Posts Tagged 'Baltic'

Intra-specific variation of ocean acidification effects in marine mussels and oysters: integrative physiological studies on tissue and organism responses

Uptake of increasing anthropogenic CO2 emissions by ocean surface waters is causing an increase of seawater PCO2 accompanied by a decrease of seawater pH and carbonate ion concentrations. This process, termed ocean acidification (OA), is predicted to negatively affect many marine organisms with likely consequences for marine ecosystems and the services they provide. Calcifying mussels and oysters, and particularly their early life stages, are predicted to be among the most OA sensitive taxa, as OA interferes with the calcification process. In addition, mussels and oysters possess a relatively low ability to compensate for CO2 induced disturbances in extracellular body fluid pH with potential physiological downstream effects such as elevated metabolic maintenance costs. As mussels and oysters are key habitat forming organisms in many highly productive temperate coastal communities, negative OA effects may translate into deleterious effects at an ecosystem scale. In particular, the relative long generation time of most marine bivalves raises the concern that the rapid rate at which OA occurs may outpace species’ ability to genetically adapt, leaving pre-existing genetic variation as a potential key to species resilience under OA. Against this backdrop, this thesis contributes to the understanding of physiological mechanisms that underpin and define the OA vulnerability of ecologically and economically important mussels and oysters. Thereby, emphasis was placed on investigating intra-specific variance as a proxy for potential adaptive capacities. Kiel Fjord is located in the Western Baltic Sea and is characterised by strong seasonal and diurnal fluctuations in seawater PCO2. These fluctuations are caused by upwelling events of acidified bottom waters with peak PCO2 values (>2300 μatm) that are already by far exceeding those projected for open ocean surface waters by the end of this century. Despite these unfavourable conditions, blue mussels (Mytilus edulis) dominate the benthic community, which makes this population particularly interesting in the context of metabolic adaptation to OA. Consequently, a long-term multi-generation CO2 acclimation experiment with different family lines of M. edulis from Kiel Fjord formed the first part of this thesis. Offspring of 16 different family lines were transferred to three different PCO2 conditions, representing present and predicted PCO2 levels in Kiel Fjord (700 μatm (control), 1120 μatm (intermediate) and 2400 μatm (high)). Larval survival rates were substantially different between family lines at the highest PCO2 level. Based on these differences, families were classified as either ‘tolerant’ (i.e. successful settlement at all PCO2 levels) or ‘sensitive’ (i.e. successful settlement only at control and intermediate PCO2 level). Subsequently, the offspring were raised for over one year at respective PCO2 levels, followed by measurements of physiological parameters at the whole-animal, tissue (gill and outer mantle) and biochemical level (key metabolic enzymes). The results revealed that routine metabolic rates (RMR) and summed tissue respiration were increased in tolerant families at intermediate PCO2, indicating elevated homeostatic costs. However, this higher energy demand at the intermediate PCO2 level was not accompanied by a simultaneous increase in energy assimilation (i.e. clearance rates (CR)), indicating an incipient imbalance in energy demand and supply. Consequently, RMRs at the highest PCO2 were not different to control RMRs but associated with reduced CRs, which correlated with a lower gill metabolic scope, reduced gill mitochondrial capacities (lower capacities for citrate synthase (CS) and cytochrome c oxidase (COX)) as well as an increased capacity for anaerobic energy production (lower ratio of pyruvate kinase to phospoenolpyruvate carboxykinase). In conjunction with a lower COX to CS ratio observed in outer mantle tissue, this suggested a CO2-induced shift of metabolic pathways in tolerant families at the highest PCO2 level. By contrast, sensitive families had an unchanged RMR, tissue respiration and CR at the intermediate CO2. However, a higher control RMR in sensitive than tolerant families at similar CR suggested a lower, CO2 independent metabolic efficiency in sensitive families. This was also reflected in their lower gill mitochondrial scope at control conditions compared to tolerant families. These findings suggested that sensitive families lack the metabolic scope to cover OA induced higher maintenance costs and have to rely on energy reallocation and thus, energy trade-offs which may also have prevented survival at the highest experimental PCO2 level. Accordingly, investigations of 3-hydroxyacyl-CoA dehydrogenase (HADH) capacities, which catalyses a key step in lipid oxidation, suggested an increased reliance on lipids as metabolic fuel in sensitive families at elevated PCO2. If this was also prevalent during the larval phase, a quicker depletion of lipid reserves before completion of metamorphosis may have contributed to the higher larval mortality at the highest PCO2 treatment in sensitive compared to tolerant mussels. The second part of the thesis aimed to clarify whether a higher OA tolerance in Sydney rock oysters (Saccostrea glomerata) is directly correlated with an increased capacity to compensate for CO2 induced extracellular acid-base disturbances, and whether such a capacity is driven by higher metabolic and ion-regulatory costs at the tissue level. Earlier studies focusing on two different populations of Sydney rock oysters demonstrated that oysters that were selectively bred for increased growth and disease resistance (‘selected oysters’) have a higher CO2 resilience compared to the wild population (‘wild oysters’). To unravel the underlying physiological mechanisms, oysters of both populations were acclimated at control and elevated PCO2 (1100 μatm) levels for seven weeks, followed by determinations of extracellular acidbase parameters (pHe, PeCO2, [HCO3 -]e), tissue respiration and indirect determination of energy demands of major ion regulatory transport proteins. Indeed, at elevated PCO2, wild oysters had a lower pHe and an increased PeCO2 whereas extracellular acid-base status of selected oysters remained unaffected. However, differing pHe values between oyster types were not driven by elevated metabolic costs of major ion regulators at tissue level. Selected oysters rather exhibited an increased systemic capacity to eliminate metabolic CO2, which likely came through higher and energetically more efficient filtration rates and associated facilitation of gas exchange, suggesting that effective filtration and CO2 resilience might be positively correlated traits in oysters. In conclusion, the findings of this thesis contribute to the growing evidence that ongoing OA will likely impair the physiology of marine mussels and oysters with potentially associated downstream consequences for the respective ecosystems. However, the results also suggest adaptive capacities in both species studied. The higher CO2 resilience of selected Sydney rock oysters was expressed within the – in evolutionary terms – rapid time span of only a few generations of selective breeding, which indicates that rapid adaptation to OA may be possible in marine bivalves. The observed intra-specific variation of OA responses in blue mussels suggests standing genetic variation within this population, which is likely to be key for the persistence of populations under rapidly occurring OA. However, as global change is not limited to OA, future research will have to reassess potential resilience traits and adaptive capacities to OA when combined with changes in other environmental drivers.

Continue reading ‘Intra-specific variation of ocean acidification effects in marine mussels and oysters: integrative physiological studies on tissue and organism responses’

Could ocean acidification influence epiphytism? A comparison of carbon-use strategies between Fucus vesiculosus and its epiphytes in the Baltic Sea

Reduced seawater pH due to elevated carbon dioxide (CO2), a process known as ocean acidification (OA), is a globally significant environmental issue. OA is predicted to influence a range of ecosystem processes, but little is known about how changing seawater carbon chemistry could influence the extent and impacts of epiphytism. In the brackish Baltic Sea, increased epiphytism is associated with coastal eutrophication and the potential for OA to interact with this relationship remains unclear. This study focuses on slow-growing perennial algae Fucus vesiculosus—which is one of the most important habitat-forming species in the Baltic Sea—and two of its most common and abundant filamentous epiphytes Ceramium tenuicorne and Pylaiella littoralis. Material for this study was collected from Estonian coastal waters. The aim of the research was to determine which carbon acquisition strategies these species possess, which could indicate how they respond to predicted changes in seawater chemistry due to elevated CO2. Carbon-use strategies in macroalgae were determined by analyzing natural carbon isotope signatures (δ13C), pH drift experiments, and photosynthesis vs. dissolved inorganic carbon (P vs. DIC) curves. Our results showed that although F. vesiculosus and its filamentous epiphytes all possess a carbon concentrating mechanism (CCM), the potential species-specific variation in the CCMs operation will favor C. tenuicorne over F. vesiculosus and P. littoralis in a future high CO2 world.

Continue reading ‘Could ocean acidification influence epiphytism? A comparison of carbon-use strategies between Fucus vesiculosus and its epiphytes in the Baltic Sea’

Season affects strength and direction of the interactive impacts of ocean warming and biotic stress in a coastal seaweed ecosystem

The plea for using more “realistic,” community‐level, investigations to assess the ecological impacts of global change has recently intensified. Such experiments are typically more complex, longer, more expensive, and harder to interpret than simple organism‐level benchtop experiments. Are they worth the extra effort? Using outdoor mesocosms, we investigated the effects of ocean warming (OW) and acidification (OA), their combination (OAW), and their natural fluctuations on coastal communities of the western Baltic Sea during all four seasons. These communities are dominated by the perennial and canopy‐forming macrophyte Fucus vesiculosus—an important ecosystem engineer Baltic‐wide. We, additionally, assessed the direct response of organisms to temperature and pH in benchtop experiments, and examined how well organism‐level responses can predict community‐level responses to the dominant driver, OW. OW affected the mesocosm communities substantially stronger than acidification. OW provoked structural and functional shifts in the community that differed in strength and direction among seasons. The organism‐level response to OW matched well the community‐level response of a given species only under warm and cold thermal stress, that is, in summer and winter. In other seasons, shifts in biotic interactions masked the direct OW effects. The combination of direct OW effects and OW‐driven shifts of biotic interactions is likely to jeopardize the future of the habitat‐forming macroalga F. vesiculosus in the Baltic Sea. Furthermore, we conclude that seasonal mesocosm experiments are essential for our understanding of global change impact because they take into account the important fluctuations of abiotic and biotic pressures.

Continue reading ‘Season affects strength and direction of the interactive impacts of ocean warming and biotic stress in a coastal seaweed ecosystem’

Ocean acidification may threaten a unique seaweed community and associated industry in the Baltic Sea

Wild harvest of seaweed supports small-scale, high-value industries in a number of regions in the world. Information is lacking on how increasing carbon dioxide (CO2) concentrations in seawater could impact seaweeds in wild harvest situations. This study focuses on a mixed unattached loose-lying red algal community of Furcellaria lumbricalis in close association with Coccotylus truncatus that is found in the West Estonian Archipelago Sea, NE Baltic Sea. In Estonian coastal waters, the wild harvest of F. lumbricalis started in 1960s and it has since been used as raw material for furcellaran production. The aim of this study was to determine how ocean acidification may impact the balance of these two red algal species in the community. Mechanistic assessment of the carbon physiology of F. lumbricalis and C. truncatus was used to predict productivity and competitive interactions between these species in a high-CO2 world. Carbon use strategies in macroalgae were determined by analysing the natural abundances of carbon isotopes (δ13C), pH drift experiments and photosynthesis vs. dissolved inorganic carbon (DIC) curves. Photosynthesis of F. lumbricalis (operating with a CO2 concentrating mechanism or CCM) performed worse along the broader range of DIC concentrations compared to C. truncatus (non-CCM), especially those characterized under future climate conditions. Therefore, changing seawater carbon chemistry through ocean acidification has the potential to influence the balance of F. lumbricalis and C. truncatus in the community and the efficiency of the wild harvest of this community and the quality of product provided.

Continue reading ‘Ocean acidification may threaten a unique seaweed community and associated industry in the Baltic Sea’

Sensitivities to global change drivers may correlate positively or negatively in a foundational marine macroalga

Ecological impact of global change is generated by multiple synchronous or asynchronous drivers which interact with each other and with intraspecific variability of sensitivities. In three near-natural experiments, we explored response correlations of full-sibling germling families of the seaweed Fucus vesiculosus towards four global change drivers: elevated CO2 (ocean acidification, OA), ocean warming (OW), combined OA and warming (OAW), nutrient enrichment and hypoxic upwelling. Among families, performance responses to OA and OW as well as to OAW and nutrient enrichment correlated positively whereas performance responses to OAW and hypoxia anti-correlated. This indicates (i) that families robust to one of the three drivers (OA, OW, nutrients) will also not suffer from the two other shifts, and vice versa and (ii) families benefitting from OAW will more easily succumb to hypoxia. Our results may imply that selection under either OA, OW or eutrophication would enhance performance under the other two drivers but simultaneously render the population more susceptible to hypoxia. We conclude that intraspecific response correlations have a high potential to boost or hinder adaptation to multifactorial global change scenarios.

Continue reading ‘Sensitivities to global change drivers may correlate positively or negatively in a foundational marine macroalga’

Seasonal interactive effects of pCO2 and irradiance on the ecophysiology of brown macroalga Fucus vesiculosus L.

Stochastic upwelling of seawater in the Baltic Sea from the deep, anoxic bottoms may bring low-pH water rich in CO2 close to the surface. Such events may become more frequent with climate change and ongoing ocean acidification (OA). Photoautotrophs, such as macroalgae, which are important foundation species, have been proposed to benefit from increased carbon availability due to reduced energetic cost in carbon acquisition. However, the exact effects of CO2 fertilization may depend on the ambient light environment, as photosynthesis rates depend on available irradiance. In this experimental study, interacting effects of CO2 addition and irradiance on the habitat-forming macroalga Fucus vesiculosus were investigated during two seasons – winter and summer – in the northern Baltic Sea. Growth rates remained unaffected by CO2 or irradiance during both seasons, suggesting that direct effects of elevated CO2 on mature F. vesiculosus are small. Increases in CO2 affected algal elemental ratios by increasing carbon and decreasing nitrogen content, with resulting changes in the C:N ratio, but only in winter. In summer, chlorophyll a content increased under low irradiance. Increases in CO2 caused a decline in light-harvesting efficiency (decrease in Fv/Fm and α) under high irradiance in summer, and conversely increased α under low irradiance. High irradiance caused increases in the maximum relative electron transport rate (rETRmax) in summer, but not in winter. Differences between winter and summer indicate that F. vesiculosus responses to CO2 and irradiance are season-specific. Increases in carbon content during winter could indicate slightly positive effects of CO2 addition in the long run if the extra carbon gained may be capitalized in growth. The results of this study suggest that increases in CO2, either through upwelling or OA, may have positive effects on F. vesiculosus, but these effects are probably small.

Continue reading ‘Seasonal interactive effects of pCO2 and irradiance on the ecophysiology of brown macroalga Fucus vesiculosus L.’

Limited response of a spring bloom community inoculated with filamentous cyanobacteria to elevated temperature and pCO2

Temperature and CO2 levels are projected to increase in the future, with consequences for carbon and nutrient cycling in brackish environments, such as the Baltic Sea. Moreover, filamentous cyanobacteria are predicted to be favored over other phytoplankton groups under these conditions. Under a 12-day outdoor experiment, we examined the effect on a natural phytoplankton spring bloom community of elevated temperature (from 1°C to 4°C) and elevated pCO2 (from 390 to 970 μatm). No effects of elevated pCO2 or temperature were observed on phytoplankton biovolumes, but a significantly higher photosystem II activity was observed at elevated temperature after 9 days. In addition, three species of diazotrophic filamentous cyanobacteria were inoculated to test their competitive capacity under spring bloom conditions. The toxic cyanobacterium Nodularia spumigena exhibited an average specific growth rate of 0.10 d−1 by the end of the experiment, indicating potential prevalence even during wintertime in the Baltic Sea. Generally, none of the inoculated cyanobacteria species were able to outcompete the natural phytoplankton species at temperatures ≤4°C. No direct effects were found on heterotrophic bacteria. This study demonstrates the highly efficient resistance towards short-term (12 days) changes in abiotic factors by the natural Baltic Sea spring bloom community.

Continue reading ‘Limited response of a spring bloom community inoculated with filamentous cyanobacteria to elevated temperature and pCO2’


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Ocean acidification in the IPCC AR5 WG II

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