Seawater carbonate chemistry and calcification rate of cold-water coral Lophelia pertusa during experiments, 2011

Ocean acidity has increased by 30% since preindustrial times due to the uptake of anthropogenic CO2 and is projected to rise by another 120% before 2100 if CO2 emissions continue at current rates. Ocean acidification is expected to have wide-ranging impacts on marine life, including reduced growth and net erosion of coral reefs. Our present understanding of the impacts of ocean acidification on marine life, however, relies heavily on results from short-term CO2 perturbation studies. Here we present results from the first long-term CO2 perturbation study on the dominant reef-building cold-water coral Lophelia pertusa and relate them to results from a short-term study to compare the effect of exposure time on the coral's responses. Short-term (one week) high CO2 exposure resulted in a decline of calcification by 26-29% for a pH decrease of 0.1 units and net dissolution of calcium carbonate. In contrast, L. pertusa was capable to acclimate to acidified conditions in long-term (six months) incubations, leading to even slightly enhanced rates of calcification. Net growth is sustained even in waters sub-saturated with respect to aragonite. Acclimation to seawater acidification did not cause a measurable increase in metabolic rates. This is the first evidence of successful acclimation in a coral species to ocean acidification, emphasizing the general need for long-term incubations in ocean acidification research. To conclude on the sensitivity of cold-water coral reefs to future ocean acidification further ecophysiological studies are necessary which should also encompass the role of food availability and rising temperatures.

Seawater carbonate chemistry during experiments with Stylophora pistillata, 2008

The decrease in the saturation state of seawater, following seawater acidification, is believed to be the main factor leading to a decrease in the calcification of marine organisms. To provide a physiological explanation for this phenomenon, the effect of seawater acidification was studied on the calcification and photosynthesis of the scleractinian tropical coral Stylophora pistillata. Coral nubbins were incubated for 8 days at three different pH (7.6, 8.0, and 8.2). To differentiate between the effects of the various components of the carbonate chemistry (pH, CO32, HCO3, CO2), tanks were also maintained under similar pH, but with 2-mM HCO3 added to the seawater. The addition of 2-mM bicarbonate significantly increased the photosynthesis in S. pistillata, suggesting carbon-limited conditions. Conversely, photosynthesis was insensitive to changes in pH and pCO2. Seawater acidification decreased coral calcification by ca. 0.1-mg CaCO3 g-1 d-1 for a decrease of 0.1 pH units. This correlation suggested that seawater acidification affected coral calcification by decreasing the availability of the CO32 substrate for calcification. However, the decrease in coral calcification could also be attributed either to a decrease in extra- or intracellular pH or to a change in the buffering capacity of the medium, impairing supply of CO32 from HCO3.

Seawater carbonate chemistry, mortality and hatching rate, size and mass of Clupea harengus during experiments, 2011

Due to atmospheric accumulation of anthropogenic CO2 the partial pressure of carbon dioxide (pCO2) in surface seawater increases and the pH decreases. This process known as ocean acidification might have severe effects on marine organisms and ecosystems. The present study addresses the effect of ocean acidification on early developmental stages, the most sensitive stages in life history, of the Atlantic herring (Clupea harengus L.). Eggs of the Atlantic herring were fertilized and incubated in artificially acidified seawater (pCO2 1260, 1859, 2626, 2903, 4635 µatm) and a control treatment (pCO2 480 µatm) until the main hatch of herring larvae occurred. The development of the embryos was monitored daily and newly hatched larvae were sampled to analyze their morphometrics, and their condition by measuring the RNA/DNA ratios. Elevated pCO2 neither affected the embryogenesis nor the hatch rate. Furthermore the results showed no linear relationship betweenpCO2 and total length, dry weight, yolk sac area and otolith area of the newly hatched larvae. For pCO2 and RNA/DNA ratio, however, a significant negative linear relationship was found. The RNA concentration at hatching was reduced at higher pCO2 levels, which could lead to a decreased protein biosynthesis. The results indicate that an increased pCO2 can affect the metabolism of herring embryos negatively. Accordingly, further somatic growth of the larvae could be reduced. This can have consequences for the larval fish, since smaller and slow growing individuals have a lower survival potential due to lower feeding success and increased predation mortality. The regulatory mechanisms necessary to compensate for effects of hypercapnia could therefore lead to lower larval survival. Since the recruitment of fish seems to be determined during the early life stages, future research on the factors influencing these stages are of great importance in fisheries science.

Response of benthic foraminifera to ocean acidification in their natural sediment environment: a long-term culturing experiment

Calcifying foraminifera are expected to be endangered by ocean acidification; however, the response of a complete community kept in natural sediment and over multiple generations under controlled laboratory conditions has not been constrained to date. During 6 months of incubation, foraminiferal assemblages were kept and treated in natural sediment with pCO2-enriched seawater of 430, 907, 1865 and 3247 µatm pCO2. The fauna was dominated by Ammonia aomoriensis and Elphidium species, whereas agglutinated species were rare. After 6 months of incubation, pore water alkalinity was much higher in comparison to the overlying seawater. Consequently, the saturation state of Omega calc was much higher in the sediment than in the water column in nearly all pCO2 treatments and remained close to saturation. As a result, the life cycle (population density, growth and reproduction) of living assemblages varied markedly during the experimental period, but was largely unaffected by the pCO2 treatments applied. According to the size-frequency distribution, we conclude that foraminifera start reproduction at a diameter of 250 µm. Mortality of living Ammonia aomoriensis was unaffected, whereas size of large and dead tests decreased with elevated pCO2 from 285 µm (pCO2 from 430 to 1865 µatm) to 258 µm (pCO2 3247 µatm). The total organic content of living Ammonia aomoriensis has been determined to be 4.3% of CaCO3 weight. Living individuals had a calcium carbonate production rate of 0.47 g/m**2/a, whereas dead empty tests accumulated a rate of 0.27 g /m**2/a. Although Omega calc was close to 1, approximately 30% of the empty tests of Ammonia aomoriensis showed dissolution features at high pCO2 of 3247 µatm during the last 2 months of incubation. In contrast, tests of the subdominant species, Elphidium incertum, stayed intact. Our results emphasize that the sensitivity to ocean acidification of the endobenthic foraminifera Ammonia aomoriensis in their natural sediment habitat is much lower compared to the experimental response of specimens isolated from the sediment.

Seawater carbonate chemistry, shell linear extension and calcification during calcein staining and 45Ca experiments with pteropod Limacina helicina, 2009

Thecosome pteropods (shelled pelagic molluscs) can play an important role in the food web of various ecosystems and play a key role in the cycling of carbon and carbonate. Since they harbor an aragonitic shell, they could be very sensitive to ocean acidification driven by the increase of anthropogenic CO2 emissions. The impact of changes in the carbonate chemistry was investigated on Limacina helicina, a key species of Arctic ecosystems. Pteropods were kept in culture under controlled pH conditions corresponding to pCO2 levels of 350 and 760 µatm. Calcification was estimated using a fluorochrome and the radioisotope 45Ca. It exhibits a 28 % decrease at the pH value expected for 2100 compared to the present pH value. This result supports the concern for the future of pteropods in a high-CO2 world, as well as of those species dependent upon them as a food resource. A decline of their populations would likely cause dramatic changes to the structure, function and services of polar ecosystems.

Seawater carbonate chemistry and benthic marine community during experiments, 2011

Ocean acidification is predicted to impact all areas of the oceans and affect a diversity of marine organisms. However, the diversity of responses among species prevents clear predictions about the impact of acidification at the ecosystem level. Here, we used shallow water CO2 vents in the Mediterranean Sea as a model system to examine emergent ecosystem responses to ocean acidification in rocky reef communities. We assessed in situ benthic invertebrate communities in three distinct pH zones (ambient, low, and extreme low), which differed in both the mean and variability of seawater pH along a continuous gradient. We found fewer taxa, reduced taxonomic evenness, and lower biomass in the extreme low pH zones. However, the number of individuals did not differ among pH zones, suggesting that there is density compensation through population blooms of small acidification-tolerant taxa. Furthermore, the trophic structure of the invertebrate community shifted to fewer trophic groups and dominance by generalists in extreme low pH, suggesting that there may be a simplification of food webs with ocean acidification. Despite high variation in individual species' responses, our findings indicate that ocean acidification decreases the diversity, biomass, and trophic complexity of benthic marine communities. These results suggest that a loss of biodiversity and ecosystem function is expected under extreme acidification scenarios.

Seawater carbonate chemistry, processes and elements during experiments with planktonic foraminifera Orbulina universa, 2004

We investigate the sensitivity of U/Ca, Mg/Ca, and Sr/Ca to changes in seawater [CO3[2-]] and temperature in calcite produced by the two planktonic foraminifera species, Orbulina universa and Globigerina bulloides, in laboratory culture experiments. Our results demonstrate that at constant temperature, U/Ca in O. universa decreases by 25 +/- 7% per 100 µmol [CO3[2-]] kg**-1, as seawater [CO3[2-]] increases from 110 to 470 µmol kg**-1. Results from G. bulloides suggest a similar relationship, but U/Ca is consistently offset by ~+40% at the same environmental [CO3[2-]]. In O. universa, U/Ca is insensitive to temperature between 15°C and 25°C. Applying the O. universa relationship to three U/Ca records from a related species, Globigerinoides sacculifer, we estimate that Caribbean and tropical Atlantic [CO3[2-]] was 110 +/- 70 µmol kg**-1 and 80 +/- 40 µmol kg**-1 higher, respectively, during the last glacial period relative to the Holocene. This result is consistent with estimates of the glacial-interglacial change in surface water [CO3[2-]] based on both modeling and on boron isotope pH estimates. In settings where the addition of U by diagenetic processes is not a factor, down-core records of foraminiferal U/Ca have potential to provide information about changes in the ocean's carbonate concentration.

Geochemistry of hydrothermal solutions, ores, and biota from hydrothermal fields at the East Pacific Rise axis near 9°50'N

Hydrothermal solutions were examined in a circulation system that started to develop after the 1991 volcanic eruption in the axial segment of the EPR between 9°45'N and 9°52'N. Within twelve years after this eruption, diffusion outflow of hot fluid from fractures in basaltic lavas gave way to focused seeps of hot solutions through channels of hydrothermal sulfide edifices. An example of the field Q demonstrates that from 1991 to 2003 H2S concentrations decreased from 86 to 1 mM/kg, and the Fe/H2S ratio simultaneously increased by factor 1.7. This fact can explain disappearance of microbial mats that were widespread within the fields before 1991. S isotopic composition of H2S does not depend on H2S concentration. This fact testifies rapid evolution of the hydrothermal system in the early years of its evolution. Carbon in CH4 from hot fluid sampled in 2003 is richer in 12C isotope than carbon in fluid from the hydrothermal field at 21°N EPR. It suggests that methane comes to the Q field from more than one source. Composition of particulate matter in hydrothermal solutions indicates that it was contributed by biological material. Experimental solutions with labeled substrates (t<70°C) show evidence of active processes of methane oxidation and sulfate reduction. Our results indicate that, during 12-year evolution of the hydrothermal system, composition of its solutions evolved and approached compositions of solutions in mature hydrothermal systems of the EPR.

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.

Short- versus long-term responses to changing CO2 in a coastal dinoflagellate bloom

Increasing pCO2 (partial pressure of CO2 ) in an "acidified" ocean will affect phytoplankton community structure, but manipulation experiments with assemblages briefly acclimated to simulated future conditions may not accurately predict the long-term evolutionary shifts that could affect inter-specific competitive success. We assessed community structure changes in a natural mixed dinoflagellate bloom incubated at three pCO2 levels (230, 433, and 765 ppm) in a short-term experiment (2 weeks). The four dominant species were then isolated from each treatment into clonal cultures, and maintained at all three pCO2 levels for approximately 1 year. Periodically (4, 8, and 12 months), these pCO2 -conditioned clones were recombined into artificial communities, and allowed to compete at their conditioning pCO2 level or at higher and lower levels. The dominant species in these artificial communities of CO2 -conditioned clones differed from those in the original short-term experiment, but individual species relative abundance trends across pCO2 treatments were often similar. Specific growth rates showed no strong evidence for fitness increases attributable to conditioning pCO2 level. Although pCO2 significantly structured our experimental communities, conditioning time and biotic interactions like mixotrophy also had major roles in determining competitive outcomes. New methods of carrying out extended mixed species experiments are needed to accurately predict future long-term phytoplankton community responses to changing pCO2 .

Mineralogy and interstitial-water chemistry of ODP Leg 101 sites

Concentrations of dissolved Ca2+, Sr2+, Mg2+, SO4[2-], and alkalinity were measured in pore waters squeezed from sediments taken from ODP Holes 626C and 626D in the Florida Straits; Holes 627A and 627B, 628A, and 630A and 630C north of Little Bahama Bank; Holes 631 A, 632A and 632B, and 633A in Exuma Sound; and Holes 634A and 635A and 635B in Northeast Providence Channel. These data are compared with the mineralogy and strontium content of the sediments from which the waters were squeezed. Contrasts in the geochemical profiles suggest that significantly different processes govern pore-water signatures at each group of sites. In Little Bahama Bank, strong positive Ca2+ gradients are correlated with weak negative Mg2+ profiles. These trends are analogous to those seen at DSDP sites where carbonate deposits immediately overlie mafic basement, but as the depth to basement may be in excess of 5000 m, we suggest that diffusion gradients are initiated by an underlying sedimentary unit. In contrast, Ca2+ and Mg2+ gradients in Exuma Sound are not developed to any appreciable extent over similar thicknesses of sediment. We suggest that the pore-water chemistry in these deposits is principally controlled by diagenetic reactions occurring within each sequence. The location and extent of carbonate diagenesis can be estimated from dissolved Sr2+ profiles. In Little Bahama Bank and Exuma Sound, Sr2+ concentrations reach a maximum value of between 700 and 1000 µmol/L. Although the depths at which these concentrations are achieved are different for the two areas, the corresponding age of the sediment at the dissolved Sr2+ maximum is similar. Consequently, the diffusive flux of Sr2+ and the calculated rates of recrystallization in the two areas are likewise of a similar magnitude. The rates of recrystallization we calculate are lower than those found in some DSDP pelagic sites. As the waters throughout most of the holes are saturated with respect to SrSO4, celestite precipitation may cause erroneously low Sr2+ production rates and, consequently, low calculated rates of recrystallization. We therefore encourage only the discriminate use of Sr2+ profiles in the quantification of diagenetic processes.

The effects of elevated CO2 on the growth and toxicity of field populations and cultures of the saxitoxin-producing dinoflagellate, Alexandrium fundyense

The effects of coastal acidification on the growth and toxicity of the saxitoxin-producing dinoflagellate Alexandrium fundyense were examined in culture and ecosystem studies. In culture experiments, Alexandrium strains isolated from Northport Bay, New York, and the Bay of Fundy, Canada, grew significantly faster (16-190%; p < 0.05) when exposed to elevated levels of PCO2 ( 90-190 Pa=900-1900 µatm) compared to lower levels ( 40 Pa=400 µatm). Exposure to higher levels of PCO2 also resulted in significant increases (71-81%) in total cellular toxicity (fg saxitoxin equivalents/cell) in the Northport Bay strain, while no changes in toxicity were detected in the Bay of Fundy strain. The positive relationship between PCO2 enrichment and elevated growth was reproducible in natural populations from New York waters. Alexandrium densities were significantly and consistently enhanced when natural populations were incubated at 150 Pa PCO2 compared to 39 Pa. During natural Alexandrium blooms in Northport Bay, PCO2 concentrations increased over the course of a bloom to more than 170 Pa and were highest in regions with the greatest Alexandrium abundances, suggesting Alexandrium may further exacerbate acidification and/or be especially adapted to these acidi-fied conditions. The co-occurrence of Alexandrium blooms and elevated PCO2 represents a previously unrecognized, compounding environmental threat to coastal ecosystems. The ability of elevated PCO2 to enhance the growth and toxicity of Alexandrium indicates that acidification promoted by eutrophication or climate change can intensify these, and perhaps other, harmful algal blooms.

Experimental ocean acidification alters the allocation of metabolic energy

Energy is required to maintain physiological homeostasis in response to environmental change. Although responses to environmental stressors frequently are assumed to involve high metabolic costs, the biochemical bases of actual energy demands are rarely quantified. We studied the impact of a near-future scenario of ocean acidification [800 µatm partial pressure of CO2 (pCO2)] during the development and growth of an important model organism in developmental and environmental biology, the sea urchin Strongylocentrotus purpuratus. Size, metabolic rate, biochemical content, and gene expression were not different in larvae growing under control and seawater acidification treatments. Measurements limited to those levels of biological analysis did not reveal the biochemical mechanisms of response to ocean acidification that occurred at the cellular level. In vivo rates of protein synthesis and ion transport increased 50% under acidification. Importantly, the in vivo physiological increases in ion transport were not predicted from total enzyme activity or gene expression. Under acidification, the increased rates of protein synthesis and ion transport that were sustained in growing larvae collectively accounted for the majority of available ATP (84%). In contrast, embryos and prefeeding and unfed larvae in control treatments allocated on average only 40% of ATP to these same two processes. Understanding the biochemical strategies for accommodating increases in metabolic energy demand and their biological limitations can serve as a quantitative basis for assessing sublethal effects of global change. Variation in the ability to allocate ATP differentially among essential functions may be a key basis of resilience to ocean acidification and other compounding environmental stressors.