ALBERT

Description: Euphausiids constitute a major biomass component in shelf ecosystems and play a fundamental role in the rapid vertical transport of carbon from the ocean surface to the deeper layers during their daily vertical migration (DVM). DVM depth and migration patterns depend on oceanographic conditions with respect to temperature, light and oxygen availability at depth, factors that are highly dependent on season in most marine regions. Here we introduce a global krill respiration ANN (artificial neural network) model including the effect of latitude (LAT), the day of the year (DoY), and the number of daylight hours (DLh), in addition to the basal variables that determine ectothermal oxygen consumption (temperature, body mass and depth). The newly implemented parameters link space and time in terms of season and photoperiod to krill respiration. The ANN model showed a better fit (r2 = 0.780) when DLh and LAT were included, indicating a decrease in respiration with increasing LAT and decreasing DLh. We therefore propose DLh as a potential variable to consider when building physiological models for both hemispheres. For single Euphausiid species investigated in a large range of DLh and DoY, we also tested the standard respiration rate for seasonality with Multiple Linear Regression (MLR) and General Additive model (GAM). GAM successfully integrated DLh (r2 = 0.563) and DoY (r2 = 0.572) effects on respiration rates of the Antarctic krill, Euphausia superba, yielding the minimum metabolic activity in mid-June and the maximum at the end of December. We could not detect DLh or DoY effects in the North Pacific krill Euphausia pacifica, and our findings for the North Atlantic krill Meganyctiphanes norvegica remained inconclusive because of insufficient seasonal data coverage. We strongly encourage comparative respiration measurements of worldwide Euphausiid key species at different seasons to improve accuracy in ecosystem modeling.

Description: Euphausiids constitute a major biomass component in shelf ecosystems and play a fundamental role in the rapid vertical transport of carbon from the ocean surface to the deeper layers during their daily vertical migration (DVM). DVM depth and migration patterns depend on oceanographic conditions with respect to temperature, light and oxygen availability at depth, factors that are highly dependent on season in most marine regions. Here we introduce a global krill respiration ANN (Artificial Neural Network) model including the effect of latitude (LAT), the day of the year (DoY), and the number of daylight hours (DLh), in addition to the basal variables that determine ectothermal oxygen consumption (temperature, body mass and depth). The newly implemented parameters link space and time in terms of season and photoperiod to krill respiration. The ANN model showed a better fit (r2=0.780) when DLh and LAT were included, indicating a decrease in respiration with increasing LAT and decreasing DLh. We therefore propose DLh as a potential variable to consider when building physiological models for both hemispheres. For single Euphausiid species investigated in a large range of DLh and DoY, we also tested the standard respiration rate for seasonality with Multiple Linear Regression (MLR) and General Additive model (GAM). GAM successfully integrated DLh (r2= 0.563) and DoY (r2= 0.572) effects on respiration rates of the Antarctic krill, Euphausia superba, yielding the minimum metabolic activity in mid-June and the maximum at the end of December. We could not detect DLh or DoY effects in the North Pacific krill Euphausia pacifica, and our findings for the North Atlantic krill Meganyctiphanes norvegica remained inconclusive because of insufficient seasonal data coverage. We strongly encourage comparative respiration measurements of worldwide Euphausiid key species at different seasons to improve accuracy in ecosystem modelling.

Description: The Antarctic krill, Euphausia superba, has evolved seasonal rhythms of physiology and behaviour to survive under the extreme photoperiodic conditions in the Southern Ocean. However, the molecular mechanisms generating these rhythms remain far from understood. The aim of this study was to investigate seasonal differences in gene expression in three different latitudinal regions (South Georgia, South Orkneys/Bransfield Strait, Lazarev Sea) and to identify genes with potential regulatory roles in the seasonal life cycle of Antarctic krill. The RNA-seq data were analysed (a) for seasonal differences between summer and winter krill sampled from each region, and (b) for regional differences within each season. A large majority of genes showed an up-regulation in summer krill in all regions with respect to winter krill. However, seasonal differences in gene expression were less pronounced in Antarctic krill from South Georgia, most likely due to the milder seasonal conditions of the lower latitudes of this region, with a less extreme light regime and food availability between summer and winter. Our results suggest that in the South Orkneys/Bransfield Strait and Lazarev Sea region, Antarctic krill entered a state of metabolic depression and regressed development (winter quiescence) in winter. Moreover, seasonal gene expression signatures seems to be driven by a photoperiodic timing system that may adapt the flexible behaviour and physiology of Antarctic krill to the highly seasonal environment according to the latitudinal region. However, at the lower latitude South Georgia region, food availability might represent the main environmental cue influencing seasonal physiology.

Description: Antarctic krill up- and down-regulate their metabolism as a strategy to cope with the strong seasonal environmental fluctuations in the Southern Ocean. In this study, we investigate the impact of this light- and temperature dependent metabolic regulation on growth, reproduction and winter survival of krill. Therefore, we advance a bioenergetic growth model of krill by adding a data-derived scaling function of krill activity. With SERBIK (SEasonally Regulated BIoenergetic Krill growth model), we conduct a numerical experiment which tests the impact of such scaling on krill life history under two different winter food conditions: In the first scenario, we simulate life history of krill when winter food availability is low; in the second scenario, winter food availability is increased within realistic ranges. The results demonstrate that the scaling of metabolism is especially important during low food winters. Reducing metabolism during winter permits individuals to grow to larger body length, reproduce successfully and release a greater number of eggs. It further significantly reduces withinyear size fluctuations caused by starvation during months with low food availability. Finally, SERBIK can be used in future spatial modelling studies which include movement of krill along latitudinal gradients and thus spatiotemporal gradients in light- and temperature.

Description: The Antarctic krill Euphausia superba experiences almost all marine photic environments throughout its life cycle. Antarctic krill eggs hatch in the aphotic zone up to 1000 m depth and larvae develop on their way to the ocean surface (development ascent) and are exposed to different quality (wavelength) and quantity (irradiance) of light. Adults show a daily vertical migration pattern, moving downward during the day and upward during the night within the top 200 m of the water column. Seawater acts as a potent chromatic filter and animals have evolved different opsin photopigments to perceive photons of specific wavelengths. We have investigated the transcriptome of E. superba and, using a candidate gene approach, we identified six novel opsins. Five are r-type visual opsins: four middle-wavelength-sensitive (EsRh2, EsRh3, EsRh4 and EsRh5) and one long-wavelengthsensitive (EsRh6). Moreover, we have identified a non-visual opsin, the EsPeropsin. All these newly identified opsin genes were significantly expressed in compound eyes and brain, while only EsPeropsin and EsRh2 were clearly detected also in the abdomen. A temporal modulation in the transcription of these novel opsins was found, but statistically significant oscillations were only observed in EsRrh3 and EsPeropsin. Our results contribute to the dissection of the complex photoreception systemof E. superba, which enables this species to respond to the daily and seasonal changes in irradiance and spectral composition in the Southern Ocean.

Description: The Southern Ocean ecosystem is characterized by extreme seasonal changes in environmental factors such as day length, sea ice extent and food availability. The key species Antarctic krill (Euphausia superba) has evolved metabolic and behavioural seasonal rhythms to cope with these seasonal changes. We investigate the switch between a physiological less active and active period for adult krill, a rhythm which seems to be controlled by internal biological clocks. These biological clocks can be synchronized by environmental triggers such as day length and food availability. They have evolved for particular environmental regimes to synchronize predictable seasonal environmental changes with important life cycle functions of the species. In a changing environment the time when krill is metabolically active and the time of peak food availability may not overlap if krill's seasonal activity is solely determined by photoperiod (day length). This is especially true for the Atlantic sector of the Southern Ocean where the spatio-temporal ice cover dynamics are changing substantially with rising average temperatures. We developed an individual-based model for krill to explore the impact of photoperiod and food availability on the growth and demographics of krill. We simulated dynamics of local krill populations (with no movement of krill assumed) along a south-north gradient for different triggers of metabolic activity and different levels of food availability below the ice. We also observed the fate of larval krill which cannot switch to low metabolism and therefore are likely to overwinter under ice. Krill could only occupy the southern end of the gradient, where algae bloom only lasts for a short time, when alternative food supply under the ice was high and metabolic activity was triggered by photoperiod. The northern distribution was limited by lack of overwintering habitat for krill larvae due to short duration of sea ice cover even for high food content under the ice. The variability of the krill's length-frequency distributions varied for different triggers of metabolic activity, but did not depend on the sea ice extent. Our findings suggest a southward shift of krill populations due to reduction in the spatial sea ice extent, which is consistent with field observations. Overall, our results highlight the importance of the explicit consideration of spatio-temporal sea ice dynamics especially for larval krill together with temporal synchronization through internal clocks, triggered by environmental factors (photoperiod and food) in adult krill for the population modelling of krill.

Description: 〈jats:title〉ABSTRACT〈/jats:title〉 〈jats:sec〉 〈jats:title /〉 〈jats:p〉 With almost a quadrillion individuals, the Antarctic krill processes five million tons of organic carbon every day during austral summer. This high carbon flux requires a broad range of hydrolytic enzymes to decompose the diverse food-derived biopolymers. While krill itself possesses numerous such enzymes, it is unclear, to what extent the endogenous microbiota contribute to the hydrolytic potential of the gut environment. Here we applied amplicon sequencing, shotgun metagenomics, cultivation, and physiological assays to characterize the krill gut microbiota. The broad bacterial diversity (273 families, 919 genera, and 2,309 species) also included a complex potentially anaerobic sub-community. Plate-based assays with 198 isolated pure cultures revealed widespread capacities to utilize lipids (e.g., tributyrin), followed by proteins (casein) and to a lesser extent by polysaccharides (e.g., alginate and chitin). While most isolates affiliated with the genera 〈jats:italic〉Pseudoalteromonas〈/jats:italic〉 and 〈jats:italic〉Psychrobacter〈/jats:italic〉 , also 〈jats:italic〉Rubritalea〈/jats:italic〉 spp. (Verrucomicrobia) were observed. The krill gut microbiota growing on marine broth agar plates possess 13,012 predicted hydrolyses; 15-fold more than previously predicted from a transcriptome-proteome compendium of krill. Cultivation-independent and -dependent approaches indicated members of the families 〈jats:italic〉Flavobacteriaceae〈/jats:italic〉 and 〈jats:italic〉Pseudoalteromonadaceae〈/jats:italic〉 to dominate the capacities for lipid/protein hydrolysis and to provide a plethora of carbohydrate-active enzymes, sulfatases, and laminarin- or porphyrin-depolymerizing hydrolases. Notably, also the potential to hydrolyze plastics such as polyethylene terephthalate and polylactatide was observed, affiliating mostly with Moraxellaceae. Overall, this study shows extensive microbial diversity in the krill gut, and suggests that the microbiota likely play a significant role in the nutrient acquisition of the krill by enriching its hydrolytic enzyme repertoire. 〈/jats:p〉 〈jats:sec〉 〈jats:title〉IMPORTANCE〈/jats:title〉 〈jats:p〉 The Antarctic krill ( 〈jats:italic〉Euphausia superba〈/jats:italic〉 ) is a keystone species of the Antarctic marine food web, connecting the productivity of phyto- and zooplankton with the nutrition of the higher trophic levels. Accordingly, krill significantly contributes to biomass turnover, requiring the decomposition of seasonally varying plankton-derived biopolymers. This study highlights the likely role of the krill gut microbiota in this ecosystem function by revealing the great number of diverse hydrolases that microbes contribute to the krill gut environment. The here resolved repertoire of hydrolytic enzymes could contribute to the overall nutritional resilience of krill and to the general organic matter cycling under changing environmental conditions in the Antarctic sea water. Furthermore, the krill gut microbiome could serve as a valuable resource of cold-adapted hydrolytic enzymes for diverse biotechnological applications. 〈/jats:p〉 〈/jats:sec〉 〈/jats:sec〉