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Ocean warming and acidification are two important environmental drivers affecting marine organisms. Organisms living at high latitudes might be especially threatened in near future, as current environmental changes are larger and occur faster. Therefore, we investigated the effect of hypercapnia on thermal tolerance and physiological performance of sub-Arctic Mytilus edulis from the White Sea. Mussels were exposed (2 weeks) to 390 µatm (control) and 1120 µatm CO2 (year 2100) before respiration rate (MO2), anaerobic metabolite (succinate) level, haemolymph acid-base status and intracellular pH (pHi) were determined during acute warming (10-28 °C, 3 °C over night). In normocapnic mussels, warming induced MO2 to rise exponentially until it levelled off beyond a breakpoint temperature of 20.5 °C. Concurrently, haemolymph PCO2 rose significantly > 19 °C followed by a decrease in PO2 indicating the pejus temperature (TP, onset of thermal limitation). Succinate started to accumulate at 28 °C under normocapnia defining the critical temperature (TC). pHi was maintained during warming until it dropped at 28 °C, in line with the concomitant transition to anaerobiosis. At acclimation temperature, CO2 had only a minor impact. During warming, MO2 was stimulated by CO2 resulting in an elevated breakpoint of 25.8 °C. Nevertheless, alterations in haemolymph gases (> 16 °C) and the concomitant changes of pHi and succinate level (25 °C) occurred at lower temperature under hypercapnia versus normocapnia indicating a downward shift of both thermal limits TP and TC by CO2. Compared to temperate conspecifics, sub-Arctic mussels showed an enhanced thermal sensitivity, exacerbated further by hypercapnia, indicating their potential vulnerability to environmental changes projected for 2100. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Ongoing climate change is predicted to affect individual organisms during all life stages, thereby affecting populations of a species, communities and the functioning of ecosystems. These effects of climate change can be direct, through changing water temperatures and associated phenologies, the lengths and frequency of hypoxia events, through ongoing ocean acidification trends or through shifts in hydrodynamics and in sea level. In some cases, climate interactions with a species will also, or mostly, be indirect and mediated through direct effects on key prey species which change the composition and dynamic coupling of food webs. Thus, the implications of climate change for marine fish populations can be seen to result from phenomena at four interlinked levels of biological organization: (1) organismal-level physiological changes will occur in response to changing environmental variables such as temperature, dissolved oxygen and ocean carbon dioxide levels. An integrated view of relevant effects, adaptation processes and tolerance limits is provided by the concept of oxygen and capacity-limited thermal tolerance (OCLT). (2) Individual-level behavioural changes may occur such as the avoidance of unfavourable conditions and, if possible, movement into suitable areas. (3) Population-level changes may be observed via changes in the balance between rates of mortality, growth and reproduction. This includes changes in the retention or dispersion of early life stages by ocean currents, which lead to the establishment of new populations in new areas or abandonment of traditional habitats. (4) Ecosystem-level changes in productivity and food web interactions will result from differing physiological responses by organisms at different levels of the food web. The shifts in biogeography and warming-induced biodiversity will affect species productivity and may, thus, explain changes in fisheries economies. This paper tries to establish links between various levels of biological organization by means of addressing the effective physiological principles at the cellular, tissue and whole organism levels. [ABSTRACT FROM AUTHOR]

Dissolved oxygen (DO) concentration in the water column is an environmental parameter that is crucial for the successful development of many pelagic organisms. Hypoxia tolerance and threshold values are species- and stage-specific and can vary enormously. While some fish species may suffer from oxygen values of less than 3 mL O2 L-1 through impacted growth, development and behaviour, other organisms such as euphausiids may survive DO levels as low as 0.1 mL O2 L-1. A change in the average or the range of DO may have significant impacts on the survival of certain species and hence on the species composition in the ecosystem with consequent changes in trophic pathways and productivity. Evidence for the deleterious effects of oxygen depletion on pelagic species is scarce, particularly in terms of the effect of low oxygen on development, recruitment and patterns of migration and distribution. While planktonic organisms have to cope with variable DOs and exploit adaptive mechanisms, nektonic species may avoid areas of unfavourable DO and develop adapted migration strategies. Planktonic organisms may only be able to escape vertically, above or beneath the Oxygen Minimum Zone (OMZ). In shallow areas only the surface layer can serve as a refuge, but in deep waters many organisms have developed vertical migration strategies to use, pass through and cope with the OMZ. This paper elucidates the role of DO for different taxa in the pelagic realm and the consequences of low oxygen for foodweb structure and system productivity. We describe processes in two contrasting systems, the semi-enclosed Baltic Sea and the coastal upwelling system of the Benguela Current to demonstrate the consequences of increasing hypoxia on ecosystem functioning and services. [ABSTRACT FROM AUTHOR]

The concept of oxygen- and capacity-dependent thermal tolerance in aquatic ectotherms has successfully explained climate-induced effects of rising temperatures on species abundance in the field. Oxygen supply to tissues and the resulting aerobic performance characters thus form a primary link between organismal fitness and its role and functioning at the ecosystem level. The thermal window of performance in water breathers matches their window of aerobic scope. Loss of performance reflects the earliest level of thermal stress, caused by hypoxaemia and the progressive mismatch of oxygen supply and demand at the borders of the thermal envelope. Oxygen deficiency elicits the transition to passive tolerance and associated systemic and cellular stress signals like hormonal responses or oxidative stress as well as the use of protection mechanisms like heat shock proteins at thermal extremes. Thermal acclimatization between seasons or adaptation to a climate regime Involves shifting thermal windows and adjusting window widths. The need to specialize on a limited temperature range results from temperature- dependent trade-offs at several hierarchical levels, from molecular structure to whole-organism functioning, and may also support maximized energy efficiency. Various environmental factors like CO2 (ocean acidification) and hypoxia Interact with these principal relationships. Existing knowledge suggests that these factors elicit metabolic depression supporting passive tolerance to thermal extremes. However, they also exacerbate hypoxaemla, causing a narrowing of thermal performance windows and prematurely leading the organism to the limits of its thermal acclimation capacity. The conceptual analysis suggests that the relationships between energy turnover, the capacities of activity and other functions and the width of thermal windows may lead to an. Integrative understanding of specialization on climate and, as a thermal matrix, of sensitivity to climate change and,the factors Involved. Such functional relationships might also relate to climate-Induced changes in species Interactions and, thus, community responses at the ecosystem level. [ABSTRACT FROM AUTHOR]

Future ocean acidification has the potential to adversely affect many marine organisms. A growing body of evidence suggests that many species could suffer from reduced fertilization success, decreases in larval- and adult growth rates, reduced calcification rates, and even mortality when being exposed to near-future levels (year 2100 scenarios) of ocean acidification. Little research focus is currently placed on those organisms/taxa that might be less vulnerable to the anticipated changes in ocean chemistry; this is unfortunate, as the comparison of more vulnerable to more tolerant physiotypes could provide us with those physiological traits that are crucial for ecological success in a future ocean. Here, we attempt to summarize some ontogenetic and lifestyle traits that lead to an increased tolerance towards high environmental pCO2. In general, marine ectothermic metazoans with an extensive extracellular fluid volume may be less vulnerable to future acidification as their cells are already exposed to much higher pCO2 values (0.1 to 0.4 kPa, ca. 1000 to 3900μatm) than those of unicellular organisms and gametes, for which the ocean (0.04 kPa, ca. 400μatm) is the extracellular space. A doubling in environmental pCO2 therefore only represents a 10% change in extracellular pCO2 in some marine teleosts. High extracellular pCO2 values are to some degree related to high metabolic rates, as diffusion gradients need to be high in order to excrete an amount of CO2 that is directly proportional to the amount of O2 consumed. In active metazoans, such as teleost fish, cephalopods and many brachyuran crustaceans, exercise induced increases in metabolic rate require an efficient ion-regulatory machinery for CO2 excretion and acid-base regulation, especially when anaerobic metabolism is involved and metabolic protons leak into the extracellular space. These ion-transport systems, which are located in highly developed gill epithelia, form the basis for efficient compensation of pH disturbances during exposure to elevated environmental pCO2. Compensation of extracellular acid-base status in turn may be important in avoiding metabolic depression. So far, maintained "performance" at higher seawater pCO2 (>0.3 to 0.6 kPa) has only been observed in adults/juveniles of active, high metabolic species with a powerful ion regulatory apparatus. However, while some of these taxa are adapted to cope with elevated pCO2 during their regular embryonic development, gametes, zygotes and early embryonic stages, which lack specialized ion-regulatory epithelia, may be the true bottleneck for ecological success - even of the more tolerant taxa. Our current understanding of which marine animal taxa will be affected adversely in their physiological and ecological fitness by projected scenarios of anthropogenic ocean acidification is quite incomplete. While a growing amount of empirical evidence from CO2 perturbation experiments suggests that several taxa might react quite sensitively to ocean acidification, others seem to be surprisingly tolerant. However, there is little mechanistic understanding on what physiological traits are responsible for the observed differential sensitivities (see reviews of Seibel and Walsh, 2003; Pörtner et al., 2004; Fabry et al., 2008; Pörtner, 2008). This leads us to the first very basic question of how to define general CO2 tolerance on the species level. [ABSTRACT FROM AUTHOR]

Future scenarios for the oceans project combined developments of CO2 accumulation and global warming and their impact on marine ecosystems. The synergistic impact of both factors was addressed by studying the effect of elevated CO2 concentrations on thermal tolerance of the coldeurythermal spider crab Hyas araneus from the population around Helgoland. Here ambient temperatures characterize the southernmost distribution limit of this species. Animals were exposed to present day normocapnia (380 ppm CO2), CO2 levels expected towards 2100 (710 ppm) and beyond (3000 ppm). Heart rate and haemolymph PO2 (PeO2) were measured during progressive short term cooling from 10 to 0°C and during warming from 10 to 25°C. An increase of PeO2 occurred during cooling, the highest values being reached at 0°C under all three CO2 levels. Heart rate increased during warming until a critical temperature (Tc) was reached. The putative Tc under normocapnia was presumably >25°C, from where it fell to 23.5°C under 710 ppm and then 21.1°C under 3000 ppm. At the same time, thermal sensitivity, as seen in the Q10 values of heart rate, rose with increasing CO2 concentration in the warmth. Our results suggest a narrowing of the thermal window of Hyas araneus under moderate increases in CO2 levels by exacerbation of the heat or cold induced oxygen and capacity limitation of thermal tolerance. [ABSTRACT FROM AUTHOR]

Dissolved oxygen (DO) concentration in the water column is an environmental parameter that is crucial for the successful development of many pelagic organisms. Hypoxia tolerance and threshold values are species- and stage-specific and can vary enormously. While some fish species may suffer from oxygen values of less than 3 ml L-1 and show impact on growth, development and behaviour, other organisms such as euphausiids may survive DO levels as low as 0.1 ml L-1. A change in the average or the minimum or maximum DO in an area may have significant impacts on the survival of certain species and hence on the species composition in the ecosystem with consequent changes in trophic pathways and productivity. Evidence of the deleterious effects of oxygen depletion on species of the pelagic realm is scarce, particularly in terms of the effect of low oxygen on development, recruitment and patterns of migration and distribution. While planktonic organisms have to cope with different DOs and find adaptive mechanisms, nektonic species may avoid areas of inconvenient DO and develop adapted migrational strategies. Planktonic organisms may only be able to escape vertically, above or beneath the Oxygen Minimum Zone (OMZ). In shallow areas only the surface layer can serve as a refuge, in deep waters many organisms have developed vertical migration strategies to use, pass and cope with the OMZ. This paper elucidates the role of DO for different taxa in the pelagic realm and the consequences of low oxygen for foodweb structure and system productivity. [ABSTRACT FROM AUTHOR]

Future ocean acidification has the potential to adversely affect many marine organisms. A growing body of evidence suggests that many species could suffer from reduced fertilization success, decreases in larval- and adult growth rates, reduced calcification rates, metabolic depression and even mortality when being exposed to nearfuture levels (year 2100 scenarios) of ocean acidification. Little research focus is currently placed on those organisms/taxa that might be less vulnerable to the anticipated changes in ocean chemistry; this is unfortunate, as the comparison of more vulnerable to more tolerant physiotypes could provide us with those physiological traits that are crucial for ecological success in a future ocean. Here, we attempt to summarize some ontogenetic and lifestyle traits that lead to an increased tolerance towards high environmental pCO2. In general, marine ectothermic metazoans with an extensive extracellular fluid volume may be less vulnerable to future acidification as their cells are already exposed to much higher pCO2 values (0.1 to 0.4 kPa, 1000 to 4000 μatm) than those of unicellular organisms and gametes, for which the ocean (0.04 kPa, 400 μatm) is the extracellular space. A doubling in environmental pCO2 therefore only represents a 10% change in extracellular pCO2 in some marine teleosts. High extracellular pCO2 values are to some degree related to high metabolic rates, as diffusion gradients need to be high in order to excrete an amount of CO2 that is directly proportional to the amount of O2 consumed. In active metazoans, such as teleost fish, cephalopods and many brachyuran crustaceans, exercise induced increases in metabolic rate require an efficient ion-regulatory machinery for CO2 excretion and acid-base regulation, especially when anaerobic metabolism is involved and metabolic protons leak into the extracellular space. These ion-transport systems, which are located in highly developed gill epithelia, form the basis for efficient compensation of pH disturbances during exposure to elevated environmental pCO2. Compensation of extracellular acid-base status in turn may be extremely important in avoiding metabolic depression. So far, maintained "performance" at higher seawater pCO2 (>0.3 to 0.6 kPa) has only been observed in adults/juveniles of active, high metabolic species with a powerful ion regulatory apparatus. However, while some of these taxa are adapted to cope with elevated pCO2 during their regular embryonic development, unicellular gametes, which lack specialized ion-regulatory epithelia, may be the true bottleneck for ecological success - even of the more tolerant taxa. [ABSTRACT FROM AUTHOR]

Future scenarios project combined developments of elevated CO2 concentrations and global warming and their impact on marine ecosystems. The synergistic impact of both factors was addressed by studying the effect of CO2 accumulation on thermal tolerance of the cold-eurythermal spider crab Hyas araneus. Animals were exposed to present day normocapnia (380 ppmCO2), CO2 levels expected towards 2100 (710 ppm) and beyond (3000 ppm). Heart rate and haemolymph PO2 PeO2) were measured during progressive short term cooling from 10 to 0°C and during warming from 10 to 25°C. An increase of PeO2 occurred during cooling with highest values reached at 0°C under all three CO2 levels. Heart rate increased during warming until a critical temperature (Tc) was reached. The putative Tc under normocapnia was presumably >25°C, from where it fell to 23.5°C under 710 ppm and then 21.1°C under 3000 ppm. At the same time, thermal sensitivity, as seen in the Q10 values of heart rate, rose with increasing CO2 concentration in the warmth. Our results suggest a narrowing of the thermal window of Hyas araneus under moderate increases in CO2 levels by exacerbation of the heat or cold induced oxygen and capacity limitation of thermal tolerance. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Exposure of ectothermic organisms to variations in temperatures causes a transient mismatch between energy supply and demand, which needs to be compensated for during acclimation. Adenosine accumulation from ATP breakdown indicates such an imbalance and its reversal reflects a restoration of energy status. We monitored adenosine levels in blood serum and liver of common eelpout (Zoarces viviparus) during cold exposure in vivo. Furthermore, we tested its effect on the pattern of thermal acclimation in hepatocytes isolated from cold- (4°C) versus warm- (11°C) exposed fish. Adenosine levels increased during cold exposure in vivo and reached a transient maximum after 24 h in serum, but remained permanently elevated in liver. Whole animal cold acclimation induced a rise of liver citrate synthase activity by 44±15%, but left cytochrome c oxidase activity (COX) and RNA expression of the respective genes unchanged. Cold incubation of hepatocytes from warm-acclimated fish failed to cause an increase of mitochondrial enzyme activities despite increased COX4 mRNA levels. Conversely, warm acclimation of hepatocytes from cold-acclimated fish reduced both enzyme activities and COX2 and COX4 mRNA levels by 26-37%. Adenosine treatment of both warm- and cold-acclimated hepatocytes suppressed COX activities but activated COX mRNA expression. These effects were not receptor mediated. The present findings indicate that adenosine has the potential to regulate mitochondrial functioning in vivo, albeit the pathways resulting in the contrasting effects on expression and activity need to be identified. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Adjustments in mitochondrial properties and capacities are crucial in acclimatization to seasonal cold as well as in evolutionary cold adaptation of marine ectotherms. To examine whether gene expression mechanisms contribute to different settings of aerobic capacities in populations of cod (Gadus morhua) along a latitudinal cline, maximum activities of key enzymes of mitochondrial metabolism and their respective mRNA levels were compared in white muscle and liver of cold (4°C) and warm (10°C) acclimated individuals from cod populations of the North Sea and the Barents Sea, respectively. In white muscle, cold acclimation caused a parallel increase in citrate synthase (CS) and in cytochrome c oxidase (COX) activities, but with a much larger effect in the cold eurythermal Arctic population. In liver, cold acclimation was accompanied by increments in CS activities, but differences between populations were minor. Overall COX activities in liver were not affected by cold acclimation, but were higher in the cold adapted population. In both populations increments in muscle CS capacities were tightly correlated with elevated mRNA levels, suggesting transcriptional control of citrate synthase levels in muscle. In liver, CS mRNA levels differed between populations but were not affected by cold acclimation, so that post-transcriptional control may contribute to elevated functional levels in this tissue. Mitochondrial-encoded COX2 mRNA levels were not limiting for functional activities in both tissues, in favour of post-transcriptional control or limitations by other transcripts of the COX complex. Altogether, the differentiation in gene expression between both populations was more strongly expressed at 4°C. The comparison of functional levels and transcript levels may reflect genetic differentiation at functional sites, in line with genetic differences between the two populations previously established by non-coding genetic markers. [ABSTRACT FROM AUTHOR]

The levels of heat-shock proteins of the 70 kDa family (Hsp70s) were measured in different soft tissues of Atlantic cod Gadus morhua from different locations and after exposure to various thermal conditions: acute temperature increments (1° C day−1), mid-term (73 days at 4–15° C) and long-term thermal acclimation (278 days at 8–15° C), and seasonal and latitudinal temperature variations (field samples). Tissue specific distribution patterns of Hsp70s were observed: liver > gills > red blood cells > brain > white muscle. Thus, different tissues may have required different levels of protection by Hsp70s, and possibly this was related to the rate of protein synthesis. There were no differences in tissue Hsp70s between Arctic cod populations (Arctic, i.e. Barents and White Seas, Norwegian coast, and North or Baltic Seas). No changes in Hsp70s levels were observed in response to temperature variation of any intensity (acute fluctuation or seasonal and latitudinal) within the range of physiological temperatures (4–15° C) in wild and laboratory Atlantic cod. This confirms previous observations that changes in Hsp70 caused by such temperature variation are often small in fishes. Probably, the constitutive level of Hsp70s in Atlantic cod was high enough to overcome potentially harmful effects of temperature variations within the physiological range. A suppressing effect of high temperature (15° C) has already been observed at a systematic level (as reduced rate of somatic growth), whereas it is not reflected in modified Hsp70s. Therefore, Hsp70s apparently played a secondary role in defining thermal tolerance limits in Atlantic cod. These conclusions are in line with a recent concept of thermal tolerance which indicated that the first line of thermal limitation in the cold and warm is a loss in aerobic scope. [ABSTRACT FROM AUTHOR]

Previous studies in marine ectotherms from a latitudinal cline have led to the hypothesis that eurythermal adaptation to low mean annual temperatures is energetically costly. To obtain more information on the trade-offs and with that the constraints of thermal adaptation, mitochondrial functions were studied in subpolar lugworms (Arenicola marina L.) adapted to summer cold at the White Sea and were compared with those in boreal specimens from the North Sea, either acclimatized to summer temperatures or to winter cold. During summer, a comparison of mitochondria from subpolar and boreal worms revealed higher succinate oxidation rates and reduced Arrhenius activation energies (Ea ) in state 3 respiration at low temperatures, as well as higher proton leakage rates in subpolar lugworms. These differences reflect a higher aerobic capacity in subpolar worms, which is required to maintain motor activity at low but variable environmental temperatures--however, at the expense of an elevated metabolic rate. The lower activity of citrate synthase (CS) found in subpolar worms may indicate a shift in metabolic control within mitochondria. In contrast, acclimatization of boreal lugworms to winter conditions elicited elevated mitochondrial CS activities in parallel with enhanced mitochondrial respiration rates. With falling acclimation temperatures, the significant Arrhenius break temperature in state 3 respiration (11°C) became insignificant (5°C) or even disappeared (0°C) at lower levels of Arrhenius activation energies in the cold, similar to a phenomenon known from hibernating vertebrates. The efficiency of aerobic energy production in winter mitochondria rose as proton leakage in relation to state 3 decreased with cold acclimation, indicated by higher respiratory control ratio values and increased adenosine diphosphate/oxygen (ADP/O) ratios. These transitions indicate reduced metabolic flexibility, possibly paralleled by a loss in aerobic scope and metabolic... [ABSTRACT FROM AUTHOR]

The cold-stenothermal freshwater gadid Lota lota inhabiting the potamic regions of lowland rivers in central Europe, is exposed to summer temperatures up to 25 °C, which is far above the thermal preferendum of this species. Oxygen consumption rates, determined in field catches sampled at different times of the year, revealed that the basal metabolic rate is depressed during summer when water temperatures are high (152±16 μmol O2 100 g-1 h-1at 22 °C in July compared to 250±33 μmol O2 100 g-1 h-1 at 6 °C in November). This observation led us to investigate whether the observed depression of the metabolic rate is caused by oxygen limitation due to thermal impairment of the ventilatory system, as has been observed in other species. Determination of anaerobic end products (lactate and succinate) in the liver tissue of fish caught at different sampling dates did not show an accumulation of anaerobic end products during the summer, indicating no oxygen limitation. Measurements of enzyme activities in the white musculature and liver suggest that enzymes involved in aerobic metabolism were down-regulated during summer, which may have contributed to the observed reduction of metabolic rate. [ABSTRACT FROM AUTHOR]

Adjustments in mitochondrial properties and capacities are crucial in acclimatization to seasonal cold and in evolutionary cold adaptation of marine ectotherms. Although long-term compensatory increments in aerobic capacity of fish tissues have frequently been described in response to cold, much less is known about transitional phases and gene expression patterns involved. We investigated the time course of adjustment to acute cold in liver of eurythermal eelpout Zoarces viviparus. Whereas citrate synthase (CS) activity rose progressively in liver, cytochrome c oxidase (COX) activity was not altered during cold acclimation. Species-specific RNA probes were used to determine mRNA levels. CS mRNA (nuclear encoded) displayed a delayed, transient increase in response to cold, such that transcript levels did not parallel the change in enzyme activity. The enzyme activities and mRNA levels in the confamilial Antarctic Pachycara brachycephalum indicate cold compensation of CS activity in this cold-adapted species. The ratio of CS and COX activities was elevated in acclimation and adaptation to cold, indicating enhanced citrate synthesis over respiratory chain capacities in cold-adapted liver mitochondria. This may support enhanced lipid synthesis typically found in cold. The ratio of enzyme activity and transcript levels differed largely between Z. viviparus populations from the Baltic and North Seas, indicating the influence of unidentified parameters other than temperature. Transcript levels may not be tightly correlated with enzyme activities during thermal adaptation and thereafter. The time course of the acclimation process indicates that regulation at the translational and posttranslational levels predominates in adjustment to moderate thermal challenges. [ABSTRACT FROM AUTHOR]

The translational system was isolated from the gills of the Antarctic scallop Adamussium colbecki (Smith) and the European scallop Aequipecten opercularis (Linnaeus) for in vitro protein synthesis capacities (μg protein mg FW-1 day-1) and the translational capacities of RNA (kRNA in vitro mg protein mg RNA-1 day-1). In vitro protein synthesis capacity in the cold-adapted pectinid at 0 °C was similar to the one found in the temperate scallop at 25 °C. These findings might reflect cold compensated rates in Adamussium colbecki, partly explainable by high tissue levels of RNA. Cold-compensated in vitro protein synthesis capacities may further result from increments in the translational capacity of RNA. The thermal sensitivity of the translation machinery was slightly different in the two species, with significantly lower levels of Arrhenius activation energies Ea and Q10 in Adamussium colbecki in the temperature range 0–15 °C. Reduced protein synthesis and translational capacities were found in vitro in gills of long-term aquarium-maintained Adamussium colbecki and were accounted for by a loss of protein synthesis machinery, i.e. a reduction in RNA levels, as well as a decrease in the amount of protein synthesized per milligram of RNA (RNA translational capacity, kRNA in vitro). Such changes may involve food uptake or mirror metabolic depression strategies, like those occurring during winter. Consequences of high in vitro RNA translational capacities found in the permanently cold-adapted species are discussed in the context of seasonal food availability and growth rates at high latitudes. [ABSTRACT FROM AUTHOR]

We investigated the temperature dependence of some physiological parameters of common eelpout (Zoarces viviparus) from different locations (North Sea, Baltic Sea and Norwegian Sea) on acclimation temperature (3 °C and 12 °C) and acute temperature variation. The lethal limit of 12 °C-acclimated eelpout was determined as the critical thermal maximum [loss of equilibrium (LE) and onset of muscular spasms (OS)] and it was found to be 26.6 °C for LE and 28.8 °C for OS for all populations. However, these parameters do not have any relevant ecological interpretation. We therefore investigated the effect of gradually increased water temperature on standard metabolic rate (measured as resting oxygen consumption Mo2) and critical oxygen concentration ([O2]c) of eelpouts. Acclimation to low temperature (3 °C) resulted in partial compensation of Mo2, paralleled by a decrease of activation energy for Mo2 (from 82 kJ mol–1 at 12 °C to about 50 kJ mol–1 at 3 °C) in North Sea and Baltic Sea eelpouts. At the same time, Norwegian eelpout showed no acclimation of oxygen demand to warm temperature (12 °C) at all. The scope for eelpout aerobic metabolism shrank considerably with increased acclimation temperature, as [O2]c approached water oxygen concentrations. At 22.5±1 °C the [O2]c reached air saturation, which is equivalent to the upper critical temperature (TcII) and at this temperature the aerobic scope for the metabolism completely disappeared. In line with previous insight, the comparative analysis of the temperature dependence of Mo2 of Z. viviparus from different populations suggests that a pejus (sub-critical) temperature for this species is about 13–15 °C. In conclusion, the capacity to adjust aerobic metabolism relates to thermal tolerance and the bio-geographical distribution of the species. Global warming would thus be likely to cause a shift in the distribution of this species to the North. [ABSTRACT FROM AUTHOR]

The effect of acute increase in temperature on oxygen partial pressure (PO[SUB2]) was measured in)the gill arches of Atlantic cod Gadus morhua between 10 and 19 °C by use of oxygen microoptodes. Oxygen saturation of the gill blood under control conditions varied between 90 and 15% reflecting a variable percentage of arterial or venous blood in accordance with the position of each optode in the gill arch. The data obtained suggested that arterial Po[SUB2] remained more or less constant and arterial oxygen uptake did not become limiting during warming. A progressive drop in venous Po[SUB2], however, was observed at > 10 °C indicating that excessive oxygen uptake from the blood is not fully compensated for by circulatory performance, until finally, Po[SUB2] levels fully collapse. In a second set of experiments energy and acid-base status of white) muscle of Atlantic cod in vivo was measured by magnetic resonance ([SUP31]P-NMR) spectroscopy in unanaesthetized and unimmobilized fish in the temperature range between 13 and 21 °C. A (decrease in white muscle intracellular pH (pH[SUBi]) with temperature occurred between 10 and 16 °C (ΔpH per °C = -0.025 per °C). In white muscle temperature changes had no influence on high-energy phosphates such as phosphocreatine (PCr) or ATP except during exposure to high critical temperatures (> 16 °C), indicating that white muscle energy status appears to be relatively insensitive to thermal stress if compared to the thermal sensitivity of the whole animal. The data were consistent with the hypothesis of an oxygen limitation of thermal tolerance in animals, which is set by limited capacity of oxygen supply mechanisms. In the case of Atlantic cod circulatory rather than ventilatory performance may be the first process to cause oxygen deficiency during heat stress. [ABSTRACT FROM AUTHOR]

Activities of citrate synthase (CS), cytochrome c oxidase (CCO) and the electron transport system (ETS) were investigated in white muscle and liver of laboratory-maintained cod, Gadus morhua, from the North Sea, Norwegian coast and Barents Sea for an analysis of temperature acclimation and adaptation in aerobic metabolism. Cold acclimation within each population led to elevated activities of CS, CCO and ETS in white muscle. In liver, however, only North Sea cod showed cold-compensated CS activities, with CCO and ETS unchanged. In contrast, cold-acclimated Norwegian cod displayed unchanged enzyme activities, and Arctic cod showed elevated activities for CS, but decreased activities for CCO and no change in ETS. Between-population comparisons revealed clear evidence for permanent cold adaptation in white muscle of northern (Norwegian coast and Barents Sea) compared to boreal (North Sea) cod populations, reflected by higher activities of CS, CCO and ETS at the same acclimation temperature. Cold-compensated, mass-specific enzyme activities in liver were found in northern compared to boreal cod for CS and CCO, however, for ETS, after warm acclimation only. When evaluated as capacity in total liver, such activities were only found in northern populations in the case of CS at all temperatures, and for CCO and ETS at 15°C only. Hepatosomatic index (IH) and liver lipid contents were highest in North Sea cod, with similar IH but lower lipid contents in cold- versus warm-acclimated animals. An acclimation effect on IH was found in Norwegian cod only, with higher IH but unchanged lipid contents in the cold. In conclusion, permanent cold adaptation of muscle aerobic metabolism prevails in cod populations at higher latitudes, which is in line with permanently elevated rates of oxygen consumption observed in a parallel study. These differences reflect higher maintenance costs in cold-adapted versus cold-acclimated cod. [ABSTRACT FROM AUTHOR]

Among brooding species, passive and active means to provide oxygen to embryos can be observed. Among passive oxygen providers, lower oxygen availability in the center than at the periphery of embryo masses seems to delay development of inner embryos. We investigated the differences in patterns of oxygen supply to the periphery and the center of embryo masses in two active oxygen providers, the brachyuran crabs Cancer setosus and Homalaspis plana, and evaluated the consequences on: (1) the proportion of time that early- and late-stage embryos were exposed to low or high oxygen partial pressure (PO2), (2) oxygen consumption of the embryos from the center (inner) and the periphery (outer) of the embryo mass at those PO2 levels that the embryos experience throughout development, and (3) development of inner and outer embryos. We found that oxygen availability in the embryo masses of brachyuran crabs exhibited dramatic contrasts between the periphery and the center during early development and that these differences decreased throughout embryonic development. These dissimilar patterns of oxygen availability produced differences in the proportion of the time that the embryos were exposed to high and low PO2 levels throughout development. PO2 affected oxygen consumption of the inner and outer embryos in the same fashion, but the oxygen demand of inner embryos was lower. Furthermore, development of inner embryos was delayed, in comparison to outer embryos of the same female. We suggest that the asynchrony in the development of inner embryos, in comparison to outer embryos, is due to oxygen limitation, since oxygen availability affects embryonic oxygen consumption. The differences between development of inner and outer embryos is relatively small, when compared to other marine invertebrates, probably because female crabs are able to adjust oxygen supply to the embryos according their needs, while passive oxygen providers are not. However, active oxygen provision may affect investment in reproduction. Our results could have important implications both on studies of larval development and survival and in understanding the life-history tradeoffs of aquatic invertebrates. [ABSTRACT FROM AUTHOR]

Investment in reproduction ranges from gamete production to active parental care, and marine invertebrates span this range. However, the cost of parental care has not yet been systematically quantified, nor incorporated into life history studies of marine invertebrates, in contrast to most other animal taxa. Since oxygen is a limiting factor in egg masses of marine invertebrates, we studied patterns of oxygen partial pressure over time in embryo masses of Brachyuran crabs, and correlated these results with the cost of providing oxygen to the embryos. We found that: (1) oxygen is limiting in the embryo masses, (2) female crabs show an active brooding behaviour that we think helps to provide oxygen to the embryo mass, and (3) there is a substantial parental investment associated with brooding behaviours. Oxygen limitation and parental investment seem to be associated in many taxa of marine invertebrates, and we suggest that oxygen provision to the embryos may be a critical factor determining parental investment in this group. [ABSTRACT FROM AUTHOR]

Anaerobic metabolism and changes in the osmotic concentration of extravisceral fluid were studied in the White Sea periwinkles (Littorina littorea, Littorina saxatilis and Littorina obtusata) during freshwater exposure. Resistance to hypoosmotic stress increased in the order: L. obtusata < L. saxatilis < L. littorea. Our data suggest that osmotic shock is not a primary reason for mortality of the periwinkles under these conditions. During environmental anaerobiosis, considerable succinate accumulation (up to 10–19 μmol g-1 wet weight), and depletion of phosphagen and ATP pools were found in the studied species. Other metabolic end products (alanopine, strombine, lactate, acetate or propionate) were not detected. Succinate accumulation and net ATP breakdown were the fastest in the least resistant species, L. obtusata, and slowest in the most resistant, L. littorea. Rate of ATP turnover decreased during freshwater exposure in L. littorea and L. saxatilis, but not in L. obtusata. Our data suggest that differential resistance of three studied Littorina spp. to extreme hypoosmotic stress may be related to their different abilities to reduce metabolic rate and ATP turnover during sustained anoxia. Species-specific variations in anaerobic capacity of Littorina spp. are discussed in relation to their vertical distribution, size and ecology. [ABSTRACT FROM AUTHOR]

Parameters of acid-base and energy status were studied by in vivo 31P-nuclear magnetic resonance spectroscopy in three White Sea Littorina spp. (L.littorea, L. saxatilis and L. obtusata) during prolonged anaerobiosis in freshwater. Intracellular pH decreased significantly, especially during the early period of anaerobiosis, but later the decrease in intracellular pH slowed down considerably, suggesting a capacity for intracellular pH regulation in all three species. There was a trend for intracellular pH to fall most rapidly in the least freshwater-resistant species, L. obtusata, as compared to the most resistant, L. littorea. Non-bicarbonate, non-phosphate buffer values estimated by the homogenate technique were similar in the three studied species (28–37 mmol pH-1 kg-1 wet weight) and did not change during freshwater exposure. The CaCO3 buffer value of the foot tissues was considerably higher (171–218 mmol pH-1 kg-1 wet weight) and decreased significantly during freshwater exposure. The contribution of the multiple tissue buffering systems to intracellular pH regulation in Littorina spp. shifts between different stages of freshwater exposure. Initially, the non-bicarbonate, non-phosphate tissue buffering system seems to be of major importance for metabolic proton buffering at intracellular pH between 7.5 and 7.0. During later stages of anaerobiosis and at lower intracellular pH, the CaCO3 buffer is involved in proton buffering. Decrease in the CaCO3 buffer value during freshwater exposure was in quantitative agreement with the amount of metabolic protons buffered, thus suggesting that CaCO3 tissue stores may serve as a major buffering system during prolonged anaerobiosis in Littorina spp. [ABSTRACT FROM AUTHOR]

The thermal sensitivity of mitochondrial function was investigated in the stenothermal Antarctic fish Lepidonotothen nudifrons. State 3 respiration increases with increasing temperature between 0 °C and 18 °C with a Q10 of 2.43–2.63. State 4 respiration in the presence of oligomycin, an inhibitor of mitochondrial ATP synthase, quantifies the leakage of protons through the inner mitochondrial membrane, which causes oxygen consumption without concomitant ATP production. This parameter shows an unusually high Q10 of 4.21 ± 0.42 (0–18 °C), which indicates that proton leakage does not depend merely on ion diffusion but is an enzyme-catalysed process. The differential thermal sensitivity of oxidative phosphorylation (=state 3) and proton leakage (=state 4 in the presence of oligomycin) leads to progressive uncoupling of the mitochondria and decreased efficiency of oxidative phosphorylation under in vivo conditions if the body temperature of L. nudifrons increases. [ABSTRACT FROM AUTHOR]

Changes in oxygen consumption, ammonia excretion and in the acid-base and energy status of various tissues were investigated in the cold stenothermal Antarctic bivalve, Limopsis marionensis, and compared to similar data in the limpet, Nacella concinna, for an assessment of thermal sensitivity. Oxygen consumption of L. marionensis varied between −1.5 and 2°C with a Q10 of 2.2. Ammonia excretion could only be detected in animals exposed to elevated temperature for periods in excess of 45 days and close to death and it is interpreted as the onset of protein and amino acid catabolism with starvation under temperature stress. In L. marionensis any change in temperature as well as starvation stress at constant temperature induced a decrease in phospho- l-arginine and ATP levels. However, only temperature stress resulted in a drop in the Gibb's free energy change of ATP hydrolysis. Intracellular pH rose in all tissues during upward or downward temperature changes of only 1.5 or 2°C for 24 h with a concomitant trend to accumulate succinate and acetate in the tissues. These changes are seen to reflect disturbances of the tissue acid-base and energy status with any under- or overshoot in aerobic metabolic rate during a temperature decrease or increase. Elevated temperature at 2°C during 2 weeks of incubation resulted in continued net ATP depletion, at low levels of ATP free energy. This indicates long-term stress, which was also mirrored in the inability to establish a new steady-state mean rate of oxygen consumption. Incubation at even higher temperatures of 4 and 7°C led to an aggravation of energetic stress and transition to an intracellular acidosis, as well as a fall in oxygen consumption. In N. concinna a drop in energy levels was also visible at 2°C but was compensated for during long-term incubation. In conclusion, L. marionensis will be able to compensate for a temperature change only in a very narrow range whereas the thermal tolerance window is much wider in N. concinna. The inability of the metabolic rate to rise continually and the concomitant transition to anaerobic metabolism and long-term energetic stress characterize the upper critical temperature. Stenothermality is discussed, not only as reflecting the permanent and very stable low temperature in the natural environment, but also regarding dif- ferences in the level of activity and aerobic scope. [ABSTRACT FROM AUTHOR]

Temperature dependent changes in the mode of energy metabolism and in acid-base status were studied in the range from −1.7 to 26 °C in two populations of Arenicola marina collected in summer as well as in winter from intertidal flats of the North Sea (boreal) and the White Sea (subpolar). Extreme temperatures led to an accumulation of anaerobic end products, indicating the existence of both a low and a high critical temperature, beyond which anaerobic metabolism becomes involved in energy production. In summer animals from the North Sea the high critical temperature was found at temperatures above 20 °C, and the low critical temperature below 5 °C. Latitudinal or seasonal cold adaptation lead to a more or less parallel shift of both high and low critical temperature values to lower values. Between critical temperatures intracellular pH declined with rising temperature. Slopes varied between −0.012 and −0.022 pH- units/°C. In summer animals from the North Sea, the slope was slightly less than in White Sea animals, but differences appeared independent of the season. However, slopes were no longer linear beyond critical temperatures. A drop in intracellular pH at low temperatures coincided with the accumulation of volatile fatty acids in the body wall tissue of North Sea animals. A failure of active pHi adjustment is held responsible for the reduced ΔpHi/ΔT at temperatures above the high critical temperature. Extracellular pH was kept constant over the whole temperature range investigated. The ability of North Sea animals to adapt to temperatures beyond the critical temperature is poor compared to White Sea specimens. The larger range of temperature fluctuations at the White Sea is seen as a reason for the higher adaptational capacity of the subpolar animals. A hypothesis is proposed that among other mechanisms critical temperature values are set by an adjustment of mitochondrial density and thus, aerobic capacity. [ABSTRACT FROM AUTHOR]

Seven polymorphic microsatellite loci are described in the marine intertidal gastropod Littorina saxatilis. Preliminary data on allelic variation of these loci in a White Sea population of L. saxatilis are presented in order to assess their potential utility as high-resolution genetic markers for this model species. [ABSTRACT FROM AUTHOR]