Hyperparasitism by Paragordius varius (Nematomorpha: Gordiida) larva of Monostome redia (Trematoda: Digenea)

Gordiid larvae enter and encyst within a large variety of aquatic vertebrates and invertebrates. Cysts of a single species of gordiid have been found in such diverse hosts as insect larvae, snails, crustaceans, fish, and many others. One particularly surprising host report was made nearly 100 yr ago, i.e., cysts within adult trematodes. The origin of these cysts remained unclear, but it was suggested that the gordiid larvae gained entry into the adult flukes via their definitive, vertebrate host. In the present study, the finding is reported of gordiid cysts within monostome redia within a Physa gyrina snail. This is the first time gordiid cysts have been reported within redia, and this provides an alternative explanation for the presence of cysts within adult trematodes.     

Characterization of 15 polymorphic microsatellite markers in the freshwater snail Aplexa marmorata (Mollusca, Gastropoda)

We describe 15 new variable microsatellites in the freshwater snail Aplexa (Physa) marmorata, as well as conditions for multiplexing and simultaneous genotyping sets of loci. Two to 11 alleles were detected per locus over the five populations studied, and up to eight alleles per locus and population. The observed heterozygosity was extremely low, most probably as the consequence of high selfing rates. The genetic differentiation among populations was large. These markers will constitute efficient tools for investigating the population structure in fragmented habitats. Cross-species amplification in three Physa species (P. acuta, P. fontinalis and P. gyrina) was rather unsuccessful.     

Predator–prey interactions between crayfish (Orconectes juvenilis) and snails (Physa gyrina) are affected by spatial scale and chemical cues

Abstract. We conducted two experiments to compare the behaviors of predatory crayfish ( Orconectes juvenilis ) and snail prey ( Physa gyrina ), with and without exposure to injured-snail cues, in seven different sizes of arena. Individuals in both O. juvenilis and P. gyrina are known to obtain information about predation events from chemical cues present in the water, but the effect of spatial scale on this information or the ensuing interaction has not been previously investigated. We found that crayfish were generally more active in larger arenas, but their response to injured-snail cues decreased as arena size increased. Snails responded to injured-snail cue by seeking refuge, but responded less strongly in larger arenas. Less time was required for crayfish to capture a snail in the presence of injured-snail cue, and there was an interaction between cue and arena size evident in middle-sized arenas. Consumption time was lower in the presence of injured-snail cue but was unaffected by arena size. Based on these results, we conclude that the effects of injured-snail cue are mediated by spatial scale, and this affects the outcome of this predator–prey interaction. Spatial scale should be considered in the design and interpretation of other predator–prey studies, particularly when sensory or motility differences exist between predators and prey that affect how each gathers information about the environment.     

Populations of the European freshwater pulmonate Physa acuta are not reproductively isolated from American Physa heterostropha or Physa integra

Abstract. It has long been speculated that Physa acuta , a pulmonate snail widespread and invasive in fresh waters of the old world, may have originated in North America. But the identification of a new-world cognate has been complicated by the confused systematics and taxonomy of the Physidae in America. More than 40 species of physids are currently recognized in the United States, many with variable and overlapping morphology. We have previously established that premating reproductive isolation is negligible among physid snails. Here we report the results from no-choice crosses each involving 2 populations of the widespread American species Physa heterostropha and Physa integra , both with each other and with P. acuta , designed to compare measures of reproductive success between species and between populations within species. Samples of P. acuta were collected from France and Ireland, P. heterostropha from eastern Pennsylvania and South Carolina, and P. integra from southern Indiana and northern Michigan. The 6 intrapopulation controls varied quite significantly in their survival, age at first reproduction, parental fecundity, F₁ viability, and F₁ fertility under our culture conditions. Measures of survival and reproduction in the 6 interpopulation crosses were generally intermediate, but in no case significantly worse than the more poorly performing control. Thus we were unable to detect evidence of reproductive isolation among our 6 populations of snails from 2 continents. All should be referred to the oldest available nomen, P. acuta .     

北京发现尖膀胱螺

尖膀胱螺(Physa acuta)为世界性广布淡水螺类,系多种重要寄生虫如广州管圆线虫(Angiostrongylis cantonensis)和卷棘口吸虫(Echinostoma revolutum)的中间宿主.在20世纪70年代淡水贝类调查中,北京地区未曾发现该螺的分布.2007年笔者在北京地区开展主要寄生虫流行病学调查,在朝阳公园、玉渊潭、圆明园及奥运村附近河流中相继发现该螺,此螺入侵北京地区并大量孳生应引起相关部门高度重视.     

Ecological factors in schistosome transmission, and an environmentally benign method for controlling snails in a recreational lake with a record of schistosome dermatitis

The avian schistosomes, Trichobilharzia stagnicolae, T. physellae and Gigantobilharzia sp., that cause Schistosome Dermatitis (Swimmers' Itch) in humans were studied in the laboratory and at Cultus Lake, British Columbia, Canada in relation to the biology and behavior of their intermediate snail hosts, Stagnicola catascopium, Physa sp. and Gyraulus parvus, respectively, and their definite bird hosts. Wind-driven, surface currents were measured. Populations of snails, close to host-bird roosting logs had a very high prevalence of schistosome infections. An experiment that mechanically disturbed the epilithic habitat of the snails using a boat-mounted rototiller or a tractor and rake, eliminated almost all of the snails if the disturbance was done in areas of high snail concentration in shallow areas of the lake during the breeding and early development phase of the snail. It is proposed that the incorporation of snail habitat disturbance into management programs is an effective way to control Schistosome Dermatitis.     

Evolutionary relationships among the Schistosomatidae (Platyhelminthes:Digenea) and an Asian origin for Schistosoma

Schistosome blood flukes parasitize birds, mammals, and crocodilians and are responsible for causing one of the great neglected diseases of humanity, schistosomiasis. A phylogenetic study of 10 schistosome genera using approximately 1,100 bases of the large subunit of the nuclear ribosomal gene complex revealed 2 major clades. One clade is entirely mammalian and includes the genera Schistosoma and Orientobilharzia. A close examination of relationships in this group suggests that the medically important Schistosoma arose in Asia and not in Africa as generally presumed and is paraphyletic. The second clade is primarily avian, consisting of 6 genera of exclusively avian parasites and 2 genera of North American mammal flukes. These results indicate a secondary host capture of mammals on the North American continent. This study provides little evidence concerning the ancestral molluscan or vertebrate schistosome host but does demonstrate that host switching has been an important feature of schistosome evolution. Evidence also indicates that the reduced sexual dimorphism characteristic of some avian schistosomes is derived evolutionarily.     

Why are there few algae on snail shells? The effects of grazing,nutrients and shell chemistry on the algae on shells of Helisoma trivolvis

1. Freshwater snails often lack visible growths of algae on their shells. We tested three possible mechanisms that may account for this (grazing, snail-derived nutrients and chemical defences), using the ramshorn snail Helisoma trivolvis .
2. The experiments were carried out in floating plastic enclosures in a pond and comprised seven treatments. Grazing treatments were: a lone snail (ungrazed, as self-grazing does not occur), Helisoma with conspecifics, Helisoma with the co-occurring pond snail Physa sp., empty shells with Helisoma , and ungrazed empty shells. Nutrient effects were possible in all treatments with occupied shells (lone snail; Helisoma with conspecifics, and with Physa ) versus absent in other treatments. Testing for chemical defences compared algae on fresh empty shells, weathered shells (outer organic periostracum layer absent) and boiled fresh shells (with denaturization of susceptible proteins).
3. Diatoms dominated algal assemblages on snail shells. Although the upright diatom Gomphonema gracile was abundant on all shells, it was dominant on the shells of snails housed with other snail grazers (either Helisoma or Physa ).
4. Only the lone snail (nutrients but no grazing) showed higher algal biomass, so presumably any nutrient effect in the treatments with grazers was masked. Both Helisoma and Physa were observed apparently grazing on Helisoma shells, and consequently algal biomass in multi-snail treatments was similar to that on empty shells. Scanning electron microscopy revealed that algal density was highest near the aperture of live snails, but not empty shells; this is consistent with a nutrient addition effect. There was no evidence of chemical defences against algal growth.
5. In soft-bottomed freshwater habitats with abundant snails, shells of living snails provide nutrient-augmented substrata that may indirectly boost overall snail production.     

Intermediate host availability masks the strength of experimentally-derived colonisation patterns in echinostome trematodes

A fundamental goal of parasite evolutionary ecology is to elucidate patterns of host use and determine the underlying mechanisms of parasite colonisation. In order to distinguish the relative contributions of host encounter rates and host compatibility to infection outcomes, we compared host use in both field and experimental laboratory settings. Two years of bi-weekly snail sampling at a freshwater pond demonstrated fluctuating availability among three potential second intermediate snail host species and suggested that two trematode species (Echinostoma revolutum and Echinoparyphium sp.) did not colonise the three potential snail host species, Lymnaea elodes, Physa gyrina and Helisoma trivolvis, differentially. However, a series of experimental infections demonstrated that both parasites colonised H. trivolvis more so than the other two host species. Thus, more echinostome parasites utilised snail hosts that cannot serve as their first intermediate host. In experimental infections, host size and vagility were not strong determinants of infection. By utilising field and laboratory approaches, we were able to compare the strength of host compatibility under controlled conditions with patterns of infection in nature. Based on the results from these studies, it appears that host encounter is the primary mechanism dictating infection outcomes in the field.     

Comparative life histories of two diplostomid trematodes, Ornithodiplostomum ptychocheilus and Posthodiplostomum minimum

A lack of comparative studies limits our understanding of interspecific variation in parasite life histories, especially for species that incorporate asexual and resting stages into their life cycles. Ornithodiplostomum ptychocheilus and Posthodiplostomum minimum are sympatric diplostomid trematodes that share the same first (pond snail, Physa gyrina) and second (fathead minnow, Pimephales promelas) intermediate hosts within lakes in Alberta, Canada. Interspecific differences in the body sizes of adults, cercariae, and metacercariae were evaluated from parasites recovered from laboratory-raised chickens, snails, and minnows, respectively. Differences in adult worm fecundity and cercariae production were estimated from chickens and snails exposed to known numbers of parasite larvae. Posthodiplostomum minimum adults, metacercariae, and eggs were 97, 86, and 5% larger than those of 0. ptychocheilus, respectively. The average numbers of eggs produced per day by P. minimum was 3 times higher than for O. ptychocheilus, and P. minimum produced approximately twice as many cercariae per day. Although the larger of the 2 species had higher egg and cercariae production, conclusions regarding covariation between adult worm size and the reproductive rates of other life cycle stages require further studies on related species.     

Trade-offs in the evolution of virulence in an indirectly transmitted macroparasite

The adaptive trade-off theory for the evolution and maintenance of parasite virulence requires that virulence be genetically correlated with other fitness characteristics of the parasite. Many theoretical models rely on a positive correlation between virulence and transmissibility. They assume that high parasite replication rates are associated with a high probability of transmission (and, hence, increased parasite fitness), but also with high levels of damage to the host (high virulence). Schistosomes are macroparasites with an indirect life cycle involving a mammalian and a molluscan host. Here we demonstrate, through the development of five substrains, a genetic basis for schistosome virulence. We used these substrains further in order to investigate the presence of parasite fitness traits that were genetically correlated with virulence. High virulence in the (mouse) definitive host was, as predicted, positively correlated with parasite replication. In contrast, in the (snail) intermediate host high virulence was associated with low parasite replication rates. Variation in infectivity to and parasite replication in the definitive host was suggested as a compensating mechanism for the maintenance of virulence in the snail host. This is the first report of a trade-off in parasite reproductive success across hosts in an indirectly transmitted macroparasite.     

Influence of lake trophic status on the occurrence of microcystin-LR in the tissue of pulmonate snails

1. To assess the influence of lake trophic status on the occurrence of microcystin-LR (MCLR) in the tissue of resident pulmonate snails, we sampled seven lakes of varying primary productivity in Alberta, Canada. Parameters associated with productivity were measured every 2 weeks from mid-May through mid-September 1995. Phytoplankton and resident pulmonate snails were collected and analysed for MCLR concentration via high-performance liquid chromatography.
2. For all species of gastropod (i.e. Lymnaea stagnalis , Helisoma trivolvis and Physa gyrina ), the concentration of MCLR in the tissue was correlated ( P  ≤ 0.03) with toxin in the phytoplankton, but not with extracellular aqueous microcystin ( P > 0.28). The concentrations of toxin in the tissues of L. stagnalis and P. gyrina were also correlated with the relative abundance of Microcystis spp. ( P < 0.01).
3. Given that Microcystis spp. abundance and MCLR concentration within phytoplankton are correlated with indicators of productivity, we conclude that trophic status is important in influencing the occurrence and concentration of MCLR in pulmonate snails.     

Prevalence of schistosome infections within molluscan populations: observed patterns and theoretical predictions

The paper draws together a large and scattered body of empirical evidence concerning the prevalence of snail infection with schistosome parasites in field situations, the duration of the latent period of infection in snails (and its dependence on temperature), and the mortality rates of infected and uninfected snails in field and laboratory conditions. A review and synthesis of quantitative data on the population biology of schistosome infections within the molluscan host is attempted and observed patterns of infection are compared with predictions of a schistosomiasis model developed by May (1977) which incorporates differential snail mortality (between infected and uninfected snails) and latent periods of infection. It is suggested that the low levels of prevalence within snail populations in endemic areas of schistosomiasis are closely associated with high rates of infected snail mortality and the duration of the latent period of infection within the mollusc. In certain instances, the expected life-span of an infected snail may be less than the duration of the latent period of infection. Such patterns generate very low levels of parasite prevalence. A new age prevalence model for schistosome infections within snail populations is developed and its predictions compared with observed patterns. The implications of this study of observed and predicted patterns of snail infection within molluscan populations are discussed in relation to the overall transmission dynamics of schistosomiasis.     

Mate choice, frequency dependence, and the maintenance of resistance to parasitism in a simultaneous hermaphrodite

Biomphalaria glabrata are simultaneous hermaphroditic freshwatersnails that act as intermediate hosts for the macroparasitictrematode Schistosoma mansoni, a causative agent of schistosomiasis.Heritability and strain-specificity of both snail resistanceand susceptibility to schistosome infection have been demonstrated,genetic variability for which is maintained, in part, throughtrade-offs between high fitness costs associated with infectionand those associated with resistance. However, despite sucha high cost of resistance and a low prevalence of infectionin natural snail populations, genes for resistance are maintainedwithin snail populations over successive generations, includingin the complete absence of parasite pressure in laboratory populations.This may be indicative of alternative benefits of resistancegenes, in addition to parasite defense, such as differentialmating success between genotypes. Here we examined the mateand gender choice of snails across a multi-factorial range ofpotential partner combinations. These included host-resistanceor susceptibility genotype, host genotype frequency within thepopulation, current parasite infection status, and parasitegenotype. We demonstrate recognition and discrimination by hostsnails depending on host and/or parasite genotype for each ofthese factors. In particular, our results suggest that a raremating advantage to resistant genotypes may be a potential explanationfor the maintenance of highly costly resistance genes withinintermediate host populations under conditions of low or zeroparasite pressure.     

Solving cryptogenic histories using host and parasite molecular genetics: the resolution of Littorina littorea's North American origin

Even after decades of investigation using multiple sources of evidence, the natural histories of some species remain unclear (i.e. cryptogenic). A key example is Littorina littorea , the most abundant intertidal snail in northeastern North America. Native to Europe, the snail's ecological history in North America has been debated for over 100 years with no definitive resolution. To resolve its cryptogenic status, we used molecular genetics from a novel combination of the snail and a highly associated trematode parasite, Cryptocotyle lingua . Based on mitochondrial sequences of 370 L. littorea and 196 C. lingua individuals, our results demonstrate a significant reduction in genetic diversity in North America vs. Europe, North American haplotypes nested within European haplotypes, and mean divergence estimates of ~500 years ago from Europe for both host and parasite — thus supporting a recent introduction of both host and parasite to North America from Europe. Our study therefore resolves not only a specific cryptogenic history, but it also demonstrates the success of our approach generally and could be used in resolving difficult invasion histories worldwide.     

Toxic effects of cadmium and zinc on the transmission of Echinoparyphium recurvatum cercariae

The toxicity of cadmium, zinc and Cd/Zn mixtures to the transmission of Echinoparyphium recurvatum (Digenea: Echinostomatidae) cercariae into the snail second intermediate hosts was investigated at concentrations ranging from 100 microg l-1 to 10 000 microg l-1 in both soft and hard water. A differential response in the infectivity of metal-exposed cercariae into Lymnaea peregra and Physa fontinalis was demonstrated which was dependent on the snail species being infected. Exposure of L. peregra, P. fontinalis, and L. stagnalis to heavy metals caused a differing susceptibility to E. recurvatum cercariae depending on the snail species being exposed. The mechanism and effects of metal toxicity, together with the importance of the parasite/host strain on cercarial transmission are discussed.     

Bioenergetic theory predicts infection dynamics of human schistosomes in intermediate host snails across ecological gradients

Epidemiological dynamics depend on the traits of hosts and parasites, but hosts and parasites are heterogeneous entities that exist in dynamic environments. Resource availability is a particularly dynamic and potent environmental driver of within‐host infection dynamics (temporal patterns of growth, reproduction, parasite production and survival). We developed, parameterised and validated a model for resource‐explicit infection dynamics by incorporating a parasitism module into dynamic energy budget theory. The model mechanistically explained the dynamic multivariate responses of the human parasite Schistosoma mansoni and its intermediate host snail to variation in resources and host density. At the population level, feedbacks mediated by resource competition could create a unimodal relationship between snail density and human risk of exposure to schistosomes. Consequently, weak snail control could backfire if reductions in snail density release remaining hosts from resource competition. If resource competition is strong and relevant to schistosome production in nature, it could inform control strategies.     

尖膀胱螺的生物学特征及光照对其生长繁殖影响的初步研究

尖膀胱螺Physa acuta是世界广泛分布的淡水螺类,在我国属外来入侵种.本文着重对该螺的形态特征、繁殖、发育及光照对螺生长和繁殖发育的影响等进行了实验观察和研究.尖膀胱螺个体中等,左旋,壳口无厣.喜聚居于水生植物较多、营养丰富的淡水区域.尖膀胱螺为雌雄同体,异体受精,有时也可自体受精.性成熟个体在条件适宜的情况下(温度、食物和光照)每天都可产卵,卵孵化率近100％.尖膀胱螺胚胎发育共经历卵裂期、囊胚期、原肠期、担轮幼虫期、面盆幼虫期、仔螺形成及出膜期7个时期.一般情况下,螺卵7～10 d可孵化,60 d即可性成熟.25℃、24 h连续光照条件下,尖膀胱螺体重明显增加,40 d就可以性成熟并产卵.     

Schistosoma mansoni: TGF-beta signaling pathways

Schistosome parasites have co-evolved an intricate relationship with their human and snail hosts as well as a novel interplay between the adult male and female parasites. We review the role of the TGF-beta signaling pathway in parasite development, host-parasite interactions and male-female interactions. The data to date support multiple roles for the TGF-beta signaling pathway throughout schistosome development, in particular, in the tegument which is at the interface with the host and between the male and female schistosome, development of vitelline cells in female worms whose genes and development are regulated by a stimulus from the male schistosome and embryogenesis of the egg. The human ligand TGF-beta1 has been demonstrated to regulate the expression of a schistosome target gene that encodes a gynecophoric canal protein in the schistosome worm itself. Studies on signaling in schistosomes opens a new era for investigation of host-parasite and male-female interactions.