Laboratory and field investigations of salmon lice [Lepeophtheirus salmonis (Krøyer)] infestation on Atlantic salmon (Salmo salar L.) post-smolts

Hatchery‐reared 1‐year‐old Atlantic salmon post‐smolts (Salmo salar L.), artificially infected with salmon lice [Lepeophtheirus salmonis (Krøyer)] copepodids, were found to suffer from primary alterations (increased cortisol levels) at early lice stages. Secondary alterations, such as osmotic stress (increased chloride levels), first occurred after the preadult stages of the lice appeared. Fish with the highest salmon lice infections died throughout the experiment. Seven years of field investigation of Trondheimsfjorden showed that Atlantic salmon post‐smolts descending coastal waters can become heavily infected with salmon lice. The migrating post‐smolts were only infected with the chalimus stages, showing that the fish had only recently left the rivers. The infection level, however, varied considerably between the years, and, in 1998, the infection was higher than previous years. The experimental results have been combined with the field data to appraise the consequences of the infection.     

Reducing sea lice (Lepeophtheirus salmonis) infestation of farmed Atlantic salmon (Salmo salar L.) through functional feeds

Health diets for Atlantic salmon have become an important component of the integrated pest management strategies targeting sea lice. A challenge trial was performed to examine the effect of supplementing salmon diets with either immunostimulants or essential oils. One control and four experimental diets containing immunostimulants or natural identical extracts were fed to Atlantic salmon in triplicate tanks for 4 weeks before challenging the fish with the sea lice copepodids. Prevalence of infection was 100%, and the mean abundance of infection was 21.2. The lowest mean lice count of 17 per fish (P < 0.05) was found in the group fed a mix of natural identical plant extracts (PX I). This represents a 20% reduction in infection, showing the potential for health diets to be employed as a tool to help control sea lice. To gain an understanding of the mechanisms of action underlying this protection, fish fed the control diet and fish fed the PX I diet were compared using quantitative histology of the epidermis and proteomic analysis of epidermal mucus. No significant differences were seen in the thickness of the epidermis or mucous cell percentage area, but differences in expression were seen for a number of proteins, including heat shock proteins, in epidermal mucus.     

Susceptibility,behaviour, and retention of the parasitic salmon louse (Lepeophtheirus salmonis) differ with Atlantic salmon population origin

Atlantic salmon populations across the world have diverse ecological and evolutionary histories, from wild anadromous or landlocked, to domestication and genetic modification. The natural host behaviours confer protection from infestation by ectoparasitic salmon lice Lepeophtheirus salmonis, yet whether genetic origin results in different behaviours and thus susceptibility to infestation is unknown. In common garden experiments, we tested antiparasite behaviours, susceptibility and retention of salmon lice in wild anadromous, wild landlocked, domesticated and genetically modified domesticated strains. Within domesticated strains, we tested two infestation histories (previously infested and naïve) and a new phenotype (albino colouring). Farmed stocks initially acquired 24%–44% higher levels of parasite density than the wild and landlocked strains. Burst swimming and displacement behaviours were higher in the domesticated groups, and jumping was more prevalent in the domesticated strains. At 34 days post‐infestation, domesticated strains and the wild anadromous strain did not differ significantly from each other; however, landlocked salmon had increased infestation levels considerably. Domesticated strains lost ~20% (±9.9%–16.5%; 95% CI) of their initial parasite load, while parasite load increased by 5.5% (±30.1%) for wild salmon and 20.1% (±28.5%) in landlocked salmon. This study provides early evidence for diverged host–parasite interactions associated with domestication in this system.     

Synergistic osmoregulatory dysfunction during salmon lice (Lepeophtheirus salmonis) and infectious hematopoietic necrosis virus co‐infection in sockeye salmon (Oncorhynchus nerka) smolts

While co‐infections are common in both wild and cultured fish, knowledge of the interactive effects of multiple pathogens on host physiology, gene expression and immune response is limited. To evaluate the impact of co‐infection on host survival, physiology and gene expression, sockeye salmon Oncorhynchus nerka smolts were infected with the salmon louse Lepeophtheirus salmonis (V?/SL+), infectious hematopoietic necrosis virus (IHNV; V+/SL?), both (V+/SL+), or neither (V?/SL?). Survival in the V+/SL+ group was significantly lower than the V?/SL? and V?/SL+ groups (p = 0.024). Co‐infected salmon had elevated osmoregulatory indicators and lowered haematocrit values as compared to the uninfected control. Expression of 12 genes associated with the host immune response was analysed in anterior kidney and skin. The only evidence of L. salmonis‐induced modulation of the host antiviral response was down‐regulation of mhc I although the possibility of modulation cannot be ruled out for mx‐1 and rsad2. Co‐infection did not influence the expression of genes associated with the host response to L. salmonis. Therefore, we conclude that the reduced survival in co‐infected sockeye salmon resulted from the osmoregulatory consequences of the sea lice infections which were amplified due to infection with IHNV.     

A review of the use of ivermectin as a treatment for sea lice [Lepeophtheirus salmonis (Krøyer) and Caligus elongatus Nordmann] infestation in farmed Atlantic salmon (Salmo salar L.)

Ectoparasitic sea lice, Lepeophtheirus salmonis (Krøyer) and Caligus elongatus (Nordmann) browse on the skin of farmed Atlantic salmon, Salmo salar L.. The resulting lesions cause stress and increase the susceptibility of the fish to secondary infections. In extreme infestations, fish can suffer from osmoregulatory failure and death. The most immediate treatment for the relief of sea lice infestations at fish farms is the use of chemotherapeutants, either by bath or oral administration. One compound which has been investigated for use as a chemotherapeutant is ivermectin. Ivermectin is a member of the avermectin group of macrocyclic lactones. The avermectins are neurotoxins, which have been used successfully in the treatment of helminthic parasitic infections in a number of terrestrial farm animals and also in the treatment of river blindness in humans. Owing to the low solubility of the compound, ivermectin has been administered as an oral treatment with the feed and has been found to be effective for the treatment of sea lice on Atlantic salmon. Ivermectin is poorly absorbed by fish with a high percentage of the administered dose being excreted in the faeces. The highest concentrations of the absorbed ivermectin were found in lipid‐rich organs. The ivermectin remained in the tissues of the treated fish for a prolonged period of time and was excreted mainly in the unchanged form. Ivermectin can reach the marine environment via excretion from the bile, unabsorbed via the fish faeces and by uneaten food pellets and has a strong affinity to lipid, soil and organic matter. Risk assessments have shown that ivermectin is likely to accumulate in the sediments and that the species therein would be more at risk than the species in the pelagic environment. Ivermectin has been shown to be toxic to some benthic infaunal species in single species tests, but there is no evidence that treatment of fish with ivermectin has affected multispecies benthic communities in the field situation. This review paper ends with details of the development and the status of the use of ivermectin in the treatment of sea lice infestations at Scottish fish farms.     

Evaluation of sea lice abundance levels on farmed Atlantic salmon (Salmo salar L.) located in the Broughton Archipelago of British Columbia from 2003 to 2005

Salmon farming began in British Columbia (BC) in the 1970s and in 2006, aquaculture represented BC's largest agricultural export. Along with this growth in production has been a growth in controversy, including the concern that sea lice originating from Atlantic salmon farms negatively impact wild juvenile pink salmon in the Broughton Archipelago. To understand the dynamic interaction between wild and farmed fish, data for on‐farm abundance of sea lice are required. In this study, 33 000 Atlantic salmon from 20 active farms were examined over 3 years. Two species of lice were found: Lepeophtheirus salmonis and Caligus clemensi. Inter‐annual and seasonal variations in abundance levels occurred with lower levels of L. salmonis in 2003 compared with 2004 and 2005, while C. clemensi levels were highest in 2003. The abundance of L. salmonis was greater on older farmed fish. The findings are compared with European and eastern Canadian sea lice reports, and possible sources of sea lice on farmed salmon are discussed.     

How are the salmon lice (Lepeophtheirus salmonis Krøyer, 1837) in Atlantic salmon farming affected by different control efforts: A case study of an intensive production area with coordinated production cycles and changing delousing practices in 2013–2018

The aims of the present study were to describe the salmon lice (Lepeophtheirus salmonis Krøyer, 1837) situation in an intensive salmon production area in mid‐Norway and to consider implications of changing practices of how salmon lice infestation can be controlled. The results in this study suggest that there are steps that can be carried out to keep salmon lice under control even during years when the temperature facilitates a quick salmon lice development. The present work indicates that the use of cleaner fish can delay the time it takes adult female lice to reach 0.1 per salmon in the beginning of a production cycle. It suggests that the timing of cleaner fish deployment into salmon cages can influence its effectiveness in controlling salmon lice. It also gives caution to letting salmon lice develop unchecked, even at levels far below the current lice limit, because of the difficulties to control salmon lice when the external infection pressure is too high. This study took place during a rapid change in delousing methods, in an area with coordinated salmon production. Despite its exploratory nature, this study offers insights into the salmon lice fluctuations in relation to efforts aimed at controlling it.     

A screening of medicinal compounds for their effect on egg strings and nauplii of the salmon louse Lepeophtheirus salmonis (Krøyer)

Egg strings and nauplii of the salmon louse Lepeophtheirus salmonis were exposed to a variety of medicinal compounds at 50 mg L^?1 for 30 min in two experiment series. This medicine concentration was selected as a starting point for a screening series. Hatching of egg strings and development to copepodid larvae were monitored in one experiment, and the survival and development of nauplii were monitored in the other. Two compounds, emamectin benzoate and cypermethrin, inhibited hatching effectively. Several compounds affected nauplii, either directly or through inhibiting development to the infective stage. A total of 50 mg L^?1 of azamethiphos, cypermethrin, emamectin benzoate and propoxur was lethal to >70% of the larvae. Diflubenzuron, fenoxycarb, pymetrozine, pyriprole and tebufenozide diminished the ability of nauplii developing to copepodids.     

Mapping quantitative trait loci for infectious salmon anaemia resistance in a North American strain of Atlantic salmon

Infectious salmon anaemia (ISA) is a highly virulent viral disease of Atlantic salmon that causes massive economic losses to infected aquaculture operations. Our goal was to detect and map quantitative trait loci (QTL) that confer resistance to ISA in an admixed commercial strain of Atlantic salmon that was largely founded from the Saint John River (SJR) in North America. Full‐sibling families were challenged with a virulent strain of ISA virus. Mortality was tracked during two annual trials with individual fish that survived to the end of the trial being classified as ‘resistant’, and those that died were classified as ‘susceptible’. Ten families with intermediate levels of mortality and an average size of 54.2 individuals were chosen for genotyping with a 50K SNP array designed for the SJR strain. Single nucleotide polymorphisms that were segregating within families were first used to make a composite 11K female linkage map that was then used to find the positions of QTL for ISA resistance using a half‐sib model. The dam‐based HS model detected a total of three QTL for ISA resistance including an experiment‐wide significant QTL on Ssa25 that accounted for 8.3% of the phenotypic variance and chromosome‐wide significant QTL on Ssa03 and on Ssa04 that accounted for 6.0% and 6.6% respectively. We conclude that classic linkage mapping within families continues to be an important method of detecting QTL for oligogenic traits in strains founded from multiple populations. Single nucleotide polymorphisms with moderate trait effects are being used to select within families for more ISA‐resistant strains of Atlantic salmon.     

Safety and efficacy of emamectin benzoate administered in-feed to Atlantic salmon, Salmo salar L., smolts in freshwater, as a preventative treatment against infestations of sea lice, Lepeophtheirus salmonis (Krøyer)

The safety and efficacy of emamectin benzoate, administered in-feed to Atlantic salmon smolts, Salmo salar L., held in freshwater, was evaluated as a preventative treatment against sea lice, Lepeophtheirus salmonis, following transfer of fish to seawater.

In the safety study, salmon smolts held in freshwater were fed with diets containing emamectin benzoate at nominal doses of 0 (control), 50 (recommended dose) and 250 (5× recommended dose) μg kg⁻¹ fish day⁻¹ for 7 days (days 0–6). Actual dose rates, based on measured concentrations of emamectin benzoate in feed, differences in fish weight, and feed consumed, were 0, 54, and 272 μg kg⁻¹ day⁻¹, respectively. On day 9, fish were transferred to seawater and observed for 14 days. No differences in feeding response, coordination, behaviour, gross and histological appearance were observed between control fish and those that received 54 μg kg⁻¹ day⁻¹. Among smolts that received 272 μg kg⁻¹ day⁻¹, approximately 50% exhibited darker coloration, and one fish (1%) exhibited uncoordinated swimming behaviour. No pathognomonic signs of emamectin benzoate toxicity were identified.

In the efficacy study, smolts held in freshwater were fed an unmedicated ration (control group) or emamectin benzoate at 50 μg kg⁻¹ day⁻¹ (treated group) for 7 days (days 0–6). On day 9, fish were re-distributed to eight seawater tanks, each holding 30 control and 30 treated fish. On days 28, 56, 77 and 109, respectively, control and treated fish in two tanks were challenged with L. salmonis copepodites. When lice in each group reached chalimus stage IV, fish were sampled and the numbers of lice were recorded. Fish challenged at day 109 were sampled for the second time when lice were at the adult stage. Efficacy was calculated as the reduction in the mean number of lice on treated fish relative to the mean on control fish. Treatment with emamectin benzoate resulted in an efficacy of 85.0–99.8% in fish challenged at days 28–77, from the start of treatment, and lice counts were significantly lower (P<0.001) on treated fish than on controls. When fish challenged at day 109 were sampled at day 128, efficacy was 44.3%, but survival of chalimus to adult lice on treated fish was lower, and at day 159, efficacy had increased to 73%. These results demonstrate that treatment of salmon smolts with emamectin benzoate in freshwater was well tolerated and highly effective in preventing sea lice infestation following transfer of fish to seawater.     

An assessment of salmon farms and wild salmonids as sources of Lepeophtheirus salmonis (Krøyer) copepodids in the water column in Loch Torridon, Scotland

Wild salmonids and farmed salmon can both be sources of Lepeophtheirus salmonis (Krøyer, 1838) larvae . Farmed salmon smolts free of L. salmonis infections are stocked in sea cages and may subsequently contract L. salmonis infections, probably from wild fish. The contribution of gravid L. salmonis at Atlantic salmon, Salmo salar L., farms to populations of L. salmonis larvae in the water column has in the past been based on estimated parameters, such as louse fecundity. This present study augments these calculations by combining empirical data on densities of infective L. salmonis copepodids in the field with estimates of the number of gravid L. salmonis on farmed and wild salmonids in Loch Torridon. Data collected between 2002 and 2007 show a significant correlation between mean densities of L. salmonis copepodids recovered in the water column and the numbers of gravid L. salmonis at the local salmon farms. Generally, the farms with greatest numbers of salmon were observed to have stronger correlations with densities of copepodids in the water than the farms with fewer fish. The study suggests that louse management approaches, e.g. treatment trigger levels, need to take account of individual farm biomass, or numbers of fish. This study highlights the importance of control of L. salmonis on salmon farms for the co-existence of both wild salmonid populations and the aquaculture industry.     

Stable isotope profiling for large‐scale evaluation of feed efficiency in Atlantic salmon (Salmo salar)

In growing animals, individual variation in feed efficiency may arise from individual differences in growth rate and protein metabolism. Over a period of time, these factors will affect the ratio between ‘new’ vs. ‘old’ protein, which can be quantified using isotope profiling. The aim of this study was to investigate the relationship between relative weight gain and atom percentage excess ¹⁵N in the muscle, liver and mid‐intestine. A 50‐day experiment was conducted with a total of 375 fish initially fed a standard diet, subsequently replaced by one out of five experimental diets, enriched with ¹⁵N. In general, fast‐growing fish are expected to have a better feed efficiency, and the results show that this is captured by isotope profiling in liver and muscle tissues. Furthermore, individual variation in isotope content, that is relative fraction of ‘new’ protein, among fish with comparable growth rates was observed, most expressed around ~50% isotope saturation, indicating differences in protein degradation and replacement not attributed to growth. The results suggest that isotope profiles can be used as individual indicator traits for feed efficiency and that inclusion levels of stable isotopes of 1%–2% gave the most reliable results.     

Rapid compositional analysis of Atlantic salmon (Salmo salar) using visible‐near infrared reflectance spectroscopy

Rapid measurement of salmon flesh quality parameters (>400 samples day^?1) was demonstrated in the laboratory and remotely at industrial sites. Visible‐near infrared spectroscopy (VNIRS) was applied to predict astaxanthin (AX) and fat content in farmed Atlantic salmon. Fish were sampled from thirteen batches (1–6 kg whole weight, containing 2.3–16.3% fat and 1.2–12.5 μg g^?1 AX), and models validated on small (average ± SD: 1.4 ± 0.4 kg) and large fish (4.2 ± 0.9 kg). Both constituents were well predicted in minced Norwegian Quality Cutlet (NQC) samples (r² ≥ 0.86; standard error of prediction (SEP) ≤0.7% for fat and ≤0.7 μg g^?1 for AX). Comparable metrics were observed for AX prediction in whole NQCs (r² = 0.80–0.88; SEP 0.7 μg g^?1). Fat was better predicted in small fish than large fish for whole NQCs (r² = 0.82, SEP 1.0% cf r² = 0.59, SEP = 0.59%) and non‐destructive scanning through the skin of whole, gutted fish (r² = 0.77, SEP = 1.2% cf r² = 0.49, SEP = 1.5%). Models were also developed for screening polyunsaturated fatty acid (PUFA) concentrations, e.g. in NQCs for docosahexaenoic acid (r² = 0.73) and n‐3:n‐6 PUFA (r² = 0.89).     

Retrospective analysis of Atlantic salmon (Salmo salar) marine growth and condition in the northwest Atlantic based on tag‐recovery data

The life history of North American Atlantic salmon (Salmo salar) is characterized by extensive round‐trip migrations between freshwater rearing habitats and marine feeding grounds off the coasts of Canada and Greenland. Growth is rapid during the marine migration, and growth rate and condition factor may be indicators of salmon health during this period. Growth data were evaluated from a tag‐recovery program conducted from 1969 to 1991 using hatchery‐reared Atlantic salmon smolts released in the Penobscot River, Maine, U.S.A. Information from recaptures of 3167 salmon that were at large in the marine environment for 1 month to 3 yr was analyzed. Length–weight measurements coupled with time‐at‐large data were used to estimate von Bertalanffy and allometric growth parameters specific to the marine phase. Variations in growth and condition factor in relation to smolt age, release date, and temperature conditions in the northwest Atlantic were also examined. The von Bertalanffy k parameter declined with ordinal release date, indicating faster growth rates during the first year of smolts released earlier in the spring. The 2‐yr‐old smolts had a larger k than 1‐yr‐old smolts, although 1‐yr‐old smolts grew to a larger asymptotic size. Sea surface temperature had variable effects on growth parameters and condition factor, with temperature at the beginning of the migration and in overwintering habitat during the first year at sea having the greatest influence on length–weight relationships. Determining the mechanisms that influence growth of individuals during the marine phase will help elucidate the factors responsible for historic growth trends, establishing a baseline for current research.     

Variation in sensitivity of sea lice [Lepeophtheirus salmonis (Krøyer)] to dichlorvos on Irish salmon farms in 1991–92

Variation in sensitivity of Lepeophtheirus salmonis (Krøyer) to dichlorvos in static toxicity tests was recorded on salmon farms in Ireland. Five hour LC50s ranged from 60 to 202 μg l ^?1 at 15 °C for five farms in Autumn 1991 and 76–151 μg l^?1 at 10 °C for four farms in Spring of 1992. Resistance ratios, calculated using the most sensitive sites as baseline, were low and ranged from 1.32 to 3.27 in 1991 and 1.28–1.98 in 1992. Repeat sampling on three farms in 1991 and 1992 also indicated short‐term changes in sensitivity to dichlorvos within farms.     

Resistance of sea lice, Lepeophtheirus salmonis (Krøyer), to hydrogen peroxide on farmed Atlantic salmon, Salmo salar L.

Hydrogen peroxide has been the only medicine used to treat salmon infected with sea lice [Lepeophtheirus salmonis (Krøyer)] on many farms in Scotland since 1992 and reports have suggested reduced treatment efficacy. The present study tests the sensitivity of sea lice to exposure to hydrogen peroxide under farm conditions and also in vivo by comparing lice from a farm with suspected resistance and lice from a farm that had not been pre‐exposed to hydrogen peroxide. In bin treatments on fish from a farm treated with hydrogen peroxide on 41 occasions, numbers of ovigerous lice declined by only 15% and 16% in two replicates and other mobile stages by 25%. Where hydrogen peroxide had not been used, ovigerous females declined by 90% and 87% and other mobile lice stages by 97% and 99%. These trials and observed poor efficacy of cage treatments, for example only 7.5% reduction in lice numbers when fish were treated with 2000 p.p.m. for 20 mins, indicated resistance of lice to hydrogen peroxide. The mechanisms involved in the development of resistance, possible genetic selection for lice with reduced carapace permeability or detoxifying enzymes such as catalase, or tolerance through induction by subtherapeutic exposure are reviewed. Implications for lice control strategies relying on hydrogen peroxide are discussed.     

Effect of the arachidonic acid/vitamin E interaction on the immune response of juvenile Atlantic salmon (Salmo salar) challenged against Piscirickettsia salmonis

Atlantic salmon (Salmo salar) were fed 6 experimental diets containing three levels of arachidonic acid (ARA) (0.18 g kg^?1, 0.28 g kg^?1 and 0.63 g kg^?1 for low, medium and high levels, respectively) and two levels of vitamin E (150 and 730 mg kg^?1 for low and high levels, respectively). At the end of the experimental period, fatty acids in the liver and immunity markers (lysozyme activity, respiratory burst and phagocytic activity) were determined and fish subjected to a challenge test against the salmonid rickettsial syndrome (SRS) pathogen. ARA, vitamin E or their interaction did not exert an effect on fish performance, whereas ARA alone clearly increased the deposition of ARA. Dietary vitamin E only enhanced liver vitamin E deposition, while the interaction of ARA and vitamin E influenced lysozyme activity and EPA/ARA ratio pointing out the effect of both nutrients on the fish immune system and metabolism. Only the medium concentration contributed to reducing mortality when the fish were exposed to the SRS pathogen. In conclusion, different levels of supplementation with ARA and vitamin E in the diet had no effect on productivity, but did have effects on immune markers and cumulative mortality when fish were exposed to the SRS pathogen.