Concentration of carbon dioxide is rising with constantly progressing industrialization. The major sources of anthropogenic CO2 are combustion of fossil fuels, transport and industrial processes. This gas is present not only in the atmosphere, but also in water. The oceans play an extremely important role in absorbing CO2 from the atmosphere, and that process leads to a decrease in water pH, which causes changes in the environment and all the organisms present within. Acidification of the oceans is a process in which the pH value of water is reduced by increased partial pressure of CO2 in the water. It is expected to observe a decrease in the pH of the open oceans by 0.35 units over the next hundred years. Reactions of marine organisms to ambient condition changes are specific for particular types or species. Effect of water acidification may affect organisms positively, negatively or may not have an impact at all. Organisms with calcareous shells or frustules are especially vulnerable to ocean acidification. Carbonic acid occurring as a result of the increasing CO2 concentration in water will dissolve the shells built of calcium carbonate. This review focuses on the excessive amount of carbon dioxide in the atmosphere and the impact of this phenomenon on the marine ecosystems.
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Posts Tagged 'Baltic'
Ocean acidification: environmental issue and its impact on marine life
Published 5 February 2014 Science ClosedTags: Baltic, review
Biogeochemical control of the coupled CO2–O2 system of the Baltic Sea: a review of the results of Baltic-C
Published 17 January 2014 Science ClosedTags: Baltic, biogeochemistry, chemistry, modeling, regionalmodeling
Past, present, and possible future changes in the Baltic Sea acid–base and oxygen balances were studied using different numerical experiments and a catchment–sea model system in several scenarios including business as usual, medium scenario, and the Baltic Sea Action Plan. New CO2 partial pressure data provided guidance for improving the marine biogeochemical model. Continuous CO2 and nutrient measurements with high temporal resolution helped disentangle the biogeochemical processes. These data and modeling indicate that traditional understandings of the nutrient availability–organic matter production relationship do not necessarily apply to the Baltic Sea. Modeling indicates that increased nutrient loads will not inhibit future Baltic Sea acidification; instead, increased mineralization and biological production will amplify the seasonal surface pH cycle. The direction and magnitude of future pH changes are mainly controlled by atmospheric CO2 concentration. Apart from decreasing pH, we project a decreasing calcium carbonate saturation state and increasing hypoxic area.
Effect of carbon dioxide-induced water acidification on the physiological processes of the Baltic isopod Saduria entomon
Published 16 January 2014 Science ClosedTags: Baltic, biological response, crustaceans, laboratory, mortality, performance, physiology
Survival, behavior, hemolymph pH, osmolality, and chloride ion concentration as well the total metabolic rate (heat dissipation rate) of the isopod Saduria entomon from the brackish Baltic Sea were investigated after exposure to carbon dioxide-induced water acidification (pH, 7.5, 7.0, and 6.5; control pH, 8.2) keeping other parameters constant (temperature, 10°C; salinity, 7). The short-term (12 h per each pH treatment) exposure to carbon dioxide-induced water acidification did not cause significant changes (P < 0.05) in the resting metabolic rate or the scope of activity in S. entomon; however, high interindividual variability was observed. The 2-wk exposure to lowered pH values did not affect either the activity of the isopods or their survival rate significantly (P > 0.05), which was greater than 90% in all pH treatments. The hemolymph pH increased significantly (P < 0.05) with a decrease of water pH from a control pH of 8.2 down to a pH of 7.0. Hemolymph osmolality increased significantly (P < 0.05) at pH 7.5, but exposure to pH 7.0 did not cause further increase in this parameter. Reduction of water pH did not affect the hemolymph chloride ion concentration. Obtained results indicated that S. entomon is adapted to large fluctuations of carbon dioxide levels in the water primarily to compensate for acid—base disturbances without additional energetic costs.
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Impact of CO2 enrichment on organic matter dynamics during nutrient induced coastal phytoplankton blooms
Published 15 January 2014 Science ClosedTags: abundance, Baltic, biogeochemistry, biological response, field, mesocosms, phytoplankton, prokaryotes
A mesocosm experiment was conducted to investigate the impact of rising fCO2 on the build-up and decline of organic matter during coastal phytoplankton blooms. Five mesocosms (∼38 m³ each) were deployed in the Baltic Sea during spring (2009) and enriched with CO2 to yield a gradient of 355–862 µatm. Mesocosms were nutrient fertilized initially to induce phytoplankton bloom development. Changes in particulate and dissolved organic matter concentrations, including dissolved high-molecular weight (>1 kDa) combined carbohydrates, dissolved free and combined amino acids as well as transparent exopolymer particles (TEP), were monitored over 21 days together with bacterial abundance, and hydrolytic extracellular enzyme activities. Overall, organic matter followed well-known bloom dynamics in all CO2 treatments alike. At high fCO2, higher ΔPOC:ΔPON during bloom rise, and higher TEP concentrations during bloom peak, suggested preferential accumulation of carbon-rich components. TEP concentration at bloom peak was significantly related to subsequent sedimentation of particulate organic matter. Bacterial abundance increased during the bloom and was highest at high fCO2. We conclude that increasing fCO2 supports production and exudation of carbon-rich components, enhancing particle aggregation and settling, but also providing substrate and attachment sites for bacteria. More labile organic carbon and higher bacterial abundance can increase rates of oxygen consumption and may intensify the already high risk of oxygen depletion in coastal seas in the future.
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Projected marine climate change: effects on copepod oxidative status and reproduction
Published 17 December 2013 Science ClosedTags: Baltic, biological response, crustaceans, laboratory, multiple factors, pathogens, physiology, reproduction, temperature, zooplankton
Zooplankton are an important link between primary producers and fish. Therefore, it is crucial to address their responses when predicting effects of climate change on pelagic ecosystems. For realistic community-level predictions, several biotic and abiotic climate-related variables should be examined in combination. We studied the combined effects of ocean acidification and global warming predicted for year 2100 with toxic cyanobacteria on the calanoid copepod, Acartia bifilosa. Acidification together with higher temperature reduced copepod antioxidant capacity. Higher temperature also decreased egg viability, nauplii development, and oxidative status. Exposure to cyanobacteria and its toxin had a negative effect on egg production but, a positive effect on oxidative status and egg viability, giving no net effects on viable egg production. Additionally, nauplii development was enhanced by the presence of cyanobacteria, which partially alleviated the otherwise negative effects of increased temperature and decreased pH on the copepod recruitment. The interactive effects of temperature, acidification, and cyanobacteria on copepods highlight the importance of testing combined effects of climate-related factors when predicting biological responses.
Habitat traits and food availability determine the response of marine invertebrates to ocean acidification
Published 2 December 2013 Science ClosedTags: Baltic, biological response, crustaceans, morphology, multiple factors, North Atlantic, nutrients, reproduction, survival
Energy availability and local adaptation are major components in mediating the effects of ocean acidification (OA) on marine species. In a long-term study, we investigated the effects of food availability and elevated pCO2 (~ 400, 1000 and 3000 μatm) on growth of newly settled Amphibalanus (Balanus) improvisus to reproduction, and on their offspring. We also compared two different populations, which were presumed to differ in their sensitivity to pCO2 due to differing habitat conditions: Kiel Fjord, Germany (Western Baltic Sea) with naturally strong pCO2 fluctuations, and the Tjärnö Archipelago, Sweden (Skagerrak) with far lower fluctuations. Over 20 weeks, survival, growth, reproduction and shell strength of Kiel barnacles were all unaffected by elevated pCO2, regardless of food availability. Moulting frequency and shell corrosion increased with increasing pCO2 in adults. Larval development and juvenile growth of the F1 generation were tolerant to increased pCO2, irrespective of parental treatment. In contrast, elevated pCO2 had a strong negative impact on survival of Tjärnö barnacles. Specimens from this population were able to withstand moderate levels of elevated pCO2 over 5 weeks when food was plentiful but showed reduced growth under food limitation. Severe levels of elevated pCO2 negatively impacted growth of Tjärnö barnacles in both food treatments. We demonstrate a conspicuously higher tolerance to elevated pCO2 in Kiel barnacles than in Tjärnö barnacles. This tolerance was carried-over from adults to their offspring. Our findings indicate that populations from fluctuating pCO2 environments are more tolerant to elevated pCO2 than populations from more stable pCO2 habitats. We furthermore provide evidence that energy availability can mediate the ability of barnacles to withstand moderate CO2 stress. Considering the high tolerance of Kiel specimens and the possibility to adapt over many generations, near future OA alone does not seem to present a major threat for A. improvisus.
Modelling the dissolved inorganic carbon system in the Baltic Sea
Published 17 October 2013 Science ClosedTags: Baltic, biogeochemistry, chemistry, multiple factors, nutrients, temperature
Oceans are capable of storing part of the emitted anthropogenic carbon dioxide (CO2) due to the formation of carbonic acid and subsequent dissociation. CO2 is also assimilated by biota and the inorganic carbon system is thus coupled to biogeochemical processes. Naturally, it is a substantial improvement of model realism if the inorganic carbon system is fully coupled to biogeochemistry in numerical models. The focus of this thesis has been to improve the accuracy of pH and the partial pressure of CO2 (pCO2) computations for a marine environment like the Baltic Sea, but the knowledge we have gained is generally applicable. A model system has been developed to consider several environmental threats simultaneously (eutrophication, acidification, and climate change) and the model skill has been certified by using objective skill metrics. To improve the coupling between biogeochemical processes and the dissolved inorganic carbon system, generation and depletion of total alkalinity (AT) due to several biogeochemical reactions was added to the model. In situ generation of AT was found to be important, specifically in regions with permanent or periodic anoxia, as the major AT changes were coupled to oxidation–reduction (redox) reactions. Without adding AT from these processes, the correct pH could not be calculated in anoxic waters and the mean volume AT content was found to be too low. The improvements were put to use when several environmental threats were evaluated simultaneously in a study of possible future changes in the Baltic Sea pH and oxygen balances. A coupled model for the catchment and sea was set up and forced by meteorological and hydrological datasets and scenarios. The results showed that increased nutrient loads will not inhibit future Baltic Sea acidification, but the seasonal pH cycle will be amplified by increased biological production and mineralization. The study indicated future acidification of the whole Baltic Sea and that the main factor controlling the direction and magnitude of the change was the atmospheric CO2 concentration. Through a previous investigation of the sensitivity of Baltic Sea surface water pH it was found that increased atmospheric CO2 can affect pH also through changing the river chemistry, especially with regard to AT. The latter could severely impact water in the northern Baltic region. To improve modelled seasonal pCO2 variations dissolved organic matter was added to a numerical model. The modelled phytoplankton were allowed to utilize the dissolved organic nutrients and the biological drawdown of CO2 in the Eastern Gotland basin was much improved by this. When phytoplankton used the organic nutrients the CO2 assimilation was higher during the summer months and the partial pressure of CO2 decreased by ~200 µatm in the Eastern Gotland Basin as a result. In the Bothnian Bay, both the duration and magnitude of CO2 assimilation was doubled when phytoplankton utilized dissolved organic nutrients.
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Effects of seawater pCO2 and temperature on shell growth, shell stability, condition and cellular stress of Western Baltic Sea Mytilus edulis (L.) and Arctica islandica (L.)
Published 6 September 2013 Science ClosedTags: Baltic, biological response, laboratory, mollusks, morphology, mortality, multiple factors, performance, physiology, temperature
Acidification of the World’s oceans may directly impact reproduction, performance and shell formation of marine calcifying organisms. In addition, since shell production is costly and stress in general draws on an organism’s energy budget, shell growth and stability of bivalves should indirectly be affected by environmental stress. The aim of this study was to investigate whether a combination of warming and acidification leads to increased physiological stress (lipofuscin accumulation and mortality) and affects the performance [shell growth, shell breaking force, condition index (Ci)] of young Mytilus edulis and Arctica islandica from the Baltic Sea. We cultured the bivalves in a fully-crossed 2-factorial experimental setup (seawater (sw) pCO2 levels “low”, “medium” and “high” for both species, temperature levels 7.5, 10, 16, 20 and 25 °C for M. edulis and 7.5, 10 and 16 °C for A. islandica) for 13 weeks in summer. Mytilus edulis and A. islandica appeared to tolerate wide ranges of sw temperature and pCO2. Lipofuscin accumulation of M. edulis increased with temperature while the Ci decreased, but shell growth of the mussels only sharply decreased while its mortality increased between 20 and 25 °C. In A. islandica, lipofuscin accumulation increased with temperature, whereas the Ci, shell growth and shell breaking force decreased. The pCO2 treatment had only marginal effects on the measured parameters of both bivalve species. Shell growth of both bivalve species was not impaired by under-saturation of the sea water with respect to aragonite and calcite. Furthermore, independently of water temperatures shell breaking force of both species and shell growth of A. islandica remained unaffected by the applied elevated sw pCO2 for several months. Only at the highest temperature (25 °C), growth arrest of M. edulis was recorded at the high sw pCO2 treatment and the Ci of M. edulis was slightly higher at the medium sw pCO2 treatment than at the low and high sw pCO2 treatments. The only effect of elevated sw pCO2 on A. islandica was an increase in lipofuscin accumulation at the high sw pCO2 treatment compared to the medium sw pCO2 treatment. Our results show that, despite this robustness, growth of both M. edulis and A. islandica can be reduced if sw temperatures remain high for several weeks in summer. As large body size constitutes an escape from crab and sea star predation, this can make bivalves presumably more vulnerable to predation—with possible negative consequences on population growth. In M. edulis, but not in A. islandica, this effect is amplified by elevated sw pCO2. We follow that combined effects of elevated sw pCO2 and ocean warming might cause shifts in future Western Baltic Sea community structures and ecosystem services; however, only if predators or other interacting species do not suffer as strong from these stressors.
Effects of carbon dioxide on biomass and species composition of a natural Baltic Sea spring bloom community
Published 28 August 2013 Science ClosedTags: abundance, Baltic, biological response, community composition, laboratory, phytoplankton
Carbon dioxide (CO2) emissions contribute to an increased mean temperature of the Earth and ocean acidification. The environmental changes give great concern for biodiversity and future environmental sustainability. Microalgae can possibly be used to recycle CO2 emissions and the biomass could be used for production of high value products like, healthy human food or biofuels. The aim of this study was to examine the effect of carbon dioxide on algae biomass production and species composition of a natural spring bloom community (NC) from the Baltic Sea. Spring blooms are dominated by diatoms which could be a good candidate for CO2 assimilation. The NC was exposed to CO2 gas and compared with NC without added CO2 sources (Air control). The NC was cultivated under controlled laboratory conditions with daily sampling for chlorophyll a and pH measurement. Species composition was investigated by microscope. Low pH reduced CO2 assimilation of the NC but was compensated for since no effect of CO2 could be seen on biomass production. Additionally CO2 had no effect on species composition indicating the species in the NC to be resistant to pH fluctuations. A clear shift in species composition could be seen over time. The diatoms dominated at experiment end confirming that they could potentially be used for algae cultivation to recycle CO2.
Extreme variations of pCO2 and pH in a macrophyte meadow of the Baltic Sea in summer: evidence of the effect of photosynthesis and local upwelling
Published 23 August 2013 Science ClosedTags: Baltic, biogeochemistry, chemistry, field
The impact of ocean acidification on benthic habitats is a major preoccupation of the scientific community. However, the natural variability of pCO2 and pH in those habitats remains understudied, especially in temperate areas. In this study we investigated temporal variations of the carbonate system in nearshore macrophyte meadows of the western Baltic Sea. These are key benthic ecosystems, providing spawning and nursery areas as well as food to numerous commercially important species. In situ pCO2, pH (total scale), salinity and PAR irradiance were measured with a continuous recording sensor package dropped in a shallow macrophyte meadow (Eckernförde bay, western Baltic Sea) during three different weeks in July (pCO2 and PAR only), August and September 2011.The mean (± SD) pCO2 in July was 383±117 µatm. The mean (± SD) pCO2 and pHtot in August were 239±20 µatm and 8.22±0.1, respectively. The mean (± SD) pCO2 and pHtot in September were 1082±711 µatm and 7.83±0.40, respectively. Daily variations of pCO2 due to photosynthesis and respiration (difference between daily maximum and minimum) were of the same order of magnitude: 281±88 µatm, 219±89 μatm and 1488±574 µatm in July, August and September respectively. The observed variations of pCO2 were explained through a statistical model considering wind direction and speed together with PAR irradiance. At a time scale of days to weeks, local upwelling of elevated pCO2 water masses with offshore winds drives the variation. Within days, primary production is responsible. The results demonstrate the high variability of the carbonate system in nearshore macrophyte meadows depending on meteorology and biological activities. We highlight the need to incorporate these variations in future pCO2 scenarios and experimental designs for nearshore habitats.
Maintenance of coelomic fluid pH in sea urchins exposed to elevated CO2: the role of body cavity epithelia and stereom dissolution
Published 22 August 2013 Science ClosedTags: Baltic, biological response, dissolution, echinoderms, growth, laboratory, morphology, North Atlantic, physiology
Experimental ocean acidification leads to a shift in resource allocation and to an increased [HCO3 −] within the perivisceral coelomic fluid (PCF) in the Baltic green sea urchin Strongylocentrotus droebachiensis. We investigated putative mechanisms of this pH compensation reaction by evaluating epithelial barrier function and the magnitude of skeleton (stereom) dissolution. In addition, we measured ossicle growth and skeletal stability. Ussing chamber measurements revealed that the intestine formed a barrier for HCO3 − and was selective for cation diffusion. In contrast, the peritoneal epithelium was leaky and only formed a barrier for macromolecules. The ossicles of 6 week high CO2-acclimatised sea urchins revealed minor carbonate dissolution, reduced growth but unchanged stability. On the other hand, spines dissolved more severely and were more fragile following acclimatisation to high CO2. Our results indicate that epithelia lining the PCF space contribute to its acid–base regulation. The intestine prevents HCO3 − diffusion and thus buffer leakage. In contrast, the leaky peritoneal epithelium allows buffer generation via carbonate dissolution from the surrounding skeletal ossicles. Long-term extracellular acid–base balance must be mediated by active processes, as sea urchins can maintain relatively high extracellular [HCO3 −]. The intestinal epithelia are good candidate tissues for this active net import of HCO3 − into the PCF. Spines appear to be more vulnerable to ocean acidification which might significantly impact resistance to predation pressure and thus influence fitness of this keystone species.
Elevated CO2 levels do not affect the shell structure of the bivalve Arctica islandica from the Western Baltic
Published 8 August 2013 Science ClosedTags: Baltic, biological response, growth, laboratory, mollusks, morphology
Shells of the bivalve Arctica islandica are used to reconstruct paleo-environmental conditions (e.g. temperature) via biogeochemical proxies, i.e. biogenic components that are related closely to environmental parameters at the time of shell formation. Several studies have shown that proxies like element and isotope-ratios can be affected by shell growth and microstructure. Thus it is essential to evaluate the impact of changing environmental parameters such as highpCO2 and consequent changes in carbonate chemistry on shell properties to validate these biogeochemical proxies for a wider range of environmental conditions. Growth experiments withArctica islandica from the Western Baltic Sea kept under different pCO2 levels (from 380 to 1120 µatm) indicate no affect of elevated pCO2 on shell growth or crystal microstructure, indicating that A. islandica shows an adaptation to a wider range of pCO2 levels than reported for other species. Accordingly, proxy information derived from A. islandica shells of this region contains no pCO2 related bias.
Differential responses of calcifying and non-calcifying epibionts of a brown macroalga to present-day and future upwelling pCO2
Published 5 August 2013 Science ClosedTags: algae, annelids, Baltic, biological response, bryozoa, growth, laboratory, morphology, reproduction
Seaweeds are key species of the Baltic Sea benthic ecosystems. They are the substratum of numerous fouling epibionts like bryozoans and tubeworms. Several of these epibionts bear calcified structures and could be impacted by the high pCO2 events of the late summer upwellings in the Baltic nearshores. Those events are expected to increase in strength and duration with global change and ocean acidification. If calcifying epibionts are impacted by transient acidification as driven by upwelling events, their increasing prevalence could cause a shift of the fouling communities toward fleshy species. The aim of the present study was to test the sensitivity of selected seaweed macrofoulers to transient elevation of pCO2 in their natural microenvironment, i.e. the boundary layer covering the thallus surface of brown seaweeds. Fragments of the macroalga Fucus serratus bearing an epibiotic community composed of the calcifiers Spirorbis spirorbis (Annelida) and Electra pilosa (Bryozoa) and the non-calcifier Alcyonidium hirsutum (Bryozoa) were maintained for 30 days under three pCO2 conditions: natural 460±59 µatm, present-day upwelling1193±166 µatm and future upwelling 3150±446 µatm. Only the highest pCO2 caused a significant reduction of growth rates and settlement of S. spirorbis individuals. Additionally, S. spirorbis settled juveniles exhibited enhanced calcification of 40% during daylight hours compared to dark hours, possibly reflecting a day-night alternation of an acidification-modulating effect by algal photosynthesis as opposed to an acidification-enhancing effect of algal respiration. E. pilosa colonies showed significantly increased growth rates at intermediate pCO2 (1193 µatm) but no response to higher pCO2. No effect of acidification on A. hirsutum colonies growth rates was observed. The results suggest a remarkable resistance of the algal macro-epibionts to levels of acidification occurring at present day upwellings in the Baltic. Only extreme future upwelling conditions impacted the tubeworm S. spirorbis, but not the bryozoans.
Community interactions dampen acidification effects in a coastal plankton system
Published 29 July 2013 Science ClosedTags: abundance, Baltic, crustaceans, physiology, phytoplankton, reproduction, zooplankton
Changing seawater chemistry towards reduced pH as a result of increasing atmospheric carbon dioxide (CO2) is affecting oceanic organisms, particularly calcifying species. Responses of non-calcifying consumers are highly variable and mainly mediated through indirect ocean acidification effects induced by changing the biochemical content of their prey, as shown within single species and simple 2-trophic level systems. However, it can be expected that indirect CO2 impacts observed at the single species level are compensated at the ecosystem level by species richness and complex trophic interactions. A dampening of CO2-effects can be further expected for coastal communities adapted to strong natural fluctuations in pCO2, typical for productive coastal habitats. Here we show that a plankton community of the Kiel Fjord was tolerant to CO2 partial pressure (pCO2) levels projected for the end of this century (<1400 µatm), and only subtle differences were observed at the extremely high value of 4000 µatm. We found similar phyto- and microzooplankton biomass and copepod abundance and egg production across all CO2 treatment levels. Stoichiometric phytoplankton food quality was minimally different at the highest pCO2 treatment, but was far from being potentially limiting for copepods. These results are in contrast to studies that include only a single species, which observe strong indirect CO2 effects for herbivores and suggest limitations of biological responses at the level of organism to community. Although this coastal plankton community was highly tolerant to high fluctuations in pCO2, increase in hypoxia and CO2 uptake by the ocean can aggravate acidification and may lead to pH changes outside the range presently experienced by coastal organisms.
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Impact of elevated pCO2 on benthic foraminifera from the southwestern Baltic Sea
Published 17 July 2013 Science ClosedTags: Baltic, biological response, calcification, chemistry, dissolution, field, laboratory, morphology, mortality, multiple factors, protists, reproduction, salinity, temperature
Increasing atmospheric CO2 concentrations have a strong impact on the marine carbonate chemistry leading to a phenomenon called ocean acidification. Excess CO2 dissolves in the surface water of the ocean, thereby the seawater pCO2 increases, whereas the [CO3 2-] and pH decrease. Reduced CO3 2- concentrations may affect marine, especially calcifying, organisms such as benthic foraminifera, in that their ability to form calcareous tests might be affected. In comparison to open oceans, water pCO2 levels are often not in equilibrium with the atmosphere in coastal regions, which are characterized by high CO2 variability during the seasonal cycle. This has also been observed for the southwestern Baltic, an eutrophic marginal sea, where bacterial degradation of large amounts of organic matter cause O2 depletion and CO2 enrichment in the bottom water.
In the frame of this thesis, the impact of elevated pCO2, temperature and salinity changes on the survival and calcification ability of the benthic foraminiferal species Ammonia aomoriensis was investigated in mid-term and long-term laboratory experiments. Under laboratory conditions, foraminifera were either isolated from the sediment or remained in their natural microhabitat. Further, the natural carbonate system variability and its impact on foraminiferal communities were monitored in a one-year field study.
Specimens of Ammonia aomoriensis were isolated from their natural sediment. They exhibited reduced survival and growth rates with increasing pCO2 of up to 3130 μatm under laboratory conditions. At pCO2 levels above 1800 μatm, dissolution caused a decrease of test diameter, and at the highest pCO2, only the inner organic lining remained. Testing the combined effects of ocean acidification, temperature and salinity on living Ammonia aomoriensis, a significant reduction of test diameter was observed at a pCO2 >1200 μatm (Ωcalc<1). Tests were mainly affected by undersaturation of calcite. This effect was partly compensated by a temperature rise, which increased Ωcalc and led to lower test degradation. In contrast, salinity did not have a significant effect on test growth. These results revealed that Ammonia ammoriensis exhibited a high sensitivity to elevated pCO2 and accompanying calcium carbonate undersaturation when the specimens were kept without their protective sedimentary habitat.
During the field survey, large seasonal fluctuations of pCO2 from 465 up to 3429 μatm were encountered in the bottom water of Flensburg Fjord in the southwestern Baltic Sea. The pCO2 in the sediment pore water reached even higher values ranging from 1244 to 3324 μatm. However, and as a consequence of higher alkalinity (AT), the calcium carbonate saturation state of the sediment pore water remained slightly supersaturated with respect to calcite for most of the year. Accordingly, during the monitoring period, no dynamic responses of foraminiferal population density and diversity to elevated sediment pore water pCO2 were recognized.
Benthic foraminifera may indeed cope with a high sediment pore water pCO2 as long as the sediment pore water remains calcite supersaturated. This evidence from the field study was also supported by the results of a long-term laboratory experiment, in which a complete foraminiferal fauna in their natural sediment was exposed to elevated pCO2 levels. Similar to field observations, the sediment pore water exhibited higher alkalinity and consequently higher saturation state of Ωcalc in comparison to the overlying seawater. Thereby the sediment chemistry created a microhabitat, which sustained the growth and development of the foraminiferal community even at highly elevated pCO2. The dominant species Ammonia aomoriensis exhibited growth and several reproduction events during the incubation time. Nevertheless, dissolution was observed on dead, empty tests of Ammonia aomoriensis, whereas tests of the second-ranked species Elphidium incertum stayed intact at high pCO2 and Ωcalc<1. This species-specific response could be due to differences in elemental composition and ultrastructure of the test walls. Benthic foraminifera in their natural, sedimentary habitat tolerate elevated pCO2 under laboratory conditions and the current high sedimentary pore water pCO2, which prevails in the southwestern Baltic Sea. In this environment, organic-rich mud influences the carbonate chemistry, and thereby provides a suitable habitat for benthic foraminifera. Consequently, the calcifying Ammonia aomoriensis plays an important role in benthic carbonate production and accumulation in this area. These results emphasize the importance of understanding the carbonate chemistry in the natural environment of benthic foraminifera, which depends upon sediment composition and remineralization processes.
It is expected that enhanced future CO2 uptake in the water column will cause a further rise of sedimentary pore water pCO2 levels. As a consequence, undersaturation with respect to calcite will occur more frequently even in the sediment. This will most probably affect the dominant species Ammonia aomoriensis, which might induce changes in the benthic foraminiferal communities and their carbonate production in the southwestern Baltic Sea.
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Effects of reduced pH on Macoma balthica larvae from a system with naturally fluctuating pH-dynamics
Published 8 July 2013 Science ClosedTags: Baltic, biological response, laboratory, mollusks, morphology, survival
Ocean acidification is causing severe changes in the inorganic carbon balance of the oceans. The pH conditions predicted for the future oceans are, however, already regularly occurring in the Baltic Sea, and the system might thus work as an analogue for future ocean acidification scenarios. The characteristics of the Baltic Sea with low buffering capacity and large natural pH fluctuations, in combination with multiple other stressors, suggest that OA effects may be severe, but remain largely unexplored. A calcifying species potentially affected by low pH conditions is the bivalve Macoma balthica (L.). We investigated larval survival and development of M. balthica by exposing the larvae to a range of pH levels: 7.2, 7.4, 7.7 and 8.1 during 20 days in order to learn what the effects of reduced pH are on the larval biology and thus also potentially for the population dynamics of this key species. We found that even a slight pH decrease causes significant negative changes during the larval phase, both by slowing growth and by decreasing survival. The growth was slower in all reduced pH treatments compared to the control treatment. The size of 250 µm that is considered indicative to imminent settling in our system was reached by 22% of the larvae grown in control conditions after 20 days, whereas in all reduced pH treatments the size of 250 µm was reached by only 7–14%. The strong impact of ocean acidification on larvae is alarming as slowly growing individuals are exposed to higher predation risk in response to the longer time they are required to spend in the plankton, further decreasing the ecological competence of the species.
Results of laboratory and field experiments of the direct effect of increasing CO2 on net primary production of macroalgal species in brackish-water ecosystems
Published 22 May 2013 Science ClosedTags: algae, Baltic, biological response, field, laboratory, mesocosms, primary production
Studies on the effects of increasing acidification on marine communities have been previously mostly carried out in truly marine areas whereas brackish-water ecosystems such as the Baltic Sea have been less studied. The current study analyses how acidification induced by elevated atmospheric carbon dioxide affects the photosynthetic net production of different macroalgal species in the brackish Baltic Sea. Research methods include sets of laboratory and field experiments carried out in shallow coastal brackish waters. The aim of the laboratory experiments was to develop the necessary techniques and experience for the mesocosm experiments. Laboratory experiments were carried out using specimens of the red alga Furcellaria lumbricalis collected from Kakumäe Bay. The mesocosm experiments were conducted in Kõiguste Bay during the field season of 2011. Separate mesocosms were operated in each set with different CO2 concentrations and a control treatment in natural conditions. Field experiments were carried out with three species representing three different morphological and ecological groups: Ulva intestinalis, a fast-growing green alga; Fucus vesiculosus, a perennial brown alga with a slow metabolism; and Furcellaria lumbricalis, a perennial red alga. Photosynthetic activity was used as the response variable. In the laboratory decreasing pH increased the net primary production of F. lumbricalis with the lowest net primary production values measured at pH 8.0 and the highest at pH 6.5. Results of the field experiments indicated that increased CO2 levels in seawater favoured photosynthetic activity of the macroalgae U. intestinalis and F. lumbricalis, but F. vesiculosus showed no response to elevated CO2. Elevated CO2 levels are suggested to favour the production of fast-growing filamentous species, which thus may indirectly enhance the effect of eutrophication in the shallow coastal brackish waters.
A potential tool for high-resolution monitoring of ocean acidification
Published 2 May 2013 Science ClosedTags: Baltic, chemistry, field, methods, North Atlantic
Current anthropogenic carbon dioxide emissions generate besides global warming unprecedented acidification rates of the oceans. Recent evidence indicates the possibility that ocean acidification and low oceanic pH may be a major reason for several mass extinctions in the past. However, a major bottleneck for research on ocean acidification is long-term monitoring and the collection of consistent high-resolution pH measurements. This study presents a low-power (< 1 W) small sample volume (25 μl) semiconductor based fluorescence method for real-time ship-board pH measurements at high temporal and spatial resolution (approximately 15 s and 100 m between samples). A 405 nm light emitting diode and the blue and green channels from a digital camera was used for swift detection of fluorescence from the pH sensitive dye 6,8-Dihydroxypyrene-1,3-disulfonic acid in real-time. Main principles were demonstrated by automated continuous measurements of pH in the surface water across the Baltic Sea and the Kattegat region with a large range in salinity (∼ 3–30) and temperature (∼ 0–25 °C). Ship-board precision of salinity and temperature adjusted pH measurements were estimated as low as 0.0001 pH units.
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Sex in murky waters: anthropogenic disturbance of sexual selection in pipefish
Published 27 March 2013 Science ClosedTags: Baltic, biological response, fish, laboratory, North Atlantic, reproduction
Animals experience variation in their environment because of natural changes. However, due to anthropogenic disturbance, the speed and severity of these changes have recently increased. This thesis investigates how reproductive behaviours may be affected by human induced environmental change. In specific, I investigate how visual and chemical changes in the aquatic environment, caused by eutrophication, affect mating systems and sexual selection in fish. Broad-nosed- and straight-nosed pipefish, which both have been studied in detail for a long period, were used as model organisms. These two species are particularly suitable model organisms since they perform complex courtship behaviours, including the advertisement of ornaments and a nuptial dance. Further, two distinct populations were studied, one on the Swedish west coast and one in the Baltic Sea, as these two locations vary in the degree and extent of environmental disturbance, in particular turbidity. I found that changes in the visual environment had no impact on the development of female sexual ornaments in these sex-role reversed pipefishes, but it hampered adaptive mate choice. Turbidity also had a negative effect on reproductive success in the Baltic Sea population. Changes in the chemical environment in the form of increased pH reduced the probability to mate, while hypoxia did not alter mating propensity. However, hypoxic water delayed the onset of both courting and mating. Hence, human induced change in aquatic environments may alter the processes of sexual selection and population dynamics.
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Response of Nodularia spumigena to pCO2 – Part 3: Turnover of phosphorus compounds (update)
Published 6 March 2013 Science ClosedTags: Baltic, biogeochemistry, biological response, growth, laboratory, molecular biology, prokaryotes
Diazotrophic cyanobacteria form extensive summer blooms in the Baltic Sea driving the surrounding surface waters into phosphate limitation. One of the main bloom-forming species is the heterocystous cyanobacterium Nodularia spumigena. N. spumigena exhibits accelerated uptake of phosphate through the release of the extracellular enzyme alkaline phosphatase whose activity also serves as an indicator of the hydrolysis of dissolved organic phosphorus (DOP). The present study investigated the utilisation of DOP and its compounds (e.g., ATP) by N. spumigena during growth under different CO2 concentrations, in order to estimate potential consequences of ocean acidification on the cell’s supply with phosphorus (P). Cell growth, the phosphorus pool, and four DOP compounds (ATP, DNA, RNA, and phospholipids) were determined in three setups with different CO2 concentrations (average 341 μatm, 399 μatm, and 508 μatm) during a 15-day batch experiment. The results showed stimulated growth of N. spumigena and a rapid depletion of dissolved inorganic phosphorus (DIP) in all pCO2 treatments. DOP uptake was enhanced by a factor of 1.32 at 399 μatm and of 2.25 at 508 μatm compared to the lowest CO2 concentration. Among the measured DOP compounds, none was found to accumulate preferentially during the incubation or in response to a specific pCO2 treatment. However, at the beginning 61.9 ± 4.3% of total DOP were not characterised but comprised the most utilised fraction. This is demonstrated by the decrement of this fraction to 27.4 ± 9.9% of total DOP during the growth phase with a preference at high pCO2. Our results indicate a stimulated growth of diazotrophic cyanobacteria at increasing CO2 concentrations which is accompanied by increasing utilisation of DOP as an alternative P source.


