ALBERT

Description: Climate change has significant implications for biodiversity and ecosystems. With slow progress towards reducing greenhouse gas emissions, climate engineering (or ‘geoengineering’) is receiving increasing attention for its potential to limit anthropogenic climate change and its damaging effects. Proposed techniques, such as ocean fertilization for carbon dioxide removal or stratospheric sulfate injections to reduce incoming solar radiation, would significantly alter atmospheric, terrestrial and marine environments, yet potential side-effects of their implementation for ecosystems and biodiversity have received little attention. A literature review was carried out to identify details of the potential ecological effects of climate engineering techniques. A group of biodiversity and environmental change researchers then employed a modified Delphi expert consultation technique to evaluate this evidence and prioritize the effects based on the relative importance of, and scientific understanding about, their biodiversity and ecosystem consequences. The key issues and knowledge gaps are used to shape a discussion of the biodiversity and ecosystem implications of climate engineering, including novel climatic conditions, alterations to marine systems and substantial terrestrial habitat change. This review highlights several current research priorities in which the climate engineering context is crucial to consider, as well as identifying some novel topics for ecological investigation.

Description: Marine teleost fish sustain compensation of extracellular pH after exposure to hypercapnia by means of efficient ion and acid-base regulation. Elevated rates of ion and acid-base regulation under hypercapnia may be stimulated further by elevated temperature. Here, we characterized the regulation of transepithelial ion transporters (NKCC1, NBC1, SLC26A6, NHE1 and 2) and ATPases (Na(+)/K(+) ATPase and V-type H(+) ATPase) in gills of Atlantic cod (Gadus morhua) after 4 weeks of exposure to ambient and future PCO2 levels (550 μatm, 1200 μatm, 2200 μatm) at optimum (10 °C) and summer maximum temperature (18 °C), respectively. Gene expression of most branchial ion transporters revealed temperature- and dose-dependent responses to elevated PCO2. Transcriptional regulation resulted in stable protein expression at 10 °C, whereas expression of most transport proteins increased at medium PCO2 and 18 °C. mRNA and protein expression of distinct ion transport proteins were closely co-regulated, substantiating cellular functional relationships. Na(+)/K(+) ATPase capacities were PCO2 independent, but increased with acclimation temperature, whereas H(+) ATPase capacities were thermally compensated but decreased at medium PCO2 and 10 °C. When functional capacities of branchial ATPases were compared with mitochondrial F1Fo ATP-synthase strong correlations of F1Fo ATP-synthase and ATPase capacities generally indicate close coordination of branchial aerobic ATP demand and supply. Our data indicate physiological plasticity in the gills of cod to adjust to a warming, acidifying ocean within limits. In light of the interacting and non-linear, dose-dependent effects of both climate factors the role of these mechanisms in shaping resilience under climate change remains to be explored.

Description: Ocean acidification and associated changes in seawater carbonate chemistry negatively influence calcification processes and depress metabolism in many calcifying marine invertebrates. We present data on the cephalopod mollusc Sepia officinalis, an invertebrate that is capable of not only maintaining calcification, but also growth rates and metabolism when exposed to elevated partial pressures of carbon dioxide (pCO(2)). During a 6 wk period, juvenile S. officinalis maintained calcification under similar to 4000 and similar to 6000 ppm CO2, and grew at the same rate with the same gross growth efficiency as did control animals. They gained approximately 4%, body mass daily and increased the mass of their calcified cuttlebone by over 500 %. We conclude that active cephalopods possess a certain level of pre-adaptation to long-term increments in carbon dioxide levels. Our general understanding of the mechanistic processes that limit calcification must improve before we can begin to predict what effects future ocean acidification will have on calcifying marine invertebrates.

Description: Changes in seawater carbonate chemistry that accompany ongoing ocean acidification have been found to affect calcification processes in many marine invertebrates. In contrast to the response of most invertebrates, calcification rates increase in the cephalopod Sepia officinalis during long-term exposure to elevated seawater pCO2. The present trial investigated structural changes in the cuttlebones of S. officinalis calcified during 6 weeks of exposure to 615 Pa CO2. Cuttlebone mass increased sevenfold over the course of the growth trail, reaching a mean value of 0.71 ± 0.15 g. Depending on cuttlefish size (mantle lengths 44–56 mm), cuttlebones of CO2-incubated individuals accreted 22–55% more CaCO3 compared to controls at 64 Pa CO2. However, the height of the CO2-exposed cuttlebones was reduced. A decrease in spacing of the cuttlebone lamellae, from 384 ± 26 to 195 ± 38 μm, accounted for the height reduction The greater CaCO3 content of the CO2-incubated cuttlebones can be attributed to an increase in thickness of the lamellar and pillar walls. Particularly, pillar thickness increased from 2.6 ± 0.6 to 4.9 ± 2.2 μm. Interestingly, the incorporation of non-acid-soluble organic matrix (chitin) in the cuttlebones of CO2-exposed individuals was reduced by 30% on average. The apparent robustness of calcification processes in S. officinalis, and other powerful ion regulators such as decapod cructaceans, during exposure to elevated pCO2 is predicated to be closely connected to the increased extracellular [HCO3 −] maintained by these organisms to compensate extracellular pH. The potential negative impact of increased calcification in the cuttlebone of S. officinalis is discussed with regard to its function as a lightweight and highly porous buoyancy regulation device. Further studies working with lower seawater pCO2 values are necessary to evaluate if the observed phenomenon is of ecological relevance.

Description: Climate change with increasing temperature and ocean acidification (OA) poses risks for marine ecosystems. According to Portner and Farrell [1], synergistic effects of elevated temperature and CO(2)-induced OA on energy metabolism will narrow the thermal tolerance window of marine ectothermal animals. To test this hypothesis, we investigated the effect of an acute temperature rise on energy metabolism of the oyster, Crassostrea gigas chronically exposed to elevated CO(2) levels (partial pressure of CO(2) in the seawater similar to 0.15 kPa, seawater pH similar to 7.7). Within one month of incubation at elevated PCO(2) and 15 degrees C hemolymph pH fell (pH(e) = 7.1 +/- 0.2 (CO(2)-group) vs. 7.6 +/- 0.1 (control)) and P(e)CO(2) values in hemolymph increased (0.5 +/- 0.2 kPa (CO(2)-group) vs. 0.2 +/- 0.04 kPa (control)). Slightly but significantly elevated bicarbonate concentrations in the hemolymph of CO(2)-incubated oysters ([HCO(3)(-)](e) = 1.8 +/- 0.3 mM (CO(2)-group) vs. 1.3 +/- 0.1 mM (control)) indicate only minimal regulation of extracellular acid-base status. At the acclimation temperature of 15 degrees C the OA-induced decrease in pHe did not lead to metabolic depression in oysters as standard metabolism rates (SMR) of CO(2)-exposed oysters were similar to controls. Upon acute warming SMR rose in both groups, but displayed a stronger increase in the CO(2)-incubated group. Investigation in isolated gill cells revealed a similar temperature-dependence of respiration between groups. Furthermore, the fraction of cellular energy demand for ion regulation via Na(+)/K(+)-ATPase was not affected by chronic hypercapnia or temperature. Metabolic profiling using (1)H-NMR spectroscopy revealed substantial changes in some tissues following OA exposure at 15 degrees C. In mantle tissue alanine and ATP levels decreased significantly whereas an increase in succinate levels was observed in gill tissue. These findings suggest shifts in metabolic pathways following OA-exposure. Our study confirms that OA affects energy metabolism in oysters and suggests that climate change may affect populations of sessile coastal invertebrates such as mollusks.

Description: The cuttlefish Sepia officinalis expresses several hemocyanin isoforms with potentially different pH optima, indicating their reliance on efficient pH regulation in the blood. Ongoing ocean warming and acidification could influence the oxygen-binding properties of respiratory pigments in ectothermic marine invertebrates. This study examined whether S. officinalis differentially expresses individual hemocyanin isoforms to maintain optimal oxygen transport during development and acclimation to elevated seawater pCO2 and temperature. Using quantitative PCR, we measured relative mRNA expression levels of three different hemocyanin isoforms in several ontogenetic stages (embryos, hatchlings, juveniles, and adults), under different temperatures and elevated seawater pCO2. Our results indicate moderately altered hemocyanin expression in all embryonic stages acclimated to higher pCO2, while hemocyanin expression in hatchlings and juveniles remained unaffected. During the course of development, total hemocyanin expression increased independently of pCO2 or thermal acclimation status. Expression of isoform 3 is reported for the first time in a cephalopod in this study and was found to be generally low but highest in the embryonic stages (0.2% of total expression). Despite variable hemocyanin expression, hemolymph total protein concentrations remained constant in the experimental groups. Our data provide first evidence that ontogeny has a stronger influence on hemocyanin isoform expression than the environmental conditions chosen, and they suggest that hemocyanin protein abundance in response to thermal acclimation is regulated by post-transcriptional/translational rather than by transcriptional modifications

Description: The ongoing process of ocean acidification already affects marine life, and according to the concept of oxygen and capacity limitation of thermal tolerance, these effects may be intensified at the borders of the thermal tolerance window. We studied the effects of elevated CO2 concentrations on clapping performance and energy metabolism of the commercially important scallop Pecten maximus. Individuals were exposed for at least 30 days to 4 °C (winter) or to 10 °C (spring/summer) at either ambient (0.04 kPa, normocapnia) or predicted future PCO2 levels (0.11 kPa, hypercapnia). Cold-exposed (4 °C) groups revealed thermal stress exacerbated by PCO2 indicated by a high mortality overall and its increase from 55 % under normocapnia to 90 % under hypercapnia. We therefore excluded the 4 °C groups from further experimentation. Scallops at 10 °C showed impaired clapping performance following hypercapnic exposure. Force production was significantly reduced although the number of claps was unchanged between normocapnia- and hypercapnia-exposed scallops. The difference between maximal and resting metabolic rate (aerobic scope) of the hypercapnic scallops was significantly reduced compared with normocapnic animals, indicating a reduction in net aerobic scope. Our data confirm that ocean acidification narrows the thermal tolerance range of scallops resulting in elevated vulnerability to temperature extremes and impairs the animal’s performance capacity with potentially detrimental consequences for its fitness and survival in the ocean of tomorrow.

Description: Early life stages of marine crustaceans respond sensitively to elevated seawater P CO 2 . However, the underlying physiological mechanisms have not been studied well. We therefore investigated the effects of elevated seawater P CO 2 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 P CO 2 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 CO 2 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 CO 2 concentration. The present study indicates that larval development under elevated seawater P CO 2 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 P CO 2 level

Description: Population substructure of the cuttlefish Sepia officinalis (Mollusca: Cephalopoda), as investigated by genetic variation of microsatellite loci, has been reported to be extensively variable around the Iberian Peninsula (Pérez-Losada et al., 2002) and panmictic in the semienclosed Adriatic Sea (Garoia et al., 2004). Yet, no verified genetic information on population substructuring existed for the northern distribution range of this species in the English Channel and Bay of Biscay.So far, reproductive and migration behaviour and in-vitro oxygen binding properties of haemocyanin have suggested separate populations in the English Channel and the Bay of Biscay. However, extensive examination of genetic variation at seven microsatellite loci indicated no significant differences between cuttlefishes from both areas. The revealed degree of genetic variation at the screened loci is comparable to the panmictic population of S. officinalis in the Adriatic Sea (Garoia et al., 2004). S. officinalis in the English Channel and the Bay of Biscay should thus also be regarded as a panmictic population.Garoia, F et al. (2004). Heredity, 93: 166-174.Pérez-Losada, M et al. (2002). Heredity, 89(6): 417-424.

Description: Activities of the antioxidative enzymes superoxide dismutase (SOD), catalase, glutathione peroxidase (GPX) and glutathione reductase (GR) were measured in the cephalopods Sepia officinalis and Lolliguncula brevis. Maximal enzyme activities were higher in gill tissue than in the mantle musculature of both species. Activities were generally lower in tissues of L. brevis than in S. officinalis. Comparison with other ectothermic animals showed both cephalopod species to have a low enzymatic antioxidative status despite their high metabolic rate. Furthermore, changes in antioxidative enzyme activities were measured in the cuttlefish S. officinalis with increasing age. The concentrations of malondialdehyde (MDA) and lipofuscin were determined as indicators of lipid peroxidation. Investigated animals were between 1.5 months and over 12 months old. Changes of antioxidative enzyme activities with age were not uniform. SOD and GPX activities increased with age, while catalase activity declined. In contrast, GR activity remained almost unchanged in all age groups. The low level of antioxidative defense might allow for the significant age-induced rise in MDA levels in gills and mantle musculature and for the increase in lipofuscin levels in mantle and brain tissue. It might thereby contribute to increased oxidative damage and a short life span in these cephalopods.