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Competing Interests: Competing interests The authors declare no competing or financial interests.

A decline in size is increasingly recognised as a major response by ectothermic species to global warming. Mechanisms underlying this phenomenon are poorly understood but could include changes in energy balance of consumers, driven by declines in prey size coupled with increased energy demands due to warming. The sardine Sardina pilchardus is a prime example of animal shrinking, European populations of this planktivorous fish are undergoing profound decreases in body condition and adult size. This is apparently a bottom-up effect coincident with a shift towards increased reliance on smaller planktonic prey. We investigated the hypothesis that foraging on smaller prey would lead to increased rates of energy expenditure by sardines, and that such expenditures would be exacerbated by warming temperature. Using group respirometry we measured rates of energy expenditure indirectly, as oxygen uptake, by captive adult sardines offered food of two different sizes (0.2 or 1.2 mm items) when acclimated to two temperatures (16 °C or 21 °C). Energy expenditure during feeding on small items was tripled at 16 °C and doubled at 21 °C compared to large items, linked to a change in foraging mode between filter feeding on small or direct capture of large. This caused daily energy expenditure to increase by ~10 % at 16 °C and ~40 % at 21 °C on small items, compared to large items at 16 °C. These results support that declines in prey size coupled with warming could influence energy allocation towards life-history traits in wild populations. This bottom-up effect could partially explain the shrinking and declining condition of many small pelagic fish populations and may be contributing to the shrinking of other fish species throughout the marine food web. Understanding how declines in prey size can couple with warming to affect consumers is a crucial element of projecting the consequences for marine fauna of ongoing anthropogenic global change., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier B.V. All rights reserved.)

Atmospheric particulate matter (APM) produced by the steel industry comprises a complex mixture of particles that includes a wide variety of metals and metallic nanoparticles. These particles settle out onto areas surrounding the industries. There is evidence that this 'settleable' APM (SePM) may cause air-to-water cross-contamination with significant effects on aquatic biota. Recent investigations have reported sublethal impacts on the gill structure and blood oxygen-carrying capacity of fishes, which raises the hypothesis that there will be consequences for gas exchange capacity and ability to support aerobic activities. Therefore, we investigated the effects of an environmentally relevant level of SePM contamination on swimming performance and associated aerobic metabolic rates in Nile tilapia, Oreochromis niloticus. Short-term exposure (96 h) to SePM reduced critical swimming speed, energetic efficiency of aerobic swimming, standard metabolic rate, maximum metabolic rate, and aerobic scope. The compromised swimming performance could have adverse ecological effects by limiting foraging ability, predator evasion, territorial protection, and migration. The impairments to aerobic capacity could also affect overall fish performance by influencing long-term energy balance and allocation to growth and reproduction. Thus, despite being sublethal, SePM contamination is considerably debilitating, and if its limiting effects are not compensated for in the longer term, this may reduce the survival and fitness of fish populations., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier Inc. All rights reserved.)

The Atlantic bluefin tuna (ABFT) is a highly prized species of large pelagic fish. Studies of their environmental physiology may improve understanding and management of their populations, but this is difficult for mature adults because of their large size. Biologging of heart rate holds promise in investigating physiological responses to environmental conditions in free-swimming fishes but it is very challenging to anesthetize large ABFT for invasive surgery to place a tag in the body cavity near to the heart. We describe a novel method for rapid deployment of a commercially available heart-rate tag on ABFT, using an atraumatic trocar to implant it in the musculature associated with the cleithrum. We performed three sequential experiments to show that the tagging method (1) is consistently repeatable and reliable, (2) can be used successfully on commercial fishing boats and does not seem to affect fish survival, and (3) is effective for long-term deployments. In experiment 3, a tag logged heart rate over 80 days on a 60-kg ABFT held in a farm cage. The logged data showed that heart rate was sensitive to prevailing seasonal temperature and feeding events. At low temperatures, there were clear responses to feeding but these all disappeared above a threshold temperature of 25.5°C. Overall, the results show that our method is simple, rapid, and repeatable, and can be used for long-term experiments to investigate physiological responses by large ABFT to environmental conditions., (© 2023 The Authors. Journal of Fish Biology published by John Wiley & Sons Ltd on behalf of Fisheries Society of the British Isles.)

Eight juvenile European seabass were exposed to two thermal ramping protocols with different levels of aerobic activity and tolerance endpoint: the critical thermal maximum for swimming (CTSmax) while exercising aerobically until fatigue and the critical thermal maximum (CTmax) under static conditions until loss of equilibrium (LOE). In the CTSmax protocol, warming caused a profound increase in the rate of oxygen uptake (ṀO2), culminating in a gait transition from steady aerobic towards unsteady anaerobic swimming, then fatigue at 30.3±0.4°C (mean±s.e.m.). Gait transition and fatigue presumably indicate an oxygen limitation, which reflects the inability to meet the combined demands of swimming plus warming. The CTmax protocol also elicited an increase in ṀO2, culminating in LOE at 34.0±0.4°C, which is significantly warmer than fatigue at CTSmax. The maximum ṀO2 achieved in the CTmax protocol was, however, less than 30% of that achieved in the CTSmax protocol. Therefore, the static CTmax did not exploit full cardiorespiratory capacity for oxygen supply, indicating that LOE was not caused by systemic oxygen limitation. Consequently, systemic oxygen supply can be significant for tolerance of acute warming in seabass but this depends upon the physiological context and the endpoint used., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2023. Published by The Company of Biologists Ltd.)

Critical thermal maxima methodology (CTM) has been used to infer acute upper thermal tolerance in fishes since the 1950s, yet its ecological relevance remains debated. In this study, the authors synthesize evidence to identify methodological concerns and common misconceptions that have limited the interpretation of critical thermal maximum (CT max ; value for an individual fish during one trial) in ecological and evolutionary studies of fishes. They identified limitations of, and opportunities for, using CT max as a metric in experiments, focusing on rates of thermal ramping, acclimation regimes, thermal safety margins, methodological endpoints, links to performance traits and repeatability. Care must be taken when interpreting CTM in ecological contexts, because the protocol was originally designed for ecotoxicological research with standardized methods to facilitate comparisons within study individuals, across species and contexts. CTM can, however, be used in ecological contexts to predict impacts of environmental warming, but only if parameters influencing thermal limits, such as acclimation temperature or rate of thermal ramping, are taken into account. Applications can include mitigating the effects of climate change, informing infrastructure planning or modelling species distribution, adaptation and/or performance in response to climate-related temperature change. The authors' synthesis points to several key directions for future research that will further aid the application and interpretation of CTM data in ecological contexts., (© 2023 The Authors. Journal of Fish Biology published by John Wiley & Sons Ltd on behalf of Fisheries Society of the British Isles.)

Aquatic ecosystems can exhibit seasonal variation in resource availability and animals have evolved to cope with the associated caloric restriction. During winter in the NW Mediterranean Sea, the European sardine Sardina pilchardus naturally experiences caloric restriction owing to a decrease in the diversity and quantity of plankton. However, ongoing global warming has had deleterious effects on plankton communities such that food shortages may occur throughout the year, especially under warm conditions in the summer. We investigated the interactive effects of temperature and food availability on sardine metabolism by continuously monitoring whole-animal respiration of groups of control (fed) and food-deprived sardines over a 60-day experiment in winter (12°C) or summer (20°C) conditions under natural photoperiod. In addition, we measured mitochondrial respiration of red muscle fibres, biometric variables and energy reserves of individuals sampled at 30 and 60 days. This revealed that winter food deprivation elicits energy saving mechanisms at whole animal and cellular levels by maintaining a low metabolism to preserve energy reserves, allowing high levels of survival. By contrast, despite energy saving mechanisms at the mitochondrial level, whole animal metabolic rate was high during food deprivation in summer, causing increased consumption of energy reserves at the muscular level and high mortality after 60 days. Furthermore, a 5-day re-feeding did not improve survival, and mortalities continued, suggesting that long-term food deprivation at high temperatures causes profound stress in sardines that potentially impairs nutrient absorption., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2023. Published by The Company of Biologists Ltd.)

Aerobic metabolism generates 15-20 times more energy (ATP) than anaerobic metabolism, which is crucial in maintaining energy budgets in animals, fueling metabolism, activity, growth and reproduction. For ectothermic water-breathers such as fishes, low dissolved oxygen may limit oxygen uptake and hence aerobic metabolism. Here, we assess, within a phylogenetic context, how abiotic and biotic drivers explain the variation in hypoxia tolerance observed in fishes. To do so, we assembled a database of hypoxia tolerance, measured as critical oxygen tensions (P crit ) for 195 fish species. Overall, we found that hypoxia tolerance has a clear phylogenetic signal and is further modulated by temperature, body mass, cell size, salinity and metabolic rate. Marine fishes were more susceptible to hypoxia than freshwater fishes. This pattern is consistent with greater fluctuations in oxygen and temperature in freshwater habitats. Fishes with higher oxygen requirements (e.g. a high metabolic rate relative to body mass) also were more susceptible to hypoxia. We also found evidence that hypoxia and warming can act synergistically, as hypoxia tolerance was generally lower in warmer waters. However, we found significant interactions between temperature and the body and cell size of a fish. Constraints in oxygen uptake related to cellular surface area to volume ratios and effects of viscosity on the thickness of the boundary layers enveloping the gills could explain these thermal dependencies. The lower hypoxia tolerance in warmer waters was particularly pronounced for fishes with larger bodies and larger cell sizes. Previous studies have found a wide diversity in the direction and strength of relationships between P crit and body mass. By including interactions with temperature, our study may help resolve these divergent findings, explaining the size dependency of hypoxia tolerance in fish., (© 2022 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

It has been proposed that larger individuals within fish species may be more sensitive to global warming, as a result of limitations in their capacity to provide oxygen for aerobic metabolic activities. This could affect size distributions of populations in a warmer world but evidence is lacking. In Nile tilapia Oreochromis niloticus (n=18, mass range 21-313 g), capacity to provide oxygen for aerobic activities (aerobic scope) was independent of mass at an acclimation temperature of 26°C. Tolerance of acute warming, however, declined significantly with mass when evaluated as the critical temperature for fatigue from aerobic swimming (CTSmax). The CTSmax protocol challenges a fish to meet the oxygen demands of constant aerobic exercise while their demands for basal metabolism are accelerated by incremental warming, culminating in fatigue. CTSmax elicited pronounced increases in oxygen uptake in the tilapia but the maximum rates achieved prior to fatigue declined very significantly with mass. Mass-related variation in CTSmax and maximum oxygen uptake rates were positively correlated, which may indicate a causal relationship. When fish populations are faced with acute thermal stress, larger individuals may become constrained in their ability to perform aerobic activities at lower temperatures than smaller conspecifics. This could affect survival and fitness of larger fish in a future world with more frequent and extreme heatwaves, with consequences for population productivity., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2022. Published by The Company of Biologists Ltd.)

Atmospheric particulate matter (APM) emitted by iron ore processing industries has a complex composition, including diverse metallic particles and nanoparticles. Settleable APM (SePM) causes air to water cross-contamination and has recently been demonstrated to have harmful sublethal impacts on fish, eliciting stress responses, affecting the immune system, and reducing blood oxygen-carrying capacity. These findings imply potential consequences for fish aerobic performance and energy allocation, particularly in their ability to tolerate respiratory challenges such as aquatic hypoxia. To assess that potential limitation, we analyzed metabolic, cardiorespiratory, and morphological alterations after exposing tilapia, Oreochromis niloticus, to an environmentally relevant concentration of SePM (96 h) and progressive hypoxia. The contamination initiated detectable gill damage, reducing respiratory efficiency, increasing ventilatory effort, and compromising fish capacity to deal with hypoxia. Even in normoxia, the resting respiratory frequency was elevated and limited respiratory adjustments during hypoxia. SePM increased O 2 crit from 26 to 34% of O 2 (1.84 to 2.76 mg O 2 ·L -1 ). Such ventilatory inefficacy implies higher ventilatory cost with relevant alterations in energy allocation. Progression in gill damage might be problematic and cause: infection, blood loss, ion imbalance, and limited cardiorespiratory performance. The contamination did not cause immediate lethality but may threaten fish populations due to limitations in physiological performance. This was the first investigation to evaluate the physiological responses of fish to hypoxia after SePM contamination. We suggest that the present level of environmental SePM deserves attention. The present results demonstrate the need for comprehensive studies on SePM effects in aquatic fauna., (Copyright © 2022 Elsevier Inc. All rights reserved.)

Steel industry emissions of atmospheric particulate matter are responsible for air to water cross-contamination, which deposits metal/metalloid contaminants in aquatic ecosystems. This source of contamination has not been considered in most of the environmental monitoring protocols. Settleable atmospheric particulate matter (SePM) collected in an area of steel industry influence was used to analyze the sublethal effects on the hematological and innate immunological variables in Nile tilapia (Oreochromis niloticus) after short-term exposure (96 h). Blood samples were analyzed to evaluate the oxygen-carrying transport capacity, innate immune activity and stress biomarkers after exposure to ecologically relevant concentration of SePM. The exposure reduced blood oxygen-carrying capacity by lessening hematocrit, hemoglobin, erythrocyte, and mean corpuscular hemoglobin concentration. Compensatory increments in mean corpuscular volume and mean corpuscular hemoglobin have also been observed. The contaminant impacted the immune system by reducing the number of leukocytes, thrombocytes, and monocytes, total plasma protein, leukocyte respiratory activity, and by increasing lysozyme concentration. Furthermore, the contaminant caused endocrine stress response, raising plasma cortisol and glucose. Therefore, the alterations caused by SePM threatened the capacity of sustaining aerobic metabolism, impaired the immune system, and changed the energy allocation due to both stress response and immune effect. This may have important implications for the impact of SePM on aquatic ecosystems. Future investigations should assess SePM impact on general physiology and aerobic performance, especially to face common ecological challenges such as hypoxia and sustained swimming. These results point out the need to develop proper protocols to address the air-to-water cross-contamination risks by iron ore processing industries., (Copyright © 2022 Elsevier Inc. All rights reserved.)

Social rank in a structured society has been linked to basal levels of glucocorticoids in various species, with dominant individuals generally presenting lower levels than subordinates. The biotic and abiotic factors influencing glucocorticoids levels across social ranks are still, however, unclear in fishes. We investigated the influences of group size, fish size, sex, age, and reproduction type, plus water salinity and temperature, on the basal levels of cortisol, the major stress hormone in fishes. A phylogenetically controlled meta-analysis was performed on data from 72 studies over 22 species of fishes. As expected, dominants generally exhibited lower levels of cortisol than subordinates. More importantly, the strength of the correlation between cortisol and rank was modulated by three main factors, group size, environmental temperature, and fish size. Differences in basal cortisol between dominants and subordinates were significantly greater in small groups (dyadic contexts) when compared to larger groups. Differences between dominants and subordinates were also greater in temperate regions when compared to the tropics, and in species with larger body size. These results provide valuable insights into the links among hierarchy, stress and metabolism in fishes., (Copyright © 2021 Elsevier Inc. All rights reserved.)

We review the use of complex physiological traits, of tolerance and performance, as biomarkers of the toxicological effects of contaminants in subtropical and tropical freshwater fishes. Such traits are growing in relevance due to climate change, as exposure to contaminants may influence the capacity of fishes to tolerate and perform in an increasingly stressful environment. We review the evidence that the critical oxygen level, a measure of hypoxia tolerance, provides a valuable biomarker of impacts of diverse classes of contaminants. When coupled with measures of cardiorespiratory variables, it can provide insight into mechanisms of toxicity. The critical thermal maximum, a simple measure of tolerance of acute warming, also provides a valuable biomarker despite a lack of understanding of its mechanistic basis. Its relative ease of application renders it useful in the rapid evaluation of multiple species, and in understanding how the severity of contaminant impacts depends upon prevailing environmental temperature. The critical swimming speed is a measure of exercise performance that is widely used as a biomarker in temperate species but very few studies have been performed on subtropical or tropical fishes. Overall, the review serves to highlight a critical lack of knowledge for subtropical and tropical freshwater fishes. There is a real need to expand the knowledge base and to use physiological biomarkers in support of decision making to manage tropical freshwater fish populations and their habitats, which sustain rich biodiversity but are under relentless anthropogenic pressure., (© 2021 Wiley Periodicals LLC.)

How ectothermic animals will cope with global warming is a critical determinant of the ecological impacts of climate change. There has been extensive study of upper thermal tolerance limits among fish species but how intraspecific variation in tolerance may be affected by habitat characteristics and evolutionary history has not been considered. Intraspecific variation is a primary determinant of species vulnerability to climate change, with implications for global patterns of impacts of ongoing warming. Using published critical thermal maximum (CT max ) data on 203 fish species, we found that intraspecific variation in upper thermal tolerance varies according to a species' latitude and evolutionary history. Overall, tropical species show a lower intraspecific variation in thermal tolerance than temperate species. Notably, freshwater tropical species have a lower variation in tolerance than freshwater temperate species, which implies increased vulnerability to impacts of thermal stress. The extent of variation in CT max among fish species has a strong phylogenetic signal, which may indicate a constraint on evolvability to rising temperatures in tropical fishes. That is, in addition to living closer to their upper thermal limits, tropical species may have higher sensitivity and lower adaptability to global warming compared to temperate counterparts. This is evidence that freshwater tropical fish communities, worldwide, are especially vulnerable to ongoing climate change., (© 2021. The Author(s).)

Global warming is causing profound modifications of aquatic ecosystems and one major outcome appears to be a decline in adult size of many fish species. Over the last decade, sardine populations in the Gulf of Lions (NW Mediterranean Sea) have shown severe declines in body size and condition as well as disappearance of the oldest individuals, which could not be related to overfishing, predation pressure or epizootic diseases. In this study, we investigated whether this situation reflects a bottom-up phenomenon caused by reduced size and availability of prey that could lead to energetic constraints. We fed captive sardines with food items of two different sizes eliciting a change in feeding mode (filter-feeding on small items and directly capturing larger ones) at two different rations for several months, and then assessed their muscle bioenergetics to test for changes in cellular function. Feeding on smaller items was associated with a decline in body condition, even at high ration, and almost completely inhibited growth by comparison to sardines fed large items at high ration. Sardines fed on small items presented specific mitochondrial adjustments for energy sparing, indicating a major bioenergetic challenge. Moreover, mitochondria from sardines in poor condition had low basal oxidative activity but high efficiency of ATP production. Notably, when body condition was below a threshold value of 1.07, close to the mean observed in the wild, it was directly correlated with basal mitochondrial activity in muscle. The results show a link between whole-animal condition and cellular bioenergetics in the sardine, and reveal physiological consequences of a shift in feeding mode. They demonstrate that filter-feeding on small prey leads to poor growth, even under abundant food and an increase in the efficiency of ATP production. These findings may partially explain the declines in sardine size and condition observed in the wild., (© 2021 British Ecological Society.)

Interest in the measurement of metabolic rates is growing rapidly, because of the importance of metabolism in advancing our understanding of organismal physiology, behaviour, evolution and responses to environmental change. The study of metabolism in aquatic animals is undergoing an especially pronounced expansion, with more researchers utilising intermittent-flow respirometry as a research tool than ever before. Aquatic respirometry measures the rate of oxygen uptake as a proxy for metabolic rate, and the intermittent-flow technique has numerous strengths for use with aquatic animals, allowing metabolic rate to be repeatedly estimated on individual animals over several hours or days and during exposure to various conditions or stimuli. There are, however, no published guidelines for the reporting of methodological details when using this method. Here, we provide the first guidelines for reporting intermittent-flow respirometry methods, in the form of a checklist of criteria that we consider to be the minimum required for the interpretation, evaluation and replication of experiments using intermittent-flow respirometry. Furthermore, using a survey of the existing literature, we show that there has been incomplete and inconsistent reporting of methods for intermittent-flow respirometry over the past few decades. Use of the provided checklist of required criteria by researchers when publishing their work should increase consistency of the reporting of methods for studies that use intermittent-flow respirometry. With the steep increase in studies using intermittent-flow respirometry, now is the ideal time to standardise reporting of methods, so that - in the future - data can be properly assessed by other scientists and conservationists., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2021. Published by The Company of Biologists Ltd.)

Animal mortality is difficult to observe in marine systems, preventing a mechanistic understanding of major drivers of fish population dynamics. In particular, starvation is known to be a major cause of mortality at larval stages, but adult mortality is often unknown. In this study, we used a laboratory food-deprivation experiment, on wild caught sardine Sardina pilchardus from the Gulf of Lions. This population is interesting because mean individual phenotype shifted around 2008, becoming dominated by small, young individuals in poor body condition, a phenomenon that may result from declines in energy availability. Continuous monitoring of body mass loss and metabolic rate in 78 captive food-deprived individuals revealed that sardines could survive for up to 57 days on body reserves. Sardines submitted to long-term caloric restriction prior to food-deprivation displayed adaptive phenotypic plasticity, reducing metabolic energy expenditure and enduring starvation for longer than sardines that had not been calorie-restricted. Overall, entry into critical fasting phase 3 occurred at a body condition of 0.72. Such a degree of leanness has rarely been observed over 34 years of wild population monitoring. Still, the proportion of sardines below this threshold has doubled since 2008 and is maximal in January and February (the peak of the reproductive season), now reaching almost 10 % of the population at that time. These results indicate that the demographic changes observed in the wild may result in part from starvation-related adult mortality at the end of the winter reproductive period, despite adaptive plastic responses., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

Gilthead seabream were equipped with intraperitoneal biologging tags to investigate cardiac responses to hypoxia and warming, comparing when fish were either swimming freely in a tank with conspecifics or confined to individual respirometers. After tag implantation under anaesthesia, heart rate (fH) required 60 h to recover to a stable value in a holding tank. Subsequently, when undisturbed under control conditions (normoxia, 21°C), mean fH was always significantly lower in the tank than in the respirometers. In progressive hypoxia (100% to 15% oxygen saturation), mean fH in the tank was significantly lower than in the respirometers at oxygen levels down to 40%, with significant bradycardia in both holding conditions below this level. Simultaneous logging of tri-axial body acceleration revealed that spontaneous activity, inferred as the variance of external acceleration (VARm), was low and invariant in hypoxia. Warming (21 to 31°C) caused progressive tachycardia with no differences in fH between holding conditions. Mean VARm was, however, significantly higher in the tank during warming, with a positive relationship between VARm and fH across all temperatures. Therefore, spontaneous activity contributed to raising fH of fish in the tank during warming. Mean fH in respirometers had a highly significant linear relationship with mean rates of oxygen uptake, considering data from hypoxia and warming together. The high fH of confined seabream indicates that respirometry techniques may bias estimates of metabolic traits in some fishes, and that biologging on free-swimming fish will provide more reliable insight into cardiac and behavioural responses to environmental stressors by fish in their natural environment., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2021. Published by The Company of Biologists Ltd.)

Feed efficiency (FE) is the amount of body weight gain for a given feed intake. Improving FE through selective breeding is key for sustainable finfish aquaculture but its evaluation at individual level is technically challenging. We therefore investigated whether individual routine metabolic rate (RMR) was a predictor of individual FE in the European sea bass Dicentrarchus labrax, a major species in European mariculture. The European sea bass has three genetically distinct populations across its geographical range, namely Atlantic (AT), West Mediterranean (WM), and East Mediterranean (EM). We compared FE and RMR of fish from these three populations at 18 or 24 °C. We held 200 fish (62 AT, 66 WM, and 72 EM) in individual aquaria and fed them from ad libitum down to fasting. FI was assessed for an ad libitum feeding rate and for a fixed restricted ration (1% of metabolic body weight·day-1, with metabolic body weight = body weight0.8). After being refed 12 wk in a common tank, individual RMR was measured over 36 h by intermittent flow respirometry. There was a significant effect of temperature whereby fish at 18 °C had greater mean FE (P < 0.05) and lower RMR (P < 0.001). There was also a significant effect of population, where AT fish had lower FE (P < 0.05) and greater RMR (P < 0.001) than WM and EM, at both temperatures. Despite these differences in temperature and population means, individual FE and RMR were not significantly correlated (P > 0.05). Therefore, although the results provide evidence of an association between metabolic rate and FE, RMR was not a predictor of individual FE, for reasons that require further investigation., (© The Author(s) 2021. Published by Oxford University Press on behalf of the American Society of Animal Science. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Intraspecific variation in key traits such as tolerance of warming can have profound effects on ecological and evolutionary processes, notably responses to climate change. The empirical evidence for three primary elements of intraspecific variation in tolerance of warming in fishes is reviewed. The first is purely mechanistic that tolerance varies across life stages and as fishes become mature. The limited evidence indicates strongly that this is the case, possibly because of universal physiological principles. The second is intraspecific variation that is because of phenotypic plasticity, also a mechanistic phenomenon that buffers individuals' sensitivity to negative impacts of global warming in their lifetime, or to some extent through epigenetic effects over successive generations. Although the evidence for plasticity in tolerance to warming is extensive, more work is required to understand underlying mechanisms and to reveal whether there are general patterns. The third element is intraspecific variation based on heritable genetic differences in tolerance, which underlies local adaptation and may define long-term adaptability of a species in the face of ongoing global change. There is clear evidence of local adaptation and some evidence of heritability of tolerance to warming, but the knowledge base is limited with detailed information for only a few model or emblematic species. There is also strong evidence of structured variation in tolerance of warming within species, which may have ecological and evolutionary significance irrespective of whether it reflects plasticity or adaptation. Although the overwhelming consensus is that having broader intraspecific variation in tolerance should reduce species vulnerability to impacts of global warming, there are no sufficient data on fishes to provide insights into particular mechanisms by which this may occur., (© 2020 Fisheries Society of the British Isles.)

Oxygen uptake, heart rate and contraction frequencies of slow oxidative (SO) and fast glycolytic (FG) muscle were measured simultaneously in gilthead seabream Sparus aurata submitted to stepwise increases in current speed in a swimming respirometer. Variation in oxygen uptake was closely related to variation in heart rate, over initial steps these rose in concert with an increase in contraction frequency of SO muscle. There was an asymptote in oxygen uptake and heart rate at high speeds that reflected a transition from exclusive use of aerobic SO muscle to a combination of SO and anaerobic FG muscle, and which preceded fatigue., (© 2020 The Fisheries Society of the British Isles.)

Warming of aquatic environments as a result of climate change is already having measurable impacts on fishes, manifested as changes in phenology, range shifts and reductions in body size. Understanding the physiological mechanisms underlying these seemingly universal patterns is crucial if we are to reliably predict the fate of fish populations with future warming. This includes an understanding of mechanisms for acute thermal tolerance, as extreme heatwaves may be a major driver of observed effects. The hypothesis of gill oxygen limitation (GOL) is claimed to explain asymptotic fish growth, and why some fish species are decreasing in size with warming; but its underlying assumptions conflict with established knowledge and direct mechanistic evidence is lacking. The hypothesis of oxygen- and capacity-limited thermal tolerance (OCLTT) has stimulated a wave of research into the role of oxygen supply capacity and thermal performance curves for aerobic scope, but results vary greatly between species, indicating that it is unlikely to be a universal mechanism. As thermal performance curves remain important for incorporating physiological tolerance into models, we discuss potentially fruitful alternatives to aerobic scope, notably specific dynamic action and growth rate. We consider the limitations of estimating acute thermal tolerance by a single rapid measure whose mechanism of action is not known. We emphasise the continued importance of experimental physiology, particularly in advancing our understanding of underlying mechanisms, but also the challenge of making this knowledge relevant to the more complex reality., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2021. Published by The Company of Biologists Ltd.)

In some fishes, the ability to breathe air has evolved to overcome constraints in hypoxic environments but comes at a cost of increased predation. To reduce this risk, some species perform group air breathing. Temperature may also affect the frequency of air breathing in fishes, but this topic has received relatively little research attention. This study examined how acclimation temperature and acute exposure to hypoxia affected the air-breathing behaviour of a social catfish, the bronze corydoras Corydoras aeneus , and aimed to determine whether individual oxygen demand influenced the behaviour of entire groups. Groups of seven fish were observed in an arena to measure air-breathing frequency of individuals and consequent group air-breathing behaviour, under three oxygen concentrations (100%, 60% and 20% air saturation) and two acclimation temperatures (25 and 30°C). Intermittent flow respirometry was used to estimate oxygen demand of individuals. Increasingly severe hypoxia increased air breathing at the individual and group levels. Although there were minimal differences in air-breathing frequency among individuals in response to an increase in temperature, the effect of temperature that did exist manifested as an increase in group air-breathing frequency at 30°C. Groups that were more socially cohesive during routine activity took more breaths but, in most cases, air breathing among individuals was not temporally clustered. There was no association between an individual's oxygen demand and its air-breathing frequency in a group. For C . aeneus , although air-breathing frequency is influenced by hypoxia, behavioural variation among groups could explain the small overall effect of temperature on group air-breathing frequency., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2020. Published by The Company of Biologists Ltd.)

We investigated whether fatigue from sustained aerobic swimming provides a sub-lethal endpoint to define tolerance of acute warming in fishes, as an alternative to loss of equilibrium (LOE) during a critical thermal maximum (CT max ) protocol. Two species were studied, Nile tilapia ( Oreochromis niloticus ) and pacu ( Piaractus mesopotamicus ). Each fish underwent an incremental swim test to determine gait transition speed ( U GT ), where it first engaged the unsteady anaerobic swimming mode that preceded fatigue. After suitable recovery, each fish was exercised at 85% of their own U GT and warmed 1°C every 30 min, to identify the temperature at which they fatigued, denoted as CT swim Fish were also submitted to a standard CT max , warming at the same rate as CT swim , under static conditions until LOE. All individuals fatigued in CT swim , at a mean temperature approximately 2°C lower than their CT max Therefore, if exposed to acute warming in the wild, the ability to perform aerobic metabolic work would be constrained at temperatures significantly below those that directly threatened survival. The collapse in performance at CT swim was preceded by a gait transition qualitatively indistinguishable from that during the incremental swim test. This suggests that fatigue in CT swim was linked to an inability to meet the tissue oxygen demands of exercise plus warming. This is consistent with the oxygen and capacity limited thermal tolerance (OCLTT) hypothesis, regarding the mechanism underlying tolerance of warming in fishes. Overall, fatigue at CT swim provides an ecologically relevant sub-lethal threshold that is more sensitive to extreme events than LOE at CT max ., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2020. Published by The Company of Biologists Ltd.)

Hypoxia and mercury contamination often co-occur in tropical freshwater ecosystems, but the interactive effects of these two stressors on fish populations are poorly known. The effects of mercury (Hg) on recorded changes in the detailed form of the electrocardiogram (ECG) during exposure to progressive hypoxia were investigated in two Neotropical freshwater fish species, matrinxã, Brycon amazonicus and traíra, Hoplias malabaricus. Matrinxã were exposed to a sublethal concentration of 0.1 mg L -1 of HgCl 2 in water for 96 h. Traíra were exposed to dietary doses of Hg by being fed over a period of 30 days with juvenile matrinxãs previously exposed to HgCl 2 , resulting in a dose of 0.45 mg of total Hg per fish, each 96 h. Both species showed a bradycardia in progressive hypoxia. Hg exposure impaired cardiac electrical excitability, leading to first-degree atrioventricular block, plus profound extension of the ventricular action potential (AP) plateau. Moreover, there was the development of cardiac arrhythmias and anomalies such as occasional absence of QRS complexes, extra systoles, negative Q-, R- and S-waves (QRS complex), and T wave inversion, especially in hypoxia below O 2 partial pressures (PO 2 ) of 5.3 kPa. Sub-chronic dietary Hg exposure induced intense bradycardia in normoxia in traira, plus lengthening of ventricular AP duration coupled with prolonged QRS intervals. This indicates slower ventricular AP conduction during ventricular depolarization. Overall, the data indicate that both acute waterborne and sub-chronic dietary exposure (trophic level transfer), at sublethal concentrations of mercury, cause damage in electrical stability and rhythm of the heartbeat, leading to myocardial dysfunction, which is further intensified during hypoxia. These changes could lead to impaired cardiac output, with consequences for swimming ability, foraging capacity, and hence growth and/or reproductive performance.

Air-breathing and amphibious fishes are essential study organisms to shed insight into the required physiological shifts that supported the full transition from aquatic water-breathing fishes to terrestrial air-breathing tetrapods. While the origin of air-breathing in the evolutionary history of the tetrapods has received considerable focus, much less is known about the evolutionary physiology of air-breathing among fishes. This review summarizes recent advances within the field with specific emphasis on the cardiorespiratory regulation associated with air-breathing and terrestrial excursions, and how respiratory physiology of these living transitional forms are affected by development and personality. Finally, we provide a detailed and re-evaluated model of the evolution of air-breathing among fishes that serves as a framework for addressing new questions on the cardiorespiratory changes associated with it. This review highlights the importance of combining detailed studies on piscine air-breathing model species with comparative multi-species studies, to add an additional dimension to our understanding of the evolutionary physiology of air-breathing in vertebrates., (© 2019 Scandinavian Physiological Society. Published by John Wiley & Sons Ltd.)

The European City Fish project aimed to develop a generic methodology for ecological risk assessment for urban rivers. Since traditional methods only consider a small fraction of substances present in the water cycle, biological effect monitoring is required for a more reliable assessment of the pollution status. A major challenge for environmental risk assessment (ERA) is the application of adverse outcome pathways (AOP), i.e. the linking of pollutant exposure via early molecular and biochemical changes to physiological effects and, ultimately, effects on populations and ecosystems. We investigated the linkage between responses at these different levels. Many AOP aspects were investigated, from external and internal exposure to different classes of micropollutants, via molecular key events (MKE) the impacts on organs and organisms (fish physiology), to changes in the population dynamics of fish. Risk assessment procedures were evaluated by comparing environmental quality standards, bioassay responses, biomarkers in caged and feral fish, and the impact on fish populations. Although no complete AOP was observed, indirect relationships linking pollutant exposure via MKE to impaired locomotion were demonstrated at the most polluted site near a landfill for chemical waste. The pathway indicated that several upstream key events requiring energy for stress responses and toxic defence are likely to converge at a single common MKE: increased metabolic demands. Both fish biomarkers and the bioanalytical SIMONI strategy are valuable indicators for micropollutant risks to fish communities., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2019 Elsevier B.V. All rights reserved.)

We investigated links between swimming behavior and muscle bioenergetics in two emblematic Mediterranean fish species that have very different ecologies and activity levels. European sardines Sardina pilchardus are pelagic, they swim aerobically, school constantly and have high muscle fat content. Gilthead seabream Sparus aurata are bentho-pelagic, they show discontinuous spontaneous swimming patterns and store less fat in their muscle. Estimating the proportion of red and white muscle phenotypes, sardine exhibited a larger proportion of red muscle (~10% of the body mass) compared to gilthead seabream (~5% of the body mass). We firstly studied red and white muscle fiber bioenergetics, using high-resolution respirometers, showing a 4-fold higher oxidation capacity for red compared to white muscle. Secondly, we aimed to compare the red muscle ability to oxidize either lipids or carbohydrates. Sardine red muscle had a 3-fold higher oxidative capacity than gilthead seabream and a greater capacity to oxidize lipids. This study provides novel insights into physiological mechanisms underlying the different lifestyles of these highly-prized species., (Copyright © 2019 Elsevier Inc. All rights reserved.)

This study investigated the effect of microcystin-LR (MC-LR) on in vivo cardiorespiratory function and on tissue biomarkers of oxidative stress in gills and liver of the trahira, a neotropical freshwater fish. Trahira were treated with an intraperitoneal injection of 100 µg MC-LR.kg -1 body mass or a saline, with the toxic effects of MC-LR then evaluated after 48 h. Rates of oxygen uptake (V̇O 2 ) did not differ significantly between Control and the exposed group (Mcys), but exposure to MC-LR significantly reduced O 2 extraction in the Mcys group at all O 2 tensions. This was associated with higher gill ventilation volume (V̇ G ) in the Mcys group at all O 2 tensions except 140 and 120 mmHg, and a higher tidal volume (V T ) of the Mcys group at all tensions except 140 mmHg. Heart rate was also higher in the Mcys group, significantly so at an O 2 tension of 40 mmHg. In the liver of trahira, exposure to MC-LR has significant effects on antioxidant defense systems, inducing a significant increase in the activity of the (GPx) glutathione peroxidase enzyme (100%) and in the reduced glutathione (GSH) content (70%) compared to the control group, but no effects on superoxide dismutase (SOD), catalase (CAT) and glutathione S-transferase (GST) enzymes. The liver showed no oxidative damage, when measured as lipid peroxidation (LPO) levels and protein carbonyl (PC) content. In the gills SOD and GPx enzyme activity increased significantly in the Mcys group (98% and 73% respectively) compared to the controls, although GSH, CAT and GST did not differ between groups. There was also no significant difference in GSH in this tissue. Levels of lipid peroxidation in the gills were 53% higher in the Mcys group, although carbonyl protein levels did not differ. In conclusion, these data show that MC-LR leads to development of hyperventilation and increased activity of the detoxification system and that this species was able to compensate the deleterious effects of microcystin on its vital functions. The antioxidant defense in the liver was able to contain the propagation of LPO and prevent the oxidation of proteins, although the gills of the fishes exposed to MC-LR were not able to contain the formation of reactive oxygen species and LPO, which led to the establishment of oxidative stress which impaired gill function., (Copyright © 2019 Elsevier Inc. All rights reserved.)

This review considers the environmental and systemic factors that can stimulate air-breathing responses in fishes with bimodal respiration, and how these may be controlled by peripheral and central chemoreceptors. The systemic factors that stimulate air-breathing in fishes are usually related to conditions that increase the O 2 demand of these animals (e.g. physical exercise, digestion and increased temperature), while the environmental factors are usually related to conditions that impair their capacity to meet this demand (e.g. aquatic/aerial hypoxia, aquatic/aerial hypercarbia, reduced aquatic hidrogenionic potential and environmental pollution). It is now well-established that peripheral chemoreceptors, innervated by cranial nerves, drive increased air-breathing in response to environmental hypoxia and/or hypercarbia. These receptors are, in general, sensitive to O 2 and/or CO 2 /H + levels in the blood and/or the environment. Increased air-breathing in response to elevated O 2 demand may also be driven by the peripheral chemoreceptors that monitor O 2 levels in the blood. Very little is known about central chemoreception in air-breathing fishes, the data suggest that central chemosensitivity to CO 2 /H + is more prominent in sarcopterygians than in actinopterygians. A great deal remains to be understood about control of air-breathing in fishes, in particular to what extent control systems may show commonalities (or not) among species or groups that have evolved air-breathing independently, and how information from the multiple peripheral (and possibly central) chemoreceptors is integrated to control the balance of aerial and aquatic respiration in these animals., (Copyright © 2018 Elsevier GmbH. All rights reserved.)

Group-living is widespread among animals and comes with numerous costs and benefits. To date, research examining group-living has focused on trade-offs surrounding foraging, while other forms of resource acquisition have been largely overlooked. Air-breathing has evolved in many fish lineages, allowing animals to obtain oxygen in hypoxic aquatic environments. Breathing air increases the threat of predation, so some species perform group air-breathing, to reduce individual risk. Within species, individual air-breathing can be influenced by metabolic rate as well as personality, but the mechanisms of group air-breathing remain unexplored. It is conceivable that keystone individuals with high metabolic demand or intrinsic tendency to breathe air may drive social breathing, especially in hypoxia. We examined social air-breathing in African sharptooth catfish Clarias gariepinus, to determine whether individual physiological traits and spontaneous tendency to breathe air influence the behaviour of entire groups, and whether such influences vary in relation to aquatic oxygen availability. We studied 11 groups of four catfish in a laboratory arena and recorded air-breathing behaviour, activity and agonistic interactions at varying levels of hypoxia. Bimodal respirometry was used to estimate individual standard metabolic rate (SMR) and the tendency to utilize aerial oxygen when alone. Fish took more air breaths in groups as compared to when they were alone, regardless of water oxygen content, and displayed temporally clustered air-breathing behaviour, consistent with existing definitions of synchronous air-breathing. However, groups displayed tremendous variability in surfacing behaviour. Aggression by dominant individuals within groups was the main factor influencing air-breathing of the entire group. There was no association between individual SMR, or the tendency to obtain oxygen from air when in isolation, and group air-breathing. For C. gariepinus, synchronous air-breathing is strongly influenced by agonistic interactions, which may expose subordinate individuals to risk of predation. Influential individuals exerted an overriding effect on risk-taking by the entire group, for reasons independent of their physiological oxygen requirements. Overall, this illustrates that social context can obscure interactions between an individual's physiological and behavioural traits and their tendency to take risks to obtain resources., (© 2017 The Authors. Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.)

Some recent modelling papers projecting smaller fish sizes and catches in a warmer future are based on erroneous assumptions regarding (i) the scaling of gills with body mass and (ii) the energetic cost of 'maintenance'. Assumption (i) posits that insurmountable geometric constraints prevent respiratory surface areas from growing as fast as body volume. It is argued that these constraints explain allometric scaling of energy metabolism, whereby larger fishes have relatively lower mass-specific metabolic rates. Assumption (ii) concludes that when fishes reach a certain size, basal oxygen demands will not be met, because of assumption (i). We here demonstrate unequivocally, by applying accepted physiological principles with reference to the existing literature, that these assumptions are not valid. Gills are folded surfaces, where the scaling of surface area to volume is not constrained by spherical geometry. The gill surface area can, in fact, increase linearly in proportion to gill volume and body mass. We cite the large body of evidence demonstrating that respiratory surface areas in fishes reflect metabolic needs, not vice versa, which explains the large interspecific variation in scaling of gill surface areas. Finally, we point out that future studies basing their predictions on models should incorporate factors for scaling of metabolic rate and for temperature effects on metabolism, which agree with measured values, and should account for interspecific variation in scaling and temperature effects. It is possible that some fishes will become smaller in the future, but to make reliable predictions the underlying mechanisms need to be identified and sought elsewhere than in geometric constraints on gill surface area. Furthermore, to ensure that useful information is conveyed to the public and policymakers about the possible effects of climate change, it is necessary to improve communication and congruity between fish physiologists and fisheries scientists., (© 2017 John Wiley & Sons Ltd.)

Individual variation in sub-lethal sensitivity to the organophosphate pesticide trichlorfon was investigated in Nile tilapia, using critical swimming speed (U crit ) as an indicator. Tilapia exposed for 96h to 500μgl -1 trichlorfon at 26°C (Tcfn group, n=27) showed a significant decline in mean U crit , compared to their own control (pre-exposure) performance in clean water (-14.5±2.3%, mean±SEM), but also compared to a Sham group (n=10) maintained for 96h in clean water. Individuals varied in their relative sensitivity to the pesticide, with the decline in U crit after exposure varying from 1 to 41%. The U crit of the Tcfn group did not recover completely after 96h in clean water, remaining 9.4±3.2% below their own control performance. The decline in performance was associated with a significant increase in net cost of aerobic swimming, of +28.4±6.5% at a sustained speed of 2bodylengthss -1 , which translated into a significant decline in swimming efficiency (E swim ) of -17.6±4.0% at that speed. Within the Tcfn group, individual E swim was a strong positive determinant of individual U crit across all trials, and a strong negative determinant of individual% decline in U crit after pesticide exposure (P<0.001, linear mixed effect models). Trichlorfon had no effects on standard metabolic rate or active metabolic rate (AMR) but, nonetheless, individual U crit in all trials, and% decline in U crit after exposure, were strongly associated with individual AMR (positively and negatively, respectively, P<0.001). Individual U crit under control conditions was also a strong positive determinant of U crit after trichlorfon exposure (P<0.001), but not of the% decline in U crit performance. In conclusion, the OP pesticide impaired U crit performance by reducing E swim but individual tilapia varied widely in their relative sensitivity. Intrinsic individual physiology determined effects of the pesticide on performance and, in particular, good swimmers remained better swimmers after exposure., (Copyright © 2017 The Author(s). Published by Elsevier B.V. All rights reserved.)

Clarias gariepinus is a facultative air-breathing catfish that exhibits changes in heart rate (ƒ H ) associated with air-breaths (AB). A transient bradycardia prior to the AB is followed by sustained tachycardia during breath-hold. This study evaluated air-breathing and cardiac responses to sustained aerobic exercise in juveniles (total length~20cm), and how exercise influenced variations in f H associated with AB. In particular, it investigated the role of adrenergic and cholinergic control in cardiac responses, and effects of pharmacological abolition of this control on air-breathing responses. Sustained exercise at 15, 30 and 45cms -1 in a swim tunnel caused significant increases in f AB and f H , from approximately 5breathsh -1 and 60heartbeatsmin -1 at the lowest speed, to over 60breathsh -1 and 100beatsmin -1 at the highest, respectively. There was a progressive decline in the degree of variation in f H , around each AB, as f AB increased with exercise intensity. Total autonomic blockade abolished all variation in f H during exercise, and around each AB, but f AB responses were the same as in untreated animals. Cardiac responses were exclusively due to modulation of inhibitory cholinergic tone, which varied from >100% at the lowest speed to <10% at the highest. Cholinergic blockade had no effect on f AB compared to untreated fish. Excitatory β-adrenergic tone was approximately 20% and did not vary with swimming speed, but its blockade increased f AB at all speeds, compared to untreated animals. This reveals complex effects of autonomic control on air-breathing during exercise in C. gariepinus, which deserve further investigation., (Copyright © 2016 Elsevier Inc. All rights reserved.)

The state of the art of research on the environmental physiology of marine fishes is reviewed from the perspective of how it can contribute to conservation of biodiversity and fishery resources. A major constraint to application of physiological knowledge for conservation of marine fishes is the limited knowledge base; international collaboration is needed to study the environmental physiology of a wider range of species. Multifactorial field and laboratory studies on biomarkers hold promise to relate ecophysiology directly to habitat quality and population status. The 'Fry paradigm' could have broad applications for conservation physiology research if it provides a universal mechanism to link physiological function with ecological performance and population dynamics of fishes, through effects of abiotic conditions on aerobic metabolic scope. The available data indicate, however, that the paradigm is not universal, so further research is required on a wide diversity of species. Fish physiologists should interact closely with researchers developing ecological models, in order to investigate how integrating physiological information improves confidence in projecting effects of global change; for example, with mechanistic models that define habitat suitability based upon potential for aerobic scope or outputs of a dynamic energy budget. One major challenge to upscaling from physiology of individuals to the level of species and communities is incorporating intraspecific variation, which could be a crucial component of species' resilience to global change. Understanding what fishes do in the wild is also a challenge, but techniques of biotelemetry and biologging are providing novel information towards effective conservation. Overall, fish physiologists must strive to render research outputs more applicable to management and decision-making. There are various potential avenues for information flow, in the shorter term directly through biomarker studies and in the longer term by collaborating with modellers and fishery biologists.

Glyphosate-based herbicides are widely used in agriculture and are commonly found in water bodies. Roundup Original(®) (RO) contains an isopropylamine glyphosate (GLY) salt containing the surfactant POEA, while Roundup Transorb R(®) (RTR) contains a potassium salt of GLY with unknown surfactants. Both contain different compositions of so-called "inert" ingredients, more toxic than glyphosate. Amphibian tadpoles often experience variations in O2 availability in their aquatic habitats; an ability to tolerate hypoxia can condition their survival and fitness. We evaluated the impacts of sublethal concentrations of GLY (1 mg L(-1)), RO (1 mg L(-1) GLY a.e) and RTR (1 mg L(-1) GLY a.e) on metabolic rate (V·O2 - mLO2 Kg1 h(-1)) of bullfrog tadpoles during normoxia and graded hypoxia, and related this to morphology of their skin, their major site of gas exchange. In control (CT) V·O2 remained unaltered from normoxia until 40 mmHg, indicating a critical O2 tension between 40 and 20 mmHg. GLY significantly reduced V·O2, possibly due to epidermal hypertrophy, which increased O2 diffusion distance to O2 uptake. In contrast, RTR increased V·O2 during hypoxia, indicating an influence of "inert" compounds and surfactants. V·O2 of RO did not differ from CT, suggesting that any increase in V·O2 caused by exposure was antagonized by epidermal hypertrophy. Indeed, all herbicides caused marked alterations in skin morphology, with cell and epithelium wall presenting hyperplasia or hypertrophy and chromatid rupture. In summary, GLY, RO and RTR exert different effects in bullfrog tadpoles, in particular the surfactants and inert compounds appear to influence oxygen uptake., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

Hypoxia is a common occurrence in aquatic habitats, and it is becoming an increasingly frequent and widespread environmental perturbation, primarily as the result of anthropogenic nutrient enrichment and climate change. An in-depth understanding of the hypoxia tolerance of fishes, and how this varies among individuals and species, is required to make accurate predictions of future ecological impacts and to provide better information for conservation and fisheries management. The critical oxygen level (P crit) has been widely used as a quantifiable trait of hypoxia tolerance. It is defined as the oxygen level below which the animal can no longer maintain a stable rate of oxygen uptake (oxyregulate) and uptake becomes dependent on ambient oxygen availability (the animal transitions to oxyconforming). A comprehensive database of P crit values, comprising 331 measurements from 96 published studies, covering 151 fish species from 58 families, provides the most extensive and up-to-date analysis of hypoxia tolerance in teleosts. Methodologies for determining P crit are critically examined to evaluate its usefulness as an indicator of hypoxia tolerance in fishes. Various abiotic and biotic factors that interact with hypoxia are analysed for their effect on P crit, including temperature, CO2, acidification, toxic metals and feeding. Salinity, temperature, body mass and routine metabolic rate were strongly correlated with P crit; 20% of variation in the P crit data set was explained by these four variables. An important methodological issue not previously considered is the inconsistent increase in partial pressure of CO2 within a closed respirometer during the measurement of P crit. Modelling suggests that the final partial pressure of CO2 reached can vary from 650 to 3500 µatm depending on the ambient pH and salinity, with potentially major effects on blood acid-base balance and P crit itself. This database will form part of a widely accessible repository of physiological trait data that will serve as a resource to facilitate future studies of fish ecology, conservation and management.

Ecosystem-based approaches to fisheries management (EAFMs) have emerged as requisite for sustainable use of fisheries resources. At the same time, however, there is a growing recognition of the degree of variation among individuals within a population, as well as the ecological consequences of this variation. Managing resources at an ecosystem level calls on practitioners to consider evolutionary processes, and ample evidence from the realm of fisheries science indicates that anthropogenic disturbance can drive changes in predominant character traits (e.g. size at maturity). Eco-evolutionary theory suggests that human-induced trait change and the modification of selective regimens might contribute to ecosystem dynamics at a similar magnitude to species extirpation, extinction and ecological dysfunction. Given the dynamic interaction between fisheries and target species via harvest and subsequent ecosystem consequences, we argue that individual diversity in genetic, physiological and behavioural traits are important considerations under EAFMs. Here, we examine the role of individual variation in a number of contexts relevant to fisheries management, including the potential ecological effects of rapid trait change. Using select examples, we highlight the extent of phenotypic diversity of individuals, as well as the ecological constraints on such diversity. We conclude that individual phenotypic diversity is a complex phenomenon that needs to be considered in EAFMs, with the ultimate realization that maintaining or increasing individual trait diversity may afford not only species, but also entire ecosystems, with enhanced resilience to environmental perturbations. Put simply, individuals are the foundation from which population- and ecosystem-level traits emerge and are therefore of central importance for the ecosystem-based approaches to fisheries management.