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.     

Investigations on the role of the salmon louse,Lepeophtheirus salmonis (Caligidae), as a vector in the transmission of Aeromonas salmonicida subsp. salmonicida

A bacteria–parasite challenge model was used to study the role of sea lice, Lepeophtheirus salmonis (Copepoda), as a vector of Aeromonas salmonicida subsp. salmonicida. Three hypotheses were tested: (i) L. salmonis can acquire A. salmonicida subsp. salmonicida via water bath exposure; (ii) L. salmonis can acquire the bacteria via parasitizing infected Atlantic salmon, Salmo salar; and (iii) L. salmonis can transmit the bacteria to naïve Atlantic salmon via parasitism. Adult L. salmonis exposed to varying A. salmonicida subsp. salmonicida suspensions (10¹–10⁷ cells mL^?1) for 1.0, 3.0 or 6.0 h acquired the bacteria externally (12.5–100%) and internally (10.0–100%), with higher prevalences associated with the highest concentrations and exposures. After exposure to 10⁷ cells mL^?1, viable A. salmonicida subsp. salmonicida could be isolated from the external carapace of L. salmonis for 120 h. Lepeophtheirus salmonis also acquired the bacteria externally and internally from parasitizing infected fish. Bacterial transmission was observed only when L. salmonis had acquired the pathogen internally via feeding on ‘donor fish’ and then by parasitizing smaller (<50 g) ‘naive’ fish. Under specific experimental conditions, L. salmonis can transfer A. salmonicida subsp. salmonicida via parasitism; however, its role as a mechanical or biological vector was not defined.     

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.     

Disruption of host‐seeking behaviour by the salmon louse,Lepeophtheirus salmonis,using botanically derived repellents

The potential for developing botanically derived natural products as novel feed‐through repellents for disrupting settlement of the salmon louse, Lepeophtheirus salmonis (Caligidae) upon farmed Atlantic salmon, Salmo salar, was investigated using an established laboratory vertical Y‐tube behavioural bioassay for assessing copepodid behaviour. Responses to artificial sea water conditioned with the odour of salmon, or to the known salmon‐derived kairomone component, α‐isophorone, in admixture with selected botanical materials previously known to interfere with invertebrate arthropod host location were recorded. Materials included oils extracted from garlic, Allium sativum (Amaryllidaceae), rosemary, Rosmarinus officinalis (Lamiaceae), lavender, Lavandula angustifolia (Lamiaceae), and bog myrtle, Myrica gale (Myricaceae), and individual components (diallyl sulphide and diallyl disulphide from garlic; allyl, propyl, butyl, 4‐pentenyl and 2‐phenylethyl isothiocyanate from plants in the Brassica genus). Removal of attraction to salmon‐conditioned water (SCW) or α‐isophorone was observed when listed materials were presented at extremely low parts per trillion (ppt), that is picograms per litre or 10^?12 level. Significant masking of attraction to SCW was observed at a level of 10 ppt for diallyl disulphide and diallyl sulphide, and allyl isothiocyanate and butyl isothiocyanate. The potential of very low concentrations of masking compounds to disrupt Le. salmonis copepodid settlement on a host fish has been demonstrated in vitro.     

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.     

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.     

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.     

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.     

The Three‐spined Stickleback,Gasterosteus aculeatus Linnaeus 1758, plays a minor role as a host of Lepeophtheirus salmonis (Krøyer 1837) in the Gulf of Maine

The sea louse, Lepeophtheirus salmonis (Krøyer 1837), is a significant parasite of farmed salmon throughout the Northern Hemisphere. Management of on‐farm louse populations can be improved by understanding the role that wild fish play in sustaining and providing refuge for the local population of sea lice. In this study, 1,064 sticklebacks were captured. Of these animals, 176 individuals were carrying a total of 238 sea lice, yielding a prevalence and intensity of 16.5% and 1.4 lice per fish, respectively. Detailed examination of the sea lice on the three‐spined sticklebacks captured in Cobscook Bay found two L. salmonis individuals using three‐spined sticklebacks as hosts. A 2012 survey of wild fish in Cobscook Bay, Maine, found multiple wild hosts for Caligus elongatus (von Nordmann 1832), including three‐spined sticklebacks (Gasterosteus aculeatus L.), but no L. salmonis were found in this earlier study.     

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.     

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 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).     

Sea lice, Lepeophtheirus salmonis, transfer between wild sympatric adult and juvenile salmon on the north coast of British Columbia, Canada

We examine sea lice, Lepeophtheirus salmonis , on juvenile and adult salmon from the north coast of British Columbia between 2004 and 2006 in an area that does not at present contain salmon farms. There is a pronounced zonation in the abundance of L. salmonis on juvenile pink salmon, Oncorhynchus gorbuscha , in the Skeena and Nass estuaries. Abundances in the proximal and distal zones of these estuaries are 0.01 and 0.05 respectively. The outer zones serve as feeding and staging areas for the pink salmon smolts. Returning Chinook, Oncorhynchus tshawytscha , and coho salmon, Oncorhynchus kisutch , concentrate in these areas. We collected data in 2006 to examine whether L. salmonis on returning adult salmon are an important source of the sea lice that appear on juvenile pink salmon. Nearly all (99%) of the sea lice on returning Chinook and over 80% on coho salmon were L. salmonis. Most of the L. salmonis were motile stages including many ovigerous females. There was a sharp increase in the abundance of sea lice on juvenile pink salmon smolts between May and July 2006 near the sites of adult captures. As there are no salmon farms on the north coast, few sticklebacks, Gasterosteus aculeatus , and very few resident salmonids until later in the summer, it seems that the most important reservoir of L. salmonis under natural conditions is returning adult salmon. This natural source of sea lice results in levels of abundance that are one or two orders of magnitude lower than those observed on juvenile pink salmon in areas with salmon farms such as the Broughton Archipelago.     

Impacts of parasites on marine survival of Atlantic salmon: a meta‐analysis

Parasites can, in theory, have large impacts on the survival of fish populations. One method to evaluate such impacts on anadromous species is to apply manipulative field experiments in which parallel groups of antiparasitically treated and non‐treated fish are simultaneously released and then subsequently recaptured as returning adults. A systematic review and meta‐analysis on all such Norwegian studies on Salmo salar provided a data set for the time period 1996 to 2011 on 118 release groups comprising 657 624 fish released and 3989 recaptured. The overall risk ratio (RR) was estimated to be 1.18 (95% CI: 1.07–1.30). The effect varied strongly between groups, (Higgins I² = 40.1%). Over 70% of this heterogeneity could be explained by the release location, time period and baseline survival. The most important predictor variable was baseline survival. In groups with low recapture in the control group (low baseline survival), the effect of treatment was high (RR = 1.7), while in groups with high recapture in the control group (high baseline survival), there was no effect of treatment (RR ~ 1.00). The most prevalent parasite in the region affected by the drugs administered was Lepeophtheirus salmonis. Hence, the meta‐analysis supports the hypothesis that L. salmonis contributes to the mortality of S. salar during outward migration. However, the effect of treatment was not consistent, but was evidently strongly modulated by other risk factors. The results suggest that the population‐level effects of parasites cannot be estimated independently of other factors affecting the marine survival of S. salar.     

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.     

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.