Seawater carbonate chemistry and processes during experiments with Emiliania huxleyi (PML B92/11), 2000

We have measured the stable carbon isotopic composition of bulk organic matter (POC), alkenones, sterols, fatty acids, and phytol in the coccolithophorid Emiliania huxleyi grown in dilute batch cultures over a wide range of CO2 concentrations (1.1-53.5 micromol L-1). The carbon isotope fractionation of POC (POC) varied by ca. 7 per mil and was positively correlated with aqueous CO2 concentration [CO2aq]. While this result confirms general trends observed for the same alga grown in nitrogen-limited chemostat cultures, considerable differences were obtained in absolute values of POC and in the slope of the relationship of POC with growth rate and [CO2aq]. Also, a significantly greater offset was obtained between the delta13C of alkenones and bulk organic matter in this study compared with previous work (5.4, cf. 3.8 per mil). This suggests that the magnitude of the isotope offset may depend on growth conditions. Relative to POC, individual fatty acids were depleted in 13C by 2.3 per mil to 4.1 per mil, phytol was depleted in 13C by 1.9 per mil, and the major sterol 24-methylcholesta-5,22E-dien-3beta-ol was depleted in 13C by 8.5 per mil. This large spread of delta13C values for different lipid classes in the same alga indicates the need for caution in organic geochemical studies when assigning different sources to lipids that might have delta13C values differing by just a few per mil. Increases in [CO2aq] led to dramatic increases in the alkenone contents per cell and as a proportion of organic carbon, but there was no systematic effect on values of U37k- used for reconstructions of paleo sea surface temperature.

Seawater carbonate chemistry and Ruditapes decussatus biological processes during experiments, 2011

We investigated the effects of ocean acidification on juvenile clams Ruditapes decussatus (average shell length 10.24 mm) in a controlled CO2 perturbation experiment. The carbonate chemistry of seawater was manipulated by diffusing pure CO2, to attain two reduced pH levels (by -0.4 and -0.7 pH units), which were compared to unmanipulated seawater. After 75 days we found no differences among pH treatments in terms of net calcification, size or weight of the clams. The naturally elevated total alkalinity of local seawater probably contributed to buffer the effects of increased pCO2 and reduced pH. Marine organisms may, therefore, show diverse responses to ocean acidification at local scales, particularly in coastal, estuarine and transitional waters, where the physical-chemical characteristics of seawater are most variable. Mortality was significantly reduced in the acidified treatments. This trend was probably related to the occurrence of spontaneous spawning events in the control and intermediate acidification treatments. Spawning, which was unexpected due to the small size of the clams, was not observed for the pH -0.7 treatment, suggesting that the increased survival under acidified conditions may have been associated with a delay in the reproductive cycle of the clams. Future research about the impacts of ocean acidification on marine biodiversity should be extended to other types of biological and ecological processes, apart from biological calcification.

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 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.

Wet tundra polygon centers and distance between polygon centroids on Samoylov Island, Lena Delta, Siberia, Russia, with links to shape files

Subgrid processes occur in various ecosystems and landscapes but, because of their small scale, they are not represented or poorly parameterized in climate models. These local heterogeneities are often important or even fundamental for energy and carbon balances. This is especially true for northern peatlands and in particular for the polygonal tundra, where methane emissions are strongly influenced by spatial soil heterogeneities. We present a stochastic model for the surface topography of polygonal tundra using Poisson-Voronoi diagrams and we compare the results with available recent field studies. We analyze seasonal dynamics of water table variations and the landscape response under different scenarios of precipitation income. We upscale methane fluxes by using a simple idealized model for methane emission. Hydraulic interconnectivities and large-scale drainage may also be investigated through percolation properties and thresholds in the Voronoi graph. The model captures the main statistical characteristics of the landscape topography, such as polygon area and surface properties as well as the water balance. This approach enables us to statistically relate large-scale properties of the system to the main small-scale processes within the single polygons.

Seawater carbonate chemistry and biological processes during experiments with spider crab Hyas araneus, 2011

The combined effects of ocean warming and acidification were compared in larvae from two populations of the cold-eurythermal spider crab Hyas araneus, from one of its southernmost populations (around Helgoland, southern North Sea, 54°N, habitat temperature 3-18°C; collection: January 2008, hatch: January-February 2008) and from one of its northernmost populations (Svalbard, North Atlantic, 79°N, habitat temperature 0-6°C; collection: July 2008, hatch: February-April 2009). Larvae were exposed to temperatures of 3, 9 and 15°C combined with present-day normocapnic (380 ppm CO2) and projected future CO2 concentrations (710 and 3,000 ppm CO2). Calcium content of whole larvae was measured in freshly hatched Zoea I and after 3, 7 and 14 days during the Megalopa stage. Significant differences between Helgoland and Svalbard Megalopae were observed at all investigated temperatures and CO2 conditions. Under 380 ppm CO2, the calcium content increased with rising temperature and age of the larvae. At 3 and 9°C, Helgoland Megalopae accumulated more calcium than Svalbard Megalopae. Elevated CO2 levels, especially 3,000 ppm, caused a reduction in larval calcium contents at 3 and 9°C in both populations. This effect set in early, at 710 ppm CO2 only in Svalbard Megalopae at 9°C. Furthermore, at 3 and 9°C Megalopae from Helgoland replenished their calcium content to normocapnic levels and more rapidly than Svalbard Megalopae. However, Svalbard Megalopae displayed higher calcium contents under 3,000 ppm CO2 at 15°C. The findings of a lower capacity for calcium incorporation in crab larvae living at the cold end of their distribution range suggests that they might be more sensitive to ocean acidification than those in temperate regions.

Seawater carbonate chemistry, fertilization rate and biological processes of sea urchin Paracentrotus lividus during experiments, 2011

The effect of pH ranging from 8.0 to 6.8 (total scale - pHT) on fertilization, cleavage and larval development until pluteus stage was assessed in an intertidal temperate sea urchin. Gametes were obtained from adults collected in two contrasting tide pools, one showing a significant nocturnal pH decrease (lowest pHT = 7.4) and another where pH was more stable (lowest pHT = 7.8). The highest pHT at which significant effects on fertilization and cleavage were recorded was 7.6. On the contrary, larval development was only affected below pHT 7.4, a value equal or lower than that reported for several subtidal species. This suggests that sea urchins inhabiting stressful intertidal environments produce offspring that may better resist future ocean acidification. Moreover, at pHT 7.4, the fertilization rate of gametes whose progenitors came from the tide pool with higher pH decrease was significantly higher, indicating a possible acclimatization or adaptation of gametes to pH stress.

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 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 processes during an experiment with coral reef, 2000

The concentration of CO2 in the atmosphere is projected to reach twice the preindustrial level by the middle of the 21st century. This increase will reduce the concentration of [CO3]2- of the surface ocean by 30% relative to the preindustrial level and will reduce the calcium carbonate saturation state of the surface ocean by an equal percentage. Using the large 2650 m3 coral reef mesocosm at the BIOSPHERE-2 facility near Tucson, Arizona, we investigated the effect of the projected changes in seawater carbonate chemistry on the calcification of coral reef organisms at the community scale. Our experimental design was to obtain a long (3.8 years) time series of the net calcification of the complete system and all relevant physical and chemical variables (temperature, salinity, light, nutrients, Ca2+,pCO2, TCO2, and total alkalinity). Periodic additions of NaHCO3, Na2CO3, and/or CaCl2 were made to change the calcium carbonate saturation state of the water. We found that there were consistent and reproducible changes in the rate of calcification in response to our manipulations of the saturation state. We show that the net community calcification rate responds to manipulations in the concentrations of both Ca2+ and [CO3]2- and that the rate is well described as a linear function of the ion concentration product, [Ca2+]0.69[[CO3]2-]. This suggests that saturation state or a closely related quantity is a primary environmental factor that influences calcification on coral reefs at the ecosystem level. We compare the sensitivity of calcification to short-term (days) and long-term (months to years) changes in saturation state and found that the response was not significantly different. This indicates that coral reef organisms do not seem to be able to acclimate to changing saturation state. The predicted decrease in coral reef calcification between the years 1880 and 2065 A.D. based on our long-term results is 40%. Previous small-scale, short-term organismal studies predicted a calcification reduction of 14-30%. This much longer, community-scale study suggests that the impact on coral reefs may be greater than previously suspected. In the next century coral reefs will be less able to cope with rising sea level and other anthropogenic stresses.

Chemistry and biological processes during experiments with spider crab Hyas araneus

The combined effects of ocean warming and acidification were compared in larvae from two popula- tions of the cold-eurythermal spider crab Hyas araneus, from one of its southernmost populations (around Helgo- land, southern North Sea, 54°N, habitat temperature 3-18°C; collection: January 2008, hatch: January-February 2008) and from one of its northernmost populations (Svalbard, North Atlantic, 79°N, habitat temperature 0-6°C; collection: July 2008, hatch: February-April 2009). Larvae were exposed to temperatures of 3, 9 and 15°C combined with present-day normocapnic (380 ppm CO2) and projected future CO2 concentrations (710 and 3,000 ppm CO2). Calcium content of whole larvae was measured in freshly hatched Zoea I and after 3, 7 and 14 days during the Megalopa stage. Significant differences between Helgoland and Svalbard Megalopae were observed at all investigated temperatures and CO2 condi- tions. Under 380 ppm CO2, the calcium content increased with rising temperature and age of the larvae. At 3 and 9°C, Helgoland Megalopae accumulated more calcium than Svalbard Megalopae. Elevated CO2 levels, especially 3,000 ppm, caused a reduction in larval calcium contents at 3 and 9°C in both populations. This effect set in early, at 710 ppm CO2 only in Svalbard Megalopae at 9°C. Fur- thermore, at 3 and 9°C Megalopae from Helgoland replenished their calcium content to normocapnic levels and more rapidly than Svalbard Megalopae. However, Svalbard Megalopae displayed higher calcium contents under 3,000 ppm CO2 at 15°C. The findings of a lower capacity for calcium incorporation in crab larvae living at the cold end of their distribution range suggests that they might be more sensitive to ocean acidification than those in temperate regions.

Seawater carbonate chemistry and biological processes during experiments with the spider crab Hyas araneus in Kongsfjorden, Svalbard, 2009

With global climate change, ocean warming and acidification occur concomitantly. In this study, we tested the hypothesis that increasing CO2 levels affect the acid-base balance and reduce the activity capacity of the Arctic spider crab Hyas araneus, especially at the limits of thermal tolerance. Crabs were acclimated to projected oceanic CO2 levels for 12 days (today: 380, towards the year 2100: 750 and 1,120 and beyond: 3,000 ?atm) and at two temperatures (1 and 4 °C). Effects of these treatments on the righting response (RR) were determined (1) at acclimation temperatures followed by (2) righting when exposed to an additional acute (15 min) heat stress at 12 °C. Prior to (resting) and after the consecutive stresses of combined righting activity and heat exposure, acid-base status and lactate contents were measured in the haemolymph. Under resting conditions, CO2 caused a decrease in haemolymph pH and an increase in oxygen partial pressure. Despite some buffering via an accumulation of bicarbonate, the extracellular acidosis remained uncompensated at 1 °C, a trend exacerbated when animals were acclimated to 4 °C. The additional combined exposure to activity and heat had only a slight effect on blood gas and acid-base status. Righting activity in all crabs incubated at 1 and 4 °C was unaffected by elevated CO2 levels or acute heat stress but was significantly reduced when both stressors acted synergistically. This impact was much stronger in the group acclimated at 1 °C where some individuals acclimated to high CO2 levels stopped responding. Lactate only accumulated in the haemolymph after combined righting and heat stress. In the group acclimated to 1 °C, lactate content was highest under normocapnia and lowest at the highest CO2 level in line with the finding that RR was largely reduced. In crabs acclimated to 4 °C, the RR was less affected by CO2 such that activity caused lactate to increase with rising CO2 levels. In line with the concept of oxygen and capacity limited thermal tolerance, all animals exposed to temperature extremes displayed a reduction in scope for performance, a trend exacerbated by increasing CO2 levels. Additionally, the differences seen between cold- and warm-acclimated H. araneus after heat stress indicate that a small shift to higher acclimation temperatures also alleviates the response to temperature extremes, indicating a shift in the thermal tolerance window which reduces susceptibility to additional CO2 exposure.