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PubMed1991

Ultrastructure and microanalyses of the protoscolex hooks of Echinococcus granulosus.

The rostellar distal cytoplasm of Echinococcus granulosus protoscoleces is characterized by extensive basal membrane infolding, prominent hemidesmosomes and is subtended by a lamina reticularis with microfibrils of approximately 10 nm diameter that occasionally show a 55 nm banding periodicity. The rostellar hooks, in 2 rows, each have a blade, guard and handle region and possess a central amorphous pulp, a middle microfibrillar medulla with microfibrils of approximately 4 nm diameter, and a complex outer cortex in all but the proximal region of the guard and the base of the handle. In these regions additional material, of similar electron density to the medulla, but lacking the fibrillar substructure, occurs and gives the areas a lobed appearance. Energy-dispersive X-ray microanalysis of whole hooks demonstrated the presence of sulphur and trace quantities of phosphorus. X-ray near-edge absorption spectra resembled those of cystine, feather and hair and showed the sulphur to be predominantly in the form of disulphide linkages. X-ray diffraction patterns of whole hook preparations revealed 2 diffuse rings with equatorial spacings of 7.99 A and 15.22 A, thus differing from vertebrate keratins.

Animals↗
PubMed1992

Developmental changes of Echinococcus multilocularis metacestodes revealed by tegumental ultrastructure and lectin-binding sites.

Ultrastructural investigations (SEM, TEM) combined with lectin-binding analysis, have revealed concurrent modifications in tegumentary structure and surface glycoconjugates during the establishment and differentiation of Echinococcus multilocularis metacestodes in jirds. The laminated layer, which is amorphous and rich in polysaccharides when initially secreted by the young cyst, takes on a different appearance and has a different glycoconjugate composition according to whether the cyst becomes fertile or sterile. The laminated layer of fertile cysts transforms into a microfibrillar matrix, the protein content of which may increase while sugar content decreases during protoscolex differentiation. Independently of this structure, brood capsules, from which arise protoscoleces, are formed by invagination of the cyst tegument. The intense secretion of glycoconjugates from the brood capsule wall during invagination may serve to interact with host factors passing through the laminated layer. The combined use of ultrastructural study and lectin labelling has allowed the demonstration of an ultrastructural and biochemical gradient of differentiation of the protoscolex. Seven stages of differentiation have been described. The possibility that the excreted-secreted tegumentary glycoconjugates, revealed by lectin labelling during protoscolex differentiation, might be the gradual biochemical expression of one or several stimuli implicated in the phenomenon of protoscolex maturation, is discussed.

Animals↗
PubMed1989

Genetic heterogeneity within Echinococcus granulosus: isolates from different hosts and geographical areas characterized with DNA probes.

A segment of the ribosomal RNA gene of Schistosoma mansoni and a DNA fragment specific to Echinococcus granulosus, cloned in plasmids, have been used as DNA probes to assess the extent of genetic variability within E. granulosus and some distinct strains have been identified. The DNA analysis, involving restriction endonuclease digestion and Southern blot hybridization with the probes, did not demonstrate any significant genetic variation within the U.K. horse/dog or sheep/dog strains but confirmed the distinctiveness of the two strains shown in previous studies. The sheep/dog strain was shown to be cosmopolitan in its distribution and fertile bovine material originating from the United Kingdom, Kenya, Spain and India conformed to this strain by DNA hybridization. In contrast, cattle isolates from Holland produced markedly different DNA hybridization banding profiles indicating that cattle can harbour more than one strain of E. granulosus. Similarly, it was shown that goats can harbour two different strains of E. granulosus, the sheep/dog strain and a form which infects camels. The strain of E. granulosus infecting equines in Spain and Ireland is genetically identical to that infecting horses in the United Kingdom. There is also a different strain infecting pigs in Poland and Yugoslavia. This pig/dog strain appears to be very similar genetically to the forms of E. granulosus which use camels and goats as intermediate hosts and is similar, though not identical, to the variant infecting Dutch cattle. It has been shown that E. granulosus material, fixed for a prolonged period in ethanol, or lyophilized, is amenable to DNA analysis and that it is possible to characterize the DNA of a single adult worm.

Animals↗
PubMed1990

Regulation of macrophage-mediated larvicidal activity in Echinococcus granulosus and Mesocestoides corti (Cestoda) infection in mice.

Killing of metacestodes by normal or post-infection macrophages and the regulation of this activity by cytokines were studied in vitro. The protoscolecidal activity of normal macrophages against Echinococcus granulosus was inhibited by a product of naive T-enriched lymphocytes co-cultured with protoscoleces (PSC). By contrast, supernates from co-cultures of Mesocestoides corti tetrathyridia (MCT) and T-enriched or B-enriched normal lymphocytes increased killing of MCT by normal macrophages. Larvicidal activity (against both PSC and MCT) was enhanced by high concentrations of macrophage-activating factors produced by Con A-stimulated rat lymphocytes (Con A-LK), but was reduced by low concentrations of these factors. Activation by synergism between Con A-LK and recombinant interferon-gamma(r. IFN-gamma) was demonstrated in macrophage-mediated killing of MCT at high effector to target ratio. Cytokine-activation of normal or post-MCT infection macrophages was compared. Macrophages from both 8 and 20 week post-infection mice were refractory to lymphokines from lymphocyte-MCT cultures and displayed greatly reduced killing of MCT. Macrophage activation by Con A-LK and r.IFN-gamma was also impaired, implying a general defect in the ability of these post-infection macrophages to respond to macrophage activating signals. The data indicate that two different mechanisms may exist by which metacestodes regulate potentially larvicidal effector mechanisms. E. granulosus can elicit the production of lymphokines suppressive for PSC killing, whereas M. corti appears directly to induce a refractory state in effector macrophages.

Animals↗
PubMed1990

Rostellar hook morphology of Echinococcus granulosus (Batsch, 1786) from natural and experimental Australian hosts, and its implications for strain recognition.

An analysis of the rostellar hooks of Australian isolates of Echinococcus granulosus revealed that there was less variation in larval (metacestode) than adult characters and that metacestode characters could be measured directly from adult worms. A factor analysis indicated that two factors, one representing a contrast between number of hooks and their length, and the other representing blade lengths, were sufficient to account for 87.5% of the variance in metacestode hook measurements. These results indicate that rostellar hook morphology is not useful for discriminating strains of E. granulosus in Australia. The Tasmanian and mainland domestic strains were found to be indistinguishable using rostellar morphology. Although many of the isolates from sylvatic hosts differed from those from domestic hosts, there was not a clear separation as would be expected if two distinct strains existed. Evidence was presented to show that the morphological differences seen in sylvatic hosts could be attributed to host-induced effects, and that the previously accepted existence of two mainland strains should be investigated further.

Animals↗
PubMed1990

Genetic diversity and genetic differentiation in Echinococcus granulosus (Batsch, 1786) from domestic and sylvatic hosts on the mainland of Australia.

Enzyme electrophoresis was used to examine genetic variation within and between populations of Echinococcus granulosus from domestic and sylvatic hosts in western and eastern Australia. Substantial genetic diversity was found within all populations. There was no evidence, however, of genetic differentiation between populations from different hosts or geographic areas. When isolates were grouped into previously described domestic or sylvatic strains on the basis of rostellar hook morphology, most (94%) of the genetic variation occurred within, rather than between strains. These results conflict with the currently accepted theory of separate domestic and sylvatic strains of E. granulosus on the mainland of Australia.

Alleles↗
PubMed1986

Echinococcus granulosus: absorption of cycloleucine and alpha-aminoisobutyric acid by protoscoleces.

Protoscoleces of Echinococcus granulosus absorb the amino acids cycloleucine and alpha-aminoisobutyric acid (AIB) by a combination of mediated uptake and diffusion. After correcting for the latter, values for Kt and Vmax of 0.124 mM and 0.947 nmoles/mg protein/2 min for cycloleucine were calculated; corresponding values for AIB were 0.039 mM and 0.139 nmoles/mg protein/2 min. Both amino acids were accumulated against a concentration gradient and a comparison of Kt and Ki values determined in mutual inhibition experiments suggested that both cycloleucine and AIB share a common uptake locus (loci). Cycloleucine uptake was pH-dependent and could be inhibited by a variety of other amino acids. Neither D- nor L-proline inhibited cycloleucine absorption but D-methionine, D-alanine, D-leucine, D-valine and D-serine were much more effective inhibitors than their L-counterparts.

Absorption↗
PubMed1986

In vitro development of hydatid cysts from posterior bladders and ruptured brood capsules of equine Echinococcus granulosus.

A variety of morphological forms developed during monophasic culture of ruptured brood capsules of equine Echinococcus granulosus. Attached to some protoscoleces were small posterior bladders, with a surface ultrastructure of truncated microtriches, and these never developed a laminated layer. In contrast, small free vesicles of similar size and ultrastructure, but devoid of an attached protoscolex, produced a layer by 20-30 days. The surface ultrastructure of both these forms is similar to that of the protoscolex attachment stalk and the brood capsule lining, and posterior bladders and small free vesicles are considered to be derived from portions of the stalk +/- part of the brood capsule lining retained at protoscolex release. Also present were large vesicles interpreted from their size and surface anatomy as ruptured, everted brood capsules. When devoid of attached protosoleces these formed cysts each with a laminated layer, whereas similar vesicles with several attached protoscoleces did not develop a layer, or only very occasionally after 70 or more days. In contrast, a few large vesicles with a single attached protoscolex produced a laminated layer. The possibility that protoscolex attachment to stalk remnants or brood capsules inhibits the cystic development of these structures is discussed.

Animals↗
PubMed1986

Echinococcus granulosus infection in mice: host responses during primary and secondary infection.

The reaction of Balb/c mice to primary and secondary subcutaneous infection with Echinococcus granulosus protoscoleces (PSC) is described. From 3 to 14 days following primary exposure to PSC, draining lymph nodes increase in weight and there is expansion of T and B lymphocyte populations, enhancement of in vitro lymphocyte transformational responses and production of PSC-specific IgM and IgE antibodies. Despite the persistence of viable PSC in host tissues, lymphocyte responses decline to pre-infection values over the period 3-8 weeks post-infection. Secondary exposure to PSC immediately induces lymphoproliferation, enhancement of transformational responses, production of IgE antibody and encapsulation of PSC by inflammatory cells. Although specific antibody levels remain high until at least 8 weeks after challenge infection, lymphocyte activity begins to decline after 4 days and is profoundly suppressed by 10 days. Parasite viability appears to be significantly reduced in secondary, as opposed to primary, infection and is associated with the accumulation of large numbers of eosinophils, mast cells and macrophages in infected tissues.

Animals↗
PubMed1986

Population dynamics in echinococcosis and cysticercosis: biological parameters of Echinococcus granulosus in dogs and sheep.

The numerical distributions of Echinococcus granulosus in an experimental dog population are described. At all dose rates of protoscoleces from 10 to 175000 the distribution of worms was over-dispersed. Host age had no effect. There was a direct proportionality between the infective-stage density and rate of infection, and between the latter and the index of clumping. The worm burdens were significantly higher in the proximal than distal portions of the small intestine. Lengths of the 3- and 4-segmented worms increased from 4 to 10 and 4 to 8 weeks of age, respectively. Thereafter apolysis was asynchronous and could not be determined. Eggs were first detected in the faeces at 6 weeks and the mean age at oogenesis was 7.26 weeks. Retarded growth of the whole population of worms was observed in some dogs. For the first few infections, worm burdens varied widely in the same dog, but by the 6th infection 50% of the dog population had developed a relative insusceptibility to infection. Growth or oogenesis of the worms were not affected. A short-acting immune response was artificially induced in some dogs following the parenteral injection of activated embryos of E. granulosus, Taenia hydatigena, T. ovis, T. multiceps, T. pisiformis and T. serialis. The response affected either the number of worms established, growth or oogenesis or all three parameters. There was a strong positive correlation between numbers and lengths of worms in dogs with acquired and induced immunity, indicating that no 'crowding' effects were involved. In sheep populations the mean number of cysts which established was directly proportional to the number of eggs given, implying that there was no negative feedback mechanism operating at this stage of the life-cycle. The distribution of the larval population in sheep was over-dispersed and the index of clumping increased with the size of the egg dose from 25 to 2500 eggs. Protoscoleces were first observed in cysts at 2 years and the proportion producing them increased with age, with an estimate of 50% of cysts containing protoscoleces at 6.29 years. No deaths were observed in dogs or sheep even when high parasite burdens were present, implying that E. granulosus does not regulate the population of its hosts.

Age Factors↗
PubMed1986

Population dynamics in echinococcosis and cysticercosis: mathematical model of the life-cycle of Echinococcus granulosus.

A mathematical model of the life-cycle of Echinococcus granulosus in dogs and sheep in New Zealand is constructed and used to discuss previously published experimental and survey data. The model is then used to describe the dynamics of transmission of the parasite, and the means by which it may be destabilized. It is found that under the conditions that prevailed in New Zealand during the late 1950s, at the time of surveys of this parasite, the dog-sheep life-cycle was not regulated by any effective density-dependent constraint. In contrast there was evidence for an effective acquisition of immunity to reinfection by cattle. The long time to maturity of the cyst in sheep, together with the practice of feeding aged sheep to dogs, provides a time delay in the intermediate host. By comparison, the time to maturity of the adult stage in dogs is short, but it is of sufficient magnitude to be a key factor in the destabilization of the system by a regular dog-dosing programme. The model used to describe the life-cycle is a linear integrodifferential equation of the Volterra type. Such equations are intrinsically unstable in that a small perturbation in parameters can drive a previous equilibrium solution to zero. At the time of the surveys, the value of the basic reproductive rate, R0, was close to 1, and it has since been reduced below 1 by control measures.

Age Factors↗
PubMed1993

Detection of Echinococcus multilocularis DNA in fox faeces using DNA amplification.

In order to identify Echinococcus multilocularis DNA in fox faeces for epidemiological purposes, we have developed a new method to prepare DNA suitable for PCR amplification. DNA isolation from fox excrement was performed according to a novel procedure involving lysis in KOH, phenol-chloroform extraction and a purification step on a matrix (Prep-A-Gene). The target sequence for amplification was the E. multilocularis U1 snRNA gene. PCR products were indistinguishable for 32 different E. multilocularis isolates and no signal was observed after ethidium bromide staining with DNAs from other tapeworm species, including E. granulosus. The sensitivity of amplification was monitored by the addition of E. multilocularis DNA or eggs to faeces free of E. multilocularis and was estimated to be 1 egg per 4 g of faeces. PCR products were blotted onto nylon membranes and hybridized with an internal oligonucleotide probe in order to confirm the results. Twenty nine faecal samples from foxes shot in Franche-Comté (East France) were tested. Out of 10 samples from foxes in which no E. multilocularis adult worms could be observed after necropsy, 7 were PCR positive, showing that the PCR test is more sensitive than microscopical examination. Out of 19 samples from foxes harbouring E. multilocularis adult worms, 18 were PCR positive. The remaining PCR-negative sample could be due either to the misidentification of the species of adult worm (E. granulosus and E. multilocularis), or to DNA variation between different isolates of E. multilocularis. Further work in the field should be initiated in order to confirm these results.

Animals↗
PubMed1994

Molecular genetic characterization of the cervid strain ('northern form') of Echinococcus granulosus.

We have used a number of molecular genetic approaches to characterize the cervid strain ('northern form') of Echinococcus granulosus. PCR-RFLP analysis of the nuclear ITS1 region of the rDNA repeat can readily distinguish the cervid form from other strains of E. granulosus. The complexity of the RFLP patterns obtained suggests that a number of distinct ITS1 types are present in this strain which may represent an inter-strain E. granulosus hybrid. Mitochondrial CO1 sequence of the cervid genotype was ambiguous at 18 positions and closely resembles a cluster of previously characterized E. granulosus genotypes, G1 (common, domestic sheep)/G2 (Tasmanian sheep)/G3 (buffalo). In contrast, mitochondrial ND1 sequence, although unique, suggests that the cervid form is most similar to strains represented by the G6 (camel)/G7 (pig) genotypes. We assume that the CO1 and ND1 sequences obtained for the cervid genotype are linked in a single mitochondrial genome although this is difficult to explain if conventional molecular genetics of mitochondrial DNA are assumed. Based on its unique ND1 sequence and ITS1 PCR-RFLP pattern, the cervid strain appears to represent a distinct genotype (designated G8) of E. granulosus.

Amino Acid Sequence↗
PubMed1994

Immunocytochemical localization of serotonin (5-HT) in the nervous system of the hydatid organism, Echinococcus granulosus (Cestoda, Cyclophyllidea).

The localization and distribution of the serotoninergic components of the nervous system in the hydatid organism, Echinococcus granulosus, were determined by immunocytochemical techniques in conjunction with confocal scanning laser microscopy (CSLM). The distribution of serotonin immunoreactivity (IR) paralleled that previously described for cholinesterase activity, although it was more widespread. Nerve cell bodies and nerve fibres immunoreactive for 5-HT were present throughout the central nervous system (CNS), occurring in the paired lateral, posterior lateral and rostellar ganglia, their connecting commissures and nerve rings in the scolex and in the ten longitudinal nerve cords that run posteriorly throughout the body of the worm. A large population of nerve cell bodies was associated with the lateral nerve cords. In the peripheral nervous system (PNS), immunoreactive nerve fibres occurred in well-developed nerve plexuses innervating the somatic musculature and the musculature of the rostellum and suckers. The genital atrium and associated reproductive ducts were richly innervated with serotoninergic nerve cell bodies and nerve fibres.

Animals↗
PubMed1993

The potential of vervet monkeys (Cercopithecus aethiops) and baboons (Papio anubis) as models for the study of the immunology of Echinococcus granulosus infections.

Nine vervet monkeys and nine baboons were infected with eggs of Echinococcus granulosus per os. Six of the vervets and one of the baboons possessed hydatid cysts at autopsy, 15-28 months post-infection. The sequential IgG response to hydatid fluid and protoscolex antigens showed considerable inter-animal variation. Infected vervets and baboons became seropositive after an average of 8 months post-infection. Considerable fluctuation in the IgG response was observed, particularly to the hydatid fluid antigen which, in humans, may contribute to the existence of a significant proportion of seronegative individuals. Vervets, in particular, may be useful to study immunological events associated with exposure, development and resolution of hydatid disease in outbred human populations.

Animals↗
PubMed1990

Development of larval Echinococcus granulosus confronted with chicken heart tissue in vitro.

Vesiculated protoscoleces (VP) were produced by culturing freshly collected protoscoleces from Echinococcus granulosus horse liver hydatids in RPMI 1640 monophasic medium at 37 degrees C for 18 days. Half of the VP were used as such, the other half used after killing them by freeze-thawing. Nine-day-old chicken heart fragments (CHF) were cultured in MEM at 37 degrees C for 72 h. Subsequently, CHF were put together with live and dead VP, respectively, for up to 53 days, on a semisolid medium consisting of agar, Ringer's and MEM. Time-dependent histological observations revealed that dead VP were surrounded by CHF cells. Dead VP tissue was eventually internalized and disintegrated in about 1 week. Live VP penetrated into the CHF tissue and further developed into small hydatid cysts, located within the boundaries of the experimental 'host' tissue. The amorphous-looking contact region PAP-stained positively only with anti-E. granulosus serum and not with anti-CHF serum; it was considered identical to the normal laminated layer. The invasion of VP in CHF tissue proved to be different from a tumour or a bacterial invasion: it was concluded that the confrontation of VP and CHF had resulted in an 'in vitro cohabitation' rather than in an 'in vitro infection'.

Animals↗
PubMed1993

Echinococcus multilocularis: molecular and immunochemical characterization of diagnostic antigen II/3-10.

A recombinant Echinococcus multilocularis antigen (II/3-10), which had previously been shown to exhibit immunodiagnostic characteristics highly specific for human alveolar echinococcosis, and the corresponding native parasite antigen, were further characterized with immunochemical and molecular biological methods. Immunoblot analysis using a polyclonal antiserum raised in rabbits against the recombinant protein, and subsequent Northern hybridization analysis, revealed that the native antigen was expressed by E. multilocularis at the adult as well as at the metacestode stage. In metacestodes, the antigen was shown by using indirect immunofluorescence and the same antiserum to be localized within the germinal layer and membrane structures of developing protoscolices. Electrophoretic analyses revealed remarkable differences in the apparent molecular weight of the antigen under reducing and non-reducing conditions. In further immunoblot analyses, anti-II/3-10 antibodies identified the corresponding epitopes on bands with identical M(r) in all E. multilocularis isolates investigated (European, Asian and North American). By Southern hybridization analyses of the respective gene, phylogenetically related sequences were shown to be present in other helminth species such as E. granulosus and several Taenia spp. In the same respect, immunoblotting revealed that anti-II/3-10 antibodies reacted with antigens of different M(r) from various E. granulosus isolates and some other cestode species, indicating the presence of shared and thus potentially cross-reacting epitopes. The relevance of these findings for the immunodiagnostic performance of the recombinant antigen is discussed.

Amino Acid Sequence↗
PubMed1995

A molecular phylogeny of the genus Echinococcus.

Three nucleotide data sets, two mitochondrial (COI and ND1) and one nuclear (ribosomal ITS1), have been investigated in order to resolve relationships among species and strains of the genus Echinococcus. The data have some unusual properties in that mitochondrial heteroplasmy was detected in one strain of E. granulosus, and more than one class of ITS1 sequence variant can occur in a single isolate. The data failed to support the hypothesis that E. granulosus, as it is currently viewed, is a single valid species. Rather, the strains of E. granulosus seem to comprise at least three evolutionarily diverse groups, the sheep strain group, bovine strain group and horse strain group. Molecular distances between them are comparable to, or greater than, molecular evolutionary distances observed between recognized species. The affinities of the cervid strain of E. granulosus are unclear because of ambiguous data, but this strain does not appear to be ancestral to others. E. multilocularis may not be distinct from E. granulosus. However, the remaining two species. E. vogeli and E. oligarthrus appear distinct and rather distant from the first two. Based on the results presented here, taxonomic revision of the genus is clearly warranted.

Animals↗