The calcareous tubeworm Spirorbis spirorbis is a widespread serpulid species in the Baltic Sea, where it commonly grows as an epibiont on brown macroalgae (genus Fucus). It lives within a Mg-calcite shell and could be affected by ocean acidification and temperature rise induced by the predicted future atmospheric CO2 increase. However, Spirorbis tubes grow in a chemically modified boundary layer around the algae, which may mitigate acidification. In order to investigate how increasing temperature and rising pCO2 may influence S. spirorbisshell growth we carried out four seasonal experiments in the Kiel Outdoor Benthocosms at elevated pCO2 and temperature conditions. Compared to laboratory batch culture experiments the benthocosm approach provides a better representation of natural conditions for physical and biological ecosystem parameters, including seasonal variations. We find that growth rates of S. spirorbis are significantly controlled by ontogenetic and seasonal effects. The length of the newly grown tube is inversely related to the initial diameter of the shell. Our study showed no significant difference of the growth rates between ambient atmospheric and elevated (1100 ppm) pCO2 conditions. No influence of daily average CaCO3 saturation state on the growth rates of S. spirorbis was observed. We found, however, net growth of the shells even in temporarily undersaturated bulk solutions, under conditions that concurrently favoured selective shell surface dissolution. The results suggest an overall resistance of S. spirorbis growth to acidification levels predicted for the year 2100 in the Baltic Sea. In contrast, S. spirorbis did not survive at mean seasonal temperatures exceeding 24 °C during the summer experiments. In the autumn experiments at ambient pCO2, the growth rates of juvenile S. spirorbis were higher under elevated temperature conditions. The results reveal that S. spirorbis may prefer moderately warmer conditions during their early life stages but will suffer from an excessive temperature increase and from increasing shell corrosion as a consequence of progressing ocean acidification.
Diatoms account for up to 40% of marine primary production and require silicic acid to grow and build their opal shell. On the physiological and ecological level, diatoms are thought to be resistant to, or even benefit from, ocean acidification. Yet, global-scale responses and implications for biogeochemical cycles in the future ocean remain largely unknown. Here we conducted five in situ mesocosm experiments with natural plankton communities in different biomes and find that ocean acidification increases the elemental ratio of silicon (Si) to nitrogen (N) of sinking biogenic matter by 17 ± 6 per cent under pCO2 conditions projected for the year 2100. This shift in Si:N seems to be caused by slower chemical dissolution of silica at decreasing seawater pH. We test this finding with global sediment trap data, which confirm a widespread influence of pH on Si:N in the oceanic water column. Earth system model simulations show that a future pH-driven decrease in silica dissolution of sinking material reduces the availability of silicic acid in the surface ocean, triggering a global decline of diatoms by 13–26 per cent due to ocean acidification by the year 2200. This outcome contrasts sharply with the conclusions of previous experimental studies, thereby illustrating how our current understanding of biological impacts of ocean change can be considerably altered at the global scale through unexpected feedback mechanisms in the Earth system.
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.
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.
Shallow coastal marine ecosystems are exposed to intensive warming events in the last decade, threatening keystone macroalgal species such as the bladder wrack (Fucus vesiculosus, Phaeophyceae) in the Baltic Sea. Herein, we experimentally tested in four consecutive benthic mesocosm experiments, if the single and combined impact of elevated seawater temperature (+ 5◦C) and pCO2 (1100 ppm) under natural irradiance conditions seasonally affected the photophysiological performance (i.e., oxygen production, in vivo chlorophyll a fluorescence, energy dissipation pathways and chlorophyll concentration) of Baltic Sea Fucus. Photosynthesis was highest in spring/early summer when water temperature and solar irradiance increases naturally, and was lowest in winter (December to January/February). Temperature had a stronger effect than pCO2 on photosynthetic performance of Fucus in all seasons. In contrast to the expectation that warmer winter conditions might be beneficial, elevated temperature conditions and sub-optimal low winter light conditions decreased photophysiological performance of Fucus. In summer, western Baltic Sea Fucus already lives close to its upper thermal tolerance limit and future warming of the Baltic Sea during summer may probably become deleterious for this species. However, our results indicate that over most of the year a combination of future ocean warming and increased pCO2 will have slightly positive effects for Fucus photophysiological performance.
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.
We studied the response of the heterotrophic flagellate (HF) community to the combined impact of warming and ocean acidification in a mesocosm experiment with a plankton community from the western Baltic Sea. We performed a quantitative analysis of the response at the level of total biomass and size classes and a semi-quantitative one at the level of individual taxa. Total biomass of HF was significantly lower under higher temperatures while there was no significant effect of CO2. The mean biomass of the picoflagellates did not respond to temperature while the three nanoflagellate size classes (class limits 3, 5, 8, 15 ?m) responded negatively to warming while not responding to CO2. The taxon-level results indicate that heterotrophic flagellates do not form a homogenous trophic guild, as often assumed in pelagic food web studies. Instead, the heterotrophic flagellates formed a ?food web within the food web?. There was a pronounced succession of flagellates leading from a dominance of bacterivores and colloidal matter feeders before the phytoplankton bloom to omnivorous feeders preying upon phytoplankton and heterotrophic flagellates during and after the bloom. This complex intraguild predation patterns probably dampened the response to experimental treatments.
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