Susceptibility of eye fluke-infected fish to predation by bird hosts.

Host manipulation by trophically transmitted parasites may predispose infected hosts to predation and in this way enhance parasite transmission. In most study systems, however, the evidence comes from laboratory studies, and therefore knowledge of the effect of manipulation on parasite transmission efficiency in the wild is still limited. Here we examined the effect ofDiplostomum spathaceum(Trematoda) eye flukes on the susceptibility of fish intermediate hosts to predation by bird definitive hosts. Our earlier studies have shown that the parasite alters fish phenotype and increases their susceptibility to artificial predation under laboratory conditions. In the present field study, we allow…

The structure of parasite component communities in brackish water fishes of the northeastern Baltic Sea

We used nestedness analysis to seek non-random patterns in the structure of component communities of metazoan parasites collected from 31 sympatric fish species from the northeastern Bothnian Bay, the most oligohaline area of the Baltic Sea. Only 8 marine parasite species were found among the 63 species recorded, although some marine fish species reproduce in the bay and others occasionally visit the area. Marine parasite species can utilize both freshwater and marine fish species as intermediate or final hosts, and marine fish can harbour freshwater parasite species. This exchange of parasite species between marine and freshwater fish has probably resulted from ecological factors acting ov…

Influence of rearing conditions on Flavobacterium columnare infection of rainbow trout, Oncorhynchus mykiss (Walbaum).

The influence of rearing conditions on Flavobacterium columnare infection of rainbow trout, Oncorhynchus mykiss (Walbaum), was studied experimentally in the laboratory and at a fish farm. In experiment I, the effect of parasitic infection on columnaris disease was studied using F. columnare carrier fish. The fish were exposed to Diplostomum spathaceum cercariae and a set of other stressors in order to induce clinical columnaris infection. Parasitic infection and other stressors failed to induce the disease. Disease occurred when the fish were challenged with F. columnare, but D. spathaceum infection did not enhance the severity of the infection. In experiment II, the influence of rearing de…

On the infracommunity structure of adult cestodes in freshwater fishes.

The distribution–co-occurrence and exchange of adult cestode species in two fish communities (the Bothnian Bay and Lake Yli-Kitka both in Finland) was studied. Coexistence of two or more mature cestode species in the same fish host population was zero for all fish species studied (33) except pike in the Bothnian Bay and whitefish in the lake. It was found that 60% of the fish species studied in the Bothnian Bay and 80% of the fish species studied from Lake Yli-Kitka harboured only 1 mature cestode species. Exchange of adult cestode species between the different fish species in these two fish communities was found to be as rare as coexistence. The infra-community structure of adult cestodes …

Parasite assemblages of crucian carp (Carassius carassius) – is depauperate composition explained by lack of parasite exchange, extreme environmental conditions or host unsuitability?

Recent studies on parasite communities have considered the factors which contribute to variability in parasite infections among populations of one host species. This paper examined parasite assemblages of crucian carp (Carassius carassius) in a natural set-up of two distinctive environments inhabited by the same host species and differing in respect to composition of other fish species: ponds, where extreme conditions prevent other fish species from occurring and lakes, where crucian carp coexist with other fish species. Our aim was to evaluate the significance of parasite exchange, environmental conditions and host suitability for the depauperate parasite assemblages observed in pond cruci…

Immunostimulants in prevention of columnaris disease of rainbow trout, Oncorhynchus mykiss (Walbaum).

Dactylogyrids on the gills of roach in central Finland: Features of infection and species composition

Monogenean parasites were examined from the gills of 660 roach (Rutilus rutilus) in four interconnected lakes in Central Finland between February and November 1986 and in three of the same lakes between February and December 1988. One of the lakes is eutrophic and polluted due to a paper and pulp mill, one is oligotrophic and in a natural state, and the other two lakes are eutrophic. The prevalence of Dactylogyrus infection was always high. Differences between the lakes and the years were observed in the intensity of infection, which was significantly higher in the polluted lake. The intensity was also higher in older fish. Nine Dactylogyrus species were found, and of these D. crucifer and …

Acanthocephalan size and sex affect the modification of intermediate host colouration

Parasitology, 136 (8)

Seasonal studies of the biology ofAchtheres percarumin perch,Perca fluviatilis, from four Finnish lakes over a 3-year period

Achtheres percarum infection on the gills of perch Perca fluviatilis was studied from four lakes in central Finland in monthly or bimonthly samples of fish during a 3-year period. The highest prevalence and intensity of infection (30%, 0.7 copepods/fish) were found in the oligotrophic, unpolluted Lake Peurunka and the lowest (6%, 0.1 copepods/fish) in the eutrophic and polluted Lake Vatia. According to hierarchical loglinear models the prevalence of A. percarum infection was related to the lake, but not to the year. However, when logit models were constructed such that two ‘seasons’ (‘cold’ and ‘warm’ periods) were included, it was apparent that the year also influenced the prevalence of A.…

Immunoreactivity of Roach,Rutilus rutilus,Following Laboratory Exposure to Bleached Pulp and Paper Mill Effluents

Abstract In order to study immunomodulation, controlled laboratory experiments were carried out with roach ( Rutilus rutilus ) exposed to bleached kraft mill effluent (BKME) or spent bleach liquor (SBL) from two pulp and paper mills using elementary chlorine and chlorine dioxide for bleaching. The total number of immunoglobulin-secreting cells (ISC) and the number of specific antibody-secreting cells (ASC) were determined by ELISPOT assay. Exposure to BKME resulted in decreased numbers of ISC in the spleen. To study the capability of response against foreign antigens the fish were immunized with bovine γ-globulin. Exposure to BKME or SBL reduced antigen-specific ASC response in fish. Moreov…

Seasonality of two gill monogeneans from two freshwater fish from an oligotrophic lake in northeast Finland.

The seasonal occurrence of Dactylogyrus amphibothrium and Discocotyle sagittata from the gills of ruffe (Gymnocephalus cernuus) and whitefish (Coregonus acronius), respectively was studied in Lake Yli-Kitka, a large oligotrophic lake in Northeast Finland. The lake, located near the Arctic Circle, is ice-covered for 7-8 months of the year. The prevalence of D. amphibothrium infection was 70.7% and remained high throughout all size-classes of fish. The length distribution and developmental stages of the worms indicated two generations per year. The overwintering generation produces a summer generation which lives for only a few weeks and matures in July. Discocotyle sagittata has only one gen…

Seasonal dynamics of egg laying and egg-laying strategy of the ectoparasite Argulus coregoni (Crustacea: Branchiura).

Substrate preferences, spatial aggregation patterns and seasonal dynamics in the egg laying of ectoparasiticArgulus coregoniwere studied at a commercial fish farm in Finland. Pilot experiments showed thatA. coregonifemales selected specific types of substrates for egg laying. Significantly moreA. coregonieggs were laid on dark substrates than on light ones suggesting the use of visual cues. Therefore, egg-laying plates of dark colour were constructed for further experiments. MostA. coregonieggs were deposited in locations in shadow and in the deepest water in a 2 m deep farming canal. Relatively more eggs were laid on bottom stones situated near each egg-laying trap than on artificial egg-l…

Furcocercous cercariae (Trematoda) from freshwater snails in Central Finland

AbstractA total of eight species of furcocercous cercariae of four families (Strigeidae, Diplostomidae, Schistosomatidae and Sanguinicolidae), were found in 2005 in Lake Konnevesi in Central Finland in four snail species (Valvata macrostoma, Lymnaea stagnalis, Bathyomphalus contortus and Planorbarius corneus). Australapatemon burti (Miller, 1923), Australapatemon sp., Cotylurus brevis Dubois et Rausch, 1950, Cercaria spinulosa Ginetsinskaya, 1959 and Sanguinicola sp. are new species records for Finland. Ichthyocotylurus variegatus (Creplin, 1825) and Bilharziella polonica (Kowalewski, 1895) were first recorded as cercariae in Finland. The most common cercariae were A. burti (prevalence 13.3…

Ecological divergence of closely related Diplostomum (Trematoda) parasites.

Parasite life-cycles present intriguing model systems to study divergence in resource use and ecology between parasite taxa. In ecologically similar taxa, consistent selective forces may lead to convergence of life-history traits, but resource overlap and similarity of life-cycles may also promote divergence between the taxa in (1) use of host species or (2) specific niche within a host. We studied the life-history characteristics of 2 sympatric species of Diplostomum parasites, D. spathaceum and D. gasterostei, concentrating particularly on differences in intermediate host use and characteristics of the infective stages between the species. This group of trematodes is a notoriously difficu…

Infracommunity level aggregation in the monogenean communities of crucian carp (Carassius carassius).

Aggregation is one of the distinctive features in parasite-host relationships, which has generally been studied at the level of host communities. Parasite aggregation at the infracommunity level may nevertheless be important for intraspecific interactions such as parasite mating success and opportunities for cross-fertilization. In the present paper, we studied the infracommunity aggregation of 3 highly abundantDactylogyrus(Monogenea) species occurring on the gills of crucian carp (Carassius carassius). In line with the previous work on monogenean communities, we observed no competition between the species. At the species level, parasites were distributed unevenly on the gills showing aggre…

Parasite communities as indicators of recovery from pollution: parasites of roach (Rutilus rutilus) and perch (Perca fuviatilis) in central Finland.

We compared parasite communities in fish taken from a polluted lake (L. Vatia) and two control lakes before (1986) and after (1995) nine years of markedly reduced chemical and nutrient loading from a pulpmill in central Finland. Discriminant analyses of the 1995 data, using a function based on the 1986 data, showed that the parasite communities in the fish from the two control lakes had changed relatively little, whereas those from L. Vatia had converged on those from the mesotrophic control lake, indicating substantial recovery from the effects of pollution. Only a few species of parasites provided evidence for recovery. These were anodontid glochidia, which had increased markedly in perch…

Seasonal changes in host phenotype manipulation by an acanthocephalan: time to be transmitted?

Parasitology, 136 (2)

Differences in parasite susceptibility and costs of resistance between naturally exposed and unexposed host populations

It is generally assumed that resistance to parasitism entails costs. Consequently, hosts evolving in the absence of parasites are predicted to invest less in costly resistance mechanisms than hosts consistently exposed to parasites. This prediction has, however, rarely been tested in natural populations. We studied the susceptibility of three naïve, three parasitized and one recently isolated Asellus aquaticus isopod populations to an acanthocephalan parasite. We found that parasitized populations, with the exception of the isopod population sympatric with the parasite strain used, were less susceptible to the parasite than the naïve populations. Exposed but uninfected (resistant) isopods f…

Effects of Pulp and Paper Mill Effluent (BKME) on Physiology and Biochemistry of the Roach (Rutilus rutilus L.)

The effects of bleached kraft pulp and paper mill effluent (BKME) on the roach (Rutilus rutilus L.) were studied under experimental and natural field conditions. In the acute experiment (72 h exposure to the concentrated BKME), the roach suffered from a general stress syndrome, characterized by a significant increase of cortisol and blood glucose, as well as a significant decrease of leucocrit and total plasma protein. In three weeks' exposure in a polluted and an unpolluted lake and in fish caught from the same lakes, the more specific effects of BKME treatments appeared. During the three weeks' exposure, slight hyperglycaemia as well as a decrease in a transaminase activity (GPT) and incr…

Development of monogenean communities on the gills of roach fry (Rutilus rutilus).

The formation and development of monogenean communities on the gills of roach fry was followed in 1992 from early June to October (size range 9 to 47 mm). Roach fry (n=291) were sampled weekly from the small, humic River Rutajoki in central Finland. A further 209 roach fry were reared in a fish farm supplied by water from the river. Four Dactylogyrus species were found: D. nanus, D. crucifer, D. micracanthus and D. suecicus. Other species found on the gills were Gyrodactylus sp. and Paradiplozoon homoion. The first Dactylogyrus juvenile occurred on a 12 mm long fish fry in late June and the first adult (D. nanus) 1 week later in Tank 1. D. nanus was also the most common parasite in the rive…

Eye fluke-induced cataracts in natural fish populations: is there potential for host manipulation?

SUMMARYManipulation of host phenotype (e.g. behaviour, appearance) is suggested to be a common strategy to enhance transmission in trophically transmitted parasites. However, in many systems, evidence of manipulation comes exclusively from laboratory studies and its occurrence in natural host populations is poorly understood. Here, we examined the potential for host manipulation by Diplostomum eye flukes indirectly by quantifying the physiological effects of parasites on fish. Earlier laboratory studies have shown that Diplostomum infection predisposes fish to predation by birds (definitive hosts of the parasites) by reducing fish vision through cataract formation. However, occurrence of ca…

Chondroitin AC lyase activity is related to virulence of fish pathogenic Flavobacterium columnare.

The virulence of eight Flavobacterium columnare strains was studied to find correlations between several virulence-related factors and virulence. Virulence was tested in vivo using rainbow trout, Oncorhynchus mykiss (Walbaum). Suggested virulence-related factors such as production of the degradative enzyme chondroitin lyase, plasmid occurrence and adhesion capability were studied in vitro. Infection with the four most virulent strains resulted in 95-100% mortality within 114 h. Chondroitin lyase activity was found to be significantly related to the virulence of the strains at 25 degrees C and it was also shown to be temperature-dependent, being higher at 25 degrees C than at 20 degrees C. V…

Phenotypic variation in infectivity of Diplostomum spathaceum cercariae within a population.

The present study examined phenotypic variation in infectivity of Diplostomum spathaceum (Trematoda) cercariae within a natural population. Twelve infected Lymnaea stagnalis were collected from the field, and the infectivity of cercariae from individual snails was assessed under constant laboratory conditions. At a water temperature of 16.3 C, the mean infectivity of cercariae from the snails varied between 55.5% and 87.5%. Depending on the source of variation, this may have important ecological and evolutionary implications for both natural parasite populations and those occurring in aquaculture.

Humoral response of roach (Rutilus rutilus) to digenean Rhipidocotyle fennica infection

The humoral immune response of roach (Rutilus rutilus) to cercariae of the digenean trematode, Rhipidocotyle fennica, was studied. Antibodies against R. fennica were found in wild roach in lakes where fish are infected by the parasite. Antibody levels were higher in sera collected in September than in sera collected in June, due to infection of R. fennica during the late summer. In experimental aquarium studies, roach immunized with homogenized cercariae produced antibodies against R. fennica. An especially strong response was elicited by infecting fish with living cercariae emerging from infected clams. The specificity of the antibodies, as shown in Western blots, was different between fis…

Experimental study on the influence of paper and pulp mill effluent on the gill parasite communities of roach (Rutilus rutilus)

SUMMARYRoach (n= 81) caught on 23 May, 1992 from oligotrophic, unpolluted Lake Peurunka were kept in cages over a 2-week period before moving half of the fish to nearby Lake Vatia, which is influenced by pulp mill effluents. Before moving the fish gill parasites were examined from 9 fish; 5Dactylogyrusspecies,Gyrodactylussp. andParadiplozoon homoionwere found, the main components of the infracommunities being dactylogyrids. Afterwards, 5 fish from each lake were studied weekly between 1 July and 17 August. Metazoan parasites were recorded from gill arches divided into 4 sections along the dorso-ventral axis.D. crucifer, D. nanus, D. micracanthusandD. suecicusoccurred in both lakes throughou…

Dactylogyrus (Monogenea) communities on the gills of roach in three lakes in Central Finland

SUMMARYDactylogyrus (Monogenea) species were studied on the gills of roach (Rutilus rutilus) from a total of 293 fish in 3 interconnected lakes in Central Finland. One of the lakes is eutrophic and polluted by a paper and pulp mill, one is eutrophic and one is oligotrophic. Nine Dactylogyrus species were found. The structure of the monogenean communities was analysed at the component and infracommunity level and the Dactylogyrus fauna of the lakes was compared. Although it was found that the component communities tended to be very similar there were also differences. The abundance and diversity were highest in the polluted lake; the infracommunities of this lake were considerably more often…

Fish population size, and not density, as the determining factor of parasite infection: a case study.

The diversity and abundance of parasites vary widely among populations of the same host species. These infection parameters are, to some extent, determined by characteristics of the host population or of its habitat. Recent studies have supported predictions derived from epidemiological models regarding the influence of host population density: parasite abundance and parasite species richness are expected to increase with increasing host population density, at least for directly transmitted parasites. Here, we test this prediction using a natural system in which populations of the crucian carp, Carassius carassius (L.), occur alone, with no other fish species, in a series of 9 isolated pond…

Aggregation ofArgulus coregoni(Crustacea: Branchiura) on rainbow trout (Oncorhynchus mykiss): a consequence of host susceptibility or exposure?

By sampling individual rainbow trout,Oncorhynchus mykiss, at a fish farm we showed thatArgulus coregoniwere aggregated within their host population. The relative significance of susceptibility and exposure generating the observed pattern was tested using experimental infections. We examined, whether rainbow trout developed protective resistance mechanisms against the louse following a challenge infection and if there was variation between individual trout in their susceptibility toA. coregonimetanauplii. Fish were exposed to 20A. coregonifor 5, 25, 50, 85 or 120 min and the numbers attaching recorded. Three weeks later, developing argulids were removed and the experiment repeated with a sta…

Molecular diversity and growth features of Flavobacterium columnare strains isolated in Finland

ABSTRACT: Columnaris disease caused by Flavobacterium columnare is a problem in fish farmingworldwide. During the last 15 yr, outbreaks have started to emerge in Finland. Flavobacteriumcolumnare Type Strain NCIMB 2248 T and 30 Finnish F. columnare isolates were studied usinganalysis of 16S rDNA by restriction-fragment length polymorphism (16S RFLP), length heterogeneityanalysis of polymerase chain reaction (LH-PCR) products, automated ribosomal intergenic spaceranalysis (ARISA), and 16S rDNA sequence analysis. All isolates fell into RFLP Genomovar I and hadthe same length in the LH-PCR analysis. Based on ARISA, 8 genetically different strains wereselected for further analyses. The growth of…

Segregation and co-occurrence of larval cestodes in freshwater fishes in the Bothnian Bay, Finland

SUMMARYTwo autogenic (Triaenophorus crassus and T. nodulosus) and four allogenic (Diphyllobothrium latum, D. dendriticum, D. ditremum and Schistocephalus solidus) larval cestode species were found in 13 out of 31 fish species studied from the Bothnian Bay, NE Baltic. Gasterosteus aculeatus was the most heavily infected fish with 4 larval cestode species; for two of them (D. ditremum and S. solidus) the three-spined stickleback was found to be the required fish intermediate host. Among allogenic cestode species, those restricted to different definitive host species segregated their larval population in relation to the fish host, while, for example, D. ditremum and S. solidus, both maturing i…

Experimental evidence for a hierarchy of mate- and host-induced cues in a fish ectoparasite, Argulus coregoni (Crustacea: Branchiura)

Argulus coregoni is an ectoparasite primarily infesting freshwater salmonids. Sexually reproducing parasites such as A. coregoni are confronted with a dilemma between finding a mate and the costs involved in doing so; if mating partners are unavailable on a host, by leaving to search for a mate on a new host, the parasite is exposed to risks such as predation and energy loss. The utilization of chemical cues could enhance the probability of finding a host and/or a suitable mating partner and thus decrease the level of costs associated with detachment from the host. In this study we constructed a Y-maze arena to determine if adult A. coregoni respond to mate- and host-related chemical cues. …

Parasite-induced aggression and impaired contest ability in a fish host

Abstract Background Success of trophically transmitted parasites depends to a great extent on their ability to manipulate their intermediate hosts in a way that makes them easier prey for target hosts. Parasite-induced behavioural changes are the most spectacular and diverse examples of manipulation. Most of the studies have been focused on individual behaviour of hosts including fish. We suggest that agonistic interactions and territoriality in fish hosts may affect their vulnerability to predators and thus the transmission efficiency of trophically transmitted parasites. The parasite Diplostomum spathaceum (Trematoda) and juvenile rainbow trout, Oncorhynchus mykiss, were used to study whe…

Perch (Perca fluviatilis L.) parasites reflect ecosystem conditions: a comparison of a natural lake and two acidic reservoirs in Finland.

Parasite communities of perch were studied in a natural lake (Vetamajarvi) and two reservoirs (the Kyrkosjarvi and Liikapuro reservoirs) located in Western Finland. All water bodies studied are small, shallow and humic. However, the reservoirs are more acidic than the lake (pH 5.9 and 5.3 vs. 6.4). Altogether, 18 parasite species were found, but the component communities were reduced in the harsh conditions of the reservoirs (12 and six species) as compared with the lake (17 species). In addition, the mean number of metazoan parasite specimens per fish was markedly lower in the reservoirs (64.3 and 14.3 specimens) than in the lake (116.1 specimens). Our prediction on the depauperation of pa…

Host searching in Argulus foliaceus L. (Crustacea: Branchiura): the role of vision and selectivity.

In laboratory experiments, the swimming behaviour of the ectoparasite Argulus foliaceus and its infection rates on juvenile perch (Perca fluviatilis) and roach (Rutilus rutilus) were examined. The highest infection rate and a preference for perch juveniles were obtained in darkness, the lowest infection rate and a lack of preference in the light, when aquaria with glass walls (high reflectivity) were used. In the light, when aquaria were lined with black plastic (low reflectivity) an intermediate level of infection for perch and the highest for roach was recorded. Under such conditions roach were significantly more heavily infected than perch; an attack rate 4 times greater was recorded for…

The influence of food competition and host specificity on the transmission of Triaenophorus crassus (Cestoda) and Cystidicola farionis (Nematoda) to Coregonus lavaretus and Coregonus albula (Pisces:Coregonidae) in Finland.

Abstract As a superior competitor for planktonic food, vendace ( Coregonus albula ), when abundant, is expected to displace whitefish ( Coregonus lavaretus ) from feeding on plankton and to force it to rely more on benthic food. The predicted result would be a reduced abundance of the copepod-transmitted cestode Triaenophorus crassus in whitefish, but an increase in the abundance of the nematode Cystidicola farionis transmitted via benthic amphipods. We studied the occurrence of both parasites in whitefish during 1991–1996 in three interconnected areas at Lake Saimaa, Finland, where the densities of the vendace stocks varied due to natural fluctuation in year-class strengths. In accordance …

Intensive fish farming and the evolution of pathogen virulence: the case of columnaris disease in Finland.

Ecological changes affect pathogen epidemiology and evolution and may trigger the emergence of novel diseases. Aquaculture radically alters the ecology of fish and their pathogens. Here we show an increase in the occurrence of the bacterial fish disease Flavobacterium columnare in salmon fingerlings at a fish farm in northern Finland over 23 years. We hypothesize that this emergence was owing to evolutionary changes in bacterial virulence. We base this argument on several observations. First, the emergence was associated with increased severity of symptoms. Second, F. columnare strains vary in virulence, with more lethal strains inducing more severe symptoms prior to death. Third, more vir…

Intestinal distribution and fecundity of two species ofDiplostomumparasites in definitive hosts

This paper investigated the intestinal distribution and fecundity of 2 species ofDiplostomumparasites,D. spathaceumandD. pseudospathaceum, in 2 species of definitive hosts, herring gull (Larus argentatus) and common gull (L. canus), using both empirical field data and experimental infections. At the level of individual hosts, the parasite species occupied different parts within the intestine, but the fecundity of the worms, measured as the number of eggs in the uterus, did not differ between the parasite species except in wild common gulls. Interestingly, egg numbers in individual hosts were positively correlated between the parasite species suggesting that some birds provided better resour…

Henneguya zschokkei (Myxozoa) infection in cultured whitefish: Age-dependence, seasonality and distribution within host

Abstract Whitefish ( Coregonus lavaretus ) is an important species for European fisheries and aquaculture. Recently, the myxosporean Henneguya zschokkei has been observed to infect the muscles of whitefish in fish farms. Plasmodia of H. zschokkei prevent marketing of the fillet and cause economic losses. The factors associated with occurrence of the parasite in fish farms have not been investigated previously. We studied age-dependence, seasonality, and within-host distribution of H. zschokkei plasmodia in farmed whitefish by examining a total of 1599 fish. Distribution of plasmodia within the fish was not uniform. When the fillet was divided into parts, the number of plasmodia was positive…

Characterisation of the external features ofSchistocephalus solidus (M�ller, 1776) (Cestoda) from different geographical regions and an assessment of the status of the Baltic ringed sealPhoca hispida botnica (Gmelin) as a definitive host

A comparative study of some morphological (segment number, scolex morphology and biometry, length and weight) and biological (maturation in different hosts) features ofSchistocephalus solidus plerocercoids and adults from different geographical regions (Baltic Sea and the British Isles) was carried out. The length of the plerocercoids fromGasterosteus aculeatus was shown to be the variable that best correlated with segment number. A very clear bimodal distribution of segment numbers separated the majority of British and Baltic plerocercoids (British n=21, mean length 25.48, SD 5.63, range 14–34 mm; mean segment number 66.33, SD 8.68, range 51–86. Baltic n=30, mean length 33.23, SD 4.64, ran…

The Effect of Echinorhynchus borealis (Acanthocephala) Infection on the Anti-Predator Behavior of a Benthic Amphipod

In benthic habitats, predators can generally not be detected visually, so olfaction may be particularly important for inducing anti-predation behaviors in prey organisms. Manipulative parasites infecting benthic hosts could suppress these responses so as to increase the probability of predation and thus trophic transmission. We studied how infection with the acanthocephalan Echinorhynchus borealis affects the response of the benthic amphipod Pallasea quadrispinosa to water conditioned by burbot (Lota lota), the parasite's definitive host. In normal lake water, refuge use by infected and uninfected amphipods was similar, but when exposed to burbot-conditioned water, uninfected amphipods spen…

Effect of Pseudomonas sp. MT5 baths on Flavobacterium columnare infection of rainbow trout and on microbial diversity on fish skin and gills

Use of Pseudomonas sp. strain MT5 to prevent and treat Flavobacterium columnare infection was studied in 2 experiments with fingerling rainbow trout Oncorhynchus mykiss. In the first experiment, length heterogeneity analysis of PCR-amplified DNA fragments (LH-PCR) was used to assess the effect of antagonistic baths on the microbial diversity of healthy and experimentally infected fish. In the 148 samples studied, no difference was found between bathed and unbathed fish, and 3 fragment lengths were detected most frequently: 500 (in 75.7% of the samples), 523 (62.2%) and 517 bp (40.5%). The species contributing to these fragment sizes were Pseudomonas sp., Rhodococcus sp. and F. columnare, re…

Being successful in the world of narrow opportunities: transmission patterns of the trematode Ichthyocotylurus pileatus.

SUMMARYParasites with complex life cycles face 2 major challenges for transmission in northern latitudes. They have to cope with the general unpredictability associated with the series of transmission events required for completion of the cycle, and transmission has to be completed within a narrow temporal window because of strong seasonality. Despite this, some parasites show high transmission success, suggesting the operation of effective transmission mechanisms. We explored the transmission of Ichthyocotylurus pileatus (Trematoda) from its snail (Valvata macrostoma) to fish (Perca fluviatilis) hosts by examining some key characteristics in the dynamics of the cercarial emergence from sna…

Accumulation of plerocercoids of Triaenophorus crassus in the second intermediate host Coregonus lavaretus and their effect on growth of the host

Patterns of accumulation of Triaenophorus crassus in its second intermediate host whitefish Coregonus lavaretus s.l. were studied between 1991 and 1996 from two host populations in two separate areas of Lake Saimaa, Finland. Whitefish were infected commonly with several T. crassus plerocercoids and the parasites were aggregated into the oldest hosts. In one host population the annual parasite accumulation was 0·9 parasites in all host age groups between 3 and 8 years. In the other host population the annual accumulation was 1·6 parasites in 3–5-year-old fish, but increased up to 3 to 4 parasites per year in fish over 5 years old. The increase did not coincide with the period of maturation o…

Within-lake dynamics in the similarity of parasite assemblages of perch (Perca fluviatilis).

Although parasite communities have been studied extensively in recent years, spatial and temporal variation in factors affecting the communities has received less attention. This paper examined the similarity of parasite assemblages of perch (Perca fluviatilis) in 18 locations within a single lake in relation to geographical distance and temporal dynamics in the host and parasite populations. We expected that in the present study-scale where distinct but potentially interacting host subpopulations could occur, similarity of the assemblages could be affected by seasonal dynamics in host movements particularly during the spawning period. Parasite species showed differences in infection levels…

Multidimensionality and intra-individual variation in host manipulation by an acanthocephalan

Parasitology, 135 (5)

Spatial and temporal structure of the trematode component community inValvata macrostoma(Gastropoda, Prosobranchia)

SUMMARYWe conducted the first comprehensive study on the spatiotemporal structure of trematode communities in the large-mouthed valve snail,Valvata macrostoma. A total of 1103 snails were examined monthly between May and October 2007 from Lake Konnevesi, Central Finland, from a shallow (1–2 m deep) and an offshore site (5–6 m deep), located ca. 50–70 m apart. Snails were infected by 10 trematode species. The species composition and prevalence were strikingly different between the sites with high species diversity in the shallow site (all 10 species; total prevalence of sporocysts/rediae 12·1%, metacercariae 55·4%) compared to the deeper site (3 species; prevalence 15·0% and 1·9%, respective…

Interspecific associations among larval helminths in fish

Various processes can generate associations between the larvae of different helminth species in their fish intermediate or paratenic host. We investigated the pairwise associations among larval helminth species in eight different fish populations, using two different coefficients of associations, in order to determine in what situations they are strongest. All helminth species included use the fish studied as either their second intermediate host or their paratenic host, and are acquired by the fish when it ingests an infected first intermediate host. The intensity of infection correlated positively with fish length for most helminth species. Pairs of species which both exhibited positive c…

Population structure and recruitment of the ectoparasite Argulus coregoni Thorell (Crustacea: Branchiura) on a fish farm

The population structure and recruitment of Argulus coregoni was monitored at a Finnish fish farm during the open water periods of 1999 and 2001 by weekly sampling of attached argulids. In 2001 the numbers of rainbow trout examined increased in the autumn when the A. coregoni population was declining. When the water temperature exceeded 10 degrees C, at the end of May, A. coregoni egg hatching commenced. A mean number of 98 (S.D. +/- 5.4) juvenile A. coregoni was recorded on each fish, before the start of female egg laying in July 1999. The abundance of lice was lower in 2001. The main recruitment of A. coregoni juveniles occurred in early summer, but the hatching of eggs continued until Se…

Climate warming and disease risks in temperate regions – Argulus coregoni and Diplostomum spathaceum as case studies

AbstractThe link between climate changes and disease risks from various pathogens has been increasingly recognized. The effect of climatic factors on host–parasite population dynamics is particularly evident in northern latitudes where the occurrence and transmission of parasites are strongly regulated by seasonality-driven changes in environmental temperatures. Shortened winter periods would increase growth potential of many parasite populations. The ways in which climate warming could affect life history dynamics of the directly transmitted crustacean ectoparasite Argulus coregoni and complex life cycle trematode Diplostomum spathaceum, which frequently cause problems in northern fish far…

Behavioural adaptations of argulid parasites (Crustacea: Branchiura) to major challenges in their life cycle.

Fish lice (Argulus spp.) are obligate ectoparasites, which contrary to most aquatic parasites, retain the ability to swim freely throughout the whole of their life. In fish farms, they can quickly increase in numbers and without effective control cause argulosis, which results in the reduced growth and survival of their fish hosts. The morphology of Argulus spp, including their sensory organs, is suitable for both parasitism and free-swimming. By spending a considerable amount of time away from their host, these parasites risk being excessively dispersed, which could endanger mating success. Here we present a review of recent studies on the behaviour of Argulus spp, especially the aggregati…

Patterns of cercarial production from Diplostomum spathaceum: terminal investment or bet hedging?

In the production of the infective cercariae of trematodes, the terminal investment hypothesis of life-history theory predicts that the rate of host exploitation and cercarial production should increase during the period of cercarial shedding since the reproductive value of the parasite decreases during this period. In contrast, a bet hedging hypothesis that focuses on the success of transmission when host contact rate is variable predicts that cercarial production should decrease in an attempt to keep the host alive for longer and thus would increase the probability of successful transmission. We examined these two hypotheses under laboratory conditions and recorded the production ofDiplos…

Delayed transmission of a parasite is compensated by accelerated growth.

Compensatory or ‘catch-up’ growth following prolonged periods of food shortages is known to exist in many free-living animals. It is generally assumed that growth rates under normal circumstances are below maximum because elevated rates of growth are costly. The present paper gives experimental evidence that such compensatory growth mechanisms also exist in parasitic species. We explored the effect of periodic host unavailability on survival, infectivity and growth of the fish ectoparasiteArgulus coregoni. Survival and infectivity ofA. coregonimetanauplii deprived of a host for selected time periods were age dependent, which indicates that all metanauplii carry similar energy resources for …

Interspecific and intraspecific interactions in the monogenean communities of fish: a question of study scale?

SUMMARYMonogenean communities of fish have generally been considered non-interactive as negative interspecific interactions have rarely been reported. Most of the earlier studies on monogenean communities, however, have been conducted not only in systems with relatively low parasite abundances but, more importantly, at study scales where microhabitat-level interactions between the parasites are easily overlooked. We examined the communities of 3 abundant Dactylogyrus (Monogenea) species on the gills of crucian carp (Carassius carassius) by analysing the interactions at the scale of individual gill filaments, where interactions between the species, if any, should most likely take place. Cont…

Eye fluke-induced cataract formation in fish: quantitative analysis using an ophthalmological microscope

We examined the parasite-induced cataract formation in rainbow trout (Oncorhynchus mykiss) using slit-lamp microscopy to determine the relationship between cataract intensity and number of Diplostomum spathaceum parasites that were established in the lens. Cataract intensity increased significantly with parasite burden, but was also affected by the pattern of exposure to the parasite cercariae. The slit-lamp methodology proved useful in scoring the cataracts since it provides a 3-dimensional view into the lens and gives an actual picture of the location and intensity of the cataracts, which allows detailed investigations of mechanisms underlying cataract formation in various fish species. P…

Aeromonas salmonicida in Finland: pathological problems associated with atypical and typical strains

. Aeromonas salmonicida was studied at fish farms producing salmonid smolts in northern and central Finland from 1982 onwards. Aeromonas salmonicida subsp. salmonicida has been causing epizootics in salmon, Salmo salar L., and trout, S. trutta L., at two coastal farms in northern Finland sinee 1986, involving 1–29% mortality in the fish-rearing units affceted. The disease causes more serious losses of sea trout yearlings and brood fish than of salmon. The achromogenic atypical A. salmonicida proved to be the most common bacterial disease in brown and sea trout at one farm in northern Finland throughout the period, causing constant heavy losses, mainly of fingerlings, especially in 1982–1986…

The impact of variable hatching rates on parasite control: a model of an aquatic ectoparasite in a Finnish fish farm

Summary 1 Many parasites of medical and veterinary importance infect via free-living stages that undergo a period of development in the environment before seeking a host. Frequently, considerable variation is observed in the development rates of these stages, such that some individuals may be considered to occupy distinct subpopulations, similar to a plant's seed bank. Under certain conditions, these subpopulations may act as a reservoir for the parasite, buffering it from the impact of management strategies and reducing control success. 2 We assessed the impact of intraspecific variation in development rates on parasite control by developing a population dynamic model of the ectoparasite A…

Factors affecting abundance of Triaenophorus infection in Cyclops strenuus, and parasite-induced changes in host fitness.

Factors affecting the abundance of Triaenophorus crassus and Triaenophorus nodulosus procercoids in their copepod first intermediate host, Cyclops strenuus, and effects of infection on feeding behaviour, reproduction and survival of the host were studied experimentally. When exposed to the same number of coracidia, copepods harboured considerably less procercoids in the trials where ciliates or Artemia salina nauplii were given as alternative food items. The prevalence of infection was higher in adult copepods as compared with copepodite stage IV and stage V, and higher in stage V than in stage IV. The prevalences in adult females and males did not differ significantly from each other. The …

Stability of Corynosoma populations with fluctuating population densities of the seal definitive host.

In theory there should be a strong coupling between host and parasite population sizes. Here, we investigated population size and structure in 3 species of acanthocephalans,Corynosoma semerme,C. strumosumandC. magdaleni, in ringed seals (Phoca hispida) from the Bothnian Bay over a period of more than 20 years. During this period, seal numbers first decreased markedly and then increased steadily; at the same time, a paratenic fish host particularly important forC. strumosumhas gradually disappeared from the bay due to decreasing salinity. We found no evidence that the mean abundance of any of the 3 acanthocephalan species changed significantly over time, nor was there any relationship betwee…

QUALITATIVE AND QUANTITATIVE BEHAVIORAL TRAITS IN A COMMUNITY OF FURCOCERCARIAE TREMATODES: TOOLS FOR SPECIES SEPARATION?

Many trematode cercariae show distinct behavioral features, which have commonly been used in species identification in combination with morphological characteristics. However, information regarding cercariae behavior has often not been quantified in detail, or it is scattered in the literature, which is why the appropriate level of precision in behavioral identity, particularly in groups of cercariae species showing considerable morphological overlap, has not been properly established. In this study, we investigated one such group, the furcocercariae trematodes, by studying their behavior in a community consisting of 8 species (Diplostomum pseudospathaceum, Ichthyocotylurus variegatus, Coty…

A link between ectoparasite infection and susceptibility to bacterial disease in rainbow trout

Rainbow trout, Oncorhynchus mykiss, were infected concomitantly with Argulus coregoni and Flavobacterium columnare and their survival was compared with that of fish infected with either the parasite or the bacterium alone. The mortality of fish challenged with A. coregoni was negligible while infection with F. columnare alone led to significantly lower survival. However, compared with single infections, the mortality was significantly higher and the onset of disease condition was earlier among fish, which were concomitantly infected by A. coregoni and F. columnare. This data presents, for the first time, experimental support for the hypothesis that an ectoparasite infection increases suscep…

Dactylogyrus (Monogenea) infections on the gills of roach (Rutilus rutilus L.) experimentally exposed to pulp and paper mill effluent

Experimental exposure to bleached kraft pulp and paper mill effluent (BKME) at a concentration of 10% significantly reduced the abundance and mean number of species of Dactylogyrus on the gills of naturally infected roach, Rutilus rutilus, over a 3 week period. Seven Dactylogyrus species were recorded which differed widely in their susceptibility to the effluent. The experiment coincided with a natural spring peak in dactylogyrid infections on roach. BMKE exposure did not prevent parasite reproduction but post-larval abundance was significantly higher on control than effluent-exposed hosts. There was no evidence of a BKME-induced shift in microhabitat distribution of dactylogyrids. Elevated…

Molecular systematics in the acanthocephalan genus Echinorhynchus (sensu lato) in northern Europe

SUMMARYNew biological species and high levels of inter- and intraspecific genetic divergence were discovered in an allozyme study of some North European members of the acanthocephalan genus Echinorhynchus (sensu lato), parasites of fish and malacostracan crustaceans. (i) A strong differentiation between the marine E. gadi and the fresh- and brackish-water E. salmonis (genetic identity I ≃ 0) supports a generic distinction between these taxa; however, the subdivision would not entirely concur with the concepts of Echinorhynchus (sensu stricto) and Metechinorhynchus suggested earlier. (ii) Samples of E. gadi from the Baltic, Norwegian and North Seas included three distinct, partially sympatri…

Transmission, infectivity and survival of Diplostomum spathaceum cercariae

The transmission dynamics of the cercariae of Diplostomum spathaceum were investigated under laboratory conditions using cercariae collected from naturally infected Lymnaea stagnalis. Cercariae were kept in a constant temperature of 20 °C and the survival and infectivity to naïve young rainbow trout recorded at 3-h intervals until few cercariae were alive. Mortality initially remained constant but increased rapidly after 20 h. While a model of constant mortality fitted the survival data, an age-dependent model provided a better fit and implied that cercariae tended to carry similar quantities of resources and once these were exhausted the cercariae died. Cercarial infectivity also showed an…

Efficacy of emamectin benzoate in the control of Argulus coregoni (Crustacea: Branchiura) on rainbow trout Oncorhynchus mykiss.

Efficacy of in-feed treatment with emamectin benzoate (Slice) for the control of ectoparasitic Argulus coregoni on rainbow trout Oncorhynchus mykiss was tested under laboratory and field conditions. In both experiments fish were fed with fish feed to deliver a therapeutic dose of 0 (control) or 50 microg emamectin benzoate kg(-1) d(-1) (treatment) for a period of 7 d. After 3 d of challenge with A. coregoni in the laboratory, the infestation level in treated fish was lower than that observed in the controls (p0.001). Efficacy of 100% against newly hatched A. coregoni metanauplii and adults and 80% against juveniles was observed. In the field, trial medication was undertaken at 2 sections on…

Diplostomum spathaceum metacercarial infection and colour change in salmonid fish

Colour changes in two salmonid fish, the salmon (Salmo salar) and sea trout (S. trutta), were examined in relation to infection with the trematode Diplostomum spathaceum. This parasite had no effect on the rate of colour change in these fish, although species specific differences in colour adjustment times were observed. Increasing asymmetry in parasite numbers between the right and left eye, which could lead to the retention of vision in one eye, nevertheless tended to reduce the colour change time in salmon with moderate infection (P=0.08). This first experimental attempt to examine colour changes in fish in relation to eye fluke infections provides grounds for future investigations. The …

Host specificity of Argulus coregoni (Crustacea: Branchiura) increases at maturation.

SUMMARYWe tested the hypothesis that host specificity in ectoparasites does not depend exclusively on the features of the host but also on surrounding habitats, using 2 fish ectoparasites, Argulus coregoni and A. foliaceus (Crustacea: Branchiura), occurring sympatrically in Finnish lakes. Although these parasites are considered to be of low specificity, we found that the larger of the 2 species, A. coregoni developed a pronounced preference for salmonid hosts at the beginning of maturation (defined by the presence of copulating specimens). Argulus foliaceus infects a much wider range of fish hosts. We showed that specialization of A. coregoni on salmonids does not necessarily result from in…

The impact of sexual selection on Corynosoma magdaleni (Acanthocephala) infrapopulations in Saimaa ringed seals (Phoca hispida saimensis)

In free-living animals sexual selection is a central force shaping the spatial distribution of individuals in a population as well as sexual size dimorphism. We studied the influence of sexual selection on spatial distribution, female-to-male body size ratio, and female mating success of acanthocephalans in a natural host population of Saimaa ringed seal (Phoca hispida saimensis) harbouring a single intestinal helminth species, Corynosoma magdaleni. The acanthocephalans were always found along the full length of the small intestine; however, the site selection varied among the individual seals according to the age of the infection. The distribution of male acanthocephalans was not random wi…

Parasite resistance and avoidance behaviour in preventing eye fluke infections in fish.

This paper examines the efficiency of acquired resistance in protecting the fish host, rainbow trout (Oncorhynchus mykiss), against the trematode parasite Diplostomum spathaceum, and the hypothesis that fish recognize areas where infective stages are aggregated and show avoidance behaviour. We found that when fish with a low level of infection were held in restricted cages in natural conditions they became infected and developed cataracts as a result of this infection. This suggests that acquired resistance is insufficient in protecting fish against the parasite or the deleterious effects of infection in conditions where fish could not avoid the parasite. Behavioural experiments in the labo…

Treatment of columnaris disease of rainbow trout: low pH and salt as possible tools?

The impact of salt and low pH on columnaris disease of fish was studied. Survival of Flavobacterium columnare after exposure to either 4% NaCl (pH 7.2) or pH 5.0, pH 4.86 or pH 4.6 for 15 min or 1 h was studied in vitro. All conditions significantly reduced the numbers of viable bacterial cells. The effects of salt (4 and 2%) and acidic baths (pH 4.6) were studied in 2 experiments in vivo with rainbow trout Oncorhynchus mykiss infected with F. columnare. Both salt and acidic baths failed to prevent fish mortality; the overall mortality reached 100% in all groups. However, according to survival analysis, the mortality rate was lower in fish treated with 4% salt baths compared to a control gr…

Bioelectronic monitoring of parasite-induced stress in brown trout and roach

A non-contact bioelectronic monitoring system was used to record changes in heart rate, ventilation rate and swimming activity in brown trout Salmo trutta and roach Rutilus rutilus, following exposure to two species of cercariae of digenean trematodes: Diplostomum pseudospathaceum which is a common parasite in the lens of fishes and xiphidiocercariae of Plagiorchis elegans, a parasite of anatid birds, both of which have the snail Lymnaea stagnalis as their first intermediate host. Swimming activity increased significantly in roach exposed to Diplostomum cercariae at densities as low as 3.8l−1 and remained elevated for 24–36 h post exposure. Brown trout showed no response when exposed to low…

Light-mediated host searching strategies in a fish ectoparasite, Argulus foliaceus L. (crustacea: branchiura).

Argulus foliaceus, an obligate fish ectoparasite, can search for its hosts in both light and dark conditions and uses vision in the light. We have examined what searching mode is used at night, when the infection rate was at its highest, and which stimuli produced by the fish are most important. A change of illumination produced a clear difference in the searching behaviour of adult Argulus females. The mean swimming speed and the area explored were 3-4 times higher in the dark, when the parasite employed a cruising search strategy. This changed to an ambush (hover-and-wait) strategy in the light. The swimming activity is accompanied by changes in metabolic costs; the activity of the electr…

Gill anomalies of perch and roach from four lakes differing in water quality

This study examined gill anomalies of two common freshwater fish species, roach Rutilus rutilus and perch Perca fluviatilis, collected over five seasons in 1989–1990 from four lakes in central Finland. The lakes differed both in water quality and in fish parasite species composition. Particular attention was paid to the differences in chloride cell proliferation. Gill anomalies were generally more common and abundant in roach than in perch. Chloride cell proliferation was the most frequent histological change in roach but was that least often found in perch. Most of the changes were systemic and light in severity. In perch an unidentified lesion of noncellular eosinophilic inclusions in an …

Fauna Europaea: Helminths (Animal Parasitic)

The Laotian Rock Rat Laonastes aenigmamus Jenkins, Kilpatrick, Robinson & Timmins, 2005 was originally discovered in Lao People's Democratic Republic in 2005. This species has been recognized as the sole surviving member of the otherwise extinct rodent family Diatomyidae. Laonastes aenigmamus was initially reported only in limestone forests of Khammouane Province, Central Lao. A second population was recently discovered in Phong Nha Ke Bang National Park (PNKB NP), Quang Binh Province, Central Vietnam in 2011. The confirmed distribution range of L. aenigmamus in Vietnam is very small, approximately 150 km , covering low karst mountains in five communes of Minh Hoa District, Quang Binh Provi…