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Population genetics and dynamics at short spatial scale in Bulinus truncatus, the intermediate host of Schistosoma haematobium, in Morocco.

The population biology of the schistosome-vector snail Bulinus truncatus was studied in an irrigation system near Marrakech, Morocco using both genetic and demographic approaches. The population genetic survey was conducted in 4 sites, 2 sites being sampled on 2 separate occasions. Individuals were genotyped at 6 microsatellite loci. No variability was found at 4 loci, and the 2 other loci had less than 4 alleles. The differentiation, both spatial and temporal, among populations was extremely weak. The demographic survey was conducted using 2 capture-mark-recapture analyses in 2 separate sites, the first in 1999 and the second in 2000. The second analysis permitted the estimation of parameters based on recent methodological developments (multisite models). Although these studies provided information on several traits, we report here on dispersal only. Both analyses showed that individual dispersal is of the order of a few hundreds of metres per reproductive life, that is the scale of the whole irrigation area. Both the genetic and demographic studies indicated that this area harbours a single--or no more than a few--populations of B. truncatus. This has implications for our understanding of the coevolutionary process between snails and flukes.

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Compatibility of Schistosoma haematobium, S. intercalatum and their hybrids with Bulinus truncatus and B. forskalii.

Schistosoma haematobium and S. intercalatum readily hybridize with each other producing generations of viable hybrid offspring. Experiments were designed to investigate the infectivity and viability of the S. haematobium x S. intercalatum F1 and F2 hybrid larvae in their two intermediate snail hosts compared with the parental species. Analysis of the data obtained suggested that the S. haematobium male x S. intercalatum female F1 hybrid miracidia were more infective to Bulinus truncatus than to B.forskalii, and also more infective to B. truncatus compared with the parental S. haematobium miracidia. This hybrid was also observed to have a greater cercarial productivity from both intermediate hosts and these cercariae were shown to be more infectious and to have a longer longevity compared with the cercariae of S. haematobium, S. intercalatum and the S. haematobium female x S. intercalatum male F1 hybrid cercariae. The S. haematobium female x S. intercalatum male F1 hybrid was shown not to be very successful in all stages of the investigations. The results indicate that the S. haematobium male x S. intercalatum female F1 hybrid may have many reproductive advantages over the reciprocal hybrid and the parental schistosome species. The significance of the results is discussed in relation to the epidemiological consequences occurring in Loum, Cameroon, and other areas where S. haematobium and S. intercalatum are sympatric and able to hybridize.

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Spatial and temporal variation of life-history traits documented using capture-mark-recapture methods in the vector snail Bulinus truncatus.

The population biology of the schistosome-vector snail Bulinus truncatus was studied in an irrigation area near Marrakech, Morocco, using demographic approaches, in order to estimate life-history parameters. The survey was conducted using 2 capture-mark-recapture analyses in 2 separate sites from the irrigation area, the first one in 1999 and the second one in 2000. Individuals larger than 5 mm were considered. The capture probability varied through time and space in both analyses. Apparent survival (from 0.7 to 1 per period of 2-3 days) varied with time and space (a series of sinks was considered), as well as a square function of size. These results suggest variation in population intrinsic rate of increase. They also suggest that results from more classical analyses of population demography, aiming, for example at estimating population size, should be interpreted with caution. Together with other results obtained in the same irrigation area, they also lead to some suggestions for population control.

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Host-parasite population genetics: a cross-sectional comparison of Bulinus globosus and Schistosoma haematobium.

The genetic population structures of the freshwater snail Bulinus globosus and its trematode parasite Schistosoma haematobium from 8 river sites in the Zimbabwean highveld were compared using randomly amplified DNA (RAPD) markers. There was significant variability between snail populations collected at different sites, but schistosome populations only showed differentiation at a wider geographical scale (between 2 non-connected river systems). For snails, genetic distance was better correlated with proximity along rivers than absolute geographical separation. In contrast, schistosome genetic distance was better correlated with absolute geographical separation than proximity along rivers. These results are consistent with different dispersal mechanisms for snails and schistosomes and the implications for host-parasite coevolution are discussed.

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Analysis of mating systems in the schistosome-vector hermaphrodite snail Bulinus globosus by DNA fingerprinting.

Bulinus globosus, one of the intermediate hosts for Schistosoma, is a hermaphrodite freshwater snail. Multilocus DNA fingerprinting was applied in order to investigate the mating system of this facultatively self-fertilizing species. By analysing DNA fingerprints of eight broods, we have shown that selfed and outcrossed individuals could be unambiguously distinguished by comparison with their parent fingerprints. First, selfed offspring patterns shared a higher number of bands with their mother than did outcrossed ones. Secondly, at least three paternal bands could be detected per outcrossed offspring pattern.

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Population genetic structure of the schistosome-vector snail Bulinus globosus: examining the role of genetic drift, migration and human activities.

Bulinus globosus, one of the intermediate hosts of the genus Schistosoma, is an hermaphrodite freshwater snail species. This species occupies different kinds of environments in Africa, which are generally subjected to large variations in water availability. The mating system of this species is outcrossing, although selfing has been suggested for one population. Here we investigate the genetic structure of populations of B. globosus in natural and human-modified environments from Niger and Ivory Coast using protein electrophoresis. Our goal is to analyse the respective role of the environment (genetic drift and gene flow), the mating system and human activities. Our results indicate a low intrapopulation polymorphism. A large differentiation among populations occurs on the scale of the whole study area whereas neighbouring populations can show little differentiation. These results point to the role of genetic drift and gene flow as the main factors shaping the genetic structure, with human activities playing a role occasionally.

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Ontogenetic reaction norm for binary traits: the timing of phallus development in the snail Bulinus truncatus.

The ontogenetic trajectory of plastic binary traits may provide valuable insights into their evolutionary rate of change. In this paper, the timing of the plastic response of a temperature-dependent sexual polymorphism, aphally, is investigated in the freshwater snail Bulinus truncatus. Aphally is defined as the loss of the male copulatory organ in otherwise hermaphroditic animals. Individuals from two inbred lines were switched at various times during their early development between 25 and 30 degrees C, and their phally status ascertained, in order to evaluate the parameters characterising the ontogenetic reaction norm of aphally to temperature. A series of nested models including parameters for the onset, offset, and the intensity of the response to temperature were fitted to the data, allowing for a wide range of reaction norms. One genotype did not show any variation in aphally ratio with switching temperature, while a switch-point model (onset and offset corresponding to the same developmental point in time) best fitted the second genotype. The results suggest that the plasticity of aphally is expressed before eggs hatch. Their consequences on the evolution of aphally are discussed. More generally, the methodology proposed here can be used to analyse variation in ontogenetic parameters of discrete traits.

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Anthraquinones in Rheum palmatum and Rumex dentatus (Polygonaceae), and phorbol esters in Jatropha curcas (Euphorbiaceae) with molluscicidal activity against the schistosome vector snails Oncomelania, Biomphalaria and Bulinus.

Hot water extracts of Rheum plamatum and Rheum dentatus (from China) showed molluscicidal activity against the snails Oncomelania hupensis, Biomphalaria glabrata and Bulinus globosus, which are vectors of Schistosoma japonicum, S. mansoni and S. haematobium, respectively. Activity was correlated with antraquinones which were identified by HPLC: rhein and chrysophanol-anthron were most active (> 50% dead snails after 2 days in a 0.03% solution). Molluscicidal activity was intermediate with Rheum-emodin and physcion and was not detectable with cinnamic acid or Aloe-emodin. The snail O. hupensis tended to be more sensitive for several compounds than B. glabrata. Extracts of Jatropha curcas seeds (from Mali) showed molluscicidal activity against both B. glabrata and O. hupensis, the latter being the more sensitive snail. The activity was associated with phorbol esters extracted from Jatropha oil. Of the pure phorbol esters tested, 4 beta-phorbol-13-decanoate killed both snail species at a concentration of 0.001% (10 p.p.m). As Jatropha is locally grown in Mali for other purposes, it might potentially be exploited for schistosomiasis control.

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[Development of the sporocysts of Schistosoma bovis after microsurgical transplantation in Bulinus truncatus].

Microsurgical transplantations of Schistosoma bovis daughter sporocysts to uninfected Bulinus truncatus leads to a dedifferenciation of transplanted sporocysts producing an additional generation of daughter sporocysts. These ones settle into the whole digestive gland and produce infective cercariae. For all the parasitosis, a part of the daughter sporocysts born from the transplanted ones, remains in the head-foot region and are in their turn responsible for an active sporocystogeny. Data demonstrate that the cercarial production dynamics is at least in part in relation with the intramollusc dynamics of sporocyst populations. Success of microsurgical transplantation of S. bovis sporocysts permits cloning perspective of this species.

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Observations on the compatibility between Bulinus spp. and Schistosoma haematobium in the Senegal River basin.

Snail-infection experiments were carried out with a number of different species and populations of Bulinus and isolates of Schistosoma haematobium. The parasites came from six localities in the Senegal River basin (SRB), in the Lower Valley (Mbodiene), Middle Valley (Podor, Diatar and Nguidjilone), and Upper Valley (Aroundou and Galladé). Isolates of S. haematobium from the Middle and Upper Valleys all showed some compatibility with laboratory-bred B. truncatus from Mali, but none of these isolates was compatible with laboratory-bred B. truncatus originating from Senegal. Schistosoma haematobium from Diatar (Middle Valley) was compatible with B. senegalensis, whereas S. haematobium from Mbodiene (Lower Valley), which is naturally transmitted by B. globosus, was incompatible with B. senegalensis and B. truncatus. These data demonstrate that different isolates of S. haematobium from different regions of the SRB exhibit distinct intermediate-host specificities, which in turn will have an effect on the epidemiology of the disease, including the periods of transmission. It is apparent that, in addition to B. senegalensis and B. globosus, B. truncatus, the most widespread bulinid snail in the SRB, may be playing a role in the epidemiology of urinary schistosomiasis. This conclusion has obvious implications for the future spread of urinary schistosomiasis in the SRB. Chemical and physical measurements from assorted habitats along the SRB, including pH, temperature, salinity, conductivity, and resistivity, are also reported.

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Bulinus senegalensis (Mollusca: Planorbidae) in northern Nigeria.

Surveys in the South Chad Irrigation Project Area have revealed the presence of substantial populations of the important potential snail host for Schistosoma haematobium, Bulinus senegalensis. This is the first confirmed record of this species outside the Senegambian region and the habitats in which it occurs differ from those previously reported. The implications of this finding for the epidemiology of S. haematobium in the Sahel region are discussed.

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Observations on the growth and population dynamics of Bulinus rohlfsi in an outdoor laboratory at Volta Lake, Ghana.

Two cohorts of Bulinus rohlfsi were reared separately in an outdoor laboratory at Volta Lake, Ghana. One group was kept in lake water with a mud substratum, the other in lake water without mud. Both cohorts were able to tolerate water temperatures ranging from 22.0 to 35.5 degrees C, but growth and survival were significantly greater for the cohort with mud. The overall intrinsic rate of natural increase, r, per fortnight was 1.096 for the snails having the mud substratum and 1.642 for the cohort not having it. The 'r' values were strongly influenced by high early fecundity of the snails in lake water only and low early fecundity of the snails with mud. Oviposition among the latter cohort was suppressed at times of high turbidity in the water above the mud substratum, caused primarily by suspended clay.

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Laboratory selection for increased tolerance to niclosamide in Bulinus truncatus (Gastropoda: Planorbidae) from Iran.

Survivors of Bulinus truncatus from one exposure to niclosamide (Bayluscide) were used to establish the next generation in a five generation long laboratory selection for increased tolerance to niclosamide. In a comparison of the tolerance to the molluscicide between two original and two selected strains about 150 snails of each strain, divided into four size-groups, were exposed to aliquots of the same niclosamide solution on the same day and during the same time of the day. It was shown that five generations of relatively mild selection (30-50% survival) resulted in a significant increased tolerance to niclosamide in snails with only one exposure before start of the selection procedure. A comparison with published data indicated that the unselected strains did not change their tolerance to niclosamide during seven years of laboratory breeding. The strain exposed to niclosamide once in each of 16 generations showed an avoidance reaction to the chemical by crawling out of the water.

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The effect of schistosome infection on the mortality rates of Bulinus globosus and Biomphalaria pfeifferi.

Schistosoma infections of Bulinus globosus increased the per capita mortality rate in the laboratory from 0.231 to 0.406 per week. Schistosoma mansoni infections of Biomphalaria pfeifferi increased the per capita mortality rate in the laboratory from 0.055 to 0.177 per week. Calculations based on data from previous work by another author indicate an increase in the mortality rate of B. pfeifferi associated with prepatent S. mansoni infection.

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Interference of Echinoparyphium elegans with the host-parasite system Bulinus truncatus-Schistosoma bovis in natural conditions.

Histological observations of Bulinus truncatus naturally infected by both Schistosoma bovis and Echinoparyphium elegans in Sardinia (Italy) have shown that E. elegans is pathogenic to the mollusc and dominant to the schistosome. The larval stages, rediae and metacercariae, acted as compressors. Rediae, mainly the young and very motile ones, also behaved as predators. The organs of the mollusc were variably affected; the ovotestis totally disappeared. The larval development of the schistosome was disturbed; cercariogenesis was strongly inhibited and gave way to high sporocystogenesis. The intensity of this interference depended on the density of echinostome larvae present.

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Further studies on the susceptibility of Bulinus africanus to infection with Schistosoma haematobium.

Genetic manipulation of the intermediate host snails of schistosomiasis has been proposed as a possible method of reducing the rate at which the parasite is transmitted to the final host. This technique is based on the finding that snail-schistosome compatibility is variable, and that refractory snails could be introduced into natural habitats in an attempt to change existing highly-susceptible populations into non-susceptible ones. In our search for such a refractory snail population, offspring from eight different Bulinus africanus populations were infected with Schistosoma haematobium, isolated from school children in the Nelspruit district. A great variation in minimum prepatent period was recorded (33-55 days), while the infected snails surviving that period ranged from 22-89%. A significant difference was found between the infection rates of B. africanus from Newlands in Natal and those of the other seven populations from the Eastern Transvaal Lowveld. The former population could be regarded as partially refractory, and none of the other populations proved to be completely refractory, to infection.

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Gross histopathological effects of an extract of Agave attenuata on the epithelium of the digestive tract of Bulinus africanus.

A lethal concentration of a crude, aqueous extract of Agave attenuata was applied as a contact poison to Bulinus africanus, the intermediate host of Schistosoma haematobium, for a 24-h period. The gross histopathological effects of the extract on the epithelium of the digestive tract were then studied. A graded series of cellular injuries to the epithelial layer was observed along the length of the tract. These included the loss of cilia and brush border, disruption of the epithelial layer, cellular vacuolation, swelling and rupture, and the discharge of secretory products from mucous gland cells. The results of the microscopy show that epithelial tissue is probably a primary target of the molluscicide. The cytological injuries induced by extracts of A. attenuata indicate that the molluscicide acts by disrupting the osmoregulatory mechanisms of the epithelial cells, but further, detailed studies are required to confirm this.

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