General responses of Thoracosphaera heimii grown under a range of pCO2

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.

Alkenone stable carbon isotopes, planktic foraminfera stable carbon isotopes, alkenone unsaturation indices and calculated sea surface temperatures and atmospheric pCO2 for ODP Site 165-999

Temperature reconstructions indicate that the Pliocene was ~3 °C warmer globally than today, and several recent reconstructions of Pliocene atmospheric CO2 indicate that it was above pre-industrial levels and similar to those likely to be seen this century. However, many of these reconstructions have been of relatively low temporal resolution, meaning that these records may have failed to capture variations associated with the 41 Kyr glacial-interglacial cycles thought to operate in the Pliocene. Here we present a new, high temporal resolution alkenone carbon isotope based record of pCO2 spanning 2.8 to 3.3 million years ago from ODP Site 999. Our record is of high enough resolution (~19 Kyrs) to resolve glacial-interglacial changes beyond the intrinsic uncertainty of the proxy method. The record suggests that Pliocene CO2 levels were relatively stable, exhibiting variation less than 55 ppm. We perform sensitivity studies to investigate the possible effect of changing sea surface temperature, which highlights the importance of accurate and precise SST reconstructions for alkenone palaeobarometry, but demonstrate that these uncertainties do not affect our conclusions of relatively stable pCO2 levels during this interval.

Impact of high pCO2 and warmer temperatures on the process of silica biomineralization in the sponge Mycale grandis

Siliceous sponges have survived pre-historical mass extinction events caused by ocean acidification and recent studies suggest that siliceous sponges will continue to resist predicted increases in ocean acidity. In this study, we monitored silica biomineralization in the Hawaiian sponge Mycale grandis under predicted pCO2 and sea surface temperature scenarios for 2100. Our goal was to determine if spicule biomineralization was enhanced or repressed by ocean acidification and thermal stress by monitoring silica uptake rates during short-term (48 h) experiments and comparing biomineralized tissue ratios before and after a long-term (26 d) experiment. In the short-term experiment, we found that silica uptake rates were not impacted by high pCO2 (1050 µatm), warmer temperatures (27°C), or combined high pCO2 with warmer temperature (1119 µatm; 27°C) treatments. The long-term exposure experiments revealed no effect on survival or growth rates of M. grandis to high pCO2 (1198 µatm), warmer temperatures (25.6°C), or combined high pCO2 with warmer temperature (1225 µatm, 25.7°C) treatments, indicating that M. grandis will continue to prosper under predicted increases in pCO2 and sea surface temperature. However, ash-free dry weight to dry weight ratios, subtylostyle lengths, and silicified weight to dry weight ratios decreased under conditions of high pCO2 and combined pCO2 warmer temperature treatments. Our results show that rising ocean acidity and temperature have marginal negative effects on spicule biomineralization and will not affect sponge survival rates of M. grandis.

Impact of atmospheric pCO2, seawater temperature, and calcification rate on the delta 18O and delta 13C composition of echinoid calcite (Echinometra viridis)

The tropical echinoid Echinometra viridis was reared in controlled laboratory experiments at temperatures of approximately 20°C and 30°C to mimic winter and summer temperatures and at carbon dioxide (CO2) partial pressures of approximately 487 ppm-v and 805 ppm-v to simulate current and predicted-end-of-century levels. Spine material produced during the experimental period and dissolved inorganic carbon (DIC) of the corresponding culture solutions were then analyzed for stable oxygen (delta 18Oe, delta 18ODIC) and carbon (The tropical echinoid Echinometra viridis was reared in controlled laboratory experiments at temperatures of approximately 20°C and 30°C to mimic winter and summer temperatures and at carbon dioxide (CO2) partial pressures of approximately 487 ppm-v and 805 ppm-v to simulate current and predicted-end-of-century levels. Spine material produced during the experimental period and dissolved inorganic carbon (DIC) of the corresponding culture solutions were then analyzed for stable oxygen (delta18Oe, delta18ODIC) and carbon (delta13Ce, delta13CDIC) isotopic composition. Fractionation of oxygen stable isotopes between the echinoid spines and DIC of their corresponding culture solutions (delta18O = delta18Oe - delta18ODIC) was significantly inversely correlated with seawater temperature but not significantly correlated with atmospheric pCO2. Fractionation of carbon stable isotopes between the echinoid spines and DIC of their corresponding culture solutions (Delta delta13C = delta13Ce - delta13CDIC) was significantly positively correlated with pCO2 and significantly inversely correlated with temperature, with pCO2 functioning as the primary factor and temperature moderating the pCO2-delta13C relationship. Echinoid calcification rate was significantly inversely correlated with both delta18O and delta13C, both within treatments (i.e., pCO2 and temperature fixed) and across treatments (i.e., with effects of pCO2 and temperature controlled for through ANOVA). Therefore, calcification rate and potentially the rate of co-occurring dissolution appear to be important drivers of the kinetic isotope effects observed in the echinoid spines. Study results suggest that echinoid delta18O monitors seawater temperature, but not atmospheric pCO2, and that echinoid delta13C monitors atmospheric pCO2, with temperature moderating this relationship. These findings, coupled with echinoids' long and generally high-quality fossil record, supports prior assertions that fossil echinoid delta18O is a viable archive of paleo-seawater temperature throughout Phanerozoic time, and that delta13C merits further investigation as a potential proxy of paleo-atmospheric pCO2. However, the apparent impact of calcification rate on echinoid delta18O and delta13C suggests that paleoceanographic reconstructions derived from these proxies in fossil echinoids could be improved by incorporating the effects of growth rate.13Ce, delta13CDIC) isotopic composition. Fractionation of oxygen stable isotopes between the echinoid spines and DIC of their corresponding culture solutions (delta18O = delta18Oe - delta18ODIC) was significantly inversely correlated with seawater temperature but not significantly correlated with atmospheric pCO2. Fractionation of carbon stable isotopes between the echinoid spines and DIC of their corresponding culture solutions (delta13C = delta13Ce - delta13CDIC) was significantly positively correlated with pCO2 and significantly inversely correlated with temperature, with pCO2 functioning as the primary factor and temperature moderating the pCO2-delta13C relationship. Echinoid calcification rate was significantly inversely correlated with both delta18O and delta13C, both within treatments (i.e., pCO2 and temperature fixed) and across treatments (i.e., with effects of pCO2 and temperature controlled for through ANOVA). Therefore, calcification rate and potentially the rate of co-occurring dissolution appear to be important drivers of the kinetic isotope effects observed in the echinoid spines. Study results suggest that echinoid delta18O monitors seawater temperature, but not atmospheric pCO2, and that echinoid delta13C monitors atmospheric pCO2, with temperature moderating this relationship. These findings, coupled with echinoids' long and generally high-quality fossil record, supports prior assertions that fossil echinoid delta18O is a viable archive of paleo-seawater temperature throughout Phanerozoic time, and that delta13C merits further investigation as a potential proxy of paleo-atmospheric pCO2. However, the apparent impact of calcification rate on echinoid delta18O and delta13C suggests that paleoceanographic reconstructions derived from these proxies in fossil echinoids could be improved by incorporating the effects of growth rate.

Echinometra sea urchins acclimatised to elevated pCO2 at volcanic vents outperform those under present-day pCO2 conditions

Rising atmospheric CO2 concentrations will significantly reduce ocean pH during the 21st century (ocean acidification, OA). This may hamper calcification in marine organisms such as corals and echinoderms, as shown in many laboratory-based experiments. Sea urchins are considered highly vulnerable to OA. We studied an Echinometra species on natural volcanic CO2 vents in Papua New Guinea, where they are CO2-acclimatized and also subjected to secondary ecological changes from elevated CO2. Near the vent site, the urchins experienced large daily variations in pH (> 1 unit) and pCO2 (> 2000 ppm) and average pH values (pHT 7.73) much below those expected under the most pessimistic future emission scenarios. Growth was measured over a 17-month period using tetracycline tagging of the calcareous feeding lanterns. Average-sized urchins grew more than twice as fast at the vent compared with those at an adjacent control site, and assumed larger sizes at the vent compared to the control site and two other sites at another reef near-by. A small reduction in gonad weight was detected at the vents, but no differences in mortality, respiration, or degree of test calcification were detected between urchins from vent and control populations. Thus, urchins did not only persist but actually 'thrived' under extreme CO2 conditions. We suggest an ecological basis for this response: increased algal productivity under increased pCO2 provided more food at the vent, resulting in higher growth rates. The wider implication of our observation is that laboratory studies on non-acclimatized specimens, which typically do not consider ecological changes, can lead to erroneous conclusions on responses to global change.