Description: Temora longicornis, a dominant calanoid copepod species in the North Sea, is characterised by low lipid reserves and high biomass turnover rates. To survive and reproduce successfully, this species needs continuous food supply and thus requires a highly flexible digestive system to exploit various food sources. Information on the capacity of digestive enzymes is scarce and therefore the aim of our study was to investigate the enzymatic capability to respond to quickly changing nutritional conditions. We conducted two feeding experiments with female T. longicornis from the southern North Sea off Helgoland. In the first experiment in 2005, we tested how digestive enzyme activities and enzyme patterns as revealed by substrate SDS-PAGE (sodium dodecylsulfate polyacrylamide gel electrophoresis) responded to changes in food composition. Females were incubated for three days fed ad libitum with either the heterotrophic dinoflagellate Oxyrrhis marina or the diatom Thalassiosira weissflogii. At the beginning and at the end of the experiment, copepods were deep-frozen for analyses. The lipolytic enzyme activity did not change over the course of the experiment but the enzyme patterns did, indicating a distinct diet-induced response. In a second experiment in 2008, we therefore focused on the enzyme patterns, testing how fast changes occur and whether feeding on the same algal species leads to similar patterns. In this experiment, we kept the females for 4 days at surplus food while changing the algal food species daily. At day 1, copepods were offered O. marina. On day 2, females received the cryptophycean Rhodomonas baltica followed by T. weissflogii on day 3. On day 4 copepods were again fed with O. marina. Each day, copepods were frozen for analysis by means of substrate SDS-PAGE. This showed that within 24 h new digestive enzymes appeared on the electrophoresis gels while others disappeared with the introduction of a new food species, and that the patterns were similar on day 1 and 4, when females were fed with O. marina. In addition, we monitored the fatty acid compositions of the copepods, and this indicated that specific algal fatty acids were quickly incorporated. With such short time lags between substrate availability and enzyme response, T. longicornis can successfully exploit short-term food sources and is thus well adapted to changes in food availability, as they often occur in its natural environment due seasonal variations in phyto- and microzooplankton distribution.

Description: A pronounced seasonal variability in primary productivity is the crucial factor in determining the life strategies of many polar zooplankton species, whereas oligotrophic tropical oceans experience little change with a low productivity. Lipid accumulation represents an important energetic adaptation in pelagic organisms to cope with the pronounced seasonal productivity in polar oceans. We compare important global euphausiid species from the Arctic via the tropics to the Antarctic, focussing on the genera Euphausia and Thysanoessa. While the Antarctic krill Euphausia superba is the species with the highest biomass of all metazoans on earth, krill species are clearly less relevant in other oceans. In polar krill species total lipid accumulation is usually very pronounced and may be utilised for metabolic maintenance during overwintering or for reproductive processes in spring. Most polar krill species store large amounts of the unusual depot lipid phosphatidylcholine (lecithin), a polar lipid with highly unsaturated fatty acids. However, they exhibit strong differences in their neutral lipid compounds, which may either consist of wax esters or triacylglycerols, but also of both types of lipid classes. Their major end-products of the fatty acid and fatty alcohol biosynthesis are quite different and usually species-specific. In contrast, tropical euphausiids do not rely on lipids as energy reserves and exhibit the usual lipid and fatty acid compositions of biomembranes with high amounts of eicosapentaenoic acid (EPA) and docosahexaenoic acid (DHA) and eicosapentaenoic acid (EPA). In polar euphausiids, the major end-products of the fatty acid and fatty alcohol biosynthesis are quite different and usually species-specific. The ecophysiological implications of these deviating lipid characteristics may determine biogeographical zonation patterns and affect the vulnerability of polar krill species to global warming.

Description: In the framework of the GENUS –“Geochemistry and Ecology of the Namibian Upwelling System” research program, trophic interactions and carbon pathways throughout the food web of the coastal upwelling system are being quantified. In contrast to earlier studies, special focus is been given to lower trophic levels in higher taxonomic resolution. Energy demands of various zooplankton taxa, including copepods, euphausiids, decapods and fish larvae, have been quantified with standardized methodology via optode respirometry. Dietary spectra and trophic levels were analyzed by trophic biomarker approaches based on fatty acid composition and stable isotopes (15N, 13C), respectively. All empirical data are assembled for an Ecopath with Ecosim (EwE) food-web model. The EwE model distinguishes between shelf and offshore communities. The conceptual food‐web model consists of three groups of primary producers, i.e. diatoms, dinoflagellates, and cyanobacteria, as well as many consumers such as Calanoides carinatus as the key herbivorous copepod in the Benguela upwelling system, other copepods, Euphausia hanseni, other zooplankton, and pelagic fishes including sardine, anchovy, and horse mackerel. Empirical data show that zooplankton and particularly copepods encompass a wider range of trophic levels from herbivory to secondary or even tertiary consumers (δ15N from 4 to 12‰), while anchovy had rather low δ15N of about 7‰. Respiration rates and metabolic activities of copepods could be parameterized for the model by an energy budget approach based on ambient temperature, body mass, and activity level. Calanoid copepods consumed 78mg C m‐2 d‐1 in shelf regions and 21mg C m‐2 d‐1 in oceanic regions. Locally, C. carinatus could remove up to 90% of the diatom biomass per day. The community consumption of pelagic decapods ranged from 7 mg C m‐2 d‐1 to 〉20mg C m‐2 d‐1 with highest values in the northernmost part of the study area. Overall, pelagic decapods apparently play a more prominent role in the northern Benguela ecosystem than previously assumed and may exert a substantial predation pressure on calanoid copepods. GENUS results emphasize that the trophic interactions within zooplankton and lower trophic levels are more complex than just linking primary producers with pelagic fish and should be taken into account in the process of developing realistic food‐web models of coastal upwelling systems. Keywords: foodweb, zooplankton, trophic interactions, energy flux

Description: The goal of the GENUS project (Geochemistry and Ecology of the Namibian Upwelling System) is to analyse the interrelationships between climate change, oceanic nutrients, greenhouse gases and the ecosystem structure in the coastal upwelling area off Namibia. The biological/ecological work package focused on the structure of the Northern Benguela Upwelling System (NBUS) and its energy flows under changing environmental conditions. Physiological constraints and adaptations were detected in several taxonomic groups investigated within the project, such as several copepod species, euphausiids and fish larvae. Temperature and oxygen distribution in the water column were identified as main drivers in modulating the distribution and ecology of many species. The extension and position of the Oxygen Minimum Zone (OMZ) seems to have a significant impact on the life cycles, vertical distribution and trophic condition of various species. In copepods we find a variety of adaptation mechanisms. While some species avoid the OMZ, others use it for resting and predator avoidance during daytime. Such vertical migrations contribute significantly to vertical carbon flux. The same holds true for decapods (4.4 mg C m‐2 d‐1). Respiration rates of 16 copepod species were determined to average 54.6 ±32.8 ml O2 d‐1 gDM‐1. Calanoides carinatus diapausing C5 stages reduced respiration at depth by 82% compared to surface activity. Further adaptations were found in euphausiid species: While Euphausia hanseni is capable to use the OMZ as a retreat by reducing its metabolic activity at lower temperatures and unfavourable trophic conditions, Nematoscelis megalops generally maintains a low level metabolism adapted to a constant life in the OMZ, and avoids crossing the thermocline. Special features of early life stages of Trachurus capensis, the fish species actually showing highest commercial landings, were analysed to elucidate their potential advantages in life performance, compared to other small pelagic species such as sardines or anchovies. The species showed short‐term hypoxia tolerance down to 30% oxygen saturation and even survived 10% saturation. Combined with the ability to switch from smaller to larger copepod prey and to surpass vulnerable early stages much faster than competitor species, this could explain the dominance of Trachurus capensis in the NBUS. In addition, competitors and predators of fish larvae such as jellyfish or chaetognaths showed little or no response to low oxygen concentrations gaining advantage over e.g. sardines and anchovies. Isotope analyses of various pelagic species and their food revealed a complex picture of the trophic levels of species including developmental stages and provide the basis for trophic flow models. The results will also serve to calibrate carbon dynamics and nutrient flux models that were developed in another GENUS work package. Data will be compared with nutrient distribution patterns and dynamics that may influence primary production and impact zooplankton distribution and higher trophic levels such as fish, seabirds or mammals. Results clearly show that continuing ocean warming coupled with expansion of the OMZ may alter horizontal and vertical distribution of species and the food web structure of the ecosystem. Keywords: Benguela Current, OMZ, Pelagic Ecosystem, Physiology