Seawater carbonate chemistry, calcification and Zn uptake during experiments with coral Stylophora pistillata, 2011

Many studies have investigated the effect of an increase in pCO2 on coral calcification and photosynthesis but the physiological consequences are still relatively speculative. We investigated the effects of ocean acidification on zinc incorporation and gross calcification in the scleractinian coral Stylophora pistillata. Zn is an essential element for health and growth of corals. Colonies were maintained at normal pHT (8.1) and at two low-pH conditions (7.8 and 7.5) during 5 weeks. Corals were exposed to 65Zn dissolved in seawater to assess uptake rates of this element. After 5 weeks, 65Zn activity measured in the whole coral and in the two compartments: tissue and skeleton, differed significantly between pH conditions with concentration factors higher at pHT 8.1, compared to lower pH. Zn is therefore taken less efficiently by corals at reduced pH. Their gross calcification, as measured by 45Ca incorporation, photosynthesis and photosynthetic efficiency did not change with pH even at the lowest level.

Seawater carbonate chemistry and calcification rate in the Bermuda reef community, 2010

Despite the potential impact of ocean acidification on ecosystems such as coral reefs, surprisingly, there is very limited field data on the relationships between calcification and seawater carbonate chemistry. In this study, contemporaneous in situ datasets of seawater carbonate chemistry and calcification rates from the high-latitude coral reef of Bermuda over annual timescales provide a framework for investigating the present and future potential impact of rising carbon dioxide (CO2) levels and ocean acidification on coral reef ecosystems in their natural environment. A strong correlation was found between the in situ rates of calcification for the major framework building coral species Diploria labyrinthiformis and the seasonal variability of [CO32-] and aragonite saturation state omega aragonite, rather than other environmental factors such as light and temperature. These field observations provide sufficient data to hypothesize that there is a seasonal "Carbonate Chemistry Coral Reef Ecosystem Feedback" (CREF hypothesis) between the primary components of the reef ecosystem (i.e., scleractinian hard corals and macroalgae) and seawater carbonate chemistry. In early summer, strong net autotrophy from benthic components of the reef system enhance [CO32-] and omega aragonite conditions, and rates of coral calcification due to the photosynthetic uptake of CO2. In late summer, rates of coral calcification are suppressed by release of CO2 from reef metabolism during a period of strong net heterotrophy. It is likely that this seasonal CREF mechanism is present in other tropical reefs although attenuated compared to high-latitude reefs such as Bermuda. Due to lower annual mean surface seawater [CO32-] and omega aragonite in Bermuda compared to tropical regions, we anticipate that Bermuda corals will experience seasonal periods of zero net calcification within the next decade at [CO32-] and omega aragonite thresholds of ~184 micro moles kg-1 and 2.65. However, net autotrophy of the reef during winter and spring (as part of the CREF hypothesis) may delay the onset of zero NEC or decalcification going forward by enhancing [CO32-] and omega aragonite. The Bermuda coral reef is one of the first responders to the negative impacts of ocean acidification, and we estimate that calcification rates for D. labyrinthiformis have declined by >50% compared to pre-industrial times.

Seawater carbonate chemistry and calcification during experiments with coral communities, 2000

Previous studies have demonstrated that coral and algal calcification is tightly regulated by the calcium carbonate saturation state of seawater. This parameter is likely to decrease in response to the increase of dissolved CO2 resulting from the global increase of the partial pressure of atmospheric CO2. We have investigated the response of a coral reef community dominated by scleractinian corals, but also including other calcifying organisms such as calcareous algae, crustaceans, gastropods and echinoderms, and kept in an open-top mesocosm. Seawater pCO2 was modified by manipulating the pCO2 of air used to bubble the mesocosm. The aragonite saturation state (omega arag) of the seawater in the mesocosm varied between 1.3 and 5.4. Community calcification decreased as a function of increasing pCO2 and decreasing omega arag. This result is in agreement with previous data collected on scleractinian corals, coralline algae and in a reef mesocosm, even though some of these studies did not manipulate CO2 directly. Our data suggest that the rate of calcification during the last glacial maximum might have been 114% of the preindustrial rate. Moreover, using the average emission scenario (IS92a) of the Intergovernmental Panel on Climate Change, we predict that the calcification rate of scleractinian-dominated communities may decrease by 21% between the pre-industrial period (year 1880) and the time at which pCO2 will double (year 2065).

Seawater carbonate chemistry and growth, reproduction of calcifying and non-calcifying epibionts of a brown macroalga in a laboratory experiment

Seaweeds are key species of the Baltic Sea benthic ecosystems. They are the substratum of numerous fouling epibionts like bryozoans and tubeworms. Several of these epibionts bear calcified structures and could be impacted by the high pCO2 events of the late summer upwellings in the Baltic nearshores. Those events are expected to increase in strength and duration with global change and ocean acidification. If calcifying epibionts are impacted by transient acidification as driven by upwelling events, their increasing prevalence could cause a shift of the fouling communities toward fleshy species. The aim of the present study was to test the sensitivity of selected seaweed macrofoulers to transient elevation of pCO2 in their natural microenvironment, i.e. the boundary layer covering the thallus surface of brown seaweeds. Fragments of the macroalga Fucus serratus bearing an epibiotic community composed of the calcifiers Spirorbis spirorbis (Annelida) and Electra pilosa (Bryozoa) and the non-calcifier Alcyonidium hirsutum (Bryozoa) were maintained for 30 days under three pCO2 conditions: natural 460±59 µatm, present-day upwelling1193±166 µatm and future upwelling 3150±446 µatm. Only the highest pCO2 caused a significant reduction of growth rates and settlement of S. spirorbis individuals. Additionally, S. spirorbis settled juveniles exhibited enhanced calcification of 40% during daylight hours compared to dark hours, possibly reflecting a day-night alternation of an acidification-modulating effect by algal photosynthesis as opposed to an acidification-enhancing effect of algal respiration. E. pilosa colonies showed significantly increased growth rates at intermediate pCO2 (1193 µatm) but no response to higher pCO2. No effect of acidification on A. hirsutum colonies growth rates was observed. The results suggest a remarkable resistance of the algal macro-epibionts to levels of acidification occurring at present day upwellings in the Baltic. Only extreme future upwelling conditions impacted the tubeworm S. spirorbis, but not the bryozoans.

Seawater carbonate chemistry, fertilization and early development of the coral eating crown-of-thorns starfish, Acanthaster planci in a laboratory experiment

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.

Incorporation of Mg and Sr in calcite of cultured benthic foraminifera (Heterostegina depressa and Ammonia tepida) and seawater carbonate chemistry, 2010

We investigated the effect of the calcium concentration in seawater and thereby the calcite saturation state (omega) on the magnesium and strontium incorporation into benthic foraminiferal calcite under laboratory conditions. For this purpose individuals of the shallow-water species Heterostegina depressa (precipitating high-Mg calcite, symbiont-bearing) and Ammonia tepida (low-Mg calcite, symbiont-barren) were cultured in media under a range of [Ca2+], but similar Mg/Ca ratios. Trace element/Ca ratios of newly formed calcite were analysed with Laser Ablation Inductively Coupled Plasma Mass Spectrometry (LA-ICP-MS) and normalized to the seawater elemental composition using the equation DTE=(TE/Cacalcite)/(TE/Caseawater). The culturing study shows that DMg of A. tepida significantly decreases with increasing omega at a gradient of -4.3x10-5 per omega unit. The DSr value of A. tepida does not change with omega, suggesting that fossil Sr/Ca in this species may be a potential tool to reconstruct past variations in seawater Sr/Ca. Conversely, DMg of H. depressa shows only a minor decrease with increasing omega, while DSr increases considerably with omega at a gradient of 0.009 per omega unit. The different responses to seawater chemistry of the two species may be explained by a difference in the calcification pathway that is, at the same time, responsible for the variation in the total Mg incorporation between the two species. Since the Mg/Ca ratio in H. depressa is 50-100 times higher than that of A. tepida, it is suggested that the latter exhibits a mechanism that decreases the Mg/Ca ratio of the calcification fluid, while the high-Mg calcite forming species may not have this physiological tool. If the dependency of Mg incorporation on seawater [Ca2+] is also valid for deep-sea benthic foraminifera typically used for paleostudies, the higher Ca concentrations in the past may potentially bias temperature reconstructions to a considerable degree. For instance, 25 Myr ago Mg/Ca ratios in A. tepida would have been 0.2 mmol/mol lower than today, due to the 1.5 times higher [Ca2+] of seawater, which in turn would lead to a temperature underestimation of more than 2 °C.

Seawater carbonate chemistry, nutrients, chlorophyll and diatom Proboscia alata during experiments, 2012

The combined effects of different light and aqueous CO2 conditions were assessed for the Southern Ocean diatom Proboscia alata(Brightwell) Sundström in laboratory experiments. Selected culture conditions (light and CO2(aq)) were representative for the natural ranges in the modern Southern Ocean. Light conditions were 40 (low) and 240 (high) µmol photons/m**2/s. The three CO2(aq) conditions ranged from 8 to 34 µmol/kg CO2(aq) (equivalent to a pCO2 from 137 to 598 µatm, respectively). Clear morphological changes were induced by these different CO2(aq) conditions. Cells in low [CO2(aq)] formed spirals, while many cells in high [CO2(aq)] disintegrated. Cell size and volume were significantly affected by the different CO2(aq) concentrations. Increasing CO2(aq) concentrations led to an increase in particulate organic carbon concentrations per cell in the high light cultures, with exactly the opposite happening in the low light cultures. However, other parameters measured were not influenced by the range of CO2(aq) treatments. This included growth rates, chlorophyll aconcentration and photosynthetic yield (FV/FM). Different light treatments had a large effect on nutrient uptake. High light conditions caused an increased nutrient uptake rate compared to cells grown in low light conditions. Light and CO2 conditions co-determined in various ways the response of P. alata to changing environmental conditions. Overall P. alata appeared to be well adapted to the natural variability in light availability and CO2(aq) concentration of the modern Southern Ocean. Nevertheless, our results showed that P. alata is susceptible to future changes in inorganic carbon concentrations in the Southern Ocean.

Seawater carbonate chemistry and pteropod Limacina helicina antarctica shell dissolution during experiments, 2012

Anthropogenic ocean acidification is likely to have negative effects on marine calcifying organisms, such as shelled pteropods, by promoting dissolution of aragonite shells. Study of shell dissolution requires an accurate and sensitive method for assessing shell damage. Shell dissolution was induced through incubations in CO2 enriched seawater for between 4 and 14 days. We describe a procedure that allows the level of dissolution to be assessed and classified into three main types: Type I with partial dissolution of the prismatic layer; Type II with exposure of underlying crossed-lamellar layer, and Type III, where crossed-lamellar layer shows signs of dissolution. Levels of dissolution showed a good correspondence to the incubation conditions, with the most severe damage found in specimens held for 14 d in undersaturated condition (Ohm ~ 0.8). This methodology enables the response of small pelagic calcifiers to acidified conditions to be detected at an early stage, thus making pteropods a valuable bioindicator of future ocean acidification.

Seawater carbonate chemistry and 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.