactive effects for both algae, but revealed additive effects for H. opuntia where the combination of both factors reduced dark calcification by 162 % compared to controls. Such species-specific differences in treatment responses indicate H. opuntia is more susceptible to a combination of high DIC and DOC than H. macroloba. From an ecological perspective, results further suggest a reduction of primary production for Halimeda-dominated benthic reef communities under high DOC concentrations and additional decreases of carbonate accretion under elevated DIC concentrations, where H. opuntia dominates the benthic community. This may reduce biogenic carbonate sedimentation rates and hence the buffering capacity against further ocean acidification.     

One‐year experiment on the physiological response of the Mediterranean crustose coralline alga,Lithophyllum cabiochae,to elevated pCO2 and temperature

The response of respiration, photosynthesis, and calcification to elevated pCO₂ and temperature was investigated in isolation and in combination in the Mediterranean crustose coralline alga Lithophyllum cabiochae. Algae were maintained in aquaria during 1 year at near‐ambient conditions of irradiance, at ambient or elevated temperature (+3°C), and at ambient (ca. 400 μatm) or elevated pCO₂ (ca. 700 μatm). Respiration, photosynthesis, and net calcification showed a strong seasonal pattern following the seasonal variations of temperature and irradiance, with higher rates in summer than in winter. Respiration was unaffected by pCO₂ but showed a general trend of increase at elevated temperature at all seasons, except in summer under elevated pCO₂. Conversely, photosynthesis was strongly affected by pCO₂ with a decline under elevated pCO₂ in summer, autumn, and winter. In particular, photosynthetic efficiency was reduced under elevated pCO₂. Net calcification showed different responses depending on the season. In summer, net calcification increased with rising temperature under ambient pCO₂ but decreased with rising temperature under elevated pCO₂. Surprisingly, the highest rates in summer were found under elevated pCO₂ and ambient temperature. In autumn, winter, and spring, net calcification exhibited a positive or no response at elevated temperature but was unaffected by pCO₂. The rate of calcification of L. cabiochae was thus maintained or even enhanced under increased pCO₂. However, there is likely a trade‐off with other physiological processes. For example, photosynthesis declines in response to increased pCO₂ under ambient irradiance. The present study reports only on the physiological response of healthy specimens to ocean warming and acidification, however, these environmental changes may affect the vulnerability of coralline algae to other stresses such as pathogens and necroses that can cause major dissolution, which would have critical consequence for the sustainability of coralligenous habitats and the budgets of carbon and calcium carbonate in coastal Mediterranean ecosystems.     

Amelioration of ocean acidification and warming effects through physiological buffering of a macroalgae

Concurrent anthropogenic global climate change and ocean acidification are expected to have a negative impact on calcifying marine organisms. While knowledge of biological responses of organisms to oceanic stress has emerged from single‐species experiments, these do not capture ecologically relevant scenarios where the potential for multi‐organism physiological interactions is assessed. Marine algae provide an interesting case study, as their photosynthetic activity elevates pH in the surrounding microenvironment, potentially buffering more acidic conditions for associated epiphytes. We present findings that indicate increased tolerance of an important epiphytic foraminifera, Marginopora vertebralis, to the effects of increased temperature (±3°C) and pCO₂ (~1,000 µatm) when associated with its common algal host, Laurencia intricata. Specimens of M. vertebralis were incubated for 15 days in flow‐through aquaria simulating current and end‐of‐century temperature and pH conditions. Physiological measures of growth (change in wet weight), calcification (measured change in total alkalinity in closed bottles), photochemical efficiency (Fv/Fm), total chlorophyll, photosynthesis (oxygen flux), and respiration were determined. When incubated in isolation, M. vertebralis exhibited reduced growth in end‐of‐century projections of ocean acidification conditions, while calcification rates were lowest in the high‐temperature, low‐pH treatment. Interestingly, association with L. intricata ameliorated these stress effects with the growth and calcification rates of M. vertebralis being similar to those observed in ambient conditions. Total chlorophyll levels in M. vertebralis decreased when in association with L. intricata, while maximum photochemical efficiency increased in ambient conditions. Net production estimates remained similar between M. vertebralis in isolation and in association with L. intricata, although both production and respiration rates of M. vertebralis were significantly higher when associated with L. intricata. These results indicate that the association with L. intricata increases the resilience of M. vertebralis to climate change stress, providing one of the first examples of physiological buffering by a marine alga that can ameliorate the negative effects of changing ocean conditions.     

Increased temperature mitigates the effects of ocean acidification on the calcification of juvenile Pocillopora damicornis,but at a cost

This study tested the interactive effects of increased seawater temperature and CO₂ partial pressure (pCO₂) on the photochemistry, bleaching, and early growth of the reef coral Pocillopora damicornis. New recruits were maintained at ambient or high temperature (29 or 30.8 °C) and pCO₂ (~ 500 and ~ 1100 μatm) in a full-factorial experiment for 3 weeks. Neither a sharp decline in photochemical efficiency (Fv/Fm) nor evident bleaching was observed at high temperature and/or high pCO₂. Furthermore, elevated temperature greatly promoted lateral growth and calcification, while polyp budding exhibited temperature-dependent responses to pCO₂. High pCO₂ depressed calcification by 28% at ambient temperature, but did not impact calcification at 30.8 °C. Interestingly, elevated temperature in concert with high pCO₂ significantly retarded the budding process. These results suggest that increased temperature can mitigate the adverse effects of acidification on the calcification of juvenile P. damicornis, but at a substantial cost to asexual budding.

     

Nutrient availability affects the response of the calcifying chlorophyte Halimeda opuntia (L.) J.V. Lamouroux to low pH

Atmospheric carbon dioxide emissions cause a decrease in the pH and aragonite saturation state of surface ocean water. As a result, calcifying organisms are expected to suffer under future ocean conditions, but their physiological responses may depend on their nutrient status. Because many coral reefs experience high inorganic nutrient loads or seasonal changes in nutrient availability, reef organisms in localized areas will have to cope with elevated carbon dioxide and changes in inorganic nutrients. Halimeda opuntia is a dominant calcifying primary producer on coral reefs that contributes to coral reef accretion. Therefore, we investigated the carbon and nutrient balance of H. opuntia exposed to elevated carbon dioxide and inorganic nutrients. We measured tissue nitrogen, phosphorus and carbon content as well as the activity of enzymes involved in inorganic carbon uptake and nitrogen assimilation (external carbonic anhydrase and nitrate reductase, respectively). Inorganic carbon content was lower in algae exposed to high CO₂, but calcification rates were not significantly affected by CO₂ or inorganic nutrients. Organic carbon was positively correlated to external carbonic anhydrase activity, while inorganic carbon showed the opposite correlation. Carbon dioxide had a significant effect on tissue nitrogen and organic carbon content, while inorganic nutrients affected tissue phosphorus and N:P ratios. Nitrate reductase activity was highest in algae grown under elevated CO₂ and inorganic nutrient conditions and lowest when phosphate was limiting. In general, we found that enzymatic responses were strongly influenced by nutrient availability, indicating its important role in dictating the local responses of the calcifying primary producer H. opuntia to ocean acidification.     

Larvae of the coral eating crown‐of‐thorns starfish,Acanthaster planci in a warmer‐high CO2 ocean

Outbreaks of crown‐of‐thorns starfish (COTS), Acanthaster planci, contribute to major declines of coral reef ecosystems throughout the Indo‐Pacific. As the oceans warm and decrease in pH due to increased anthropogenic CO₂ production_, coral reefs are also susceptible to bleaching, disease and reduced calcification. The impacts of ocean acidification and warming may be exacerbated by COTS predation, but it is not known how this major predator will fare in a changing ocean. Because larval success is a key driver of population outbreaks, we investigated the sensitivities of larval A. planci to increased temperature (2–4 °C above ambient) and acidification (0.3–0.5 pH units below ambient) in flow‐through cross‐factorial experiments (3 temperature × 3 pH/pCO₂ levels). There was no effect of increased temperature or acidification on fertilization or very early development. Larvae reared in the optimal temperature (28 °C) were the largest across all pH treatments. Development to advanced larva was negatively affected by the high temperature treatment (30 °C) and by both experimental pH levels (pH 7.6, 7.8). Thus, planktonic life stages of A. planci may be negatively impacted by near‐future global change. Increased temperature and reduced pH had an additive negative effect on reducing larval size. The 30 °C treatment exceeded larval tolerance regardless of pH. As 30 °C sea surface temperatures may become the norm in low latitude tropical regions, poleward migration of A. planci may be expected as they follow optimal isotherms. In the absence of acclimation or adaptation, declines in low latitude populations may occur. Poleward migration will be facilitated by strong western boundary currents, with possible negative flow‐on effects on high latitude coral reefs. The contrasting responses of the larvae of A. planci and those of its coral prey to ocean acidification and warming are considered in context with potential future change in tropical reef ecosystems.     

Microenvironmental changes support evidence of photosynthesis and calcification inhibition in Halimeda under ocean acidification and warming

The effects of elevated CO₂ and temperature on photosynthesis and calcification of two important calcifying reef algae (Halimeda macroloba and Halimeda cylindracea) were investigated with O₂ microsensors and chlorophyll a fluorometry through a combination of two pCO₂ (400 and 1,200 μatm) and two temperature treatments (28 and 32 °C) equivalent to the present and predicted conditions during the 2100 austral summer. Combined exposure to pCO₂ and elevated temperature impaired calcification and photosynthesis in the two Halimeda species due to changes in the microenvironment around the algal segments and a reduction in physiological performance. There were no significant changes in controls over the 5-week experiment, but there was a 50–70 % decrease in photochemical efficiency (maximum quantum yield), a 70–80 % decrease in O₂ production and a threefold reduction in calcification rate in the elevated CO₂ and high temperature treatment. Calcification in these species is closely coupled with photosynthesis, such that a decrease in photosynthetic efficiency leads to a decrease in calcification. Although pH seems to be the main factor affecting Halimeda species, heat stress also has an impact on their photosystem II photochemical efficiency. There was a strong combined effect of elevated CO₂ and temperature in both species, where exposure to elevated CO₂ or temperature alone decreased photosynthesis and calcification, but exposure to both elevated CO₂ and temperature caused a greater decline in photosynthesis and calcification than in each stress individually. Our study shows that ocean acidification and ocean warming are drivers of calcification and photosynthesis inhibition in Halimeda. Predicted climate change scenarios for 2100 would therefore severely affect the fitness of Halimeda, which can result in a strongly reduced production of carbonate sediments on coral reefs under such changed climate conditions.     

Effects of elevated pCO2 on the metabolism of a temperate rhodolith Lithothamnion corallioides grown under different temperatures

Coralline algae are considered among the most sensitive species to near future ocean acidification. We tested the effects of elevated pCO₂ on the metabolism of the free‐living coralline alga Lithothamnion corallioides (“maerl”) and the interactions with changes in temperature. Specimens were collected in North Brittany (France) and grown for 3 months at pCO₂ of 380 (ambient pCO₂), 550, 750, and 1000 μatm (elevated pCO₂) and at successive temperatures of 10°C (ambient temperature in winter), 16°C (ambient temperature in summer), and 19°C (ambient temperature in summer +3°C). At each temperature, gross primary production, respiration (oxygen flux), and calcification (alkalinity flux) rates were assessed in the light and dark. Pigments were determined by HPLC. Chl a, carotene, and zeaxanthin were the three major pigments found in L. corallioides thalli. Elevated pCO₂ did not affect pigment content while temperature slightly decreased zeaxanthin and carotene content at 10°C. Gross production was not affected by temperature but was significantly affected by pCO₂ with an increase between 380 and 550 μatm. Light, dark, and diel (24 h) calcification rates strongly decreased with increasing pCO₂ regardless of the temperature. Although elevated pCO₂ only slightly affected gross production in L. corallioides, diel net calcification was reduced by up to 80% under the 1,000 μatm treatment. Our findings suggested that near future levels of CO₂ will have profound consequences for carbon and carbonate budgets in rhodolith beds and for the sustainability of these habitats.     

Effects of increased pCO2 on zinc uptake and calcification in the tropical coral Stylophora pistillata

Zinc (Zn) is an essential element for corals. We investigated the effects of ocean acidification on zinc incorporation, photosynthesis, and gross calcification in the scleractinian coral Stylophora pistillata. Colonies were maintained at normal pH_T (8.1) and at two low-pH conditions (7.8 and 7.5) for 5 weeks. Corals were exposed to ⁶⁵Zn dissolved in seawater to assess uptake rates. After 5 weeks, corals raised at pH_T (8.1) exhibited higher ⁶⁵Zn activity in the coral tissue and skeleton, compared with corals raised at a lower pH. Photosynthesis, photosynthetic efficiency, and gross calcification, measured by ⁴⁵Ca incorporation, were however unchanged even at the lowest pH.     

Species‐specific responses to climate change and community composition determine future calcification rates of Florida Keys reefs

Anthropogenic climate change compromises reef growth as a result of increasing temperatures and ocean acidification. Scleractinian corals vary in their sensitivity to these variables, suggesting species composition will influence how reef communities respond to future climate change. Because data are lacking for many species, most studies that model future reef growth rely on uniform scleractinian calcification sensitivities to temperature and ocean acidification. To address this knowledge gap, calcification of twelve common and understudied Caribbean coral species was measured for two months under crossed temperatures (27, 30.3 °C) and CO₂ partial pressures (pCO₂) (400, 900, 1300 μatm). Mixed‐effects models of calcification for each species were then used to project community‐level scleractinian calcification using Florida Keys reef composition data and IPCC AR5 ensemble climate model data. Three of the four most abundant species, Orbicella faveolata, Montastraea cavernosa, and Porites astreoides, had negative calcification responses to both elevated temperature and pCO₂. In the business‐as‐usual CO₂ emissions scenario, reefs with high abundances of these species had projected end‐of‐century declines in scleractinian calcification of >50% relative to present‐day rates. Siderastrea siderea, the other most common species, was insensitive to both temperature and pCO₂ within the levels tested here. Reefs dominated by this species had the most stable end‐of‐century growth. Under more optimistic scenarios of reduced CO₂ emissions, calcification rates throughout the Florida Keys declined <20% by 2100. Under the most extreme emissions scenario, projected declines were highly variable among reefs, ranging 10–100%. Without considering bleaching, reef growth will likely decline on most reefs, especially where resistant species like S. siderea are not already dominant. This study demonstrates how species composition influences reef community responses to climate change and how reduced CO₂ emissions can limit future declines in reef calcification.     

Combined effects of global climate change and nutrient enrichment on the physiology of three temperate maerl species

Made up of calcareous coralline algae, maerl beds play a major role as ecosystem engineers in coastal areas throughout the world. They undergo strong anthropogenic pressures, which may threaten their survival. The aim of this study was to gain insight into the future of maerl beds in the context of global and local changes. We examined the effects of rising temperatures (+3°C) and ocean acidification (?0.3 pH units) according to temperature and pH projections (i.e., the RCP 8.5 scenario), and nutrient (N and P) availability on three temperate maerl species (Lithothamnion corallioides, Phymatolithon calcareum, and Lithophyllum incrustans) in the laboratory in winter and summer conditions. Physiological rates of primary production, respiration, and calcification were measured on all three species in each treatment and season. The physiological response of maerl to global climate change was species‐specific and influenced by seawater nutrient concentrations. Future temperature–pH scenario enhanced maximal gross primary production rates in P. calcareum in winter and in L. corallioides in both seasons. Nevertheless, both species suffered an impairment of light harvesting and photoprotective mechanisms in winter. Calcification rates at ambient light intensity were negatively affected by the future temperature–pH scenario in winter, with net dissolution observed in the dark in L. corallioides and P. calcareum under low nutrient concentrations. Nutrient enrichment avoided dissolution under future scenarios in winter and had a positive effect on L. incrustans calcification rate in the dark in summer. In winter conditions, maximal calcification rates were enhanced by the future temperature–pH scenario on the three species, but P. calcareum suffered inhibition at high irradiances. In summer conditions, the maximal calcification rate dropped in L. corallioides under the future global climate change scenario, with a potential negative impact on CaCO₃ budget for maerl beds in the Bay of Brest where this species is dominant. Our results highlight how local changes in nutrient availability or irradiance levels impact the response of maerl species to global climate change and thus point out how it is important to consider other abiotic parameters in order to develop management policies capable to increase the resilience of maerl beds under the future global climate change scenario.     

Distributional patterns of mobile fauna associated with Halimeda on the Tiahura coral-reef complex (Moorea,French Polynesia)

This study deals with the mobile fauna living associated with the turfs of three Halimeda species [H. incrassata, H. opuntia (three forms) and H. macroloba] from the Tiahura Reef complex. Ten 0.05 m² test areas of each Halimeda form have been randomly sampled from each geomorphological reef unit in order to obtain the specific abundance and biomass of different faunistic groups and species living within Halimeda populations. Then the raw data have been classified using the correspondence factor analysis to highlight the relationships between faunistic groups, or species, and the different Halimeda forms studied. The distributional patterns of the faunal communities seems to be controlled by the morphological features of the host-alga. The complex arborescent structure of the rhyzophytic H. incrassata species shelters fauna three times higher in abundance than other Halimeda heads. This fauna is mostly composed of a highly rich community of microcrustaceans and a very diversified community of Syllidae polychaetes. H. macroloba living on the outer reef flat retains a lot of small Nereidae polychaetes and a rich microgastropod assemblage. Dense H. opuntia tussocks (type A) on the fringing-reef and isolated H. opuntia fronds of pendulous chains (type B) on the barrier reef as well as beds of scattered H. opuntia (type C) on the outer reef flat provide environment for larger animals, and according to their zone of life, they retain very different faunal associations. Some ophiuroids, and some fish, collected in H. opuntia A and C, are newly reported from the Society Islands, French Polynesia, and the Pacific province. One Brachyurid species is presumably undescribed.     

Photosynthesis and calcification in the calcifying algae Halimeda discoidea studied with microsensors

With microsensors, we measured the steady‐state microprofiles of O₂, pH and Ca²⁺ on the topside of young segments of Halimeda discoidea, as well as the surface dynamics upon light–dark shifts. The effect of several inhibitors was studied. The steady‐state measurements showed that under high light intensity, calcium and protons were taken up, while O₂ was produced. In the dark, O₂ was consumed, the pH decreased to below seawater level and Ca²⁺ uptake was reduced to 50%. At low light intensity (12 mmol photons m^‐2 s^‐1), Ca²⁺ efflux was observed. Upon light–dark shifts, a complicated pattern of both the pH and calcium surface dynamics was observed. Illumination caused an initial pH decrease, followed by a gradual pH increase: this indicated that the surface pH of H. discoidea is determined by more than one light‐induced process. When photosynthesis was inhibited by dichlorophenyl dimethyl urea (DCMU), a strong acidification was observed upon illumination. The nature and physiological function of this putative pump is not known. The calcium dynamics followed all pH dynamics closely, both in the presence and absence of DCMU. The Ca‐channel blockers verapamil and nifedipine had no effect on the Ca²⁺ dynamics and steady‐state profiles. Thus, in H. discoidea, calcification is not regulated by the alga, but is a consequence of pH increase during photosynthesis. Acetazolamide had no effect on photosynthesis, whereas ethoxyzolamide inhibited photosynthesis at higher light intensities. Therefore, all carbonic anhydrase activity is intracellular. Carbonic anhydrase is required to alleviate the CO₂ limitation. Calcification cannot supply sufficient protons and CO₂ to sustain photosynthesis.     

Stress-tolerant corals of Florida Bay are vulnerable to ocean acidification

In situ calcification measurements tested the hypothesis that corals from environments (Florida Bay, USA) that naturally experience large swings in pCO₂ and pH will be tolerant or less sensitive to ocean acidification than species from laboratory experiments with less variable carbonate chemistry. The pCO₂ in Florida Bay varies from summer to winter by several hundred ppm roughly comparable to the increase predicted by the end of the century. Rates of net photosynthesis and calcification of two stress-tolerant coral species, Siderastrea radians and Solenastrea hyades, were measured under the prevailing ambient chemical conditions and under conditions amended to simulate a pH drop of 0.1–0.2 units at bimonthly intervals over a 2-yr period. Net photosynthesis was not changed by the elevation in pCO₂ and drop in pH; however, calcification declined by 52 and 50 % per unit decrease in saturation state, respectively. These results indicate that the calcification rates of S. radians and S. hyades are just as sensitive to a reduction in saturation state as coral species that have been previously studied. In other words, stress tolerance to temperature and salinity extremes as well as regular exposure to large swings in pCO₂ and pH did not make them any less sensitive to ocean acidification. These two species likely survive in Florida Bay in part because they devote proportionately less energy to calcification than most other species and the average saturation state is elevated relative to that of nearby offshore water due to high rates of primary production by seagrasses.     

Adaptive evolution in the coccolithophore Gephyrocapsa oceanica following 1,000 generations of selection under elevated CO2

Coccolithophores are important oceanic primary producers not only in terms of photosynthesis but also because they produce calcite plates called coccoliths. Ongoing ocean acidification associated with changing seawater carbonate chemistry may impair calcification and other metabolic functions in coccolithophores. While short‐term ocean acidification effects on calcification and other properties have been examined in a variety of coccolithophore species, long‐term adaptive responses have scarcely been documented, other than for the single species Emiliania huxleyi. Here, we investigated the effects of ocean acidification on another ecologically important coccolithophore species, Gephyrocapsa oceanica, following 1,000 generations of growth under elevated CO₂ conditions (1,000 μatm). High CO₂‐selected populations exhibited reduced growth rates and enhanced particulate organic carbon (POC) and nitrogen (PON) production, relative to populations selected under ambient CO₂ (400 μatm). Particulate inorganic carbon (PIC) and PIC/POC ratios decreased progressively throughout the selection period in high CO₂‐selected cell lines. All of these trait changes persisted when high CO₂‐grown populations were moved back to ambient CO₂ conditions for about 10 generations. The results suggest that the calcification of some coccolithophores may be more heavily impaired by ocean acidification than previously predicted based on short‐term studies, with potentially large implications for the ocean's carbon cycle under accelerating anthropogenic influences.     

Interactive effects of ocean acidification and warming on coral reef associated epilithic algal communities under past,present-day and future ocean conditions

Epilithic algal communities play critical ecological roles on coral reefs, but their response to individual and interactive effects of ocean warming (OW) and ocean acidification (OA) is still largely unknown. We investigated growth, photosynthesis and calcification of early epilithic algal community assemblages exposed for 6 months to four temperature profiles (−1.1, ±0.0, +0.9, +1.6 °C) that were crossed with four carbon dioxide partial pressure (pCO₂) levels (360, 440, 650, 940 µatm), under flow-through conditions and natural light regimes. Additionally, we compared the cover of heavily calcified crustose coralline algae (CCA) and lightly calcified red algae of the genus Peyssonnelia among treatments. Increase in cover of epilithic communities showed optima under moderately elevated temperatures and present pCO₂, while cover strongly decreased under high temperatures and high-pCO₂ conditions, particularly due to decreasing cover of CCA. Similarly, community calcification rates were strongly decreased at high pCO₂ under both measured temperatures. While final cover of CCA decreased under high temperature and pCO₂ (additive negative effects), cover of Peyssonnelia spp. increased at high compared to annual average and moderately elevated temperatures. Thus, cover of Peyssonnelia spp. increased in treatment combinations with less CCA, which was supported by a significant negative correlation between organism groups. The different susceptibility to stressors most likely derived from a different calcification intensity and/or mineral. Notably, growth of the epilithic communities and final cover of CCA were strongly decreased under reduced-pCO₂ conditions compared to the present. Thus, CCA may have acclimatized from past to present-day pCO₂ conditions, and changes in carbonate chemistry, regardless in which direction, negatively affect them. However, if epilithic organisms cannot further acclimatize to OW and OA, the interacting effects of both factors may change epilithic communities in the future, thereby likely leading to reduced reef stability and recovery.

     

Halimeda contribution to organic and inorganic production in a Tahitian reef system

Of the seven species of Halimeda inhabiting a lagoon on Moorea island, three representing 10% of the algal covering and averaging 111 g of dry weight m^-2, have been studied in the course of a year. The biomass, measured bimonthly, stresses a slight seasonal variability in the species life. The main decrease was reported for H. opuntia after a fruiting event, which happened in October. The primary production was assessed, in situ, periodically over a year, by measuring oxygen variations in enclosures. Either expressed on specific-weight basis or in area units, the highest primary productions were recorded for H. opuntia. Productions and biomasses vary simultaneously during the year. The three species produce all together about 6 g C m^-2y^-1. The growth rate of the sand-dwelling H. incrassata f. ovata was followed during the year by staining, in the field, individuals with alizarin Red-S. The average rates measured were 3.3 segments ind^-1d^-1 and 0.17 gdw d^-1m^-2. The contribution of the three species to the carbonate budget of the reef was estimated by total alkalinity measurements during 24-h cycles. H. opuntia had the highest CaCO₃ production. For the three species studied, CaCO₃ production of 1.4 kg m^-2y^-1, which could correspond to a 0.4mm/year accretion of the studied reefal system, was estimated.     

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.

     

The physiological response of marine diatoms to ocean acidification: differential roles of seawater pCO2 and pH

Although increasing the pCO₂ for diatoms will presumably down‐regulate the CO₂‐concentrating mechanism (CCM) to save energy for growth, different species have been reported to respond differently to ocean acidification (OA). To better understand their growth responses to OA, we acclimated the diatoms Thalassiosira pseudonana, Phaeodactylum tricornutum, and Chaetoceros muelleri to ambient (pCO₂ 400 μatm, pH 8.1), carbonated (pCO₂ 800 μatm, pH 8.1), acidified (pCO₂ 400 μatm, pH 7.8), and OA (pCO₂ 800 μatm, pH 7.8) conditions and investigated how seawater pCO₂ and pH affect their CCMs, photosynthesis, and respiration both individually and jointly. In all three diatoms, carbonation down‐regulated the CCMs, while acidification increased both the photosynthetic carbon fixation rate and the fraction of CO₂ as the inorganic carbon source. The positive OA effect on photosynthetic carbon fixation was more pronounced in C. muelleri, which had a relatively lower photosynthetic affinity for CO₂, than in either T. pseudonana or P. tricornutum. In response to OA, T. pseudonana increased respiration for active disposal of H⁺ to maintain its intracellular pH, whereas P. tricornutum and C. muelleri retained their respiration rate but lowered the intracellular pH to maintain the cross‐membrane electrochemical gradient for H⁺ efflux. As the net result of changes in photosynthesis and respiration, growth enhancement to OA of the three diatoms followed the order of C. muelleri > P. tricornutum > T. pseudonana. This study demonstrates that elucidating the separate and joint impacts of increased pCO₂ and decreased pH aids the mechanistic understanding of OA effects on diatoms in the future, acidified oceans.