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Seawater carbonate chemistry and carbonate load of seagrass leaves

Seagrass meadows play a significant role in the formation of carbonate sediments, serving as a substrate for carbonate-producing epiphyte communities. The magnitude of the epiphyte load depends on plant structural and physiological parameters, related to the time available for epiphyte colonization. Yet, the carbonate accumulation is likely to also depend on the carbonate saturation state of seawater (Omega) that tends to decrease as latitude increases due to decreasing temperature and salinity. A decrease in carbonate accumulation with increasing latitude has already been demonstrated for other carbonate producing communities. The aim of this study was to assess whether there was any correlation between latitude and the epiphyte carbonate load and net carbonate production rate on seagrass leaves. Shoots from 8 different meadows of the Zostera genus distributed across a broad latitudinal range (27 °S to up to 64 °N) were sampled along with measurements of temperature and Omega. The Omega within meadows significantly decreased as latitude increased and temperature decreased. The mean carbonate content and load on seagrass leaves ranged from 17 % DW to 36 % DW and 0.4-2.3 mg CO3/cm**2, respectively, and the associated mean carbonate net production rate varied from 0.007 to 0.9 mg CO3/cm**2/d. Mean carbonate load and net production rates decreased from subtropical and tropical, warmer regions towards subpolar latitudes, consistent with the decrease in Omega. These results point to a latitudinal variation in the contribution of seagrass to the accumulation of carbonates in their sediments which affect important processes occurring in seagrass meadows, such as nutrient cycling, carbon sequestration and sediment accretion.

Seawater carbonate chemistry and in situ and laboratory measurements of calcification

Ocean acidification (OA) is generally assumed to negatively impact calcification rates of marine organisms. At a local scale however, biological activity of macrophytes may generate pH fluctuations with rates of change that are orders of magnitude larger than the long-term trend predicted for the open ocean. These fluctuations may in turn impact benthic calcifiers in the vicinity. Combining laboratory, mesocosm and field studies, such interactions between OA, the brown alga Fucus vesiculosus, the sea grass Zostera marina and the blue mussel Mytilus edulis were investigated at spatial scales from decimetres to 100s of meters in the western Baltic. Macrophytes increased the overall mean pH of the habitat by up to 0.3 units relative to macrophyte- free, but otherwise similar, habitats and imposed diurnal pH fluctuations with amplitudes ranging from 0.3 to more than 1 pH unit. These amplitudes and their impact on mussel calcification tended to increase with increasing macrophyte biomass to bulk water ratio. At the laboratory and mesocosm scales, biogenic pH fluc- tuations allowed mussels to maintain calcification even under acidified conditions by shifting most of their calcification activity into the daytime when biogenic fluctuations caused by macrophyte activity offered temporal refuge from OA stress. In natural habitats with a low biomass to water body ratio, the impact of biogenic pH fluctuations on mean calcification rates of M. edulis was less pronounced. Thus, in dense algae or seagrass habitats, macrophytes may mitigate OA impact on mussel calcification by raising mean pH and providing temporal refuge from acidification stress.

Seawater carbonate chemistry and growth and carbon metabolism of the seaweed Fucus vesiculosus in the western Baltic Sea

Warming and acidification of the oceans as a consequence of increasing CO2-concentrations occur at large scales. Numerous studies have shown the impact of single stressors on individual species. However, studies on the combined effect of multiple stressors on a multi-species assemblage, which is ecologically much more realistic and relevant, are still scarce. Therefore, we orthogonally crossed the two factors warming and acidification in mesocosm experiments and studied their single and combined impact on the brown alga Fucus vesiculosus associated with its natural community (epiphytes and mesograzers) in the Baltic Sea in all seasons (from April 2013 to April 2014). We superimposed our treatment factors onto the natural fluctuations of all environmental variables present in the Benthocosms in so-called delta-treatments. Thereby we compared the physiological responses of F. vesiculosus (growth and metabolites) to the single and combined effects of natural Kiel Fjord temperatures and pCO2 conditions with a 5 °C temperature increase and/or pCO2 increase treatment (1100 ppm in the headspace above the mesocosms). Responses were also related to the factor photoperiod which changes over the course of the year. Our results demonstrate complex seasonal pattern. Elevated pCO2 positively affected growth of F. vesiculosus alone and/or interactively with warming. The response direction (additive, synergistic or antagonistic), however, depended on season and daylength. The effects were most obvious when plants were actively growing during spring and early summer. Our study revealed for the first time that it is crucial to always consider the impact of variable environmental conditions throughout all seasons. In summary, our study indicates that in future F. vesiculosus will be more affected by detrimental summer heat-waves than by ocean acidification although the latter consequently enhances growth throughout the year. The mainly negative influence of rising temperatures on the physiology of this keystone macroalga may alter and/or hamper its ecological functions in the shallow coastal ecosystem of the Baltic Sea.

Seawater carbonate chemistry and gastric pH homeostasis and larval recruitment in the sea star Asterias rubens

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.

Langzeitenwicklung und Populationsdynamik des Makrozoobenthos im Vorstrand der Insel Norderney

Seit 1977 werden in Wassertiefen zwischen 3 und 20 m in monatlichen Abstaenden die ansaessigen benthischen Lebensgemeinschaften sowie die Populationsdynamik ausgewaehlter Arten untersucht. Parallel dazu laufen sedimentologische sowie physikalisch-chemische Messungen.

Seawater carbonate chemistry and biometry and dissolution features of the benthic foraminifer Ammonia aomoriensis in a laboratory experiment

Culturing experiments were performed with the benthic foraminifer Ammonia aomoriensis from Flensburg Fjord, western Baltic Sea. The experiments simulated a projected rise in atmospheric CO2 concentrations. We exposed specimens to 5 seawater pCO2 levels ranging from 618 µatm (pH 7.9) to 3130 µatm (pH 7.2) for 6 wk. Growth rates and mortality differed significantly among pCO2 treatments. The highest increase of mean test diameter (19%) was observed at 618 µatm. At partial pressures >1829 µatm, the mean test diameter was observed to decrease, by up to 22% at 3130 µatm. At pCO2 levels of 618 and 751 µatm, A. aomoriensis tests were found intact after the experiment. The outer chambers of specimens incubated at 929 and 1829 µatm were severely damaged by corrosion. Visual inspection of specimens incubated at 3130 µatm revealed wall dissolution of all outer chambers, only their inner organic lining stayed intact. Our results demonstrate that pCO2 values of >=929 µatm in Baltic Sea waters cause reduced growth of A. aomoriensis and lead to shell dissolution. The bottom waters in Flensburg Fjord and adjacent areas regularly experience pCO2 levels in this range during summer and fall. Increasing atmospheric CO2 concentrations are likely to extend and intensify these periods of undersaturation. This may eventually slow down calcification in A. aomoriensis to the extent that net carbonate precipitation terminates. The possible disappearance of this species from the Baltic Sea and other areas prone to seasonal undersaturation would likely cause significant shifts in shallow-water benthic ecosystems in the near future.

Seawater carbonate chemistry and seasonal variations of Fucus vesiculosus fertility in the western Baltic Sea

Ocean warming and acidification may substantially affect the reproduction of keystone species such as Fucus vesiculosus (Phaeophyceae). In four consecutive benthic mesocosm experiments, we compared the reproductive biology and quantified the temporal development of Baltic Sea Fucus fertility under the single and combined impact of elevated seawater temperature and pCO2 (1100 ppm). In an additional experiment, we investigated the impact of temperature (0–25°C) on the maturation of North Sea F. vesiculosus receptacles. A marked seasonal reproductive cycle of F. vesiculosus became apparent in the course of 1 year. The first appearance of receptacles on vegetative apices and the further development of immature receptacles of F. vesiculosus in autumn were unaffected by warming or elevated pCO2. During winter, elevated pCO2 in both ambient and warmed temperatures increased the proportion of mature receptacles significantly. In spring, warming and, to a lesser extent, elevated pCO2 accelerated the maturation of receptacles and advanced the release of gametes by up to 2 weeks. Likewise, in the laboratory, maturation and gamete release were accelerated at 15–25°C relative to colder temperatures. In summary, elevated pCO2 and/or warming do not influence receptacle appearance in autumn, but do accelerate the maturation process during spring, resulting in earlier gamete release. Temperature and, to a much lesser extent, pCO2 affect the temporal development of Fucus fertility. Thus, rising temperatures will mainly shift or disturb the phenology of F. vesiculosus in spring and summer, which may alter and/or hamper its ecological functions in shallow coastal ecosystems of the Baltic Sea.

Seawater carbonate chemistry and skeletal integrity of a marine keystone predator (Asterias rubens)

The current increase in atmospheric CO2 concentration induces changes in the seawater carbonate system resulting in decreased pH and calcium carbonate saturation state, a phenomenon called ocean acidification (OA). OA has long been considered as a major threat to echinoderms because their extensive endoskeleton is made of high‑magnesium calcite, one of the most soluble forms of calcium carbonate. Numerous studies addressed this question in sea urchins, but very few questioned the impact of OA on the sea star skeleton, although members of this taxon do not compensate their extracellular pH, contrary to most sea urchins. In the present study, adults of the common sea star, Asterias rubens from Kiel Fjord, a site experiencing natural acidification events exceeding pCO2 levels of 2500 μatm, were chronically exposed to different levels of simulated ocean acidification (pHT-SW 8.0, 7.4, 7.2), encompassing present and future conditions, for the duration of 109 days. Corrosion and mechanical properties of skeletal elements were studied using scanning electron microscopy, three-point bending tests as well as nanoindentation. The spines were significantly corroded at pHT-SW 7.4 and below while the ambulacral plates were only affected at pHT-SW 7.2. Nanoindentation of newly formed spines and ambulacral plates did not reveal significant CO2-induced differences in skeleton hardness or elasticity across treatments. Results of three-point bending tests revealed significantly reduced characteristic strength and fracture force of ambulacral plates from the median arm segment at pHT-SW 7.4 and below. These plates are those supporting the tube feet involved in the opening of bivalves during feeding and in the animal attachment to the substrate. Under reduced seawater pH, this might result in fracture of sea star plates during predation on mussel. The present results predict a possible impact of ocean acidification on the skeletal integrity of a marine keystone predator.

Seawater carbonate chemistry and growth, survival and Chlorophyll a fluorescence parameters of Fucus vesiculosus L.(Phaeophyceae) in a seasonally fluctuating environment

Global change exposes brown algal Fucus vesiculosus populations to increasing temperature and pCO2, which may threaten individuals, in particular the early life-stages. Genetic diversity of F. vesiculosus populations is low in the Baltic compared to Atlantic populations. This might jeopardise their potential for adaptation to environmental changes. Here, we report on the responses of early life-stage F. vesiculosus to warming and acidification in a near-natural scenario maintaining natural and seasonal variation (spring 2013–2014) of the Kiel Fjord in the Baltic Sea, Germany (54°27ʹN, 10°11ʹW). We assessed how stress sensitivity differed among sibling groups and how genetic diversity of germling populations affected their stress tolerance. Warming increased growth rates of Fucus germlings in spring and in early summer, but led to higher photoinhibition in spring and decreased their survival in late summer. Acidification increased germlings' growth in summer but otherwise showed much weaker effects than warming. During the colder seasons (autumn and winter), growth was slow while survival was high compared to spring and summer, all at ambient temperatures. A pronounced variation in stress response among genetically different sibling groups (full-sib families) suggests a genotypic basis for this variation and thus a potential for adaptation for F. vesiculosus populations to future conditions. Corroborating this, survival in response to warming in populations with higher diversity was better than the mean survival of single sibling groups. We conclude that impacts on early life-stages depend on the combination of stressors and season and that genetic variation is crucial for the tolerance to global change stress.

Seawater carbonate chemistry and benthic foraminifera Ammonia sp. uranium incorporation during experiments, 2013

The chemical and isotopic composition of foraminiferal shells (so-called proxies) reflects the physico-chemical properties of the seawater. In current day paleoclimate research, the reconstruction of past seawater carbonate system to infer atmospheric CO2 concentrations is one of the most pressing challenges and a variety of proxies have been investigated, such as foraminiferal U/Ca. Since in natural seawater and traditional CO2 perturbation experiments, the carbonate system parameters co-vary, it is not possible to determine the parameter of the carbonate system causing e.g. changes in U/Ca, complicating the use of the latter as a carbonate system proxy. We overcome this problem, by culturing the benthic foraminifer Ammonia sp. at a range of carbonate chemistry manipulation treatments. Shell U/Ca values were determined to test sensitivity of U incorporation to various parameters of the carbonate system. We argue that CO3 is the parameter affecting the U/Ca ratio and consequently, the partitioning coefficient for U in Ammonia sp DU. We can confirm the strong potential of foraminiferal U/Ca as a CO3 proxy.

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