Sterol, n-alcohol and n-alkane composition of living fen plants

In groundwater-fed fen peatlands, the surface biomass decays rapidly and, as a result, highly humified peat is formed. A high degree of humification constrains palaeoecological studies because reliable identification of plant remains is hampered. Organic geochemistry techniques as a means of identifying historical plant communities have been successfully applied tobog peat. The method has also been applied to fen peat, but without reference to the composition of fen plants. We have applied selected organic geochemistry methods to determine the composition of the neutral lipid fractions from 12 living fen plants, to investigate the potential for the distributions to characterize and separate different fen plants and plant groups. Our results show correspondence with previous studies, e.g. C23 and C25n-alkanes dominating Sphagnum spp. and C27 to C31 alkanes dominating vascular plants. However, we also found similarities in n-alkane distributions between Sphagnum spp. and the below ground parts of some vascular plants. We tested the efficiency of different n-alkane ratios to separate species and plant groups. The ratios used for bog studies (e.g. n-C23/n-C25 and n-C23/n-C29) did not work as consistently for fen plants. Some differences in sterol distribution were found between vascular plants and mosses; in general vascular plants had a higher concentration of sterols. When distributions of n-alkanes, n-alkane ratios and sterols were all included as variables, redundancy analysis (RDA) separated different plant groups into their own clusters. Our results imply that the pattern for bog biomarkers cannot directly be applied to fen environments. Nevertheless, they encourage further testing to determine whether or not the identification of plant groups, plants or plant parts from highly humified peat is possible by applying fen species-specific biomarker proxies.

Seasonal changes in the D/H ratio of fatty acids of pelagic microorganisms

Culture studies of microorganisms have shown that the hydrogen isotopic composition of fatty acids depends on their metabolism, but there are only few environmental studies available to confirm this observation. Here we studied the seasonal variability of the deuterium/hydrogen (D/H) ratio of fatty acids in the coastal Dutch North Sea and compared this with the diversity of the phyto- and bacterioplankton. Over the year, the stable hydrogen isotopic fractionation factor epsilon between fatty acids and water ranged between -172 per mil and -237 per mil, the algal-derived polyunsaturated fatty acid nC20:5 being the most D-depleted and nC18:0 the least D-depleted fatty acid. The D-depleted nC20:5 is in agreement with culture studies, which indicates that photoautotrophic microorganisms produce fatty acids which are significantly depleted in D relative to water. The epsilon-lipid/water of all fatty acids showed a transient shift towards increased fractionation during the spring phytoplankton bloom, indicated by increasing chlorophyll a concentrations and relative abundance of the nC20:5 PUFA, suggesting increased contributions of photoautotrophy. Time periods with decreased fractionation (less negative epsilon-lipid/water values) can be explained by an increased contribution by heterotrophy to the fatty acid pool. Our results show that the hydrogen isotopic composition of fatty acids is a useful tool to assess the community metabolism of coastal plankton.

Planktic and benthic 14C reservoir ages, calibrated by a suite of 14C plateaus in the glacial-to-deglacial Suigetsu atmospheric 14C record

We here present a compilation of planktic and benthic 14C reservoir ages for the Last Glacial Maximum (LGM) and early deglacial from 11 key sites of global ocean circulation in the Atlantic and Indo-Pacific Ocean. The ages were obtained by 14C plateau tuning, a robust technique to derive both an absolute chronology for marine sediment records and a high-resolution record of changing reservoir/ventilation ages (Delta14C values) for surface and deep waters by comparing the suite of planktic 14C plateaus of a sediment record with that of the atmospheric 14C record (Sarnthein et al., 2007, doi:10.1029/173GM13). Results published thus far used as atmospheric 14C reference U/Th-dated corals, the Cariaco planktic record, and speleothems (Fairbanks et al., 2005, doi:10.1016/j.quascirev.2005.04.007; Hughen et al., 2006, doi:10.1016/j.quascirev.2006.03.014; Beck et al., 2001, doi:10.1023/A:1008175728826). We have now used the varve-counted atmospheric 14C record of Lake Suigetsu terrestrial macrofossils (Ramsey et al., 2012, doi:10.1126/science.1226660) to recalibrate the boundary ages and reservoir ages of the seven published records directly to an atmospheric 14C record. In addition, the results for four new cores and further planktic results for four published records are given. Main conclusions from the new compilation are: (1) The Suigetsu atmospheric 14C record on its varve counted time scale reflects all 14C plateaus, their internal structures and relative length previously identified, but implies a rise in the average 14C plateau age by 200-700 14C yr during LGM and early deglacial times. (2) Based on different 14C ages of coeval atmospheric and planktic 14C plateaus, marine surface water Delta14C may have temporarily dropped to an equivalent of ~0 yr in low-latitude lagoon waters, but reached >2500 14C yr both in stratified subpolar waters and in upwelled waters such as in the South China Sea. These values differ significantly from a widely assumed constant global planktic Delta14C value of 400 yr. (3) Suites of deglacial planktic Delta14C values are closely reproducible in 14C records measured at neighboring core sites. (4) Apparent deep-water 14C ventilation ages (equivalents of benthic Delta14C), deduced from the sum of planktic Delta14C and coeval benthic-planktic 14C differences, vary from 500 up to >5000 yr in LGM and deglacial ocean basins.

Seawater carbonate chemistry, biomass and calcification of Porites spp. corals during experiments, 2011

I tested the hypothesis that the effects of high pCO2 and temperature on massive Porites spp. (Scleractinia) are modified by heterotrophic feeding (zooplanktivory). Small colonies of massive Porites spp. from the back reef of Moorea, French Polynesia, were incubated for 1 month under combinations of temperature (29.3°C vs. 25.6°C), pCO2 (41.6 vs. 81.5 Pa), and feeding regimes (none vs. ad libitum access to live Artemia spp.), with the response assessed using calcification and biomass. Area-normalized calcification was unaffected by pCO2, temperature, and the interaction between the two, although it increased 40% with feeding. Biomass increased 35% with feeding and tended to be higher at 25.6°C compared to 29.3°C, and as a result, biomass-normalized calcification statistically was unaffected by feeding, but was depressed 12-17% by high pCO2, with the effect accentuated at 25.6°C. These results show that massive Porites spp. has the capacity to resist the effects on calcification of 1 month exposure to 81.5 Pa pCO2 through heterotrophy and changes in biomass. Area-normalized calcification is sustained at high pCO2 by a greater biomass with a reduced biomass-normalized rate of calcification. This mechanism may play a role in determining the extent to which corals can resist the long-term effects of ocean acidification.

Effects of increased CO2 on fish gill and plasma proteome

Ocean acidification and warming are both primarily caused by increased levels of atmospheric CO2, and marine organisms are exposed to these two stressors simultaneously. Although the effects of temperature on fish have been investigated over the last century, the long-term effects of moderate CO2 exposure and the combination of both stressors are almost entirely unknown. A proteomics approach was used to assess the adverse physiological and biochemical changes that may occur from the exposure to these two environmental stressors. We analysed gills and blood plasma of Atlantic halibut (Hippoglossus hippoglossus) exposed to temperatures of 12°C (control) and 18°C (impaired growth) in combination with control (400 µatm) or high-CO2 water (1000 µatm) for 14 weeks. The proteomic analysis was performed using two-dimensional gel electrophoresis (2DE) followed by Nanoflow LC-MS/MS using a LTQ-Orbitrap. The high-CO2 treatment induced the up-regulation of immune system-related proteins, as indicated by the up-regulation of the plasma proteins complement component C3 and fibrinogen beta chain precursor in both temperature treatments. Changes in gill proteome in the high-CO2 (18°C) group were mostly related to increased energy metabolism proteins (ATP synthase, malate dehydrogenase, malate dehydrogenase thermostable, and fructose-1,6-bisphosphate aldolase), possibly coupled to a higher energy demand. Gills from fish exposed to high-CO2 at both temperature treatments showed changes in proteins associated with increased cellular turnover and apoptosis signalling (annexin 5, eukaryotic translation elongation factor 1 gamma, receptor for protein kinase C, and putative ribosomal protein S27). This study indicates that moderate CO2-driven acidification, alone and combined with high temperature, can elicit biochemical changes that may affect fish health.

1H NMR Metabolomics Reveals Contrasting Response by Male and Female Mussels Exposed to Reduced Seawater pH, Increased Temperature, and a Pathogen

Human activities are fundamentally altering the chemistry of the world's oceans. Ocean acidification (OA) is occurring against a background of warming and an increasing occurrence of disease outbreaks, posing a significant threat to marine organisms, communities, and ecosystems. In the current study, 1H NMR spectroscopy was used to investigate the response of the blue mussel, Mytilus edulis, to a 90-day exposure to reduced seawater pH and increased temperature, followed by a subsequent pathogenic challenge. Analysis of the metabolome revealed significant differences between male and female organisms. Furthermore, males and females are shown to respond differently to environmental stress. While males were significantly affected by reduced seawater pH, increased temperature, and a bacterial challenge, it was only a reduction in seawater pH that impacted females. Despite impacting males and females differently, stressors seem to act via a generalized stress response impacting both energy metabolism and osmotic balance in both sexes. This study therefore has important implications for the interpretation of metabolomic data in mussels, as well as the impact of environmental stress in marine invertebrates in general.

Effects of ocean acidification on sponge communities

The effects of ocean acidification on lower invertebrates such as sponges may be pronounced because of their low capacity for acid-base regulation. However, so far, most studies have focused on calcifiers. We present the first study of the effects of ocean acidification on the Porifera. Sponge species composition and cover along pH gradients at CO2 vents off Ischia (Tyrrhenian Sea, Italy) was measured at sites with normal pH (8.1-8.2), lowered pH (mean 7.8-7.9, min 7.4-7.5) and extremely low pH (6.6). There was a strong correlation between pH and both sponge cover and species composition. Crambe crambe was the only species present in any abundance in the areas with mean pH 6.6, seven species were present at mean pH 7.8-7.9 and four species (Phorbas tenacior, Petrosia ficiformis, Chondrilla nucula and Hemimycale columella) were restricted to sites with normal pH. Sponge percentage cover decreased significantly from normal to acidified sites. No significant effect of increasing CO2 levels and decreasing pH was found on spicule form in Crambe crambe. This study indicates that increasing CO2 concentrations will likely affect sponge community composition as some demosponge species appear to be more vulnerable than others. Further research into the mechanisms by which acidification affects sponges would be useful in predicting likely effects on sessile marine communities.

Selective mortality associated with variation in CO2 tolerance in a marine fish

Predicted future CO2 levels can affect reproduction, growth, and behaviour of many marine organisms. However, the capacity of species to adapt to predicted changes in ocean chemistry is largely unknown. We used a unique field-based experiment to test for differential survival associated with variation in CO2 tolerance in a wild population of coral-reef fishes. Juvenile damselfish exhibited variation in their response to elevated (700 µatm) CO2 when tested in the laboratory and this influenced their behaviour and risk of mortality in the wild. Individuals that were sensitive to elevated CO2 were more active and move further from shelter in natural coral reef habitat and, as a result, mortality from predation was significantly higher compared with individuals from the same treatment that were tolerant of elevated CO2. If individual variation in CO2 tolerance is heritable, this selection of phenotypes tolerant to elevated CO2 could potentially help mitigate the effects of ocean acidification.

Seawater carbonate chemistry, nitrogen concentration and macro community analyse, 2011

Rising anthropogenic CO2 emissions acidify the oceans, and cause changes to seawater carbon chemistry. Bacterial biofilm communities reflect environmental disturbances and may rapidly respond to ocean acidification. This study investigates community composition and activity responses to experimental ocean acidification in biofilms from the Australian Great Barrier Reef. Natural biofilms grown on glass slides were exposed for 11 d to four controlled pCO2 concentrations representing the following scenarios: A) pre-industrial (~300 ppm), B) present-day (~400 ppm), C) mid century (~560 ppm) and D) late century (~1140 ppm). Terminal restriction fragment length polymorphism and clone library analyses of 16S rRNA genes revealed CO2-correlated bacterial community shifts between treatments A, B and D. Observed bacterial community shifts were driven by decreases in the relative abundance of Alphaproteobacteria and increases of Flavobacteriales (Bacteroidetes) at increased CO2 concentrations, indicating pH sensitivity of specific bacterial groups. Elevated pCO2 (C + D) shifted biofilm algal communities and significantly increased C and N contents, yet O2 fluxes, measured using in light and dark incubations, remained unchanged. Our findings suggest that bacterial biofilm communities rapidly adapt and reorganize in response to high pCO2 to maintain activity such as oxygen production.

Effects of CO2 enrichment on photosynthesis, growth, and nitrogen metabolism of the seagrass Zostera noltii

Seagrass ecosystems are expected to benefit from the global increase in CO2 in the ocean because the photosynthetic rate of these plants may be Ci-limited at the current CO2 level. As well, it is expected that lower external pH will facilitate the nitrate uptake of seagrasses if nitrate is cotransported with H+ across the membrane as in terrestrial plants. Here, we investigate the effects of CO2 enrichment on both carbon and nitrogen metabolism of the seagrass Zostera noltii in a mesocosm experiment where plants were exposed for 5 months to two experimental CO2 concentrations (360 and 700 ppm). Both the maximum photosynthetic rate (Pm) and photosynthetic efficiency (a) were higher (1.3- and 4.1-fold, respectively) in plants exposed to CO2-enriched conditions. On the other hand, no significant effects of CO2 enrichment on leaf growth rates were observed, probably due to nitrogen limitation as revealed by the low nitrogen content of leaves. The leaf ammonium uptake rate and glutamine synthetase activity were not significantly affected by increased CO2 concentrations. On the other hand, the leaf nitrate uptake rate of plants exposed to CO2-enriched conditions was fourfold lower than the uptake of plants exposed to current CO2 level, suggesting that in the seagrass Z. noltii nitrate is not cotransported with H+ as in terrestrial plants. In contrast, the activity of nitrate reductase was threefold higher in plant leaves grown at high-CO2 concentrations. Our results suggest that the global effects of CO2 on seagrass production may be spatially heterogeneous and depend on the specific nitrogen availability of each system. Under a CO2 increase scenario, the natural levels of nutrients will probably become limiting for Z. noltii. This potential limitation becomes more relevant because the expected positive effect of CO2 increase on nitrate uptake rate was not confirmed.

Seawater carbonate chemistry, biomass and metabolic rates (leucine incorporation, CO2 fixation and respiration) of Rhodobacteraceae (strain MED165) and Flavobacteriaceae (strain MED217) in a laboratory experiment

Experimental results related to the effects of ocean acidification on planktonic marine microbes are still rather inconsistent and occasionally contradictory. Moreover, laboratory or field experiments that address the effects of changes in CO2 concentrations on heterotrophic microbes are very scarce, despite the major role of these organisms in the marine carbon cycle. We tested the direct effect of an elevated CO2 concentration (1000 ppmv) on the biomass and metabolic rates (leucine incorporation, CO2 fixation and respiration) of 2 isolates belonging to 2 relevant marine bacterial families, Rhodobacteraceae (strain MED165) and Flavobacteriaceae (strain MED217). Our results demonstrate that, contrary to some expectations, high pCO2 did not negatively affect bacterial growth but increased growth efficiency in the case of MED217. The elevated partial pressure of CO2 (pCO2) caused, in both cases, higher rates of CO2 fixation in the dissolved fraction and, in the case of MED217, lower respiration rates. Both responses would tend to increase the pH of seawater acting as a negative feedback between elevated atmospheric CO2 concentrations and ocean acidification.