Thermal effects on the dynamics of infection with metacercariae of Diplostomum baeri and D. spathaceum were monitored between May and September, in a year-class of perch Perca fluviatilis, in an artificial lake receiving warm water discharges from a nuclear power station and in an unheated reference site, for a two year period. In the heated area the prevalence of infection of the retinal form, D. baeri, was always 100%, whereas in the unheated site there was an increase from 93% in May to 100% in June 1986. The relative density of D. baeri was found to increase gradually during the first summer in both areas, although the accumulation rate of metacercariae was significantly increased in the heated area. In August of the first year the relative density of D. baeri peaked in the heated area, whereas it continued to increase in the unheated control. However, in September of the following year, the relative densities were at the same level in both thermal regimes. A concomitant decline in the degree of overdispersion of metacercariae within the host population was observed in the heated area as the population density of metacercariae decreased, whereas the index of dispersion remained at the same level throughout the study in the unheated area. However, it was not possible to sample perch in the unheated area between October 1986 and May 1987 and changes in the parasite population could have occurred during this period. The prevalence and relative density of D. spathaceum, the lens form, was on the contrary low, especially in the unheated site where it was recorded only occasionally. The infection of D. baeri exhibited a convex pattern in both thermal regimes, although the peak infection was noticed earlier in the heated area. Regulation of the parasite infrapopulation may have been achieved by the combined effects of a decreased transmission rate of cercariae with increasing age of the host, the mortality of metacercariae as a natural termination of the life span and to selective predation of heavily infected hosts. However, regardless of an increased accumulation rate of metacercariae in the heated area, the relative densities of D. baeri became equal in both thermal regimes at the end of the study. The mortality rate of metacercariae in the heated area was therefore presumably increased as compared with the unheated reference site.

Experiments were performed to study the effect of age and lifespan on the infection success of Diplostomum spathaceum miracidia. Specimens of Lymnaea peregra and L. stagnalis were individually exposed to single miracidia of D. spathaceum of different ages under conditions of constant illumination at 20°C. The life-span of the free-swimming miracidium of D. spathaceum is 24 h at 20°C. It was observed that as the miracidium ages, its ability to successfully infect snails declines rapidly.

The effects of water temperature on the ability of Diplostomum spathaceum miracida to infect and establish patent infections in Lymnaea peregra and L. stagnalis were investigated. Snails were infected over a range of temperatures (6–20°C) and kept thereafter at 20°C or were infected at 20°C and kept at either 14, 20, or 25°C. Infection success was determined after 8 weeks by either observing cercarial shedding or examining snail viscera for sporocysts. The establishment of miracidia declined at lower water temperatures despite maintenance for 8 weeks at 20°C while exposure of snails to miracidia at 20°C and maintenance at different temperatures had little apparent effect. Infection success under these conditions was related more to the numbers of miracidia to which the snails were exposed. However, under this latter experimental regime, the time taken for the infection to become patent clearly depended upon maintenance temperature.

The infectivity of Diplostomum spathaceum (Digenea: Trematoda) cercariae to rainbow trout and the efficacy of the diplostomule migration to the lens following different routes of administration was examined. The optimum age of infectivity for cercariae was between 0–5 h after liberation from the snail and for intraperitoneally injected diplostomules, 5 h post-transformation in vitro through fish skin. After exposure of the entire fish body or head to cercariae, metacercariae first appeared in the lens at 5 h and their numbers gradually increased until 22 h. Following exposure of the tail region of rainbow trout to cercariae, metacercariae first appeared in the lens at 14 h. Significantly more metacercariae established in the lens of fish following exposure of the fish head compared with the tail region; 40% of penetrating cercariae reached the lens of fish following exposure of the head or entire body, 20% of cercariae or diplostomules injected either intraperitoneally, intramuscularly or intracardially reached the lens while only 5% of cercariae established as metacercariae following exposure of the tail region.

Cercariae of Diplostomum spathaceum penetrate the skin of fish, and then migrate along blood vessels and tissues towards the head and the eye-lens. We studied their orientation behaviour in tail fins of guppies and in chemical concentration gradients within agar-filled choice chambers. In fins, they entered veins and orientated cranially, independent of the blood flow and living cells. In choice chambers, they were attracted by a small molecular fraction of fish serum, D-glucose (at 1, 10, and 1000 μm), D-mannose, D-maltotriose and Cl-ions, whereas D-glucosamine repelled them (even at 1·0 nm). Amino acids were not attractive, but arginine in tetrapeptides attracted at concentrations as low as 1 μm and melatonin at 0·4–4·3 pm. We suggest a preliminary model for the behaviour of diplostomula in fish fins and attracting (+) or repelling (−) host cues: (1) migration towards deeper skin layers and avoidance of skin surface, cues: Cl-ions (+ and −), glucose (+), glucosamine (−), light radiation (−); (2) orientation in cranial direction, cue: Cl-ions (+); (3) localization of blood vessels, cues: glucose (+), arginine-residues (+); (4) localization of the retina, cue: melatonin (+). A comparison with the navigation mechanisms of tissue-migrating schistosomules and hookworm larvae reveals an enormous diversity of strategies.

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 of Diplostomum 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 allowed wild birds to feed on fish from cages placed into a lake, and found that predation vulnerability of infected fish did not differ from that of controls. However, we suggest that the experimental set-up likely affected the result because the cages allowed gulls, which caught most of the fish in the study, to feed on fish in an easy, unnatural manner by standing on the edges of the cages. Nevertheless, considerable predation was observed, which provides important initial evidence of how this question should be addressed in the wild.

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 crucian carp. As predicted, crucian carp had more diverse parasite species composition in lakes, which supports the hypotheses of increased opportunities for parasite exchange with other fish species and better environmental conditions for the parasites. However, for instance, diplostomids were found only from few fish individuals in very low numbers. Experimental exposure trials with the eye fluke Diplostomum spathaceum in the laboratory indicated that crucian carp were totally resistant to infection, which is exceptional and suggests that the low number of these parasites in crucian carp was due to physiological unsuitability of the host. To obtain a better understanding on the mechanisms underlying the formation of parasite assemblages in these fish, further studies on the relationships between the unique physiology of crucian carp and infection success by other parasite species are needed.

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. Potential applications of the method in fish farming and parasitological studies are discussed.

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 laboratory showed that fish reacted to the parasite cercariae by avoiding the infection source, which decreased the rate of parasite establishment. We conclude that by using a combination of behavioural avoidance and physiological resistance, fish could defend against the parasite more effectively.

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 of Diplostomum spathaceum cercariae from naturally infected snail hosts, Lymnaea stagnalis. The average number of cercariae produced per day decreased as the snail host approached death counter to the terminal investment hypothesis. The finding supports the prediction of the bet hedging hypothesis and implies that the pattern of cercarial production may be explained by reduced virulence of the parasite within the snails to ensure extended total production time of cercariae. Nevertheless, survival of infected snails was still lower than uninfected snails suggesting that ultimately the infection still increased snail mortality rate. Cercarial production varied between days but was not cyclic, probably because of the physiology of the sporocysts within snails. Fewer cercariae were released at night, which may increase transmission efficiency to diurnally-active fish hosts. The mechanisms associated with daily cercarial production are discussed.

The toxicity of cadmium and zinc at concentrations ranging from 0·1 to 10 000 μg/l to the life-span of decaudised cercarial bodies (cercariae that have shed their tails) of Diplostomum spathaceum was investigated. The effects of metal exposure at 3 temperatures (12, 20, and 25 °C) and 3 levels of water hardness (distilled water, soft water and hard water) were studied. In general, under most experimental conditions increasing metal concentrations reduced the life-span of decaudised cercariae. Increasing water hardness and decreasing water temperature caused an increase in the life-span of both control and metal exposed decaudised cercariae. However, at certain isolated metal concentrations, associated with a specific level of water hardness and temperature, increased survival above controls occurred. Differences in the relative toxicity of cadmium and zinc were dependent on the environmental conditions of exposure. The decaudised cercarial life-span under metal exposure was found to be generally independent of the overall cercarial survival and tail loss in most experimental conditions. Prolonged exposure to cadmium and zinc caused changes in the decaudised cercarial life-span when compared to individuals decaudised during the initial 24 h exposure period to those which were decaudised during the final 24 h period of cercarial survival. The validity of studying the decaudised cercarial life-span as an indicator of ‘fitness’ of larvae to migrate through the tissues of the target fish host, in terms of glycogen utilization, was assessed for those cercariae decaudised during the initial 24 h exposure period only. A limited reduction in the decaudised cercarial life-span during this period compared to controls was recorded, which may possibly indicate a reduced penetration ‘fitness’ of cercariae exposed to cadmium and zinc. The importance and relevance of these findings to parasite migration and establishment in the fish host are discussed.

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 age-dependent pattern although infectivity tended (P=0·09) to increase with age over the first 6 h of life and then fall. The per capita transmission rate of cercariae was investigated by experimentally infecting rainbow trout under standardized conditions, first with an increasing cercarial density and second, by keeping density constant but increasing numbers of cercariae. The per capita transmission rate was frequency dependent and averaged 0·341/h (±0·036).

The effect of cadmium and zinc at concentrations ranging from 0.1 to 10000 μg/l on tail loss in cercariae of Diplostomum spathaceum was investigated at 3 temperatures (12, 20 and 25 °C) and 3 levels of water hardness (distilled water, soft water and hard water). Increasing tail loss over time was found to be linked with a parallel decrease in cercarial survival in controls. Exposure to the heavy metals induced, especially at high concentrations, a change in the relationship between cercarial tail loss and survival, causing either stimulation or inhibition of tail loss dependent on the individual toxic exposure. Under most environmental conditions the rate of tail loss over time was reduced by increasing metal concentrations. Inhibition of tail loss occurred in a limited number of both control and metal-exposed cercariae, with a number of low metal concentrations inducing greater inhibition than in controls. Stimulation of tail loss causing an increased tail loss rate above controls also occurred at certain high metal concentrations. Increasing water hardness and decreasing water temperature caused a reduced tail loss rate over time in both control and metal-exposed cercariae. However, with decreasing temperature a reduced rate of tail loss over time in metal-exposed cercariae compared to controls occurred at some low metal concentrations. When tail loss was compared against cercarial death of the experimental population toxic exposure induced changes in the parallel relationship of these parameters, dependent on individual metal concentration, water temperature and hardness. Differences in the relative effects of cadmium and zinc on cercariae were dependent on the environmental conditions of exposure. Both metals showed limited effects during the period of maximum cercarial infectivity (0–5 h). The mechanisms and importance of metal toxicity to cercarial tail loss are discussed.