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A monoclonal antibody against a glycolipid SEGLx from Spirometra erinaceieuropaei plerocercoid.

A mouse monoclonal antibody AK97 (IgM) was established against a new type of glycosphingolipid, SEGLx, isolated from plerocercoids of tapeworm, Spirometra erinaceieuropaei. The chemical structure of SEGLx (Gal beta1-4(Fuc alpha1-3)(Glc beta1-3Gal beta1-ceramide) had been previously characterized. The specificity of AK97 was determined by thin-layer chromatography-immunostaining and enzyme-linked immunosorbent assay. AK97 was found to be directed to SEGLx and GalSEGLx (Gal beta1-4(Fuc alpha1-3)Glc beta1-3(Gal beta1-6)Gal beta1-ceramide) and also showed cross-reactivity with the stage specific embryonic antigen-1 (SSEA-1), the epitope being defined to be the non-reducing terminal trisaccharide sequence. On immunohistochemical examination, AK97 predominantly stained the tegument, the external surfaces of worms which have a brush border-like organization. Based on the immunohistochemical findings for the staining liability as to organic solvents and the results of Western blot analysis of the plerocercoid glycoproteins, it was proved that the antigens in the tapeworm were glycolipids. Considering that the tapeworm is in direct contact with its host's tissue through the tegument, the membrane surface of which is exposed to the external environment, it is suspected that SEGLx and GalSEGLx on the tegument play functionally important roles in the host parasite interaction.

Animals↗

Control of Echinococcus granulosus in Uruguay: evaluation of different treatment intervals for dogs.

This study attempted control of transmission of Echinococcus granulosus from dogs to sheep in different areas in the Department of Florida, Uruguay, by treating dogs with praziquantel at intervals of 6, 12 and 16 weeks. The 6-week interval was based on the prepatent period of infection with E. granulosus, the 12- and 16-week intervals were based on the rate of reinfection with tapeworms in dogs in the area. Dogs had become reinfected with E. granulosus between 2 and 4 months after treatment, whereas they became reinfected with the Taenia spp. tapeworms within 2 months of treatment. One year after the start of treatments sentinel lambs were born and grazed the farms in the three treatment areas. Approximately, 15 months later when the sentinel lambs were killed and examined for parasites the six weekly treatments had stopped the transmission of E. granulosus to the sentinel lambs. Treatment of dogs at 12- and 16-week intervals failed to stop transmission of E. granulosus but both the numbers of farms and the numbers of sheep infected with E. granulosus were lower where dogs received 12 weekly treatments compared with dogs receiving 16 weekly treatments and a fourth area where dogs had received no treatments (chi(2)P=0.002). Lambs continued to become infected with the Taenia spp. tapeworms in all the areas. Control was complicated by large changes in the dog population. From a starting population of 1164 dogs in the three treated areas, 832 new dogs, most of these adult hunting dogs, entered the population and 793 dogs were lost from the population.

Animals↗

Reliability of coprological diagnosis of Anoplocephala perfoliata infection.

Three coprological methods were tested to establish the reliability of in vivo diagnosis of Anoplocephala perfoliata. A total of 107 faecal samples were analyzed, and the presence of tapeworms were confirmed postmortem in 24 animals with burdens that ranged from 1 to 248 worms; most of them (71%) with less than 100 parasites. Best results were obtained with a combination of two sedimentation/flotation methods, detecting only half the parasitized animals (54% sensitivity). No relationship could be established between tapeworm burden and egg detection, but results indicate that coprological methods have a lower likelihood of diagnosing cestode infection when horses have less than 100 tapeworms.

Animals↗

Life-cycle of Bothriocephalus claviceps, a specific parasite of eels.

The life-cycle of the tapeworm Bothriocephalus claviceps (Goeze, 1782) (Cestoda: Pseudophyllidea), a specific parasite of eels (Anguilla spp.), was studied under experimental conditions. It was demonstrated that the parasite completed its development within 4 months at the temperature range of 22-24 degrees C. Embryonic development was finished in 2 days at this temperature, when coracidia spontaneously hatched. The rate of their development was controlled by water temperature, with delayed formation of coracidia at lower temperatures (8 days at 10-12 degrees C). At 2-4 and 6 degrees C, development did not take place but the eggs remained viable; at 33 degrees C the eggs died. The procercoids developed in copepods of the species Macrocyclops albidus, M. fuscus, Megacyclops viridis, Cyclops strenuus, C. vicinus and Acanthocyclops vernalis (Copepoda: Cyclopidae); their development was finished after 8-12 days at 22-24 degrees C. The definitive hosts, eels, acquired infection after ingestion of infected copepods; the prepatent period of the parasite was more than three months. Perch (Perca fluviatilis) and guppies (Poecilia reticulata) were shown to serve as paratenic hosts of the parasite; in guppies, the tapeworms survived up to 14 days after exposure and they developed similarly to tapeworms in the definitive host.

Anguilla↗

Seasonal abundance of Proteocephalus ambloplitis (Cestoidea: Proteocephalidea) from largemouth bass living in a heated reservoir.

Proteocephalus ambloplitis was recovered from largemouth bass inhabiting different thermal regimes, ranging from normal to greater than 10 degrees C above normal, on the Energy Research and Development Administration's (ERDA) Savannah River Plant near Aiken, South Carolina. Possible alterations in the mechanisms initiating plerocercoid migration as proposed by Fischer & Freeman (1969) were noted. According to these authors, a rise in temperature from 4 to 7 degrees C and above stimulated migration of parenteric plerocercoids into the gut. The lowest water temperature recorded for the reservoir for the entire sampling period was 8.0 degrees C. Theoretically, this should have terminated the life-cycle of this tapeworm since migration temperature was never attained. At Savannah River, the appearance of adult tapeworms coincided with a decrease in water temperature, while the appearance of adult tapeworms in Canada was correlated with an increase in water temperature. It is postulated that the decline in water temperature in southern latitudes and the increase in water temperature in northern latitudes initiates the same response, that is, migration of plerocercoids from parenteric to enteric sites where maturation to the adult form ensues.

Animals↗

Hymenolepis diminuta and H. microstoma: uptake of cyclosporin A and drug binding to parasite cyclophilins.

Cyclosporin A (CsA) acts as a powerful immunosuppressant through its binding to the cytosolic isomerase, cyclophilin (CyP), forming a complex which inhibits the phosphatase activity of calcineurin. The drug is also selectively anti-parasitic but its mode of action remains unknown. The mouse tapeworm, Hymenolepis microstoma is sensitive to CsA, but the rat tapeworm, H. diminuta is not susceptible either in rats, mice or in vitro. Using these two tapeworm models, the uptake and binding of CsA were examined in relation to parasite cyclophilins. Uptake and compartmentalization of the drug were markedly different in the two species: H. microstoma takes up more drug than does H. diminuta and sequesters more drug into intracellular compartments. Characterization of cyclophilins using both CsA binding and isomerase activity assays reveals that H. microstoma possesses two cyclophilin isoforms (M(r) 17,700 and 21,400) with isomerase activity that is inhibited by CsA. using identical assays, we have been unable to demonstrate CsA-binding proteins or CsA-sensitive isomerase activity in H. diminuta. These data suggest that the anthelmintic action of CsA relates in some way to the presence and function of parasite cyclophilins.

Amino Acid Isomerases↗

Genotyping of human cystic echinococcosis in Xinjiang, PR China.

The Xinjiang Uygur Autonomous Region, multi-ethnic province in northwestern China, is one of the most important foci of human cystic echinococcosis (CE) in the world. Two Echinococcus granulosus genotypes (G1 and G6) are known to infect the intermediate hosts in this area but, to date, the source of the human infection remains unclear. The current study aimed to genetically analyse 67 hydatid cysts removed from 47 CE patients for which epidemiological, clinical and serological data were also recorded. Mitochondrial cox 1 gene sequencing suggested that the E. granulosus G1 genotype is the major source of infection (45/47 CE patients). Nevertheless, for the first time in China, 2 patients were found with hydatid cysts of the G6 genotype. In addition, 45 E. granulosus gravid tapeworms, isolated from 13 dogs, were genotyped. The majority of adult worms (42/45) exhibited the G1 genotype, whereas 3 adult tapeworms with the G6 genotype were found in one dog, that also harboured E. granulosus tapeworms of the G1 genotype. This sympatric occurrence of G1 and G6 genotypes of E. granulosus, not only in the same area but also in the same definitive host, raises the interesting question of putative genetic recombination between these E. granulosus genotypes.

Adolescent↗

Recent developments in research into the Cyathostominae and Anoplocephala perfoliata.

Intestinal helminths are an important cause of equine disease. Of these parasites, the Cyathostominae are the commonest group that infect horses. These nematodes consist of a complex tribe of 51 species, although individual horses tend to harbour 10 or so common species, in addition to a few rarer species. The Cyathostominae can be extremely pathogenic, and high levels of infection result in clinical symptoms ranging from chronic weight loss to colic, diarrhoea and death. As part of their life cycle, immature cyathostomins penetrate the large intestinal wall, where they can enter a state of inhibited larval development. These larvae can exist in this state for months to years, after which they subsequently re-emerge. If larvae re-emerge in large numbers (i.e. several million), severe pathological consequences ensue. The inhibited larvae are also relatively refractory to several of the currently available anthelmintics, so that horses treated previously with anthelmintics can still carry life-threatening burdens of these parasitic stages. Little is known about the cyathostomin larvae during their mucosal phase, and current research efforts are focused on investigating the biology of these stages. Much of the research described here highlights this area of research and details studies aimed at investigating the host immune responses that the mucosal larvae invoke. As part of this research effort, molecular tools have been developed to facilitate the identification of larval and egg stages of cyathostomins. These molecular tools are now proving very useful in the investigation of the relative contributions that individual, common cyathostomin species make to the pathology and epidemiology of mixed helminth infections. At the more applied level, research is also in progress to develop an immunodiagnostic test that will allow numbers of mucosal larvae to be estimated. This test utilises antigen-specific IgG(T) serum antibody responses as markers of infection. As anthelmintic resistance will be the major constraint on the future control of the Cyathostominae, researchers are now actively investigating this area and studies aimed at elucidating the molecular mechanisms of drug resistance are described. Another parasite which has assumed a clinically important role in horses is the tapeworm, Anoplocephala perfoliata. This parasite is prevalent world-wide and has been shown to be a significant cause of equine colic. Because previous methods of estimating the infection intensity of tapeworm were inaccurate, recent research has been directed at developing an immunodiagnostic ELISA for these cestodes. Specific IgG(T) responses to antigens secreted by adult tapeworms have been shown to provide a reasonable indication of infection intensity. An ELISA based on these responses is now commercially available. The steps involved in the development of this ELISA are described here. In addition to these recent advances in research, this review also outlines the principle areas for future research into these important equine parasites.

Animals↗

Hymenolepis microstoma: direct life cycle in immunodeficient mice.

The mouse bile duct tapeworm Hymenolepis microstoma requires beetles as the obligatory intermediate host. However, when congenitally athymic NMRI-nu mice were infected with the mature tapeworm and allowed to eat their own faeces with tapeworm eggs, the oncospheres penetrated the intestinal tissue and developed to cysticercoids. After excysting, growth to adult worms occurs in the lumen of the small intestine and bile duct. Furthermore, the same happened when NMRI-nu mice, non-obese diabetic severe combined immunodeficiency (NOD/Shi-scid) mice and NOD/Shi-scid, IL-2 Rgamma(null) (NOG) mice were orally inoculated with shell-free eggs of this parasite. Differences between the cysticercoids of H. microstoma and H. nana developed in the mouse intestinal tissues were: (i) the time course for the development of fully matured cysticercoids of H. microstoma in mice was about 11 days but only 4 days for H. nana; and (ii) cysticercoids of H. microstoma developed in mice had a tail while those of H. nana had none.

Animals↗

Taeniasis unresponsive to a single dose of niclosamide: case report of persistent infection with Taenia saginata and a review of therapy.

A patient infected in Ethiopia with the beef tapeworm Taenia saginata was unsuccessfully treated with 2 g of niclosamide on four separate occasions over four months. clinical and parasitologic cures were effected by a regimen of 2 g of niclosamide given on three consecutive days. Niclosamide is the drug of choice for treatment of T. saginata and Taenia solium (pork tapeworm) infection; cure rates are approximately 90%. It is not absorbable and thus is nontoxic. Alternative treatments of taeniasis vary in the degree of safety; tin compounds and Aspidium extracts often are toxic. Paromomycin sulfate, which is also nonabsorbable, is effective, although adverse gastrointestinal effects are common with multiple-dose regimens. A relatively safe therapy is with mebendazole, a broad-spectrum anthelmintic agent, although efficacy has varied in different trials. A number of benzimidazoles, as well as nitazoxamide, have been effective against tapeworms. Praziquantel is the best alternative to niclosamide in terms of cost, efficacy, availability, and safety.

Adult↗

Analysis of antibody responses to Hymenolepis nana infection in mice by the enzyme-linked immunosorbent assay and immunoprecipitation.

Serum antibody responses in two strains of mice infected with embryonated eggs of Hymenolepis nana were analysed by the enzyme-linked immunosorbent assay (ELISA) and immunoprecipitation (IP) using sodium deoxycholate (DOC)-solubilized antigens prepared from embryonated eggs (eggs), mouse-derived cysticercoids (cysts) and adult tapeworms with immature segments only (adults). Highly susceptible dd mice, which harbour mature tapeworms for a long period (greater than 70 days), produced high levels of antibodies to all three different stages of H. nana. BALB/c mice, almost all of which expel adult tapeworms by 30 days after infection, produced high levels of antibody against egg antigens only. The high antibody titres to cyst and adult antigens in dd mice did not lead to expulsion of the worms. However, worms are rejected early in BALB/c mice when there is little or no detectable serum antibody. The antibody responses to eggs seen in BALB/c mice which had long since shed their adult worms were probably due to ingestion of eggs from faeces of other infected mice. Antibodies to eggs were not detected in BALB/c mice which were initially inoculated with eggs (day 0) and then treated with praziquantel on day 6 after the tissue phase of infection only. The different antibody responses to egg antigens and the other two antigens (cyst and adult) in BALB/c mice suggest a difference in antigen specificity between eggs and both cysts and adults. A major antigen component with Mr 32,000 appears to be specific to the egg (or oncosphere) stage of H. nana. Antibody to this major component of eggs was absorbed only with intact eggs, but not with intact cysts nor adults with immature segments only, so that the antigen appears to be on the surface of the oncosphere.

Animals↗

Costly resistance to parasitism: evidence from simultaneous quantitative trait loci mapping for resistance and fitness in Tribolium castaneum.

Information on the molecular basis of resistance and the evolution of resistance is crucial to an understanding of the appearance, spread, and distribution of resistance genes and of the mechanisms of host adaptation in natural populations. One potential important genetic constraint for the evolution of resistance is fitness cost associated with resistance. To determine whether host resistance to parasite infection is associated with fitness costs, we conducted simultaneous quantitative trait loci (QTL) mapping of resistance to parasite infection and fitness traits using the red flour beetle (Tribolium castaneum) and the tapeworm parasite (Hymenolepis diminuta) system in two independent segregating populations. A genome-wide QTL scan using amplified fragment length polymorphism (AFLP) markers revealed three QTL for beetle resistance to tapeworm infection. These three QTL account for 44-58% variance in beetle infection intensity. We identified five QTL for fecundity and five QTL for egg-to-adult viability, which accounted for 36-57% and 36-49%, respectively, of the phenotypic variance in fecundity and egg-to-adult viability. The three QTL conferring resistance were colocalized with the QTL affecting beetle fitness. The genome regions that contain the QTL for parasite resistance explained the majority of the variance in fecundity and egg-to-adult viability in the mapping populations. Colocalization of QTL conferring resistance to parasite infection and beetle fitness may result from the pleiotropic effects of the resistance genes on host fitness or from tight linkages between resistance genes and adverse deleterious mutations. Therefore, our results provide evidence that the genome regions conferring resistance to tapeworm infection are partially responsible for fitness costs in the resistant beetle populations.

Animals↗

First evidence of the presence of microtriches in the Gyrocotylidea.

The Gyrocotylidea, a small and enigmatic group of intestinal parasites of chimaeras, has been considered to be related either to the Monogenea, or, more frequently, to the most primitive monozoic tapeworms (Cestoda), i.e., the Amphilinidea and Caryophyllidea. The present study, based on transmission electron microscopical observations of a species of Gyrocotyle from the rabbit fish, Chimaera monstrosa, in the North Atlantic, demonstrates for the first time the presence of microtriches as surface structures of gyrocotylideans. Because microtriches are considered to be an autapomorphy of tapeworms (Cestoda), in which they differ from other Neodermata (Monogenea and Trematoda), the present data represent another source of evidence in support of a close relationship between the gyrocotylideans and the tapeworms sensu stricto (Eucestoda). Simple morphology, small size, and shape uniformity of the microtriches of Gyrocotyle sp. may indicate they represent an original (plesiomorphic) form that then evolved in more derived cestode groups into a variety of types present mainly on the scolex. The microtriches of Gyrocotyle sp. resemble those found in caryophyllidean, spathebothriidean, pseudophyllidean, and trypanorhynch cestodes, which are considered to represent the most basal groups of the Eucestoda.

Animals↗

Factors associated with development of ileal impaction in horses with surgical colic: 78 cases (1986-2000).

Deal impaction is prevalent in the south-eastern USA, where feeding of Coastal Bermuda hay has been implicated as a risk factor. Alternatively, infection with the tapeworm Anoplocephala perfoliata has been identified as a risk factor for ileal impaction in the UK. We hypothesised that feeding Coastal Bermuda hay and failure to administer routinely an anthelmintic with efficacy against tapeworms would place horses at risk of developing ileal impaction in the USA. Seventy-eight horses, with surgically confirmed ileal impaction and 100 horses admitted for colic that did not have an ileal impaction, were selected retrospectively for logistic regression analysis. Using odds ratios (OR) as an index of risk, feeding Coastal Bermuda hay (OR = 2.9) and failure to administer a pyrantel salt within 3 months of admission (OR = 3.1) placed horses at risk of development of ileal impaction. This study confirms the belief that feeding Coastal Bermuda hay places horses at risk of ileal impaction, although the quality of the hay may also play a role. Periodic administration of anthelmintics with efficacy against tapeworms should be considered to reduce risk of ileal impaction.

Animal Feed↗

Morphologic descriptions of Taenia asiatica sp. n.

Among taeniid tapeworms infecting humans through pork or beef, Taenia solium Linnaeus 1758 and Taenia saginata Goeze 1782 have already been known. Based on the morphologic characteristics of adult and metacestodes of Asian Taenia saginata, the third kind of human taeniid tapeworm known to distribute in Asian countries, a new species name of Taenia asiatica is proposed. In addition to the known biology in their intermediate hosts, T. asiatica was different morphologically from Taenia saginata Goeze 1782 in having the unarmed rostellum on the scolex of adult, the large number of 'uterine twigs' and the existence of 'posterior protuberance'. These structures in the gravid proglottids were used as taxonomic keys in taeniid tapeworms for the first time. T. asiatica metacestode (Cysticercus viscerotropica) was different morphologically from T. saginata metacestode (Cysticercus bovis) in having wartlike formations on the external surface of the bladder wall.

Adult↗

Prevalence and risk factors for Taenia solium taeniasis and cysticercosis in humans and pigs in a village in Morelos, Mexico.

In a Mexican village in which Taenia solium infection was known to be endemic, we selected a cluster sample of 368 households (21% of the total) for demographic, environmental, and diagnostic surveys, and medical histories for taeniasis and cysticercosis. Coproparasitologic studies of 1,531 participants revealed infection by Taenia sp. in four (0.3%) individuals; however, 5.8% of the respondents reported a history of having passed tapeworm proglottids in feces. Of 1,552 human serum specimens, 10.8% tested positive in the cysticercosis immunoblot assay. Seropositivity increased with age and reached a maximum in subjects ages 46-55 years. Risk factors associated with seropositivity included a history of passing tapeworm proglottids, frequent consumption of pork, and poor personal and household hygiene (P less than 0.05). A history of seizures was also significantly associated with seropositivity (P less than 0.05); approximately one-third of persons with such histories were seropositive. Of 571 pigs examined by tongue inspection, 23 (4.0%) had cysticerci; infection rates increased with the age of pigs, and were higher in pigs that habitually ran loose or were fed human feces (P less than 0.05). Goodness of fit analysis confirmed that seropositive persons (but not infected pigs) were significantly clustered within households, particularly, in households in which a member reported a history of having passed tapeworm proglottids. The results of this study have identified community behavioral and environmental practices that must be modified to prevent continued transmission of cysticercosis and taeniasis.

Adolescent↗

Pancreatic duct hyperplasia in a raccoon (Procyon lotor) caused by Atriotaenia procyonis.

A raccoon (Procyon lotor) live-trapped in Philadelphia, Pennsylvania was heavily infected with the tapeworm Atriotaenia procyonis. Histopathologic examination of the duodenum revealed this tapeworm attached to the mucosa; the mucosa was mildly hyperplastic without a significant inflammatory infiltrate. This tapeworm was seen also within the proximal pancreatic duct and histopathologic lesions associated with its presence consisted of marked hyperplasia of mucosal cells, causing the formation of an extensive papillary projection into the duct lumen. Inflammatory cellular reaction was minimal, consisting primarily of lymphocytes, and was seen in the mucosa and submucosa of the duct. The presence of A. procyonis in the pancreas of this raccoon is considered to be an aberrant location for this parasite and has not been reported previously.

Animals↗

Life cycles of species of Proteocephalus, parasites of fishes in the Palearctic region: a review.

The life cycles of species of Proteocephalus Weinland, 1858 (Cestoda: Proteocephalidea) parasitizing fishes in the Palearctic Region are reviewed on the basis of literary data and personal experimental observations, with special attention being paid to the development within the intermediate and definitive hosts. Planktonic crustaceans, diaptomid or cyclopid copepods (Copepoda), serve as the only intermediate hosts of all Proteocephalus species considered. A metacestode, or procercoid, develops in the body cavity of these planktonic crustaceans and the definitive host, a fish, becomes infected directly after consuming them. No previous reports of the parenteral location of metacestodes within the second intermediate host as it is in the Nearctic species P. ambloplitis have been recorded. Thus, the life cycles of Proteocephalus tapeworms resemble in their general patterns those of some pseudophyllidean cestodes such as Eubothrium or Bothriocephalus, differing from the latter in the presence of a floating eggs instead of possessing an operculate egg from which a ciliated, freely swimming larva, a coracidium, is liberated. The scolex of Proteocephalus is already formed at the stage of the procercoid within the copepod intermediate host; in this feature, proteocephalideans resemble caryophyllidean rather than pseudophyllidean cestodes. The morphology of procercoids of individual species is described with respect to the possibility of their differentiation and data on the spectrum of intermediate hosts are summarized. Procercoids of most taxa have a cercomer, which does not contain embryonic hooks in contrast to most pseudophyllidean cestodes. The role of invertebrates (alder-fly larvae-Megaloptera) and small prey fishes feeding upon plankton in the transmission of Proteocephalus tapeworms still remains unclear but these hosts are likely to occur in the life cycle. Data on the establishment of procercoids in definitive hosts, morphogenesis of tapeworms within fish hosts, and the length of the prepatent period are still scarce and new observations are needed. Whereas extensive information exists on the development of P. longicollis (syns. P. exiguus and P. neglectus), almost no data are available on the ontogeny of other taxa, in particular those occurring in brackish waters (P. gobiorum, P. tetrastomus). The morphology of P. cernuae and P. osculatus procercoids from experimentally infected intermediate hosts is described for the first time.

Animals↗