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The exposure of organisms to multiple stressors can lead to special effects, which result e.g. from interactions between stressors, from interactions with the organisms or from temporal patterns of stress intensities. In my thesis, I propose a concept to quantify such kinds of cumulative effects by the utilization of literature data from experiments. Further, I compare the results with a summation of the effects of each of the occurring stressors. The concept comprises a network model, a Dynamic Energy Budget Model, and a matrix model. The application of the concept for modeling cumulative effects of anthropogenic pressures on Mytilus edulis and on seagrass habitats shows that cumulative effects can have a significant effect on ecosystem components and that the magnitude of a cumulative effect can change over time. The concept is applicable for environmental assessments such as needed for the Marine Strategy Framework Directive.

Every year billions of vertebrates travel over considerable distances. In this thesis, I presented a comprehensive overview on the theories on orientation and on the compasses and map senses used in vertebrate orientation. My own experiments aimed to provide new insights on the compass systems used by two vertebrate species with very distinctive orientational needs - migratory songbirds and coral reef fish larvae. In coral reef fish, I demonstrated that O. doederleini larvae possess a magnetic compass that might help them return to their natal reefs. In my main projects, I described why an invasive electrophysiological approach was unfit to unequivocally demonstrate polarization sensitivity in domestic chicken and in three songbird species, I was able to experimentally suggest the inability of birds to detect polarized light information. In the outlook of this thesis, I presented a so far undescribed novel optical property of the third eye lid in birds. I elaborated the potential implications of my discovery for a completely novel view on polarized light sensitivity in birds.

Settlement of pelagic coral larvae is commonly induced by chemical cues that originate from biofilms and coralline algae. These natural settlement cues initiate signal pathways leading to attachment and metamorphosis of the coral larva. In order to investigate the settlement process and its natural inducers, it is necessary to gain a better understanding of these signal pathways. At present, the pathways and neurotransmitters involved in this signal transduction are still widely unknown. In this study, we exposed larvae of the brooding coral Leptastrea purpurea to five neuroactive compounds known to be present in cnidarians, and K+ Ions. All compounds were applied at different dilutions and settlement behavior of the larvae was documented over 48 h. Dopamine, glutamic acid and epinephrine significantly induced settlement in the coral larvae. The highest observed metamorphosis response was 54% in 10−5 M dopamine. Serotonin, L-DOPA and K+ ions did not have an influence on settlement behavior in our experiments. Exposing larvae to settlement-inducing neurotransmitters and thus bypassing the initial induction could be utilized in coral aquaculture. The active neurotransmitters should be used to further study the settlement process in L. purpurea in greater detail. Their role and relevance should also be assessed for other coral species as they may represent or reveal a universal inducer for coral settlement.

Gas exchange across the air-water interface is strongly influenced by the uppermost water layer (

Plant performance is correlated with element concentrations in plant tissue, which may be impacted by adverse chemical soil conditions. Antibiotics of veterinary origin can adversely affect plant performance. They are released to agricultural fields via grazing animals or manure, taken up by plants and may be stored, transformed or sequestered by plant metabolic processes. We studied the potential effects of three antibiotics (penicillin, sulfadiazine, and tetracycline) on plant element contents (macro- and microelements). Plant species included two herb species (Brassica napus and Capsella bursa-pastoris) and two grass species (Triticum aestivum and Apera spica-venti), representing two crop species and two noncrop species commonly found in field margins, respectively. Antibiotic concentrations were chosen as to reflect in vivo situations, that is, relatively low concentrations similar to those detected in soils. In a greenhouse experiment, plants were raised in soil spiked with antibiotics. After harvest, macro- and microelements in plant leaves, stems, and roots were determined (mg/g). Results indicate that antibiotics can affect element contents in plants. Penicillin exerted the greatest effect both on element contents and on scaling relationships of elements between plant organs. Roots responded strongest to antibiotics compared to stems and leaves. We conclude that antibiotics in the soil, even in low concentrations, lead to low-element homeostasis, altering the scaling relationships between roots and other plant organs, which may affect metabolic processes and ultimately the performance of a plant.

The copepod Calanus finmarchicus has an ecological key position in the northern Atlantic pelagic food web and its life is characterized by diel and seasonal rhythmicity. Neither diel nor seasonal rhythmicity of C. finmarchicus are understood with regard to their mechanistic regulation. Endogenous clock systems are central in controlling rhythms in various terrestrial species, but have hardly been investigated in marine organisms. This thesis shows that C. finmarchicus possesses an endogenous circadian clock, that regulated 24h rhythms of gene expression, metabolic activity and vertical migration behavior. The thesis further suggests that clock-based day length measurement and an endogenous annual clock is involved in the regulation of seasonal rhythmicity. The findings on C. finmarchicus’ timing systems are further related to the extreme light conditions in polar environments, discussing potential effects of climate chance on the copepods rhythmicity and biology.

Coral reefs are experiencing increasing anthropogenic impacts that result in substantial declines of reef-building corals and a change of community structure towards other benthic invertebrates or macroalgae. Reefs around Zanzibar are exposed to untreated sewage and runoff from the main city Stonetown. At many of these sites, sponge cover has increased over the last years. Sponges are one of the top spatial competitors on reefs worldwide. Their success is, in part, dependent on their strong chemical defenses against predators, microbial attacks and other sessile benthic competitors. This is the first study that investigates the bioactive properties of sponge species in the Western Indian Ocean region. Crude extracts of the ten most dominant sponge species were assessed for their chemical defenses against 35 bacterial strains (nine known as marine pathogens) using disc diffusion assays and general cytotoxic activities were assessed with brine shrimp lethality assays. The three chemically most active sponge species were additionally tested for their allelopathic properties against the scleractinian coral competitor Porites sp.. The antimicrobial assays revealed that all tested sponge extracts had strong antimicrobial properties and that the majority (80%) of the tested sponges were equally defended against pathogenic and environmental bacterial strains. Additionally, seven out of ten sponge species exhibited cytotoxic activities in the brine shrimp assay. Moreover, we could also show that the three most bioactive sponge species were able to decrease the photosynthetic performance of the coral symbionts and thus were likely to impair the coral physiology.

Dissolved organic sulfur (DOS) is the largest pool of organic sulfur in the oceans, and as such it is an important component of the global sulfur cycle. DOS in the ocean is resistant against microbial degradation and turns over on a millennium time scale. However, sources and mechanisms behind its stability are largely unknown. Here, we hypothesize that in sulfate-reducing sediments sulfur is abiotically incorporated into dissolved organic matter (DOM) and released to the ocean. We exposed natural seawater and the filtrate of a plankton culture to sulfidic conditions. Already after 1-h at 20°C, DOS concentrations had increased 4-fold in these experiments, and 14-fold after 4 weeks at 50°C, indicating that organic matter does not need long residence times in natural sulfidic environments to be affected by sulfurization. Molecular analysis via ultrahigh-resolution mass spectrometry showed that sulfur was covalently and unselectively bound to DOM. Experimentally produced and natural DOS from sediments were highly similar on a molecular and structural level. By combining our data with published benthic DOC fluxes we estimate that 30–200 Tg DOS are annually transported from anaerobic and sulfate reducing sediments to the oceans. Uncertainties in this first speculative assessment are large. However, this first attempt illustrates that benthic DOS flux is potentially one order of magnitude larger than that via rivers indicating that this could balance the estimated global net removal of refractory DOS.

Heterotrophic bacterioplankton communities play an important role in organic matter processing in the oceans worldwide. In order to investigate the significance of distinct phylogenetic bacterial groups it is not only important to assess their quantitative abundance but also their growth dynamics in relation to the entire bacterioplankton. Therefore bacterial abundance, biomass production and the composition of the entire and cell-proliferating bacterioplankton community were assessed in North Sea surface waters between the German Bight and 58°N in early summer by applying catalyzed reporter deposition (CARD-FISH) and bromodeoxyuridine fluorescence in situ hybridization (BrdU-FISH). Bacteroidetes and the Roseobacter group dominated the cell-proliferating fraction with 10–55 and 8–31% of total BrdU-positive cells, respectively. While Bacteroidetes also showed high abundances in the total bacterial fraction, roseobacters constituted only 1–9% of all cells. Despite abundances of up to 55% of total bacterial cells, the SAR11 clade constituted

The periwinkle Littorina saxatilis has repeatedly evolved both a small, fragile and globose "wave ecotype" confined to wave-swept shores and a large, robust and elongated "crab ecotype" found in nearby crab-rich but less-exposed shores. This phenotypic divergence is assumed to reflect, in some part, local adaptation to wave exposure, but this hypothesis has received incomplete experimental testing. Here, we report a test of the prediction that the wave ecotype has a higher capacity to resist water flow than the crab ecotype. We sampled snails along a crab-wave transect and measured their resistance to dislodgement in a high-speed water flume with water speeds that match those of breaking waves in a range of relevant field conditions. Snails from the wave environment were consistently more resistant to water flow than snails from the crab environment and high resistance was positively correlated with the surface area of the foot and the area of the outer aperture contour both relative to shell size, and to the extent of lateral shell compression. In a separate experiment, we found that snails raised in still water in a common garden showed higher resistance to water flow if originating from a wave environment than from a crab environment, and this was true both at juvenile (2 weeks) and adult (10 months) developmental stages. This result suggests genetic control of a distinct "wave adapted" phenotype, likely to be maintained under strong divergent selection between the two adjacent habitats.

The effect-response framework states that plant functional traits link the abiotic environment to ecosystem functioning. One ecosystem property is the body size of the animals living in the system, which is assumed to depend on temperature or resource availability, among others. For primary consumers, resource availability may directly be related to plant traits, while for secondary consumers the relationship is indirect. We used plant traits to describe resource availability along an elevational gradient on Mount Kilimanjaro, Tanzania. Using structural equation models, we determined the response of plant traits to changes in precipitation, temperature and disturbance with and assessed whether abiotic conditions or community-weighted means of plant traits are stronger predictors of the mean size of bees, moths, frugivorous birds, and insectivorous birds. Traits indicating tissue density and nutrient content strongly responded to variations in precipitation, temperature and disturbance. They had direct effects on pollination and fruit traits. However, the average body sizes of the animal groups considered could only be explained by temperature and habitat structure, not by plant traits. Our results demonstrate a strong link between traits and the abiotic environment, but suggest that temperature is the most relevant predictor of mean animal body size. Community-weighted means of plant traits and body sizes appear unsuitable to capture the complexity of plant-animal interactions.

Annually recurring phytoplankton spring blooms are characteristic of temperate coastal shelf seas. During these blooms, environmental conditions, including nutrient availability, differ considerably from non-bloom conditions, affecting the entire ecosystem including the bacterioplankton. Accordingly, the emerging ecological niches during bloom transition are occupied by different bacterial populations, with Roseobacter RCA cluster and SAR92 clade members exhibiting high metabolic activity during bloom events. In this study, the functional response of the ambient bacterial community toward a Phaeocystis globosa bloom in the southern North Sea was studied using metaproteomic approaches. In contrast to other metaproteomic studies of marine bacterial communities, this is the first study comparing two different cell lysis and protein preparation methods [using trifluoroethanol (TFE) and in-solution digest as well as bead beating and SDS-based solubilization and in-gel digest (BB GeLC)]. In addition, two different mass spectrometric techniques (ESI-iontrap MS and MALDITOF MS) were used for peptide analysis. A total of 585 different proteins were identified, 296 of which were only detected using the TFE and 191 by the BB GeLC method, demonstrating the complementarity of these sample preparation methods. Furthermore, 158 proteins of the TFE cell lysis samples were exclusively detected by ESI-iontrap MS while 105 were only detected using MALDI-TOF MS, underpinning the value of using two different ionization and mass analysis methods. Notably, 12% of the detected proteins represent predicted integral membrane proteins, including the difficult to detect rhodopsin, indicating a considerable coverage of membrane proteins by this approach. This comprehensive approach verified previous metaproteomic studies of marine bacterioplankton, e.g., detection of many transport-related proteins (17% of the detected proteins). In addition, new insights into e.g., carbon and nitrogen metabolism were obtained. For instance, the C1 pathway was more prominent outside the bloom and different strategies for glucose metabolism seem to be applied under the studied conditions. Furthermore, a higher number of nitrogen assimilating proteins were present under non-bloom conditions, reflecting the competition for this limited macro nutrient under oligotrophic conditions. Overall, application of different sample preparation techniques as well as MS methods facilitated a more holistic picture of the marine bacterioplankton response to changing environmental conditions. peerReviewed

The sea-surface microlayer (SML) forms the uppermost boundary layer between atmosphere and ocean, and has distinctive physico-chemical and biological features compared to the underlying water. First findings on metabolic contributions of microorganisms to gas exchange processes across the SML raised the need for new in situ technologies to explore plankton-oxygen turnover in this special habitat. Here, we describe an inexpensive research tool, the Surface In Situ Incubator (SISI), which allows simultaneous incubations of the SML, and water samples from 1 m and 5 m, at the respective depths of origin. The SISI is deployed from a small boat, seaworthy up to 5 bft (Beaufort scale), and due to global positioning system (GPS) tracking, capable of drifting freely for hours or days. We tested the SISI by applying light/dark bottle incubations in the Baltic Sea and the tropical Pacific Ocean under various conditions to present first data on planktonic oxygen turnover rates within the SML, and two subsurface depths. The SISI offers the potential to study plankton-oxygen turnover within the SML under the natural influence of abiotic parameters, and hence, is a valuable tool to routinely monitor their physiological role in biogeochemical cycling and gas exchange processes at, and near, the sea surface.

In this thesis the competitive ability of two different potentially harmful dinoflagellates, Lingulodinium polyedrum and Alexandrium catenella, was investigated under different nutrient conditions and grazing control by zooplankton consumers. I particularly focused on the species-specific adaptive strategies mixotrophy and allelopathy. My results indicated that both strategies can be important for reducing competitors either through phagotrophic feeding or by lytic activity. Beside these benefits, both strategies also affected the cellular nutrient concentrations of the dinoflagellates, which indicated initial metabolic ‘costs’ for switching nutritional modes from photosynthesis to phagotrophic feeding, before having a benefit from ingested prey. Furthermore, the competitive success of L. polyedrum was potentially facilitated by selective zooplankton grazing on smaller prey in a more complex natural plankton community. My study emphasizes that bloom forming dinoflagellates may use very different strategies to become dominant in a plankton community, entailing different trophic interactions leading to altered food web dynamics.

Sponges (phylum Porifera) are important members of almost all aquatic ecosystems, and are renowned for hosting often dense and diverse microbial communities. While the specificity of the sponge microbiota seems to be closely related to host phylogeny, the environmental factors that could shape differences within local sponge-specific communities remain less understood. On tropical coral reefs, sponge habitats can span from shallow areas to deeper, mesophotic sites. These habitats differ in terms of environmental factors such as light, temperature, and food availability, as well as anthropogenic impact. In order to study the host specificity and potential influence of varying habitats on the sponge microbiota within a local area, four tropical reef sponges, Rhabdastrella globostellata, Callyspongia sp., Rhaphoxya sp., and Acanthella cavernosa, were collected from exposed shallow reef slopes and a deep reef drop-off. Based on 16S rRNA gene pyrosequencing profiles, beta diversity analyses revealed that each sponge species possessed a specific microbiota that was significantly different to those of the other species and exhibited attributes that are characteristic of high- and/or lowmicrobial- abundance sponges. These findings emphasize the influence of host identity on the associated microbiota. Dominant sponge- and seawater-associated bacterial phyla were Chloroflexi, Cyanobacteria, and Proteobacteria. Comparison of individual sponge taxa and seawater samples between shallow and deep reef sites revealed no significant variation in alpha diversity estimates, while differences in microbial beta diversity (variation in community composition) were significant for Callyspongia sp. sponges and seawater samples. Overall, the sponge-associated microbiota is significantly shaped by host identity across all samples, while the effect of habitat differentiation seems to be less predominant in tropical reef sponges.

The interplay between hydrodynamic motion and nutrient availability influences the distribution of plankton that makes up the basis of the aquatic food web. Horizontal stirring by mesoscale structures like vortices and wakes redistributes nutrients and enhances primary production, and can also initiate plankton blooms and affect the competition and coexistence of different plankton species. Vertical upwelling in connection with and Ekman flow and strong mesoscale activity occurs in the region the Canary Islands in the Atlantic ocean. The residence time of the plankton species in the structures in the wake is determined by the chaotic saddle, the vorticity and other nonlinear effects in the flow. The long residence times in the vicinity of the island and the hydrodynamic forcing and confinement of plankton in the vortices initiate localized plankton blooms.

Distinctive, patchy abundance patterns of spionid polychaetes i.e. Polydora cornuta and Streblospio benedicti with indirect larval development were found in sediments of tidal flats in the German Wadden Sea. These patterns were not explicable through grain size or other sediment parameters. This led to the hypotheses that active site selection and initial settlement of larvae were triggered by sediment associated microbial cues of i.e. biofilms. This thesis addressed the first time questions about different biofilm components e.g. diatoms, bacteria or their epipolysaccharides or halogenated metabolites as potential settlement cues for soft sediment settlers. In single-choice and multiple-choice assays this hypothesis was tested with larvae of P. cornuta and S. benedicti obtained from laboratory stock cultures which provide a sufficient amount of larvae. Larvae of both species were able to accept or reject sediments in different sediment qualities.

Marine macroalgae are often devoid of micro- and macro-colonizers, suggesting some sort of host-derived antifouling defense. Previous studies mainly focused on tissue extracts of entire algae against few bacterial isolates, thereby not only neglecting ecologically relevant bacteria but also natural delivery mechanisms of algal antimicrobial agents. In my study, I investigated antimicrobial effects of waterborne algal metabolites utilizing culture-independent techniques against planktonic bacteria and fungi in presence and absence of macroalgae. I also studied the inhibitory effects of macroalgae against diatoms. Results revealed that macroalgae affected planktonic bacteria qualitatively and quantitatively by waterborne algal metabolites. Fungal community pattern was also affected in the presence of algae, but with a different specificity of algae. Furthermore, algal extracts exhibited a distinct pattern of activities against epiphytic diatoms.