Seawater carbonate chemistry and biological processes of Emiliania huxleyi (strains RCC1256 and NZEH) during experiments, 2011

With respect to their sensitivity to ocean acidification, calcifiers such as the coccolithophore Emiliania huxleyi have received special attention, as the process of calcification seems to be particularly sensitive to changes in the marine carbonate system. For E. huxleyi, apparently conflicting results regarding its sensitivity to ocean acidification have been published (Iglesias-Rodriguez et al., 2008a; Riebesell et al., 2000). As possible causes for discrepancies, intra-specific variability and different effects of CO2 manipulation methods, i.e. the manipulation of total alkalinity (TA) or total dissolved inorganic carbon (DIC), have been discussed. While Langer et al. (2009) demonstrate a high degree of intra-specific variability between strains of E. huxleyi, the question whether different CO2 manipulation methods influence the cellular responses has not been resolved yet. In this study, closed TA as well as open and closed DIC manipulation methods were compared with respect to E. huxleyi's CO2-dependence in growth rate, POC- and PIC-production. The differences in the carbonate chemistry between TA and DIC manipulations were shown not to cause any differences in response patterns, while the latter differed between open and closed DIC manipulation. The two strains investigated showed different sensitivities to acidification of seawater, RCC1256 being more negatively affected in growth rates and PIC production than NZEH.

Seawater carbonate chemistry and reproduction of a coral reef fish in a laboratory experiment

Ocean acidification is predicted to negatively impact the reproduction of many marine species, either by reducing fertilization success or diverting energy from reproductive effort. While recent studies have demonstrated how ocean acidification will affect larval and juvenile fishes, little is known about how increasing partial pressure of carbon dioxide (pCO2) and decreasing pH might affect reproduction in adult fishes. We investigated the effects of near-future levels of pCO2 on the reproductive performance of the cinnamon anemonefish, Amphiprion melanopus, from the Great Barrier Reef, Australia. Breeding pairs were held under three CO2 treatments [Current-day Control (430 µatm), Moderate (584 µatm) and High (1032 µatm)] for a 9-month period that included the summer breeding season. Unexpectedly, increased CO2 dramatically stimulated breeding activity in this species of fish. Over twice as many pairs bred in the Moderate (67% of pairs) and High (55%) compared to the Control (27%) CO2 treatment. Pairs in the High CO2 group produced double the number of clutches per pair and 67% more eggs per clutch compared to the Moderate and Control groups. As a result, reproductive output in the High group was 82% higher than that in the Control group and 50% higher than that in the Moderate group. Despite the increase in reproductive activity, there was no difference in adult body condition among the three treatment groups. There was no significant difference in hatchling length between the treatment groups, but larvae from the High CO2 group had smaller yolks than Controls. This study provides the first evidence of the potential effects of ocean acidification on key reproductive attributes of marine fishes and, contrary to expectations, demonstrates an initially stimulatory (hormetic) effect in response to increased pCO2. However, any long-term consequences of increased reproductive effort on individuals or populations remain to be determined.

Experiment: Acidified seawater impacts sea urchin larvae pH regulatory systems relevant for calcification

Calcifying echinoid larvae respond to changes in seawater carbonate chemistry with reduced growth and developmental delay. To date, no information exists on how ocean acidification acts on pH homeostasis in echinoderm larvae. Understanding acid-base regulatory capacities is important because intracellular formation and maintenance of the calcium carbonate skeleton is dependent on pH homeostasis. Using H(+)-selective microelectrodes and the pH-sensitive fluorescent dye BCECF, we conducted in vivo measurements of extracellular and intracellular pH (pH(e) and pH(i)) in echinoderm larvae. We exposed pluteus larvae to a range of seawater CO(2) conditions and demonstrated that the extracellular compartment surrounding the calcifying primary mesenchyme cells (PMCs) conforms to the surrounding seawater with respect to pH during exposure to elevated seawater pCO(2). Using FITC dextran conjugates, we demonstrate that sea urchin larvae have a leaky integument. PMCs and spicules are therefore directly exposed to strong changes in pH(e) whenever seawater pH changes. However, measurements of pH(i) demonstrated that PMCs are able to fully compensate an induced intracellular acidosis. This was highly dependent on Na(+) and HCO(3)(-), suggesting a bicarbonate buffer mechanism involving secondary active Na(+)-dependent membrane transport proteins. We suggest that, under ocean acidification, maintained pH(i) enables calcification to proceed despite decreased pH(e). However, this probably causes enhanced costs. Increased costs for calcification or cellular homeostasis can be one of the main factors leading to modifications in energy partitioning, which then impacts growth and, ultimately, results in increased mortality of echinoid larvae during the pelagic life stage.

Seawater carbonate chemistry and calcification at Anse des Cuivres, NW Mediterranean, 2004

The community metabolism of a shallow infralittoral ecosystem dominated by the calcareous macroalgae Corallina elongata was investigated in Marseilles (NW Mediterranean), by monitoring hourly changes of seawater pH and total alkalinity over 6 d in February 2000. Fair weather conditions prevailed over the study period as indicated by oceanographic (temperature, salinity, and current velocity and direction) and meteorological variables, which validated the standing water hypothesis. This temperate ecosystem exhibited high community gross primary production (GPP = 519 ± 106 mmol C m-2 d-1, n = 6) and also supported high rates of community respiration (R). As a result, the system was slightly autotrophic (net community production, NCP = 20 mmol C m-2 d-1), with a GPP/R ratio of 1.06. NCP exhibited circadian variations with 2- to 3-fold changes in community respiration, both in the light and in the dark. Rates of net community calcification also exhibited circadian variations, with positive rates (up to 24 mmol CaCO3 m-2 h-1) for irradiance values >300 W m-2 (about 1380 µmol photon m-2 s-1). Below this irradiance threshold, net community dissolution prevailed. Daily net calcification (G) was on average 8 mmol CaCO3 m-2 d-1. CO2 fluxes generated by primary production, respiration, and calcification suggest that the study site was a potential atmospheric CO2 sink of 15 mmol CO2 m-2 d-1 at the time of measurement.

Seawater carbonate chemistry and growth rate during experiments with coral Acropora cervicornis, 2005

The growth rate of Acropora cervicornis branch tips maintained in the laboratory was measured before, during, and after exposure to elevated nitrate (5 and 10 µM NO3-), phosphate (2 and 4 µM P-PO43) and/or pCO2 (CO2 ~700 to 800 µatm). The effect of increased pCO2 was greater than that of nutrient enrichment alone. High concentrations of nitrate or phosphate resulted in significant decreases in growth rate, in both the presence and absence of increased pCO2. The effect of nitrate and phosphate enrichment combined was additive or antagonistic relative to nutrient concentration and pCO2 level. Growth rate recovery was greater after exposure to increased nutrients or CO2 compared to increased nutrients and CO2. If these results accurately predict coral response in the natural environment, it is reasonable to speculate that the survival and reef-building potential of this species will be significantly negatively impacted by continued coastal nutrification and projected pCO2 increases.

Surface seawater carbonate chemistry, nutrients and phytoplankton community composition on a transect between North Sea and Arctic Ocean, 2008

This data was collected during the 'ICE CHASER' cruise from the southern North Sea to the Arctic (Svalbard) in July-Aug 2008. This data consists of coccolithophore abundance, calcification and primary production rates, carbonate chemistry parameters and ancillary data of macronutrients, chlorophyll-a, average mixed layer irradiance, daily irradiance above the sea surface, euphotic and mixed layer depth, temperature and salinity.

Seawater carbonate chemistry, physiology and skeletal mineralogy of coralline alga Corallina elongata in a laboratory experiment

Marine organisms inhabiting environments where pCO2/pH varies naturally are suggested to be relatively resilient to future ocean acidification. To test this hypothesis, the effect of elevated pCO2 was investigated in the articulated coralline red alga Corallina elongata from an intertidal rock pool on the north coast of Brittany (France), where pCO2 naturally varied daily between 70 and 1000 µatm. Metabolism was measured on algae in the laboratory after they had been grown for 3 weeks at pCO2 concentrations of 380, 550, 750 and 1000 µatm. Net and gross primary production, respiration and calcification rates were assessed by measurements of oxygen and total alkalinity fluxes using incubation chambers in the light and dark. Calcite mol % Mg/Ca (mMg/Ca) was analysed in the tips, branches and basal parts of the fronds, as well as in new skeletal structures produced by the algae in the different pCO2 treatments. Respiration, gross primary production and calcification in light and dark were not significantly affected by increased pCO2. Algae grown under elevated pCO2 (550, 750 and 1000 µatm) formed fewer new structures and produced calcite with a lower mMg/Ca ratio relative to those grown under 380 µatm. This study supports the assumption that C. elongata from a tidal pool, where pCO2 fluctuates over diel and seasonal cycles, is relatively robust to elevated pCO2 compared to other recently investigated coralline algae.

Seawater carbonate chemistry and 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., Omega 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.

Seawater carbonate chemistry and calcification during tropical reef studies at Younge reef (Great Barrier Reef), 1996

Community metabolism was investigated using a Lagrangian flow respirometry technique on 2 reef flats at Moorea (French Polynesia) during austral winter and Yonge Reef (Great Barrier Reef) during austral summer. The data were used to estimate related air-sea CO2 disequilibrium. A sine function did not satisfactorily model the diel light curves and overestimated the metabolic parameters. The ranges of community gross primary production and respiration (Pg and R; 9 to 15 g C m-2 d-1) were within the range previously reported for reef flats, and community net calcification (G; 19 to 25 g CaCO3 m-2 d-1) was higher than the 'standard' range. The molar ratio of organic to inorganic carbon uptake was 6:1 for both sites. The reef flat at Moorea displayed a higher rate of organic production and a lower rate of calcification compared to previous measurements carried out during austral summer. The approximate uncertainty of the daily metabolic parameters was estimated using a procedure based on a Monte Carlo simulation. The standard errors of Pg,R and Pg/R expressed as a percentage of the mean are lower than 3% but are comparatively larger for E, the excess production (6 to 78%). The daily air-sea CO2 flux (FCO2) was positive throughout the field experiments, indicating that the reef flats at Moorea and Yonge Reef released CO2 to the atmosphere at the time of measurement. FCO2 decreased as a function of increasing daily irradiance.

Planktic foraminifera isotopes, temperature and d18O of seawater at the central Walvis Ridge for Termination II (MIS 6 to 5), sediment core 64PE-174P13

Salty and warm Indian Ocean waters enter the South Atlantic via the Agulhas leakage, south of Africa. Model simulations and proxy evidence of Agulhas leakage strengthening during glacial terminations led to the hypothesis that it was an important modulator of the Atlantic Ocean circulation. Yet, the fate of the leakage salinity and temperature anomalies remains undocumented beyond the southern tip of Africa. Downstream of the leakage, new paleoceanographic evidence from the central Walvis Ridge (southeast Atlantic) shows that salinity increased at the thermocline, and less so at the surface, during glacial termination II. Thermocline salinity change coincided with higher frequency of Agulhas rings passage at the core location and with salinity maxima in the Agulhas leakage area, suggesting that leakage waters were incorporated in the Atlantic circulation through the thermocline. Hydrographic changes at the Walvis Ridge and in the leakage area display a distinct two-step structure, with a reversal at ca. 134 ka. This matched a wet interlude within the East Asia weak monsoon interval of termination II, and a short-lived North Atlantic warming. Such concurrence points to a Bølling-Allerød-like recovery of the Atlantic circulation amidst termination II, with a northward shift of the Intertropical Convergence Zone and Southern Hemisphere westerlies, and attendant curtailment of the interocean connection south of Africa.

Continuous measurement of carbon dioxid partial pressure in seawater in the region of the Iceland-Faroe Ridge (Table 1)

During the 14th expedition of the research vessel "Meteor" from the 2nd of July to the 7th of August 1968 continously recording instruments for measuring the CO2 partial pressure of seawater and atmospheric CO2 were developped by the Meteorological Institute, University of Frankfurt/M. During the Faroer expedition instrumental constants, such as relative and absolute accuracy, inertia and solvent power were tested. The performance of discontinous analyses of water samples was adopted to shipboard conditiones and correction factors depending on water volume, depth of sampling and water temperature were measured. After having computed average values of the continous records (atmosp. CO2 content, CO2 partial pressure, water temperature) geographical distribution, diurnal variation and dependence of diurnal averages were tested. At four different locations CO2 partial pressure was measured in various depths. During the voyage from the Faroer islands to Helgoland the measured concentrations of atmospheric CO2 content and CO2 partial pressure were tested with respect to a correlation of the geographical latitude.

The morphological response of Emiliania huxleyi to seawater carbonate chemistry changes: an inter-strain comparison

Four strains of the coccolithophore Emiliania huxleyi (RCC1212, RCC1216, RCC1238, RCC1256) were grown in dilute batch culture at four CO2 levels ranging from ~200 µatm to ~1200 µatm. Coccolith morphology was analyzed based on scanning electron micrographs. Three of the four strains did not exhibit a change in morphology over the CO2 range tested. One strain (RCC1256) displayed an increase in the percentage of malformed coccoliths with increasing CO2 concentration. We conclude that the sensitivity of the coccolith-shaping machinery to carbonate chemistry changes is strain-specific. Although it has been shown before that carbonate chemistry related changes in growth- and calcification rate are strain-specific, there seems to be no consistent correlation between coccolith morphology and growth or calcification rate. We did not observe an increase in the percentage of incomplete coccoliths in RCC1256, indicating that the coccolith-shaping machinery per se is affected by acidification and not the signalling pathway that produces the stop-signal for coccolith growth.

Seawater carbonate chemistry, calcification and survival of coral recruits in a laboratory experiment

Manipulative studies have demonstrated that ocean acidification (OA) is a threat to coral reefs, yet no experiments have employed diurnal variations in pCO2 that are ecologically relevant to many shallow reefs. Two experiments were conducted to test the response of coral recruits (less than 6 days old) to diurnally oscillating pCO2; one exposing recruits for 3 days to ambient (440 µatm), high (663 µatm) and diurnally oscillating pCO2 on a natural phase (420-596 µatm), and another exposing recruits for 6 days to ambient (456 µatm), high (837 µatm) and diurnally oscillating pCO2 on either a natural or a reverse phase (448-845 µatm). In experiment I, recruits exposed to natural-phased diurnally oscillating pCO2 grew 6-19% larger than those in ambient or high pCO2. In experiment II, recruits in both high and natural-phased diurnally oscillating pCO2 grew 16 per cent larger than those at ambient pCO2, and this was accompanied by 13-18% higher survivorship; the stimulatory effect on growth of oscillatory pCO2 was diminished by administering high pCO2 during the day (i.e. reverse-phased). These results demonstrate that coral recruits can benefit from ecologically relevant fluctuations in pCO2 and we hypothesize that the mechanism underlying this response is highly pCO2-mediated, night-time storage of dissolved inorganic carbon that fuels daytime calcification.

Seawater carbonate chemistry and growth, calcification of Thoracosphaera heimii in a laboratory experiment

Ocean acidification is considered a major threat to marine ecosystems and may particularly affect calcifying organisms such as corals, foraminifera and coccolithophores. Here we investigate the impact of elevated pCO2 and lowered pH on growth and calcification in the common calcareous dinoflagellate Thoracosphaera heimii. We observe a substantial reduction in growth rate, calcification and cyst stability of T. heimii under elevated pCO2. Furthermore, transcriptomic analyses reveal CO2 sensitive regulation of many genes, particularly those being associated to inorganic carbon acquisition and calcification. Stable carbon isotope fractionation for organic carbon production increased with increasing pCO2 whereas it decreased for calcification, which suggests interdependence between both processes. We also found a strong effect of pCO2 on the stable oxygen isotopic composition of calcite, in line with earlier observations concerning another T. heimii strain. The observed changes in stable oxygen and carbon isotope composition of T. heimii cysts may provide an ideal tool for reconstructing past seawater carbonate chemistry, and ultimately past pCO2. Although the function of calcification in T. heimii remains unresolved, this trait likely plays an important role in the ecological and evolutionary success of this species. Acting on calcification as well as growth, ocean acidification may therefore impose a great threat for T. heimii.