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.     

High susceptibility of Neotricula aperta gamma-strain from Krakor and Sdau in Cambodia to Schistosoma mekongi from Khong Island in Laos

Neotricula aperta gamma-strain snails collected from Krakor and Sdau in Cambodia were found to have the same or higher susceptibility to Schistosoma mekongi as N. aperta originally isolated from Khong in Laos. Infection rates of N. aperta gamma-strain snails exposed to 3 miracidia at week 8 were: Khong gamma-strain, 22.6%; Krakor gamma-strain, 33.3%; and Sdau gamma-strain, 67.4%. At week 10, the Sdau gamma-strain showed the highest infection rate of 83.3%. We thus found significantly high susceptibility of the Sdau gamma-strain to S. mekongi originally isolated from Khong. However, in another experiment, susceptibility of the Sdau gamma-strain was rather comparable to that of Khong and Krakor gamma-strain. We also found no significant differences in infection and survival rates between the Khong and Krakor gamma-strain when the snails were exposed to 3 or 6 miracidia. This is the first report to confirm the high susceptibility in the laboratory of N. aperta gamma-strain snails from endemic areas in Cambodia to S. mekongi originally isolated from Laos. The high susceptibility of N. aperta gamma-strain snails to S. mekongi in distant areas may be an important factor in the endemic transmission of human schistosomiasis.     

Schistosomiasis mansoni in areas of low transmission: epidemiological characterization of Venezuelan foci

Severe schistosomiasis is a rare event in Venezuela nowadays, after a successful national campaign by the Schistosomiasis Control Program. Unfortunately, this program has practically disappeared, and snail surveillance in field is not a priority, anymore. Thus, schistosomiasis has become a neglected disease in this country. However, surveys in different populations from the endemic area have shown particular epidemiological features described herein. In five communities we evaluated 2,175 persons and searched for the presence of Biomphalaria glabrata snails. Some markers were used for classifying schistosomiasis foci: mean age of the persons with Schistosoma mansoni eggs in the stools, serological tests, presence of B. glabrata snails, and intensity of infection. Places without B. glabrata snails and with few schistosomiasis cases were defined as "past transmission sites"; a site with abundant snails but few cases was defined as "potential risk"; "new transmission" foci were characterized by the presence of infected snails and young people passing eggs in the stools. A "re-emergent" focus has shared these last features, showing in addition a place where schistosomiasis had been reported before. Recent evidences of active transmission with the increasing dispersion of B. glabrata snails, point out the necessity for the re-establishment of the Schistosomiasis Control Program in Venezuela.     

Index of potential contamination for schistosomiasis

Schistosomiasis occurs under a wide variety of climatic, geographical and other conditions. The basic reason for the transmission of schistosomes is, however, the low level of sanitation in endemic areas with the result that faeces or urine, or both, containing schistosome eggs get into water that contains freshwater snails susceptible to infection. Unfortunately, the degree of contamination of natural habitats with schistosome ova is difficult to estimate; consequently, there are few data on how much contamination, relative to a defined snail population density, is necessary to initiate or maintain transmission of schistosomiasis. This article discusses a little-used method to estimate the degree of contamination or transmission - the index of potential contamination (IPC). The expected changes in IPC after praziquantel treatment will be considered, as well as the level of reduction in IPC that might be needed to affect transmission or reduce morbidity.     

Metabolic depression during aestivation does not involve remodelling of membrane fatty acids in two Australian frogs

Changes in membrane lipid composition (membrane remodelling) have been associated with metabolic depression in some aestivating snails but has not been studied in aestivating frogs. This study examined the membrane phospholipid composition of two Australian aestivating frog species Cyclorana alboguttata and Cyclorana australis. The results showed no major membrane remodelling of tissue in either frog species, or in mitochondria of C. alboguttata due to aestivation. Mitochondrial membrane remodelling was not investigated in C. australis. Where investigated in C. alboguttata, total protein and phospholipid content, and citrate synthase (CS) and cytochrome c oxidase (CCO) activities in tissues and mitochondria mostly did not change with aestivation in liver. In skeletal muscle, however, CS and CCO activities, mitochondrial and tissue phospholipids, and mitochondrial protein decreased with aestivation. These decreases in muscle indicate that skeletal muscle mitochondrial content may decrease during aestivation. Na⁺K⁺ATPase activity of both frog species showed no effect of aestivation. In C. alboguttata different fat diets had a major effect on both tissue and mitochondrial phospholipid composition indicating an ability to remodel membrane composition that is not utilised in aestivation. Therefore, changes in lipid composition associated with some aestivating snails do not occur during aestivation in these Australian frogs.     

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.     

Human schistosomiasis in wetlands in southern Africa

This review describes the types of habitats in which human schistosomiasis is transmitted in wetlands across southern Africa. The snails that serve as the parasites?? intermediate hosts are tolerant of water with a wide range of chemical composition but are restricted by current velocity, temperature and habitat permanence. Human contact with the water in snail habitats is crucial to the establishment of transmission and children are more frequently and more severely infected than adults. Swimming is the contact activity that contributes most to transmission. Case studies illustrating the roles of high temperatures, hydrology and over-fishing in the epidemiology of the disease in the region are also reviewed.     

Heat shock proteins and survival strategies in congeneric land snails (Sphincterochila) from different habitats

Polmunate land snails are subject to stress conditions in their terrestrial habitat, and depend on a range of behavioural, physiological and biochemical adaptations for coping with problems of maintaining water, ionic and thermal balance. The involvement of the heat shock protein (HSP) machinery in land snails was demonstrated following short-term experimental aestivation and heat stress, suggesting that land snails use HSPs as part of their survival strategy. As climatic variation was found to be associated with HSP expression, we tested whether adaptation of land snails to different habitats affects HSP expression in two closely related Sphincterochila snail species, a desert species Sphincterochila zonata and a Mediterranean-type species Sphincterochila cariosa. Our study suggests that Sphincterochila species use HSPs as part of their survival strategy following desiccation and heat stress, and as part of the natural annual cycle of activity and aestivation. Our studies also indicate that adaptation to different habitats results in the development of distinct strategies of HSP expression in response to stress, namely the reduced expression of HSPs in the desert-inhabiting species. We suggest that these different strategies reflect the difference in heat and aridity encountered in the natural habitats, and that the desert species S. zonata relies on mechanisms and adaptations other than HSP induction thus avoiding the fitness consequences of continuous HSP upregulation.     

Genotyping natural infections of Schistosoma mansoni in Biomphalaria alexandrina from Damietta, Egypt, with comparisons to natural snail infections from Kenya

The distribution of Schistosoma genotypes among individuals in snail populations provides insights regarding the dynamics of transmission and compatibility between schistosome and snail hosts. A survey of Biomphalaria alexandrina from Damietta (Nile Delta, Egypt), an area subjected to persistent schistosomiasis control efforts, provided only 17 snails infected with Schistosoma mansoni (6.1% overall prevalence), each shown by microsatellite analysis to have a single genotype infection. By contrast, recent studies of uncontrolled S. mansoni transmission foci in Kenya revealed that 4.3% Biomphalaria pfeifferi and 20-25% Biomphalaria sudanica snails had multiple genotype infections. Compared with the 3 Kenyan populations, the Egyptian population of S. mansoni also showed a lesser degree of genetic variability and was genetically differentiated from them. We suggest that tracking of genotype diversity in infected snails could be further developed to serve as an additional and valuable independent indicator of efficacy of schistosomiasis control in Egypt and elsewhere.     

Host--parasite relationships of Schistosoma japonicum in mammalian hosts.

Control of schistosomiasis caused by Schistosoma japonicum has been severely hindered by the fact that several non-human mammalian species, including domesticated as well as wild animals, serve as zoonotic carriers of this infection. For effective control, it is imperative that the full host spectrum of this infection is understood. Although about 46 species of mammals are known to carry natural infection with S. japonicum, only a few might be of potential threat to human infection. Generally, in an endemic area, transmission of schistosomiasis to human depends largely on the availability and abundance of permissive hosts. Another important factor that needs to be taken into consideration in developing control measures against S. japonicum is potential strain differences. This review collates pertinent host-parasite relationship of S. japonicum in mammals in an endemic area and assesses the epidemiological significance of these findings for human infection.     

Relictual physiological ecology in the threatened land snail Codringtonia helenae: A cause for decline in a changing environment?

Land snails often exhibit intra-annual cycles of activity interspersed by periods of dormancy (hibernation/aestivation), accompanied by a range of behavioural and physiological adaptations to ensure their survival under adverse environmental conditions. These adaptations are useful to understand species-specific habitat requirements and to predict their response to environmental changes. We examined the seasonal physiological and biochemical composition patterns of the threatened land snail Codringtonia helenae, endemic to Greece, in relation to its behavioural ecology and climatic conditions. Fuel reserves (glycogen, lipids, proteins) and water were accumulated prior to aestivation, but subsequently were rapidly depleted. LDH exhibited substantial rise during aestivation, suggesting that anaerobic pathways may provide additional energy. The major outcome of our study was the unambiguous discrimination of the four life-cycle periods. Most remarkable was the clear distinction of the aestivation period, with hibernation, the other dormancy period, showing similarity with the two active periods but not with aestivation. We observed disassociation between behavioural and physiological responses and climatic conditions. The physiological responses of C. helenae were effective during hibernation, but only partly compensate the effect of adverse conditions during aestivation, since its aestivating behaviour is occasional and time limited. Perhaps, the behavioural ecology of Codringtonia is relictual and shaped during past environmental conditions. This constitutes an important extinction threat considering the current climatic trends and the deterioration of the habitat of that species due to human activities.     

Genetic diversity and recruitment pattern of Schistosoma mansoni in a Biomphalaria glabrata snail population: a field study using random-amplified polymorphic DNA markers.

Random-amplified polymorphic DNA markers have been used to assess the amount and the distribution of the genetic diversity of Schistosoma mansoni within a natural population of Biomphalaria glabrata at a transmission site of the murine schistosomiasis focus of Guadeloupe. Despite high infection rate and heavy schistosome load within the definitive hosts (Ratus rattus), prevalences within intermediate snails ranged from 0.2 to 4.8%. Whatever the transmission season may be (rainy vs. dry), most of the infected snails were spatially aggregated and 88.4% of them harbored a single parasite genotype indicative of a monomiracidial infection; 4.7% had dual sex infections and a parasite intensity not exceeding 3 miracidia per snail. A substantial resistance level toward the parasite and recruitment regulatory process within snails may explain in part the observed low parasite prevalences and intensities. Considering such a distribution pattern of larval S. mansoni genetic diversity among B. glabrata, mobility of the definitive hosts, or rapid turnover of infected snails, or both, are required to maintain genetic heterogeneity within adult schistosome populations.     

Three Gorges Dam and Its Impact on the Potential Transmission of Schistosomiasis in Regions along the Yangtze River

Large-scale hydroprojects have a propensity for incurring schistosomiasis epidemics by altering the environments of their vicinities. As the construction of the Three Gorges Dam, one of the world’s largest hydroprojects to date, draws near its conclusion, an assessment of the dam’s capacity in causing schistosomiasis becomes more urgent and pressing. This article reviews recent investigations into the possible effects of the dam on schistosomiasis in the Three Gorges region and areas along the Yangtze downstream from the dam. Data used in this article were extracted from peer-reviewed papers found in PubMed, Chinese Journal of Parasitology and Parasitic Diseases, and Chinese Journal of Schistosomiasis Control. Results indicate that the Three Gorges Dam is capable of inducing a wide variety of environmental and ecological changes both within the Three Gorges region and in downstream areas. These changes, however, carry ambivalent implications for the reproduction of Oncomelania snails and the spreading of schistosome infections. Furthermore, major changes in the demographics and agricultural practices of the Three Gorges and downstream Yangtze areas caused by the dam could also exert significant influence on the transmission of schistosomiasis in these regions. Major conclusions of this review include the need for further ecological simulations of the Three Gorges Dam and the need for deploying monitoring and intervention systems to provide successful prophylaxis of the Three Gorges Dam-associated schistosomiasis emergence.     

First report on the presence of Biomphalaria straminea in the municipality of Jaboticatubas,State of Minas Gerais,Brazil

This is the first report on occurrence of Biomphalaria straminea in the district of S?o José de Almeida (municipality of Jaboticatubas) State of Minas Gerais, Brazil. The presence of B. glabrata and B. tenagophila had already been reported in this area. Such municipality is part of the metropolitan region of Belo Horizonte and comprises 60% of the Tourist Complex of Serra do Cipó. Since the 1950s throughout the 1990s, a schistosomiasis prevalence ranging from 15 to 40% has been observed. Although no B. straminea specimen has been found naturally infected in the region, descendants of these snails collected in the area, showed to be experimentally susceptible to Schistosoma mansoni infection reaching rates from 14.6 to 28.6%. Even not being found naturally infected, in the State of Minas Gerais, the possibility that the species B. straminea may keep endemicity foci of schistosomiasis should be regarded, as in the Northeastern region of Brazil where the high density of this planorbid and the social-economic and sanitary conditions enable to the transmission.     

Natural history of Theba pisana in Israel (Pulmonata: Helicidae)

Theba pisana in southern Israel is a semelparous species with an annual life cycle. Young snails hatch in early winter, grow rapidly in spring and by summer they reach, first adult shell-size, then adult shell-weight. Shell-size is negatively related to density, suggesting that, as in some other landsnails, there may be a density-dependent inhibition of growth. Summer forces the snails into 3–4 months of aestivation.
After the first winter rains the snails, awakened perhaps by the sudden drop in barometric pressure, emerge from aestivation. Only now are most of the spermatozoa and mature oocytes formed (maturation of gametes actually starts some two weeks before the first rains), and the snails copulate. No evidence of precocious breeding was found. The snails sometimes dig a few "sham" holes, perhaps to discourage egg-eating predators, before finally depositing their eggs (mean clutch size: 80 eggs) in the last hole.
In mid-winter, the snails die. In a habitat where the sheltering conditions of the vegetation offer the snails more protection, the dying off process of the population is stretched over a few more months. This could, perhaps, be the first step towards the evolution of a bi-annual cycle, such as is typical of T. pisana in some other areas of its distribution.     

Identification of a genetic marker associated with the resistance to Schistosoma mansoni infection using random amplified polymorphic DNA analysis

In schistosomiasis, the host/parasite interaction remains not completely understood. Many questions related to the susceptibility of snails to infection by respective trematode still remain unanswered. The control of schistosomiasis requires a good understanding of the host/parasite association. In this work, the susceptibility/resistance to Schistosoma mansoni infection within Biomphalaria alexandrina snails were studied starting one month post infection and continuing thereafter weekly up to 10 weeks after miracidia exposure. Genetic variations between susceptible and resistant strains to Schistosoma infection within B. alexandrina snails using random amplified polymorphic DNA analysis technique were also carried out. The results showed that 39.8% of the examined field snails were resistant, while 60.2% of these snails showed high infection rates.In the resistant genotype snails, OPA-02 primer produced a major low molecular weight marker 430 bp. Among the two snail strains there were interpopulational variations, while the individual specimens from the same snail strain, either susceptible or resistant, record semi-identical genetic bands. Also, the resistant character was ascendant in contrast to a decline in the susceptibility of snails from one generation to the next.     

The relationship between genetic variability and the susceptibility of Biomphalaria alexandrina snails to Schistosoma mansoni infection

In the present study, Biomphalaria snails collected from five Egyptian governorates (Giza, Fayoum, Kafr El-Sheikh, Ismailia and Damietta), as well as reference control Biomphalaria alexandrina snails from the Schistosome Biological Supply Center (SBSC) (Theodor Bilharz Research Institute, Egypt), were subjected to species-specific polymerase chain reaction (PCR) assays to identify the collected species. All of the collected snails were found to be B. alexandrina and there was no evidence of the presence of Biomphalaria glabrata. Randomly amplified polymorphic DNA (RAPD)-PCR assays showed different fingerprints with varying numbers of bands for the first generation (F?) of B. alexandrina snail populations (SBSC, Giza, Fayoum, Kafr El-Sheikh, Ismailia and Damietta). The primer OPA-1 produced the highest level of polymorphism and amplified the greatest number of specific bands. The estimated similarity coefficients among the B. alexandrina populations based on the RAPD-PCR profiles ranged from 0.56 (between SBSC and Ismailia snails) to 0.72 (between Ismailia and Kafr El-Sheikh snails). Experimental infection of the F? of progeny from the collected snails with Schistosoma mansoni (SBSC strain) showed variable susceptibility rates ranging from 15% in the Fayoum snail group to 50.3% in SBSC snails. A negative correlation was observed between the infection rates in the different snail groups and the distances separating their corresponding governorates from the parasite source. The infection rates of the snail groups and their similarity coefficients with SBSC B. alexandrina snails were positively correlated. The variations in the rates of infection of different B. alexandrina groups with S. mansoni, as well as the differences in the similarity coefficients among these snails, are dependent not only on the geographical distribution of the snails and the parasite, but also on the genetic variability of the snails. Introduction of this variability into endemic areas may reduce the ability of the parasite to infect local hosts and consequently reduce schistosomiasis epidemiology.     

Location of Biomphalaria glabrata (Say) by miracidia of Schistosoma mansoni Sambon in natural standing and running waters on the West Indian Island of St. Lucia

Upatham E. S. 1973. Location of Biomphalaria glabrata (Say) by miracidia of Schistosoma mansoni Sambon in natural standing and running waters on the West Indian Island of St. Lucia. International Journal of Parasitology3: 289–297. The ability of S. mansoni miracidia to locate B. glabrata in natural ditches and streams was investigated. Miracidia located and infected snails at distances of 9–14 and 97-54 in horizontally in standing and running waters respectively. In running water, no infection occurred above a velocity of 13.11 $\text{cm}\text{sec}$. In both types of habitat, infection rates in snails increased with increasing levels of miracidia but decreased as the location of caged snails moved away from the miracidial point of entry. Laboratory experi- ments showed that the number of daughter sporocysts was proportional to the number of miracidia. Judging by the number of daughter sporocysts recovered only a small percentage of miracidia succeeded in locating and penetrating snails (6.8–13-7 % and 1.4–6.2 % in standing and running waters respectively). In standing water, infection may be inhibited by the limited ability of miracidia to move horizontally. In running water, the flow extends significantly the effective scanning capacity of the miracidia, giving them a better chance of coming into contact with snails, which is of importance in the epidemiology of schistosomiasis. Owing to a con- siderable wastage of miracidia and the higher relative efficiency of miracidia at lower densities in detecting snails, control measures such as chemotherapy or provision of safe water supplies designed to lower egg output and reduce contamination may not seriously influence transmission unless S. mansoni egg production or contamination is massively reduced.     

Distributional ecology of snails (Gastropoda: Mollusca) in seasonal ponds of Tamil Nadu,South India

The present study was focused to examine diversity and distributional pattern of freshwater snails and to identify the schistosome infected snails in ten different ponds of Madurai district, Tamil Nadu, South India. Snails were collected from native substrates in ponds using dip net from Madurai North, Tamil Nadu province of south India and microscopic examination for detecting cercariae larvae in snails were assessed. In total, 1250 individuals belonging to six species under five families of freshwater snails were collected. The alpha and beta diversity analyses were calculated for estimating the diversity of snails. The cercariae infection was observed in four collected freshwater snail species. Of four infected snails, Lymnaea luteola had the highest percentage of cercariae infection. Statistical tests showed that the environmental factors of total dissolved solids, pH and salinity were significantly related with snail distribution and cercariae infection rather than spatial factor. The study demonstrated that the salinity, conductivity and total dissolved solids of pond water play a key factor for the distribution of snails and cercariae infection. The findings of our study will improve the understanding of the dynamics of schistosomiasis transmission and would ease the implementation of integrated disease control measures based on ecological knowledge of schistosomiasis vector in Madurai and Tamil Nadu zones.