Seawater carbonate chemistry and biological processes of Mytilus edulis during experiments, 2010

Marine organisms are exposed to increasingly acidic oceans, as a result of equilibration of surface ocean water with rising atmospheric CO2 concentrations. In this study, we examined the physiological response of Mytilus edulis from the Baltic Sea, grown for 2 months at 4 seawater pCO2 levels (39, 113, 243 and 405 Pa/385, 1,120, 2,400 and 4,000 µatm). Shell and somatic growth, calcification, oxygen consumption and excretion rates were measured in order to test the hypothesis whether exposure to elevated seawater pCO2 is causally related to metabolic depression. During the experimental period, mussel shell mass and shell-free dry mass (SFDM) increased at least by a factor of two and three, respectively. However, shell length and shell mass growth decreased linearly with increasing pCO2 by 6-20 and 10-34%, while SFDM growth was not significantly affected by hypercapnia. We observed a parabolic change in routine metabolic rates with increasing pCO2 and the highest rates (+60%) at 243 Pa. excretion rose linearly with increasing pCO2. Decreased O:N ratios at the highest seawater pCO2 indicate enhanced protein metabolism which may contribute to intracellular pH regulation. We suggest that reduced shell growth under severe acidification is not caused by (global) metabolic depression but is potentially due to synergistic effects of increased cellular energy demand and nitrogen loss.

Mass balance and acceleration of the Greenland ice sheet

Within the last decade, the Greenland ice sheet (GrIS) and its surroundings have experienced record high surface temperatures (Mote, 2007, doi:10.1029/2007GL031976; Box et al., 2010), ice sheet melt extent (Fettweis et al., 2011, doi:10.5194/tc-5-359-2011) and record-low summer sea-ice extent (Nghiem et al., 2007, doi:10.1029/2007GL031138). Using three independent data sets, we derive, for the first time, consistent ice-mass trends and temporal variations within seven major drainage basins from gravity fields from the Gravity Recovery and Climate Experiment (GRACE; Tapley et al., 2004, doi:10.1029/2004GL019920), surface-ice velocities from Inteferometric Synthetic Aperture Radar (InSAR; Rignot and Kanagaratnam, 2006, doi:10.1126/science.1121381) together with output of the regional atmospheric climate modelling (RACMO2/ GR; Ettema et al., 2009, doi:10.1029/2009GL038110), and surface-elevation changes from the Ice, cloud and land elevation satellite (ICESat; Sorensen et al., 2011, doi:10.5194/tc-5-173-2011). We show that changing ice discharge (D), surface melting and subsequent run-off (M/R) and precipitation (P) all contribute, in a complex and regionally variable interplay, to the increasingly negative mass balance of the GrIS observed within the last decade. Interannual variability in P along the northwest and west coasts of the GrIS largely explains the apparent regional mass loss increase during 2002-2010, and obscures increasing M/R and D since the 1990s. In winter 2002/2003 and 2008/2009, accumulation anomalies in the east and southeast temporarily outweighed the losses by M/R and D that prevailed during 2003-2008, and after summer 2010. Overall, for all basins of the GrIS, the decadal variability of anomalies in P, M/R and D between 1958 and 2010 (w.r.t. 1961-1990) was significantly exceeded by the regional trends observed during the GRACE period (2002-2011).

Accumulation rates, carbon composition and Eocene carbonate compensation depths of ODP Sites 199-1218 and 199-1219

CaCO3, Corg, and biogenic SiO2 were measured in Eocene equatorial Pacific sediments from Sites 1218 and 1219, and bulk oxygen and carbon isotopes were measured on selected intervals from Site 1219. These data delineate a series of CaCO3 events that first appeared at ~48 Ma and continued to the Eocene/Oligocene boundary. Each event lasted 1-2 m.y. and is separated from the next by a low CaCO3 interval of a similar time span. The largest of these carbonate accumulation events (CAE-3) is in Magnetochron 18. It began at ~42.2 Ma, lasted until ~40.3 Ma, and was marked by higher than average productivity. The end of CAE-3 was abrupt and was associated with a large-scale carbon transfer to the oceans prior to warming of high-latitude regions. Changes in carbonate compensation depth associated with CAE excursions were small in the early part of the middle Eocene but increased to as much as 800 m by the late middle Eocene before decreasing into the late Eocene. Oxygen isotope data indicate that the carbonate events are associated with cooling conditions and may mark small glaciations in the Eocene.

Ostracodes in bottom sediments of the Laptev Sea

This work is the first detailed description of the Late Pleistocene-Holocene and Recent Ostracoda of the Laptev Sea. A total of 45 species in 22 genera and 13 families have been identified. All these species are described monographically. Three different ecological assemblages of ostracodes corresponding to different combinations of environmental parameters have been established; they are restricted to three regions of the sea: western-central, eastern, and southern. The recent ostracode assemblages of the Laptev Sea have been compared with those from other Arctic areas and are most similar to those of the Beaufort and Kara seas. Data on recent Ostracoda are used for paleoenvironmental reconstructions on the eastern shelf and western continental slope of the Laptev Sea. For this purpose, ostracodes from five sections obtained from these parts of the sea have been examined. The oldest sediments, which are of Late Pleistocene age (15.8 cal. ka BP), have been recovered in a core from the western continental slope. These yielded five ostracode assemblages, which correspond to different paleoenvironments and replaced each other in the course of the rapid postglacial sea-level rise, thus showing variations in the Atlantic water inflow from the west and freshwater discharge from the subaerially exposed shelf. On the outer shelf of the eastern part of the sea, the rapid sea-level rise in the Early Holocene (lowermost dating 11.3 cal. ka BP) led to a rapid transition from assemblages of brackish-water nearshore environments to those of modernlike normal marine environments; modern environments were established about 8.2 cal. ka ago. Since core sections from the inner shelf correspond to the time when the level of the sea had already reached its modern values, changes in taxonomic composition of ostracode assemblages primarily mirror variations in river runoff.

Seawater carbonate chemistry and physiological and mechanical properties of the starfish Asterias rubens in a laboratory experiment

The increase in atmospheric CO2 due to anthropogenic activity results in an acidification of the surface waters of the oceans. Its impact will depend on the considered organisms and ecosystems. The intertidal may harbor organisms pre-adapted to the upcoming changes as they face tidal pH and temperature fluctuations. However, these environments will be more affected as shallow waters will face the highest decrease in seawater pH. In this context, the effects of reduced environmental pH on the physiology and tube feet mechanical properties of the intertidal starfish Asterias rubens, a top predator, were investigated during 15 and 27 days. A. rubens showed a respiratory acidosis with its coelomic fluid pH always lower than that of seawater. This acidosis was most pronounced at pH 7.4. Notwithstanding, the starfish showed no significant variations in RNA/DNA ratio of different tissues and in tube feet strength. However, respiration rates were significantly lower for individuals maintained at reduced seawater pH. Within the ocean acidification context, the present results suggest that A. rubens withstands the effects of reduced seawater pH, at least for medium term exposures.

(Appendix 1-4) Geochemistry, desorption methods, TOC, TC, and mineral content of different settings

Sorption of volatile hydrocarbon gases (VHCs) to marine sediments is a recognized phenomenon that has been investigated in the context of petroleum exploration. However, little is known about the biogeochemistry of sorbed methane and higher VHCs in environments that are not influenced by thermogenic processes. This study evaluated two different extraction protocols for sorbed VHCs, used high pressure equipment to investigate the sorption of methane to pure clay mineral phases, and conducted a geochemical and mineralogical survey of sediment samples from different oceanographic settings and geochemical regimes that are not significantly influenced by thermogenic gas. Extraction of sediments under alkaline conditions yielded higher concentrations of sorbed methane than the established protocol for acidic extraction. Application of alkaline extraction in the environmental survey revealed the presence of substantial amounts of sorbed methane in 374 out of 411 samples (91%). Particularly high amounts, up to 2.1 mmol kg**-1 dry sediment, were recovered from methanogenic sediments. Carbon isotopic compositions of sorbed methane suggested substantial contributions from biogenic sources, both in sulfate-depleted and sulfate-reducing sediments. Carbon isotopic relationships between sorbed and dissolved methane indicate a coupling of the two pools. While our sorption experiments and extraction conditions point to an important role for clay minerals as sorbents, mineralogical analyses of marine sediments suggest that variations in mineral composition are not controlling variations in quantities of sorbed methane. We conclude that the distribution of sorbed methane in sediments is strongly influenced by in situ production.

Pollen analyses of sediment core GeoB9508-5

Millennial-scale dry events in the Northern Hemisphere monsoon regions during the last Glacial period are commonly attributed to southward shifts of the Intertropical Convergence Zone (ITCZ) associated with an intensification of the northeasterly (NE) trade wind system during intervals of reduced Atlantic meridional overturning circulation (AMOC). Through the use of high-resolution last deglaciation pollen records from the continental slope off Senegal, our data show that one of the longest and most extreme droughts in the western Sahel history, which occurred during the North Atlantic Heinrich Stadial 1 (HS1), displayed a succession of three major phases. These phases progressed from an interval of maximum pollen representation of Saharan elements between ~19 and 17.4 kyr BP indicating the onset of aridity and intensified NE trade winds, followed by a millennial interlude of reduced input of Saharan pollen and increased input of Sahelian pollen, to a final phase between ~16.2 and 15 kyr BP that was characterized by a second maximum of Saharan pollen abundances. This change in the pollen assemblage indicates a mid-HS1 interlude of NE trade wind relaxation, occurring between two distinct trade wind maxima, along with an intensified mid-tropospheric African Easterly Jet (AEJ) indicating a substantial change in West African atmospheric processes. The pollen data thus suggest that although the NE trades have weakened, the Sahel drought remained severe during this time interval. Therefore, a simple strengthening of trade winds and a southward shift of the West African monsoon trough alone cannot fully explain millennial-scale Sahel droughts during periods of AMOC weakening. Instead, we suggest that an intensification of the AEJ is needed to explain the persistence of the drought during HS1. Simulations with the Community Climate System Model indicate that an intensified AEJ during periods of reduced AMOC affected the North African climate by enhancing moisture divergence over the West African realm, thereby extending the Sahel drought for about 4000 years.

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.

One size fits all: stability of metabolic scaling under warming and ocean acidification in echinoderms

Responses by marine species to ocean acidification (OA) have recently been shown to be modulated by external factors including temperature, food supply and salinity. However the role of a fundamental biological parameter relevant to all organisms, that of body size, in governing responses to multiple stressors has been almost entirely overlooked. Recent consensus suggests allometric scaling of metabolism with body size differs between species, the commonly cited 'universal' mass scaling exponent (b) of ¾ representing an average of exponents that naturally vary. One model, the Metabolic-Level Boundaries hypothesis, provides a testable prediction: that b will decrease within species under increasing temperature. However, no previous studies have examined how metabolic scaling may be directly affected by OA. We acclimated a wide body-mass range of three common NE Atlantic echinoderms (the sea star Asterias rubens, the brittlestars Ophiothrix fragilis and Amphiura filiformis) to two levels of pCO2 and three temperatures, and metabolic rates were determined using closed-chamber respirometry. The results show that contrary to some models these echinoderm species possess a notable degree of stability in metabolic scaling under different abiotic conditions; the mass scaling exponent (b) varied in value between species, but not within species under different conditions. Additionally, we found no effect of OA on metabolic rates in any species. These data suggest responses to abiotic stressors are not modulated by body size in these species, as reflected in the stability of the metabolic scaling relationship. Such equivalence in response across ontogenetic size ranges has important implications for the stability of ecological food webs.

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.

Tolerance of juvenile barnacles (Amphibalanus improvisus) to warming and elevated pCO2

We investigated the impacts of warming and elevated pCO2 on newly settled Amphibalanus improvisus from Kiel Fjord, an estuarine ecosystem characterized by significant natural pCO2 variability. In two experiments, juvenile barnacles were maintained at two temperature and three pCO2 levels (20/24°C, 700-2.140 µatm) for 8 weeks in a batch culture and at four pCO2 levels (20°C, 620-2.870 µatm) for 12 weeks in a water flow-through system. Warming as well as elevated pCO2 hardly affected growth or the condition index of barnacles, although some factor combinations led to temporal significances in enhanced or reduced growth with an increase in pCO2. While warming increased the shell strength of A. improvisus individuals, elevated pCO2 had only weak effects. We demonstrate a strong tolerance of juvenile A. improvisus to mean acidification levels of about 1,000 µatm pCO2 as is already naturally experienced by the investigated barnacle population.

Effects of ocean acidification on respiration, growth and C:N ratio of arctic Hyas araneus larvae

Early life stages of marine crustaceans respond sensitively to elevated seawater PCO2. However, the underlying physiological mechanisms have not been studied well. We therefore investigated the effects of elevated seawater PCO2 on oxygen consumption, dry weight, elemental composition, median developmental time (MDT) and mortality in zoea I larvae of the spider crab Hyas araneus (Svalbard 79°N/11°E; collection, May 2009; hatch, December 2009). At the time of moulting, oxygen consumption rate had reached a steady state level under control conditions. In contrast, elevated seawater PCO2 caused the metabolic rate to rise continuously leading to a maximum 1.5-fold increase beyond control level a few days before moulting into the second stage (zoea II), followed by a pronounced decrease. Dry weight of larvae reared under high CO2 conditions was lower than in control larvae at the beginning of the moult cycle, yet this difference had disappeared at the time of moulting. MDT of zoea I varied between 45 ± 1 days under control conditions and 42 ± 2 days under the highest seawater CO2 concentration. The present study indicates that larval development under elevated seawater PCO2 levels results in higher metabolic costs during premoulting events in zoea I. However, H. araneus zoea I larvae seem to be able to compensate for higher metabolic costs as larval MDT and survival was not affected by elevated PCO2 levels.

Seawater carbonate chemistry, clearance rates, respiration rates, condition index and cellular turnover (RNA: DNA) of Juvenile King Scallop, Pecten maximus in a laboratory experiment

The decline in ocean water pH and changes in carbonate saturation states through anthropogenically mediated increases in atmospheric CO2 levels may pose a hazard to marine organisms. This may be particularly acute for those species reliant on calcareous structures like shells and exoskeletons. This is of particular concern in the case of valuable commercially exploited species such as the king scallop, Pecten maximus. In this study we investigated the effects on oxygen consumption, clearance rates and cellular turnover in juvenile P. maximus following 3 months laboratory exposure to four pCO2 treatments (290, 380, 750 and 1140 µatm). None of the exposure levels were found to have significant effect on the clearance rates, respiration rates, condition index or cellular turnover (RNA: DNA) of individuals. While it is clear that some life stages of marine bivalves appear susceptible to future levels of ocean acidification, particularly under food limiting conditions, the results from this study suggest that where food is in abundance, bivalves like juvenile P. maximus may display a tolerance to limited changes in seawater chemistry.

Fossil chironomids of Lake Temje

A 380 cm long sediment core from Lake Temje (central Yakutia, Eastern Siberia) was studied to infer Holocene palaeoenvironmental change in the extreme periglacial setting of eastern Siberia during the last 10,000 years. Data on sediment composition were used to characterize changes in the depositional environment during the ontogenetic development of the Lake Temje. The analysis of fossil chironomid remains and statistical treatment of chironomid data by the application of a newly developed regional Russian transfer functions provided inferences of mean July air temperatures (T_July) and water depths (WD). Reconstructed WDs show minor changes throughout the core and range between 80 and 120 cm. All the fluctuations in reconstructed water depth lie within the mean error of prediction of the inference model (RMSEP = 0.35) so it is not possible to draw conclusions from the reconstructions. A qualitative and quantitative reconstruction of Holocene climate in central Yakutia recognized three stages of palaeoenvironmental changes. The early Holocene between 10 and 8 ka BP was characterized by colder-than-today and moist summer conditions. Cryotextures in the lake sediments document full freezing of the lake water during the winter time. A general warming trend started around 8.0 ka BP in concert with enhanced biological productivity. Reconstructed mean T_July were equal or up to 1.5 °C higher than today between 6.0 ka and 5.0 ka BP. During the entire late Holocene after 4.8 ka BP, reconstructed mean T_July remained below modern value. Limnological conditions did not change significantly. The inference of a mid-Holocene climate optimum supports scenarios of Holocene climatic changes in the subpolar part of eastern Siberia and indicates climate teleconnections to the North Atlantic realm.

Size matters: plasticity in metabolic scaling shows body-size may modulate responses to climate change

Variability in metabolic scaling in animals, the relationship between metabolic rate ( R) and body mass ( M), has been a source of debate and controversy for decades. R is proportional to Mb, the precise value of b much debated, but historically considered equal in all organisms. Recent metabolic theory, however, predicts b to vary among species with ecology and metabolic level, and may also vary within species under different abiotic conditions. Under climate change, most species will experience increased temperatures, and marine organisms will experience the additional stressor of decreased seawater pH ('ocean acidification'). Responses to these environmental changes are modulated by myriad species-specific factors. Body-size is a fundamental biological parameter, but its modulating role is relatively unexplored. Here, we show that changes to metabolic scaling reveal asymmetric responses to stressors across body-size ranges; b is systematically decreased under increasing temperature in three grazing molluscs, indicating smaller individuals were more responsive to warming. Larger individuals were, however, more responsive to reduced seawater pH in low temperatures. These alterations to the allometry of metabolism highlight abiotic control of metabolic scaling, and indicate that responses to climate warming and ocean acidification may be modulated by body-size.