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Farmed Atlantic salmon (Salmo salar) welfare and performance can be strongly influenced by stress episodes caused by handling during farming practices. To better understand the changes occurring after an acute stress response, we exposed a group of Atlantic salmon parr to an acute stressor, which involved netting and transferring fish to several new holding tanks. We describe a time-course response to stress by sampling parr in groups before (T0) and 10, 20, 30, 45, 60, 120, 240, 300, and 330 min post-stress. A subgroup of fish was also subjected to the same stressor for a second time to assess their capacity to respond to the same challenge again within a short timeframe (ReStressed). Fish plasma was assessed for adrenocorticotropic hormone (ACTH), cortisol, and ions levels. Mucus cortisol levels were analyzed and compared with the plasma cortisol levels. At 5 selected time points (T0, 60, 90, 120, 240, and ReStressed), we compared the head kidney transcriptome profile of 10 fish per time point. The considerably delayed increase of ACTH in the plasma (60 min post-stress), and the earlier rise of cortisol levels (10 min post-stress), suggests that cortisol release could be triggered by more rapidly responding factors, such as the sympathetic system. This hypothesis may be supported by a high upregulation of several genes involved in synaptic triggering, observed both during the first and the second stress episodes. Furthermore, while the transcriptome profile showed few changes at 60 min post-stress, expression of genes in several immune-related pathways increased markedly with each successive time point, demonstrating the role of the immune system in fish coping capacity. Although many of the genes discussed in this paper are still poorly characterized, this study provides new insights regarding the mechanisms occurring during the stress response of salmon parr and may form the basis for a useful guideline on timing of sampling protocols.

Dip nets are widely used to retrieve fish from the water but can cause injuries to the fish. The objective of this study was to document the potential effects of dip net mesh size with respect to external injuries, more specifically fin splits and scale losses, in Atlantic salmon (Salmo salar). The study included 273 post-smolt Atlantic salmon from two fish batches referred to as the Tank group (w¯¯¯ = 178 ± 36 g, n = 198) and the Cage group (w¯¯¯ = 1319 ± 334 g, n = 75). Four identical dip nets with mesh sizes of 5, 10, 15, and 20 mm, respectively, were used to net fish out of their enclosures before the external injuries were assessed by visual inspection. The results showed that the number and severity of fin splits increased with dip net mesh size, but no effect of mesh size on the percentage of scale losses was found. Dip-netting of five fish at a time instead of one with the 20 mm dip net, had no statistically significant effect on the examined injuries. It is concluded that the use of dip nets with smaller mesh sizes can be positive for fish welfare by reducing fin splitting.

A better understanding of carotenoid dynamics (transport, absorption, metabolism, and deposition) is essential to develop a better strategy to improve astaxanthin (Ax) retention in muscle of Atlantic salmon. To achieve that, a comparison of post-smolt salmon with (+ Ax) or without (− Ax) dietary Ax supplementation was established based on a transcriptomic approach targeting pyloric, hepatic, and muscular tissues. Results in post-smolts showed that the pyloric caeca transcriptome is more sensitive to dietary Ax supplementation compared to the other tissues. Key genes sensitive to Ax supplementation could be identified, such as cd36 in pylorus, agr2 in liver, or fbp1 in muscle. The most modulated genes in pylorus were related to absorption but also metabolism of Ax. Additionally, genes linked to upstream regulation of the ferroptosis pathway were significantly modulated in liver, evoking the involvement of Ax as an antioxidant in this process. Finally, the muscle seemed to be less impacted by dietary Ax supplementation, except for genes related to actin remodelling and glucose homeostasis. In conclusion, the transcriptome data generated from this study showed that Ax dynamics in Atlantic salmon is characterized by a high metabolism during absorption at pyloric caeca level. In liver, a link with a potential of ferroptosis process appears likely via cellular lipid peroxidation. Our data provide insights into a better understanding of molecular mechanisms involved in dietary Ax supplementation, as well as its beneficial effects in preventing oxidative stress and related inflammation in muscle.

It is well established that farmed Atlantic salmon (Salmo salar) need n3-highly unsaturated fatty acids (HUFA) in their diet to thrive and grow. However, the biological functions to the individual HUFAs may differ, implying that future supplementation could require fixed ratios for maximum benefit. The intestinal barrier is essential to f ish health, and any disruption of the barrier can have detrimental effects. The current experiment was designed to examine the response of the intestinal hindgut when fed a low HUFA diet with 8.5 g/kg EPA + DHA (4.5% total fatty acid) or two high HUFA diets, with either high DHA (28.7 g/kg and 5.9% total fatty acid) or high EPA (25 g/kg and 14.2% total fatty acid). The diets were fed to Atlantic salmon post-smolt over 10 weeks and thereafter exposed to 3 weeks of chronic stress. After 10 weeks of feeding there were no differences in intestinal permeability and integrity, but intestinal morphology indicated increased intestinal health in the high EPA group. Gene expression also suggest that fish fed the high EPA diet had more regulation of pathways related to protein turnover compared to the high DHA fed fish. There was also indication of lower energy utilization in the low HUFA fed fish than high HUFA. Subjecting fish to 3 weeks of chronic stress led to a reduction in transepithelial resistance, increased ion flux and active L-lysine transport across the intestinal barrier in addition to a decrease in mucosal fold, enterocyte height and supranuclear vacuole density and an increase in thickness of the intestinal muscularis. After stress, the low HUFA group showed signs of inflammation with increased infiltration of MHCII positive cells. Gene expression also showed that low HUFA fed fish had a lower response to chronic stress compared to the high HUFA groups. Comparing fish fed either high DHA or EPA exposed to chronic stress showed few physical effects, but a lower density of supranuclear vacuoles and upregulation in immune-related gene expression indicate inflammation in the high DHA group.

Increased cortisol production and oxygen consumption are commonly used as indicators for stress in most fish species. For comparative studies measuring acute stress responsiveness, it is often assumed that the cortisol is sampled at peak response. However, the effect of acclimated temperature on cortisol response is largely unknown, and so is also the dynamics of oxygen uptake following stress. In the present study we acclimated Atlantic salmon post-smolt to four different temperature within the species relevant temperature spectrum (4, 8, 12 and 17 °C), and applied an acute stressor of 60 s of confinement. Cortisol, as measured in water, was sampled before and up to 360 min following the stressor, while oxygen consumption rates was estimated per minute. Basal cortisol production increased significantly with temperature, which must be considered when calculating the cortisol response per se. The cortisol response followed a Gaussian pattern, where the amplitude (max value) increased with temperature while the curve width (time before peak and recovery) decreased. Also, the total amount of cortisol released in response to the stressor increased with temperature. The cortisol peak at 4, 8, 12 and 17 °C was respectively measured after 123.75 ± 7.2, 67.5 ± 9.7, 56.25 ± 3.75 and 33.74 ± 3.75 min. The stress induced peak in oxygen uptake increased with temperature in an exponential manner, and both peak value and the total oxygen uptake above baseline levels was three times higher at 17 than at 12 °C, while the total oxygen uptake was similar between 4, 8, 12 °C. This strong difference in oxygen uptake between 12 and 17 °C was not reflected by the cortisol levels, and suggests that salmon are extra vulnerable for creating hypoxic conditions when stressed at high temperature. The current work suggests when to sample cortisol for attaining peak values at a given acclimated temperatures in salmon post-smolt. This should allow for better comparison of cortisol values between studies.

Warm water treatment has in recent years become widely used for ridding salmonids of sea lice in aquaculture although the consequences of the treatment for fish welfare are not adequately investigated. The objective of this study was to document potential long-term welfare effects of repeated warm water treatments on Atlantic salmon (Salmo salar). Five weeks after a baseline welfare indicator scoring, non-anaesthetised Atlantic salmon ( w ¯ = 1379 ± 313 g, n = 332) were treated individually in a chamber with seawater at a temperature of 34 °C (warm water treatment) or 9 °C (control treatment) for 30 s. The treatment was repeated after 23–24 days. During the second treatment, a subset of the fish was video recorded for behavioural analysis. Seventeen to eighteen days after the second treatment, welfare indicators were again scored, and organ samples were taken for histopathological examination. The repeated warm water treatments resulted in a significantly increased prevalence and/or severity of scale losses, snout wounds, various eye problems, and active fin injuries as well as a significantly reduced specific growth rate. The fish displayed an immediate, strong behavioural reaction when exposed to warm water, which was probably the main cause of the detected injuries.

Sterile triploid Atlantic salmon (Salmo salar) show inconsistent seawater grow-out, but the reason why remains unclear. The purpose of this study was to determine the salinity optima of triploid post-smolts. Diploids and triploids were assessed for smoltification status during an underyearling smolt regime before being transferred to one of four different salinities, 0, 11, 23 and 35 ppt at 12 °C and under 24 h continuous light for 83 days. During this period, fish growth, plasma biochemistry, and production traits (vertebral deformities, ocular cataracts, sexual maturation) were monitored. Molecular biomarkers in the gill (nkaα1a, nkaα1b, nkcc1a) suggested triploids reached peak smolt earlier than diploids and began the desmoltification process before the start of the salinity treatments, however this was not reflected in gill Na+/K+-ATPase enzyme activity. At the initiation of the salinity treatments triploids were significantly larger than diploids (mean weight g ± SE: 71 ± 0.7 and 87.2 ± 0.8 for diploids and triploids, respectively) and there was a ploidy effect on post-smolt growth, with body weight showing a clearer positive trend with salinity in diploids (0 < 11 = 23 = 35 ppt) than in triploids (0 < 11 < 35 = 23 ppt) (final mean weight g ± SE: 255.2 ± 7.4, 303.9 ± 9, 313.9 ± 9 and 342.4 ± 12 for diploids and 322.9 ± 9.7, 361.7 ± 10.7, 425.9 ± 12.1, 415.2 ± 12.2 for triploids at 0, 11, 23, and 35 ppt, respectively). Plasma Na+ and Cl− increased, but plasma pH decreased, with increasing salinity in both ploidy. However, ploidy only had transient effects on plasma biochemistry depending on the salinity treatment. There was no ploidy effect on vertebral deformities (21% of both ploidy had one or more deformed vertebra). In contrast, triploids had a significantly higher prevalence of ocular cataracts (84 vs 98% in diploids and triploids, respectively) with a higher mean cataract score (mean ± SE: 1.93 ± 0.1 and 2.78 ± 0.1 for diploids and triploids, respectively), but a significantly lower prevalence of pubertal male post-smolts (15 vs 2% in diploids and triploids, respectively). Salinity treatment had no effect on vertebral deformities, cataracts, or post-smolt sexual maturation. In summary, there was a ploidy mismatch for smoltification biomarkers in the gill and salinity had a strong effect on post-smolt growth, but the effects were ploidy dependent. publishedVersion

Avoiding genetic interactions between wild and farmed Atlantic salmon is regarded as one of the major requirements for a sustainable salmon aquaculture industry. For this reason, farming functional sterile triploids has been suggested as a possible solution. However, knowledge about how triploids cope under commercial conditions is lacking. In the present study, we compared the performance of diploid and triploid Atlantic salmon among four Norwegian aquaculture companies. Diploid and triploid groups of the same genetic line were farmed in western, mid, and northern Norway under commercial conditions from seawater transfer until slaughter.Overall, triploid salmon exhibited reduced survival, higher incidence of emaciated fish, and scored, on average, a lower quality rating during primary processing. The results highlight the need for further research on how to improve the welfare and performance of triploid salmon in commercial aquaculture operations.

This chapter introduces principles for welfare assessment of farmed fish. The introduction details the developments in science and society regarding the proposition that fish have a welfare state and that welfare assessment is important. The next sections of the chapter highlight the neurological basis of fish welfare and outline the primary welfare needs of the fish; appropriate water environment, adequate nutrition, good health, the expression of (species-specific) behaviors and safety. The second half of the chapter discusses fish welfare assessment, including the selection of welfare indicators, which can be both input-based and outcome-based. It also discusses issues related to conducting on-farm welfare assessment and how to integrate welfare outcomes into an overall welfare assessment. The final section of the chapter looks at some future developments and directions related to the validation and standardization of welfare indicators, precision fish farming and new sensor-based technologies aimed for tracking and monitoring the welfare of individual fish.

Astaxanthin (Ax), the main carotenoid responsible for the distinct red flesh color in salmonids (Oncorhynchus, Salvelinus, Salmo, and Parahucho), is added to the diet of farmed fish at a substantial cost. Despite the great economical value for the salmon industry, the key molecular mechanisms involved in the regulation of muscle coloration are poorly understood. Chinook salmon (Oncorhynchus tshawytscha) represent an ideal model to study flesh coloration because they exhibit a distinct color polymorphism responsible for two color morphs, white and red flesh pigmented fish. This study was designed to identify the molecular basis for the development of red and white coloration of fish reared under the same experimental conditions and to better understand the absorption mechanism of Ax in salmonids. Pyloric caeca, liver, and muscle of both groups (n = 6 each) were selected as the most likely critical target organs to be involved respectively in the intestinal uptake, metabolism, and retention of Ax. Difference in the transcriptome profile of each tissue using next-generation sequencing technology was conducted. Ten KEGG pathways were significantly enriched for differentially expressed genes between red and white salmon pylorus tissue, while none for the transcriptome profile in the other two tissues. Differential expressed gene (DE) analyses showed that there were relatively few differences in muscle (31 DE genes, p

As a first attempt to assess bone health in cleaner fish production, wild and cultured ballan wrasse Labrus bergylta and lumpfish Cyclopterus lumpus were examined by radiology. In C. lumpus, wild fish (57%) had more vertebra deformities (≥1 deformed vertebrae) than cultured fish (2-16%). One wild C. lumpus had lordosis and another was missing the tail fin. In L. bergylta, wild fish (11%) had fewer vertebra deformities than cultured individuals (78-91%). Among the cultured L. bergylta, 17-53% of the fish had severe vertebra deformities (≥6 deformed vertebrae) with two predominate sites of location, one between vertebra 4 and 10 (S1) in the trunk, and one between 19 and 26 (S2) in the tail. Fusions dominated S1, while compressions dominated S2. Although wild L. bergylta had a low vertebra deformity level, 83% had calluses and 14% had fractures in haemal/neural spines and/or ribs. The site-specific appearance and pathology of fracture and callus in wild L. bergylta suggests these are induced by chronic mechanical stress, and a possible pathogenesis for fish hyperostosis is presented based on this notion. In conclusion, good bone health was documented in cultured C. lumpus, but cultured L. bergylta suffered poor bone health. How this affects survival, growth, swimming abilities and welfare in cultured wrasse should be further investigated. SIGNIFICANCE STATEMENT: Skeletal deformities were studied in ballan wrasse and lumpfish of both wild and cultured origin for the first time to identify potential welfare issues when deploying them as cleaner fish in salmon sea cages. While cultured lumpfish showed good bone health, cultured wrasse had a high occurrence of vertebra deformities, which is expected to impact lice eating efficiency and animal welfare negatively. These deformities are most likely induced early in development.

For a rapidly developing aquaculture industry where availability of raw materials for diet formulation are quickly changing it is fundamental to define what the dietary needs and benefits are. In farming of Atlantic salmon (Salmo salar), highly unsaturated fatty acids (HUFA) are known to be essential and a minimum dietary inclusion is needed. The aim of this study has been to map Atlantic salmon performance and midgut health towards dietary HUFAs under normal and challenging condition. Atlantic salmon post-smolt at 150 g were fed four experimental diets over a feeding period of 10 weeks up to a final weight of 377 g followed by three weeks of unpredictable chronic stress (UCS). The diets varied in their HUFA level, with either 28.7 g/kg DHA (HDHA), 25 g/kg EPA (HEPA), 21.3 g/kg ARA (HARA) or 8.5 g/kg EPA + DHA (LHUFA). After the feeding period all fish had more than doubled their weight and fish fed HARA showed a significant lower specific growth rate compared to all other diets. Diets had little effect on intestinal morphology, barrier and transport function. Intestinal polar lipid composition suggested higher lipid metabolism in fish feed LHUFA and transcriptome analysis confirm upregulated lipid metabolism and downregulated energy turn over. Comparing transcriptome profiles from HDHA to HEPA, more upregulated pathways are seen in fish fed HEPA, including metabolism and immunity. The fish feed HARA had downregulated turn-over and metabolism compared to fish fed HEPA. The UCS period resulted in zero growth and lower enterocyte height in all diet groups, and the intestinal barrier was reduced in the paracellular permeability to mannitol. Contradictory to previous studies, fish fed HDHA and HEPA responded to UCS with enhanced intestinal resistance. UCS also resulted in impaired intestinal transport by elevated ion flux and epithelial electrochemical gradient in fish fed LHUFA. Transcriptome pathway analysis indicated more cell turn over and lipid metabolism in LHUFA and HDHA. In conclusion, a low HUFA diet with 8.5 g/kg DHA + EPA can sustain growth and intestinal health under controlled condition, but the health is impaired after three weeks of chronic stress. Decreasing the n6/n3 ratio by increasing levels of EPA and DHA strengthens intestinal barrier in response to chronic stress. Increasing the ARA to HUFA ratio may have a negative effect on growth, but not on intestinal health.

Thermal delousing is a new method for removing sea lice from farmed Atlantic salmon (Salmo salar L). We investigated thermally-related tissue injuries in Atlantic salmon in a pilot laboratory trial to describe the acute effect of high water temperatures (34–38 °C). Acute tissue injuries in gills, eyes, brain and possible also nasal cavity and thymus were seen in salmon exposed to water temperatures of 34 - 38 °C in 72 to 140 s. This implies that exposing salmon to such water temperatures is a welfare risk, not only due to the direct tissue injuries that may also be dependent on exposure time, but also due to risk of thermal pain and aversion, including flight reactions. Key words: Fish, Delousing, Thermal pain, Tissue damage, Welfare

The in vivo functions of Atlantic salmon fatty acyl desaturases (fads2), Δ6fads2-a, Δ6fads2-b, Δ6fads2-c and Δ5fads2 in long chain polyunsaturated fatty acid (LC-PUFA) synthesis in salmon and fish in general remains to be elucidated. Here, we investigate in vivo functions and in vivo functional redundancy of salmon fads2 using two CRISPR-mediated partial knockout salmon, Δ6abc/5Mt with mutations in Δ6fads2-a, Δ6fads2-b, Δ6fads2-c and Δ5fads2, and Δ6bcMt with mutations in Δ6fads2-b and Δ6fads2-c. F0 fish displaying high degree of gene editing (50–100%) were fed low LC-PUFA and high LC-PUFA diets, the former containing reduced levels of eicosapentaenoic (20:5n-3) and docosahexaenoic (22:6n-3) acids but higher content of linoleic (18:2n-6) and alpha-linolenic (18:3n-3) acids, and the latter containing high levels of 20:5n-3 and 22:6n-3 but reduced compositions of 18:2n-6 and 18:3n-3. The Δ6abc/5Mt showed reduced 22:6n-3 levels and accumulated Δ6-desaturation substrates (18:2n-6, 18:3n-3) and Δ5-desaturation substrate (20:4n-3), demonstrating impaired 22:6n-3 synthesis compared to wildtypes (WT). Δ6bcMt showed no effect on Δ6-desaturation compared to WT, suggesting Δ6 Fads2-a as having the predominant Δ6-desaturation activity in salmon, at least in the tissues analyzed. Both Δ6abc/5Mt and Δ6bcMt demonstrated significant accumulation of Δ8-desaturation substrates (20:2n-6, 20:3n-3) when fed low LC-PUFA diet. Additionally, Δ6abc/5Mt demonstrated significant upregulation of the lipogenic transcription regulator, sterol regulatory element binding protein-1 (srebp-1) in liver and pyloric caeca under reduced dietary LC-PUFA. Our data suggest a combined effect of endogenous LC-PUFA synthesis and dietary LC-PUFA levels on srebp-1 expression which ultimately affects LC-PUFA synthesis in salmon. Our data also suggest Δ8-desaturation activities for salmon Δ6 Fads2 enzymes.

Cleaner fish used as a biological control agent against salmon lice is rapidly increasing in Atlantic salmon aquaculture. However, concerns have been raised about the welfare and mortality of cleaner fish in salmon cage systems, which could in turn affect their performance in controlling salmon lice. In a 4-month autumn-winter study, we monitored growth, welfare, mortality and daytime depth distribution of the most commonly used cleaner fish, farmed ballan wrasse and lumpfish, in six salmon production sea cages where thermo- and haloclines were present. Ballan wrasse did not grow (SGR: small: −0.01% day−1, large: −0.06% day−1), while lumpfish significantly doubled in size (SGR: 0.87% day−1) during the study. High losses (registered mortality + unregistered loss) were observed in both species (57 and 27% of ballan wrasse and lumpfish, respectively). The welfare status of remaining individuals generally improved over the study period, regardless of species. Brief daytime camera observations at hides found ballan wrasse were typically deeper at warmer (median 12.4 °C) more saline (median 31.7 ppt) depths, where salmon were expected to reside during day periods, compared to lumpfish generally occupying colder (median 7.3 °C), brackish (median 18.9 ppt) water in surface layers. Considerable mortalities, minimal feeding (inferred from ceased growth) by ballan wrasse and a possible mismatch in lumpfish and salmon depths (inferred from limited daytime camera observations) suggest that cleaner fish may have low long-term effectiveness against salmon lice in stratified salmon sea cages over autumn-winter. Similar studies across seasons, locations and cage types (e.g. depth-based cage technologies) are vital to understand the extent of these issues in salmon aquaculture more broadly. publishedVersion

When Atlantic salmon parr migrate from fresh water towards the sea, they undergo extensive morphological, neural, physiological and behavioural changes. Such changes have the potential to affect their responsiveness to various environmental factors that impose stress. In this study we compared the stress responses in parr and post-smolt salmon following exposure to repeated chasing stress (RCS) for three weeks. At the end of this period, all fish were challenged with a novel stressor and sampled before (T0) and after 1h (T1). Parr had a higher growth rate than post-smolts. Plasma cortisol declined in the RCS groups within the first week suggesting a rapid habituation/desensitisation of the endocrine stress axis. As a result of the desensitised HPI axis, RCS groups showed a reduced cortisol response when exposed to the novel stressor. In preoptic area (POA) crf mRNA levels were higher in all post-smolt groups compared to parr. 11βhsd2 decreased by RCS and by the novel stressor in post-smolt controls (T1), whereas no effect of either stress was seen in parr. The grs were low in all groups except for parr controls. In pituitary, parr controls had higher levels of crf1r mRNA than the other parr and post-smolt groups, whilst pomcb was higher in post-smolt control groups. Overall, 11βhsd2 transcript abundance in parr was lower than post-smolt groups; after the novel stressor pomcs, grs and mr were up-regulated in parr control (T1). In summary, we highlight differences in the central stress response between parr and post-smolt salmon following RCS.

Thermal treatment has become the most used delousing method in salmonid aquaculture. However, concerns have been raised about it being painful for the fish. We studied the behavioural response of Atlantic salmon acclimated to 8 °C when transferred to temperatures in the range 0–38 °C. Exposure time was 5 min or until they reached the endpoint of losing equilibrium and laying on their side, a sign of imminent death. At temperatures below 28 °C, none of the fish reached endpoint within the 5-min maximum. At 28 °C four of five fish reached endpoint, and fish reached endpoint more rapidly as temperature increased further. Fish transferred to temperatures above 28 °C had higher swimming speed immediately after transfer and maintained a high swimming speed until just before loss of equilibrium. Their behaviour was from the start characterised by collisions into tank walls and head shaking. Just before loss of equilibrium they started breaking the surface of the water, swimming in a circle pattern and in some instances displayed a side-wise bending of their body. In other words, salmon transferred to temperatures above 28 °C showed instant behavioural responses indicative of nociception or pain. Keywords: Behavioural response, Fish, Heated water, Nociception, Pain, Thermal delousing

A common-garden experiment was carried out to compare two genetically distinct strains of Atlantic salmon (Salmo salar) fed diets with either high (CHO) or low (NoCHO) digestible carbohydrate (starch). Twenty salmon from either a commercial farmed strain (F) or a land-locked population (G) were placed in two tanks (10 fish of each population in each tank) and fed either CHO or NoCHO feeds. At the end of the experiment fish were fasted for 8 h, euthanized and blood and liver collected. Both diet and population had an effect on circulating glucose levels with G showing hypoglycaemia and dietary starch increasing this parameter. In contrast, G showed increased plasma triacylglycerol levels regardless of dietary treatment suggesting faster conversion of glucose to triacylglycerol. This different ability to metabolize dietary starch among strains was also reflected at a molecular (gene) level as most of the metabolic pathways evaluated were mainly affected by the factor population rather than by diet. The data are promising and suggest different regulatory capacities toward starch utilization between land-locked salmon and the farmed stock. Further analyses are necessary in order to fully characterize the capacity of land-locked salmon to utilize dietary carbohydrate. © 2018 Betancor, Olsen, Marandel, Skulstad, Madaro, Tocher and Panserat. This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

Individual variation in behavior and physiological traits in a wide variety of animals has been the focus of numerous studies in recent years. In this context, early life experiences shape responses that individuals have to subsequent environments, i.e. developmental plasticity. In this experiment, we subjected 10-month old fish to an unpredictable chronic stress (UCS) regime or no stress (control) for 3weeks. These individuals then underwent the parr-smolt transformation, when salmonids become adapted for the seawater environment, and were subsequently transferred into seawater before the final sampling. Biometric data was collected at the end of each period. Sampling on the final day was conducted in order to analyze basal monoaminergic activity in the brain stem and hypothalamus, as well as gene expression of target genes in the telencephalon. We found that post-hoc sorting of individuals by their serotonergic activity (high and low) resulted in the elucidation of growth and gene expression differences. UCS groups were found to have less growth disparities throughout the experiment, compared to control fish. Furthermore, we found brain serotonergic signaling and corticotropic releasing factor binding protein expression were positively associated with brain stem serotonergic activity, which is consistent with fish showing a stress reactivity neurophysiological profile. In conclusion, we here submit evidence that sorting individuals by their basal serotonergic activity levels may be a useful tool in the study of developmental plasticity. These results may thus apply directly to improving husbandry practices in aquaculture and elucidating neural mechanisms for coping behavior.

Ongoing rapid domestication of Atlantic salmon implies that individuals are subjected to evolutionarily novel stressors encountered under conditions of artificial rearing, requiring new levels and directions of flexibility in physiological and behavioural coping mechanisms. Phenotypic plasticity to environmental changes is particularly evident at early life stages. We investigated the performance of salmon, previously subjected to an unpredictable chronic stress (UCS) treatment at an early age (10 month old parr), over several months and life stages. The UCS fish showed overall higher specific growth rates compared with unstressed controls after smoltification, a particularly challenging life stage, and after seawater transfer. Furthermore, subjecting fish to acute stress at the end of the experiment, we found that UCS groups had an overall lower hypothalamic catecholaminergic and brain stem serotonergic response to stress compared with control groups. In addition, serotonergic activity was negatively correlated with final growth rates, which implies that serotonin responsive individuals have growth disadvantages. Altogether, our results may imply that a subdued monoaminergic response in stressful farming environments may be beneficial, because in such situations individuals may be able to reallocate energy from stress responses into other life processes, such as growth. http://creativecommons.org/licenses/by/4.0/ Published by the Royal Society under the terms of the Creative Commons Attribution License http://creativecommons.org/licenses/by/4.0/, which permits unrestricted use, provided the original author and source are credited.

The possibility to prepare for and respond to challenges in a proper manner is essential to cope with a changing environment, and learning allows fish to up or downregulate the stress response based on experience. The regulation of the response to predicted needs should be easier in more predictable environments. We exposed salmon parr to chasing of either 15 s (weak stressor) or 5 min (strong stressor) twice daily for a 7-day learning period, with chasing either announced by a 30 s light signal (conditioned) or not announced (unconditioned). The behavioural response to the light signal was different between the conditioned and unconditioned groups, demonstrating that conditioned groups associated the signal with chasing. We could, however, not demonstrate any effect on the stress response of anticipation. The fish habituated to repeated stress exposures with a similar decrease in oxygen hyperconsumption in all groups. Due to habituation, possible effects of predictable announcement of a stressor on the physiological stress response may not have been expressed in this study. Plasma cortisol concentrations 1h after light signal and chasing the day after the training period was moderate in all groups although higher after 5 min chasing (13 ng ml(-1)) than 15 s chasing (7 ng ml(-1)). There was no physiological stress response after exposure to the light signal only after the learning period. We argue that the benefit of predictability of stressors is limited when the fish have no way to avoid the stressor.

Contains fulltext : 149118.pdf (Publisher’s version ) (Open Access) Combinations of stressors occur regularly throughout an animal's life, especially in agriculture and aquaculture settings. If an animal fails to acclimate to these stressors, stress becomes chronic, and a condition of allostatic overload arises with negative results for animal welfare. In the current study, we describe effects of exposing Atlantic salmon parr to an unpredictable chronic stressor (UCS) paradigm for 3 weeks. The paradigm involves exposure of fish to seven unpredictable stressors three times a day. At the end of the trial, experimental and control fish were challenged with yet another novel stressor and sampled before and 1 h after that challenge. Plasma cortisol decreased steadily over time in stressed fish, indicative of exhaustion of the endocrine stress axis. This was confirmed by a lower cortisol response to the novel stressor at the end of the stress period in chronically stressed fish compared with the control group. In the preoptic area (POA) and pituitary gland, chronic stress resulted in decreased gene expression of 11βhsd2, gr1 and gr2 in the POA and increased expression of those genes in the pituitary gland. POA crf expression and pituitary expression of pomcs and mr increased, whereas interrenal gene expression was unaffected. Exposure to the novel stressor had no effect on POA and interrenal gene expression. In the pituitary, crfr1, pomcs, 11βhsd2, grs and mr were down-regulated. In summary, our results provide a novel overview of the dynamic changes that occur at every level of the hypothalamic-pituitary gland–interrenal gland (HPI) axis as a result of chronic stress in Atlantic salmon.