Ultrasound as potential inhibitor of salmon louse infestation—A small‐scale study

The effect of ultrasound exposure during controlled infection with salmon lice, Lepeophtheirus salmonis, has been studied. Salmon were placed in tanks with salmon lice copepodids for 1 hr, while simultaneously being exposed to sound frequencies of 9.3, 21 or 54 kHz. The sound transducers were operated at maximum power levels, and the 9.3 kHz transducer generated the highest sound level (220.6 dB). Only the group exposed to 9.3 kHz displayed a significant reduction in louse infestation. However, the observed effect of ultrasound was relatively small, and in a practical implementation in sea cages, the sound intensity will be lower than that used in the experiments. It is also possible that the observed reduction in infestation is due to ultrasonic cavitation effects, which are only present at a very short range from the ultrasound source. We therefore do not consider ultrasound a feasible method for preventing attachment of salmon lice copepodids on salmon in cage farms.     

The development of the Norwegian wrasse fishery and the use of wrasses as cleaner fish in the salmon aquaculture industry

Norway leads the world aquaculture production of Atlantic salmon Salmo salar and farmed Norwegian Atlantic salmon is currently consumed around the globe. However, sea lice infestation is a major problem faced by the salmon aquaculture industry in Norway and elsewhere. The use of wild-caught cleaner fish, mainly wrasses, has been recommended over the other available methods as the most economical and environmentally friendly option to control sea lice infestation in salmon farming. Here, we review the development of the Norwegian wrasse fishery and the use of wrasses as cleaner fish. In this document, we address the sea lice problem and introduce the main wrasse species employed as cleaner fish, document the cleaning behaviour of wrasses, present the development of a new wrasse fishery associated with the salmon aquaculture industry, and finally, we identify the main challenges associated with the intensive use of wild-caught cleaner wrasses and provide some insight for future directions of the wrasse fishery and further development of aquaculture techniques to supply salmon facilities with domesticated cleaner fish.     

Modelling salmon lice,Lepeophtheirus salmonis,reproduction on farmed Atlantic salmon,Salmo salar L

The aim of this study was to model sea lice levels and the effect on reproduction by a stochastic simulation model and to evaluate the uncertainty of lice estimates based upon counts. Two empirical data sets were examined to parameterize the models. An overall fit of the data to the Poisson distribution was found and thus was used as the base of the stochastic models. In the model, salmon lice reproduction is not linear with the number of adult females and at low lice loads a smaller proportion of the adult female lice will reproduce. Depending on the variance structure, it was estimated that between 40% and 60% of the adult female lice will reproduce at an abundance of 0.5 adult females per fish. Lice counts, especially when examining few fish at low lice loads, are uncertain and at a true abundance of 0.1 one may count between 0 and 5 lice when examining 10 fish. Understanding the dynamics of sea lice reproduction is a key factor in the development of sustainable control strategies.     

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.     

Distributions of planktonic sea lice larvae Lepeophtheirus salmonis in the inter-tidal zone in Loch Torridon, Western Scotland in relation to salmon farm production cycles

This study examines the relative density of sea lice, Lepeophtheirus salmonis (Krøyer), larvae in the inter‐tidal areas of Loch Torridon, a fjordic sea loch in Western Scotland and the site of several commercial salmon farms. Samples of planktonic sea lice were collected in each spring from 1999 to 2003 near the mouth of the River Shieldaig, where sea trout have been shown to congregate in the first few days after going to sea, and from 2001 to 2003 near the mouth of the River Balgy. No larval sea lice were found near the mouth of the River Shieldaig in spring 2000 or 2002, when gravid female sea lice were absent on local fish farms. The same finding was observed in 2002 at the mouth of the River Balgy. In springs of 1999, 2001 and 2003, peak densities of larval sea lice of 33, 94 and 143 m^?3, respectively, were observed at the mouth of the river Shieldaig. At the mouth of the River Balgy, peak densities of 74 and 78 lice m^?3 were observed in 2001 and 2003 respectively. At the time these samples were collected, the two local salmon farms were in the second year of their production cycle and gravid female sea lice were present. Samples of larval sea lice were also obtained year round, at approximately weekly intervals, from a 50‐m transect at the mouth of the River Shieldaig, from March 2001 to June 2003, and compared with frequencies of gravid female sea lice on the two local salmon farms. High levels were found in the winter. No planktonic sea lice were found in this transect when gravid females were not present on the local 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.     

Differences in Lepeophtheirus salmonis abundance levels on Atlantic salmon farms in the Broughton Archipelago, British Columbia, Canada

Sea lice data collected from Atlantic salmon farms in the Broughton Archipelago between 2003 and 2005 were examined for inter-regional differences in mobile Lepeophtheirus salmonis (Krøyer) abundance using the generalized linear model procedure. Factors such as age of the salmon populations, location of farms and time of year had a significant effect on the abundance of the mobile stages of L. salmonis whereas water temperature and salinity did not. Separate evaluation of SLICE® treatment data found no significant difference in treatment frequency among the areas but did show that there were significantly lower numbers of farm treatments during the summer months when compared with other seasons. The role of environment and wild fish in influencing sea lice abundance on the farmed salmon is discussed. The findings suggest that effective management programmes for sea lice should not only be based on geographical location but should take into account other factors which could influence lice abundance levels.     

Atlantic salmon infected with salmon lice are more susceptible to new lice infections

Aggregation is commonly observed for macroparasites, but its adaptive value remains unclear. Heavy infestations intensities may lead to a decrease in some fitness‐related traits of parasites (e.g. parasite fecundity or survival). However, to a dioecious parasite, increased aggregation could also increase the chance of finding individuals of the opposite sex. In a laboratory experiment, we tested if previous experience with salmon lice (Lepeophtheirus salmonis) affected susceptibility of Atlantic salmon (Salmo salar) to later exposure to the same parasite species. We found that currently infected fish got higher intensities of new lice than naive fish. This suggests that hosts already carrying parasites are more susceptible to new lice infections. For this dioecious parasite, such positive density dependence might be adaptive, ensuring successful reproduction under conditions of low lice densities by increasing the probability of both sexes infecting the same host.     

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.     

Early‐stage sea lice recruits on Atlantic salmon are freshwater sensitive

Sea lice are significant parasites of marine and brackish farmed fishes. Freshwater bathing is a potential control option against numerous sea lice species, although has been viewed as futile against those that are capable of tolerating freshwater for extended periods. By comparing freshwater survival times across host‐attached stages of Lepeophtheirus salmonis (Krøyer), a key parasite in Atlantic salmon farming, we show the first attached (copepodid) stage undergoes 96–100% mortality after 1 h in freshwater, whereas later attached stages can tolerate up to 8 days. Thus, regular freshwater bathing methods targeting the more susceptible attached copepodid stage may successfully treat against L. salmonis and potentially other sea lice on fish cultured in marine and brackish waters.     

A note on sea lice abundance on farmed Atlantic salmon in Scotland 2011–2013: significant regional and seasonal variation

Sea lice are the most economically damaging parasite affecting marine Atlantic salmon production the dominant sector of UK aquaculture. The Scottish Salmon Producers' Organisation has published mean adult female lice abundance for four regions for 2011–2013 and for 30 areas in 2013. An analysis of these data shows that sea lice abundance is highly seasonal with values lowest in the first half of the year. Abundance also varies by region, the regional patterns vary from year‐to‐year but overall the highest abundance was found in the northern part of mainland Scotland. While most areas had low lice abundance, in a few areas the abundance was persistently above average; these areas were clustered together. There was no significant year‐on‐year trend in lice abundance, and trends appear to vary between regions. Although previous Scottish analyses have been carried out using data from a large company, this analysis cover 95% of production in all salmon farming regions of Scotland and it can be extended as more data become available.     

PCR survey for Paramoeba perurans in fauna,environmental samples and fish associated with marine farming sites for Atlantic salmon (Salmo salar L.)

Amoebic gill disease (AGD) caused by the amoeba Paramoeba perurans is an increasing problem in Atlantic salmon aquaculture. In the present PCR survey, the focus was to identify reservoir species or environmental samples where P. perurans could be present throughout the year, regardless of the infection status in farmed Atlantic salmon. A total of 1200 samples were collected at or in the proximity to farming sites with AGD, or with history of AGD, and analysed for the presence of P. perurans. No results supported biofouling organisms, salmon lice, biofilm or sediment to maintain P. perurans. However, during clinical AGD in Atlantic salmon, the amoeba were detected in several samples, including water, biofilm, plankton, several filter feeders and wild fish. It is likely that some of these samples were positive as a result of the continuous exposure through water. Positive wild fish may contribute to the spread of P. perurans. Cleaner fish tested positive for P. perurans when salmon tested negative, indicating that they may withhold the amoeba longer than salmon. The results demonstrate the high infection pressure produced from an AGD‐afflicted Atlantic salmon population and thus the importance of early intervention to reduce infection pressure and horizontal spread of P. perurans within farms.     

The settlement and reproductive success of Lepeophtheirus salmonis (Krøyer 1837; Copepoda: Caligidae) on atypical hosts

The reproductive success of Lepeophtheirus salmonis settled on host and non‐host fish has been compared. Triplicate single species tanks of Atlantic salmon, marine three‐spined sticklebacks, saithe and Atlantic cod were exposed to 10 adult female L. salmonis per tank (n=30 lice per species). Adult female L. salmonis settlement and egg string production occurred only on salmon and cod, with no egg production occurring on saithe and three‐spined sticklebacks. The number of eggs in egg strings, hatching success of eggs and the survival of all larval stages to the copepodid stage were severely affected by the species of fish on which female L. salmonis had settled. L. salmonis settled on cod produced significantly fewer eggs, lower hatching rates and lower survival rates of larvae than females on Atlantic salmon. The production of egg strings by L. salmonis females infecting cod, which successfully hatch and moult through to the infective copepodid stage, albeit in small numbers, is discussed in terms of the implications to aquaculture and salmon and cod farming scenarios.     

Does the AquasmartTM feeding system reduce sea louse [Lepeophtheirus salmonis (Krøyer)] infestation on farmed Atlantic salmon (Salmo salar L.) in winter?

Infections of sea lice [Lepeophtheirus salmonis (Krøyer)] were studied over winter on a working, commercial salmon farm in relation to two types of feeding system: a regular, pneumatic spray feeder and an irregular, ‘demand feed’ sensor system (Aquasmart^TM). Contrary to prediction, the Aquasmart^TM system alone had no significant effect on chalimus infestation compared with fish fed by pneumatic spray. However, it was associated with significantly reduced numbers of mobile stages, both at the beginning of the study (P < 0.05) and, more particularly, in conjunction with treatment of all pens using hydrogen peroxide (P < 0.001). It is suggested that feeding regime may represent a useful tool in reducing lice infestation when co‐ordinated with other methods of control, such as peroxide treatment, and that further work in this field is merited.     

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.     

Aeromonas salmonicida infection levels in pre‐ and post‐stocked cleaner fish assessed by culture and an amended qPCR assay

Due to increasing resistance to chemical therapeutants, the use of ‘cleaner fish’ (primarily wrasse, Labridae, species) has become popular in European salmon farming for biocontrol of the salmon louse, Lepeophtheirus salmonis (Krøyer). While being efficient de‐licers, cleaner fish mortality levels in salmon cages are commonly high, and systemic bacterial infections constitute a major problem. Atypical furunculosis, caused by Aeromonas salmonicida A‐layer types V and VI, is among the most common diagnoses reached in clinical investigations. A previously described real‐time PCR (qPCR), targeting the A. salmonicida A‐layer gene (vapA), was modified and validated for specific and sensitive detection of all presently recognized A‐layer types of this bacterium. Before stocking and during episodes of increased mortality in salmon cages, cleaner fish (primarily wild‐caught wrasse) were sampled and screened for A. salmonicida by qPCR and culture. Culture indicated that systemic bacterial infections are mainly contracted after salmon farm stocking, and qPCR revealed A. salmonicida prevalences of approximately 4% and 68% in pre‐ and post‐stocked cleaner fish, respectively. This underpins A. salmonicida's relevance as a contributing factor to cleaner fish mortality and emphasizes the need for implementation of preventive measures (e.g. vaccination) if current levels of cleaner fish use are to be continued or expanded.     

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.     

Relationship between sea lice levels on sea trout and fish farm activity in western Scotland

The relationship between aquaculture and infestations of sea lice on sea trout, Salmo trutta L., is controversial. Here, the association between sea lice infestations on wild sea trout and characteristics of local Atlantic salmon, Salmo salar L., farms were investigated using data collected on the Scottish west coast. The proportion of sea trout with louse burdens above a critical level was positively related to the fork length of the sea trout and the mean weight of salmon on the nearest fish farm, and negatively related to the distance to that farm. The distance to the nearest fish farm did not influence the probability of infestations above the critical level beyond 31 km although there was considerable uncertainty around this cut‐off distance (95% limits: 13–149 km). The results support a link between Atlantic salmon farms and sea lice burdens on sea trout in the west of Scotland and provide the type of information required for marine spatial planning.     

Development of a Piscirickettsia salmonis immersion challenge model to investigate the comparative susceptibility of three salmon species

Piscirickettsia salmonis, the aetiological agent of salmonid rickettsial septicaemia (SRS), is a global pathogen of wild and cultured marine salmonids. Here, we describe the development and application of a reproducible, standardized immersion challenge model to induce clinical SRS in juvenile pink (Oncorhynchus gorbuscha), Atlantic (Salmo salar) and sockeye salmon (O. nerka). Following a 1‐hr immersion in 10⁵ colony‐forming units/ml, cumulative mortality in Atlantic salmon was 63.2% while mortality in sockeye salmon was 10%. Prevalence and levels of the bacterium in kidney prior to onset of mortality were lower in sockeye compared with Atlantic or pink salmon. The timing and magnitude of bacterial shedding were estimated from water samples collected during the exposure trials. Shedding was estimated to be 82‐fold higher in Atlantic salmon as compared to sockeye salmon and peaked in the Atlantic salmon trial at 36 d post‐immersion. These data suggest sockeye salmon are less susceptible to P. salmonis than Atlantic or pink salmon. Finally, skin lesions were observed on infected fish during all trials, often in the absence of detectable infection in kidney. As a result, we hypothesize that skin is the primary point of entry for P. salmonis during the immersion challenge.