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Zeitliche Variabilität von internen Wellen und vertikaler Vermischung im Nordatlantik

Das Ziel dieses Projektes ist die Untersuchung der zeitlichen Variabilität in der Energie von internen Wellen und der Stärke von vertikaler Vermischung in Abhängigkeit des Nordatlantikstroms und dem damit verbundenen Wirbelfeld. Hierfür werden 5-6 Jahre von Strömungsmesserdaten und Temperatur/Leitfähigkeitsmessungen von drei Verankerungen entlang eines Schnittes westlich des Mittelatlantischen Rückens (MAR) sowie LADCP/CTD Daten von fünf Schifffahrten genutzt. Konkrete wissenschaftliche Ziele dabei sind:- Erstellung von Zeitserien der Energie in internen Wellen unter Benutzung der Verankerungszeitreihen von Strömung und Schichtung- Untersuchung der Zeitskalen auf denen Veränderungen in der Energie interner Wellen stattfinden. Mögliche Ursachen für Variabilität sind der Windeintrag, Position des Nordatlantikstroms und Wirbel- Identifizierung von Prozessen, welche die beobachteten internen Wellen generieren, wie z.B. Gezeiten, Stürme, Jahresgang, Wirbel, die Arme des Nordatlanikstroms (Verhältnis von lokalen zu großräumigen Erzeugungsmechanismen)- Bestimmung der Vermischungsraten (Temperaturinversionen, Thorpe Skalen, Feinstrukturparameterisierung) in Abhängigkeit der variablen Hintergrundbedingungen Hierfür werden zunächst Spektren potentieller und kinetischer Energy der internen Wellen auf ihre Abhängigkeit von veränderlichen Hintergrundbedingungen wie z.B. Wind, Gezeiten, Wirbel, Schichtung und Variabilität im Nordatlantikstrom sowieso des Einflusses der Topographie untersucht. Die instrumentelle Ausstattung der Verankerungen seit Sommer 2012 erlaubt zusätzlich die Approximation der internen Wellen durch vertikale Moden und damit verbunden die Berechnung von Energieflüssen, welche wichtige Informationen über die Menge und die Variabilität in der Energie, die in internen Wellen im Nordatlantik transportiert wird, liefern. Außerdem geben diese so gewonnenen Energieflüsse in Kombination mit der Berechnung von Ausbreitungspfaden von internen Wellen, welche am mittelatlantischen Rücken erzeugt wurden, Aufschluss über die relative Bedeutung der Topographie des MAR für die Erzeugung von internen Wellen. Beginnend vom Sommer 2015 werden die Analysen erweitert, indem Temperatur- und Druckdaten mit hoher Tiefenauflösung für die Berechnung von Thorpe Skalen und Dissipationsraten und deren zeitlichen Variabilität genutzt werden. Weitere Informationen über die zeitliche und räumliche Variabilität der Vermischungsraten im Nordatlantik werden durch die Analyse von Diffusionsraten, die anhand von LADCP/CTD Daten und einer Feinstrukturparameterisierung berechnet werden, erlangt. Dies liefert weitere Aufschlüsse über die dominanten Prozesse in der Erzeugung von internen Wellen und vertikaler Vermischung im Nordatlantik, sowie deren zeitlicher und räumlicher Variabilität.

Seawater carbonate chemistry and benthic foraminifera Ammonia sp. mass, size, and growth rate during experiments, 2013

About 30% of the anthropogenically released CO2 is taken up by the oceans; such uptake causes surface ocean pH to decrease and is commonly referred to as ocean acidification (OA). Foraminifera are one of the most abundant groups of marine calcifiers, estimated to precipitate ca. 50 % of biogenic calcium carbonate in the open oceans. We have compiled the state of the art literature on OA effects on foraminifera, because the majority of OA research on this group was published within the last three years. Disparate responses of this important group of marine calcifiers to OA were reported, highlighting the importance of a process-based understanding of OA effects on foraminifera. We cultured the benthic foraminifer Ammonia sp. under a range of carbonate chemistry manipulation treatments to identify the parameter of the carbonate system causing the observed effects. This parameter identification is the first step towards a process-based understanding. We argue that CO3 is the parameter affecting foraminiferal size-normalized weights (SNWs) and growth rates. Based on the presented data, we can confirm the strong potential of Ammonia sp. foraminiferal SNW as a CO3 proxy.

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 growth of calcifying tubeworm shells (Spirorbis spirorbis)

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.

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.

Seawater carbonate chemistry and physiological performance parameters of Carcinus maenas under respective incubation conditions

Ocean acidification causes an accumulation of CO2 in marine organisms and leads to shifts in acid-base parameters. Acid-base regulation in gill breathers involves a net increase of internal bicarbonate levels through transmembrane ion exchange with the surrounding water. Successful maintenance of body fluid pH depends on the functional capacity of ion-exchange mechanisms and associated energy budget. For a detailed understanding of the dependence of acid-base regulation on water parameters, we investigated the physiological responses of the shore crab Carcinus maenas to 4 weeks of ocean acidification [OA, P(CO2)w = 1800 µatm], at variable water bicarbonate levels, paralleled by changes in water pH. Cardiovascular performance was determined together with extra-(pHe) and intracellular pH (pHi), oxygen consumption, haemolymph CO2 parameters, and ion composition. High water P(CO2) caused haemolymph P(CO2) to rise, but pHe and pHi remained constant due to increased haemolymph and cellular [HCO3-]. This process was effective even under reduced seawater pH and bicarbonate concentrations. While extracellular cation concentrations increased throughout, anion levels remained constant or decreased. Despite similar levels of haemolymph pH and ion concentrations under OA, metabolic rates, and haemolymph flow were significantly depressed by 40 and 30%, respectively, when OA was combined with reduced seawater [HCO3-] and pH. Our findings suggest an influence of water bicarbonate levels on metabolic rates as well as on correlations between blood flow and pHe. This previously unknown phenomenon should direct attention to pathways of acid-base regulation and their potential feedback on whole-animal energy demand, in relation with changing seawater carbonate parameters.

Seawater carbonate chemistry and mass fluxes and elemental composition of particulate export in KOSMOS mesocosm experiments (2010-2014)

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.

Seawater carbonate chemistry and PI-curve parameters, non-photochemical quenching, maximum quantum yield, gross oxygen production and chlorophyll concentrations of Fucus vesiculosus in the Kiel Outdoor Benthocosms

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.

The influence of seawater pH on U/ Ca ratios in the scleractinian cold-water coral Lophelia pertusa

The increasing pCO2 in seawater is a serious threat for marine calcifiers and alters the biogeochemistry of the ocean. Therefore, the reconstruction of past-seawater properties and their impact on marine ecosystems is an important way to investigate the underlying mechanisms and to better constrain the effects of possible changes in the future ocean. Cold-water coral (CWC) ecosystems are biodiversity hotspots. Living close to aragonite undersaturation, these corals serve as living laboratories as well as archives to reconstruct the boundary conditions of their calcification under the carbonate system of the ocean. We investigated the reef-building CWC Lophelia pertusa as a recorder of intermediate ocean seawater pH. This species-specific field calibration is based on a unique sample set of live in situ collected L. pertusa and corresponding seawater samples. These data demonstrate that uranium speciation and skeletal incorporation for azooxanthellate scleractinian CWCs is pH dependent and can be reconstructed with an uncertainty of ±0.15. Our Lophelia U / Ca-pH calibration appears to be controlled by the high pH values and thus highlighting the need for future coral and seawater sampling to refine this relationship. However, this study recommends L. pertusa as a new archive for the reconstruction of intermediate water mass pH and hence may help to constrain tipping points for ecosystem dynamics and evolutionary characteristics in a changing ocean.

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.

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