Seawater carbonate chemistry, nutrients, particulate carbon and growth rate of Emiliania huxleyi (AC472), Calcidiscus leptoporus (AC370) and Syracosphaera pulchra (AC418) during experiments, 2011

The response of Emiliania huxleyi (Lohmann) W. W. Hay et H. Mohler, Calcidiscus leptoporus (G. Murray et V. H. Blackman) J. Schiller, andSyracosphaera pulchra Lohmann to elevated partial pressure of carbon dioxide (pCO2) was investigated in batch cultures. We reported on the response of both haploid and diploid life stages of these three species. Growth rate, cell size, particulate inorganic carbon (PIC), and particulate organic carbon (POC) of both life stages were measured at two different pCO2 (400 and 760 parts per million [ppm]), and their organic and inorganic carbon production were calculated. The two life stages within the same species generally exhibited a similar response to elevated pCO2, the response of the haploid stage being often more pronounced than that of the diploid stage. The growth rate was consistently higher at elevated pCO2, but the response of other processes varied among species. Calcification rate of C. leptoporusand of S. pulchra did not change at elevated pCO2, whereas it increased in E. huxleyi. POC production and cell size of both life stages of S. pulchra and of the haploid stage of E. huxleyi markedly decreased at elevated pCO2. It remained unaltered in the diploid stage of E. huxleyi and C. leptoporus and increased in the haploid stage of the latter. The PIC:POC ratio increased in E. huxleyi and was constant in C. leptoporus and S. pulchra. Elevated pCO2 has a significant effect on these three coccolithophore species, the haploid stage being more sensitive. This effect must be taken into account when predicting the fate of coccolithophores in the future ocean.

Seawater carbonate chemistry and biological processes during experiments with haploid and diploid life stages of Emiliania huxleyi, Calcidiscus leptoporus and Syracosphaera pulchra

The response of Emiliania huxleyi (Lohmann), Calcidiscus leptoporus (Murray and Blackman), and Syracosphaera pulchra (Lohmann) to elevated partial pressure of carbon dioxide (pCO2) was investigated in batch cultures. For the first time, we reported on the response of the non-calcifying (haploid) life stage of these three species. Growth rate, cell size, particulate inorganic (PIC) and particulate organic carbon (POC) of both life stages were measured at two different pCO2 (400 and 760 ppm) and their organic and inorganic carbon production calculated. The two life stages within the same species generally exhibited a similar response to elevated pCO2, the response of the haploid stage being often more pronounced than that of the diploid stage. The growth rate was consistently higher at elevated pCO2 but the response of other processes varied among species. Calcification rate of C. leptoporus and of S. pulchra did not change at elevated pCO2 while it increased in E. huxleyi. Particulate organic carbon production and cell size of both life stages of S. pulchra and of the haploid stage of E. huxleyi markedly decreased at elevated pCO2. It remained unaltered in the diploid stage of E. huxleyi and C. leptoporus and increased in the haploid stage of the latter. The PIC:POC ratio increased in E. huxleyi and was constant in C. leptoporus and S. pulchra. Elevated pCO2 has a significant effect on these three coccolithophores species, the haploid stage being more sensitive. This must be taken into account when predicting the fate of coccolithophores in the future ocean.

Seawater carbonate chemistry and processes during experiments with Crassostrea gigas, 2007

This study demonstrated that the increased partial pressure of CO2 (pCO2) in seawater and the attendant acidification that are projected to occur by the year 2300 will severely impact the early development of the oyster Crassostrea gigas. Eggs of the oyster were artificially fertilized and incubated for 48 h in seawater acidified to pH 7.4 by equilibrating it with CO2-enriched air (CO2 group), and the larval morphology and degree of shell mineralization were compared with the control treatment (air-equilibrated seawater). Only 5% of the CO2 group developed into normal 'D-shaped' veliger larvae as compared with 68% in the control group, although no difference was observed between the groups up to the trochophore stage. Thus, during embryogenesis, the calcification process appears to be particularly affected by low pH and/or the low CaCO3 saturation state of high-CO2 seawater. Veliger larvae with fully mineralized shells accounted for 30% of the CO2-group larvae, compared with 72% in the control (p < 0.005). Shell mineralization was completely inhibited in 45% of the CO2-group larvae, but only in 16% of the control (p < 0.05). Normal D-shaped veligers of the control group exhibited increased shell length and height between 24 and 48 h after fertilization, while the few D-shaped veligers of the CO2 group showed no shell growth during the same period. Our results suggest that future ocean acidification will have deleterious impacts on the early development of marine benthic calcifying organisms.

Seawater carbonate chemistry, survival, metamorphosis and respiration rate of coral Acropora digitifera during experiments, 2011

Ocean acidification may negatively impact the early life stages of some marine invertebrates including corals. Although reduced growth of juvenile corals in acidified seawater has been reported, coral larvae have been reported to demonstrate some level of tolerance to reduced pH. We hypothesize that the observed tolerance of coral larvae to low pH may be partly explained by reduced metabolic rates in acidified seawater because both calcifying and non-calcifying marine invertebrates could show metabolic depression under reduced pH in order to enhance their survival. In this study, after 3-d and 7-d exposure to three different pH levels (8.0, 7.6, and 7.3), we found that the oxygen consumption of Acropora digitifera larvae tended to be suppressed with reduced pH, although a statistically significant difference was not observed between pH conditions. Larval metamorphosis was also observed, confirming that successful recruitment is impaired when metamorphosis is disrupted, despite larval survival. Results also showed that the metamorphosis rate significantly decreased under acidified seawater conditions after both short (2 h) and long (7 d) term exposure. These results imply that acidified seawater impacts larval physiology, suggesting that suppressed metabolism and metamorphosis may alter the dispersal potential of larvae and subsequently reduce the resilience of coral communities in the near future as the ocean pH decreases.

Seawater carbonate chemistry and biological processes of Porites panamensis during experiments, 2011

Survival of coral planulae, and the successful settlement and healthy growth of primary polyps are critical for the dispersal of scleractinian corals and hence the recovery of degraded coral reefs. It is therefore important to explore how the warmer and more acidic oceanic conditions predicted for the future could affect these processes. This study used controlled culture to investigate the effects of a 1 °C increase in temperature and a 0.2-0.25 unit decrease in pH on the settlement and survival of planulae and the growth of primary polyps in the Tropical Eastern Pacific coral Porites panamensis. We found that primary polyp growth was reduced only marginally by more acidic seawater but the combined effect of high temperature and lowered pH caused a significant reduction in growth of primary polyps by almost a third. Elevated temperature was found to significantly reduce the amount of zooxanthellae in primary polyps, and when combined with lowered pH resulted in a significant reduction in biomass of primary polyps. However, survival and settlement of planula larvae were unaffected by increased temperature, lowered acidity or the combination of both. These results indicate that in future scenarios of increased temperature and oceanic acidity coral planulae will be able to disperse and settle successfully but primary polyp growth may be hampered. The recovery of reefs may therefore be impeded by global change even if local stressors are curbed and sufficient sources of planulae are available.

Wave height (regular and irregular waves) reduction in 2m water depth over a 40-m-long salt marsh test section in a large scale laboratory flume

Coastal communities around the world face increasing risk from flooding as a result of rising sea level, increasing storminess, and land subsidence. Salt marshes can act as natural buffer zones, providing protection from waves during storms. However, the effectiveness of marshes in protecting the coastline during extreme events when water levels and waves are highest is poorly understood. Here, we experimentally assess wave dissipation under storm surge conditions in a 300-m-long wave flume that contains a transplanted section of natural salt marsh. We find that the presence of marsh vegetation causes considerable wave attenuation, even when water levels and waves are high. From a comparison with experiments without vegetation, we estimate that up to 60% of observed wave reduction is attributed to vegetation. We also find that although waves progressively flatten and break vegetation stems and thereby reduce dissipation, the marsh substrate remained remarkably stable and resistant to surface erosion under all conditions.The effectiveness of storm wave dissipation and the resilience of tidal marshes even at extreme conditions suggest that salt marsh ecosystems can be a valuable component of coastal protection schemes.

Seawater carbonate chemistry, survival rate, shell size, calcification rate of the planktonic foraminifer Neogloboquadrina pachyderma (sinistral) in a laboratory experiment

The present study investigated the effects of ocean acidification and temperature increase on Neogloboquadrina pachyderma (sinistral), the dominant planktonic foraminifer in the Arctic Ocean. Due to the naturally low concentration of [CO3] 2- in the Arctic, this foraminifer could be particularly sensitive to the forecast changes in seawater carbonate chemistry. To assess potential responses to ocean acidification and climate change, perturbation experiments were performed on juvenile and adult specimens by manipulating seawater to mimic the present-day carbon dioxide level and a future ocean acidification scenario (end of the century) under controlled (in situ) and elevated temperatures (1 and 4 °C, respectively). Foraminifera mortality was unaffected under all the different experiment treatments. Under low pH, N. pachyderma (s) shell net calcification rates decreased. This decrease was higher (30 %) in the juvenile specimens than decrease observed in the adults (21 %) ones. However, decrease in net calcification was moderated when both, pH decreased and temperature increased simultaneously. When only temperature increased, a net calcification rate for both life stages was not affected. These results show that forecast changes in seawater chemistry would impact calcite production in N. pachyderma (s), possibly leading to a reduction of calcite flux contribution and consequently a decrease in biologic pump efficiency.

Seawater carbonate chemistry and particulate organic particles during an incubation experiments with natural phytoplankton community, 2002

Incubation experiments with natural phytoplankton revealed a relationship between CO2 concentration and the production of transparent exopolymer particles (TEP), with TEP production being linearly related to theoretical CO2 uptake rates. The effect of different CO2 concentrations on TEP production was examined during incubation experiments with natural phytoplankton sampled at two different locations in the central Baltic Sea in summer 1999.

Seawater carbonate chemistry and biological processes during experiments with Limacina helicina, 2009

Thecosome pteropods (pelagic mollusks) can play a key role in the food web of various marine ecosystems. They are a food source for zooplankton or higher predators such as fishes, whales and birds that is particularly important in high latitude areas. Since they harbor a highly soluble aragonitic shell, they could be very sensitive to ocean acidification driven by the increase of anthropogenic CO2 emissions. The effect of changes in the seawater chemistry was investigated on Limacina helicina, a key species of Arctic pelagic ecosystems. Individuals were kept in the laboratory under controlled pCO2 levels of 280, 380, 550, 760 and 1020 µatm and at control (0°C) and elevated (4°C) temperatures. The respiration rate was unaffected by pCO2 at control temperature, but significantly increased as a function of the pCO2 level at elevated temperature. pCO2 had no effect on the gut clearance rate at either temperature. Precipitation of CaCO3, measured as the incorporation of 45Ca, significantly declined as a function of pCO2 at both temperatures. The decrease in calcium carbonate precipitation was highly correlated to the aragonite saturation state. Even though this study demonstrates that pteropods are able to precipitate calcium carbonate at low aragonite saturation state, the results support the current concern for the future of Arctic pteropods, as the production of their shell appears to be very sensitive to decreased pH. A decline of pteropod populations would likely cause dramatic changes to various pelagic ecosystems.

Seawater carbonate chemistry and expression of hsp70, hsp90 and hsf1 in the reef coral Acropora digitifera during experiments, 2012

Ocean acidification is an ongoing threat for marine organisms due to the increasing atmospheric CO2 concentration. Seawater acidification has a serious impact on physiologic processes in marine organisms at all life stages. On the other hand, potential tolerance to external pH changes has been reported in coral larvae. Information about the possible mechanisms underlying such tolerance responses, however, is scarce. In the present study, we examined the effects of acidified seawater on the larvae of Acropora digitifera at the molecular level. We targeted two heat shock proteins, Hsp70 and Hsp90, and a heat shock transcription factor, Hsf1, because of their importance in stress responses and in early life developmental stages. Coral larvae were maintained under the ambient and elevated CO2 conditions that are expected to occur within next 100 years, and then we evaluated the expression of hsps and hsf1 by quantitative real-time polymerase chain reaction (PCR). Expression levels of these molecules significantly differed among target genes, but they did not change significantly between CO2conditions. These findings indicate that the expression of hsps is not changed due to external pH changes, and suggest that tolerance to acidified seawater in coral larvae may not be related to hsp expression.

Seawater carbonate chemistry and biological processes during experiments with phytoplankton Emiliania huxleyi (CS369), 2009

Increasing atmospheric CO2 concentration affects calcification in most planktonic calcifiers. Both reduced or stimulated calcification under high CO2 have been reported in the widespread coccolithophore Emiliania huxleyi. This might affect the response of cells to photosynthetically active radiation (PAR; 400-700 nm) and ultraviolet radiation (UVR; 280-400 nm) by altering the thickness of the coccolith layer. Here we show that in the absence of UVR, the calcification rates in E. huxleyi decrease under lowered pH levels (pHNBS of 7.9 and 7.6; pCO2 of 81 and 178 Pa or 804 and 1759 ppmv, respectively) leading to thinned coccolith layers, whereas photosynthetic carbon fixation was slightly enhanced at pH 7.9 but remained unaffected at pH 7.6. Exposure to UVR (UV-A 19.5 W m**-2, UV-B 0.67 W m**-2) in addition to PAR (88.5 W m**-2), however, results in significant inhibition of both photosynthesis and calcification, and these rates are further inhibited with increasing acidification. The combined effects of UVR and seawater acidification resulted in the inhibition of calcification rates by 96% and 99% and that of photosynthesis by 6% and 15%, at pH 7.9 and 7.6, respectively. This differential inhibition of calcification and photosynthesis leads to significant reduction of the ratio of calcification to photosynthesis. Seawater acidification enhanced the transmission of harmful UVR by about 26% through a reduction of the coccolith layer of 31%. Our data indicate that the effect of a high-CO2 and low-pH ocean on E. huxleyi (because of reduced calcification associated with changes in the carbonate system) enhances the detrimental effects of UVR on the main pelagic calcifier.

Seawater carbonate chemistry and carbon and oxygen isotopes during experiments with planktonic foraminifera Orbulina universa and Globigerina bulloides, 1997

Stable oxygen and carbon isotope measurements on biogenic calcite and aragonite have become standard tools for reconstructing past oceanographic and climatic change. In aquatic organisms, 18O/16O ratios in the shell carbonate are a function of the ratio in the sea water and the calcification temperature (Epstein et al., 1953). In contrast, 13C/12C ratios are controlled by the ratio of dissolved inorganic carbon in sea water and physiological processes such as respiration and symbiont photosynthesis (Spero et al., 1991, doi:10.1029/91PA02022). These geochemical proxies have been used with analyses of foraminifera shells to reconstruct global ice volumes (Shackleton and Opdyke, 1973, doi:10.1016/0033-5894(73)90052-5), surface and deep ocean temperatures (Broecker, 1986, doi:10.1016/0033-5894(86)90087-6; Labeyrie et al., 1987, doi:10.1038/327477a0), ocean circulation changes (Duplessy et al., 1988, doi:10.1029/PA003i003p00343) and glacial-interglacial exchange between the terrestrial and oceanic carbon pools (Sackleton, 1977). Here, we report experimental measurements on living symbiotic and non-symbiotic plankton foraminifera (Orbulina universa and Globigerina bulloides respectively) showing that the 13C/12C and 18O/16O ratios of the calcite shells decrease with increasing seawater [CO3 2-]. Because glacial-period oceans had higher pH and [CO3 2-] than today (Sanyal et al., 1995, doi:10.1038/373234a0), these new relationships confound the standard interpretation of glacial foraminiferal stable-isotope data. In particular, the hypothesis that the glacial-interglacial shift in the 13C/12C ratio was due to a transfer of terrestrial carbon into the ocean(Shackleton ,1977) can be explained alternatively by an increase in ocean alkalinity (Lea et al., 1996). A carbonate-concentration effect could also help explain some of the extreme stable-isotope variations during the Proterozoic and Phanerozoic aeons (Kaufman et al., 1993, doi:10.1016/0012-821X(93)90254-7).

Seawater carbonate chemistry and resource allocation and extracellular acid-base status in the sea urchin Strongylocentrotus droebachiensis during experiments, 2012

Anthropogenic CO2 emission will lead to an increase in seawater pCO2 of up to 80-100 Pa (800-1000 µatm) within this century and to an acidification of the oceans. Green sea urchins (Strongylocentrotus droebachiensis) occurring in Kattegat experience seasonal hypercapnic and hypoxic conditions already today. Thus, anthropogenic CO2 emissions will add up to existing values and will lead to even higher pCO2 values >200 Pa (>2000 µatm). To estimate the green sea urchins' potential to acclimate to acidified seawater, we calculated an energy budget and determined the extracellular acid base status of adult S. droebachiensis exposed to moderately (102 to 145 Pa, 1007 to 1431 µatm) and highly (284 to 385 Pa, 2800 to 3800 µatm) elevated seawater pCO2 for 10 and 45 days. A 45 - day exposure to elevated pCO2 resulted in a shift in energy budgets, leading to reduced somatic and reproductive growth. Metabolic rates were not significantly affected, but ammonium excretion increased in response to elevated pCO2. This led to decreased O:N ratios. These findings suggest that protein metabolism is possibly enhanced under elevated pCO2 in order to support ion homeostasis by increasing net acid extrusion. The perivisceral coelomic fluid acid-base status revealed that S. droebachiensis is able to fully (intermediate pCO2) or partially (high pCO2) compensate extracellular pH (pHe) changes by accumulation of bicarbonate (maximum increases 2.5 mM), albeit at a slower rate than typically observed in other taxa (10 day duration for full pHe compensation). At intermediate pCO2, sea urchins were able to maintain fully compensated pHe for 45 days. Sea urchins from the higher pCO2 treatment could be divided into two groups following medium-term acclimation: one group of experimental animals (29%) contained remnants of food in their digestive system and maintained partially compensated pHe (+2.3 mM HCO3), while the other group (71%) exhibited an empty digestive system and a severe metabolic acidosis (-0.5 pH units, -2.4 mM HCO3). There was no difference in mortality between the three pCO2 treatments. The results of this study suggest that S. droebachiensis occurring in the Kattegat might be pre-adapted to hypercapnia due to natural variability in pCO2 in its habitat. We show for the first time that some echinoderm species can actively compensate extracellular pH. Seawater pCO2 values of >200 Pa, which will occur in the Kattegat within this century during seasonal hypoxic events, can possibly only be endured for a short time period of a few weeks. Increases in anthropogenic CO2 emissions and leakages from potential sub-seabed CO2 storage (CCS) sites thus impose a threat to the ecologically and economically important species S. droebachiensis.