Mechanisms of enhanced cardiorespiratory performance under hyperoxia differ with exposure duration in yellowtail kingfish.

Hyperoxia has been shown to expand the aerobic capacity of some fishes, although there have been very few studies examining the underlying mechanisms and how they vary across different exposure durations. Here, we investigated the cardiorespiratory function of yellowtail kingfish (Seriola lalandi) acutely (~20 h) and chronically (3-5 weeks) acclimated to hyperoxia (~200% air saturation). Our results show that the aerobic performance of kingfish is limited in normoxia and increases with environmental hyperoxia. The aerobic scope was elevated in both hyperoxia treatments driven by a ~33% increase in maximum O2 uptake (MO2max), although the mechanisms differed across treatments. Fish acutely transferred to hyperoxia primarily elevated tissue O2 extraction, while increased stroke volume-mediated maximum cardiac output was the main driving factor in chronically acclimated fish. Still, an improved O2 delivery to the heart in chronic hyperoxia was not the only explanatory factor as such. Here, maximum cardiac output only increased in chronic hyperoxia compared with normoxia when plastic ventricular growth occurred, as increased stroke volume was partly enabled by an ~8%-12% larger relative ventricular mass. Our findings suggest that hyperoxia may be used long term to boost cardiorespiratory function potentially rendering fish more resilient to metabolically challenging events and stages in their life cycle.

Thermal constraints on exercise and metabolic performance do not explain the use of dormancy as an overwintering strategy in the cunner (Tautogolabrus adspersus).

Winter cold slows ectotherm physiology, potentially constraining activities and ecological opportunities at poleward latitudes. Yet, many fishes are winter-active, facilitated by thermal compensation that improves cold performance. Conversely, winter-dormant fishes (e.g. cunner, Tautogolabrus adspersus) become inactive and non-feeding overwinter. Why are certain fishes winter-dormant? We hypothesized that winter dormancy is an adaptive behavioural response arising in poleward species that tolerate severe, uncompensated constraints of cold on their physiological performance. We predicted that below their dormancy threshold of 7--8°C, exercise and metabolic performance of cunner are greatly decreased, even after acclimation (i.e. shows above-normal, uncompensated thermal sensitivity, Q10>1-3). We measured multiple key performance metrics (e.g. C-start maximum velocity, chase swimming speed, aerobic scope) in cunner after acute exposure to 26-2°C (3°C intervals using 14°C-acclimated fish) or acclimation (5-8 weeks) to 14-2°C (3°C intervals bracketing the dormancy threshold). Performance declined with cooling, and the acute Q10 of all six performance rate metrics was significantly greater below the dormancy threshold temperature (Q10,acute8-2°C=1.5-4.9, mean=3.3) than above (Q10,acute14-8°C=1.1-1.9, mean=1.5), inferring a cold constraint. However, 2°C acclimation (temporally more relevant to seasonal cooling) improved performance, abolishing the acute constraint (Q10,acclimated8-2°C=1.4-3.0, mean=2.0; also cf. Q10,acclimated14-8°C=1.2-2.9, mean=1.7). Thus, dormant cunner show partial cold-compensation of exercise and metabolic performance, similar to winter-active species. However, responsiveness to C-start stimuli was greatly cold-constrained even following acclimation, suggesting dormancy involves sensory limitation. Thermal constraints on metabolic and exercise physiology are not significant drivers of winter dormancy in cunner. In fact, compensatory plasticity at frigid temperatures is retained even in a dormant fish.

Coronary circulation enhances the aerobic performance of wild Pacific salmon.

Female Pacific salmon often experience higher mortality than males during their once-in-a-lifetime up-river spawning migration, particularly when exposed to secondary stressors (e.g. high temperatures). However, the underlying mechanisms remain unknown. One hypothesis is that female Pacific salmon hearts are more oxygen-limited than those of males and are less able to supply oxygen to the body's tissues during this demanding migration. Notably, female hearts have higher coronary blood flow, which could indicate a greater reliance on this oxygen source. Oxygen limitations can develop from naturally occurring coronary blockages (i.e. coronary arteriosclerosis) found in mature salmon hearts. If female hearts rely more heavily on coronary blood flow but experience similar arteriosclerosis levels as males, they will have disproportionately impaired aerobic performance. To test this hypothesis, we measured resting (RMR) and maximum metabolic rate (MMR), aerobic scope (AS) and acute upper thermal tolerance in coho salmon (Oncorhynchus kisutch) with an intact or artificially blocked coronary oxygen supply. We also assessed venous blood oxygen and chemistry (cortisol, ions and metabolite concentrations) at different time intervals during recovery from exhaustive exercise. We found that coronary blockage impaired MMR, AS and the partial pressure of oxygen in venous blood (PvO2) during exercise recovery but did not differ between sexes. Coronary ligation lowered acute upper thermal tolerance by 1.1°C. Although we did not find evidence of enhanced female reliance on coronary supply, our findings highlight the importance of coronary blood supply for mature wild salmon, where migration success may be linked to cardiac performance, particularly during warm water conditions.

Hyperoxia disproportionally benefits the aerobic performance of large fish at elevated temperature.

Increasing evidence shows that larger fish are more vulnerable to acute warming than smaller individuals of the same species. This size-dependency of thermal tolerance has been ascribed to differences in aerobic performance, largely owing to a decline in oxygen supply relative to demand. To shed light on these ideas, we examined metabolic allometry in 130 rainbow trout ranging from 12 to 358 g under control conditions (17°C) and in response to acute heating (to 25°C), with and without supplemental oxygen (100% versus 150% air saturation). Under normoxia, high temperature caused an average 17% reduction in aerobic scope compared with 17°C. Aerobic performance disproportionally deteriorated in bigger fish as the scaling exponent (b) for aerobic scope declined from b=0.87 at 17°C to b=0.74 at 25°C. Hyperoxia increased maximum metabolic rate and aerobic scope at both temperatures and disproportionally benefited larger fish at 25°C as the scaling exponent for aerobic scope was reestablished to the same level as at 17°C (b=0.86). This suggests that hyperoxia may provide metabolic refuge for larger individuals, allowing them to sustain aerobic activities when facing acute warming. Notably, the elevated aerobic capacity afforded by hyperoxia did not appear to improve thermal resilience, as mortality in 25°C hyperoxia (13.8%, n=4) was similar to that in normoxia (12.1%, n=4), although we caution that this topic warrants more targeted research. We highlight the need for mechanistic investigations of the oxygen transport system to determine the consequences of differential metabolic scaling across temperature in a climate warming context.

Thermal performance curves for aerobic scope and specific dynamic action in a sexually dimorphic piscivore: implications for a warming climate.

Digestion can make up a substantial proportion of animal energy budgets, yet our understanding of how it varies with sex, body mass and ration size is limited. A warming climate may have consequences for animal growth and feeding dynamics that will differentially impact individuals in their ability to efficiently acquire and assimilate meals. Many species, such as walleye (Sander vitreus), exhibit sexual size dimorphism (SSD), whereby one sex is larger than the other, suggesting sex differences in energy acquisition and/or expenditure. Here, we present the first thorough estimates of specific dynamic action (SDA) in adult walleye using intermittent-flow respirometry. We fed male (n=14) and female (n=9) walleye two ration sizes, 2% and 4% of individual body mass, over a range of temperatures from 2 to 20°C. SDA was shorter in duration and reached higher peak rates of oxygen consumption with increasing temperature. Peak SDA increased with ration size and decreased with body mass. The proportion of digestible energy lost to SDA (i.e. the SDA coefficient) was consistent at 6% and was unrelated to temperature, body mass, sex or ration size. Our findings suggest that sex has a negligible role in shaping SDA, nor is SDA a contributor to SSD for this species. Standard and maximum metabolic rates were similar between sexes but maximum metabolic rate decreased drastically with body mass. Large fish, which are important for population growth because of reproductive hyperallometry, may therefore face a bioenergetic disadvantage and struggle most to perform optimally in future, warmer waters.

Hyperoxia Does Not Improve the Acute Upper Thermal Tolerance of a Tropical Marine Fish (Lutjanus apodus).

Fish can experience hyperoxia in shallow environments due to photosynthetic activity, and this has been suggested to provide them with a metabolic refuge during acute warming. However, this hypothesis has never been tested on a tropical marine species. Thus, we fitted 29°C-acclimated wild schoolmaster snapper (Lutjanus apodus; a species known to experience diel hyperoxia in mangrove creeks and coastal waters) with Transonic® flow probes, and exposed them to an acute increase in temperature (at 1°C h-1) in respirometers under normoxia and hyperoxia (150% air saturation), until their critical thermal maximum (CTmax). The CTmax of both groups was ∼39°C, and no differences in maximum cardiac function were recorded as the fish were warmed. However, temperature-induced factorial aerobic scope was significantly greater in fish tested under hyperoxia. These data suggest that hyperoxia will not protect coastal tropical fish species during marine heat waves, despite its effects on metabolic scope / capacity.

Physiologically-informed predictions of climate warming effects on native and non-native populations of blue catfish.

Blue catfish Ictalurus furcatus has been widely introduced throughout the United States to enhance recreational fisheries. Its success in both its native and non-native range, especially in the context of climate change, will be influenced by its thermal performance. We conducted a laboratory experiment to investigate the responses of wild-captured, subadult blue catfish to temperatures ranging from 7 °C to 38 °C. Blue catfish had relatively low standard metabolic rates, indicating low energetic demands, and hence an ability to survive well even during low-food conditions. Metabolic scope and food consumption rate increased with temperature, with metabolic scope peaking at 29.1 °C, and consumption rate peaking at 32 °C. Body condition remained high up to 32 °C, but decreased drastically thereafter, suggesting limitations in maintaining metabolism through food consumption at temperatures >32 °C; blue catfish cannot survive in such habitats indefinitely. Yet, many fish were able to survive temperatures as high as 38 °C for 5 days, suggesting that acute and occasionally chronic heat waves will not limit this species. Using these results, we also predicted the performance of blue catfish under prevailing conditions and under climate warming at seven locations throughout their current range in the U.S. We found that some blue catfish populations in southern and southeastern areas will likely experience temperatures above the optimal temperature for extended periods due to climate change, thus limiting potential habitat availability for this species. But, many non-native populations, especially those in northern areas such as Idaho, North Dakota, and northern California, may benefit from the expected warmer temperatures during spring and fall.

Comparing methods for determining the metabolic capacity of lumpfish (Cyclopterus lumpus Linnaeus 1758).

Lumpfish (Cyclopterus lumpus) mortalities have been reported during the summer at some North Atlantic salmon cage-sites where they serve as "cleaner fish." To better understand this species' physiology and whether limitations in their metabolic capacity and thermal tolerance can explain this phenomenon, we compared the aerobic scope (AS) of 6°C-acclimated lumpfish (~50 g and 8.8 cm in length at the beginning of experiments) when all individuals (N = 12) were given a chase to exhaustion, a critical swim speed (Ucrit) test, and a critical thermal maximum (CTMax) test (rate of warming 2°C h-1). The Ucrit and CTMax of the lumpfish were 2.36 ± 0.08 body lengths per second and 20.6 ± 0.3°C. The AS of lumpfish was higher during the Ucrit test (206.4 ± 8.5 mg O2 kg-1 h-1) versus that measured in either the CTMax test or after the chase to exhaustion (141.0 ± 15.0 and 124.7 ± 15.5 mg O2 kg-1 h-1, respectively). Maximum metabolic rate (MMR), AS, and "realistic" AS (ASR) measured using the three different protocols were not significantly correlated, indicating that measurements of metabolic capacity using one of these methods cannot be used to estimate values that would be obtained using another method. Additional findings include that (1) the lumpfish's metabolic capacity is comparable to that of Atlantic cod, suggesting that they are not as "sluggish" as previously suggested in the literature, and (2) their CTMax (20.6°C when acclimated to 6°C), in combination with their recently determined ITMax (20.6°C when acclimated to 10°C), indicates that high sea-cage temperatures are unlikely to be the primary cause of lumpfish mortalities at salmon sea-cages during the summer.

The allometric scaling of oxygen supply and demand in the California horn shark, Heterodontus francisci.

The gill surface area of aquatic ectotherms is thought to be closely linked to the ontogenetic scaling of metabolic rate, a relationship that is often used to explain and predict ecological patterns across species. However, there are surprisingly few within-species tests of whether metabolic rate and gill area scale similarly. We examined the relationship between oxygen supply (gill area) and demand (metabolic rate) by making paired estimates of gill area with resting and maximum metabolic rates across ontogeny in the relatively inactive California horn shark, Heterodontus francisci. We found that the allometric slope of resting metabolic rate was 0.966±0.058 (±95% CI), whereas that of maximum metabolic rate was somewhat steeper (1.073±0.040). We also discovered that the scaling of gill area shifted with ontogeny: the allometric slope of gill area was shallower in individuals <0.203 kg in body mass (0.564±0.261), but increased to 1.012±0.113 later in life. This appears to reflect changes in demand for gill-oxygen uptake during egg case development and immediately post hatch, whereas for most of ontogeny, gill area scales in between that of resting and maximum metabolic rate. These relationships differ from predictions of the gill oxygen limitation theory, which argues that the allometric scaling of gill area constrains metabolic processes. Thus, for the California horn shark, metabolic rate does not appear limited by theoretical surface-area-to-volume ratio constraints of gill area. These results highlight the importance of data from paired and size-matched individuals when comparing physiological scaling relationships.

Oxygen limitation may affect the temperature and size dependence of metabolism in aquatic ectotherms.

Both oxygen and temperature are fundamental factors determining metabolic performance, fitness, ecological niches, and responses of many aquatic organisms to climate change. Despite the importance of physical and physiological constraints on oxygen supply affecting aerobic metabolism of aquatic ectotherms, ecological theories such as the metabolic theory of ecology have focused on the effects of temperature rather than oxygen. This gap currently impedes mechanistic models from accurately predicting metabolic rates (i.e., oxygen consumption rates) of aquatic organisms and restricts predictions to resting metabolism, which is less affected by oxygen limitation. Here, we expand on models of metabolic scaling by accounting for the role of oxygen availability and temperature on both resting and active metabolic rates. Our model predicts that oxygen limitation is more likely to constrain metabolism in larger, warmer, and active fish. Consequently, active metabolic rates are less responsive to temperature than are resting metabolic rates, and metabolism scales to body size with a smaller exponent whenever temperatures or activity levels are higher. Results from a metaanalysis of fish metabolic rates are consistent with our model predictions. The observed interactive effects of temperature, oxygen availability, and body size predict that global warming will limit the aerobic scope of aquatic ectotherms and may place a greater metabolic burden on larger individuals, impairing their physiological performance in the future. Our model reconciles the metabolic theory with empirical observations of oxygen limitation and provides a formal, quantitative framework for predicting both resting and active metabolic rate and hence aerobic scope of aquatic ectotherms.

Thermal optimality and physiological parameters inferred from experimental studies scale latitudinally with marine species occurrences.

Ocean warming is expected to occur due to anthropogenic climate change bringing a spatial shift of marine communities. Experimental data that characterize the aerobic power budget via an aerobic scope, thermal metabolic scope, or thermal preferences have been proposed as tools that can describe species distribution since they characterize species fitness or performance under different temperatures. This study tested the potential relationship between observed occurrences and different physiological studies in the Americas for 11 commercially important species in Mexico. Projections were also developed for Mexico's exclusive economic zone under different climate warming scenarios. The physiological data were fitted from optimum up to pejus temperatures and projected to sea surface temperatures for present (2003-2014) and Representative Concentration Pathway (RCP) scenarios (RCP 2.6, RCP 4.5, RCP 6.0, and RCP 8.5) for the period 2040-2050 and 2090-2100. For species with wide distributions in the Americas, the number of occurrences reported decreases at higher latitudes related to the decrease in species performance calculated from laboratory experiments. In addition, higher species occurrences are usually reported around optimum temperatures. Overall, the results suggest that pejus temperatures likely restrict latitudinal distribution, at least for widely distributed taxons. Regarding Mexican projections, the results varied widely by species. For example, in the Atlantic Ocean, Octopus maya and Panulirus argus are vulnerable to warming scenarios, while Centropomus undecimalis is not. Interestingly, northern Campeche Bank, the Gulf of California, and Western Baja California may act as thermal refugia for marine species indicating they could be assigned as protected areas to support fisheries throughout the Mexican exclusive economic zone. This research adds to the increasing evidence of the relationship between thermal niche and wild population distribution.

Aerobic response to thermal stress across ontogeny and habitats in a teleost fish.

Near-future climate change projections predict an increase in sea surface temperature that is expected to have significant and rapid effects on marine ectotherms, potentially affecting a number of critical life processes. Some habitats also undergo more thermal variability than others, and the inhabitants therefore must be more tolerant to acute periods of extreme temperatures. Mitigation of these outcomes may occur through acclimation, plasticity or adaptation, although the rate and extent of a species' ability to adjust to warmer temperatures is largely unknown, specifically as it pertains to effects on various performance metrics in fishes that inhabit multiple habitats throughout ontogenetic stages. Here, the thermal tolerance and aerobic performance of schoolmaster snapper (Lutjanus apodus Walbaum, 1792) collected from two different habitats were experimentally assessed under different warming scenarios (temperature treatments = 30, 33, 35, 36°C) to assess vulnerability to an imminently changing thermal habitat. Larger subadult and adult fish collected from a 12 m deep coral reef exhibited a lower critical thermal maximum (CTmax ) compared to smaller juvenile fish collected from a 1 m deep mangrove creek. However, the CTmax of the creek-sampled fish was only 2°C above the maximum water temperature measured in the habitat from which they were collected, compared to a CTmax that was 8°C higher in the reef-sampled fish, resulting in a wider thermal safety margin at the reef site. A generalized linear model showed a marginally significant effect of temperature treatment on resting metabolic rate (RMR), but there were no effects of any of the tested factors on maximum metabolic rate or absolute aerobic scope. Post hoc tests revealed that RMR was significantly higher for creek-collected fish at the 36°C treatment and significantly higher for reef-collected fish at 35°C. Swimming performance [measured by critical swimming speed] was significantly lower at the highest temperature treatment for creek-collected fish and trended down with each successive increase in temperature treatment for reef-collected fish. These results show that metabolic rate and swimming performance responses to thermal challenges are somewhat consistent across collection habitats, and this species may be susceptible to unique types of thermal risk depending on its habitat. We show the importance of intraspecific studies that couple habitat profiles and performance metrics to better understand possible outcomes under thermal stress.

Physiological effects of temperature on Greenland halibut Reinhardtius hippoglossoides shows high vulnerability of Arctic stenotherms to global warming.

Global warming affects the metabolism of ectothermic aquatic breathers forcing them to migrate and undergo high-latitudinal distribution shifts to circumvent the temperature-induced mismatch between increased metabolic demand and reduced water oxygen availability. Here the authors examined the effects of temperature on oxygen consumption rates in an Arctic stenotherm, the Greenland halibut Reinhardtius hippoglossoides, and calculated the optimal temperature for maximum aerobic scope, AS(Topt,AS ), which was found to be 2.44°C. They also investigated cardiac performance as limiting the oxygen transport chain at high temperatures by measuring maximum heart rate (fHmax ) over acute temperature increases and found various metrics related to fHmax to be at least 3.2°C higher than Topt,AS . The authors' measured Topt,AS closely reflected in situ temperature occurrences of Greenland halibut from long-term tagging studies, showing that AS of the species is adapted to its habitat temperature, and is thus a good proxy for the species' sensitivity to environmental warming. The authors did not find a close connection between fHmax and Topt,AS , suggesting that cardiac performance is not limiting for the oxygen transport chain at high temperatures in this particular Arctic stenotherm. The authors' estimate of the thermal envelope for AS of Greenland halibut was from -1.89 to 8.07°C, which is exceptionally narrow compared to most other species of fish. As ocean temperatures increase most rapidly in the Arctic in response to climate change, and species in these areas have limited possibility for further poleward-range shifts, these results suggest potential severe effects of global warming on Arctic stenotherms, such as the Greenland halibut. The considerable economic importance of the species raises concerns for future fisheries and species conservation of Arctic stenotherms in the Northern Hemisphere.

Ontogeny and ecological significance of metabolic rates in sea turtle hatchlings.

BACKGROUND: Sea turtle hatchlings must avoid numerous predators during dispersal from their nesting beaches to foraging grounds. Hatchlings minimise time spent in predator-dense neritic waters by swimming almost continuously for approximately the first 24 h post-emergence, termed the 'frenzy'. Post-frenzy, hatchling activity gradually declines as they swim in less predator-dense pelagic waters. It is well documented that hatchlings exhibit elevated metabolic rates during the frenzy to power their almost continuous swimming, but studies on post-frenzy MRs are sparse. RESULTS: We measured the frenzy and post-frenzy oxygen consumption of hatchlings of five species of sea turtle at different activity levels and ages to compare the ontogeny of mass-specific hatchling metabolic rates. Maximal metabolic rates were always higher than resting metabolic rates, but metabolic rates during routine swimming resembled resting metabolic rates in leatherback turtle hatchlings during the frenzy and post-frenzy, and in loggerhead hatchlings during the post-frenzy. Crawling metabolic rates did not differ among species, but green turtles had the highest metabolic rates during frenzy and post-frenzy swimming. CONCLUSIONS: Differences in metabolic rate reflect the varying dispersal stratagems of each species and have important implications for dispersal ability, yolk consumption and survival. Our results provide the foundations for links between the physiology and ecology of dispersal of sea turtles.

Metabolic rate increases with acclimation temperature and is associated with mitochondrial function in some tissues of threespine stickleback.

The metabolic rate (MO2) of eurythermal fishes changes in response to temperature, yet it is unclear how changes in mitochondrial function contribute to changes in MO2. We hypothesized that MO2 would increase with acclimation temperature in the threespine stickleback (Gasterosteus aculeatus) in parallel with metabolic remodeling at the cellular level but that changes in metabolism in some tissues, such as liver, would contribute more to changes in MO2 than others. Threespine stickleback were acclimated to 5, 12 and 20°C for 7 to 21 weeks. At each temperature, standard and maximum metabolic rate (SMR and MMR, respectively), and absolute aerobic scope (AAS) were quantified, along with mitochondrial respiration rates in liver, oxidative skeletal and cardiac muscles, and the maximal activity of citrate synthase (CS) and lactate dehydrogenase (LDH) in liver, and oxidative and glycolytic skeletal muscles. SMR, MMR and AAS increased with acclimation temperature, along with rates of mitochondrial phosphorylating respiration in all tissues. Low SMR and MMR at 5°C were associated with low or undetectable rates of mitochondrial complex II activity and a greater reliance on complex I activity in liver, oxidative skeletal muscle and heart. SMR was positively correlated with cytochrome c oxidase (CCO) activity in liver and oxidative muscle, but not mitochondrial proton leak, whereas MMR was positively correlated with CCO activity in liver. Overall, the results suggest that changes in MO2 in response to temperature are driven by changes in some aspects of mitochondrial function in some, but not all, tissues of threespine stickleback.

Characterizing the hypoxic performance of a fish using a new metric: PAAS-50.

The hypoxic constraint on peak oxygen uptake (MO2,peak) was characterized in rainbow trout over a range of ambient oxygen tensions with different testing protocols and statistical models. The best-fit model was selected using both statistical criteria (R2 and AIC) and the model's prediction of three anchor points for hypoxic performance: critical PO2 (Pcrit), maximum MO2 and a new metric, the minimum PO2 that supports 50% of absolute aerobic scope (PAAS-50). The best-fitting model was curvilinear using five strategically selected PO2 values. This model predicted PAAS-50 as 70 mmHg (coefficient of variation, CV=9%) for rainbow trout. Thus, while a five-point hypoxic performance curve can characterize the limiting effects of hypoxia in fish, as envisaged by Fry over 75 years ago, PAAS-50 is a promising metric to compare hypoxic constraints on performance in a standardized manner both within and across fish species.

Energetic costs of ectoparasite infection in Atlantic salmon.

Parasites are widespread in nature, where they affect the energy budget of hosts, and depending on the imposed pathogenic severity, this may reduce host fitness. However, the energetic costs of parasite infections are rarely quantified. In this study, we measured metabolic rates in recently seawater adapted Atlantic salmon (Salmo salar) infected with the ectoparasitic copepod Lepeophtheirus salmonis and used an aerobic scope framework to assess the potential ecological impact of this parasite-host interaction. The early chalimus stages of L. salmonis did not affect either standard or maximum metabolic rates. However, the later mobile pre-adult stages caused an increase in both standard and maximum metabolic rate yielding a preserved aerobic scope. Notably, standard metabolic rates were elevated by 26%, presumably caused by increased osmoregulatory burdens and costs of mobilizing immune responses. The positive impact on maximum metabolic rates was unexpected and suggests that fish are able to transiently overcompensate energy production to endure the burden of parasites and thus allow for continuation of normal activities. However, infected fish are known to suffer reduced growth, and this suggests that a trade-off exists in acquisition and assimilation of resources despite an uncompromised aerobic scope. As such, when assessing impacts of environmental or biotic factors, we suggest that elevated routine costs may be a stronger predictor of reduced fitness than the available aerobic scope. Furthermore, studying the effects on parasitized fish in an ecophysiological context deserves more attention, especially considering interacting effects of other stressors in the Anthropocene.

Experimental hyperoxia (O2 supersaturation) reveals a gill diffusion limitation of maximum aerobic performance in fish.

Several studies have demonstrated that hyperoxia increases the maximal O2 consumption rate (MO2max) in fish, but exactly how this occurs remains to be explained. Here, we tested the hypothesis that hyperoxia improves arterial oxygenation in rainbow trout during exhaustive exercise. We demonstrate a 35% higher MO2max in hyperoxia (200% air saturation) relative to normoxia, which was achieved through a combined 15% increase in cardiac output due to elevated peak heart rate, and a 19% increase of the arterial-venous (A-V) O2 content difference. While arterial O2 partial pressure (PaO2) and O2 saturation of haemoglobin declined post-exhaustive exercise in normoxia, this did not occur in hyperoxia. This protective effect of hyperoxia on arterial oxygenation led to a 22% higher arterial O2 content post-exhaustive exercise, thereby allowing a higher A-V O2 content difference. These findings indicate that MO2max is gill diffusion limited in exhaustively exercised rainbow trout. Moreover, as previous studies in salmonids have demonstrated collapsing PaO2 in normoxia at maximal swimming speed and at acutely high temperatures, a diffusion limitation may constrain MO2 in other situations eliciting peak metabolic demand. These findings, along with the fact that hyperoxia increases MO2max in several other fishes, suggest that gill diffusion limitations of MO2max may be widespread in fishes.

Aerobic scope in fishes with different lifestyles and across habitats: Trade-offs among hypoxia tolerance, swimming performance and digestion.

Exercise and aerobic scope in fishes have attracted scientists' attention for several decades. While it has been suggested that aerobic scope may limit behavioral expression and tolerance to environmental stressors in fishes, the exact importance of aerobic scope in an ecological context remains poorly understood. In this review, we examine the ecological relevance of aerobic scope by reconsidering and reanalyzing the existing literature on Chinese freshwater fishes across a wide-range of habitats and lifestyles. The available evidence suggests that natural selection in fast-flowing aquatic habitats may favor species with a high aerobic scope and anaerobic capacity for locomotion, whereas in relatively slow-flowing habitats, hypoxia tolerance may be favored at the cost of reduced locomotor capacity. In addition, while physical activity can usually cause fishes from fast-flowing habitats to reach their aerobic metabolic ceiling (i.e., maximum metabolic rate), possibly due to selection pressure on locomotion, most species from slow-flowing habitats can only reach their metabolic ceiling during digestion, either alone or in combination with physical activity. Overall, we suggest that fish exhibit a continuum of metabolic types, from a 'visceral metabolic type' with a higher digestive performance to a 'locomotion metabolic type' which appears to have reduced capacity for digestion but enhanced locomotor performance. Generally, locomotor-type species can either satisfy the demands of their high swimming capacity with a high oxygen uptake capacity or sacrifice digestion while swimming. In contrast, most visceral-type species show a pronounced decrease in swimming performance while digesting, probably owing to conflicts within their aerobic scope. In conclusion, the ecological relevance of aerobic scope and the consequent effects on other physiological functions are closely related to habitat and the lifestyle of a given species. These results suggest that swimming performance, digestion and hypoxia tolerance might coevolve due to dependence on metabolic traits such as aerobic scope.

Prolonged cortisol elevation alters whole body and tissue metabolism in rainbow trout (Oncorhynchus mykiss).

Chronic elevation of circulating cortisol is known to have deleterious effects on fish, but information about the consequences of prolonged cortisol elevation on the metabolism of fish is scarce. To test the effects of chronic cortisol elevation on the aerobic performance of rainbow trout, we examined how two severities of chronically elevated plasma cortisol levels affected the oxygen uptake during rest and after exhaustive exercise using a high (HC) and a medium cortisol (MC) treatment. High cortisol doses significantly affected standard (SMR) and maximum metabolic rates (MMR) compared to control fish. In comparison, the medium cortisol treatment elevated maximum metabolic rates (MMR) but did not significantly influence SMR compared to a sham group (S) and control group (C). The medium cortisol treatment resulted in a significantly increased metabolic scope due to an elevation of MMR, an effect that was abolished in the HC group due to co-occuring elevations in SMR. The elevated SMR of the HC-treated fish could be explained by increased in vitro oxygen uptake rates (MO2) of specific tissues, indicating that the raised basal metabolism was caused, in part, by an increase in oxygen demand of specific tissues. Haematological results indicated an increased reliance on anaerobic metabolic pathways in cortisol-treated fish under resting conditions.