The Interaction between Water Currents and Salmon Swimming Behaviour in Sea Cages

Positioning of sea cages at sites with high water current velocities expose the fish to a largely unknown environmental challenge. In this study we observed the swimming behaviour of Atlantic salmon (Salmo salar L.) at a commercial farm with tidal currents altering between low, moderate and high velocities. At high current velocities the salmon switched from the traditional circular polarized group structure, seen at low and moderate current velocities, to a group structure where all fish kept stations at fixed positions swimming against the current. This type of group behaviour has not been described in sea cages previously. The structural changes could be explained by a preferred swimming speed of salmon spatially restricted in a cage in combination with a behavioural plasticity of the fish.     

Evaluating the effect of clustering when monitoring the abundance of sea lice populations on farmed Atlantic salmon

Using cluster random sampling theory and empirical estimates of the intra‐class correlations for sea lice Lepeophtheirus salmonis abundances, methods on how best to sample Atlantic salmon Salmo salar from cages on farms were derived. Estimates of intra‐class correlations for the abundance of the chalimus and mobile sea lice stages on Atlantic salmon in Scottish farms are given. These correlations were higher for mobile stages than for chalimus, and they had a substantive effect on increasing the number of cages and fish to be sampled for all sea lice stages. An important finding is that it is better to have a procedure that randomly samples a large number of cages using a small number of fish from each. This finding and the cluster random sampling approach have relevance to the monitoring of all marine species being farmed in cages or tanks.     

Serological evidence of infectious salmon anaemia virus (ISAV) infection in farmed fishes,using an indirect enzyme-linked immunosorbent assay (ELISA)

Antibody detection tests are rarely used for diagnostic purposes in fish diseases. Infectious salmon anaemia (ISA) caused by ISA virus (ISAV) is an emerging disease of Atlantic salmon Salmo salar L. The virus has also been isolated from diseased coho salmon Oncorhynchus kisutch in Chile. An indirect enzyme-linked immunosorbent assay (ELISA) that should facilitate serodiagnosis of ISAV infection, the study of epidemiology, and the control of ISA in farmed fishes has been developed using purified ISAV as the coating antigen, and monoclonal antibodies that detect fish immunoglobulins bound to the antigen on the plate. Application of the test to a random sample of farmed Atlantic salmon from the Bay of Fundy, New Brunswick, Canada, positively identified 5 of the 7 ISAV RT-PCR-positive fish, and all 10 RT-PCR-negative fish were also negative in the ELISA. Some RT-PCR-negative fish had an elevated non-specific antibody reactivity suggestive of chronic infection or resistance to ISAV. This test was also able to detect 11 of the 14 coho salmon pooled serum samples from a clinically affected farm in Chile that were positive by the virus neutralization (VN) test, and 2 of the 4 VN-negative samples. We conclude that this ELISA would be suitable as a routine test for ISAV infection or for assessing ISAV vaccine efficacy before placing smolts in sea cages, and for testing fishes in sea cages to detect level of resistance to ISA. The assay enables vaccination in combination with depopulation control methods.     

Influence of aperiodic summer droughts on leaf litter breakdown and macroinvertebrate assemblages: testing the <Emphasis Type="Italic">drying memory</Emphasis> in a Central Apennines River (Aterno River,Italy)

Management of multiple exploited stocks of anadromous salmonids in large catchments requires understanding of movement and catchment use by the migrating fish and of their harvesting. The spawning migration of sea trout (Salmo trutta) and Atlantic salmon (Salmo salar) was studied in the River Tweed, UK, using acoustic telemetry to complement exploitation rate data and to quantify catchment penetration. Salmon (n = 79) and sea trout (n = 65) were tagged in the tidal-influenced Tweed in summer–autumn. No tagged salmon left the river before spawning, but 3% (2010) and 8% (2011) of pre-spawning sea trout dropped out. Combined tag regurgitation/fish mortality in salmon was 12.5%, while trout mortality was 6% (2010) and 0% (2011). The estimated spawning positions of salmon and sea trout differed; tagged salmon were mostly in the main channel while trout occurred mostly in the upper Tweed and tributaries. Early fish migrated upstream slower than later fish, but sea trout moved through the lower-middle river more quickly than salmon, partly supporting the hypothesis that the lower exploitation rate in autumn of trout (1 vs 3.3% for salmon) there is generated by differences in migration behaviour.     

Genome‐wide SNP analysis reveals a genetic basis for sea‐age variation in a wild population of Atlantic salmon (Salmo salar)

Delaying sexual maturation can lead to larger body size and higher reproductive success, but carries an increased risk of death before reproducing. Classical life history theory predicts that trade‐offs between reproductive success and survival should lead to the evolution of an optimal strategy in a given population. However, variation in mating strategies generally persists, and in general, there remains a poor understanding of genetic and physiological mechanisms underlying this variation. One extreme case of this is in the Atlantic salmon (Salmo salar), which can show variation in the age at which they return from their marine migration to spawn (i.e. their ‘sea age’). This results in large size differences between strategies, with direct implications for individual fitness. Here, we used an Illumina Infinium SNP array to identify regions of the genome associated with variation in sea age in a large population of Atlantic salmon in Northern Europe, implementing individual‐based genome‐wide association studies (GWAS) and population‐based F_ST outlier analyses. We identified several regions of the genome which vary in association with phenotype and/or selection between sea ages, with nearby genes having functions related to muscle development, metabolism, immune response and mate choice. In addition, we found that individuals of different sea ages belong to different, yet sympatric populations in this system, indicating that reproductive isolation may be driven by divergence between stable strategies. Overall, this study demonstrates how genome‐wide methodologies can be integrated with samples collected from wild, structured populations to understand their ecology and evolution in a natural context.     

The formation of non-anadromous populations of Atlantic salmon, Salmon salar L., in Europe

Comparison of the different populations of non-anadromous Atlantic salmon in Europe has revealed some common traits. The non-anadromous populations probably became isolated from the anadromous salmon populations during a period of rapid land-upheaval after the last ice age, about 10 000 years ago. The oceanic temperature in the eastern part of the Atlantic was especially low in this period. This probably diminished the advantage gained from the anadromous habit. During this period the low temperatures in the rivers probably increased the frequency of females which matured without migrating seawards. A large number of individuals in the different populations seems to have been favourable for the formation and survival of such races. The occurrence of a lake in such a river system would probably ease the transition to non-anadromous behaviour, because the salmon would not then be forced to adjust its entire life-history when isolated (the salmon could spawn in the river in the usual manner and the smolts then migrate downstream into lakes). Larger lakes would also seem to infer problems for the navigation of the smolts towards the outlet. The only known exception is the river-living land-locked salmon population found in the River Namsen, Norway.
The factors mentioned above have led to an increase in the probability of occurrence and thriving of mature females without a sea migration or to a decrease in the competitive advantage of anadromous behaviour. The above mentioned similarities can explain the seemingly casual distribution of the non-anadromous populations of Atlantic salmon in Europe.     

A note on pre-spawning maternal cortisol exposure in farmed Atlantic salmon and its impact on the behaviour of offspring in response to a novel environment

This study was designed to assess whether pre-spawning maternal cortisol administration and an additional stressor, here modelled by mild hyperthermia experienced during incubation, affected the behavioural responses of farmed Atlantic salmon (Salmo salar) offspring when exposed to a novel environment. Mature females were injected prior to stripping with 0, 50 or 100 mg/kg cortisol. After fertilization the eggs were kept at incubation temperatures of either 8 or 10 °C. Four months after hatching, the fish were introduced to a novel environment, i.e. an aquarium (38 l). In each trial two individuals originating from different exposure groups were video recorded and their activity (number of turns, time spent not swimming, time spent at the bottom) and feeding behaviour were measured. Maternal cortisol increased time spent non-swimming and also time spent at the bottom, while cortisol and temperature together additionally affected number of turns. To the best of our knowledge this study is the first in fish to demonstrate a possible link between the maternal endocrinological state at spawning and the offspring's behavioural reactivity to novel situations.     

Factors influencing return rate and marine residence duration in sea trout populations in Central Norway

Brown trout (Salmo trutta) display extensive plasticity in marine migratory behaviours, with marine migrations considered to be an adaptive strategy which enables sea trout to maximize growth and reproductive potential. However, marine migrations are not without associated costs, including threats posed by ever-increasing salmon lice (Lepeophtheirus salmonis) infestations. In the present study, we used passive integrated transponder technology to characterize variability in sea trout migration behaviour amongst three catchments situated in a region of intensive salmon farming in central Norway. Specifically, we investigate how lice infestation, out-migration date and body size alter sea trout return rate and marine residence duration during the first out-migration to sea from each catchment. Distinct catchment-specific differences in sea trout out-migration size and the number of cohorts were observed, but larger body size did not guarantee the successful return of migrating trout. The marine residence duration of individuals that successfully returned to freshwater was positively correlated with lice infestation risk, suggesting for these individuals the lethal infestation threshold had not been reached. Our results also suggest that sea trout populations from lotic-dominated catchments are potentially at greater risk from size-related threats to their survival encountered during their marine migrations than sea trout from lentic-dominated catchments. The variability in sea trout migratory behaviour amongst catchments observed here emphasizes the challenges fisheries managers face when deciding the best actions to take to protect the anadromous portion of brown trout populations.     

Response of downstream migrant juvenile Pacific salmonids to accelerating flow and overhead cover

Mechanical structures designed to divert or guide juvenile migrant salmon at dams often work less effectively than expected. This likely results from a lack of understanding of fish behaviour. In order to address this, a series of experiments was conducted at McNary dam, USA, to assess the influence of hydraulic transition and overhead cover on the behaviour of seaward migrating juvenile Pacific salmon. Fish passing through a flume encountered a choice of route that varied based on hydraulic factors and overhead cover. Direct observation revealed that individuals elicited strong avoidance behaviour when they encountered areas where hydraulic conditions changed rapidly or were covered. Our findings have implications relevant to fish pass design and culvert restoration in Europe. Critically, improvement of current fish pass design and development of alternative mechanisms that effectively block fish access to, and divert fish away from poor passage routes, requires consideration of the behavioural component of fish migration. Guest editors: R. L. Welcomme & G. Marmulla Hydropower, Flood Control and Water Abstraction: Implications for Fish and Fisheries     

Enhancing data mobilisation through a centralised data repository for Atlantic salmon (Salmo salar L.): Providing the resources to promote an ecosystem-based management framework.

Data and knowledge mobilisation are significant challenges in ecology and resource management, with the journey from data collection through to management action often left incomplete due to difficulties sharing information across diverse and dispersed communities. This disconnect between science and management must be resolved if we are to successfully tackle the increasing impact of human activity on our ecosystems. Across their North Atlantic range, Atlantic salmon (Salmo salar L.) populations are in steep decline in many areas and urgent actions are required to curb this decline. Being commercially important this species has been subject to intense research, but management action often suffers from both a lack of access to this knowledge resource and support for its integration into effective management strategies. To respond to this challenge, the science and management communities must place higher priority on mobilising existing and emerging knowledge sources to inform current and future resource use and mitigation strategies. This approach requires a more complete picture of the current salmon ecology data and knowledge landscape, new mechanisms to enable data mobilisation and re-use, and new research to describe and parameterise the responses of wild populations to habitat changes. Here we present a unique interface for registering and linking data resources relevant to the Atlantic salmon life cycle that can address the data mobilisation aspect of these challenges. The Salmon Ecosystem Data Hub is a salmon-specific metadata catalogue, natively interoperable with many existing data portals, which creates a low resistance pathway to maximise visibility of data relevant to Atlantic salmon. This includes the capacity to annotate datasets with life-stage domains and variable classes, thereby permitting dispersed data to be formally contextualised and integrated to support hypotheses specific to scenario-based modelling and decision-making. The alignment and mobilisation of data within the Salmon Ecosystem Data Hub will help advance the development of appropriate environmentally driven forecast models and an ecosystem-based management approach for Atlantic salmon that optimises future management strategies.     

Evolutionary mechanisms in behaviour: An intraspecific genetic approach

The study of the evolutionary mechanisms in behaviour and of the biological grounds of individual variability may be based on the comparative phylogenetic approach or on an intraspecific genetic analysis. The discontinuous behavioural progression evident across species is due to the assumption that the phylogenetic “scale” may be adopted in place of the phylogenetic trees and to the fact that today's existing species have evolved in parallel and may not be used to represent an evolutionary sequence. However, despite the existence of different motor and perceptual abilities the comparative approach has shown that many analogies exist between some basic brain mechanisms across species.The existence of outstanding individual differences within the same species must be regarded as a powerful potential tool since quantitative differences between the behaviours of different individuals belonging to the same species might later result in qualitatively different phenotypes. By using different strains and mutations of mice and different genetic approaches it has been possible to assess the mode of inheritance, to calculate estimates of heritability and of the number of segregating units for different behavioural traits ranging from avoidance to maze learning and activity. The existence of clear genetic correlations between some of these behavioural patterns has also allowed the identification of some single-gene mutants affecting these traits and the characterization of a gene responsible for a major effect on activity. This psychogenetic approach shows that there are behaviour differences which are the products of evolutionary adaptive processes and that the knowledge of the genetic systems that underlie these differences is a basic step for understanding the brain mechanisms in Man.     

Fatty acid compositions of gonadal material and diets of the sea urchin, Psammechinus miliaris: trophic and nutritional implications

The fatty acid compositions of gonadal material was examined for the sea urchin Psammechinus miliaris (Gmelin) held in aquaria and fed either salmon feed pellets or the macroalga, Laminaria saccharina for 18 months. Gonadal material was also examined from P. miliaris collected from four field sites, including commercial scallop lines encrusted with the mussel, Mytilus edulis, sea cages stocked with Atlantic salmon Salmo salar and two intertidal sea-loch sites, characterised by either a fine mud or a macroalgal substratum. The fatty acid compositions of known and potential dietary material was examined. The proportions of certain fatty acids in the gonads of P. miliaris were significantly affected by diet type and location. Docosahexaenoic acid (DHA) 22:6 n-3 was significantly higher in the gonads of the sea urchins fed salmon feed in aquaria and collected from the salmon cages and scallop lines than in the gonads of the sea urchins fed L. saccharina in aquaria and collected from the intertidal sea loch sites. The salmon feed and the mussel tissue also contained a high proportion of this fatty acid. Stearidonic acid 18:4 n-3 and arachidonic acid 20:4 n-6, however, were found in significantly higher proportions than DHA in the gonads of the sea urchins fed L. saccharina and collected from the two intertidal sea-loch sites. L. saccharina was also found to contain high proportions of stearidonic and arachidonic acid. The gonads of the sea urchins collected from the intertidal site, characterised by a mud substratum, and from the scallop lines were found to contain a lower 18:1 n-9/18:1 n-7 ratio and a higher proportion of branched and odd-chained fatty acids, signifying a high dietary bacterial input, than the sea urchins held in the aquaria and collected from the salmon cage. 20:2 and 22:2 non-methylene-interrupted dienoic fatty acids (NMIDs) were found in P. miliaris fed diets lacking these fatty acids suggesting de novo biosynthesis. These results, therefore, suggest that the proportions/ratios of certain fatty acids in the gonads of P. miliaris could be used to give an indication of the predominant diet type of this species in the wild.     

Population and size-specific distribution of Atlantic salmon Salmo salar in the Baltic Sea over five decades

Population-specific assessment and management of anadromous fish at sea requires detailed information about the distribution at sea over ontogeny for each population. However, despite a long history of mixed-stock sea fisheries on Atlantic salmon, Salmo salar, migration studies showing that some salmon populations feed in different regions of the Baltic Sea and variation in dynamics occurs among populations feeding in the Baltic Sea, such information is often lacking. Also, current assessment of Baltic salmon assumes equal distribution at sea and therefore equal responses to changes in off-shore sea fisheries. Here, we test for differences in distribution at sea among and within ten Atlantic salmon Salmo salar populations originating from ten river-specific hatcheries along the Swedish Baltic Sea coast, using individual data from >125,000 tagged salmon, recaptured over five decades. We show strong population and size-specific differences in distribution at sea, varying between year classes and between individuals within year classes. This suggests that Atlantic salmon in the Baltic Sea experience great variation in environmental conditions and exploitation rates over ontogeny depending on origin and that current assessment assumptions about equal exploitation rates in the offshore fisheries and a shared environment at sea are not valid. Thus, our results provide additional arguments and necessary information for implementing population-specific management of salmon, also when targeting life stages at sea.     

Randomized clinical trial to investigate the effectiveness of teflubenzuron for treating sea lice on Atlantic salmon

A double-blind, randomized control clinical trial was performed to investigate the effectiveness of teflubenzuron in controlling sea lice Lepeophtheirus salmonis on farmed Atlantic salmon Salmo salar. A total of 40 sea cages from 3 commercial cage sites in Atlantic Canada were used in this Good Clinical Practice (GCP) trial. The teflubenzuron was administered in the feed at a dosage of 10 mg kg(-1) biomass d(-1) for 7 d. Medicated and control cages were matched by site, cage size, and pre-treatment mean lice counts using cages as the unit of concern. Post-treatment lice counts and staging of developmental stages were performed at 1 and 2 wk after the end of treatment. Chalimus stages in medicated cages were significantly lower than in control cages at 1 wk (79% reduction in mean lice counts, p < 0.001), and at 2 wk (53% reduction, p < 0.001). Mobile (pre-adult and adult) stages were also significantly reduced in medicated cages at 1 wk (69% reduction, p < 0.01), and at 2 wk (40% reduction, p < 0.01) post-treatment, respectively. Teflubenzuron was proven effective for reducing lice burdens on salmon despite the low parasite levels experienced during the trial and the recruitment of lice from the untreated cages. The use of cage as the unit of concern was an important design component of this trial.     

Population dynamics and harvesting of semelparous species with phenotypic and genotypic variability in reproductive age

We study the evolution of polymorphic life histories in anadromous semelparous salmon and the effects of harvesting. We derive dynamic phenotypic and genetic ESS models for describing the evolutionary dynamics. We show in our deterministic analysis that polymorphisms are not possible in a panmictic random mating population. Instead, genetic or behavioral polymorphisms may be observed in populations with assortative mating systems. Positive assortative mating may be supported and generated by behavioral and phenotypic traits like male mate choice, spawning ground selection by phenotype, or within-river homing-migration-distance by size. In the case of an evolutionarily stable dimorphism, the ESS is characterized by a reproductive ideal free distribution such that at an equilibrium the individuals are indifferent from the fitness point of view between the two life histories of early and late reproduction. Different strategy models - that is, phenotypic and genetic ESS models - yield identical behavioral predictions and, consequently, genetics does not seem to play an important role in the present model. An evolutionary response to increased fishing mortality is obvious and may have resource management implications. High sea fishing mortalities drive the populations toward early spawning. Thus it is possible that unselective harvesting at sea may eliminate, depending on the biological system, behavioral polymorphisms or genetic heterozygozity and drive the population to a monomorphic one. If within-river homing migration distances depend on the size of fish, unselective harvesting at sea, or selective harvesting of spawning runs in rivers, may reduce local population sizes on spawning grounds high up rivers. Finally, harvesting in a population may cause a switch in a dominant life-history strategy in a population so that anticipated sustainable yields cannot be realized in practice.     

The effect of salmon farming on the benthos of a Scottish sea loch

The effects of waste from a salmon farm on the benthos of a fjordic sea loch on the western coast of Scotland have been studied. Within 3 m of the floating cages the sediment was highly reducing, and dissolved oxygen content of the water overlying the sediment ranged from 35 to 75% saturation. At ⩾ 15 m from the cages the sediment was oxygenated, and the dissolved oxygen content of the water overlying the sediment was 50–85% saturation. Sedimentary redox potential and dissolved oxygen content of bottom water showed a seasonal variation. The benthic fauna showed marked changes in species number, species diversity, faunal abundance, and biomass in the region of the fish farm, with four zones of effect identified. Directly beneath, and up to the edge of the cages, there was an azoic zone. A highly enriched zone, dominated byCapitella capitata (Fabricius) andScolelepis fuliginosa (Claparède), occurred from the edge of the cages out to as 8 m. A slightly enriched ″transitional ″ zone occurred at ⩽ 25 m, and a “clean” zone at distances > 25 m. This study showed that salmon farming had similar effects on the benthos as other forms of organic enrichment, but the effects were limited to a small area in the immediate vicinity of the cages     

Intestinal barrier function of Atlantic salmon (<Emphasis Type="Italic">Salmo salar</Emphasis> L.) post smolts is reduced by common sea cage environments and suggested as a possible physiological welfare indicator

Background  

Fish farmed under high intensity aquaculture conditions are subjected to unnatural environments that may cause stress. Therefore awareness of how to maintain good health and welfare of farmed fish is important. For Atlantic salmon held in sea cages, water flow, dissolved oxygen (DO) levels and temperature will fluctuate over time and the fish can at times be exposed to detrimentally low DO levels and high temperatures. This experimental study investigates primary and secondary stress responses of Atlantic salmon post smolts to long-term exposure to reduced and fluctuating DO levels and high water temperatures, mimicking situations in the sea cages. Plasma cortisol levels and cortisol release to the water were assessed as indicators of the primary stress response and intestinal barrier integrity and physiological functions as indicators of secondary responses to changes in environmental conditions.     

Quantification of piscine reovirus (PRV) at different stages of Atlantic salmon Salmo salar production

The newly described piscine reovirus (PRV) appears to be associated with the development of heart and skeletal muscle inflammation (HSMI) in farmed Atlantic salmon Salmo salar L. PRV seems to be ubiquitous among fish in Norwegian salmon farms, but high viral loads and tissue distribution support a causal relationship between virus and disease. In order to improve understanding of the distribution of PRV in the salmon production line, we quantified PRV by using real-time PCR on heart samples collected at different points in the life cycle from pre-smolts to fish ready for slaughter. PRV positive pre-smolts were found in about 36% of the freshwater cohorts and a general increase in viral load was observed after their transfer to seawater. A reduction in viral loads was recorded when fish approached slaughter (18 mo in sea cages). Sequencing of positive samples did not support the hypothesis that outbreaks are caused by the spreading of a particular (virulent) strain of PRV.     

Relevance of incubation temperature for Vibrio salmonicida vaccine production

AIMS: To investigate the relationships between water temperature, bacterial growth, virulence and antigen expression in Vibrio salmonicida, the causal agent of cold water vibriosis in Atlantic salmon (Salmo salar L.). METHODS AND RESULTS: The significance of sea temperature was investigated using historical clinical and oceanographic data. An upper threshold for disease of approx. 10 degrees C was established. The effects of culture temperature and media type on bacterial growth were studied on solid and in liquid media. The highest rates of cell division were identified at 15 degrees C on solid media and 10 degrees C in liquid media. Outer membrane protein (OMP) expression and serological response in Atlantic salmon were studied using sodium dodecyl sulphate-polyacrylamide gel electrophoresis, Western blotting and enzyme-linked immunosorbent assay. A novel 76-kDa OMP produced in unshaken cultures at 10 degrees C was not found to stimulate a specific humoral response. CONCLUSIONS: Diagnostic agar plate-based incubation of suspected V. salmonicida should be carried out at 15 degrees C. High yield broth cultures for vaccine production should be incubated at 10 degrees C or lower. SIGNIFICANCE AND IMPACT OF THE STUDY: This study is, to the best of our knowledge, the first to identify different optimal temperatures in a bacterial species cultured on physically different types of media. The evidence presented suggests that V. salmonicida and possibly other bacteria destined for vaccine use in poikilothermic organisms should be cultured at temperatures consistent with that at which disease occurs.