Subordinate Fish Research Articles

11β-Hydroxysteroid dehydrogenase type 2 may mediate the stress-specific effects of cortisol on brain cell proliferation in adult zebrafish (Danio rerio).

Stress-induced increases in cortisol can stimulate or inhibit brain cell proliferation, but the mechanisms behind these opposing effects are unknown. We tested the hypothesis that 11β-hydroxysteroid dehydrogenase type 2 (Hsd11b2), a glucocorticoid-inactivating enzyme expressed in neurogenic regions of the adult zebrafish brain, mitigates cortisol-induced changes to brain cell proliferation, using one of three stress regimes: a single 1min air exposure (acute stress), two air exposures spaced 24 h apart (repeat acute stress) or social subordination (chronic stress). Plasma cortisol was significantly elevated 15 min after air exposure and recovered within 24 h after acute and repeat acute stress, whereas subordinate fish exhibited significant and sustained elevations relative to dominant fish for 24 h. Following acute stress, brain hsd11b2 transcript abundance was elevated up to 6 h after a single air exposure but was unchanged by repeat acute stress or social subordination. A sustained increase in brain Hsd11b2 protein levels occurred after acute stress, but not after repeat or chronic stress. Following acute and repeat acute stress, brain pcna transcript abundance (a marker of cell proliferation) exhibited a prolonged elevation, but was unaffected by social subordination. Interestingly, the number of telencephalic BrdU+ cells increased in fish after a single air exposure but was unchanged by repeat acute stress. Following acute and repeat acute stress, fish expressed lower brain glucocorticoid and mineralocorticoid receptor (gr and mr) transcript abundance while subordinate fish exhibited no changes. Taken together, these results demonstrate stressor-specific regulation of Hsd11b2 in the zebrafish brain that could modulate rates of cortisol catabolism contributing to observed differences in brain cell proliferation.

Submissive behaviour is affected by group size in a social fish.

For social groups to form and be stable over time, animals must develop strategies to cope with conflict among group members. Animals may behave submissively either by fleeing from an aggressor, or by signalling submission. The use of these two submissive responses may vary depending on the social and ecological context. Group size is a key aspect of social context for group living animals, as individuals in smaller groups may respond to aggression differently than those from larger groups. Here, we examine the relationship between group size and submissive behaviour in a cooperatively breeding fish, the daffodil cichlid (Neolamprologus pulcher). We found that subordinate fish showed similar levels of submission signals in response to dominant aggression in larger and smaller groups, however, subordinates from larger groups were less likely to flee from dominant aggression than those in smaller groups. Subordinates in larger groups also showed more digging behaviour which may be also used to avoid conflict with the dominant group members. Our data show that social context affects submissive behaviour in a cooperatively breeding fish.

Exploring transcriptional and post-transcriptional epigenetic regulation of crf and 11βhsd2 in rainbow trout brain during chronic social stress

Using dominance hierarchies in juvenile rainbow trout (Oncorhynchus mykiss) as a model of chronic social stress in fish, we explored whether epigenetic transcriptional and post-transcriptional mechanisms are involved in the gene expression of corticotropin-releasing factor (crf) and 11β-hydroxysteroid dehydrogenase (11βhsd2), key factors involved in the regulation of the endocrine stress axis response. In juvenile rainbow trout pairs, subordinate individuals display sustained elevation of circulating cortisol concentrations. Cortisol production is controlled by the hypothalamic-pituitary-interrenal (HPI) axis in fish and initiated by CRF release from the preoptic area (POA). Given that crf is modulated during chronic social stress, and that such stress has been implicated in the epigenetic regulation of crf in other taxa, we probed a role for epigenetic regulation of crf transcript abundance in chronically stressed rainbow trout. We also investigated the regulation of the cortisol-metabolising enzyme 11βhsd2 in the POA, which is upregulated in subordinates. The potential involvement of DNA methylation and microRNAs (miRNAs) in the regulation of crf transcript abundance was investigated during social stress in the POA of fish, as was the potential involvement of miRNAs in 11βhsd2 regulation. Although transcript abundances of crf were elevated in subordinate fish after 4 days, DNA methylation profiles within putative promoter sequences upstream of the crf gene were not significantly affected by chronic stress. An inverse relationship between crf and its predicted posttranscriptional regulator miR-103a-3p in the POA suggests that miRNAs may be involved in mediating the effects of chronic social stress on key components of the endocrine stress axis.

Negative feedback regulation in the hypothalamic-pituitary-interrenal axis of rainbow trout subjected to chronic social stress

The formation of dominance hierarchies in pairs of juvenile rainbow trout (Oncorhynchus mykiss) results in subordinate individuals exhibiting chronically elevated plasma cortisol concentrations. Cortisol levels reflect a balance between cortisol production, which is coordinated by the hypothalamic-pituitary-interrenal (HPI) axis in teleost fish, and negative feedback regulation and hormone clearance, which act to lower cortisol levels. However, the mechanisms contributing to the longer-term elevation of cortisol levels during chronic stress are not well established in fishes. The current study aimed to determine how subordinate fish maintain elevated cortisol levels, by testing the prediction that negative feedback and clearance mechanisms are impaired by chronic social stress. Plasma cortisol clearance was unchanged by social stress based on a cortisol challenge trial, hepatic abundance of the cortisol-inactivating enzyme 11-beta hydroxysteroid dehydrogenase type 2 (11βHSD2), and tissue fate of labelled cortisol. The capacity for negative feedback regulation in terms of transcript and protein abundances of corticosteroid receptors in the preoptic area (POA) and pituitary appeared stable. However, changes in 11βHSD2 and mineralocorticoid receptor (MR) expression suggest subtle regulatory changes in the pituitary that may alter negative feedback. The chronic cortisol elevation observed during social subordination likely is driven by HPI axis activation and compounded by dysregulated negative feedback.

Social status-dependent regulation and function of the somatotropic axis in juvenile rainbow trout

Juvenile rainbow trout (Oncorhynchus mykiss) develop social hierarchies when competing for resources in a constrained environment. Among the physiological consequences of social status are changes in organismal energy metabolism, which generally favour anabolic pathways in dominant fish and catabolic pathways in subordinate fish. The somatotropic axis is an important regulator of metabolism and growth that could be involved in mediating metabolic changes in response to social status in juvenile rainbow trout. Here we used juvenile trout housed either in dyads or individually (sham controls) to determine whether social status changes indices of somatotropic axis function. Although pituitary growth hormone expression (gh1 and gh2) did not differ among groups, circulating growth hormone (GH) increased ∼12-fold in subordinate fish compared to sham and dominant fish. Social status caused consistent differential expression of GH receptor paralogues in liver and muscle, two principal target tissues of GH. Compared to dominant and/or sham fish, ghra paralogue expression (ghra1 and ghra2) was lower, while ghrb1 expression was higher in subordinate fish. Across tissues, ghra paralogue expression was generally positively correlated with expression of insulin growth factors (igf1, igf2), while ghrb1 expression was positively correlated with transcript abundance of hormone sensitive lipase (hsl1). Because igf and hsl expression are subject to context-dependent GH control in rainbow trout, these results suggest that increased circulating GH in conjunction with differential expression of ghr paralogues may translate into prioritization of downstream catabolic lipolytic pathways in subordinate rainbow trout. These findings support a social context-dependent role for GH signalling in mediating metabolic changes in juvenile rainbow trout.

Regulation of cortisol production during chronic social stress in rainbow trout

Chronic stress resulting from social interactions impacts the endocrine stress response in many vertebrates, including teleost fishes. Juvenile rainbow trout held in pairs form a dominance hierarchy with the subordinate individual exhibiting chronic elevation of plasma cortisol and an attenuated cortisol response to an additional acute stressor. The current study investigated the mechanisms underlying this apparent dichotomy in cortisol production at the level of the head kidney (adrenal homolog). Following four days of chronic social stress, subordinate rainbow trout exhibited elevated plasma cortisol levels that correlated with basal cortisol production by the head kidney in vitro. Subordinate trout had higher transcript abundances of steroidogenic acute regulatory protein and cytochrome p450 side chain cleavage enzyme, which facilitate key steps in steroidogenesis, as well as two paralogs of steroidogenic factor 1. Despite elevation of basal steroidogenesis, acute cortisol production in response to ACTH (in vivo and in vitro) was lower in subordinate trout. Transcript abundances of the ACTH receptor accessory proteins were elevated in subordinate fish, but head kidney cortisol production in response to a cAMP analogue was lower than in dominant fish. Together, the data suggest that the attenuated acute cortisol response of subordinate trout reflects limitations on cortisol production downstream of cAMP signalling in steroidogenic cells of the head kidney, despite the increased basal abundance of key components of the steroidogenic pathway.

Identifying the Interaction of the Brain and the Pituitary in Social – and Reproductive – State of Tilapia by Transcriptome Analyses

Introduction: As in all vertebrates, reproduction in fish is regulated by gonadotrophin-releasing hormone (GnRH) control on gonadotrophic hormones (GtHs) activity. However, the neuroendocrine factors that promote GnRH and GtH activity are unknown. In Nile tilapia (Oreochromis niloticus), sexual activity and reproduction ability depend on social rank; only dominant males and females reproduce. Here, this characteristic of dominant fish allows us to compare brain and pituitary gene expression in animals that do and do not reproduce, aiming to reveal mechanisms that regulate reproduction. Methods: An extensive transcriptome analysis was performed, combining two sets of transcriptomes: a novel whole-brain and pituitary transcriptome of established dominant and subordinate males, together with a cell-specific transcriptome of luteinizing hormone (LH) and follicle-stimulating hormone cells. Pituitary incubation assay validated the direct effect of steroid application on chosen genes and GtH secretion. Results: In most dominant fish, as determined behaviorally, the gonadosomatic index was higher than in subordinate fish, and the leading upregulated pituitary genes were those coding for GtHs. In the brain, various neuropeptide genes, including isotocin, cholecystokinin, and MCH, were upregulated; these may be related to reproductive status through effects on behavior and feeding. In a STRING network analysis combining the two transcriptome sets, brain aromatase, highly expressed in LH cells, is the most central gene with the highest number of connections. In the pituitary incubation assay, testosterone and estradiol increased the secretion of LH and specific gene transcription. Conclusions: The close correlation between behavioral dominance and reproductive capacity in tilapia allows unraveling novel genes that may regulate the hypothalamic-pituitary-gonadal axis, highlighting aromatase as the main factor affecting the brain and pituitary in maintaining a sexually active organism.

Effects of Social Hierarchy Establishment on Stress Response and Cell Phagocytosis in Gilt-Head Sea Bream (Sparus aurata)

Social stress can affect the ability of fish to respond to various stressors, such as pathogens or environmental variations. In this paper, the effects of social stress on gilt-head bream (Sparus aurata) were investigated. To study the effects of physiological stress, we evaluated biochemical and cellular parameters, such as cortisol, glucose, lactate, osmolarity, and phagocytosis, 24 h after the establishment of social hierarchy in a group of three fish. Social hierarchy was determined and characterized by behavioral observation (aggressive acts and feeding order) of the specimens (dominant: “α”; subordinate: “β” and “γ”). After the establishment of social hierarchy, we observed that, overall, levels of plasma cortisol and other biochemical plasmatic stress markers (glucose and lactate) were higher in subordinate individuals than in dominant individuals. In addition, the modulation of phagocytic activity of the peritoneal exudate cells (PECs) demonstrated that social stress appeared to affect immune response. Finally, principal component analysis clearly separated the subordinate fish groups from the dominant groups, based on stress markers and the phagocytic activity of peritoneal exudate cells. This study contributes to current knowledge on gilt-head sea bream, helping to understand the link between social stress, behavior, and physiology of this species, relevant in the aquaculture sector, where fish are subjected to several kinds of stress.

Low concentrations of the antidepressant venlafaxine affect courtship behaviour and alter serotonin and dopamine systems in zebrafish (Danio rerio).

Venlafaxine, a serotonin-noradrenaline reuptake inhibitor, is a widely used antidepressant drug routinely detected in aquatic environments. However, its potential impact on courtship behaviour in zebrafish is unknown. We tested the hypothesis that venlafaxine disrupts brain monoamine levels and molecular responses essential for courtship behaviour in zebrafish. Zebrafish (Danio rerio) were exposed to venlafaxine (1, 10, and 100μg/L) for 20 days. We evaluated the molecular levels and neuronal basis of the effect of venlafaxine on courtship behaviour. Here, we show that venlafaxine inhibited courtship behaviour in zebrafish and increased the transcript levels of 5-ht1a and 5-ht2c while decreasing the transcript levels of genes involved in the dopaminergic system, including th1, th2, drd1b, and drd2b. Venlafaxine upregulated 5-HT levels and downregulated dopamine levels. Moreover, the subordinate fish from the venlafaxine-exposed group had significantly lower motor activity than the subordinate fish of the control group. Collectively, our results reveal that venlafaxine can disturb brain monoamine levels, affecting courtship behaviour in adult zebrafish.

Growth and Hepatic Insulin-Like Growth Factor-I Gene Expression in Dominant and Subordinate Oreochromis Niloticus L Under Limited Period of Physical Interaction

Social stress is known to regulate several aspects of the teleost physiology. This study explored the influence of limited period of physical interaction on growth and hepatic insulin-like growth factor-I (IGF-I) messenger RNA (mRNA) expression of the fish. Twenty all-male Oreochromis niloticus were isolated for 10 days and were used in a social pair study. After the social interaction was settled, dominant and subordinate individuals in a pair were reared separately in one aquarium separated by glass divider. The fish were fed at the same rate daily to remove the possible effect of nutrition. The glass divider was removed 10 min daily for social interaction. Weight was monitored on Days 2, 7 and 14 during the experimental period, then hepatic IGF-I mRNA expression was quantified. During the 14 days social experiment, mean specific growth rate of dominant fish (1.6%·day−1) was significantly higher (P<0.01) than that of the subordinate fish (0.9%·day−1). Dominant fish also had 2.5 fold significantly higher (P<0.05) mean IGF-I mRNA expression than that of subordinate fish. These indicate that even under limited period of physical contact but with period of visual communication, social status regulates growth and hepatic IGF-I gene expression in this species of fish. There was also a significant positive correlation (r = 0.52; P<0.01) between growth and IGF-I mRNA level which supports the previous studies that hepatic IGF-I gene expression has a potential utility as an instantaneous growth rate indicator for O. niloticus.

Social status modulates the behavioral and physiological consequences of a chemical pollutant in animal groups.

The social environment (i.e., the suite of social interactions that occur among individuals that can result in variation in social ranks) is a commonly overlooked aspect of biology when scientists evaluate the effects of chemical contaminants. The social environment, however, represents the arena in which individual-level performance shapes group- or population-level outcomes and may therefore mediate many of the ultimate consequences of chemicals for wildlife. Here, we evaluated the role that the social environment plays in determining the consequences of pollutant exposure. We exposed groups of juvenile brown trout (Salmo trutta) to an emerging pharmaceutical pollutant that is commonly detected in freshwaters (the benzodiazepine, oxazepam) and allowed them to form dominance hierarchies. Exposure affected dominant and subordinate fish differently, causing fish to become less aggressive at high doses and subordinate fish to become more competitively successful at low doses. These perturbations had further consequences for growth, fin damage, and survival. Exposure also modulated physiological stress in the hierarchy, and social status itself affected how much oxazepam was absorbed in tissues, potentially creating a dynamic feedback loop that further influences the asymmetric effects of exposure on differing social statuses. Many effects followed a "U-shaped" dose-response curve, highlighting the importance of nonlinear, low-dose effects. Altogether, we show that social structure in animal groups can interact with and modulate the effects of an environmental contaminant. We underscore the need to account for an organism's natural ecological context, including their social environment, in future experiments and environmental risk assessments to predict the effects of chemical contaminants on wildlife.

Stress, Subordination, and Anomalies of Feeding Across the Tree of Life: Implications for Interpreting Human Eating Disorders

Eating behaviors of animals living in naturalistic environments offer unique insights into several dysregulated eating patterns observed in humans. Social subordination is a known precipitant of hyperphagia and hypophagia in human beings, and examples of similar responses have been identified in a phylogenetically widespread range of vertebral species. This points to potentially conserved, patterned responses to animals navigating lives within social hierarchies. Self-imposed food restriction in subordinate fish and hyperphagic responses in socially subordinated bird and primate individuals may represent evolved adaptations to the stress of social subordination. As such, hyperphagic and hypophagic responses to social subordination in these species may model the natural history, neurobiology, and behavioral ecology of human dieting and bingeing more accurately than some current animal models.Phylogenetically widespread similarities in eating patterns under the stress of social subordination point to potentially shared biological benefits of these behaviors across species and the role of evolutionary trade-offs, adaptations, and other processes in shaping them. The application of a broadly comparative lens to disordered eating behaviors in other species exposes important similarities and differences between neurophysiology of eating across species. In doing so, it highlights the value of phylogenetic analyses and macroevolution as tools for identifying novel, naturally occurring models for understanding disordered human eating. Moreover, this approach introduces the intriguing possibility that human cultural influences on disordered eating may have far more ancient origins than previously considered.

Social effects on AVT and CRF systems

Stress and aggression have negative effects on fish welfare and productivity in aquaculture. Thus, research to understand aggression and stress in farmed fish is required. The neuropeptides arginine-vasotocin (AVT) and corticotropin-releasing factor (CRF) are involved in the control of stress and aggression. Therefore, we investigated the effect of agonistic interactions on the gene expression of AVT, CRF and their receptors in juvenile rainbow trout (Oncorhynchus mykiss). The social interactions lead to a clear dominant-subordinate relationship with dominant fish feeding more and being more aggressive. Subordinate fish had an upregulation of the AVT receptor (AVT-R), an upregulation of CRF mRNA levels, and higher plasma cortisol levels. The attenuating effect of AVT on aggression in rainbow trout is proposed to be mediated by AVT-R, and the attenuating effect of the CRF system is proposed to be mediated by CRF.

Neuroendocrine Mechanisms Underlying Non-breeding Aggression: Common Strategies Between Birds and Fish.

Aggression is an adaptive behavior that plays an important role in gaining access to limited resources. Aggression may occur uncoupled from reproduction, thus offering a valuable context to further understand its neural and hormonal regulation. This review focuses on the contributions from song sparrows (Melospiza melodia) and the weakly electric banded knifefish (Gymnotus omarorum). Together, these models offer clues about the underlying mechanisms of non-breeding aggression, especially the potential roles of neuropeptide Y (NPY) and brain-derived estrogens. The orexigenic NPY is well-conserved between birds and teleost fish, increases in response to low food intake, and influences sex steroid synthesis. In non-breeding M. melodia, NPY increases in the social behavior network, and NPY-Y1 receptor expression is upregulated in response to a territorial challenge. In G. omarorum, NPY is upregulated in the preoptic area of dominant, but not subordinate, individuals. We hypothesize that NPY may signal a seasonal decrease in food availability and promote non-breeding aggression. In both animal models, non-breeding aggression is estrogen-dependent but gonad-independent. In non-breeding M. melodia, neurosteroid synthesis rapidly increases in response to a territorial challenge. In G. omarorum, brain aromatase is upregulated in dominant but not subordinate fish. In both species, the dramatic decrease in food availability in the non-breeding season may promote non-breeding aggression, via changes in NPY and/or neurosteroid signaling.

Preference and Motivation Tests for Body Tactile Stimulation in Fish.

Simple SummaryBody tactile stimulation, such as human massage therapy, is a way to relieve stress in humans and other animals, therefore it could improve animal health and welfare. This physical stimulation can also be done through artificial devices, as a sensory enrichment. However, before using it in an artificial environment, it is imperative to test whether animals perceive such enrichment as positive (searching for it spontaneously) or negative (avoiding it). Here, we tested whether the Nile tilapia fish search for or avoid tactile stimulation. We used a rectangular PVC frame, filled with vertical plastic sticks sided with silicone bristles that provided tactile stimulation when fish passed through them. We carried out preference and motivation tests, in which fish could choose to cross through the device with and without tactile stimulus. The same procedure was repeated after fish were exposed to either isolation or social stress. We found that fish crossed less by tactile device than by open areas. However, as fish spontaneously crossed through the bristles, and overcame an aversive high-intensity lighted route to reach the device, we conclude that tactile stimulation is not a negative condition. Thus, further studies can be designed to test several effects of tactile stimulation on the welfare of fish.We tested whether territorial fish (Nile tilapia) perceive body tactile stimulation as a positive or negative resource. Individual male fish were placed for eight days in an aquarium containing a rectangular PVC frame, which was filled with vertical plastic sticks sided with silicone bristles in the middle of the tank. Fish passing this device received a tactile stimulus. The fish then underwent a preference test by choosing between areas half-with and half-without tactile bristles. Then, fish were submitted to a motivation test where they had to pass an aversive stimulus (bright light) to access the device. Fish were, then, paired to settle social rank, which occurs by way of fights (social stressor), and were assigned again to preference and motivation tests. A group without social stress was used as a control. Contrary to our expectations, fish preferred the area without tactile bristles, although subordinate fish reached tactile stimulation more than the dominant one. Social stress did not affect the preference and motivation, suggesting that fish do not perceive tactile stimulation as a stressor reliever. However, as fish did not avoid the stimulation, reached the device spontaneously, and faced an aversive stimulus to access it, we conclude that tactile stimulation is not a negative condition and, therefore, can be used in further studies regarding fish welfare.

Elevated cortisol lowers thermal tolerance but results in limited cardiac remodelling in rainbow trout (Oncorhynchus mykiss) experiencing chronic social stress.

Juvenile rainbow trout (Oncorhynchus mykiss) held in pairs form dominance hierarchies in which subordinate individuals experience chronic social stress accompanied by lowered thermal tolerance (assessed as the critical thermal maximum, CTmax). Here, we tested the hypothesis that chronic elevation of circulating cortisol levels reduces thermal tolerance in subordinate trout. In support of this hypothesis, subordinate trout that recovered from social stress for 48 h, a period sufficient to return cortisol to normal baseline levels, no longer showed reduced CTmax. Further, thermal tolerance was not restored in subordinates treated with cortisol during recovery from social stress. To explore possible mechanisms underlying the effect of chronic stress on CTmax, we also tested the hypothesis that chronic cortisol elevation induces cardiac remodelling in subordinate trout, as previously reported for cortisol-treated rainbow trout. Ventricle mass and cardiac hypertrophy markers were unaffected by social stress. Picrosirius Red staining revealed a trend for lower collagen levels in the ventricles of subordinate relative to dominant trout. However, collagen type I transcript and protein levels, and markers of collagen turnover were unaffected. Indicators of cardiac function, including ventricle passive stiffness and intrinsic heart rate (fH), similarly were unaffected. In vivo fH was also similar between subordinate and dominant fish. Nevertheless, in keeping with their lower CTmax, subordinate fish exhibited cardiac arrhythmia at significantly lower temperatures than dominant fish during CTmax trials. Thus, high baseline cortisol levels in subordinate trout result in lowered thermal tolerance, but 5 days of social stress did not greatly affect cardiac structure or function.

Identifying and mitigating systematic biases in fish habitat simulation modeling: Implications for estimating minimum instream flows

AbstractHabitat simulation approaches (e.g., PHABSIM) have been used to model instream flows in thousands of streams and rivers and remain the most widely implemented detailed instream flow methodology. However, recent studies suggest that conventional habitat simulation models incorporate assumptions that may systematically underestimate instream flow needs, particularly for drift‐feeding fish. These include: (i) systematic biases in velocity habitat suitability curves (HSCs) caused by territoriality where dominant individuals displace subordinate fish to lower velocity micro‐habitats at high densities, thereby inflating the fitness value of low velocities; (ii) habitat simulation models do not account for flow effects on prey flux to drift‐feeding fishes, which may decrease more rapidly with reduced flow than does available habitat; (iii) use of focal velocities to construct traditional HSCs, which systematically underestimates velocity preference within the broader foraging arena of a drift‐feeding fish, and (iv) inadvertent use of low‐velocity HSCs associated with daytime refuging behavior from predators that may underestimate the higher velocities necessary for crepuscular foraging. Collectively, these factors suggest that current and historic flow prescriptions using traditional habitat simulation methods may underestimate optimal rearing flows for salmonids and other drift‐feeding species by anywhere from 10 to 50%. This implies that traditional instream flow management may be failing to provide the intended level of protection for drift‐feeding fishes in multiple streams at landscape scales. We provide guidelines for identifying contexts where model predictions are likely to be biased and approaches for correcting them.

Chronic social stress alters protein metabolism in juvenile rainbow trout, Oncorhynchus mykiss

When confined in pairs, juvenile rainbow trout (Oncorhynchus mykiss) form dominance hierarchies in which subordinate fish exhibit characteristic physiological changes including reduced growth rates and chronically elevated plasma cortisol concentrations. We hypothesized that alterations in protein metabolism contribute to the reduced growth rate of socially stressed trout, and predicted that subordinate trout would exhibit reduced rates of protein synthesis coupled with increases in protein degradation. Protein metabolism was assessed in dominant and subordinate fish after 4 days of social interaction, and in fish that were separated after 4 days of interaction for a 4 days recovery period, to determine whether effects on protein metabolism recovered when social stress was alleviated. Protein metabolism was assessed in liver and white muscle by measuring the fractional rate of protein synthesis and markers of protein degradation. In the white muscle of subordinate fish, protein synthesis was inhibited and activities of the ubiquitin-proteasome pathway (UPP) and the autophagy lysosomal system (ALS) were elevated. By contrast, the liver of subordinate fish exhibited increased rates of protein synthesis and activation of the ALS. When allowed to recover from chronic social stress for 4 days, differences in protein metabolism observed in white muscle of subordinate fish during the interaction period disappeared. In liver, protein synthesis returned to baseline levels during recovery from social stress, but markers of protein degradation did not. Collectively, these data support the hypothesis that inhibition of muscle protein synthesis coupled with increases in muscle protein breakdown contribute to the reduced growth rates of subordinate rainbow trout.

Does shelter influence the metabolic traits of a teleost fish?

Availability of shelter is an important component of habitat selection for animals as it can influence survival (protection against harsh physical conditions and predation) and growth (energy acquisition and expenditure). Few studies address the effect of shelter on metabolic expenditures associated with non-mechanical tasks (excluding station holding or movement). The main goal of this study was to investigate the influence of shelter use on metabolic traits of smallmouth bass (Micropterus dolomieu) from two populations (Kiamika River and Lake Long). Respirometry experiments on smallmouth bass were conducted to measure standard metabolic rate (SMR), resting metabolic rate (RMR), aerobic scope (AS), recovery time (RT) and excess post-exercise oxygen consumption (EPOC) in the presence or absence of shelter. The presence of shelter did not affect most metabolic traits, except for RMR, which was reduced in the presence of shelter for Lake Long fish. The results of this study also show that larger fish had lower SMR in the presence of shelter than when it was absent. When accounting for social hierarchy, there were no differences in most metabolic traits in dominant or subordinate fish in the presence or absence of shelter, except for RT, which was significantly lower in the presence of shelter for dominant fish. These results do not support the existence of an unequivocal relationship between individual metabolic traits and the presence of shelter. If physiological motives may influence the use of shelter, sheltering in itself might not have important consequences on energy expenditures required for non-mechanical tasks.

Interspecific social dominance networks reveal mechanisms promoting coexistence in sympatric charr in Hokkaido, Japan.

Coexistence of species requires equalizing mechanisms that minimize fitness differences, which are balanced by stabilizing mechanisms that enhance negative intraspecific interactions versus interspecific ones. Here, we develop a simple theoretical framework that allows measuring the relative strength of intraspecific versus interspecific competition in dominance hierarchies. We use it to evaluate mechanisms promoting coexistence between two congeneric charr that compete for foraging positions, which strongly influence density-dependent growth and survival. Agonistic interactions (n=761) among 71 Dolly Varden Salvelinus malma and whitespotted charr Salvelinus leucomaenis were measured by snorkelling in two pools in the sympatric zone of a Hokkaido stream during two summers. Interspecific dominance hierarchies, analysed using three methods, were closely correlated with fish length but the species treated each other equally. Ranks for the most dominant fish in each pool, determined directly by knockout experiments, were also virtually identical to ranks by length. Similarly, exponential random graph modelling of the social networks provided no evidence that either species was dominant over the other. Instead, larger fish were more likely to win contests, especially over fish of the next lower ranks. These results demonstrated that the two species were nearly ecological equivalents in accessing key resources in this sympatric zone. Nearly identical growth and stable densities over 4years further supported this inference, although Dolly Varden were a minority (29% of the assemblage), a sign of some fitness difference. Detailed foraging observations coupled with two concurrent studies revealed an effective stabilizing mechanism. Dolly Varden shifted to feeding directly from the benthos when drifting invertebrates declined, a behaviour enhanced by morphological character displacement, thereby partitioning food resources and enhancing intraspecific competition while avoiding agonistic encounters with whitespotted charr. The plurality of evidence indicates that fitness differences between these ecologically equivalent species are small in this local assemblage, and balanced by resource partitioning, a modest stabilizing mechanism that promotes coexistence. The theoretical framework presented here is a useful tool to evaluate the strength of interspecific versus intraspecific competition, which combined with information on trade-offs in ecological performance can contribute to a mechanistic understanding of species coexistence.