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  • 1
    Publication Date: 2021-03-19
    Description: In estuarine coastal systems such as the Baltic Sea, mussels suffer from low salinity which limits their distribution. Anthropogenic climate change is expected to cause further desalination which will lead to local extinctions of mussels in the low saline areas. It is commonly accepted that mussel distribution is limited by osmotic stress. However, along the salinity gradient environmental conditions for biomineralization are successively becoming more adverse as a result of reduced [Ca2+] and dissolved inorganic carbon (CT) availability. In larvae, calcification is an essential process starting during early development with formation of the prodissoconch I (PD I) shell which is completed under optimal conditions within 2 days. Experimental manipulations of seawater [Ca2+] start to impair PD I formation in Mytilus larvae at concentrations below 3 mM, which corresponds to conditions present in the Baltic at salinities below 8 g kg-1. In addition, lowering dissolved inorganic carbon to critical concentrations (〈 1 mM) similarly affected PD I size which was well correlated with calculated ΩAragonite and [Ca2+][HCO3-]/[H+] in all treatments. Comparing results for larvae from the western Baltic with a population from the central Baltic revealed significantly higher tolerance of PD I formation to lowered [Ca2+] and [Ca2+][HCO3-]/[H+] in the low saline adapted population. This may result from genetic adaptation to the more adverse environmental conditions prevailing in the low saline areas of the Baltic. The combined effects of lowered [Ca2+] and adverse carbonate chemistry represent major limiting factors for bivalve calcification and can thereby contribute to distribution limits of mussels in the Baltic Sea.
    Type: Article , PeerReviewed , info:eu-repo/semantics/article
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  • 2
    Publication Date: 2021-02-08
    Description: In vivo confocal Raman microscopy (CRM), polarized light microscopy and Fourier transform infrared spectroscopy (FTIR) were used to determine if a significant amount of amorphous calcium carbonate (ACC) exists within larval shells of Baltic mytilid mussels (Mytilus edulis-like) and whether the amount of ACC varies during larval development. No evidence for ACC was found from the onset of shell deposition at 21 h post-fertilization (hpf) until 48 hpf. Larval Mytilus shells were crystalline from 21 hpf onwards and exhibited CRM and FTIR peaks characteristic of aragonite. Prior to shell deposition at 21 hpf, no evidence for carbonates was observed through in vivo CRM.We further analysed the composition of larval shells in three other bivalve species, Mercenaria mercenaria, Crassostrea gigas and Crassostrea virginica and observed no evidence for ACC, which is in contrast to previous work on the same species. Our findings indicate that larval bivalve shells are composed of crystalline aragonite and we demonstrate that conflicting results are related to sub-optimal measurements and misinterpretation of CRM spectra. Our results demonstrate that the common perception that ACC generally occurs as a stable and abundant precursor during larval bivalve calcification needs to be critically reviewed.
    Type: Article , PeerReviewed , info:eu-repo/semantics/article
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  • 3
    Publication Date: 2021-02-08
    Description: Aim: Experimental simulation of near‐future ocean acidification (OA) has been demonstrated to affect growth and development of echinoderm larval stages through energy allocation towards ion and pH compensatory processes. To date, it remains largely unknown how major pH regulatory systems and their energetics are affected by trans‐generational exposure to near‐future acidification levels. Methods: Here, we used the common sea star Asterias rubens in a reciprocal transplant experiment comprising different combinations of OA scenarios, to study trans‐generational plasticity using morphological and physiological endpoints. Results: Acclimation of adults to pHT 7.2 (pCO2 3500 μatm) led to reductions in feeding rates, gonad weight and fecundity. No effects were evident at moderate acidification levels (pHT 7.4; pCO2 2000 μatm). Parental pre‐acclimation to pHT 7.2 for 85 days reduced developmental rates even when larvae were raised under moderate and high pH conditions, whereas pre‐acclimation to pHT 7.4 did not alter offspring performance. Microelectrode measurements and pharmacological inhibitor studies carried out on larval stages demonstrated that maintenance of alkaline gastric pH represents a substantial energy sink under acidified conditions that may contribute up to 30% to the total energy budget. Conclusion: Parental pre‐acclimation to acidification levels that are beyond the pH that is encountered by this population in its natural habitat (eg, pHT 7.2) negatively affected larval size and development, potentially through reduced energy transfer. Maintenance of alkaline gastric pH and reductions in maternal energy reserves probably constitute the main factors for a reduced juvenile recruitment of this marine keystone species under simulated OA.
    Type: Article , PeerReviewed
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  • 4
    Publication Date: 2020-02-06
    Description: Ocean acidification severely affects bivalves, especially their larval stages. Consequently, the fate of this ecologically and economically important group depends on the capacity and rate of evolutionary adaptation to altered ocean carbonate chemistry. We document successful settlement of wild mussel larvae (Mytilus edulis) in a periodically CO2-enriched habitat. The larval fitness of the population originating from the CO2-enriched habitat was compared to the response of a population from a nonenriched habitat in a common garden experiment. The high CO2–adapted population showed higher fitness under elevated Pco2 (partial pressure of CO2) than the non-adapted cohort, demonstrating, for the first time, an evolutionary response of a natural mussel population to ocean acidification. To assess the rate of adaptation, we performed a selection experiment over three generations. CO2 tolerance differed substantially between the families within the F1 generation, and survival was drastically decreased in the highest, yet realistic, Pco2 treatment. Selection of CO2-tolerant F1 animals resulted in higher calcification performance of F2 larvae during early shell formation but did not improve overall survival. Our results thus reveal significant short-term selective responses of traits directly affected by ocean acidification and long-term adaptation potential in a key bivalve species. Because immediate response to selection did not directly translate into increased fitness, multigenerational studies need to take into consideration the multivariate nature of selection acting in natural habitats. Combinations of short-term selection with long-term adaptation in populations from CO2-enriched versus nonenriched natural habitats represent promising approaches for estimating adaptive potential of organisms facing global change.
    Type: Article , PeerReviewed
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  • 5
    Publication Date: 2020-06-18
    Description: Understanding mollusk calcification sensitivity to ocean acidification (OA) requires a better knowledge of calcification mechanisms. Especially in rapidly calcifying larval stages, mechanisms of shell formation are largely unexplored—yet these are the most vulnerable life stages. Here we find rapid generation of crystalline shell material in mussel larvae. We find no evidence for intracellular CaCO3 formation, indicating that mineral formation could be constrained to the calcifying space beneath the shell. Using microelectrodes we show that larvae can increase pH and [CO32−] beneath the growing shell, leading to a ~1.5-fold elevation in calcium carbonate saturation state (Ωarag). Larvae exposed to OA exhibit a drop in pH, [CO32−] and Ωarag at the site of calcification, which correlates with decreased shell growth, and, eventually, shell dissolution. Our findings help explain why bivalve larvae can form shells under moderate acidification scenarios and provide a direct link between ocean carbonate chemistry and larval calcification rate.
    Type: Article , PeerReviewed , info:eu-repo/semantics/article
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  • 6
    Publication Date: 2021-03-30
    Description: Coastal global oceans are expected to undergo drastic changes driven by climate change and increasing anthropogenic pressures in coming decades. Predicting specific future conditions and assessing the best management strategies to maintain ecosystem integrity and sustainable resource use are difficult, because of multiple interacting pressures, uncertain projections, and a lack of test cases for management. We argue that the Baltic Sea can serve as a time machine to study consequences and mitigation of future coastal perturbations, due to its unique combination of an early history of multistressor disturbance and ecosystem deterioration and early implementation of cross-border environmental management to address these problems. The Baltic Sea also stands out in providing a strong scientific foundation and accessibility to long-term data series that provide a unique opportunity to assess the efficacy of management actions to address the breakdown of ecosystem functions. Trend reversals such as the return of top predators, recovering fish stocks, and reduced input of nutrient and harmful substances could be achieved only by implementing an international, cooperative governance structure transcending its complex multistate policy setting, with integrated management of watershed and sea. The Baltic Sea also demonstrates how rapidly progressing global pressures, particularly warming of Baltic waters and the surrounding catchment area, can offset the efficacy of current management approaches. This situation calls for management that is (i) conservative to provide a buffer against regionally unmanageable global perturbations, (ii) adaptive to react to new management challenges, and, ultimately, (iii) multisectorial and integrative to address conflicts associated with economic trade-offs.
    Type: Article , PeerReviewed , info:eu-repo/semantics/article
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  • 7
    Publication Date: 2021-03-19
    Description: In coastal temperate regions such as the Baltic Sea, calcifying bivalves dominate benthic communities playing a vital ecological role in maintaining biodiversity and nutrient recycling. At low salinities, bivalves exhibit reduced growth and calcification rates which is thought to result from physiological constraints associated with osmotic stress. Calcification demands a considerable amount of energy in calcifying molluscs and estuarine habitats provide sub-optimal conditions for calcification due to low concentrations of calcification substrates and large variations in carbonate chemistry. Therefore, we hypothesize that slow growth rates in estuarine bivalves result from increased costs of calcification, rather than costs associated with osmotic stress. To investigate this, we estimated the cost of calcification for the first time in benthic bivalve life stages and the relative energy allocation to calcification in three Mytilus populations along the Baltic salinity gradient. Our results indicate that calcification rates are significantly reduced only in 6 psu populations compared to 11 and 16 psu populations, coinciding with ca. 2–3-fold higher calcification costs at low salinity and temperature. This suggests that reduced growth of Baltic Mytilus at low salinities results from increased calcification costs rather than osmotic stress related costs. We also reveal that shell growth (both calcification and shell organic production) demands 31–60% of available assimilated energy from food, which is significantly higher than previous estimates. Energetically expensive calcification represents a major constraint on growth of mytilids in the estuarine and coastal seas where warming, acidification and desalination are predicted over the next century.
    Type: Article , PeerReviewed , info:eu-repo/semantics/article
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  • 8
    Publication Date: 2019-09-23
    Description: Calanoid copepods and euphausiids are key components of marine zooplankton communities worldwide. Most euphausiids and several copepod species perform diel vertical migrations (DVMs) that contribute to the export of particulate and dissolved matter to midwater depths. In vast areas of the global ocean, and in particular in the eastern tropical Atlantic and Pacific, the daytime distribution depth of many migrating organisms corresponds to the core of the oxygen minimum zone (OMZ). At depth, the animals experience reduced temperature and oxygen partial pressure (pO2) and an increased carbon dioxide partial pressure (pCO2) compared to their near-surface nighttime habitat. Although it is well known that low oxygen levels can inhibit respiratory activity, the respiration response of tropical copepods and euphausiids to relevant pCO2, pO2 and temperature conditions remains poorly parameterized. Further, the regulation of ammonium excretion at OMZ conditions is generally not well understood. It was recently estimated that DVM-mediated ammonium supply considerably fuels bacterial anaerobic ammonium oxidation – a major loss process for fixed nitrogen in the ocean. These estimates were based on the implicit assumption that hypoxia or anoxia in combination with hypercapnia (elevated pCO2) does not result in a downregulation of ammonium excretion. Here we show that exposure to OMZ conditions can result in strong depression of respiration and ammonium excretion in calanoid copepods and euphausiids from the Eastern Tropical North Atlantic and the Eastern Tropical South Pacific. These physiological responses need to be taken into account when estimating DVM-mediated fluxes of carbon and nitrogen into OMZs.
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  • 9
    Publication Date: 2020-06-18
    Description: Carbon capture and storage is promoted as a mitigation method counteracting the increase of atmospheric CO2 levels. However, at this stage, environmental consequences of potential CO2 leakage from sub-seabed storage sites are still largely unknown. In a 3-month-long mesocosm experiment, this study assessed the impact of elevated pCO2 levels (1,500 to 24,400 μatm) on Cerastoderma edule dominated benthic communities from the Baltic Sea. Mortality of C. edule was significantly increased in the highest treatment (24,400 μatm) and exceeded 50%. Furthermore, mortality of small size classes (0–1 cm) was significantly increased in treatment levels ≥6,600 μatm. First signs of external shell dissolution became visible at ≥1,500 μatm, holes were observed at 〉6,600 μatm. C. edule body condition decreased significantly at all treatment levels (1,500–24,400 μatm). Dominant meiofauna taxa remained unaffected in abundance. Densities of calcifying meiofauna taxa (i.e. Gastropoda and Ostracoda) decreased in high CO2 treatments (〉6,600 μatm), while the non - calcifying Gastrotricha significantly increased in abundance at 24,400 μatm. In addition, microbial community composition was altered at the highest pCO2 level. We conclude that strong CO2 leakage can alter benthic infauna community composition at multiple trophic levels, likely due to high mortality of the dominant macrofauna species C. edule.
    Type: Article , PeerReviewed , info:eu-repo/semantics/article
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  • 10
    Publication Date: 2019-09-23
    Description: Larval stages are considered as the weakest link when a species is exposed to challenging environmental changes1, 2. Reduced rates of growth and development in larval stages of calcifying invertebrates in response to ocean acidification might be caused by energetic limitations3. So far no information exists on how ocean acidification affects digestive processes in marine larval stages. Here we reveal alkaline (~pH 9.5) conditions in the stomach of sea urchin larvae. Larvae exposed to decreased seawater pH suffer from a drop in gastric pH, which directly translates into decreased digestive efficiencies and triggers compensatory feeding. These results suggest that larval digestion represents a critical process in the context of ocean acidification, which has been overlooked so far.
    Type: Article , PeerReviewed , info:eu-repo/semantics/article
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