Wild salmonids and sea louse infestations on the west coast of Scotland: sources of infection and implications for the management of marine salmon farms

The sea louse (Lepeophtheirus salmonis Kr?yer) is a major health problem for both farmed and wild salmonids. This paper investigates louse epidemiology and management in the salmon-farming zone of western Scotland. Based on a review of the marine ecology of wild salmon (Salmo salar L) and sea trout (Salmo trutta L), and catch and farm production statistics, best estimates were made for numbers of wild and farm hosts present in coastal waters in March-June 2000. Applying data for ovigerous female louse infections and fecundity, the sources and risks of larval transmission to wild salmon and sea trout were modelled. Farm salmon in the second spring of production were the primary host group (98% of fish), while numbers of wild salmonids (< 1%) and escaped farm salmon (2%) were relatively insignificant. Farm salmon produced 97% of louse eggs at high levels (eight ovigerous lice per fish), and 78% at low levels (one per fish). Wild salmonids produced < 1% of eggs under both scenarios, but escaped farm salmon produced 3% and 21%, respectively. All hosts potentially cross-infect one another, but farm salmon are more likely to infect wild and farm smolts, and also other farm salmon. Monitoring of lice on sea trout in June 1998-2000 by the Association of West Coast Fisheries Trusts corroborated the model's conclusions. Localised epizootics occurred every year and coincided with the presence of ovigerous lice on local farms. In areas of mixed-year class production on farms, epizootics were evident every spring, but occurred every second spring in areas of single-year class production. In 1998-2000 at least 14-40% of sea trout were infected with potentially lethal infestations of lice. Ovigerous louse levels of < 0.005 per fish were required on farm salmon in the spring of 2000 to produce less eggs than those emitted by wild salmonids. With the industry's continued expansion, and thus increased numbers of farm salmon, a target of zero ovigerous lice will be required on farms to minimise impacts on wild salmonids. Due to the limited long-term efficacy and availability of louse medicines, management strategies are discussed which will improve control, including single-year class production over large areas, alternate S1-S1/2 smolt inputs, and 11-month production cycles.     

Temporal,environmental and management factors influencing the epidemiological patterns of sea lice (Lepeophtheirus salmonis) infestations on farmed Atlantic salmon (Salmo salar) in Scotland

In association with industry, an extensive database has been established on measurements of sea louse counts on farmed Atlantic salmon for the years 1996 to 2000 from 33 Scottish fish farms. These data include extensive counts on the sea louse species, Lepeophtheirus salmonis, at various stages of the life cycle and in particular the chalimus and mobile stages. There has been considerable speculation as to what factors might affect the abundance of sea lice, much of which is based on limited evidence. Our analyses show that there is tremendous variation in sea louse infestation patterns from year to year, whereas stock type, geographical region and coastal exposure do not appear to affect mean levels of abundance. In contrast, treatments lead to pronounced cycles of sea louse infestation with peaks and troughs at 3-week intervals, and these interventions are important if the sea louse levels on fish are to be controlled. There was no evidence of water temperature affecting the mean annual abundance of sea louse infestation.     

Development of vaccines against sea lice

A review of efforts to develop a vaccine against sea lice is presented together with analysis of the rationale behind the approaches and potential future directions. Vaccines against the caligid copepod, Lepeophtheirus salmonis, have the potential to be a cost-effective means of controlling the infection and avoid many of the disadvantages of medicine treatments. However, research towards such vaccines is in its infancy and approaches so far used have met with little or no success. Most strategies for sea louse vaccines have adopted methods used for vaccines against other ectoparasites. A vaccine against the cattle tick (Boophilus microplus) is in field use while other vaccines such as the sheep blowfly (Lucilia cuprina) vaccine are at an earlier stage of development. These haematophageous parasites ingest host antibody as part of a large blood meal which can target antigenic sites in the gut. However, the assumption that arachnid and insect physiology are directly comparable with that of sea lice is not proven, and this may partly explain the slow progress this approach has had with sea lice. Success in developing a louse vaccine will depend upon a better understanding of louse digestive biology, particularly an evaluation of whether the cattle tick model is applicable to the development of a louse vaccine. If the louse gut is to be targeted immunologically, critical antigens will need to be identified and evaluated, bearing in mind that an economic vaccine must include recombinant proteins or be a DNA vaccine. Alternatives to the louse gut as a target are also worth consideration. Antibodies could target critical host-parasite interactions that are amenable to disruption, although no such targets have been identified.     

Differential host utilisation by different life history stages of the fish ectoparasite Argulus foliaceus (Crustacea: Branchiura)

In this study we examine differences in the occurrence of life history stages of the destructive fish ectoparasite Argulus foliaceus (L., 1758) on eight fish species (stickleback, rudd, roach, gudgeon, bream, tench, crucian carp and common carp) sampled from a mixed-species recreational fishing lake on nine occasions during late spring and summer. Total numbers ofA. foliaceus, as well as the number of larval, juvenile and adult parasite stages, from each fish were recorded along with the fish species. Lice generally exhibited an aggregated distribution approximating a negative binomial distribution. Significant differences in the prevalence, intensity and intensity frequency distribution were observed between life history stages and between host species. In general, all life history stages of A. foliaceus exhibited an over-dispersed distribution. However, larval lice did show some degree of aggregation particularly within the stickleback samples. Infection data for parasite larval stages suggested that sticklebacks are more likely to be infected than other host species. For adult lice, however, carp appeared to be the main host. We propose that A. foliaceus infection characteristics are predominantly determined by the level of host exposure to the parasite and its life history stages (larval, juvenile and adult) rather than by an innate difference in host susceptibility related to individual host factors such as immune responses. We conclude that host exposure is determined by the parasite-host behavioural interplay related to species-specific ecology and behavioural traits such as microhabitat preference and normal swimming speed.     

A review of potential pathogens of sea lice and the application of cleaner fish in biological control

There are many examples of successful biological control of pest populations in aquatic environments. This approach to sea louse control has environmental benefits and is cost-effective. The range of possible pathogens of lice is reviewed and epibionts recorded from sea lice, including the monogenean Udonella caligorum and ciliates, are examined. Baculoviruses when ingested by insects form occlusion bodies resulting in severe damage to the digestive system and subsequent death, and this may be a promising approach. Cleaner wrasse (Labridae) have been stocked commercially with farmed salmon since 1989, and recent work on improving the method is reviewed. Wrasse are sourced from a wild fishery and stocked at ratios of 1 to 25-150 salmon. Over 5 million wrasse are stocked annually in Norway and c 30% of smolts in Scotland were stocked with wrasse until 1998, when an outbreak of infections salmon anaemia (ISA) deterred many farmers from transferring wild fish to cages. A case study is given showing that salmon in cages stocked with wrasse had a burden of one to eight lice through the first year compared with up to 40 lice per fish on unprotected and untreated fish. Electivity indices were used to compare the relative composition of lice developmental stages on salmon in stocked and unstocked cages, and adult male and female lice were found to comprise only 6% of the population in cages with wrasse, compared with 49% adults on fish in control cages. Measures to improve the efficacy of wrasse as a way of cleaning salmon in the second production year include the use of refuges to assist over-wintering survival, and stocking ballan wrasse. Health hygiene includes sourcing wrasse in the farm locality, testing for pathogens, vaccination of wrasse and ultimately rearing wrasse for stocking. The role of wrasse in an IPM strategy is described.     

Towards selective breeding of Atlantic salmon for sea louse resistance: approaches to identify trait markers

Several species of sea lice, in particular Lepeophtheirus salmonis (Kr?yer), affect the welfare and condition of farmed salmon, with an estimated annual cost to the Scottish industry of 15-30 Pounds million. In Atlantic salmon, some stocks show resistance to L salmonis. Such natural resistance could be utilized for stock improvement using molecular genetic technologies. The development of molecular markers linked to resistance genes, allowing the identification of resistant fish, could increase the efficacy of selective breeding programmes. Various approaches to achieve this goal are described. One way to identify genes conferring resistance is to develop screens for salmon genes that are activated upon louse infection. One such screen--differential display--requires no previous knowledge of gene sequences, involves no preconceptions about which gene families are involved and can therefore identify novel genes. Preliminary results of comparative gene expression in sea-louse-challenged and control fish illustrate the application of differential display.     

Semiochemical strategies for sea louse control: host location cues

The development of behavioural bioassays and electrophysiological recording techniques has enabled the role of semiochemicals to be investigated for the first time in Lepeophtheirus salmonis (Kr?yer). Adult male sea lice L salmonis were activated by, and attracted to, salmon-conditioned water (SCW) and SCW extract. Non-host fish odours, turbot-conditioned water (TCW) and turbot-conditioned water extract (TCW extract), elicited activation but no attraction in the lice. Solid-phase extraction techniques were developed to extract low molecular weight components of fish odours, and biological activity was shown to be confined to these. Electrophysiological activity from aesthetascs in the first antennae was found and neural responses to odours were recorded from the antennal nerves. Preliminary experiments on individual chemicals (isophorone, 1-octen-3-ol) linked to salmon revealed behavioural activation and electrophysiological responses in adult male L salmonis. Isophorone was shown to be significantly attractive to sea lice when placed in a slow-release system in a perfused tank of seawater in a choice situation. Proof of concept has been established for successfully extracting odour cues from seawater, analysing their biological activity and applying these to slow-release technologies for field trapping of lice. Future work involving linked GC-MS techniques using behaviour, electrophysiological responses, and organolepsis will establish further host location cues specific to Atlantic salmon.     

Monooxygenase mediated pyrethroid detoxification in sea lice (Lepeophtheirus salmonis)

The role of monooxygenases in detoxification of the pyrethroids cypermethrin and deltamethrin was examined. Four strains of sea lice (Lepeophtheirus salmonis Krøyer) with normal or moderately reduced sensitivity towards the pyrethroids were tested in bioassays by exposure to the pyrethroid alone and in combination with an oxygenase inhibitor, piperonyl butoxide (PBO). The normal (baseline) sensitivity was considered as the sensitivity range for the two most sensitive strains. Pre‐treatment with PBO elevated the sensitivity (P < 0.01) compared with groups exposed to the pyrethroid only. A positive, but not statistically significant, correlation between the activity of haem peroxidases and the pyrethroid concentration immobilizing 50% of the parasites was demonstrated (ρ = 0.500 for deltamethrin and ρ = 0.310 for cypermethrin). The results indicate that cytochrome P450 monooxygenases are involved in detoxification of pyrethroids in sea lice. ¹⁴C‐Deltamethrin was absorbed in a lesser amount in a group of sea lice exposed to a mixture of the compound and PBO than in a group exposed to ¹⁴C‐deltamethrin alone. A significant difference could be demonstrated both immediately after exposure (P < 0.01) and 24 h after exposure (P < 0.05). No significant differences were found between groups pre‐treated with PBO and groups exposed to ¹⁴C‐deltamethrin only. ¹⁴C‐Deltamethrin was taken up mainly through the cuticle, especially the cuticle on the extremities of the ventral surface, and subsequently distributed throughout the body of the parasite. Copyright © 2005 Society of Chemical Industry     

Trickle and single infection with Discocotyle sagittata (Monogenea: Polyopisthocotylea): effect of exposure mode on parasite abundance and development

Experimental infection of rainbow trout Oncorhynchus mykiss (Walbaum) with the monogenean Discocotyle sagittata (Leuckart, 1842) allowed comparison between trickle and single exposure, two infection modes demonstrated to occur in the wild. Both types of infection resulted in mean larval attachment success around 50%, which was significantly dependent on dose of infective larvae used (P < 0.0001), but was not affected by mode of infection (P = 0.244). Worms recovered from fish exposed to the same number of oncomiracidia but different mode of infection differed in their rate of development. The developmental stage attained by parasites was significantly affected by number of infective larvae used (P = 0.005), and by the interaction between dose and mode of infection (P = 0.026), suggesting competition among attached larvae. Statistical analysis demonstrated that in the early stages of infestation, worm distribution over the gill arches can be explained by the relative amount of water flowing over them. One, two and three months post-infection parasite numbers were comparable (P = 0.805), but their observed distribution gradually decreased in gill arches III and IV and increased in gill arch I, suggesting that parasites migrate after initial attachment. These results reproduce phenomena observed in the field, indicating that the experimental infection system could be employed to study infection dynamics and host-parasite interactions under controlled conditions.     

Medicines for sea lice

Sea louse (Family Caligidae: genera Caligus and Lepeophtheirus) infection of farmed salmonids represents a significant threat to animal welfare and undermines profitability. Lice may also act as vectors for the transmission of viral and bacterial pathogens. Pest-control programmes parallel those deployed in terrestrial livestock farming and include the use of parasiticides. The authorisation process for fish medicines varies widely between salmon farming countries and undue regulatory constraint may place farmers in one country at a competitive disadvantage. In many jurisdictions, fish are a 'minor' species and mounting demands for environmental assessment increase registration costs. A successful integrated louse-management strategy requires free access to a range of effective, chemically unrelated active ingredients deployed according to current best practice. Over-reliance on a limited number of products will lead, inevitably, to resistance, which is difficult to counter.     

Gill parasites of Cephalopholis argus (Teleostei: Serranidae) from Moorea (French Polynesia): site selection and coexistence

The distribution and coexistence of gill ectoparasites of 121 specimens of Cephalopholis argus Bloch et Schneider, caught between October 1994 and October 1995, were investigated. Adults of the monogenean Benedenia sp. and copepod Hatschekia sp., the larval caligid copepod Caligus sp. (copepodite and chalimus stages), and praniza larvae of the isopod Gnathia sp. were found. All species were aggregated within the host population. Infracommunities were poor, with only 40.5% of fish infected by two parasite species. Only two individual fish harboured all the parasite species observed at the component community level. Prevalences were less than 50% and mean intensities were low (less than 6 parasites/host). No dominant parasite species were observed in the host population. The spatial distribution of each parasite species was studied on different partitions of the gill arches. Adult parasite stages that are mobile showed much overlap in their distribution, whereas temporarily attached larvae of Caligidae were more site specific. Copepodite and chalimus larvae showed niche restriction that is probably due to gregarious behaviour. Positive associations between caligid larvae reflected intraspecific interaction for site and/or resources. Each of the Caligus sp. larval stages prefers specific sites, as do the adults, which occur exclusively in the buccal cavity of the host. Infracommunities were too poor and too few to induce processes of interspecific competition.     

The ecology of Crepidostomum species (Allocreadiidae: Digenea) in the salmonoids of Lake Yli-Kitka in northeastern Finland

Crepidostomum spp. were studied from 417 whitefish (Coregonus acronius Rapp, 1854), 220 vendace (Coregonus albula L.) and 38 brown trout (Salmo trutta m. lacustris (L.)) from Lake Yli-Kitka in northeastern Finland during 1980-1983.12.7% of the whitefish were infected with Crepidostomum with a mean intensity of 4.6 worms per infected fish. The species involvedare C. farionis (Müller, 1784) and Crepidostomum wikgreni Gibson et Valtonen, 1988, which differed from C. farionis in the number and size of the eggs. Both types were encountered in the gall-bladder and intestine, but were unable to be distinguished as immature stages. 21% of the brown trout harboured C. farionis in the intestine with a mean number of 1.5 worms per infected fish. Two C. farionis specimens occurred also in the intestine of one vendace. Whitefish smaller than 150 mm did not harbour Crepidostomum specimens, but in bigger fish the prevalence and the mean intensity of infection increased to a certain limit as the fish got larger. The prevalence of Crepidostomum infection also clearly increased during the three seasons in the three size-groups studied. The seasonal pattern of Crepidostomum in whitefish of the size greater than or equal to 150 mm was variable in that the lowest values, both in the prevalence and mean intensity of infection, occurred between September and December in 1980 and between April and August in 1982. Reinvasion was found throughout the year, although the main recruitment occurred during autumn and early winter. Eggs matured between late spring and early autumn. The smallest worms occurred mainly in the gall-bladder, and most of the gravid specimens were located in the intestine. Both the variance-to-mean ratio and the low values of the parameter k of the fitted negative binomial distribution reveal the Crepidostomum specimens are overdispersed in the whitefish of Lake Yli-Kitka.     

Spatial Aspects of Light Leaf Spot (Pyrenopeziza brassicae) Epidemic Development on Winter Oilseed Rape (Brassica napus) in the United Kingdom

ABSTRACT In microplot experiments in 1998-99 and 1999-2000, the start of light leaf spot epidemics could be predicted from weather data, using empirical equations for Pyrenopeziza brassicae apothecial (ascospore) development, ascospore infection criteria, and the latent period of P. brassicae. The dates when P. brassicae sporulation was first observed fitted predictions and initial spread of light leaf spot from an inoculum source was mostly in the prevailing wind direction, with differences between the two growing seasons attributable to differences in wind patterns. Subsequent secondary spread of disease could be predicted using temperature and rainfall data, and observations fitted predicted dates. In both 1998-99 and 1999-2000, initial spatial patterns of observed disease in January were random, because data were not significantly different from a binomial distribution (P = 0.18). Analysis of spatial data from samples in February and March indicated aggregation, because data fit was significantly different from a binomial distribution (P 相似文献   

Susceptibility to DDT and some other insecticides of head lice (Pediculus capitis De Geer) in Czechoslovakia

For testing the susceptibility of the head louse to insecticides impregnated bunches of polyamide fibres, whose diameter resembled that of human hair (0.1 mm) were used. In this manner a low mortality of control lice collected from infected persons was obtained (2.9% after 16 hours of exposure). Even after 25 years of DDT use no apparent resistance could be demonstrated in 25 tested louse populations and cross resistance to permethrin in 7 tested populations. The diagnostic concentration for resistance to pp'-DDT was established at 1%. Of the tested insecticides the highest toxicity was shown by malathion (LC50 = 0.000 018%), pirimiphos-methyl had on average a twice lower toxicity, trichlorphon 104 times lower, tetramethrin 1220 times lower, permethrin 2.5 times lower and pp'-DDT 630 times lower toxicity.     

Dispersal in patchy environments: effect on the prevalence of small mammal ectoparasites

Part-time ectoparasites on small mammals disperse via the habitat, while full-time parasites spread throughout the host population by direct contacts between host animals. It is therefore supposed that the effect of the natural environment is different for the two groups. This was studied as differences between observed and expected prevalence, the percentage of the infested host population, during different environmental conditions. Two possible mechanisms of such an effect were analysed, i.e. a) host selection and its change with host frequency and b) parasite migration and reproduction rates as reflected by the frequency distribution patterns on the hosts. As expected the prevalence of full-time ectoparasites (Anoplura and subfamily Laelapinae) could be predicted on the basis of host species frequencies during different environmental conditions, with the exception of one louse species (Hoplopleura acanthopus), because of its restricted distribution. Prediction was not possible for part-time ectoparasites (subfamily Hameogamasinae). Species of the subfamily Haemogamasinae were more catholic in host selection than species of Anoplura and subfamily Laelapinae. The haemogamasin mites changed host species to a greater extent than did Anoplura and Laelapinae. All haemogamasin mites had short-tailed frequency distribution patterns and all Anoplura and Laelapinae, except Hyperlaelaps microti, had long-tailed frequency distributions.     

Temporal and spatial distribution of glochidial larval stages of European unionid mussels (Mollusca: Unionidae) on host fishes

Glochidia are the larval stage of freshwater unionid mussels that parasitize the fins and gill apparatus of fish. A total of 22 fish species were examined for the presence of glochidia whose distribution on individual hosts was studied on three common fish species, the roach Rutilus rutilus (L.), perch Percafluviatilis L. and bitterling Rhodeus sericeus (Pallas). Between 1997 and 1999, the fish were obtained from the rivers Morava and Kyjovka and surrounding water pools in the Czech Republic. The glochidia of two genera, Unio and Anodonta, were found. Anodonta glochidia were observed on 10 fish species, Unio glochidia on 17 fish species. There was a difference in spatial distribution of glochidia on the body of the host fish. Unio glochidia were predominantly located on the gills, whereas most Anodonta glochidia were found on the fins, with the highest numbers of glochidia were observed on the margin of the pectoral fins. For the gill apparatus, Unio glochidia were found predominantly on the second and third arch. Anodonta glochidia were predominantly found during winter and spring (November-May), whereas Unio glochidia were more abundant during May and June. The number of glochidia was positively correlated with fish length in perch highly infected by Anodonta glochidia and perch infected by Unio glochidia. Of the three fish species, the highest occurrence of parasites was found on perch with fewer observed on roach. In spite of the close relationship between bitterling and unionid mussels, glochidiosis was rare on this fish species.     

Identification of cDNAs induced by the organophosphate trichlorphon in the parasitic copepod Lepeophtheirus salmonis (Copepoda; Caligidae)

Lepeophtheirus salmonis is a copepod ectoparasite of wild and farmed salmonids and can cause considerable damage to infected fish. We have examined the effect of the organophosphate trichlorphon, which was one of the early treatments for sea lice as Neguvon^® on L. salmonis. Suppression subtractive hybridisation was used to prepare a cDNA library enriched for cDNAs expressed in response to the organophosphate trichlorophon and using this technique a total of 33 cDNAs were found to be differentially regulated in response to organophosphate exposure. Sequence analysis of the cDNAs revealed that many were involved in cellular stress responses and or neurotoxicity. The expression of two of these cDNAs was confirmed to be up-regulated in response to trichlorophon.     

Determination of knockdown resistance allele frequencies in global human head louse populations using the serial invasive signal amplification reaction

BACKGROUND: Pediculosis is the most prevalent parasitic infestation of humans. Resistance to pyrethrin‐ and pyrethroid‐based pediculicides is due to knockdown (kdr)‐type point mutations in the voltage‐sensitive sodium channel α‐subunit gene. Early detection of resistance is crucial for the selection of effective management strategies. RESULTS: Kdr allele frequencies of lice from 14 countries were determined using the serial invasive signal amplification reaction. Lice collected from Uruguay, the United Kingdom and Australia had kdr allele frequencies of 100%, while lice from Ecuador, Papua New Guinea, South Korea and Thailand had kdr allele frequencies of 0%. The remaining seven countries investigated, including seven US populations, two Argentinian populations and populations from Brazil, Denmark, Czech Republic, Egypt and Israel, displayed variable kdr allele frequencies, ranging from 11 to 97%. CONCLUSION: The newly developed and validated SISAR method is suitable for accurate monitoring of kdr allele frequencies in head lice. Proactive management is needed where kdr‐type resistance is not yet saturated. Based on sodium channel insensitivity and its occurrence in louse populations resistant to pyrethrin‐ and pyrethroid‐based pediculicides, the T917I mutation appears to be a key marker for resistance. Results from the Egyptian population, however, indicate that phenotypic resistance of lice with single or double mutations (M815I and/or L920F) should also be determined. Copyright © 2010 Society of Chemical Industry     

Sea louse control in Scotland,past and present

Methods for farming the Atlantic salmon (Salmo salar L) in Scotland were developed at the Unilever Research Laboratory facility at Lochailort on the west coast of Scotland in the late 1960s. By the spring of 1972 a fledgling commercial company had been set up and Marine Harvest, as it was known, received its first year class of smolts at Lochailort. Sea lice (Lepeophtheirus salmonis Kr?yer) from passing wild fish infected the smolts but did not cause any significant damage until the summer of 1976 when intervention was required. The Scottish industry has since grown to become a vital source of employment for fragile rural economies and produced about 130,000 tonnes of salmon in 2000. This paper gives an overview of how fish welfare has been maintained by the development of treatments and management strategies aimed at minimising the impact of sea lice.