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PubMed1975

Field studies on inhibition of Haemonchus contortus.

The epidemiology of Haemonchus contortus was studied in north east England in lambs reared under field conditions with infected ewes. The parasite population was monitored by pasture larval counts, faecal egg counts and lamb postmortems. From 25 June to 11 November, eight lambs were slaughtered each month, together with eight worm-free tracer lambs. Postmortem worm counts showed that on 25 June, small numbers of adult H. contortus were present but no inhibited larvae. However, subsequently the number of worms recovered rose rapidly to 1000-5000 and the percentage inhibition increased to 57% in July, 75% in August and virtually 100% in September, October and November. On each sampling occasion no significant differences in worm count or percentage inhibition were recorded between experimental and tracer lambs. These results suggest that neither autumn climatic effects nor host immunity were responsible for inhibition in this strain of H. contortus.

Animals↗
PubMed1976

Response of Haemonchus contortus cayugensis to a change in the ratio of smooth to linguiform.

The equilibrium of vulvar phenotypes in Haemonchus contortus cayugensis was changed by infecting lambs with either a selected strain of smooth or of linguiform worms. Subsequently the lambs grazed the same pasture and observations were made on changes in worm populations in the lambs, and on pasture it appeared that the seasonal change in the relative frequencies of A and B linguiform worms was related to density-dependent factors in the host. In the spring, when lambs first become infected, the worm population is small and the majority of linguiform worms are A-type. As the season progresses the density of the parasite population increases. Once the population becomes crowded selection begins to favour the B-type females. The effects of crowding on linguiform A females can be seen as a positive correlation between the percentage of linguiform A and their weight. Population density also affected the survival of A-type worms since there was a negative correlation of percentage A-type on total worm numbers. There was a similar negative correlation between percentage A-type on total worm numbers. There was a similar negative correlation between percentage A-type and total worm numbers in both the continuously infected and tracer lambs. Population density did not affect the proportion of smooth worms. However, smooth worms increased in the tracers from approximately 10% at the beginning of the experiment to 55% at its conclusion 18 weeks later. The latter percentage is almost within range of the local subspecies, which has 60% or more smooth worms. It appears therefore that the proportion of smooth to linguiform worms is a stable equilibrium maintained by natural selection.

Animals↗
PubMed1975

Observations on the development of Haemonchus contortus in young sheep given a single infection.

Worm free lambs 2-4 months old were given a single infection of 50,000 Haemonchus contortus larvae. The changes in packed red cell volume were compared with those of uninfected sheep and the infected sheep were divided into two groups according to the rate of change of packed cell volume. In group 1, packed cell volume fell sharply from the 10th day of infection and the sheep eventually died. In group 2, the fall was less marked and the packed cell volume returned to normal after about 40 days. The rates of increase of parasite body length and dry weight were compared for the two groups. Parasites in group 1 sheep grew more slowly than their group 2 contemporaries. Accurate planning of metabolic studies on H. contortus was made possible by using this information to predict the course of an infection and the size of the worms. Parasite development was also measured in sheep for which the packed cell volume was not recorded. Computer analysis showed that adult worm length increase did not follow a simple growth pattern starting from the last ecdysis, but consisted of a rapid elongation followed by a simple negative exponential increase. Dry weight increase also followed a negative exponential during the second phase. Measurements of dry weight as a percentage of wet weight indicated that the rapid elongation was possibly linked with cell enlargement by the uptake of water.

Abomasum↗
PubMed1977

The development of Haemonchus contortus, a nematode parasite of the ovine abomasum, in the laboratory rabbit.

The development and growth of Haemonchus contortus was studied in the stomach of the laboratory rabbit. Less than 1% of the ensheathed infective larvae given to rabbits became established, whereas after exsheathment in vitro under physiological conditions, 20-30% of the larvae administered developed to the 4th stage. Following doses of up to 10000 larvae, nearly all worms were lost from the 16th to the 21st day after infection, but after doses of 50000 larvae considerable numbers of 4th-stage larvae were recovered 25 days after infection. No adult worms were recovered and no eggs were found in the faeces. Infection was accompanied by inflammation, haemorrhage, disruption and flattening of the mucus secreting cells and by mononuclear infiltration. No cellular reactions in the immediate vicinity of the nematodes in the mucosa or changes in the pH of the stomach contents were observed.

Animals↗
PubMed1979

The inheritance of thiabendazole resistance in Haemonchus contortus.

Haemonchus contortus worm populations isolated from naturally infected sheep at the Pastoral Research Laboratory, Armidale, N.S.W., were found to contain approximately 20% of worms resistant to a 50 mg/kg dose of thiabendazole. Following 3 generations of selection with 50 mg/kg thiabendazole the number of worms removed by the anthelmintic was too small to detect differences between treated and control groups. After more than 15 generations of selection, matings between males from the selected strain and non-resistant females produced resistant males and females in equal numbers. Thus, thiabendazole resistance does not appear to be sex-linked. A dose--response assay on the F2 adults indicated that worms from female resistant x male non-resistant crosses were more resistant than F2 adults of the reciprocal cross. An in vitro technique that identified thiabendazole-resistant eggs by their ability to hatch in a solution containing thiabendazole and 0.1% NaCl solution was also used to study the inheritance of resistance. F1 eggs had similar LC50's to the resistant parents. F2 and back-cross eggs from an original mating of thiabendazole-resistant females x non-resistant males had a higher LC50 than F2 and back-cross eggs from the reciprocal mating, indicating a degree of matroclinous inheritance of resistance. However, the resistant parents had tolerances to thiabendazole exceeding those of F2. F3 eggs had a resistance distribution that ranged from that of the resistant to the non-resistant parent. No significant deviation from linearity was observed in any of the dose--response lines. These results indicate that thiabendazole resistance in H. contortus worms is inherited as an autosomal and semi-dominant trait.

Animals↗
PubMed1978

Genetic resistance to helminths. The influence of breed and haemoglobin type on the response of sheep to re-infection with Haemonchus contortus.

The influence of genetic factors on acquired resistance to Haemonchus contortus infections in sheep was investigated. Animals whose primary infections were terminated with an anthelmintic failed to develop any immunity against subsequent challenge as judged by worm numbers. Nevertheless, all were better able to retard the development and reduce the fecundity and haematophagic activities of their parasite populations than animals undergoing primary infections. High levels of resistance, as judged by all these parameters, were observed in most animals when the challenge larvae were superimposed on existing worm populations. The patterns of worm establishment and disease indicated that genetic factors operated in determining resistance, since fewer worms became established and less severe clinical and pathophysiological changes were observed in Scottish Blackface than in Finn Dorset sheep with the same haemoglobin type. Similar advantages were displayed by animals with haemoglobin AA and to a lesser extent those with haemoglobin AB over haemoglobin BB types. The importance of breed was further indicated by the occurrence of 'self-cure' in the majority of the Scottish Blackfaces but in only one Finn Dorset. There was no evidence that this reaction was associated with haemoglobin type.

Animals↗
PubMed1976

Biological controls in mixed trichostrongyle infections.

The parasite reaction phenotype of 80 lambs was determined with natural infection on pasture. These lambs were then assigned to outcome groups and randomized into treatment groups. Developing some simple but unusual analytical techniques it was possible to demonstrate that mixed natural infections of trichostrongyles with the chief biomass component Haemonchus contortus cayugensis protected against superinfections with 35000 infective larvae but protected against superinfections with 1000 infective larvae per day only within the limits characteristic of the enclosing barn ecosystem. The limits of the barn ecosystem were less constraining on more diathetic sheep than the natural pasture infection. Internal evidence from the experiment suggests that the interaction of sheep X pasture X infective trichostrongylid larvae constrains the parasite biomass in the host in more diathetic animals at significantly lower levels. Anaphylactoid 'self-cure' did not occur in this experiment but something like premunition certainly did.

Animals↗
PubMed1986

In vitro demonstration of in utero larval development in an oviparous parasitic nematode: Haemonchus contortus.

The life-cycle of Haemonchus contortus, a pathogenic stomach nematode of sheep, is typical of those of the other members of the superfamily Trichostrongyloides, all of which require a period of development outside the definitive host. Classically, gravid H. contortus, known as strictly oviparous, releases her eggs into the abomasal lumen. The eggs are passed out in the faeces in which they hatch into the 1st-stage larvae and then develop into the infective 3rd-stage larvae. We have developed a method to study the fate of eggs within gravid worms. Using this procedure, we have shown that H. contortus may also exhibit viviparity with apparently normal development from the egg to the 4th larval stage taking place within the gravid female maintained in vitro. On the basis of these observations we speculate that viviparity might occur in vivo with consequent autoinfections; if so, this might explain some puzzling clinical and epidemiological features of haemonchosis, as well as the incomplete efficacy of current control measures.

Abomasum↗
PubMed1987

Vaccination of young lambs by means of a protein fraction extracted from adult Haemonchus contortus.

Lambs aged 48-150 days which had been injected with extracts enriched in a functional antigen (contortin) obtained from adult Haemonchus contortus developed specific circulating antibodies and were less susceptible to haemonchosis when challenged 1 month later with a single dose of 20,000-25,000 infective 3rd-stage larvae. Sera from 18 out of a total of 19 lambs injected with the extract contained precipitating antibodies to 2-5 components of the extract. None of these lambs died. The one extract-injected lamb which did not develop antibodies and 9 of the 13 lambs used as controls died of haemonchosis. The average weight of worms recovered was 1.45 g from the immune lambs and 5.72 g from the non-immune lambs.

Animals↗
PubMed1982

The use of faecal egg counts for estimating worm burdens in sheep infected with Haemonchus contortus.

Clun forest sheep, worm-free from birth, were given a single dose of 20 000 infective larvae of Haemonchus contortus. The total number of eggs/day in the faeces was recorded for 21 infections and data on population size, sex ratio and individual worm development were collected from 76 sheep. The female to male ratio was 1.28 +/- 0.07 (S.E.). The relation between increase in worm size and uterine egg content was linear. The number of eggs present in the uteri was found to be an accurate measure of eggs passed. It was shown that the daily faecal egg output is related to total parasite weight and is not a measure of the number of individuals present.

Animals↗
PubMed1984

Studies on the pathophysiology of chronic ovine haemonchosis in Merino and Scottish blackface lambs.

An experiment was conducted to examine, under laboratory conditions, the pathophysiology of chronic ovine haemonchosis. In the present study, groups of Merino and Scottish Blackface lambs were maintained on a low plane of nutrition and infected with 50 Haemonchus contortus larvae/kg live weight. The parasitized lambs along with respective breed controls were examined over a 27-week period. In addition to the body weight, haematological and parasitological parameters, erythrokinetic, metabolic, digestibility and nitrogen balance studies were conducted in the early stages of the infection. The results demonstrated that a low level of infection with H. contortus in lambs on a poor plane of nutrition caused the development of a normochromic normocytic anaemia which was associated with a modest but significant increase in abomasal blood loss and slightly elevated erythropoiesis in both breeds relative to the controls. The level of anaemia, hypoproteinaemia and pathophysiological changes were more marked in the Merino lambs and they also had higher faecal egg counts than the Blackface lambs.

Anemia↗
PubMed1990

Effect of selection pressure for thiabendazole resistance on fitness of Haemonchus contortus in sheep.

A strain of Haemonchus contortus showing moderate levels of resistance to thiabendazole (TBZ) (egg hatch assay LC50 2.1 microM) was subjected, in alternating sequence, to 5 in vitro and 4 in vivo selections with TBZ. This resulted in a significant increase in resistance (egg hatch assay LC50 2.8 microM). The isolates obtained after each combined in vitro and in vivo selection were compared in terms of egg hatch assay, establishment, net egg output, per capita egg output and degree of pathology caused in infected hosts. The increase in resistance was accompanied by a significant increase in larval establishment in sheep (22.8-39.2%) and was associated with more severe pathology, higher egg count and worm recovery 3 months post-infection. The resistant strains were also compared with a TBZ-susceptible strain (egg hatch assay LC50 1.0 microM). The TBZ-susceptible strain had the highest net egg output, it caused the most severe pathology and it resulted in higher worm recovery than any of the TBZ-resistant strains, although the most resistant strain was very similar to the TBZ-susceptible strain. Neither the per capita egg output nor biological features of the free-living stages were significantly affected by the degree of resistance. The implications of these observations for anthelmintic use and for management of nematode anthelmintic resistance in the field are discussed.

Animals↗
PubMed1995

Ecological, morphological and genetic characterization of sympatric Haemonchus spp. parasites of domestic ruminants in Mauritania.

The 4 species of ruminants (dromedary, zebu cattle, sheep and goat) in arid areas of Mauritania harboured Haemonchus spp. as the most frequent internal parasite. This was a rare situation where the 3 putative species, H. longistipes (dromedary), H. placei (zebu cattle) and H. contortus (sheep and goat) occurred sympatrically. The study was undertaken on hosts slaughtered at the Nouakchott abattoir, on the basis of monthly collection of worms. The environment was very unfavourable to H. placei and unfavourable to H. contortus, as intensity of infection remained low throughout the year, whereas infection in the dromedary was 10 to 20-fold higher. The survival strategies during the long, dry season were different: the surviving stages were either 4th-stage larvae in digesta (dromedaries), 4th-stage larvae either in digesta or mucosae (cattle), or 4th-stage larvae in mucosae and few adults (sheep and goats). The prolificacy of female worms, indicative of the potential to contaminate pastures, was similar for all Haemonchus spp. in the rainy season. H. longistipes behave differently during the pre-rainy season as no increase of prolificacy could be demonstrated as observed in the other species. Traits of vulvar morphology are considered as markers of ecological adaptation and were studied. The knobbed and smooth female morphs (in equal proportions) were the most frequent in H. longistipes, the knobbed morph out-numbered the other morphs in H. placei, and all 3 morphs were present in sheep and goats with the linguiform form being predominant. Genetic characterization of the 3 species was performed by means of Random Amplified Polymorphic DNA (RAPD). Three groups were obtained from analysis of these data: 1 group with individuals of H. contortus, 1 group with individuals of H. placei, and 1 group with individuals of H. longistipes. This indicated that, although the 3 species were valid, H. contortus and H. placei were more similar. Intraspecific variability was 2-fold higher in H. contortus than in the 2 other species. The ecological, morphological and genetical studies showed that H. longistipes, H. placei and H. contortus could be arranged in increasing order of variability.

Adaptation, Physiological↗
PubMed1993

Characterization of proteinases in extracts of adult Haemonchus contortus, the ovine abomasal nematode.

The degradation of several protein substrates, including the blood proteins haemoglobin, albumin and fibrinogen, by proteinases present in extracts of adult Haemonchus contortus was examined over a broad pH range. These proteinases were further characterized on the basis of substrate specificity, inhibitor sensitivity and molecular size by spectrophotometric and substrate gel analysis. The majority of the proteinases capable of degrading the blood proteins tested were active at acidic pH and could be ascribed to the cysteine proteinase class. In addition, evidence is presented that these proteinases are differentially recognized and inhibited by immune sera and that parasites capable of withstanding protective host immune responses exhibit modified expression of proteinases.

Abomasum↗
PubMed1994

Immune responses of Texel sheep to excretory/secretory products of adult Haemonchus contortus.

The excretory/secretory (E/S) products of adult Haemonchus contortus comprise of at least 15 polypeptides with molecular weights ranging from 10 to > 100 kDa. These E/S products induce an immune response in infected Texel sheep, as demonstrated by specific IgG1 levels and a significant lymphocyte proliferation index. Moreover, immunoblotting analysis revealed that sera of primary H. contortus-infected sheep specifically recognize a 24 kDa E/S product. In addition, sera of challenged sheep react strongly with a 15 kDa E/S product. The other E/S products of H. contortus showed immunoreactivity with serum samples of Haemonchus-infected sheep as well as with samples of sheep harbouring other trichostrongylid infections. These cross-reacting epitopes are the main cause of the lack of specificity of an E/S material-based ELISA. This ELISA can differentiate Haemonchus infections from Nematodirus battus infections, but not from Ostertagia circumcincta or Trichostrongylus colubriformis infections.

Animals↗
PubMed1988

The population biology of the parasitic stages of Haemonchus contortus.

The population biology of the parasitic phase of the Haemonchus contortus life-cycle is discussed within the context of a mathematical model of those processes which regulate population density. Parasite death is assumed to be attributable to at least three processes: failure of the 3rd-stage larvae (L3) to exsheath following ingestion, failure to become established following exsheathment (immune exclusion), and mortality of 4th- and 5th-stage worms on the mucosal surface. The proportion of worms which avoid immune exclusion is a declining sigmoidal function of the duration of infection, at least under conditions of continual exposure to L3 larvae. Under the same conditions, the death rate of 4th- and 5th-stage worms can be modelled as an increasing linear function of the host's experience of 3rd-stage larvae. The model provides a good description of the worm burdens observed during the course of a long-term trickle infection experiment even though a priori arguments indicate that it would be an inadequate representation of events in the field.

Animals↗