Susceptibility of Baltic and East Atlantic salmon Salmo salar stocks to Gyrodactylus salaris (Monogenea)

The susceptibility of a Baltic salmon stock Salmo salar (Indals?lv, central Sweden) to Norwegian Gyrodactylus salaris (Figga strain, central Norway) was experimentally tested and compared with previously obtained results on East Atlantic salmon (Lierelva, SE Norway). Contrary to expectation, the Baltic salmon, which had no prior exposure to this parasite strain, appeared almost as susceptible as the Norwegian salmon parr that naturally experience G. salaris-induced mortality. Individually isolated salmon of both stocks sustained G. salaris infections with little evidence of innate resistance. A few individuals of the Indals?lv stock controlled their infection from the beginning, but overall there was considerable heterogeneity in the course of infection in both stocks. On individual hosts, G. salaris growth rates declined steadily throughout the infection, a trend which was particularly marked amongst the Lierelva stock. On shoaling Lierelva fish, there was some evidence of reduced parasite population growth towards the end of the infection; this was not apparent in Indals?lv fishes. These results reflect a growing awareness that not all Baltic salmon may be resistant to Norwegian G. salaris, and that Norwegian and Baltic G. salaris strains may differ in virulence. Consequently, management decisions concerning this parasite-host system should be based upon the actual, and tested, susceptibility of stocks under consideration and not upon identification of stocks as either Atlantic or Baltic.     

High Gyrodactylus salaris infection rate in triploid Atlantic salmon Salmo salar

We describe an unusually high infection rate of Gyrodactylus salaris Malmberg in juvenile Atlantic salmon Salmo salar L. of Baltic Sea origin, which are generally believed to be more resistant to G. salaris than East Atlantic salmon populations. Based on analyses of mitochondrial (complete cytochrome oxidase 1 [CO1] gene, 1548 bp) and nuclear (ADNAM1, 435 bp; internal transcribed spacer [ITS] rDNA region, 1232 bp) DNA fragments, the closest relatives of the characterized Estonian G. salaris strain were parasites found off the Swedish west coast and in Raasakka hatchery, Iijoki (Baltic Sea, Finland). Analyses of 14 microsatellite loci of the host S. salarrevealed that approximately 40% of studied fish were triploids. We subsequently identified triploid Atlantic salmon of Baltic origin as more susceptible to G. salaris infection than their diploid counterparts, possibly due to compromised complement-dependent immune pathways in triploid salmon. This is in accordance with earlier studies that have shown elevated susceptibility of triploids to various viral or bacterial pathogens, and represents one of the first reports of increased susceptibility of triploid salmonid fish to an ectoparasite. However, further experimental work is needed to determine whether triploid Atlantic salmon is generally more susceptible to G. salaris compared to their diploid counterparts, irrespective of the particular triploidization method and population of origin.     

Identification of genetic markers associated with Gyrodactylus salaris resistance in Atlantic salmon Salmo salar

Gyrodactylus salaris Malmberg, 1957 is a freshwater monogenean ectoparasite of salmonids, first recorded in Norway in 1975 and responsible for extensive epizootics in wild Atlantic salmon Salmo salar L. The susceptibility of different populations of Atlantic salmon to G. salaris infection differs markedly, with fish from the Baltic being characterised as relatively resistant whereas those from Norway or Scotland are known to be (extremely) susceptible. Resistance to Gyrodactylus infection in salmonids has been found to be heritable and a polygenic mechanism of control has been hypothesised. The current study utilises a 'Quantitative trait loci' (QTL) screening approach in order to identify molecular markers linked to QTL influencing G. salaris resistance in B1 backcrosses of Baltic and Scottish salmon. Infection patterns in these fish exhibited 3 distinct types; susceptible (exponential parasite growth), responding (parasite load builds before dropping) and resistant (parasite load never increases). B1 backcross fish were screened at 39 microsatellite markers and single marker-trait associations were examined using general linear modelling. We identified 10 genomic regions associated with heterogeneity in both innate and acquired resistance, explaining up to 27.3% of the total variation in parasite loads. We found that both innate and acquired parasite resistance in Atlantic salmon are under polygenic control, and that salmon would be well suited to a selection programme designed to quickly increase resistance to G. salaris in wild or farmed stocks.     

Infestations of Atlantic salmon, Salmo salar, by Gyrodactylus salaris in Norwegian rivers

Over the 6 years 1980–85, 212 Norwegian rivers have been examined for occurrence of Gyrodactylus salaris: it was found in 26 rivers and six salmon hatcheries scattered throughout the country from Troms county in the north to Sogn og Fjordane in western Norway. The distribution of G. salaris is connected with the stocking of fish from infected salmon hatcheries. The populations of salmon parr have been drastically reduced in the infected rivers. In later years catches of ascending salmon in these rivers have also sharply declined: in 1984 salmon fishery losses were estimated at 250–500 t.
Gyrodactylus salaris is most probably a recent introduction to Norwegian rivers. A primary aim is to exterminate this parasite from all infected rivers and hatcheries: so far this has been accomplished in one river and one hatchery.     

The ultrastructure of hypersymbionts on the monogenean Gyrodactylus salaris infecting Atlantic salmon Salmo salar

There is increasing pressure to develop alternative control strategies against the pathogen Gyrodactylus salaris, which has devastated wild Atlantic salmon Salmo salar in Norway. Hyperparasitism is one option for biological control and electron microscopy has revealed two ectosymbionts associated with G. salaris: unidentified rod-shaped bacteria, and the protist, Ichthyobodo necator. No endosymbionts were detected. The flagellate I. necator occurred only occasionally on fish suffering costiosis, whereas bacterial infections on the tegument of G. salaris were observed throughout the year, but at variable densities. Bacteria were seldom observed attached to fish epidermis, even when individuals of G. salaris on the same host were heavily infected. Wounds on salmon epidermis caused by the feeding activity of bacteria-infected G. salaris did not appear to be infected with bacteria. On heavily infected gyrodactylids, bacteria were most abundant anteriorly on the cephalic lobes, including the sensory structures, but no damaged tissue was detected by transmission electron microscopy in the region of bacterial adherence. Furthermore, transmission and survival of infected G. salaris on wild salmon did not appear to be influenced by the bacterial infection. The lack of structural damage and impact on G. salaris biology indicates that these bacteria are not a potential agent for control of gyrodactylosis. However, this may not be the case for all gyrodactylid-bacterial interactions and a review of bacterial infections of platyhelminths is presented.     

Increased susceptibility of Atlantic salmon Salmo salar to infections with Gyrodactylus derjavini induced by dexamethasone bath treatment

Dexamethasone, a known immunosuppressant, was administered by bath or injection to Atlantic salmon Salmo salar (Conon stock) to study if this treatment could affect the susceptibility of fish to infections with a Danish strain of Gyrodactylus derjavini (Monogenea). Three groups of S. salar (Conon stock) were immersion treated either with 10, 60 or 240 microg dexamethasone l-1 water, respectively. In addition, one group (positive control) was treated intraperitoneally with 200 microg dexamethasone per fish and one negative control group was kept untreated. A single G. derjavini parasite was placed on the anal fin of each fish and the infection was subsequently monitored weekly for 6 weeks. An increase in parasite populations on the salmon was positively correlated with the amount of immunosuppressant used. Infection levels in the group immersion treated with dexamethasone (240 microg l-1 water) and in the i.p. treated positive control group were significantly higher compared to the untreated control group.     

Isolation of a novel fish thymidylate kinase gene, upregulated in Atlantic salmon (Salmo salar L.) following infection with the monogenean parasite Gyrodactylus salaris

Analysis of differential gene expression in salmon (Salmo salar) blood following infection with the monogenean parasite Gyrodactylus salaris, resulted in the isolation of a thymidylate kinase gene not previously described from fish and which showed similarity to an LPS-inducible thymidylate kinase gene isolated from mouse macrophages. This salmon TYKi-like gene may play a role in an innate generalised response to pathogen infection as it was upregulated in salmon following infection with the parasite, and also in response to injection with the immunostimulants LPS and Poly I:C, used to emulate bacterial and viral infections, respectively. The possible role of this gene in the biosynthesis of mitochondrial DNA in activated macrophages, in response to G. salaris infection is discussed.     

Comparative susceptibility of Atlantic salmon (Salmo salar) to the enteric redmouth bacterium and Aeromonas salmonicida

The bacterium causing enteric redmouth (ERM) and Aeromonas salmonicida were found to be equally pathogenic for fingerling Atlantic salmon (Salmo salar). Injection of 5 x 10(5) cells of ERM or A. salmonicida killed all salmon within 96 h. After a 30 min exposure to water-borne cells of the two test bacteria about one-half of the test salmon died within 14 days. Both ERM and A. salmonicida were transmitted horizontally. Results indicate that efforts should be made to prevent introduction of ERM into watersheds where Atlantic salmon occur.     

Genetic assessment of Atlantic salmon Salmo salar and sea trout Salmo trutta stocking in a Baltic Sea river

Microsatellite DNA variation was used to assess the outcome of stocking Atlantic salmon Salmo salar and migratory trout Salmo trutta in River Sävarå, N Sweden. No information on pre‐stocking genetic composition of S. salar and S. trutta in River Sävarå was available. In 2 year‐classes of S. salar smolt, microsatellite data indicated that post‐stocking genetic composition differed markedly (F_ST= 0·048) from the main donor strain, Byskeälven S. salar, and from other Gulf of Bothnia S. salar stocks (F_ST 0·047 and 0·132). The STRUCTURE programme failed to detect any substructuring within Sävarå salmon. It was concluded that only minor introgression estimated to a proportion of 0·11 (95% CI 0·07–0·16) has occurred in S. salar. Salmo trutta showed overall low differentiation among populations with maximum F_ST of 0·03 making analysis more cumbersome than in S. salar. Still, the SävaråS. trutta deviated significantly from potential donor populations, and STRUCTURE software supported that majority of trout in Sävarå formed a distinct genetic population. Admixture was more extensive in S. trutta and estimated to 0·17 (95% CI 0·10–0·25).     

The existence of two races of Atlantic salmon (Salmo salar L.) in the British Isles

Sera from approximately 10 000 Atlantic salmon collected from rivers in the British Isles have been analysed. Polymorphism at the transferrin locus was observed and the distribution of the transferrin alleles provides further evidence supporting the presence of two races of salmon first postulated by Payne et al. (1971 a ).     

Variation in the timing of river entry of Atlantic salmon (Salmo salar L.) in the Baltic

The timing of river entry in the Atlantic salmon is known to depend on genetic,demographic and environmental factors,but little is known about the relative magnitude of among population and among year variation and covariation in this respect in natural state Atlantic salmon rives.To investigate this,variability in the timing of river entry in three historical Finnish Atlantic salmon populations were analyzed using salmon trap data collected during 1870-1902.The analyses reveled that 1)the timing of river e...     

The susceptibility of grayling (Thymallus thymallus) to experimental infections with the monogenean Gyrodactylus salaris

The pathogenic monogenean Gyrodactylus salaris infecting Atlantic salmon (Salmo salar) is found to attach and reproduce under laboratory conditions on several species in the subfamily Salmoninae other than the Atlantic salmon. The gyrodactylid species Gyrodactylus thymalli infecting grayling (Thymallus thymallus) in another subfamily, Thymallinae, is previously said to be very similar to G. salaris based on morphometry and genetical analysis which prompted the present laboratory experiments to test the susceptibility and resistance of grayling to G. salaris. All 0+ and 1+ grayling became infected with G. salaris during the experimental infection procedure. However, both innate resistant and susceptible grayling were found. In susceptible individually isolated fish, parasite reproduction lasted for more than 35 days. Parasite reproduction also occurred among grouped grayling as judged from the duration of infection of more than 50 days. However, grayling susceptibility as judged from G. salaris reproduction, was very limited. Hence, the results indicate significant biological differences between the function of Atlantic salmon and grayling as host for G. salaris. The grayling is interpreted as unable to sustain G. salaris in nature which implies that G. thymalli is not conspecific with G. salaris. However, G. salaris dispersal by grayling cannot be excluded.     

Post-smolt migration of Atlantic salmon, Salmo salar L., from the Simojoki river to the Baltic Sea

The migration pattern of Atlantic salmon ( Salmo salar L.) post-smolts in the Baltic Sea was investigated based on tag recoveries of Carlin-tagged wild and hatchery-reared smolts released in the Simojoki river in 1972–2005. Exact date of sea entry was known only for the wild smolts. Tag recoveries of wild salmon in the estuary within 10 km from the river mouth were received on average 3.5 days (±2.0 SD) after release. Time required for emigration from the estuary was dependent on the sea surface temperature (SST) off the river ( R ² = 0.625, P = 0.004), being shorter in years with warmer than colder sea temperatures. Outside the estuary, the wild and hatchery-reared post-smolts migrated southwards along the eastern coast of the northern Gulf of Bothnia, the tag recoveries coinciding with the warm thermal zone in the SST occurring along the coastal area. After arriving in the southern Gulf of Bothnia in late summer the post-smolts mostly migrated near the western coast, reaching the Baltic main basin in late autumn. The relationships between the marine conditions and migration patterns are discussed.     

Variations of opisthaptoral hard parts of Gyrodactylus salaris Malmberg, 1957 (Monogenea: Gyrodactylidae) on parr of Atlantic Salmon Salmo salar L. in laboratory experiments

On parr of Atlantic salmon Salmo salar L. in laboratory experiments at different water temperatures, opisthaptoral hard parts of Gyrodactylus salaris Malmberg, 1957 differed in size. Their shape, on the other hand, varied only slightly. The opisthaptoral hard parts were largest at the lowest water temperatures and decreased in size with increasing water temperatures (1.5°C to 20.0°C). At both 18.0°C and 20.0°C the opisthaptoral hard parts of G. salaris were smaller than has been found under natural conditions. In the experiments, the mean size of the opisthaptor in the G. salaris micropopulation at each water temperature gradually increased or decreased compared to the mean size at the start of the experiments. At the end of the experiments the mean size of the opisthaptoral hard parts in the G. salaris micropopulations showed a significant regression to the different water temperatures.     

Seasonal dynamic of the infestation of young landlocked salmon (Salmo salar morpha sebago, Girard) by the monogenean species Gyrodactylus salaris

Seasonal dymanic of the infestation of young landlocked salmon (Salmo salar morpha sebago, Girard) by the monogenean species Gyrodactylus salaris Malberg, 1957 was studied in the river Lizhma, Karelia. It is established, that the temperature optimal for the development and reproduction of the parasite is about 3-8 degrees C. It is stated, that the abundance of G. salaris in the river Lizhma do not reach so high values, as it was observed in the regions outside the natural area of the parasite.     

Evidence for an autumn downstream migration of Atlantic salmon Salmo salar (Linnaeus) and brown trout Salmo trutta (Linnaeus) parr to the Baltic Sea

In the eastern Baltic rivers, anadromous salmonid parr are known to smoltify and migrate to the sea from March until June, depending on latitude, climate and hydrological conditions. In this study, we present the first records of autumn descent of brown trout Salmo trutta and Atlantic salmon Salmo salar from the Baltic Sea Basin. Otolith microchemistry analyses revealed that these individuals hatched in freshwater and had migrated to the brackish water shortly prior to capture. The fish were collected in 2006, 2008, 2009 and 2013 from Eru Bay (surface salinity 4.5–6.5 ‰), Gulf of Finland. This relatively wide temporal range of observations indicates that the autumn descent of anadromous salmonids is not a random event. These results imply that autumn descent needs more consideration in the context of the effective stock management, assessment and restoration of Baltic salmonid populations and their habitats.     

The susceptibility of Salvelinus fontinalis (Mitchill) to Gyrodactylus salaris Malmberg (Platyhelminthes; Monogenea) under experimental conditions

The host specificity of Gyrodaclylus Solaris is examined experimentally with respect to its ability to infect the brook trout, Salvelinus fontinalis . The parasite readily attached to and reproduced on parr of this host and infections grew for c. 20 days from first monitoring (c. 30 days from first infection) before declining. Parasites could persist on this host for up to 70 days before finally disappearing. The pattern of infection resembled that seen in many other gyrodactylid species on their normal hosts, and suggested the action of a host response, In this respect infections of G. salaris on parr of S. fontinalis , anadromous Salvelinus alpinus, Oncorhynchus mykiss, Thymallus thymallus and Baltic Salmo salar follow a normal pattern, while infections of Norwegian S. salar are unusual in a continued unchecked growth, until the host dies, under pooled laboratory conditions.