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.

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.

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.

Impacts of climate-related stressors on social group cohesion and individual sociability in fish.

Group-living in animals comes with a number of benefits associated with predator avoidance, foraging, and reproduction. A large proportion of fish species display grouping behaviour. Fish may also be particularly vulnerable to climate-related stressors including thermal variation, hypoxia, and acidification. As climate-related stressors are expected to increase in magnitude and frequency, any effects on fish behaviour may be increased and affect the ability of fish species to cope with changing conditions. Here we conduct a systematic review of the effects of temperature, hypoxia, and acidification on individual sociability and group cohesion in shoaling and schooling fishes. Searches of the published and grey literature were carried out, and studies were included or excluded based on selection criteria. Data from studies were then included in a meta-analysis to examine broad patterns of effects of climate-related stressors in the literature. Evidence was found for a reduction in group cohesion at low oxygen levels, which was stronger in smaller groups. While several studies reported effects of temperature and acidification, there was no consistent effect of either stressor on sociability or cohesion. There was some evidence that marine fishes are more strongly negatively affected by acidification compared with freshwater species, but results are similarly inconsistent and more studies are required. Additional studies of two or more stressors in combination are also needed, although one study found reduced sociability following exposure to acidification and high temperatures. Overall, there is some evidence that hypoxia, and potentially other climate-related environmental changes, impact sociability and group cohesion in fishes. This may reduce survival and adaptability in shoaling and schooling species and have further ecological implications for aquatic systems. However, this synthesis mainly highlights the need for more empirical studies examining the effects of climate-related factors on social behaviour in fishes.

Swimming behavior affects ingestion of microplastics by fish.

Microplastics (< 5 mm) are widely found in organisms and have the potential harm to ecosystems. Despite their widespread prevalence in environments, there is high individual varation in the abundance of microplastics found in individuals of the same species. In the present study, juvenile cichlid fish (Chindongo demasoni) were chosen to determine the ingestion personality for microplastics in the laboratory. The visible implant fluorescent tags were used for individual recognition. The fish were fed with microplastic fiber, pellet, and food for comparison. Our results showed that the observation of the behaviors of fish could be successfully matched with subsequent measurements for each individual through the tag method in microplastic research. The difference in the abundance of fiber (0-27 items/ind.) among fish individuals was also observed in our study. Meanwhile, the abundance of fiber showed a positive correlation with the average speed and covered area of fish, which indicates the degree of activity of fish. Moreover, fish with higher speed or a front position had higher capturing times for pellet. Our results suggest that the swimming behaviors of fish affect their ingestion of microplastics, and active fish had a higher likelihood of ingesting microplastics, which might be one of the reasons for the common phenomena, i.e., great individual differences observed in microplastic studies.

Capture and discard practises associated with an ornamental fishery affect the metabolic rate and aerobic capacity of three-striped dwarf cichlids Apistogramma trifasciata.

Fishing causes direct removal of individuals from wild populations but can also cause a physiological disturbance in fish that are released or discarded after capture. While sublethal physiological effects of fish capture have been well studied in commercial and recreational fisheries, this issue has been overlooked for the ornamental fish trade, where it is common to capture fish from the wild and discard non-target species. We examined metabolic responses to capture and discard procedures in the three-striped dwarf cichlid Apistogramma trifasciata, a popular Amazonian aquarium species that nonetheless may be discarded when not a target species. Individuals (n = 34) were tagged and exposed to each of four treatments designed to simulate procedures during the capture and discard process: 1) a non-handling control; 2) netting; 3) netting +30 seconds of air exposure; and 4) netting +60 seconds of air exposure. Metabolic rates were estimated using intermittent-flow respirometry, immediately following each treatment then throughout recovery overnight. Increasing amounts of netting and air exposure caused an acute increase in oxygen uptake and decrease in available aerobic scope. In general, recovery occurred quickly, with rapid decreases in oxygen uptake within the first 30 minutes post-handling. Notably, however, male fish exposed to netting +60 seconds of air exposure showed a delayed response whereby available aerobic scope was constrained <75% of maximum until ~4-6 hours post-stress. Larger fish showed a greater initial increase in oxygen uptake post-stress and slower rates of recovery. The results suggest that in the period following discard, this species may experience a reduced aerobic capacity for additional behavioural/physiological responses including feeding, territory defence and predator avoidance. These results are among the first to examine impacts of discard practises in the ornamental fishery and suggest ecophysiological research can provide valuable insight towards increasing sustainable practises in this global trade.

Integrating animal behaviour into research on multiple environmental stressors: a conceptual framework.

While a large body of research has focused on the physiological effects of multiple environmental stressors, how behavioural and life-history plasticity mediate multiple-stressor effects remains underexplored. Behavioural plasticity can not only drive organism-level responses to stressors directly but can also mediate physiological responses. Here, we provide a conceptual framework incorporating four fundamental trade-offs that explicitly link animal behaviour to life-history-based pathways for energy allocation, shaping the impact of multiple stressors on fitness. We first address how small-scale behavioural changes can either mediate or drive conflicts between the effects of multiple stressors and alternative physiological responses. We then discuss how animal behaviour gives rise to three additional understudied and interrelated trade-offs: balancing the benefits and risks of obtaining the energy needed to cope with stressors, allocation of energy between life-history traits and stressor responses, and larger-scale escape from stressors in space or time via large-scale movement or dormancy. Finally, we outline how these trade-offs interactively affect fitness and qualitative ecological outcomes resulting from multiple stressors. Our framework suggests that explicitly considering animal behaviour should enrich our mechanistic understanding of stressor effects, help explain extensive context dependence observed in these effects, and highlight promising avenues for future empirical and theoretical research.

Enhancing the ecological realism of evolutionary mismatch theory.

Following rapid environmental change, why do some animals thrive, while others struggle? We present an expanded, cue-response framework for predicting variation in behavioral responses to novel situations. We show how signal detection theory can be used when individuals have three behavioral options (approach, avoid, or ignore). Based on this theory, we outline predictions about which animals are more likely to make mistakes around novel conditions (i.e., fall for a trap or fail to use an undervalued resource) and the intensity of that mismatch (i.e., severe versus moderate). Explicitly considering three options provides a more holistic perspective and allows us to distinguish between severe and moderate traps, which could guide management strategies in a changing world.

The Potential for Physiological Performance Curves to Shape Environmental Effects on Social Behavior.

As individual animals are exposed to varying environmental conditions, phenotypic plasticity will occur in a vast array of physiological traits. For example, shifts in factors such as temperature and oxygen availability can affect the energy demand, cardiovascular system, and neuromuscular function of animals that in turn impact individual behavior. Here, we argue that nonlinear changes in the physiological traits and performance of animals across environmental gradients-known as physiological performance curves-may have wide-ranging effects on the behavior of individual social group members and the functioning of animal social groups as a whole. Previous work has demonstrated how variation between individuals can have profound implications for socially living animals, as well as how environmental conditions affect social behavior. However, the importance of variation between individuals in how they respond to changing environmental conditions has so far been largely overlooked in the context of animal social behavior. First, we consider the broad effects that individual variation in performance curves may have on the behavior of socially living animals, including: (1) changes in the rank order of performance capacity among group mates across environments; (2) environment-dependent changes in the amount of among- and within-individual variation, and (3) differences among group members in terms of the environmental optima, the critical environmental limits, and the peak capacity and breadth of performance. We then consider the ecological implications of these effects for a range of socially mediated phenomena, including within-group conflict, within- and among group assortment, collective movement, social foraging, predator-prey interactions and disease and parasite transfer. We end by outlining the type of empirical work required to test the implications for physiological performance curves in social behavior.

You're Just My Type: Mate Choice and Behavioral Types.

Consistent individual differences in behavior [i.e., behavioral types (BTs)], are common across the animal kingdom. Consistency can make behavior an adaptive trait for mate choice decisions. Here, we present a conceptual framework to explain how and why females might evaluate a male's BT before mating. Because BTs are consistent across time or context, a male's BT can be a reliable indicator of his potential to provide direct benefits. Heritable BTs can enable informed mate choice via indirect benefits. Many key issues regarding patterns of mate choice, including sensory biases, context dependence, and assortative mating apply to BT-dependent mate choice. Understanding the relationship between BTs and mate choice may offer insights into patterns of variation and consistency common in behavioral traits.

Environmentally relevant concentrations of bifenthrin affect the expression of estrogen and glucocorticoid receptors in brains of female western mosquitofish.

In recent decades, pyrethroid pesticides have been deemed a safer alternative to previously used pesticides. While some evidence supports this assumption in mammals and birds, exposure to certain pyrethroids can affect concentrations of hormones vital to reproduction in fish. Thus, we hypothesized that pyrethroid exposure impacts fish reproductive behavior and the expression of genes associated with reproduction. We tested our hypothesis by examining effects of the widely used pyrethroid pesticide, bifenthrin, on the reproductive behaviors of the broadly distributed livebearing western mosquitofish, Gambusia affinis. We exposed sexually mature female fish to one of five environmentally relevant concentrations of bifenthrin and conducted behavioral assays to assess reproductive, social, and space use behaviors before and after exposure. We did not detect changes in behaviors measured in response to bifenthrin. However, exposure was associated with increased expression of an estrogen receptor gene (ER-α) and glucocorticoid receptor (GR) in brain tissue at bifenthrin concentrations at concentrations of 5.90 and 24.82 ng/L, and 5.90 and 12.21 ng/L, respectively. Our study supports the perspective that the use of multiple endpoints through integrative approaches is essential for understanding the cumulative impact of pollutants. Integrating physiological, morphological, and behavioral investigations of nonlethal concentrations of pollutants like bifenthrin may heighten our potential to predict their impact on individuals, populations, and communities.