Habitat characteristics and energy budget of the ocean quahog (Arctica islandica)

We compared lifetime and population energy budgets of the extraordinary long-lived ocean quahog Arctica islandica from 6 different sites - the Norwegian coast, Kattegat, Kiel Bay, White Sea, German Bight, and off northeast Iceland - covering a temperature and salinity gradient of 4-10°C (annual mean) and 25-34, respectively. Based on von Bertalanffy growth models and size-mass relationships, we computed organic matter production of body (PSB) and of shell (PSS), whereas gonad production (PG) was estimated from the seasonal cycle in mass. Respiration (R) was computed by a model driven by body mass, temperature, and site. A. islandica populations differed distinctly in maximum life span (40 y in Kiel Bay to 197 y in Iceland), but less in growth performance (phi' ranged from 2.41 in the White Sea to 2.65 in Kattegat). Individual lifetime energy throughput, as approximated by assimilation, was highest in Iceland (43,730 kJ) and lowest in the White Sea (313 kJ). Net growth efficiency ranged between 0.251 and 0.348, whereas lifetime energy investment distinctly shifted from somatic to gonad production with increasing life span; PS/PG decreased from 0.362 (Kiel Bay, 40 y) to 0.031 (Iceland, 197 y). Population annual energy budgets were derived from individual budgets and estimates of population mortality rate (0.035/y in Iceland to 0.173/y in Kiel Bay). Relationships between budget ratios were similar on the population level, albeit with more emphasis on somatic production; PS/ PG ranged from 0.196 (Iceland) to 2.728 (White Sea), and P/B ranged from 0.203-0.285/y. Life span is the principal determinant of the relationship between budget parameters, whereas temperature affects net growth efficiency only. In the White Sea population, both growth performance and net growth efficiency of A. islandica were lowest. We presume that low temperature combined with low salinity represent a particularly stressful environment for this species.

Global macrozoobenthos production and energy budget data base V150731, with link to database

I developed a new model for estimating annual production-to-biomass ratio P/B and production P of macrobenthic populations in marine and freshwater habitats. Self-learning artificial neural networks (ANN) were used to model the relationships between P/B and twenty easy-to-measure abiotic and biotic parameters in 1252 data sets of population production. Based on log-transformed data, the final predictive model estimates log(P/B) with reasonable accuracy and precision (r2 = 0.801; residual mean square RMS = 0.083). Body mass and water temperature contributed most to the explanatory power of the model. However, as with all least squares models using nonlinearly transformed data, back-transformation to natural scale introduces a bias in the model predictions, i.e., an underestimation of P/B (and P). When estimating production of assemblages of populations by adding up population estimates, accuracy decreases but precision increases with the number of populations in the assemblage.

Impact of long-term moderate hypercapnia and elevated temperature on the energy budget of isolated gills and branchial in vivo and in vitro enzyme capacities of Atlantic cod (Gadus morhua)

Effects of severe hypercapnia have been extensively studied in marine fishes, while knowledge on the impacts of moderately elevated CO2 levels and their combination with warming is scarce. Here we investigate ion regulation mechanisms and energy budget in gills from Atlantic cod acclimated long-term to elevated PCO2 levels (2500 µatm) and temperature (18 °C). Isolated perfused gill preparations established to determine gill thermal plasticity during acute exposures (10-22 °C) and in vivo costs of Na+/K+-ATPase activity, protein and RNA synthesis. Maximum enzyme capacities of F1Fo-ATPase, H+-ATPase and Na+/K+-ATPase were measured in vitro in crude gill homogenates. After whole animal acclimation to elevated PCO2 and/or warming, branchial oxygen consumption responded more strongly to acute temperature change. The fractions of gill respiration allocated to protein and RNA synthesis remained unchanged. In gills of fish CO2-exposed at both temperatures, energy turnover associated with Na+/K+-ATPase activity was reduced by 30% below rates of control fish. This contrasted in vitro capacities of Na+/K+-ATPase, which remained unchanged under elevated CO2 at 10 °C, and earlier studies which had found a strong upregulation under severe hypercapnia. F1Fo-ATPase capacities increased in hypercapnic gills at both temperatures, whereas Na+/K+ATPase and H+-ATPase capacities only increased in response to elevated CO2 and warming indicating the absence of thermal compensation under CO2. We conclude that in vivo ion regulatory energy demand is lowered under moderately elevated CO2 levels despite the stronger thermal response of total gill respiration and the upregulation of F1Fo-ATPase. This effect is maintained at elevated temperature.

Combined effects of short-term exposure to elevated CO2 and decreased O2 on the physiology and energy budget of the thick shell mussel Mytilus coruscus

Hypoxia and ocean acidification are two consequences of anthropogenic activities. These global trends occur on top of natural variability. In environments such as estuarine areas, short-term acute pH and O2 fluctuations are occurring simultaneously. The present study tested the combined effects of short-term seawater acidification and hypoxia on the physiology and energy budget of the thick shell mussel Mytilus coruscus. Mussels were exposed for 72 h to six combined treatments with three pH levels (8.1, 7.7 and 7.3) and two dissolved oxygen (DO) levels (2 mg/L, 6 mg/L). Clearance rate (CR), food absorption efficiency (AE), respiration rate (RR), ammonium excretion rate (ER), O:N ratio and scope for growth (SFG) were significantly reduced, and faecal organic dry weight ratio (E) was significantly increased at low DO. Low pH did not lead to a reduced SFG. Interactive effects of pH and DO were observed for CR, E and RR. Principal component analysis (PCA) revealed positive relationships among most physiological indicators, especially between SFG and CR under normal DO conditions. These results demonstrate that Mytilus coruscus was sensitive to short-term (72 h) exposure to decreased O2 especially if combined with decreased pH levels. In conclusion, the short-term oxygen and pH variation significantly induced physiological changes of mussels with some interactive effects.

Differential impacts of elevated CO2 and acidosis on the energy budget of gill and liver cells from Atlantic cod

Ocean acidification impacts fish and other marine species through increased seawater PCO2 levels (hypercapnia). Knowledge of the physiological mechanisms mediating effects in various tissues of fish is incomplete. Here we tested the effects of extracellular hypercapnia and acidosis on energy metabolism of gill and liver cells of Atlantic cod. Exposure media mimicked blood conditions in vivo, either during normo- or hypercapnia and at control or acidic extracellular pH (pHe). We determined metabolic rate and energy expenditure for protein biosynthesis, Na+/K+-ATPase and H+-ATPase and considered nutrition status by measurements of metabolic rate and protein biosynthesis in media with and without free amino acids (FAA). Addition of FAA stimulated hepatic but not branchial oxygen consumption. Normo- and hypercapnic acidosis as well as hypercapnia at control pHe depressed metabolic stimulation of hepatocytes. In gill cells, acidosis depressed respiration independent of PCO2 and FAA levels. For both cell types, depressed respiration was not correlated with the same reduction in energy allocated to protein biosynthesis or Na+/K+-ATPase. Hepatic energy expenditure for protein synthesis and Na+/K+- ATPase was even elevated at acidic compared to control pHe suggesting increased costs for ion regulation and cel- lular reorganization. Hypercapnia at control pHe strongly reduced oxygen demand of branchial Na+/K+-ATPase with a similar trend for H+-ATPase. We conclude that extracellular acidosis triggers metabolic depression in gill and metabolically stimulated liver cells. Additionally, hypercapnia itself seems to limit capacities for metabolic usage of amino acids in liver cells while it decreases the use and costs of ion regulatory ATPases in gill cells.

Fluxes and meteorological observations on Spitsbergen

Arctic permafrost may be adversely affected by climate change in a number of ways, so that establishing a world-wide monitoring program seems imperative. This thesis evaluates possibilities for permafrost monitoring at the example of a permafrost site on Svalbard, Norway. An energy balance model for permafrost temperatures is developed that evaluates the different components of the surface energy budget in analogy to climate models. The surface energy budget, consisting of radiation components, sensible and latent heat fluxes as well as the ground heat flux, is measured over the course of one year, which has not been accomplished for arctic land areas so far. A considerable small-scale heterogeneity of the summer surface temperature is observed in long-term measurements with a thermal imaging system, which can be reproduced in the energy balance model. The model can also simulate the impact of different snow depths on the soil temperature, that has been documented in field measurements. Furthermore, time series of terrestrial surface temperature measurements are compared to satellite-borne measurements, for which a significant cold-bias is observed during winter. Finally, different possibilities for a world-wide monitoring scheme are assessed. Energy budget models can incorporate different satellite data sets as training data sets for parameter estimation, so that they may constitute an alternative to purely satellite-based schemes.

Field measurements of the atmosphere, ocean, sea ice and sub-ice platelet layer at Atka Bay in 2013

Ice shelves strongly interact with coastal Antarctic sea ice and the associated ecosystem by creating conditions favourable to the formation of a sub-ice platelet layer. The close investigation of this phenomenon and its seasonal evolution remain a challenge due to logistical constraints and a lack of suitable methodology. In this study, we characterize the seasonal cycle of Antarctic fast ice adjacent to the Ekström Ice Shelf in the eastern Weddell Sea. We used a thermistor chain with the additional ability to record the temperature response induced by cyclic heating of resistors embedded in the chain. Vertical sea-ice temperature and heating profiles obtained daily between November 2012 and February 2014 were analyzed to determine sea-ice and snow evolution, and to calculate the basal energy budget. The residual heat flux translated into an ice-volume fraction in the platelet layer of 0.18 ± 0.09, which we reproduced by a independent model simulation and agrees with earlier results. Manual drillings revealed an average annual platelet-layer thickness increase of at least 4m, and an annual maximum thickness of 10m beneath second-year sea ice. The oceanic contribution dominated the total sea-ice production during the study, effectively accounting for up to 70% of second-year sea-ice growth. In summer, an oceanic heat flux of 21 W/m**2 led to a partial thinning of the platelet layer. Our results further show that the active heating method, in contrast to the acoustic sounding approach, is well suited to derive the fast-ice mass balance in regions influenced by ocean/ice-shelf interaction, as it allows sub-diurnal monitoring of the platelet-layer thickness.

Solar radiation over and under sea ice during the POLARSTERN cruise ARK-XXVI/3 (TransArc) in summer 2011

Arctic sea ice has declined and become thinner and younger (more seasonal) during the last decade. One consequence of this is that the surface energy budget of the Arctic Ocean is changing. While the role of surface albedo has been studied intensively, it is still widely unknown how much light penetrates through sea ice into the upper ocean, affecting sea-ice mass balance, ecosystems, and geochemical processes. Here we present the first large-scale under-ice light measurements, operating spectral radiometers on a remotely operated vehicle (ROV) under Arctic sea ice in summer. This data set is used to produce an Arctic-wide map of light distribution under summer sea ice. Our results show that transmittance through first-year ice (FYI, 0.11) was almost three times larger than through multi-year ice (MYI, 0.04), and that this is mostly caused by the larger melt-pond coverage of FYI (42 vs. 23%). Also energy absorption was 50% larger in FYI than in MYI. Thus, a continuation of the observed sea-ice changes will increase the amount of light penetrating into the Arctic Ocean, enhancing sea-ice melt and affecting sea-ice and upper-ocean ecosystems.

Heat fluxes and surface radiation measurements from Samoylov Island, Lena Delta, Siberia, October 2007 to March 2008

In this study, we present the winter time surface energy balance at a polygonal tundra site in northern Siberia based on independent measurements of the net radiation, the sensible heat flux and the ground heat flux from two winter seasons. The latent heat flux is inferred from measurements of the atmospheric turbulence characteristics and a model approach. The long-wave radiation is found to be the dominant factor in the surface energy balance. The radiative losses are balanced to about 60 % by the ground heat flux and almost 40 % by the sensible heat fluxes, whereas the contribution of the latent heat flux is small. The main controlling factors of the surface energy budget are the snow cover, the cloudiness and the soil temperature gradient. Large spatial differences in the surface energy balance are observed between tundra soils and a small pond. The ground heat flux released at a freezing pond is by a factor of two higher compared to the freezing soil, whereas large differences in net radiation between the pond and soil are only observed at the end of the winter period. Differences in the surface energy balance between the two winter seasons are found to be related to differences in snow depth and cloud cover which strongly affect the temperature evolution and the freeze-up at the investigated pond.

Daily polynya area and ice production in the North Water polynya region

The North Water (NOW) Polynya is a regularly-forming area of open-water and thin-ice, located between northwestern Greenland and Ellesmere Island (Canada) at the northern tip of Baffin Bay. Due to its large spatial extent, it is of high importance for a variety of physical and biological processes, especially in wintertime. Here, we present a long-term remote sensing study for the winter seasons 1978/1979 to 2014/2015. Polynya characteristics are inferred from (1) sea ice concentrations and brightness temperatures from passive microwave satellite sensors (Advanced Microwave Scanning Radiometer (AMSR-E and AMSR2), Scanning Multichannel Microwave Radiometer (SMMR), Special Sensor Microwave Imager/Sounder (SSM/I-SSMIS)) and (2) thin-ice thickness distributions, which are calculated using MODIS ice-surface temperatures and European Center for Medium-Range Weather Forecasts (ECMWF) atmospheric reanalysis data in a 1D thermodynamic energy-balance model. Daily ice production rates are retrieved for each winter season from 2002/2003 to 2014/2015, assuming that all heat loss at the ice surface is balanced by ice growth. Two different cloud-cover correction schemes are applied on daily polynya area and ice production values to account for cloud gaps in the MODIS composites. Our results indicate that the NOW polynya experienced significant seasonal changes over the last three decades considering the overall frequency of polynya occurrences, as well as their spatial extent. In the 1980s, there were prolonged periods of a more or less closed ice cover in northern Baffin Bay in winter. This changed towards an average opening on more than 85% of the days between November and March during the last decade. Noticeably, the sea ice cover in the NOW polynya region shows signs of a later-appearing fall freeze-up, starting in the late 1990s. Different methods to obtain daily polynya area using passive microwave AMSR-E/AMSR2 data and SSM/I-SSMIS data were applied. A comparison with MODIS data (thin-ice thickness < 20 cm) shows that the wintertime polynya area estimates derived by MODIS are about 30 to 40% higher than those derived using the polynya signature simulation method (PSSM) with AMSR-E data. In turn, the difference in polynya area between PSSM and a sea ice concentration (SIC) threshold of 70% is fairly low (approximately 10%) when applied to AMSR-E data. For the coarse-resolution SSM/I-SSMIS data, this difference is much larger, particularly in November and December. Instead of a sea ice concentration threshold, the PSSM method should be used for SSM/I-SSMIS data. Depending on the type of cloud-cover correction, the calculated ice production based on MODIS data reaches an average value of 264.4 ± 65.1 km**3 to 275.7 ± 67.4 km**3 (2002/2003 to 2014/2015) and shows a high interannual variability. Our achieved long-term results underline the major importance of the NOW polynya considering its influence on Arctic ice production and associated atmosphere/ocean processes.

Trematomus bernacchii experimental data files

As a response to ocean warming, shifts in fish species distribution and changes in production have been reported that have been partly attributed to temperature effects on the physiology of animals. The Southern Ocean hosts some of the most rapidly warming regions on earth and Antarctic organisms are reported to be especially temperature sensitive. While cellular and molecular organismic levels appear, at least partially, to compensate for elevated temperatures, the consequences of acclimation to elevated temperature for the whole organism are often less clear. Growth and reproduction are the driving factors for population structure and abundance. The aim of this study was to assess the effect of long-term acclimation to elevated temperature on energy budget parameters in the high-Antarctic fish Trematomus bernacchii. Our results show a complete temperature compensation for routine metabolic costs after 9 weeks of acclimation to 4°C. However, an up to 84% reduction in mass growth was measured at 2 and 4°C compared with the control group at 0°C, which is best explained by reduced food assimilation rates at warmer temperatures. With regard to a predicted temperature increase of up to 1.4°C in the Ross Sea by 2200, such a significant reduction in growth is likely to affect population structures in nature, for example by delaying sexual maturity and reducing production, with severe impacts on Antarctic fish communities and ecosystems.