Effects of Schistosomatium douthitti infection on the growth, survival, and reproduction of Lymnaea catascopium

Lymnaea catascopium snails infected with Schistosomatium douthitti grew faster than uninfected control snails during the first 2 months postexposure, but thereafter grew more slowly, and by 8 months postexposure were significantly smaller. When reared in isolation, uninfected snails survived significantly longer (mean survival time, 515 days) than snails exposed to three miracidia each (400 days), which in turn survived longer than snails exposed to 10 miracidia per snail (223 days). When maintained in aquaria in contact with other snails, snails exposed to three miracidia each survived longer (227 days), but not significantly longer, than control snails (198 days). Production of large numbers of eggs by control snails grown under the latter conditions may account for their reduced survival. The ovotestes and accessory genitalia of snails infected with S. douthitti were much reduced in size in comparison with uninfected control snails. These effects were most pronounced in snails which had been infected for over 100 days. Egg production was normally totally inhibited if snails were infected before the onset of sexual maturity. If infected after the onset of maturity, eggs were produced only during the prepatent period.     

Energy allocation patterns in Biomphalaria alexandrina snails in response to cadmium exposure and Schistosoma mansoni infection

This study was aimed to investigate the effects of both parasitism and environmental stress on the growth, reproduction, and survival of Biomphalaria alexandrina snails. Resource allocation strategies may be influenced by both biotic and abiotic factors. Using the planorbid snail B. alexandrina and Schistosoma mansoni, this hypothesis was examined by raising snails fed the same diet under two stressors (infection and Cd exposure). The snails divided into four groups, uninfected, infected, Cd-exposed uninfected, and Cd-exposed infected snails. Egg production, growth, and survival of the snails were monitored over a 9-week period postinfection. Inhibition of snail reproductive activity by parasitism results in increased snail growth in the first week postinfection, termed gigantism, during which the snail is hypothesized to allocate excess energy normally used for reproduction to somatic growth. Infection status and Cd exposure had significant effects on snail growth and reproduction. The infected and Cd-exposed infected snails exhibiting reduced survival relative to snails of other treatments. It was found that parasite development influenced by Cd exposure. Results of this study suggest that energy allocation patterns are context-dependent in B. alexandrina snails, influenced by infection and Cd exposure.     

Parasitic mites influence fitness components of their host, the land snail Arianta arbustorum

Abstract. Parasites can influence the population dynamics of their hosts by affecting life-history strategies and behavior. The hematophageous mite Riccardoella limacum lives in the lung cavity of terrestrial gastropods. We used correlational and experimental approaches to investigate the influence of parasite infection on the behavior and life-history traits of the simultaneously hermaphroditic land snail Arianta arbustorum , a common host of R. limacum. Naturally infected individuals of A. arbustorum , collected in the wild, showed a decreased activity compared with uninfected snails. The reproductive output, expressed as the number of eggs deposited in a reproductive season, was reduced in mite-infected hosts. However, the hatching success of the eggs laid by parasitized snails was slightly higher than that of uninfected individuals. We also examined winter survival in 361 adults of A. arbustorum collected from four natural populations. The prevalence of mite infection ranged from 44.8% to 70.1% in three populations (snails in the fourth population were not infected). Winter survival was reduced in infected snails in two out of three populations. Furthermore, experimentally infected snails from an uninfected population showed a reduced winter survival compared with control snails. Our results indicate that parasite pressure imposed by members of R. limacum may influence life history in A. arbustorum.     

Changes in the reproductive biology of Biomphalara glabrata infected with different doses of Echinostoma paraensei miracidia

The egg-laying rate, number of egg masses, number of eggs/mass, number of eggs hatched/snail and egg viability of Biomphalaria glabrata exposed to different doses (5 and 50) of Echinostoma paraensei miracidia were analyzed as indicators of reproductive activity. Polystyrene plates were placed in aquariums containing the snails and every other day for four weeks after infection the plates were removed to count the number of egg masses and eggs laid. After this, the plates were numbered individually and placed in new aquariums free of snails and the egg masses were observed daily to determine the hatching rate. On average there was an increase in the parameters evaluated in the infected snails in relation to the controls (uninfected snails), except for egg viability, which was significantly lower in the groups infected with 50 miracidia. These findings indicate that when infected, this host snail is able to increase its reproductive activity, suggesting an ecological strategy to maintain the species.     

Plagiorchis elegans (Digenea: Plagiorchiidae) infections in Stagnicola elodes (Pulmonata: Lymnaeidae): host susceptibility, growth, reproduction, mortality, and cercarial production.

Eggs of Plagiorchis elegans were readily ingested by Stagnicola elodes of all ages, but were more infective to immature than mature snails. Infection enhanced the growth of the host in a dose-dependent manner. The number of cercariae released by immature snails increased with the age of the snail host; mature snails yielded fewer cercariae. Heavily infected snails tended to die prematurely, thereby reducing their total production of cercariae to levels below those of more lightly infected individuals. Even light infections castrated the snail host. Snails that acquired the infection as juveniles never produced eggs. Actively reproducing snails ceased egg laying within days of infection and never recovered. All parasite effects on the growth and reproduction of the snail host first manifested themselves during the early stages of infection, long before the development of daughter sporocysts and cercariae, and are therefore attributable to the effects of mother sporocysts. The study provides insight into how this natural enemy of mosquito larvae may be established in natural snail populations by means of strategically timed introductions of parasite eggs, with a goal of maximizing cercarial production for the biological control of sympatric mosquito larvae.     

Effects of diet and<Emphasis Type="Italic"> Echinostoma revolutum</Emphasis> infection on energy allocation patterns in juvenile<Emphasis Type="Italic"> Lymnaea elodes</Emphasis> snails

Resource allocation strategies may be influenced by both biotic and abiotic factors. The purpose of this study was to investigate the effects of both parasitism and diet quality on the growth, reproduction, and survival of the pond snail, Lymnaea elodes. In addition, we assessed parasite growth and reproduction. High-protein (high diet) or low-protein diets (low diet) were fed to juvenile L. elodes snails that were either exposed or sham-exposed to the castrating trematode, Echinostoma revolutum. Host growth was assessed weekly; reproduction and survival were recorded every 2-3 days. We estimated parasite development as the time to parasite release from the host (patency), and parasite reproduction as the number of larvae shed from infected snails at two time points. Diet and infection status had significant effects on snail growth. Infected snails produced few eggs and tended to grow to larger sizes than uninfected snails regardless of diet. In contrast, exposed-uninfected individuals displayed diet-dependent patterns of growth and reproduction. On the high-protein diet, uninfected and exposed-uninfected snails exhibited similar patterns of growth and reproduction, whereas in the low-diet treatment, exposed-uninfected snails exhibited reduced growth and delayed reproduction relative to uninfected individuals. Survival differed among treatments in the latter stages of the study with infected snails exhibiting reduced survival relative to snails from other treatments. Moreover, infected low-diet snails exhibited lower survival than infected high-diet snails. Parasite development and reproduction did not appear to be directly influenced by the quality of host diet. Results from this study suggest that energy allocation patterns are context-dependent in juvenile snails, influenced by parasite exposure and diet quality. Furthermore, parasite reproduction appears to depend more on host size than on the quality of host diet.     

Schistosoma mansoni: effect of infection on reproduction and gonadal growth in Biomphalaria glabrata

Sexually mature Biomphalaria glabrata were exposed to 12 miracidia of Schistosoma mansoni, and egg production of snails was monitored over a period of 5 weeks. During the study period, exposed snails grew at approximately the same rate as unexposed controls. Castration, as measured by a reduction in the mean number of eggs laid per snail occurred between 14 and 21 days postexposure (PE). The reduction in fecundity in infected snails coincided with the migration and establishment of daughter sporocysts in the digestive gland and gonad. Enumeration of individual oocytes in longitudinal sections of the ovotestis revealed that uninfected snails contained significantly more oocytes per section than infected snails at 27, 31, and 40 days PE. In addition, the mean area of gonadal sections of control snails increased over the 40-day experimental period, whereas there was no such increase in gonadal area of infected snails. These data suggest that there is an inhibition in gonadal growth in infected snails. When oocyte data were expressed in terms of mean gonadal area, the mean number of oocytes per mm2 of gonad of uninfected and infected snails did not differ significantly over the study period, except at Day 14 PE, when infected snails contained a significantly greater number of oocytes per mm2 of gonad than did uninfected controls. It is hypothesized that daughter sporocysts of S. mansoni are primarily responsible for the inhibition of host reproductive activity, and may be mediating their effects through mechanisms involved in the regulation of gonadal growth.(ABSTRACT TRUNCATED AT 250 WORDS)     

Parasite-induced changes in nitrogen isotope signatures of host tissues

To estimate isotopic changes caused by trematode parasites within a host, we investigated changes in the carbon and nitrogen isotope ratios of the freshwater snail Lymnaea stagnalis infected by trematode larvae. We measured carbon and nitrogen stable isotopes within the foot, gonad, and hepatopancreas of both infected and uninfected snails. There was no significant difference in the delta13C and delta15N values of foot and gonad between infected and uninfected snails; thus, trematode parasite infections may not cause changes in snail diets. However, in the hepatopancreas, delta15N values were significantly higher in infected than in uninfected snails. The 15N enrichment in the hepatopancreas of infected snails is caused by the higher 15N ratio in parasite tissues. Using an isotope-mixing model, we roughly estimated that the parasites in the hepatopancreas represented from 0.8 to 3.4% of the total snail biomass, including the shell.     

Schistosoma mansoni infections in neonatal Biomphalaria glabrata snails.

In areas endemic for schistosomiasis, the population dynamics of the snail intermediate hosts have a direct effect on parasite transmission. The present study focused on the potential for neonatal Biomphalaria glabrata snails to become infected with Schistosoma mansoni and to produce cercariae under various conditions. It was found that snails as small as 0.74 mm in shell diameter could survive miracidial penetration and could release cercariae when as small as 1.6 mm in diameter. Cercariae produced by small snails were equally infectious for mice when compared with those shed by larger snails. Likewise, histological examination of neonatally exposed snails revealed normally developing parasites at all stages of infection. It was found that in 2 snail populations expressing either high or low susceptibility to the parasite, peak susceptibility occurred at 25 days of age in both groups. Daily cercarial production for neonatally exposed snails was initially low but increased dramatically as the snails grew, eventually reaching values as high as 2,100 cercariae/snail/day. A moderate to high percentage of snails infected as neonates was eventually capable of simultaneously producing both eggs and cercariae. These studies emphasize the potential importance of neonatal and preadult snails in helping to maintain foci of S. mansoni infection in endemic areas.     

Lymnaea fuscus (Pfeiffer, 1821) as a potential intermediate host of Fascioloides magna in Europe

Experimental infections of two different populations of Lymnaea fuscus in France and Sweden, with a Czech isolate of Fascioloides magna were carried out to determine if this lymnaeid species enables parasite larval development. Species identification of both snail populations was performed using the morphology of the copulatory organ, and also confirmed by sequencing of the internal transcribed spacer 2 (ITS2) region of the snail genomic rDNA. Only juvenile snails measuring less than 3 mm (1–3 weeks of age) were successfully infected (the viable cercariae were recorded) and infection prevalence decreased with age, as documented by increased shell height. In both French and Swedish L. fuscus populations, prevalence ranged between 1.1% and 58.8%. The mean number of metacercariae obtained from cercariae-shedding snails was 13.7 (±11.4), while the total cercarial production noted in snails dissected at day 85 post-exposure was 147.5 (±56.6). Compared to uninfected control snails, we observed reduced growth of infected snails. Despite age-related resistance of snail to the parasite, and limited cercarial production in these experimentally infected snails, F. magna was still able to complete larval development in L. fuscus.     

Schistosoma mansoni: influence of Biomphalaria glabrata size on susceptibility to infection and resultant cercarial production

Biomphalaria glabrata snails of the same age, but different sizes, were used to determine size-related susceptibility to Schistosoma mansoni miracidial infection and the influence of snail size on total cercarial production. Snails with shell diameters from less than 5 to greater than 17 mm were individually exposed to one or several miracidia, depending on the experiment. In snails exposed to multiple numbers of miracidia, the percentage of snails which developed patent infections was lower in snails with larger shell sizes. This was also reflected by fewer primary sporocysts per infected snail found in tissues of the larger snails. Upon determining cercarial production in these groups over a 1-month period there were no statistical differences between any groups in the numbers of cercariae produced per snail. However, upon determining the number of successful primary sporocysts found in cohort snails of each size group, cercarial production increased as a function of the number of successful primary sporocysts. This was verified by examining cercarial production in various size snails with known monomiracidial infections. Our data therefore confirm and extend earlier work using snails infected with unknown numbers of miracidia and clearly show that total S. mansoni cercarial development and decreased susceptibility of snails is a direct reflection of snail size and not necessarily age of the snail.     

Pathophysiologic alterations in Biomphalaria glabrata infected with Angiostrongylus costaricensis

Hemolymph glucose, alkaline phosphatase, lactic dehydrogenase, and creatine phosphokinase in Biomphalaria glabrata infected with Angiostrongylus costaricensis were significantly higher on day 27 postinfection (PI) than in uninfected snails. Hemolymph total calcium from infected snails was less on days 6, 12, and 27 PI than that from controls. Total hemolymph protein was similar for controls and infected animals during the entire study. Throughout the study the mean number of amoebocytes/mm³ hemolymph from infected snails was significantly less than that for controls. Mean total wet weights of digestive gland and foot muscle from infected and uninfected snails was similar throughout the study. Mean μg glycogen/mg wet weight of digestive gland from infected snails was significantly greater on days 24, 27, and 28 PI than that from controls. Mean μg glycogen/mg wet weight of foot muscle from infected snails was significantly reduced between days 12 and 28 PI from that of uninfected snails. It is suggested that hemolymph glucose and digestive gland glycogen in infected snails are augmented by glycogen breakdown in the foot muscle of parasitized animals. Elevations in hemolymph enzymes are due to tissue destruction by larvae emerging from the foot muscle of infected snails. Parasite-induced derangements in shell metabolism underlie observed changes in hemolymph calcium in infected snails.     

Development of Fasciola hepatica in Lymnaea columella infected with miracidia derived from cattle and marmoset infections

The development of Fasciola hepatica from two species of definitive hosts, i.e. cattle (Bos taurus) and a marmoset (Callithrix penicillata) in the snail Lymnaea columella was determined based on the production of rediae and cercariae and snail survival rate. More rediae and cercariae at 60-74 days post-infection were produced by snails infected by cattle-derived miracidia (cattle group) than by those infected by marmoset-derived miracidia (marmoset group). Among the L. columella parasitized by the marmoset group, the survival rate and the percentage of positive snails were higher than among those parasitized by the cattle group. Eggs of F. hepatica released in cattle faeces were significantly bigger than those released in marmoset faeces. Miracidia originating from parasites that completed their development in cattle were more efficient in infecting the intermediate host. These results suggest that vertebrate-host origin influences the eggs produced by the parasite and the infection rates in the snail host L. columella.     

Cost of resistance: relationship between reduced fertility and increased resistance in a snail—schistosome host—parasite system

Natural host populations often exhibit genetic variability in resistance to parasitism. One possible mechanism for maintaining such diversity is a trade-off between fitness costs associated with resistance and fitness costs associated with parasitism. However, little is known about the nature or magnitude of these costs in animal populations. Using artificial selection experiments in a Biomphalaria glabrata–Schistosoma mansoni host–parasite system, we demonstrated that resistance and susceptibility to infection are heritable. We then investigated whether resistance had any associated costs in terms of snail reproductive success. Susceptible-selected snail lines showed significantly higher fertility (number of offspring produced) than resistant-selected or unselected control snail lines, irrespective of current infection status. There were no consistent differences between snail lines in fecundity, proportion of abnormal egg masses produced, or mean number of eggs per egg mass. Mortality rate was higher among infected than uninfected snails. These results are consistent with snails incurring costs of resistance to schistosome infection in the absence of the parasite.     

Echinostoma liei miracidia and Biomphalariaglabrata snails: Effect of egg age,habitat heterogeneity,water quality and volume on infectivity

Infectivity of Echinostoma liei miracidia to NIH albino Biomphalaria glabrata declines significantly from 62% with eggs incubated for 10–24 days to 3% for eggs incubated for 30–42 days. In mass exposures of 25 snails to 125 miracidia in 1 liter of water infectivity was high (54–66 %) and not affected by the presence of lettuce, plastic sheets, chalk, detritus or snail-conditioned water. In distilled water or snail-conditioned water the proportion of infected snails exposed singly to five miracidia per snail in 5 ml was not significantly different from the results of mass exposures of 25 snails in 1 liter to the same snail: miracidia ratio. Some evidence is presented suggesting that infected snails are less likely to suffer mortality than uninfected snails during the first 7–10 days post-exposure.The results suggest that Echinostoma liei miracidial searching efficiency is robust in volumes of at least 1 liter and in a heterogeneous habitat. These aspects enhance the competitive potential of echinostomes as possible biological control agents for Schistosoma mansoni.     

Altered behavior of the snail Biomphalaria glabrata as a result of infection with Schistosoma mansoni

In this comparative behavioral study, the effect of infection with Schistosoma mansoni on its snail intermediate host Biomphalaria glabrata was investigated. Three groups of snails were compared for their activity: (1) uninfected, (2) infected with male parasites, and (3) infected with female parasites. In solitary movement trials, uninfected snails traveled greater distances at faster rates, explored more surface area, and had shorter rest periods than snails infected with either male or female schistosomes. In Y-maze experiments designed to determine attraction, the uninfected snails more often and more quickly moved toward other snails than the infected individuals. Snails from all 3 groups were more attracted to infected individuals than to uninfected ones. There was no difference in attraction toward snails infected with male or female parasites. These experiments provide evidence that behavioral alterations as a result of infection may lead to aggregation of infected snails in the field. We propose that such an effect may result in enhanced parasite transmission.     

Altered feeding response as a cause for the altered heartbeat rate and locomotor activity of Schistosoma mansoni-infected biomphalaria glabrata

Biomphalaria glabrata infected with Schistosoma mansoni for 33 days fed more often than uninfected snails. Whereas uninfected snails had nocturnal increases in feeding, snails with a 33-day-old infection of S. mansoni fed as often during the day as in the night. Using direct observation and film analysis, we found that feeding increased the heartbeat rate and locomotor activity of B. glabrata. When snails were allowed to feed ad lib., infected snails had higher heartbeat rates than uninfected snails both during the day (P less than 0.01) and the night (P less than 0.001). However, when the snails were deprived of food for 24 hr, infected snails had slightly higher heartbeat rates than uninfected snails only during the day (P less than 0.05). There was no difference between the heartbeat rates of feeding, infected snails and the heartbeat rates of uninfected snails that were starved for 8 hr, and then allowed to feed. Uninfected snails had nocturnal increases in heartbeat rate regardless of feeding schedule, but infected snails had greater nighttime heartbeat rate than daytime heartbeat rate only when they were not allowed to feed. Infected snails had less nocturnal locomotor activity than uninfected snails when feeding, but there was no difference between the locomotor activity of infected and uninfected snails when the snails were deprived of food for 24 hr. Absence of food also resulted in an increased nighttime to daytime ratio of locomotor activity of infected snails. These results suggest that the increased heartbeat rate and altered rhythms of heartbeat rate and locomotor activity in B. glabrata infected with S. mansoni for 33 days were caused by the altered feeding response of these snails.     

Effects of diet on the development of Schistosoma mansoni in Biomphalaria glabrata and on the neutral lipid content of the digestive gland-gonad complex of the snail

In a previous study, when the snail Biomphalaria glabrata was infected with Schistosoma mansoni and maintained on a diet of hen's egg yolk, it produced fully developed cercariae in about one-half the time taken by snails fed Romaine lettuce. Increased lipids were also noted in the snails fed the yolk diet. The purpose of the present study was to further investigate nutritional effects of a high-lipid diet on larval schistosome development and to reexamine the time to cercarial patency in infected snails maintained on either the yolk or lettuce diet and to use high-performance thin-layer chromatography (HPTLC) to analyze the neutral lipids in the digestive gland-gonad complex (DGG) of snails maintained on both diets. Infected snails maintained at 26 C and fed either diet produced fully developed cercariae by 4 wk postinfection (PI). Likewise, infected snails maintained at 23 C and fed either diet produced fully developed cercariae by 6 wk PI. The contention that the yolk diet enhanced the time to cercarial patency was not confirmed. The HPTLC analysis of neutral lipids showed that the DGG of infected snails fed the yolk diet contained significantly greater amounts of free sterols and cholesteryl esters but not triacylglycerols than that of the infected snails fed the lettuce diet.     

Self-fertilization and cross-fertilization in the land snail Arianta arbustorum (Mollusca, Pulmonata: Helicidae)

The simultaneously hermaphroditic land snail Arianta arbustorum (L.) (Pulmonata: Helicidae) was long believed to be self-incompatible. In the present study, however, uniparental reproduction was observed under laboratory conditions. Snails from two natural populations and from laboratory culture reared in isolation from the subadult stage produced fertile eggs. Over a period of three years after maturation, 38.5% of the unmated snails produced fertile eggs, mostly in the second and third years instead of the first year. Egg production and hatching success (the proportion of eggs that hatched) were lower in unmated snails than in mated snails. The number of hatchlings produced by unmated snails was 1–2% of that produced by mated snails. Hatchlings from unmated snails survived as well as those from mated snails under laboratory conditions. The result indicate that A. arbustorum can self-fertilize but with a great fitness reduction.