Gyrodactylus salaris infecting allopatric Arctic charr Salvelinus alpinus fry: an experimental study of host survival

Artificially reared Arctic charr Salvelinus alpinus fry (post yolk‐sac stage) from Skibotnelva in northern Norway were exposed to infection with the monogenean ectoparasite Gyrodactylus salaris to determine if the parasite caused mortality in fry. About 95% of the fry became infected within the first week post‐exposure, and at the end of the experiment (77 days) all fry carried the parasite. Mortality rate was significantly higher in the groups of infected fry than in the uninfected control groups, and when the experiment was terminated c. 30% of the infected and 8% of the uninfected fish had died. Gyrodactylus salaris had a pronounced negative effect upon fry survival and caused an additive host mortality. Moreover, the concurrent temporal changes in parasite intensities and aggregation indicated that the parasite‐induced host mortality was density dependent.     

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.     

Infestations by Gyrodactylus sp. of Atlantic salmon, Salmo salar L., in Norwegian rivers

Young Atlantic salmon, Salmo salar , brown trout, Salmo trutta , and Arctic char, Salvelinus alpinus , from eight rivers in North and Mid Norway were examined for Gyrodactylus. The fish were collected from 1975 to 1980. A Gyrodactylus salaris type was observed infecting salmon from six of these rivers. No trout or char were infected. A high frequency and intensity of infection of salmon were observed in all but one of the rivers surveyed. In the Saltdalselva, only one specimen of Gyrodactylus infecting one fish was observed. No subsequent mortality of salmon was observed in this river while there were signs of a high mortality of salmon in the other rivers. The salmon parr were more frequently attacked than the fry, the mortality of salmon seemed to have happened in most of the rivers one year after the first observations of Gyrodactylus were made. The mortality of salmon and the high infection rate of Gyrodactylus in these rivers appears unique, and as far as known to the authors there are no other described cases of mortality due to Gyodactylus in Atlantic salmon in natural waters. The reasons for the outbreak of Gyrodactylus in these rivers are not known. Two theories are discussed: one that the fish were weakened by environmental factors and the other that Gyrodactylus was introduced from some of the infected salmon hatcheries in Scandinavia.     

Differences in the host resistance of Atlantic salmon, Salmo salar L., stocks to the monogenean Gyrodactylussalaris Malmberg, 1957

The susceptibility and resistance of hatchery-reared salmon parr, native to the rivers Neva (U.S.S.R. Baltic Sea), Alta (northern Norway) and Lone (western Norway) (both eastern Atlantic Ocean), to Gyrodactylus salaris from Norway, was examined. The level of resistance to the parasite was assessed from counts, made on anaesthetized salmon, ofthe numbers of G. salaris after an initial experimental exposure (2 weeks) to G. salaris-infected salmon. Three experiments, all in water at c. 12° C, were carried out: (1) 50 Alta and 50 Neva salmon, initial mean parasite intensity c. 12; (2) 50 Lone and 50 Neva salmon, initial mean parasite intensity c. 60; (3) 10 Lone and 10 Neva salmon individually isolated, initial intensity one gravid G. salaris . In both the Norwegian salmon stocks, the G. salaris infrapopulations steadily increased during the experimental period of 5 weeks, in contrast to a prominent decline in the Neva salmon stock, after, respectively: (Exp. 1) week 3, average peak intensity 32.6; (Exp. 2) week 2, average peak intensity 58.7; and (Exp. 3) week 3, average peak intensity 6.3. The hatchery-reared Baltic Neva stock demonstrated both an innate and an acquired resistance towards G. salaris , in contrast to the highly susceptible, Norwegian Alta and Lone salmon stocks.     

Differences in susceptibility of anadromous and resident stocks of Arctic charr to infections of Gyrodactylus salaris, under experimental conditions

The ability of Gyrodactylus salaris , an important pathogen of the Atlantic salmon Salmo salar , in Norway, to infect anadromous and resident stocks of the Arctic charr, Salvelinus alpinus , has been examined in the laboratory. Resident charr (Korssjoen stock) exposed to heavily infected salmon, were considered innately resistant as they lost their infections within 21 days when individually isolated. Isolated anadromous harr (Hammerfest stock) remained infected for up to 150 days, although most infections disappeared within 30–50 days. In many cases the parasite population grew initially, but growth was limited after 20–30 days and infections subsequently disappeared. At the same time, shoals of 50 anadromous charr, swimming in the tanks containing the individually isolated fish in floating cages, remained infected for up to 280 days. Charr isolated from these shoals after 115 days and subsequently monitored individually lost their infections within 30 days, although the parasite persisted within the shoals for a further 75–135 days. This suggests that G. salaris , persisted on shoaling charr despite an immune response which led to the elimination of parasites from isolated hosts. The Hammerfest stock of anadromous charr supports G. salaris , in the laboratory, and the extended period of survival on this host suggests that charr may be important in the epidemiology of G. salaris , in northern Norway.     

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.     

Seasonal variations in the prevalence and infestation intensity of Gyrodactylus salaris Malmberg, 1957 (Monogenea: Gyrodactylidae) on Atlantic salmon parr, Salmo salar L., in the River Batnfjordselva, Norway

The prevalence of Gyrodactylus suluris Malmberg. 1957 on both yearlings and older parr of Atlantic salmon ( Sulmo sulur L.) in the River Batnfjordselva was 100% through most of the year. With one exception, uninfested fish were only found in the winter and spring after the water temperature had fallen to almost 0° C for 2–3 months. In general, the abundance (i.e. the mean number of parasites per investigated fish) of G. salaris increased during the warm period of the year (summer and early autumn). Abundanceas high as 1153 and 4418 in early autumn was found on yearlings and older parr, respectively. The abundance decreased during the cold period of the year (winter and early spring), in some cases to as low as two and four G. salaris on yearlings and older parr, respectively. About 86% of the G. salaris specimens were found on the fins of the salmon parr: mainly on the dorsal fin (34.4%) and the pectoral fins (27.0%). The remainder of the parasites were distributed on the body (7.8%), the head (3.5%), and the gills (2.6%).     

Characterisation of a low pathogenic form of Gyrodactylus salaris from rainbow trout

Gyrodactylus salaris was isolated from rainbow trout in a Danish freshwater trout farm, and a laboratory population of this particular parasite form was established on rainbow trout. Challenge infections were performed using different salmonid strains and species, including East Atlantic salmon Salmo salar (from the Danish River Skjern?), Baltic salmon S. salar (from the Swedish River Ume Alv) and rainbow trout Oncorhynchus mykiss (from the Danish rainbow trout farm Fousing). These were compared to infection studies on the Norwegian Laerdalselva parasite form kept under exactly the same conditions in the laboratory. The Danish G. salaris form had low virulence towards both Atlantic and Baltic salmon, whereas rainbow trout proved susceptible to the parasite. The Danish G. salaris form was able to maintain a very low infection on East Atlantic salmon, but not on the Baltic salmon, which eliminated the infection within 2 wk. Rainbow trout developed infection intensities ranging up to several hundred parasites per host. The host colonization patterns of the parasite differed clearly from those of previous studies on microhabitats of the Norwegian form of G. salaris. A comparative study on morphological characters (opisthaptoral hard parts) from the Danish parasite form and Norwegian G. salaris showed no significant differences. Selected genes comprising internal transcribed spacers 1 and 2 (ITS), ribosomal RNA intergenic spacer (IGS) and cytochrome c oxidase subunit I (COI) regions were cloned and sequenced. Five sequenced ITS clones from 5 individuals of the Danish strain consistently revealed a single base substitution compared to ITS sequences from all other known species and strains of Gyrodactylus. Mitochondrial COI gene sequences demonstrated that the Danish G. salaris form is closely similar to the Laerdalselva parasite form found in Norway. The IGS sequences were highly variable, but very similar to those obtained from German isolates of G. salaris.     

The critical thermal limits for juvenile Arctic charr Salvelinus alpinus

The chief objective was to determine the critical thermal limits for alevins, fry and parr of Arctic charr, Salvelinus alpinus , (L.) from four races living in Windermere (northwest England). The experimental fish were reared in a hatchery but were the progeny of wild parents. As comparisons between tethal temperatures at four acclimation temperatures (5, 10, 15, 20° C) revealed few significant racial differences, the data were pooled to estimate the lethal values for survival over 7 days (incipient lethal temperature) and over only 10 min (ultimate lethal temperature) for each life stage. Upper lethal values increased with acclimation temperatures for alevins but this effect was negligible for fry and parr, Alevins were generally less tolerant than fry and parr at lower, but not higher, acclimation temperatures; e.g. after acclimation at 5° C, mean upper ultimate values were 23·3, 25·1 and 25·7° C and mean upper incipient values were 18·7, 21·5 and 21·5° C for alevins, fry and parr respectively; after acclimation at 20° C, mean upper ultimate and incipient values were 26·2, 26·1 and 26·6° C and 20·8, 20·8 and 21·6° C for alevins, fry and parr respectively. The area of the temperature tolerance polygon (expressed as ° C²) for juvenile Arctic charr is amongst the lowest recorded for salmonids; being 409, 439 and 461° C² for alevins, fry and parr respectively. These low values are due to lower upper tolerance limits, not high lower tolerance limits; the latter being close to 0° C (<1°C for parr and fry, <0·3° C for alevins) at all acclimation temperatures. Arctic charr are therefore amongst the least resistant of salmonids to high temperatures but probably the most resistant to low temperatures.     

Mitochondrial DNA variation of Gyrodactylus spp (Monogenea, Gyrodactylidae) populations infecting Atlantic salmon, grayling, and rainbow trout in Norway and Sweden

Approximately 800 bp of the mitochondrial cytochrome oxidase I (COI) gene were sequenced from 76 Gyrodactylus specimens of 32 salmonid host populations, i.e. from Salmo salar, Thymallus thymallus, and Oncorhynchus mykiss in Norway, Sweden and Latvia. The COI sequences indicated a substantial intraspecific differentiation of Gyrodactylus salaris and Gyrodactylus thymalli. In total, 12 haplotypes were identified which group into five well supported clades, three clades with parasites from Atlantic salmon and two clades with parasites from grayling. The basal nodes linking the five clades together are only weakly supported. Thus, there is no support for the monophyly of all G. salaris haplotypes and the monophyly of all G. thymalli haplotypes. The lack of monophyly of the mitochondrial haplotypes of G. salaris and G. thymalli may indicate that G. salaris and G. thymalli represent (i). two polytypic species or (ii). one polytypic species, or (iii). refer to a complex of more than two sibling species. The mtDNA data indicate multiple introductions of G. salaris and G. thymalli into Norway. A minimum of three independent introductions of G. salaris and two independent introductions of G. thymalli are supported. This is congruent with earlier hypotheses on the introduction of G. salaris and G. thymalli into Norway.     

Molecular faunistics of accidental infections of <Emphasis Type="Italic">Gyrodactylus</Emphasis> Nordmann, 1832 (Monogenea) parasitic on salmon <Emphasis Type="Italic">Salmo salar</Emphasis> L. and brown trout <Emphasis Type="Italic">Salmo trutta</Emphasis> L. in NW Russia

Salmon Salmo salar L. and brown trout S. trutta L. juveniles were examined for the presence of accidental monogenean ectoparasitic species of Gyrodactylus Nordmann, 1832 in the Baltic and White Sea basins of Russian Karelia in order to estimate the frequency of host-switching attempts on an ecological timescale. To collect phylogeographical information and for exact species identification, the parasites were characterised by nuclear internal transcribed spacer sequences of rDNA (ITS) and, for some species, also by their mitochondrial DNA (CO1 gene) sequences. Four accidental Gyrodactylus species were observed on salmon and brown trout. A few specimens of G. aphyae Malmberg, 1957, the normal host of which is the Eurasian minnow Phoxinus phoxinus (L.), were observed on lake salmon from the Rivers Kurzhma (Lake Kuito, White Sea basin) and Vidlitsa (Lake Ladoga, Baltic basin). G. lucii Kulakovskaya, 1952, a parasite of the northern pike Esox lucius L., was observed on salmon in the Kurzhma. In the River Vidlitsa, two specimens of G. papernai Ergens & Bychowsky, 1967, normally on stone loach Barbatula barbatula (L.), were found on salmon. On anadromous White Sea salmon in the River Pulonga in Chupa Bay, a few salmon parr carried small colonies of G. arcuatus Bychowsky, 1933, which were shown to have originated from the local three-spined stickleback Gasterosteus aculeatus L. consumed as prey. No specimens of Gyrodactylus salaris Malmberg, 1957 were observed, although the Pulonga is the nearest salmon spawning river to the River Keret', which is heavily infected with introduced G. salaris. In the River Satulinoja, Lake Ladoga, three specimens of G. lotae Gusev, 1953, from burbot Lota lota (L.), were collected from a single brown trout S. trutta. All nonspecific gyrodactylid infections on salmonids were judged to be temporary, because only a few specimens were observed on each of the small number of infected fishes. The prevalence of endemic G. salaris was also low, only 1% (Nfish = 296) in Lake Onega and 0.7% (Nfish = 255) in Lake Ladoga, while brown trout specific Gyrodactylus species were not observed on any of the 429 trout examined from the Ladoga basin. The host-specific and unspecific burden of Gyrodactylus spp. on these 'glacial relict' populations of salmon and brown trout was very low, suggesting a generalised resistance against the co-evolved freshwater parasite community, or some kind of 'vaccination' effect. These hypotheses deserve further testing.     

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.     

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.     

Temperature requirements of Atlantic salmon Salmo salar, brown trout Salmo trutta and Arctic charr Salvelinus alpinus: predicting the effects of climate change

Atlantic salmon Salmo salar, brown trout Salmo trutta (including the anadromous form, sea trout) and Arctic charr Salvelinus alpinus (including anadromous fish) provide important commercial and sports fisheries in Western Europe. As water temperature increases as a result of climate change, quantitative information on the thermal requirements of these three species is essential so that potential problems can be anticipated by those responsible for the conservation and sustainable management of the fisheries and the maintenance of biodiversity in freshwater ecosystems. Part I compares the temperature limits for survival, feeding and growth. Salmo salar has the highest temperature tolerance, followed by S. trutta and finally S. alpinus. For all three species, the temperature tolerance for alevins is slightly lower than that for parr and smolts, and the eggs have the lowest tolerance; this being the most vulnerable life stage to any temperature increase, especially for eggs of S. alpinus in shallow water. There was little evidence to support local thermal adaptation, except in very cold rivers (mean annual temperature <6·5° C). Part II illustrates the importance of developing predictive models, using data from a long-term study (1967-2000) of a juvenile anadromous S. trutta population. Individual-based models predicted the emergence period for the fry. Mean values over 34 years revealed a large variation in the timing of emergence with c. 2 months between extreme values. The emergence time correlated significantly with the North Atlantic Oscillation Index, indicating that interannual variations in emergence were linked to more general changes in climate. Mean stream temperatures increased significantly in winter and spring at a rate of 0·37° C per decade, but not in summer and autumn, and led to an increase in the mean mass of pre-smolts. A growth model for S. trutta was validated by growth data from the long-term study and predicted growth under possible future conditions. Small increases (<2·5° C) in winter and spring would be beneficial for growth with 1 year-old smolts being more common. Water temperatures would have to increase by c. 4° C in winter and spring, and 3° C in summer and autumn before they had a marked negative effect on trout growth.     

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.     

Susceptibility of brown trout to Gyrodactylus salaris (Monogenea) under experimental conditions

Brown trout ( Salmo trutta ) from anadromous River Lierelva, resident Lake Tunhovd, and resident Nordmarka stocks were exposed to Gyrodactylus salaris -infected salmon parr. The brown trout were fed pellets before the experiments, except for one group of the Nordmarka stock which was starved for 19 days before the experiments. The mean number of parasites declined directly and rapidly post infection for all groups of trout. There were no pronounced differences in resistance between the anadromous and the resident stocks. G. salaris infections tended to persist longer on starved than on fed trout of the Nordmarka stock. The maximum parasite persistence on trout was 50 days, and as parasite numbers increased on some fish parasite reproduction must have occurred on those trout. However, the limited susceptibility and marked innate resistance of trout to G. salaris establishment, development and reproduction, suggest parasite metapopulations will not survive on this species. Nevertheless, trout may still play a role in the dispersal of G. salaris within and between rivers.     

Host specificity dynamics: observations on gyrodactylid monogeneans

The directly transmitted viviparous gyrodactylids have high species richness but low morphological and biological diversity, and many species are recorded from only a single host. They therefore constitute a guild of species ideal for studies of the evolutionary significance of host specificity. The group has the widest host range of any monogenean family, being found on 19 orders of bony fish. However, individual species range from narrowly specific (71% of 402 described species recorded from a single host) to extremely catholic (Gyrodactylus alviga recorded from 16 hosts). Gyrodactylid-host interactions extend from 60 mya (G. lotae, G. lucii) down to 150 years (G. derjavini on Oncorhynchus mykiss). Co-evolution with the host is comparatively rare within the gyrodactylids, but host switching or ecological transfer is common, and has been facilitated by the mixing of fish strains that followed glaciation. In this review, we consider the factors responsible for gyrodactylid specificity patterns, using examples from our work on salmonid gyrodactylids including G. salaris, responsible for major epidemics on wild Atlantic salmon (Salmo salar) in Norway since 1975, and G. thymalli from grayling and G. derjavini from trout.G. salaris has a wide host range with highest population growth rates on Norwegian salmon strains. However, growth rates are variable on both host strains and species, because of the multitude of micro- and macro-environmental factors influencing parasite mortality and fecundity. A better predictor of performance is the proportion of fishes of a strain which are innately resistant to the parasite, a measure which is negatively correlated with the time to peak infection in a host strain. Population growth rate is also negatively correlated with age of infection; the initial rate, therefore, predicts best the suitability of a fish as host for G. salaris. The host response to gyrodactylids appears to be the same mechanism in all salmonids with innate resistance as one end of a spectrum, but influenced by stress and probably under polygenic control. Hybrid experiments show that performance of G. salaris on a host is heritable, and usually intermediate between that of the parents. This host response mechanism, coupled with the initial parasite population growth on a fish, determines the host specificity, i.e. whether the fish will be susceptible, a responder or innately resistant. The use of population growth rate parameters allows comparison of different hosts as a resource for a gyrodactylid. In the case of G. salaris, East Atlantic and Baltic strains of Atlantic salmon are core hosts, but other salmonids can physiologically sustain infections for considerable periods, and may be important in parasite dispersal and transmission. A further group of non-salmonid fishes are unable to sustain G. salaris reproduction, but can act as transport hosts.Population growth parameters are very labile to stressors and environmental factors, particularly temperature and salinity, and also other aspects of host ecology and water quality. These factors may also influence the spectrum of hosts that can be infected under particular conditions, and probably favoured ecological transfer of gyrodactylids between host species in periglacial conditions. G. salaris may still be undergoing post-glacial range expansion (aided by anthropogenic spread) as shown by the increase in the species range over the last 25 years. The origin of G. salaris, G. teuchis and G. thymalli is discussed in relation to glacial refugiums during the last ice age.     

Carbohydrate localization on Gyrodactylus salaris and G. derjavini and corresponding carbohydrate binding capacity of their hosts Salmo salar and S. trutta

The congeners Gyrodactylus salaris and G. derjavini are specific ectoparasites of Atlantic salmon Salmo salar and brown trout S. trutta, respectively. To elucidate the involvement of lectin-carbohydrate interactions in this host specificity, carbohydrates on the tegument of the two species and the corresponding lectin activity of their hosts have been studied. Carbohydrate composition on the tegument differed significantly between the two gyrodactylids. Three of four commercially available peroxidase-labelled lectins with primary affinity towards D-mannoside, D-GalNAc and L-fucose bound more strongly to G. derjavini than to G. salaris. Lectins with an affinity towards D-mannoside and D-GalNAc bound significantly stronger to the cephalic lobes on G. derjavini compared to the tegument and sheaths of the hamuli. One brown trout strain and three different salmon strains were tested for lectin activity in skin and plasma. Two Baltic salmon strains and one strain from the Atlantic region were included. Brown trout differed significantly from the salmon strains when skin samples were tested for D-GalNAc activity. Lectins binding to other carbohydrates showed trends for similar host differences. The implications of carbohydrate-lectin interactions for host specificity in gyrodactylids are discussed.     

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.