The effect of light on the settlement of the salmon louse, Lepeophtheirus salmonis, on Atlantic salmon, Salmo salar L

The salmon louse, Lepeophtheirus salmonis, is an ectoparasitic copepod that infests both wild and farmed salmonid fish. Salmon lice are a major disease problem in the farming of Atlantic salmon, Salmo salar L., and the possibility of salmon lice playing a role in the decline of wild anadromous stocks has also been raised. Lepeophtheirus salmonis can detect a range of stimuli (pressure/moving water, chemicals and light) in the external environment. However, the response thresholds to various stimuli, and the spatial and temporal scales over which they operate in the context of host location, are largely unknown. In this context, we attempted to determine whether salmon lice copepodids settle onto hosts more effectively, or at different locations on the fish's body, under different qualities of light. Lice settlement trials were conducted under three lighting conditions; L1: unpolarized under ultraviolet A (UVA)-through visible; L2: unpolarized without UVA (control); L3: 100% linearly polarized without UVA. A dark control was also conducted. No statistically significant difference in lice settlement was found. While changes in light intensity are involved in host detection at spatial scales on the order of metres, the results presented here suggest that it is not the primary sensory modality underlying host location at smaller spatial scales (cm to mm).     

A comparison of epidemiological patterns of salmon lice, Lepeophtheirus salmonis, infections on farmed Atlantic salmon, Salmo salar L., in Norway and Scotland

This paper examines two large national data sets collected over several years and contrasts the patterns of sea lice, Lepeophtheirus salmonis (Krøyer, 1837), infections, the use of treatments and the occurrence of chalimus peaks between Norwegian and Scottish farms. Infection levels in Scotland were significantly higher in general over the period under study. For the chalimus stage group in the first quarter of the year, Norwegian mean abundance stayed below 10 lice per m² while Scottish means reached 45 lice per m² of fish skin per m³ of water. Both countries had more chalimus in summer than at other times of year, but in the last 3 months of the year Scottish fish had, on average, two to four times as many chalimus as Norwegian fish. Peaks of chalimus abundance were more frequent in Scotland, particularly in winter, but the most prominent peaks occurred in summer in both countries. In Scotland a marked mid-year build-up of mobile pre-adult and adult stages was seen, and both countries showed a tendency for mobile counts on the second year fish to increase towards the end of the year. Scottish fish carried, on average, three times as many mobile lice per m² of skin as Norwegian fish in the last 3 months of the year. The difference in lice loads was reflected in the greater use of veterinary medicines on Scottish farms. The higher infection levels in Scotland may be due to shallower and more enclosed water bodies used for farming, smaller and shallower pens, differences in sea water temperatures or in access to appropriate medication. The results highlight the importance of ensuring that effective veterinary medicines are available in the UK for the control of infection.     

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.     

Parasite fecundity decreases with increasing parasite load in the salmon louse Lepeophtheirus salmonis infecting Atlantic salmon Salmo salar

Aggregation is common amongst parasites, where a small number of hosts carry a large proportion of parasites. This could result in density‐dependent effects on parasite fitness. In a laboratory study, we explored whether parasite load affected parasite fecundity and survival, using ectoparasitic salmon lice (Lepeophtheirus salmonis Krøyer, 1837) infecting Atlantic salmon (Salmo salar) hosts. We found a significant reduction in fecundity with higher parasite load, but no significant effect on survival. Together with previous findings, this suggests that stronger competition amongst female lice under high parasite load is a more likely explanation than increased host immune response.     

A mathematical model of the growth of sea lice, Lepeophtheirus salmonis, populations on farmed Atlantic salmon, Salmo salar L., in Scotland and its use in the assessment of treatment strategies

Sea lice are a persistent problem for farmed and wild salmonid populations. Control can be achieved through the use of veterinary medicines. A model was developed to describe the patterns of sea lice infection on salmon farms in Scotland and to predict the likely effect of various treatment strategies. This model takes into account development rates and mortality using compartments representing life history stages and external infection pressure. The national sea lice infection pattern was described using parameters representing stage survival, background infection levels and egg viability rates. The patterns observed across farms varied greatly and the model gave broad agreement to observed trends with different parameters being required in the model for sites using hydrogen peroxide and cypermethrin treatments. The parameter estimates suggest that the background infection pressure on sites where cypermethrin was administered was higher than for those using hydrogen peroxide. Both models had comparable magnitudes of sensitivity with survival from one stage to another being the most sensitive parameter, followed by feedback rates at which gravid females produce eggs, with background infection levels the least sensitive. The effect of different cypermethrin treatment strategies was assessed using the model. Increasing treatments in a production cycle gave more effective control. However, the model showed that timing of treatments is most important if sea lice are to be effectively controlled.     

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.     

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.     

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.     

Development of microsatellite PCR typing methodology for the sea louse, Lepeophtheirus salmonis (Krøyer)

In order to develop a microsatellite typing system for Lepeophtheirus salmonis (Krøyer), a DNA preparation method for individual sea lice suitable for analysis by polymerase chain reaction (PCR) was designed, and the DNA sequences of 50 L. salmonis microsatellite elements were determined. The microsatellites were composed of 60% perfect, 25% imperfect, and 15% compound repeats. Based on the flanking DNA sequences, four microsatellite‐PCR assays were optimized and used in a pilot study to analyse L. salmonis samples collected in Ireland, Norway and Scotland. Two of the microsatellite‐PCR assays targeted polymorphic loci amplifying seven and 10 alleles respectively. The results showed that microsatellite‐PCR typing could detect genetic variation both within and between the L. salmonis groups, and also was capable of amplifying group‐specific alleles.     

Changes in epidemiological patterns of sea lice infestation on farmed Atlantic salmon, Salmo salar L., in Scotland between 1996 and 2006

Analyses of a unique database containing sea lice records over an 11 year period provide evidence of changing infestation patterns in Scotland. The data, collected from more than 50 commercial Atlantic salmon farms, indicate that both species of sea lice commonly found in Scotland, Lepeophtheirus salmonis and Caligus elongatus, have declined on farms over the past decade. Reductions for both species have been particularly marked since 2001 when more effective veterinary medicines became available. Treatment data were also available in the database and these show a growing trend towards the use of the in-feed medication emamectin benzoate (Slice), particularly in the first year of the salmon production cycle. However, this trend towards single product use has not been sustained in 2006, the latest year for which data are available. There is some evidence of region to region variation within Scotland with the Western Isles experiencing higher levels of infestation. However, compared to the levels observed between 1996 and 2000, all regions have benefited from reduced lice infestation, with the overall pattern showing a particular reduction in the second and third quarters of the second year of production.     

The histopathology associated with the juvenile stages of Lepeophtheirus salmonis on the Atlantic salmon, Salmo salar L.

Abstract. Atlantic salmon infected with juvenile stages of Lepeophtheirus salmonis were obtained from sea loch cage sites on the West coast of Scotland. Attached parasites on the skin were studied using light and electron microscopy. Behavioural activity was observed in attached stages maintained on skin in vitro. Pathological changes were associated with four main areas of interaction between the parasite and the host, namely second antennal attachment, filament attachment, maxilliped activity and feeding activity. The skin changes involved were initial mechanical disruption followed by epidermal hyperplasia. The filament material was secreted into the epidermis and spread laterally without breaching the basement membrane. Surprisingly little reaction to the filament occurred, but chalimus activity resulted in chronic changes which were more marked as the later chalimus stages were reached. Fibroplasia was a feature of lesions induced by redundant filaments resulting in small nodular lesions. After erosion of these nodules, all that remained were the melanized rings characteristic of old chalimus sites.     

The host transfer ability of Lepeophtheirus salmonis (Copepoda: Caligidae) from farmed Atlantic salmon, Salmo salar L.

The ability of mobile stages of Lepeophtheirus salmonis to transfer between salmon, Salmo salar L., hosts was examined by two methods. The first of these was the removal of certain stages on farmed fish, while the second comprised the placement of mobile lice on clean fish in the laboratory. In one-sea-winter farmed salmon, 63% of males and 52% of female lice transferred to new hosts over a period of 4 days. In laboratory trials on post-smolts, 61% of males and 69% of female lice transferred to new hosts over 3 days. In another experiment, free-swimming adult male lice were added to the tank, and 90% of these were found on fish after 24 h. Immigration of lice to fish in trial cages and the large proportion of lice transferring between fish in both laboratory and farm trials indicates that L. salmonis can transfer between hosts.     

Apoptosis inhibition of Atlantic salmon (Salmo salar) peritoneal macrophages by Piscirickettsia salmonis

To improve the understanding of the piscirickettsiosis pathogenesis, the in vivo apoptosis modulation of peritoneal macrophages and lymphocytes was studied in juvenile Salmo salar intraperitoneally injected with Piscirickettsia salmonis. Five fish were sampled at post‐exposure days 1, 5, 8 (preclinical), 20 (clinical) and 40 (post‐clinical period of the disease), and the leucocytes of their coelomic washings were analysed by flow cytometry (using the JC‐1 cationic dye), TUNEL and cytology to detect apoptotic cells. A selective and temporal pattern of apoptosis modulation by P. salmonis infection was observed. Apoptosis in lymphocytes was not affected, whereas it was inhibited in macrophages but only during the preclinical stage of the induced piscirickettsiosis. Hence, it is postulated that P. salmonis inhibits macrophage apoptosis at the beginning of the disease development to survive, multiply and probably be transported inside these phagocytes; once this process is complete, macrophage apoptosis is no longer inhibited, thus facilitating the exit of the bacteria from the infected cells for continuing their life cycle.     

Experimental infection of turbot, Scophthalmus maximus (L.), by Moritella viscosa, vaccination effort and vaccine-induced side-effects

Moritella viscosa is the causative agent of winter ulcers in farmed salmonids and Atlantic cod in countries around the North Atlantic. The bacterium has also been isolated from various marine fish species. Bacterial diseases have been a limiting factor in farming of turbot, but M. viscosa has not so far been isolated. In this study, turbot was shown to be sensitive to M. viscosa infection in experimental challenges. Pathological changes in infected turbot were comparable with those previously described for winter ulcers in salmon. A multivalent commercial salmon vaccine, containing M. viscosa as one of five antigens and a mineral oil adjuvant, did not protect turbot against challenge 13 weeks post-vaccination. Weight gain of vaccinated turbot compared with controls was not reduced 7 weeks post-vaccination. Vaccination did not induce a specific anti-M. viscosa response, while elevated anti-M. viscosa antibody levels were detected both in vaccinated and unvaccinated fish 5 weeks post-challenge. The vaccine did, however, induce an antibody response against Aeromonas salmonicida, another vaccine component. Minor intra-abdominal adhesions were detected in vaccinated fish and fish injected with a mineral oil adjuvant. The measurement of various innate humoral immune parameters did not reveal significant differences between vaccinated and control groups.     

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.