Rainer Knust

Climate induced temperature effects on growth performance, fecundity and recruitment in marine fish: developing a hypothesis for cause and effect relationships in Atlantic cod (Gadus morhua) and common eelpout (Zoarces viviparus)

Effects of global warming on animal distribution and performance become visible in many marine ecosystems. The present study was designed to develop a concept for a cause and effect understanding with respect to temperature changes and to explain ecological findings based on physiological processes. The concept is based on a wide comparison of invertebrate and fish species with a special focus on recent data obtained in two model species of fish. These fish species are both characterized by northern and southern distribution limits in the North Atlantic: eelpout (Zoarces viviparus), as a typical non-migrating inhabitant of the coastal zone and the cod (Gadus morhua), as a typical inhabitant of the continental shelf with a high importance for fisheries. Mathematical modelling demonstrates a clear significant correlation between climate induced temperature fluctuations and the recruitment of cod stocks. Growth performance in cod is optimal at temperatures close to 10°C, regardless of the population investigated in a latitudinal cline. However, temperature specific growth rates decrease at higher latitudes. Also, fecundity is less in White Sea than in North and Baltic Sea cod or eelpout populations. These findings suggest that a cold-induced shift in energy budget occurs which is unfavorable for growth performance and fecundity. Thermal tolerance limits shift depending on latitude and are characterized by oxygen limitation at both low or high temperatures. Oxygen supply to tissues is optimized at low temperature by a shift in hemoglobin isoforms and oxygen binding properties to lower affinities and higher unloading potential. Protective stimulation of heat shock protein synthesis was not observed. According to a recent model of thermal tolerance the downward shift of tolerance limits during cold adaptation is associated with rising mitochondrial densities and, thus, aerobic capacity and performance in the cold, especially in eurythermal species. At the same time the costs of mitochondrial maintenance reflected by mitochondrial proton leakage should rise leaving a lower energy fraction for growth and reproduction. The preliminary conclusion can be drawn that warming will cause a northern shift of distribution limits for both species with a rise in growth performance and fecundity larger than expected from the Q10 effect in the north and lower growth or even extinction of the species in the south. Such a shift may heavily affect fishing activities in the North Sea.

The Dogger Bank: A special ecological region in the central North Sea

This paper is a review of the present state of knowledge about the ecology of the Dogger Bank, a shallow area in the central southern North Sea. The biological situation in this region is different from other regions in the North Sea; phytoplankton production occurs throughout the year, connected with low periodicity in macrofaunal abundance and condition of fish. Against prevailing opinions, the Dogger Bank, an offshore region, seems to be affected by eutrophication and pollution.

Temperature-dependent energy allocation to growth in Antarctic and boreal eelpout (Zoarcidae)

Antarctic fishes display slower annual growth rates than congeneric species from temperate zones. For an analysis of growth in relation to energy turnover, body composition was analysed in two benthic fish species to establish a whole animal energy budget. The Antarctic eelpout, Pachycara brachycephalum, was maintained at 0, 2, 4 and 6°C and the boreal eelpout, Zoarces viviparus at 4, 6, 12 and 18°C. At maximum food supply the weight gain was highest for P. brachycephalum at 4°C. Routine metabolic rate in acclimated Antarctic eelpouts did not differ between temperatures, whereas in Z. viviparus maximized growth benefited from a reduction of metabolic energy demands at 12°C. The lipid content of liver declined with increasing temperature in both species. The thermal window for growth is based on food conversion efficiency and the level of metabolic energy demand and is limited according to the level of aerobic scope available between pejus temperatures.

Impact of climate change on fishes in complex Antarctic ecosystems

Abstract Antarctic marine ecosystems are increasingly threatened by climate change and are considered to be particularly sensitive because of the adaptation of most organisms to cold and stable environmental conditions. Fishes play a central role in the Antarctic marine food web and might be affected by climate change in different ways: (i) directly by increasing water temperatures, decreasing seawater salinity and/or increasing concentrations of CO2; (ii) indirectly by alterations in the food web, in particular by changes in prey composition, and (iii) by alterations and loss of habitat due to sea ice retreat and increased ice scouring on the sea floor. Based on new data and data collected from the literature, we analyzed the vulnerability of the fish community to these threats. The potential vulnerability and acting mechanisms differ among species, developmental stages and habitats. The icefishes (family Channichthyidae) are one group that are especially vulnerable to a changing South Polar Sea, as are the pelagic shoal fish species Pleuragramma antarcticum. Both will almost certainly be negatively affected by abiotic alterations and changes in food web structure associated with climate change, the latter additionally by habitat loss. The major bottleneck for the persistence of the majority of populations appears to be the survival of early developmental stages, which are apparently highly sensitive to many types of alterations. In the long term, if climate projections are realized, species loss seems inevitable: within the demersal fish community, the loss or decline of one species might be compensated by others, whereas the pelagic fish community in contrast is extremely poor in species and dominated by P. antarcticum. The loss of this key species could therefore have especially severe consequences for food web structure and the functioning of the entire ecosystem.

The role of iceberg scours in niche separation within the Antarctic fish genus Trematomus

Abstract. Species of the Antarctic fish genus Trematomus occupy different trophic niches. It is not clear, however, whether small-scale variability in benthic community structure affects niche separation. Therefore abundance and biomass of fish were determined and stomach content and food composition were compared in areas affected by iceberg scours and unaffected areas in the Weddell Sea. Trematomus eulepidotus, T. lepidorhinus and T. scotti dominate undisturbed areas, whereas T. nicolai and especially T. pennellii dominate disturbed areas. Total stomach content and number of prey taxa per fish are higher in preferred than in non-preferred areas. These findings indicate that small-scale horizontal patterns caused by iceberg scours play a distinct role in Trematomus niche separation.

Stress response or beneficial temperature acclimation: transcriptomic signatures in Antarctic fish (Pachycara brachycephalum)

Research on the thermal biology of Antarctic marine organisms has increased awareness of their vulnerability to climate change, as a flipside of their adaptation to life in the permanent cold and their limited capacity to acclimate to variable temperatures. Here, we employed a species‐specific microarray of the Antarctic eelpout, Pachycara brachycephalum, to identify long‐term shifts in gene expression after 2 months of acclimation to six temperatures between −1 and 9 °C. Changes in cellular processes comprised signalling, post‐translational modification, cytoskeleton remodelling, metabolic shifts and alterations in the transcription as well as translation machinery. The magnitude of transcriptomic responses paralleled the change in whole animal performance. Optimal growth at 3 °C occurred at a minimum in gene expression changes indicative of a balanced steady state. The up‐regulation of ribosomal transcripts at 5 °C and above was accompanied by the transcriptomic activation of differential protein degradation pathways, from proteasome‐based degradation in the cold towards lysosomal protein degradation in the warmth. From 7 °C upwards, increasing transcript levels representing heat‐shock proteins and an acute inflammatory response indicate cellular stress. Such patterns may contribute to a warm‐induced energy deficit and a strong weight loss at temperatures above 6 °C. Together, cold or warm acclimation led to specific cellular rearrangements and the progressive development of functional imbalances beyond the optimum temperature. The observed temperature‐specific expression profiles reveal the molecular basis of thermal plasticity and refine present understanding of the shape and positioning of the thermal performance curve of ectotherms on the temperature scale.

Temperature-dependent lipid levels and components in polar and temperate eelpout (Zoarcidae)

Total lipid content, lipid classes and fatty acid composition were analysed in tissues from two eelpout species fed on the same diet, the Antarctic Pachycara brachycephalum and the temperate Zoarces viviparus, with the aim of determining the role of lipids in fishes from different thermal habitats. The lipid content increased with decreasing temperature in the liver of both species, suggesting enhanced lipid storage under cold conditions. In P. brachycephalum, lipid composition in the liver and muscle was strongly dominated by triacylglycerols between 0 and 6°C. In contrast, in the temperate species, lipid class composition changed with changes in the temperature. When acclimatized to 4 and 6°C Z. viviparus not only displayed a shift to lipid anabolism and pronounced lipid storage, as indicated by high triacylglycerol levels, but also a shift to patterns of cold adaptation, as reflected by an increased content of polyunsaturated fatty acids in the lipid extract. Unsaturated fatty acids were also abundant in the Antarctic eelpout, but when compared to Z. viviparus at the same temperatures, the latter had significantly higher ratios of polyunsaturated to saturated fatty acid levels, whereas the Antarctic eelpout showed significantly higher ratios of monounsaturated to saturated fatty acid levels. High δ-15N values of the Antarctic eelpout reflect the high trophic level of this scavenger in the Weddell Sea food web. Stable carbon values suggest that lipid-enriched prey forms a major part of its diet. The strategy to accumulate storage lipids in the cold is interpreted to be adaptive behaviour at colder temperatures and during periods of irregular, pulsed food supply.

Fluctuating asymmetry and other parameters of morphological variation of eelpout Zoarces viviparus (Zoarcidae, Teleostei) from different parts of its distributional range

Five populations of eelpout Zoarces viviparus from the White Sea, the Baltic (Gulf of Gdansk and Gulf of Finland) and the North Sea (Wadden Sea and Hafrsfjord) were studied. Using 17 bilateral meristic characters (number of holes in cranial bones) we assessed the following parameters: (i) fluctuating asymmetry (minor deviations from perfect bilateral symmetry), (ii) the factorial component of total phenotypic variance, which is a measure of within-population heterogeneity and (iii) mean values of each character. Morphological similarity among samples was only weakly correlated with geographical distances between sample locations. The magnitude of fluctuating asymmetry was higher in the samples from the White Sea and Hafrsfjord than in samples from the Wadden Sea and the Baltic Sea. The sample from the Gulf of Gdansk was characterized by the lowest fluctuating asymmetry and the highest factorial variation. We propose that among the most important factors determining fluctuating asymmetry in eelpout populations are salinity and temperature conditions and the level of genetic variation.

Temperature-dependent metabolism in Antarctic fish: Do habitat temperature conditions affect thermal tolerance ranges?

Climatic warming is most pronounced in the polar regions. For marine ectotherms such as fish, temperature is a key abiotic factor, influencing metabolic processes. Species distribution and abundance are driven by reproduction and growth, which depend on available energy exceeding baseline maintenance costs. These routine metabolic costs make up a large part of the energy expenditure. Thermal stress can increase routine metabolism, affecting an organism’s fitness. Data of routine metabolic rates of Antarctic fish are scarce, and comparability of existing data sets is often problematic due to ecological differences between species and in experimental protocols. Our objective was to compare routine metabolism and thermal sensitivity of species with similar ecotypes but different thermal environments to assess possible ecological implications of warming waters on energy expenditure in Antarctic fish, a fauna characterised by geographic isolation, endemism and putative thermal adaptation. We measured routine metabolic rates of three benthic Antarctic fish species from low- and high-Antarctic regions at habitat temperature and during acute temperature increase. Our analysis revealed differences in metabolic rates at the same temperature suggesting local adaptation to habitat temperature. Acute thermal stress induced a comparable response of metabolic rates to increasing temperature. We conclude that higher metabolic rates and thus higher energetic costs could be associated with narrower thermal windows, a potential disadvantage to the endemic high-Antarctic fish fauna facing the challenge of climate change.

A first insight into the spleen transcriptome of the notothenioid fish Lepidonotothen nudifrons: Resource description and functional overview

In this study, we describe a de novo sequencing and assembly of the spleen transcriptome of Lepidonotothen nudifrons, a notothenioid fish widely distributed around the Antarctic Peninsula and the Scotia Arc. Sequences were generated on an Illumina MiSeq system and assembled to a total of 112,477 transcripts. Putative functional annotation was possible for more than 34% of the transcripts. This data will be relevant for future studies targeting the erythrocyte turnover, oxygen transport mechanism and immune system, which are key functional traits to investigate cold adaptation and thermal sensitivity of Antarctic notothenioids.