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Epidemiological studies on equine cestodes in central Spain: infection pattern and population dynamics.

An epidemiological study on equine cestodosis was carried out in central Spain. A total of 372 digestive tracts from equids slaughtered in abattoirs located in central Spain were studied from November 2001 to May 2004. Anoplocephala perfoliata was detected in 24% of the animals and Anoplocephala magna in 18%. Individual tapeworm burden was from 1 to 491 tapeworms for A. perfoliata and from 1 to 64 tapeworms for A. magna. Low tapeworm burdens (less than 30 cestodes) predominated significantly (p<0.01) in all seasons for both species. Seasonal prevalence of infection by A. perfoliata was significantly higher (p<0.01) in autumn (37.5%) and winter (32.3%) than in spring (9.2%) and summer (10.8%). Immature tapeworms were detected throughout summer (11%), autumn (23.4%) and winter (26.6%), signalling a summer to winter risk period for tapeworm infection in our conditions. Mature non-gravid tapeworms were collected in all seasons, with a decreasing pattern from summer (89%) to spring (6.7%). Conversely, gravid tapeworms showed an increasing pattern, from 0 in summer to a maximum (93.3%) in spring. Since prevalence of infection was significantly higher in winter than in spring, winter seems to be the season when more eggs would be available to be eaten by mites. A. perfoliata infection was detected in three different periods throughout the 3 year study: autumn 2001-winter 2002, summer 2002-spring 2003 and autumn 2003-spring 2004. This epidemiological pattern seems to describe the dependence of A. perfoliata to humidity in warm dry climate. In our conditions, A. perfoliata appears to follow a pattern of having only one generation per year, with a marked dependence on humidity. According to the results, autumn rainfall would influence the length, and late spring rainfall the appearance, of each annual generation. A. magna showed a different pattern. Infection was detected throughout the whole study period. Seasonal prevalence was higher in autumn (25.5%) than in winter (14.9%), spring (12.1%) and summer (10.5%), but the differences were not statistically significant. Non-gravid A. magna tapeworms could be detected almost throughout the year but percentages were significantly higher (p<0.01) in autumn (50.2%), indicating recent ingestion, than in the other seasons (30% in summer, 12.8% in winter and 0% in spring). However, data from spring were not enough to discard the season as a risk period for A. magna infection.

Animals↗

Basic and applied immunology in cestode infections: from Hymenolepis to Taenia and Echinococcus.

In larval cestode infections, it is well established that the intermediate mammalian host infected with egg-derived metacestodes in the tissue becomes completely immune to reinfection with eggs, whereas autoinfection has been conceived to occur in Hymenolepis nana/mouse (and human) and Taenia solium/human systems when these hosts are initially infected with metacestode-derived adult tapeworms in the lumen. In this review paper, the first topic is immunobiology of H. nana/mouse system on the reinfection immunity in order to get critical information as to how the initially ingested parasite (eggs or metacestodes) can develop into adult worms and how autoinfection does or does not occur in immunocompetent mice, since H. nana can complete its whole life cycle in the mouse intestinal tissue and lumen. When mice are infected with eggs (= oncospheres) of H. nana, they become immune to challenge infections with eggs within a few days (early response) and with cysticercoids within two weeks (late response). The initially established adult worms are expelled later (worm expulsion response). When mice are infected with cysticercoids, either derived from beetles or mice, they become immune to challenge infection with cysticercoids but not with eggs. Therefore, autoinfection occurs in the intestinal tissue for the establishment of cysticercoids in the tissue but never occurs in the intestinal lumen for the establishment of adult worms in immunocompetent mice. The second topic is vaccination trial against challenge infection with eggs of Asian Taenia in pigs. Pigs vaccinated with frozen oncospheres of Asian Taenia from Taiwan or Korea or T. saginata showed very strong resistance, whereas pigs vaccinated with those of T. solium showed partial resistance only. It is suggested that Asian Taenia is much closer to T. saginata than T. solium from the immunobiological viewpoint. The third topic is immunodiagnosis of echinococcosis and cysticercosis. Immunoblot analysis has revealed that Em18 (18 kDa component of crude antigens of Echinococcus multilocularis protoscolex) and glycoproteins of T. solium cysticerci are highly specific or unique to alveolar echinococcosis and cysticercosis, respectively. The fourth topic is discussion on miscellaneous prospects including laboratory animal models for echinococcosis and cysticercosis.

Animals↗

To avoid or eliminate: cestode infections in copepods.

The outcome of a parasite infection is the result of the interaction between the host and the parasite. In the system we studied, there are 3 critical stages for the outcome of infection of the (intermediate) host, the copepod Macrocyclops albidus, with the cestode Schistocephalus solidus. During the establishment phase of the parasite, the host may firstly avoid ingesting the parasite and, secondly, may prevent the parasite from entering the body cavity and, thirdly, during the growth phase of the parasite, the host's immune system may eliminate the parasite from the body cavity. We were able to study the growth phase separately from the establishment phase. The establishment phase was influenced by characteristics of the host as well as characteristics of the parasites. Small copepods and males performed poorly; they were more often infected and had a lower survival. Parasites from different sib-groups differed in infectivity. During the growth phase some disappearance of parasites was observed. However, this could not be related to any of the studied characteristics of the host, and the sib-groups of parasites did not seem to differ in their likelihood to disappear. Instead, we suggest that disappearance of parasites, once they have entered the body cavity, may be due to intrinsic mortality of the parasites, independent of the host or the sib-group that the parasites belong to. This indicates that the crucial interactions between host and parasite determining the outcome of infection takes place in the short time-period between ingestion and penetration of the gut-wall.

Animals↗

The consequences of self-fertilization and outcrossing of the cestode Schistocephalus solidus in its second intermediate host.

Many hermaphroditic parasites reproduce by both cross-fertilization and self-fertilization. To understand the maintenance of such mixed mating systems it is necessary to compare the fitness consequences of the two reproductive modes. This has, however, almost never been done in the context of host-parasite coevolution. Here we show the consequences of outcrossing and selfing in an advanced life-stage of the cestode Schistocephalus solidus, i.e. in its second intermediate host, the three-spined stickleback (Gasterosteus aculeatus). Each juvenile stickleback was simultaneously exposed to 2 experimentally infected copepods, one harbouring outcrossed the other selfed parasites. At 60 days p.i. parasites were removed from the fish's body cavity and, with microsatellite markers, assigned to either outcrossed or selfed origin. Prevalence was not significantly higher in outcrossed parasites. However, those fish that were infected contained significantly more out-crossed than selfed parasites. Thus the probability of a selfed parasite to progress in the life-cycle is reduced in the second intermediate host. Furthermore, we found that even the multiply infected fish increased in weight during the experiment. Nevertheless, total worm weight in multiply infected fish was significantly lower than in singly infected ones, which thus might be a parasite life-history strategy.

Animals↗

Tritiated thymidine ([3H]-TdR) and immunocytochemical tracing of cellular fate within the asexually dividing cestode Mesocestoides vogae (syn. M. corti).

This report documents the presence of an active thymidine kinase (TK) system within Mesocestoides vogae tetrathyridia as quantified by tritiated thymidine ([3H]-TdR) incorporation using liquid scintillation counting. A 100-fold increase in [3H]-TdR incorporation was observed at 37 degrees C when compared with its incorporation at 0 degrees C. Thymidine's competitive analogue, BrdU, competed for sites within newly replicated DNA. Immunohistochemical trials performed here using antibodies against BrdU identified cells that have entered and passed through S-phase. Positively stained nuclei were most numerous at the anterior tip of tetrathyridia especially within the ganglia, lesser numbers of these cells occurred along the growing commissure and amongst surface tegumental cytons suggesting that stem cells do not exist in one region but are found throughout the entire body. As M. vogae has no internal organ systems the major sites for cell proliferation are those exhibiting maximal cell recruitment and undergoing tissue repair. These results show that it is possible to monitor changes in the cell recruitment pattern within this cestode. Thus use of BrdU and immunohistochemistry demonstrates how spatial arrangement and cellular reorganization can be successfully traced within M. vogae.

Animals↗

Studies on immune responses to larval cestodes in mice: a simple mechanism of non-specific immunosuppression in Mesocestoides corti-infected mice.

After intraperitoneal (i.p.) injection of sheep erythrocytes (SRBC) or dinitrophenylated Ficoll (DNP-Ficoll), mice infected with the larval cestode, Mesocestoides corti, contain at least 20x fewer antibody-secreting cells (PFC) in their spleens (or spleens plus lymph nodes) than uninfected mice. By contrast, intravenous injection of antigen leads to normal PFC responses. Results of studies on the fate of labelled syngeneic erythrocytes and foreign proteins suggest that i.p. injected materials are retained in the inflamed peritoneal cavity. Sequestration of antigen, and its subsequent local destruction, presumably accounts for the markedly suppressed systemic immune responses induced by i.p. injected antigens in M. corti-infected mice.

Animals↗

Hybridoma antibody immunoassays for the detection of parasitic infection: development of a model system using a larval cestode infection in mice.

A prototype immunodiagnostic assay has been developed using chronic infection with the larval cestode, Mesocestoides corti, as a model system in mice. The assay is highly sensitive, it appears to be absolutely specific for M. corti infection, and is based on the inhibition of binding (by sera from infected mice) of a radiolabelled anti-M. corti hybridoma antibody to a crude M. corti antigen extract. The hybridoma antibody binds to living M. corti larvae and is an IgG1 protein. In large scale experiments no false positives were detected and the only M. corti-infected mice not detected by the assay were hypothymic nude (nu/nu) mice. Only limited success has been achieved in attempts to convert the assay to one not requiring parasite antigen and based on the inhibition of binding of radiolabelled anti-parasite hybridoma antibody and a large pool of anti-idiotype antiserum. Monoclonal antibodies derived from anti-parasite antibody-secreting hybridoma cell lines will be of particular use in the development of new, highly specific, immunodiagnostic reagents for the detection of parasite infection, exposure and disease.

Animals↗

Neuroparasitic infections: cestodes, trematodes, and protozoans.

Parasitic infection of the nervous system can produce a variety of symptoms and signs. Because symptoms of infection are often mild or nonspecific, diagnosis can be difficult. Familiarity with basic epidemiological characteristics and distinguishing radiographic findings can increase the likelihood of detection and proper treatment of parasitic infection of the nervous system. This article discusses the clinical presentation, diagnosis, and treatment for some of the more common infections of the nervous system caused by cestodes, trematodes and protozoans: Echinococcus spp., Spirometra spp. (sparganosis), Paragonimus spp., Schistosoma spp., Trypanosoma spp., Naegleria fowlerii, Acanthamoeba histolytica, and Balamuthia mandrillaris.

Animals↗

Incidental coenurosis: larval cestode presenting as an axillary mass.

This article describes the occurrence of a coenurus, the encysted larval stage of a cestode of the genus Multiceps, within the axilla of a 40-year-old Nigerian woman. The patient had a history of intraductal carcinoma of the breast two years prior to the finding of an ipsilateral axillary mass. Pathologic evaluation of the biopsy specimen yielded the diagnosis of coenurosis. Only four other cases of human Multiceps infestation have been reported in the United States with fewer than 100 validated cases worldwide. Surgical removal of the larval parasite is the only known effective therapy.

Adult↗

The parasitic trematode Fasciola hepatica exhibits mammalian-type glycolipids as well as Gal(beta1-6)Gal-terminating glycolipids that account for cestode serological cross-reactivity.

Neutral glycosphingolipids from sheep-derived Fasciola hepatica liver flukes were isolated and characterized both structurally and serologically. After HPLC fractionation, glycolipids were analyzed by linkage analysis, enzymatic cleavage, and MALDI-TOF as well as electrospray ionization mass spectrometry. Obtained results revealed the presence of two types of neutral glycolipids. The first group represented mammalian-type species comprising globo- and isoglobotriaosylceramides (Gal(alpha1-4)Gal(beta1-4)Glc(1-1)ceramide and Gal(alpha1-3)Gal(beta1-4)Glc(1-1)ceramide, respectively) as well as Forssman antigen (GalNAc(alpha1-3)GalNAc(beta1-3/4)Gal(alpha1-4/3)Gal(beta1-4)Glc(1-1)ceramide). Applying Helix pomatia agglutinin, recognizing terminal alpha-linked GalNAc, to cryosections of adult flukes, the latter glycolipid could be localized to the F. hepatica gut. As Forssman antigen from the parasite and sheep host led to identical MALDI-TOF MS profiles, this glycolipid might be acquired from the definitive host. As a second group, highly antigenic glycolipids were structurally characterized as Gal(beta1-6)Gal(beta1-4)Glc(1-1)ceramide, Gal(beta1-6)Gal(alpha1-3/4)Gal(beta1-4)Glc(1-1)ceramide and Gal(beta1-6)Gal(beta1-6)Gal(alpha1-3/4)Gal(beta1-4)Glc(1-1)ceramide, the latter two structures of which exhibited both isoglobo- or globo-series core structures. Terminal Gal(beta1-6)Gal1-motifs have previously been shown to represent antigenic epitopes of neogala-series glycosphingolipids from tape worms. Using human Echinococcus granulosus infection sera, Gal(beta1-6)Gal-terminating glycolipids could be allocated to the gut in adult liver fluke cryosections. Corresponding neogala-reactive antibodies in F. hepatica infection serum were detected by their binding to E. granulosus and Taenia crassiceps neogala-glycosphingolipids. These antibodies might contribute to the known serological cross-reactivity between F. hepatica and parasitic cestode infections.

Animals↗

Mode of attachment and lesions associated with trypanorhynch cestodes in the gastrointestinal tracts of two species of sharks collected from coastal waters of Borneo.

Lesions associated with two species of tapeworms within the digestive tract of wild-caught specimens of the bull shark, Carcharhinus leucas, and the sicklefin weasel shark, Hemigaleus microstoma, from Malaysian Borneo are described. Portions of the glandular stomach and pyloric gut with parasites were removed and fixed in 10% formalin buffered in sea water. Whole mounts, histological sections of tissues with and without worms in situ, and scanning electron microscopy images of detached worms were examined. Both species of cestodes belonged to the trypanorhynch family Tentaculariidae. Heteronybelinia estigmena was found in large numbers parasitizing the pyloric gut of C. leucas; an unidentified tentaculariid was found in relatively small numbers in both the glandular stomach and pyloric gut of H. microstoma. Both species burrowed their scoleces deeply in the mucosa and attached via hooked tentacles and unciniform microtriches of the scolex. The lesions induced by the parasites were marked in both sharks and ranged from acute necrotizing to chronic granulomatous gastroenteritis. Regenerative hyperplasia and intestinal metaplasia of gastric epithelium were also present. The severity and character of pathology was causally linked to the intensity of infection, the attachment mode of the parasites, and to the anatomophysiological relationships within the gut of the host shark.

Animals↗

Cestodes.

A review has been made of the advances in knowledge on the treatment of tapeworm infections of man and animals from the time of the introduction to dichlorophen in 1956. This opened up the era of out-patient treatment for human tapeworm infections. The drugs studied have been compared, where possible, on the basis of estimates of the single dose ED90 or the number of treatments required to reach that efficacy at safe dose levels. During the period under review, niclosamide, introduced in 1960 has been regarded as the drug of choice for the treatment of human tapeworm infections with paromomycin as a possible alternative. The bunamidine salts, introduced in 1966 permitted the treatment of Echinococcus spp in dogs. Several treatments were required to achieve acceptable efficacy. In 1975 praziquantel was introduced and based on the ED90, a single dose at no more than 10mg/kg removed all tapeworms responsible for the cestode zoonoses and for which data are available with the exception of H. nana and D. latum; these require a higher dose rate. In the mid 1970s, several benzimidazoles and praziquantel were shown to have activity against metacestodes. This has opened up a new field of research promising a practical outcome. No progress has been made during the period under review in finding effective ovicides.

Animals↗

Long-term survey on intestinal nematode and cestode infections in stray puppies Ibaraki Prefecture.

We conducted the survey on intestinal nematode, mainly Toxocara canis, and cestode infections in stray puppies aged less than 5 months in Ibaraki Prefecture for 8 years from 1988 to 1995. The rate of T. canis infection stayed almost unchanged for the 8 years and ranged from 72.2 to 85.4%. Of the total 916 dogs, 732 (79.9%) harboured with T. canis. The average rates of other helminthic infections showed 24.6 (Diphylidium caninum), 2.5 (Trichuris vulpis), 1.2 (Ancylostoma caninum) and 0.7% (Spirometra erinaceieuropaei) respectively.

Animals↗

Fimbriaria fasciolaris and Cloacotaenia megalops (Eucestoda, Hymenolepididae), cestodes from Brazilian waterfowl.

Two cestode species, Fimbriaria fasciolaris (Pallas, 1781) Frölich, 1802 Cloacotaenia megalops (Nitzsch in Creplin, 1829) Wolffhügel, 1938 collected from Anas bahamensis Linné, 1758 and Amazonetta brasiliensis (Gmelin, 1758) in lagoons of the Maricá District, State of Rio de Janeiro, Brazil, are described. This is the first record of F. fasciolaris parasitizing A. bahamensis. The prevalence, intensity of infection, and mean intensity of infection for both species are given. Overdispersion distribution is reported for F. fasciolaris with 535 specimens collected in a single A. bahamensis. A key for the genera in the Fimbriariinae is presented. Anatomical features of F. fasciolaris and C. megalops are discussed.

Animals↗

The "crowding effect" in the cestode Schistocephalus solidus: density-dependent effects on plerocercoid size and infectivity.

The occurrence of the crowding effect was demonstrated in plerocercoids of the cestode Schistocephalus solidus infecting threespine stickleback Gasterosteus aculeatus from Walby and Scout lakes, Alaska. Contrary to an earlier report, relatively large numbers of parasites (>3-4 plerocercoids) were observed to grow large enough in an intermediate host fish to become competent to infect and to mature in the definitive host under any of 3 assumed threshold values and 1 scenario of graded sizes for parasite competency. In Walby Lake, intensity and host body mass were significant predictors of mean plerocercoid mass per host, whereas intensity, host body mass, and combined parasite index were significant predictors in Scout Lake. Slopes of equations expressing the relationship between mean parasite mass and intensity for both lakes were less than 1, implying that processes other than or in combination with simple resource limitation might be producing the observed crowding effect. The causal mechanism for the crowding effect could include exploitative competition, interference competition, and host immune response. There were significant differences in infection between the two lakes, including different distributions of parasite intensities among hosts and different expressions of the crowding effect; however, an explanation of the differences awaits further investigation.

Alaska↗

Reduction of egg size in natural populations of threespine stickleback infected with a cestode macroparasite.

Manifestations of infectious disease may represent host adaptations to avoid or reduce the effects of infection on host fitness, parasite manipulations that benefit the pathogen's fitness, or nonadaptive side effects of parasitism. Threespine stickleback fish (Gasterosteus aculeatus) from Alaska and the cestode macroparasite Schistocephalus solidus provide an excellent system for study of the effects of parasitism on host egg size because females in populations there are capable of producing clutches of eggs in the face of substantial infection, contrary to the inhibition of reproduction that has been observed in other stickleback populations or other species of fish. A side effect resulting in reduction of mean ovummass among infected females was predicted based on the egg production process in female stickleback, the considerable energy and resource demands of S. solidus, and the chronic and progressive nature of the effects the macroparasite should have on the host fish. In each of 9 populations of G. aculeatus representing replicate natural experiments in lakes scattered across the Matanuska-Susitna Valley and the Kenai Peninsula of south-central Alaska and among all populations combined, the mean ovum mass of infected female fish is significantly reduced in comparison with that of uninfected females taken from the same population at the same time. Reduction in mean female egg mass ranged from 8 to 32% across all populations. To examine whether reduction in mean female ovum mass was a nonadaptive side effect or an adaptation, relatively large data sets from 2 of the populations were used. Mean ovum mass of infected females was predicted to decrease directly in relation to parasite index (PI) if the diminution in mean egg mass were the result of a nonadaptive side effect resulting from host nutrient loss. Alternatively, the absence of a relationship between PI and reduction in ovum mass is predicted if decreases in mean female ovum mass result from host or parasite adaptation (or both) because lightly infected hosts should show a response similar to that of heavily infected ones. In each of the 2 populations, there is a significant, negative relationship between mean female ovum mass and PI, demonstrating a correlation between the decrease in ovum mass and the level of infection. Thus, the results are consistent with the hypothesis that the reductions in mean female egg mass represent side effects of parasitism involving nutrient theft. Moreover, the proportional decline in egg mass with increasing PI apparently differed between the 2 populations, and there was no significant relationship between mean percent decrease in mean female ovum mass and mean PI across populations. These observations suggest that unknown ecological and evolutionary factors influence the degree of reduction in mean ovum mass in a population-specific manner.

Alaska↗

The clearance of hidden cestode infection triggered by an independent activation of host defense in a teleost fish.

Parasites often elude effective recognition or attack (or both) by the host immune system, for example, though a tegument that possesses nonimmunogenic features. However, a general activation of host defense due to independent stimuli may increase immune activity to a level where such disguises are no longer effective, resulting in the clearance of an infection. We experimentally infected three-spined sticklebacks (Gasterosteus aculeatus) with the cestode Schistocephalus solidus. To independently foster a general immune response a few days later, we cut the tips of spines in some fish and sham-treated other fish. Cutting spines significantly reduced the prevalence of the infection. The injury evoked a physiological reaction that helped to clear a hidden parasite infection.

Analysis of Variance↗

Spermiogenesis and spermatozoon ultrastructure of the cestode Mosgovoyia ctenoides (Cyclophyllidea: Anoplocephalidae), an intestinal parasite of Oryctolagus cuniculus (Lagomorpha: Leporidae).

Ultrastructural characters in spermiogenesis and spermatozoa are considered important tools to elucidate the phylogenetic relationships within the Platyhelminthes. In the Anoplocephalidae, ultrastructural data refer to the spermatozoon of 14 species, whereas data on spermiogenesis refer to only 7 species. The present study focused on the spermiogenesis and spermatozoon of the anoplocephalid cestode Mosgovoyia ctenoides, as revealed by transmission electron microscopy. Type IV spermiogenesis was detected, beginning with the formation of a differentiation zone containing 2 centrioles, with a centriolar adjunct and vestigial striated rootlets. Different forms of the latter character have been described in other anoplocephalids. This study supports spermiogenesis of type IV as the most frequent in the Anoplocephalidae and confirms the presence of a centriolar adjunct in yet another type IV spermiogenesis species. The spermatozoon of M. ctenoides possesses 1 axoneme of the 9+ '1' trepaxonematan type, 2 crestlike bodies, dense plates, and granules of electron-dense cytoplasmic material, nucleus, and twisted cortical microtubules. It was again confirmed that the presence of granular material and the absence of both a periaxonemal sheath and intracytoplasmic walls are constant characters in the spermatozoa of all the Anoplocephalinae.

Animals↗