Solving the conundrum of intra-specific variation in metabolic rate: A multidisciplinary conceptual and methodological toolkit: New technical developments are opening the door to an understanding of why metabolic rate varies among individual animals of a species: New technical developments are opening the door to an understanding of why metabolic rate varies among individual animals of a species.

Researchers from diverse disciplines, including organismal and cellular physiology, sports science, human nutrition, evolution and ecology, have sought to understand the causes and consequences of the surprising variation in metabolic rate found among and within individual animals of the same species. Research in this area has been hampered by differences in approach, terminology and methodology, and the context in which measurements are made. Recent advances provide important opportunities to identify and address the key questions in the field. By bringing together researchers from different areas of biology and biomedicine, we describe and evaluate these developments and the insights they could yield, highlighting the need for more standardisation across disciplines. We conclude with a list of important questions that can now be addressed by developing a common conceptual and methodological toolkit for studies on metabolic variation in animals.

The interplay between sleep and ecophysiology, behaviour and responses to environmental change in fish.

Evidence of behavioural sleep has been observed in every animal species studied to date, but current knowledge of the behaviour, neurophysiology and ecophysiology associated with sleep is concentrated on mammals and birds. Fish are a hugely diverse group that can offer novel insights into a variety of sleep-related behaviours across environments, but the ecophysiological relevance of sleep in fish has been largely overlooked. Here, we systematically reviewed the literature to assess the current breadth of knowledge on fish sleep, and surveyed the diverse physiological effects and behaviours associated with sleep. We also discuss possible ways in which unstudied external factors may alter sleep behaviours. For example, predation risk may alter sleep patterns, as has been shown in mammalian, avian and reptilian species. Other environmental factors - such as water temperature and oxygen availability - have the potential to alter sleep patterns in fish differently than for terrestrial endotherms. Understanding the ecological influences on sleep in fish is vital, as sleep deprivation has the potential to affect waking behaviour and fitness owing to cognitive and physiological impairments, possibly affecting ecological phenomena and sensitivity to environmental stressors in ways that have not been considered.

Differential metabolic responses in bold and shy sea anemones during a simulated heatwave.

As climate change-induced heatwaves become more common, phenotypic plasticity at multiple levels is a key mitigation strategy by which organisms can optimise selective outcomes. In ectotherms, changes to both metabolism and behaviour can help alleviate thermal stress. Nonetheless, no study in any ectotherm has yet empirically investigated how changing temperatures affect among-individual differences in the associations between these traits. Using the beadlet anemone (Actinia equina), an intertidal species from a thermally heterogeneous environment, we investigated how individual metabolic rates, linked to morphotypic differences in A. equina, and boldness were related across changing temperatures. A crossed-over design and a temporal control were used to test the same individuals at a non-stressful temperature, 13°C, and under a simulated heatwave at 21°C. At each temperature, short-term repeated measurements of routine metabolic rate (RMR) and a single measurement of a repeatable boldness-related behaviour, immersion response time (IRT), were made. Individual differences, but not morphotypic differences, were highly predictive of metabolic plasticity, and the plasticity of RMR was associated with IRT. At 13°C, shy animals had the highest metabolic rates, while at 21°C, this relationship was reversed. Individuals that were bold at 13°C also exhibited the highest metabolic rates at 21°C. Additional metabolic challenges during heatwaves could be detrimental to fitness in bold individuals. Equally, lower metabolic rates at non-stressful temperatures could be necessary for optimal survival as heatwaves become more common. These results provide novel insight into the relationship between metabolic and behavioural plasticity, and its adaptive implications in a changing climate.

Genomic basis of fishing-associated selection varies with population density.

Fisheries induce one of the strongest anthropogenic selective pressures on natural populations, but the genetic effects of fishing remain unclear. Crucially, we lack knowledge of how capture-associated selection and its interaction with reductions in population density caused by fishing can potentially shift which genes are under selection. Using experimental fish reared at two densities and repeatedly harvested by simulated trawling, we show consistent phenotypic selection on growth, metabolism, and social behavior regardless of density. However, the specific genes under selection-mainly related to brain function and neurogenesis-varied with the population density. This interaction between direct fishing selection and density could fundamentally alter the genomic responses to harvest. The evolutionary consequences of fishing are therefore likely context dependent, possibly varying as exploited populations decline. These results highlight the need to consider environmental factors when predicting effects of human-induced selection and evolution.

Colonization, diversity, and seasonality of fishes at pelagic fish aggregating devices.

The pelagic zone of the ocean can be a challenging environment in which to conduct research and as a result we lack the robust baseline abundance and diversity data, compared to what is available in more accessible coastal habitats, to be able to track changes or stressors to the biota in this environment. Many large-scale fisheries target pelagic fish, and much of the information available on these species is based on fisheries-dependent data that may be biased towards hotspots and commercially valuable fishes. Here, a long-term video and visual fish survey was conducted on two subsurface moored fish aggregating devices (FADs) in the pelagic waters of the central Bahamas to determine the feasibility of using moored pelagic FADs as tools for collecting fish abundance and diversity data. A wide range of species was documented, including large migratory fish that are the focus of commercial and recreational fisheries, and smaller often overlooked species on which little abundance or seasonality information exists. We found that FADs colonize quickly and reach a peak stable (albeit seasonally cyclical) abundance and diversity within the first several months after deployment. Species richness was higher in video surveys, but abundance was higher in visual surveys, except for sharks. Our results highlight the need to tailor survey methods to fit the context and study objective, and provide further evidence for the importance of fisheries-independent data in monitoring pelagic species.

Temperature change effects on marine fish range shifts: A meta-analysis of ecological and methodological predictors.

The current effects of global warming on marine ecosystems are predicted to increase, with species responding by changing their spatial distributions. Marine ectotherms such as fish experience elevated distribution shifts, as temperature plays a key role in physiological functions and delineating population ranges through thermal constraints. Distributional response predictions necessary for population management have been complicated by high heterogeneity in magnitude and direction of movements, which may be explained by both biological as well as methodological study differences. To date, however, there has been no comprehensive synthesis of the interacting ecological factors influencing fish distributions in response to climate change and the confounding methodological factors that can affect their estimation. In this study we analyzed published studies meeting criteria of reporting range shift responses to global warming in 115 taxa spanning all major oceanic regions, totaling 595 three-dimensional population responses (latitudinal, longitudinal, and depth), with temperature identified as a significant driver. We found that latitudinal shifts were the fastest in non-exploited, tropical populations, and inversely correlated with depth shifts which, in turn, dominated at the trailing edges of population ranges. While poleward responses increased with rate of temperature change and latitude, niche was a key factor in predicting both depth (18% of variation) and latitudinal responses (13%), with methodological predictors explaining between 10% and 28% of the observed variance in marine fish responses to temperature change. Finally, we found strong geographical publication bias and limited taxonomical scope, highlighting the need for more representative and standardized research in order to address heterogeneity in distribution responses and improve predictions in face of changing climate.

A warmer environment can reduce sociability in an ectotherm.

The costs and benefits of being social vary with environmental conditions, so individuals must weigh the balance between these trade-offs in response to changes in the environment. Temperature is a salient environmental factor that may play a key role in altering the costs and benefits of sociality through its effects on food availability, predator abundance, and other ecological parameters. In ectotherms, changes in temperature also have direct effects on physiological traits linked to social behaviour, such as metabolic rate and locomotor performance. In light of climate change, it is therefore important to understand the potential effects of temperature on sociality. Here, we took the advantage of a 'natural experiment' of threespine sticklebacks from contrasting thermal environments in Iceland: geothermally warmed water bodies (warm habitats) and adjacent ambient-temperature water bodies (cold habitats) that were either linked (sympatric) or physically distinct (allopatric). We first measured the sociability of wild-caught adult fish from warm and cold habitats after acclimation to a low and a high temperature. At both acclimation temperatures, fish from the allopatric warm habitat were less social than those from the allopatric cold habitat, whereas fish from sympatric warm and cold habitats showed no differences in sociability. To determine whether differences in sociability between thermal habitats in the allopatric population were heritable, we used a common garden breeding design where individuals from the warm and the cold habitat were reared at a low or high temperature for two generations. We found that sociability was indeed heritable but also influenced by rearing temperature, suggesting that thermal conditions during early life can play an important role in influencing social behaviour in adulthood. By providing the first evidence for a causal effect of rearing temperature on social behaviour, our study provides novel insights into how a warming world may influence sociality in animal populations.

Maze design: size and number of choices impact fish performance in cognitive assays.

Although studies on fish cognition are increasing, consideration of how methodological details influence the ability to detect and measure performance is lagging. Here, in two separate experiments the authors compared latency to leave the start position, latency to make a decision, levels of participation and success rates (whether fish entered the rewarded chamber as first choice) across different physical designs. Experiments compared fish performance across (a) two sizes of T-mazes, large and standard, and a plus-maze, and (b) open choice arenas with either two or four doors. Fish in T-mazes with longer arms took longer to leave the start chamber and were less likely to participate in a trial than fish in T-mazes with shorter arms. The number of options, or complexity, in a maze significantly impacted success but did not necessarily impact behavioural measures, and did not impact the number of fish that reached a chamber. Fish in the plus-maze had similar latencies to leave the start box and time to reach any chamber as fish in the same-sized T-maze but exhibited lower overall success. Similarly, in an open choice arena, increasing the number of options - doors to potential reward chambers - resulted in lower probability of success. There was an influence of reward position in the choice arena, with rewarded chambers closest to the sides of the arena resulting in lower latencies to enter and higher probability of decision success. Together the results allow the authors to offer practical suggestions towards optimal maze design for studies of fish cognition.

Fish vulnerability to capture by trapping is modulated by individual parasite density.

Commercial fishery harvest is a powerful evolutionary agent, but we know little about whether environmental stressors affect harvest-associated selection. We test how parasite infection relates to trapping vulnerability through selective processes underlying capture. We used fish naturally infected with parasites, including trematodes causing black spots under fish skin. We first assessed how individual parasite density related to standard metabolic rate (SMR), maximum metabolic rate (MMR) and absolute aerobic scope (AAS)-then used laboratory fishing simulations to test how capture vulnerability was related to parasite density. We further explored group-trapping dynamics using experimental shoals containing varying proportions of infected fish (groups of six with either 0, 2, 4 or 6 infected individuals). At the individual level, we found a positive relationship between parasite presence and SMR, but not MMR or AAS. While we saw no relationship between individual metabolic capacity and vulnerability to trapping, we found the length of time fish spent in traps increased with increasing parasite density, a predictor of trapping-related capture probability. At the group level, the number of infected individuals in a shoal did not affect overall group trapping vulnerability. Our results suggest that parasite infection has some capacity to shift individual vulnerability patterns in fisheries, and potentially influence the evolutionary outcomes of fisheries-induced evolution.

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.

Guidelines for reporting methods to estimate metabolic rates by aquatic intermittent-flow respirometry.

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.

Social dynamics obscure the effect of temperature on air breathing in Corydoras catfish.

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.

Reduced exploration capacity despite brain volume increase in warm-acclimated common minnow.

While evidence suggests that warming may impact cognition of ectotherms, the underlying mechanisms remain poorly understood. A possible but rarely considered mechanism is that the metabolic response of ectotherms to warming is associated with changes in brain morphology and function. Here, we compared aerobic metabolism, brain volume, boldness and accuracy of maze solving of common minnows (Phoxinus phoxinus) acclimated for 8 months to either their current optimal natural (14°C) or warm (20°C) water temperature. Metabolic rates indicated increased energy expenditure in warm-acclimated fish, but also at least partial thermal compensation as warm-acclimated fish maintained high aerobic scope. Warm-acclimated fish had larger brains than cool-acclimated fish. The volume of the dorsal medulla relative to the overall brain size was larger in warm- than in cool-acclimated fish, but the proportion of other brain regions did not differ between the temperature treatments. Warm-acclimated fish did not differ in boldness but made more errors than cool-acclimated fish in exploring the maze across four trials. Inter-individual differences in the number of exploration errors were repeatable across the four trials of the maze test. Our findings suggest that in warm environments, maintaining a high aerobic scope, which is important for the performance of physically demanding tasks, can come at the cost of changes in brain morphology and impairment of the capacity to explore novel environments. This trade-off could have strong fitness implications for wild ectotherms.

Using aerobic exercise to evaluate sub-lethal tolerance of acute warming in fishes.

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 (CTmax) 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 (UGT), where it first engaged the unsteady anaerobic swimming mode that preceded fatigue. After suitable recovery, each fish was exercised at 85% of their own UGT and warmed 1°C every 30 min, to identify the temperature at which they fatigued, denoted as CTswim Fish were also submitted to a standard CTmax, warming at the same rate as CTswim, under static conditions until LOE. All individuals fatigued in CTswim, at a mean temperature approximately 2°C lower than their CTmax 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 CTswim was preceded by a gait transition qualitatively indistinguishable from that during the incremental swim test. This suggests that fatigue in CTswim 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 CTswim provides an ecologically relevant sub-lethal threshold that is more sensitive to extreme events than LOE at CTmax.

Effects of repeated daily acute heat challenge on the growth and metabolism of a cold water stenothermal fish.

Temperature is an important environmental factor influencing fish physiology that varies both spatially and temporally in ecosystems. In small north temperate zone lakes, cold water piscivores rely on nearshore prey; however, this region exceeds the optimal temperature of the foraging species during summer. To cope, piscivores make short excursions into the nearshore to feed and return to cold water to digest their meal, but the physiological impacts of these repeated acute exposures to warm water are not well understood. We exposed juvenile lake trout (Salvelinus namaycush) to treatments where they were held at ∼10°C and exposed to either 17 or 22°C for 5-10 min daily for 53 days mimicking warm-water forays. Control fish, held at an average temperature of ∼10°C but not exposed to thermal variation, consumed more food and grew slightly faster than heat challenged fish, with no clear differences in body condition, hepatosomatic index, ventricle mass, or muscle concentrations of lactate dehydrogenase and cytochrome c oxidase. Aerobic metabolic rates measured at 10°C indicated that standard metabolic rates (SMR) were similar among treatments; however, fish that were repeatedly exposed to 17°C had higher maximum metabolic rates (MMR) and aerobic scopes (AS) than control fish and those repeatedly exposed to 22°C. There were no differences in MMR or AS between fish exposed to 22°C and control fish. These results suggest that although SMR of fish are robust to repeated forays into warmer environments, MMR displays plasticity, allowing fish to be less constrained aerobically in cold water after briefly occupying warmer waters.

Anemone bleaching increases the metabolic demands of symbiont anemonefish.

Increased ocean temperatures are causing mass bleaching of anemones and corals in the tropics worldwide. While such heat-induced loss of algal symbionts (zooxanthellae) directly affects anemones and corals physiologically, this damage may also cascade on to other animal symbionts. Metabolic rate is an integrative physiological trait shown to relate to various aspects of organismal performance, behaviour and locomotor capacity, and also shows plasticity during exposure to acute and chronic stressors. As climate warming is expected to affect the physiology, behaviour and life history of animals, including ectotherms such as fish, we measured if residing in bleached versus unbleached sea anemones (Heteractis magnifica) affected the standard (i.e. baseline) metabolic rate and behaviour (activity) of juvenile orange-fin anemonefish (Amphiprion chrysopterus). Metabolic rate was estimated from rates of oxygen uptake [Formula: see text], and the standard metabolic rate [Formula: see text] of anemonefish from bleached anemones was significantly higher by 8.2% compared with that of fish residing in unbleached anemones, possibly due to increased stress levels. Activity levels did not differ between fish from bleached and unbleached anemones. As [Formula: see text] reflects the minimum cost of living, the increased metabolic demands may contribute to the negative impacts of bleaching on important anemonefish life history and fitness traits observed previously (e.g. reduced spawning frequency and lower fecundity).

Conserved but flexible modularity in the zebrafish skull: implications for craniofacial evolvability.

Morphological variation is the outward manifestation of development and provides fodder for adaptive evolution. Because of this contingency, evolution is often thought to be biased by developmental processes and functional interactions among structures, which are statistically detectable through forms of covariance among traits. This can take the form of substructures of integrated traits, termed modules, which together comprise patterns of variational modularity. While modularity is essential to an understanding of evolutionary potential, biologists currently have little understanding of its genetic basis and its temporal dynamics over generations. To address these open questions, we compared patterns of craniofacial modularity among laboratory strains, defined mutant lines and a wild population of zebrafish (Danio rerio). Our findings suggest that relatively simple genetic changes can have profound effects on covariance, without greatly affecting craniofacial shape. Moreover, we show that instead of completely deconstructing the covariance structure among sets of traits, mutations cause shifts among seemingly latent patterns of modularity suggesting that the skull may be predisposed towards a limited number of phenotypes. This new insight may serve to greatly increase the evolvability of a population by providing a range of 'preset' patterns of modularity that can appear readily and allow for rapid evolution.

Digestive and locomotor capacity show opposing responses to changing food availability in an ambush predatory fish.

Metabolic rates vary widely within species, but little is known about how variation in the 'floor' [i.e. standard metabolic rate (SMR) in ectotherms] and 'ceiling' [maximum metabolic rate (MMR)] for an individual's aerobic scope (AS) are linked with digestive and locomotor function. Any links among metabolic traits and aspects of physiological performance may also be modulated by fluctuations in food availability. This study followed changes in SMR, MMR, and digestive and locomotor capacity in southern catfish (Silurus meridionalis) throughout 15 days of food deprivation and 15 days of refeeding. Individuals downregulated SMR during food deprivation and showed only a 10% body mass decrease during this time. Whereas critical swim speed (Ucrit) was robust to food deprivation, digestive function decreased after fasting with a reduced peak oxygen uptake during specific dynamic action (SDA) and prolonged SDA duration. During refeeding, individuals displayed rapid growth and digestive function recovered to pre-fasting levels. However, refed fish showed a lower Ucrit than would be expected for their increased body length and in comparison to measures at the start of the study. Reduced swimming ability may be a consequence of compensatory growth: growth rate was negatively correlated with changes in Ucrit during refeeding. Southern catfish downregulate digestive function to reduce energy expenditure during food deprivation, but regain digestive capacity during refeeding, potentially at the cost of decreased swimming performance. The plasticity of maintenance requirements suggests that SMR is a key fitness trait for in this ambush predator. Shifts in trait correlations with food availability suggest that the potential for correlated selection may depend on context.

Aggression supersedes individual oxygen demand to drive group air-breathing in a social catfish.

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.

How does school size affect tail beat frequency in turbulent water?

The energy savings experienced by fish swimming in a school have so far been investigated in an near-idealised experimental context including a relatively laminar water flow. The effects of explicitly turbulent flows and different group sizes are yet to be considered. Our repeated-measures study is a first step in addressing both of these issues: whether schooling is more energetically economical for fish when swimming in a quantified non-laminar flow and how this might be moderated by group size. We measured tail beat frequency (tbf) in sea bass swimming in a group of 3 or 6, or singly. Video data enabled us to approximately track the movements of the fish during the experiments and in turn ascertain the water flow rates and turbulence levels experienced for each target individual. Although the fish exhibited reductions in tbf during group swimming, which may indicate some energy savings, these savings appear to be attenuated, presumably due to the water turbulence and the movement of the fish relative to each other. Surprisingly, tbf was unrelated to flow rate when the fish were swimming singly or in a group of three, and decreased with increasing flow rates when swimming in a group of six. However, the fish increased tbf in greater turbulence at all group sizes. Our study demonstrates that under the challenging and complex conditions of turbulent flow and short-term changes in school structure, group size can moderate the influences of water flow on a fish's swimming kinematics, and in turn perhaps their energy costs. SUMMARY STATEMENT: The energy savings that sea bass experience from schooling are affected by flow speed or turbulence, moderated by group size.