Coral reefs modify their seawater carbon chemistry – implications for impacts of ocean acidification

Reviews suggest that that the biogeochemical threshold for sustained coral reef growth will be reached during this century due to ocean acidification caused by increased uptake of atmospheric CO₂. Projections of ocean acidification, however, are based on air‐sea fluxes in the open ocean, and not for shallow‐water systems such as coral reefs. Like the open ocean, reef waters are subject to the chemical forcing of increasing atmospheric pCO₂. However, for reefs with long water residence times, we illustrate that benthic carbon fluxes can drive spatial variation in pH, pCO₂ and aragonite saturation state (Ω_a) that can mask the effects of ocean acidification in some downstream habitats. We use a carbon flux model for photosynthesis, respiration, calcification and dissolution coupled with Lagrangian transport to examine how key groups of calcifiers (zooxanthellate corals) and primary producers (macroalgae) on coral reefs contribute to changes in the seawater carbonate system as a function of water residence time. Analyses based on flume data showed that the carbon fluxes of corals and macroalgae drive Ω_ain opposing directions. Areas dominated by corals elevate pCO₂ and reduce Ω_a, thereby compounding ocean acidification effects in downstream habitats, whereas algal beds draw CO₂ down and elevate Ω_a, potentially offsetting ocean acidification impacts at the local scale. Simulations for two CO₂ scenarios (600 and 900 ppm CO₂) suggested that a potential shift from coral to algal abundance under ocean acidification can lead to improved conditions for calcification in downstream habitats, depending on reef size, water residence time and circulation patterns. Although the carbon fluxes of benthic reef communities cannot significantly counter changes in carbon chemistry at the scale of oceans, they provide a significant mechanism of buffering ocean acidification impacts at the scale of habitat to reef.     

Coral uptake of inorganic phosphorus and nitrogen negatively affected by simultaneous changes in temperature and pH

The effects of ocean acidification and elevated seawater temperature on coral calcification and photosynthesis have been extensively investigated over the last two decades, whereas they are still unknown on nutrient uptake, despite their importance for coral energetics. We therefore studied the separate and combined impacts of increases in temperature and pCO₂ on phosphate, ammonium, and nitrate uptake rates by the scleractinian coral S. pistillata. Three experiments were performed, during 10 days i) at three pH_T conditions (8.1, 7.8, and 7.5) and normal temperature (26°C), ii) at three temperature conditions (26°, 29°C, and 33°C) and normal pH_T (8.1), and iii) at three pH_T conditions (8.1, 7.8, and 7.5) and elevated temperature (33°C). After 10 days of incubation, corals had not bleached, as protein, chlorophyll, and zooxanthellae contents were the same in all treatments. However, photosynthetic rates significantly decreased at 33°C, and were further reduced for the pH_T 7.5. The photosynthetic efficiency of PSII was only decreased by elevated temperature. Nutrient uptake rates were not affected by a change in pH alone. Conversely, elevated temperature (33°C) alone induced an increase in phosphate uptake but a severe decrease in nitrate and ammonium uptake rates, even leading to a release of nitrogen into seawater. Combination of high temperature (33°C) and low pH_T (7.5) resulted in a significant decrease in phosphate and nitrate uptake rates compared to control corals (26°C, pH_T = 8.1). These results indicate that both inorganic nitrogen and phosphorus metabolism may be negatively affected by the cumulative effects of ocean warming and acidification.     

Productivity gains do not compensate for reduced calcification under near‐future ocean acidification in the photosynthetic benthic foraminifer species Marginopora vertebralis

Changes in the seawater carbonate chemistry (ocean acidification) from increasing atmospheric carbon dioxide (CO₂) concentrations negatively affect many marine calcifying organisms, but may benefit primary producers under dissolved inorganic carbon (DIC) limitation. To improve predictions of the ecological effects of ocean acidification, the net gains and losses between the processes of photosynthesis and calcification need to be studied jointly on physiological and population levels. We studied productivity, respiration, and abundances of the symbiont‐bearing foraminifer species Marginopora vertebralis on natural CO₂ seeps in Papua New Guinea and conducted additional studies on production and calcification on the Great Barrier Reef (GBR) using artificially enhanced pCO₂. Net oxygen production increased up to 90% with increasing pCO₂; temperature, light, and pH together explaining 61% of the variance in production. Production increased with increasing light and increasing pCO₂ and declined at higher temperatures. Respiration was also significantly elevated (~25%), whereas calcification was reduced (16–39%) at low pH/high pCO₂ compared to present‐day conditions. In the field, M. vertebralis was absent at three CO₂ seep sites at pH_Total levels below ~7.9 (pCO₂ ~700 μatm), but it was found in densities of over 1000 m^?2 at all three control sites. The study showed that endosymbiotic algae in foraminifera benefit from increased DIC availability and may be naturally carbon limited. The observed reduction in calcification may have been caused either by increased energy demands for proton pumping (measured as elevated rates of respiration) or by stronger competition for DIC from the more productive symbionts. The net outcome of these two competing processes is that M. vertebralis cannot maintain populations under pCO₂ exceeding 700 μatm, thus are likely to be extinct in the next century.     

End of the Century pCO2 Levels Do Not Impact Calcification in Mediterranean Cold-Water Corals

Ocean acidification caused by anthropogenic uptake of CO₂ is perceived to be a major threat to calcifying organisms. Cold-water corals were thought to be strongly affected by a decrease in ocean pH due to their abundance in deep and cold waters which, in contrast to tropical coral reef waters, will soon become corrosive to calcium carbonate. Calcification rates of two Mediterranean cold-water coral species, Lophelia pertusa and Madrepora oculata, were measured under variable partial pressure of CO₂ (pCO₂) that ranged between 380 µatm for present-day conditions and 930 µatm for the end of the century. The present study addressed both short- and long-term responses by repeatedly determining calcification rates on the same specimens over a period of 9 months. Besides studying the direct, short-term response to elevated pCO₂ levels, the study aimed to elucidate the potential for acclimation of calcification of cold-water corals to ocean acidification. Net calcification of both species was unaffected by the levels of pCO₂ investigated and revealed no short-term shock and, therefore, no long-term acclimation in calcification to changes in the carbonate chemistry. There was an effect of time during repeated experiments with increasing net calcification rates for both species, however, as this pattern was found in all treatments, there is no indication that acclimation of calcification to ocean acidification occurred. The use of controls (initial and ambient net calcification rates) indicated that this increase was not caused by acclimation in calcification response to higher pCO₂. An extrapolation of these data suggests that calcification of these two cold-water corals will not be affected by the pCO₂ level projected at the end of the century.     

Effects of elevated pCO2 on epilithic and endolithic metabolism of reef carbonates

We investigated effects of elevated partial pressure of CO₂ (pCO₂) on the metabolism of epilithic and endolithic phototrophic communities that colonized experimental coral blocks. Blocks of the massive coral Porites lobata were exposed to colonization by epilithic and endolithic organisms at an oceanic site in Kaneohe Bay (Hawaii) for 6 months, and then were transported to laboratory tanks. A bubbling system was used to maintain two treatments for 3 months, one at ambient pCO₂ (400 ppm) and the second at elevated pCO₂ (750 ppm). Net photosynthetic rates of epilithic communities in the high pCO₂ treatment, dominated by encrusting coralline algae, decreased by 35% while respiration rates remained constant. In contrast, metabolism of endolithic phototrophs, comprised of cyanobacteria and algae, was not significantly affected by the elevated pCO₂ even though endoliths contributed about 63% to block production.     

Interactive Effects of CO2, Temperature,Irradiance, and Nutrient Limitation on the Growth and Physiology of the Marine Diatom Thalassiosira pseudonana (Coscinodiscophyceae)

The marine diatom Thalassiosira pseudonana was grown in continuous culture systems to study the interactive effects of temperature, irradiance, nutrient limitation, and the partial pressure of CO₂ (pCO₂) on its growth and physiological characteristics. The cells were able to grow at all combinations of low and high irradiance (50 and 300 μmol photons · m⁻² · s⁻¹, respectively, of visible light), low and high pCO₂ (400 and 1,000 μatm, respectively), nutrient limitation (nitrate-limited and nutrient-replete conditions), and temperatures of 10–32°C. Under nutrient-replete conditions, there was no adverse effect of high pCO₂ on growth rates at temperatures of 10–25°C. The response of the cells to high pCO₂ was similar at low and high irradiance. At supraoptimal temperatures of 30°C or higher, high pCO₂ depressed growth rates at both low and high irradiance. Under nitrate-limited conditions, cells were grown at 38 ± 2.4% of their nutrient-saturated rates at the same temperature, irradiance, and pCO₂. Dark respiration rates consistently removed a higher percentage of production under nitrate-limited versus nutrient-replete conditions. The percentages of production lost to dark respiration were positively correlated with temperature under nitrate-limited conditions, but there was no analogous correlation under nutrient-replete conditions. The results suggest that warmer temperatures and associated more intense thermal stratification of ocean surface waters could lower net photosynthetic rates if the stratification leads to a reduction in the relative growth rates of marine phytoplankton, and at truly supraoptimal temperatures there would likely be a synergistic interaction between the stresses from temperature and high pCO₂ (lower pH).     

The Response of Antarctic Sea Ice Algae to Changes in pH and CO2

Ocean acidification substantially alters ocean carbon chemistry and hence pH but the effects on sea ice formation and the CO₂ concentration in the enclosed brine channels are unknown. Microbial communities inhabiting sea ice ecosystems currently contribute 10–50% of the annual primary production of polar seas, supporting overwintering zooplankton species, especially Antarctic krill, and seeding spring phytoplankton blooms. Ocean acidification is occurring in all surface waters but the strongest effects will be experienced in polar ecosystems with significant effects on all trophic levels. Brine algae collected from McMurdo Sound (Antarctica) sea ice was incubated in situ under various carbonate chemistry conditions. The carbon chemistry was manipulated with acid, bicarbonate and bases to produce a pCO₂ and pH range from 238 to 6066 µatm and 7.19 to 8.66, respectively. Elevated pCO₂ positively affected the growth rate of the brine algal community, dominated by the unique ice dinoflagellate, Polarella glacialis. Growth rates were significantly reduced when pH dropped below 7.6. However, when the pH was held constant and the pCO₂ increased, growth rates of the brine algae increased by more than 20% and showed no decline at pCO₂ values more than five times current ambient levels. We suggest that projected increases in seawater pCO₂, associated with OA, will not adversely impact brine algal communities.     

Symbiodinium mitigate the combined effects of hypoxia and acidification on a noncalcifying cnidarian

Anthropogenic nutrient inputs enhance microbial respiration within many coastal ecosystems, driving concurrent hypoxia and acidification. During photosynthesis, Symbiodinium spp., the microalgal endosymbionts of cnidarians and other marine phyla, produce O₂ and assimilate CO₂ and thus potentially mitigate the exposure of the host to these stresses. However, such a role for Symbiodinium remains untested for noncalcifying cnidarians. We therefore contrasted the fitness of symbiotic and aposymbiotic polyps of a model host jellyfish (Cassiopea sp.) under reduced O₂ (~2.09 mg/L) and pH (~ 7.63) scenarios in a full‐factorial experiment. Host fitness was characterized as asexual reproduction and their ability to regulate internal pH and Symbiodinium performance characterized by maximum photochemical efficiency, chla content and cell density. Acidification alone resulted in 58% more asexual reproduction of symbiotic polyps than aposymbiotic polyps (and enhanced Symbiodinium cell density) suggesting Cassiopea sp. fitness was enhanced by CO₂‐stimulated Symbiodinium photosynthetic activity. Indeed, greater CO₂ drawdown (elevated pH) was observed within host tissues of symbiotic polyps under acidification regardless of O₂ conditions. Hypoxia alone produced 22% fewer polyps than ambient conditions regardless of acidification and symbiont status, suggesting Symbiodinium photosynthetic activity did not mitigate its effects. Combined hypoxia and acidification, however, produced similar numbers of symbiotic polyps compared with aposymbiotic kept under ambient conditions, demonstrating that the presence of Symbiodinium was key for mitigating the combined effects of hypoxia and acidification on asexual reproduction. We hypothesize that this mitigation occurred because of reduced photorespiration under elevated CO₂ conditions where increased net O₂ production ameliorates oxygen debt. We show that Symbiodinium play an important role in facilitating enhanced fitness of Cassiopea sp. polyps, and perhaps also other noncalcifying cnidarian hosts, to the ubiquitous effects of ocean acidification. Importantly we highlight that symbiotic, noncalcifying cnidarians may be particularly advantaged in productive coastal waters that are subject to simultaneous hypoxia and acidification.     

Ocean acidification and warming scenarios increase microbioerosion of coral skeletons

Biological mediation of carbonate dissolution represents a fundamental component of the destructive forces acting on coral reef ecosystems. Whereas ocean acidification can increase dissolution of carbonate substrates, the combined impact of ocean acidification and warming on the microbioerosion of coral skeletons remains unknown. Here, we exposed skeletons of the reef‐building corals, Porites cylindrica and Isopora cuneata, to present‐day (Control: 400 μatm – 24 °C) and future pCO₂–temperature scenarios projected for the end of the century (Medium: +230 μatm – +2 °C; High: +610 μatm – +4 °C). Skeletons were also subjected to permanent darkness with initial sodium hypochlorite incubation, and natural light without sodium hypochlorite incubation to isolate the environmental effect of acidic seawater (i.e., Ω_aragonite <1) from the biological effect of photosynthetic microborers. Our results indicated that skeletal dissolution is predominantly driven by photosynthetic microborers, as samples held in the dark did not decalcify. In contrast, dissolution of skeletons exposed to light increased under elevated pCO₂–temperature scenarios, with P. cylindrica experiencing higher dissolution rates per month (89%) than I. cuneata (46%) in the high treatment relative to control. The effects of future pCO₂–temperature scenarios on the structure of endolithic communities were only identified in P. cylindrica and were mostly associated with a higher abundance of the green algae Ostreobium spp. Enhanced skeletal dissolution was also associated with increased endolithic biomass and respiration under elevated pCO₂–temperature scenarios. Our results suggest that future projections of ocean acidification and warming will lead to increased rates of microbioerosion. However, the magnitude of bioerosion responses may depend on the structural properties of coral skeletons, with a range of implications for reef carbonate losses under warmer and more acidic oceans.     

Combined effects of different CO2 levels and N sources on the diazotrophic cyanobacterium Trichodesmium

To predict effects of climate change and possible feedbacks, it is crucial to understand the mechanisms behind CO₂ responses of biogeochemically relevant phytoplankton species. Previous experiments on the abundant N₂ fixers Trichodesmium demonstrated strong CO₂ responses, which were attributed to an energy reallocation between its carbon (C) and nitrogen (N) acquisition. Pursuing this hypothesis, we manipulated the cellular energy budget by growing Trichodesmium erythraeum IMS101 under different CO₂ partial pressure (pCO₂) levels (180, 380, 980 and 1400 µatm) and N sources (N₂ and NO₃^?). Subsequently, biomass production and the main energy‐generating processes (photosynthesis and respiration) and energy‐consuming processes (N₂ fixation and C acquisition) were measured. While oxygen fluxes and chlorophyll fluorescence indicated that energy generation and its diurnal cycle was neither affected by pCO₂ nor N source, cells differed in production rates and composition. Elevated pCO₂ increased N₂ fixation and organic C and N contents. The degree of stimulation was higher for nitrogenase activity than for cell contents, indicating a pCO₂ effect on the transfer efficiency from N₂ to biomass. pCO₂‐dependent changes in the diurnal cycle of N₂ fixation correlated well with C affinities, confirming the interactions between N and C acquisition. Regarding effects of the N source, production rates were enhanced in NO₃^? grown cells, which we attribute to the higher N retention and lower ATP demand compared with N₂ fixation. pCO₂ effects on C affinity were less pronounced in NO₃^? users than N₂ fixers. Our study illustrates the necessity to understand energy budgets and fluxes under different environmental conditions for explaining indirect effects of rising pCO₂.     

Ocean acidification does not affect the physiology of the tropical coral Acropora digitifera during a 5-week experiment

The increase in atmospheric CO₂ concentration, which has resulted from the burning of fossil fuels, is being absorbed by the oceans and is causing ocean acidification. Ocean acidification involves the decrease of both the pH and the calcium carbonate saturation state. Ocean acidification is predicted to impact the physiology of marine organisms and reduce the calcification rates of corals. In the present study, we measured the rates of calcification, respiration, photosynthesis, and zooxanthellae density of the tropical coral Acropora digitifera under near-natural summertime temperature and sunlight for a 5-week period. We found that these key physiological parameters were not affected by both mid-CO₂ (pCO₂ = 744 ± 38, pH = 7.97 ± 0.02, Ω_arag = 2.6 ± 0.1) and high-CO₂ conditions (pCO₂ = 2,142 ± 205, pH = 7.56 ± 0.04, Ω_arag = 1.1 ± 0.2) throughout the 35 days experimental period. Additionally, there was no significant correlation between calcification rate and seawater aragonite saturation (Ω_arag). These results suggest that the impacts of ocean acidification on corals physiology may be more complex than have been previously proposed.     

Responses of Two Scleractinian Corals to Cobalt Pollution and Ocean Acidification

The effects of ocean acidification alone or in combination with warming on coral metabolism have been extensively investigated, whereas none of these studies consider that most coral reefs near shore are already impacted by other natural anthropogenic inputs such as metal pollution. It is likely that projected ocean acidification levels will aggravate coral reef health. We first investigated how ocean acidification interacts with one near shore locally abundant metal on the physiology of two major reef-building corals: Stylophora pistillata and Acropora muricata. Two pH levels (pH_T 8.02; pCO₂ 366 μatm and pH_T 7.75; pCO₂ 1140 μatm) and two cobalt concentrations (natural, 0.03 μg L^-1 and polluted, 0.2 μg L^-1) were tested during five weeks in aquaria. We found that, for both species, cobalt input decreased significantly their growth rates by 28% while it stimulated their photosystem II, with higher values of rETR_max (relative Electron Transport Rate). Elevated pCO₂ levels acted differently on the coral rETR_max values and did not affect their growth rates. No consistent interaction was found between pCO₂ levels and cobalt concentrations. We also measured in situ the effect of higher cobalt concentrations (1.06 ± 0.16 μg L^-1) on A. muricata using benthic chamber experiments. At this elevated concentration, cobalt decreased simultaneously coral growth and photosynthetic rates, indicating that the toxic threshold for this pollutant has been reached for both host cells and zooxanthellae. Our results from both aquaria and in situ experiments, suggest that these coral species are not particularly sensitive to high pCO₂ conditions but they are to ecologically relevant cobalt concentrations. Our study reveals that some reefs may be yet subjected to deleterious pollution levels, and even if no interaction between pCO₂ levels and cobalt concentration has been found, it is likely that coral metabolism will be weakened if they are subjected to additional threats such as temperature increase, other heavy metals, and eutrophication.     

The combined effects of temperature and CO2 lead to altered gene expression in Acropora aspera

This study explored the interactive effects of near-term CO₂ increases (40–90 ppm above current ambient) during a simulated bleaching event (34 °C for 5 d) of Acropora aspera by linking physiology to expression patterns of genes involved in carbon metabolism. Symbiodinium photosynthetic efficiency (F _v /F _m ) was significantly depressed by the bleaching event, while elevated pressure of CO₂ (pCO₂) slightly mitigated the effects of increased temperature on F _v /F _m during the final 4 d of the recovery period, however, did not affect the loss of symbionts. Elevated pCO₂ alone had no effect on F _v /F _m or symbiont density. Expression of targeted Symbiodinium genes involved in carbon metabolism and heat stress response was not significantly altered by either increased temperature and/or CO₂. Of the selected host genes, two carbonic anhydrase isoforms (coCA2 and coCA3) exhibited the largest changes, most notably in crossed bleaching and elevated pCO₂ treatments. CA2 was significantly down-regulated on day 14 in all treatments, with the greatest decrease in the crossed treatment (relative expression compared to control = 0.16; p < 0.05); CA3 showed a similar trend, with expression levels 0.20-fold of controls on day 14 (p < 0.05) in the elevated temperature/pCO₂ treatment. The synergistic effects of ocean acidification and bleaching were evident during this study and demonstrate that increased pCO₂ in surface waters will impact corals much sooner than many studies utilising end-of-century pCO₂ concentrations would indicate.     

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⁻.     

Interactive effects of temperature and pCO2 on sponges: from the cradle to the grave

As atmospheric CO₂ concentrations rise, associated ocean warming (OW) and ocean acidification (OA) are predicted to cause declines in reef‐building corals globally, shifting reefs from coral‐dominated systems to those dominated by less sensitive species. Sponges are important structural and functional components of coral reef ecosystems, but despite increasing field‐based evidence that sponges may be ‘winners’ in response to environmental degradation, our understanding of how they respond to the combined effects of OW and OA is limited. To determine the tolerance of adult sponges to climate change, four abundant Great Barrier Reef species were experimentally exposed to OW and OA levels predicted for 2100, under two CO₂ Representative Concentration Pathways (RCPs). The impact of OW and OA on early life‐history stages was also assessed for one of these species to provide a more holistic view of species impacts. All species were generally unaffected by conditions predicted under RCP6.0, although environmental conditions projected under RCP8.5 caused significant adverse effects: with elevated temperature decreasing the survival of all species, increasing levels of tissue necrosis and bleaching, elevating respiration rates and decreasing photosynthetic rates. OA alone had little adverse effect, even under RCP8.5 concentrations. Importantly, the interactive effect of OW and OA varied between species with different nutritional modes, with elevated pCO₂ exacerbating temperature stress in heterotrophic species but mitigating temperature stress in phototrophic species. This antagonistic interaction was reflected by reduced mortality, necrosis and bleaching of phototrophic species in the highest OW/OA treatment. Survival and settlement success of Carteriospongia foliascens larvae were unaffected by experimental treatments, and juvenile sponges exhibited greater tolerance to OW than their adult counterparts. With elevated pCO₂ providing phototrophic species with protection from elevated temperature, across different life stages, climate change may ultimately drive a shift in the composition of sponge assemblages towards a dominance of phototrophic species.     

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.     

Seasonal patterns of photosynthesis in Douglas fir seedlings during the third and fourth year of exposure to elevated CO2 and temperature

The interactive effects of elevated atmospheric CO₂ and temperature on seasonal patterns of photosynthesis in Douglas fir (Psuedotsuga menziesii (Mirb.) Franco) seedlings were examined. Seedlings were grown in sunlit chambers controlled to track either ambient (~400 p.p.m.) CO₂ or ambient +200 p.p.m. CO₂, and either ambient temperature or ambient +4 °C. Light‐saturated net photosynthetic rates were measured approximately monthly over a 21 month period. Elevated CO₂ increased net photosynthetic rates by an average of 21% across temperature treatments during both the 1996 hydrologic year, the third year of exposure, and the 1997 hydrologic year. Elevated mean annual temperature increased net photosynthetic rates by an average of 33% across CO₂ treatments during both years. Seasonal temperature changes also affected net photosynthetic rates. Across treatments, net photosynthetic rates were highest in the spring and autumn, and lowest in July, August and December–January. Seasonal increases in temperature were not correlated with increases in the relative photosynthetic response to elevated CO₂. Seasonal shifts in the photosynthetic temperature optimum reduced temperature effects on the relative response to elevated CO₂. These results suggest that the effects of elevated CO₂ on net photosynthetic rates in Douglas fir are largely independent of temperature.     

High CO2 decreases the long‐term resilience of the free‐living coralline algae Phymatolithon lusitanicum

Mäerl/rhodolith beds are protected habitats that may be affected by ocean acidification (OA), but it is still unclear how the availability of CO₂ will affect the metabolism of these organisms. Some of the inconsistencies found among OA experimental studies may be related to experimental exposure time and synergetic effects with other stressors. Here, we investigated the long‐term (up to 20 months) effects of OA on the production and calcification of the most common mäerl species of southern Portugal, Phymatolithon lusitanicum. Both the photosynthetic and calcification rates increased with CO₂ after the first 11 months of the experiment, whereas respiration slightly decreased with CO₂. After 20 months, the pattern was reversed. Acidified algae showed lower photosynthetic and calcification rates, as well as lower accumulated growth than control algae, suggesting that a metabolic threshold was exceeded. Our results indicate that long‐term exposure to high CO₂ will decrease the resilience of Phymatolithon lusitanicum. Our results also show that shallow communities of these rhodoliths may be particularly at risk, while deeper rhodolith beds may become ocean acidification refuges for this biological community.     

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.