Dynamics of infestation of tracers lambs by gastrointestinal helminths under a traditional management system in the North of Tunisia

The authors present a survey of gastrointestinal helminths of sheep on permanent pastures in the extreme north region of Tunisia (Mediterranean climate). Dynamic infestation of animals has been monitored by using batches of three tracer lambs introduced each two months during 2004 and 2005. These lambs were kept in the pens of veterinary school of Sidi Thabet (Tunisia) during three months and then necropsied. Faecal and blood samples were took from tracer lambs each two months during the whole period, and from animal flock only during 2004. The main helminth genera encountered were Trichostrongylus spp., Teladorsagia spp., Strongyloides papillosus and Anoplocephalidea; occasionaly were found Nematodirus, Oesophagostomum, Chabertia, Cooperia, Trichuris and Paramphistomum. The egg count of the ewes and lambs in the flock showed two peaks. For both ewes and lambs there is a gradual increase from January with a peak in May-June. This first peak is considered to be due to acquisition of infective larvae during the rainy and cold season, as evident from the worm burdens of tracer lambs. The second peak was exclusively observed in ewes during late autumn-early winter (November-December); it has two origins: infestation by third larvae stage and the periparturient rise. The worm burdens of tracer lambs showed that there was a gradual accumulation of nematodes from September- October, reaching a peak in March-April; a very low or naught infection is reported during the dry period (July-August). Infection by Anoplocephalidea was higher during the dry season. This study is primordial for a comprehensive control programme implementation against gastrointestinal helminths.     

Determination of the effective dose rate for monepantel (AAD 1566) against adult gastro-intestinal nematodes in sheep

Monepantel (AAD 1566) is the first compound from the recently discovered amino-acetonitrile derivative (AAD) class of anthelmintics to be developed for use in sheep. Three dose determination studies were conducted in Australia and Europe to identify the therapeutic dose of monepantel, when formulated for the oral treatment of sheep, to control adult gastro-intestinal nematodes. In each study, sheep infected with various nematode species were treated with either 1.25, 2.5 or 5.0 mg monepantel/kg bodyweight. Following euthanasia and worm counting, their worm burdens were compared with those from untreated control groups. At a dose rate of 1.25 mg/kg, monepantel showed efficacy above 91.9% against all major nematode species, with the exception of Chabertia ovina and Oesophagostomum venulosum. Efficacy against these two species was 93.6% and 94.0%, respectively, at a dose of 2.5 mg/kg. At this dose, efficacy was above 99.2% against nine other nematode species including Haemonchus contortus, Teladorsagia circumcincta, Trichostrongylus spp. and Nematodirus spp. It was concluded that 2.5 mg/kg would be a suitable dose rate for a commercial product.     

Effects on plasma pepsinogen, gastrin and pancreatic polypeptide of Ostertagia spp. transferred directly into the abomasum of sheep

Ostertagia spp. obtained at slaughter from the abomasa. of previously infected sheep and transferred via cannulae into the abomasa of previously worm-free sheep produced severe effects. This was shown with populations of mainly adult parasites given to three sheep and also when mixed populations of 4th stage larvae and adults were given to three other sheep. Although more severe effects were produced when both larval and adult worms were transferred, there were clearly effects when isolates composed predominantly of adults were transferred. Sheep of each group had reductions in food intake and elevations in plasma pepsinogen and gastrin within 24 h of transfer of the parasites. Further increase in plasma pepsinogen and gastrin occurred when abomasal pH rose 5–7 days after infection. Plasma pancreatic polypeptide varied but showed a general tendency to be markedly lower after infection. Ostertagia spp. eggs were detected in the faeces of infected sheep 48 h after transfer. The proportions of adults and inhibited larvae recovered from the abomasa were similar to those in the donor sheep, showing that inhibition, or arrested development of larvae, continued after their transfer to worm free sheep.     

Regulation of Haemonchus contortus populations in sheep exposed to continuous infection

Groups of sheep were infected three times weekly for 15 weeks with infective larvae of Haemonchus contortus at four rates ranging from 600 to 4800 larvae per week. At 3-weekly intervals, sheep from each group were killed for total worm counts after receiving a dose of radiolabelled larvae which enabled a direct measurement of establishment of incoming larvae during the continuing infection. Peak H. contortus burdens were reached between 6 and 9 weeks of infection, and were related to infection rate. In the groups receiving 2400 and 4800 larvae per week, worm numbers then declined rapidly and by the end of the experiment were lower than those in the groups given 600 or 1200 larvae per week. Establishment of incoming larvae was not influenced by infection rate, and declined from 45% in the first 4 weeks of infection to insignificant levels during the final 6 weeks. The proportion of incoming larvae arrested in their development increased as the infection progressed. It was concluded that H. contortus numbers were regulated by development of resistance to infection and by a loss of established worms which was related to the current rate of larval intake, and to the host's previous experience of infection.     

Seasonal variations in the populations of infective larvae on pasture and the numbers of nematode eggs in the faeces of farmed goats

The levels of nematode egg production in goats and the availability of infective larvae (L3) on pasture were investigated on a dairy unit in New South Wales, Australia. The output of eggs by adult goats was always above 300 epg. The profile of the graph of larval availability in herbage paralleled those for temperature and rainfall, suggesting that larval peaks occurred when the temperature and availability of moisture were optimal. The dominant genus was Trichostrongylus, followed by Haemonchus, then Ostertagia. A larger proportion of Haemonchus larvae in the cultures of faeces were collected during the summer months.     

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.     

Observations on parasitic gastroenteritis and bronchitis in grazing calves: Untreated calves

Pott J. M., Jpones R. M. and Cornwell R. L. 1978. Observations on parasitic gastroenteritis and bronchitis in grazing calves: untreated calves. International Journal for Parasitology8: 331–339. The epidemiology of parasitic gastroenteritis (p.g.e.) and incidental observations on lungworm infection were studied on two equal-sized paddocks of permanent pasture on a Kent farm in 1970. Groups of 10 Friesian calves were exposed for the first time to each paddock in May and grazed until the end of October. Faeces were monitored, body weights recorded and clinical signs observed. Herbage samples were examined for larvae throughout the year. Five ‘cumulative’ tracer calves exposed at the commencement were removed at monthly intervals and 5 ‘short term’ tracer calves exposed at successive monthly intervals for 1 month were killed for worm counts. A similar pattern of events occurred on both paddocks. Moderate herbage larval levels of Ostertagia and Cooperia fell to almost zero during April/May but rose quickly in July to high levels in August/September. Lower levels continued until the end of the year. Faecal egg excretion began in the test calves at the end of May, showed peaks in July and September and fell to low levels in October. D. viviparus larval excretion occurred in most calves with greater counts on Paddock 2. Clinical signs of p.g.e. complicated by lungworm infection occurred in August/September. Body weight gain which was initially rapid levelled off in June due partly to drought in addition to the parasitic infection. Weight loss occurred in August and there were 2 deaths on Paddock 1 and one on Paddock 2. Survivors gained weight again in September/October.Worm burdens in tracer calves showed quite heavy infections with Ostertagia and Cooperia as early as June/July. Massive infections complicated by lungworm disease accompanied clinical signs and death. There was evidence that aquisition of resistance to Cooperia and Dictyocaulus was acquired more readily than to Ostertagia. Inhibition of development of Ostertagia and Cooperia also became evident at the end of the trial period.     

Control of nematode parasites by grazing management—I. Decontamination of cattle pastures by grazing with sheep

Between December and June, six paddocks that had been contaminated naturally with infective larvae of nematode parasites of cattle, were each subjected to one of six grazing treatments. The effects of these treatments on the infectivity of the pastures were assessed by slaughter and total worm counts of calves which subsequently grazed the paddocks, and by counts of larvae on pasture samples. Grazing by sheep during autumn resulted in reduced populations of Ostertagia ostertagi and Trichostrongylus axei, both in calves and on the pasture for up to 12 months after treatment. Similar effects were noted on pasture grazed by cattle repeatedly dosed with anthelmintics, and on ungrazed pasture. There was little effect of any treatment on numbers of Cooperia oncophora. It was concluded that the level of autumn contamination was of importance in determining the numbers of O. ostertagi in the following spring and summer.     

Control of nematode parasites by grazing management—II. Decontamination of sheep and cattle pastures by varying periods of grazing with the alternate host

Commencing in December 1970, paddocks of a uniform series of sheep pastures were grazed for 6, 12 or 24 weeks by either yearling steers is or yearling ewes. Cattle pastures were treated similarly. All ewes and steers were pre-dosed with anthelmintic.At the conclusion of alternate grazing the effectiveness of the grazing treatments was evaluated by grazing each paddock for 1 month with either ten worm-free lambs (sheep pastures) or three wormfree calves (cattle pastures). The test animals were then slaughtered for total worm counts. The grazing of sheep pastures with cattle for 6, 12 or 24 weeks from December onwards resulted in reductions in numbers of Haemonchus contortus and Trichostrongylus colubriformis in test lambs. In comparison with continuous grazing by sheep, Nematodirus spp was only reduced after 24 weeks grazing by cattle. Cattle pastures grazed by sheep for 6 weeks showed no reduction in numbers of Ostertagia ostertagi or Cooperia oncophora in test calves. However after 12 weeks with sheep, numbers of O. ostertagi though not C. oncophora were reduced and after 24 weeks of alternate grazing both these species were reduced.Calves following the 6 week sheep treatments acquired both Haemonchus contortus and Trichostrongylus colubriformis and the calves from the 12 week sheep treatment paddock also carried H. contortus. For sheep the only evidence of cross-transmission was the occurrence of small numbers of Cooperia oncophora in test lambs from the 24 week cattle grazing treatment.The results provide evidence that sequential stocking with cattle and sheep in conjunction with anthelmintic treatment is an effective management strategy for preparing parasitologically safer pastures, but further information is required to determine the optimum timing of sequential stocking in farming situations.     

Host-induced effects on the morphology of Ostertagia circumcincta in grazing sheep

Waller P. J. and Thomas R. J. 1978. Host-induced effects on the morphology of Ostertagia circumcincta in grazing sheep. International Journal for Parasitology8: 365–370. Morphological changes are described in adult Ostertagia circumcincta recovered from lambs slaughtered monthly from June to November. Comparisons were made between continuously grazed principal lambs exposed to infection for increasingly long periods, and tracer Iambs exposed for only four weeks. Lambs were either slaughtered immediately off pasture or held 2 weeks under worm free conditions before slaughter. Although no resistance to establishment of infection occurred, shown by comparable worm burdens of principals and tracers, a rapid parasite population turnover was observed in the principal lambs which ceased once the lambs were removed from the infected pasture. Significantly fewer adult female Ostertagia with fully developed vulval flaps were recorded in principals compared to tracers. Such differences are considered to be primarily host induced responses which rapidly come into play, evidenced by the entire populations of developing female worms in tracers slaughtered immediately off pasture showing arrested vulval flap development after they had completed development in those tracers held for a further two weeks. A progressive reduction in the proportion of adult male worms also occurred in the principal lambs. Population density was shown to have an inverse correlation with worm size. By analogy, it is suggested that seasonal change in the presence of cuticular inflations in Haemonchus contortus females with linguiform vulval flaps may be primarily host induced, rather than a change in the relative fitness of discrete genotypes.     

Micrometeorological factors involved in development and survival of free-living stages of the sheep nematodesHaemonchus contortus andTrichostrongylus colubriformis. A review

Temperature and soil moisture are the most important factors affecting the development and survival ofHaemonchus contortus andTrichostrongylus columbriformis eggs and larvae on pasture. More than half of the eggs develop into infective larvae in the laboratory, but a very low percentage (0.03% forH. contortus) does so on pasture. There is a marked difference betweenH. contortus andT. colubriformis in survival of infective larvae.H. contortus larvae survived in the winter at Urbana poorly, whereasT. colubriformis did well. The former survived better than the latter in the spring and worse in the summer, while both survived equally well in the fall. Technics for larval recovery from pasture are not very efficient. Meteorologic conditions at ground level where the larvae exist are quite different from those in a standard weather shelter 1.6 m above the ground. Bioclimatographs in which mean monthly maximum temperatures are plotted against total monthly precipitation are fairly good in predicting the type of nematode liable to be important in a given region, but they are too simplistic to be relied on for more than approximations.     

Failure to vaccinate lambs against Haemonchus contortus with functional metabolic antigens identified by immunoelectrophoresis.

Metabolic products from Haemonchus contortus larvae cultured in vitro from the infective third to fourth stage were collected and concentrated. Chromatographic and immunoelectrophoretic analyses were made to study the numbers and activity of antigens in the metabolic products derived from the in vitro cultured larvae. Three-month-old lambs were given a series of injections of metabolic antigens with and without adjuvant at dose rates of 0·05, 0·5 and 5·0 mg antigen protein per injection. These animals and saline injected controls were each challenged with 3000 H. contortus infective larvae after the last antigen injection and killed 35 days later. No difference was seen in the faecal worm egg counts or the differential worm counts among the vaccinated and control animals. The antigen preparation of worm metabolic products conferred no resistance to challenge infection with the parasite.     

Strategies for the avoidance of faeces by grazing sheep

Experiments were conducted to investigate which environmental cues were used by sheep when discriminating against patches of pasture contaminated with faeces. The influence of the spatial distribution of contaminated patches and the parasite infection status of sheep on avoidance of contaminated patches and ingestion of parasite larvae was also investigated. In experiment 1, sheep infected with the parasite Ostertagia circumcincta were given the opportunity to graze in uncontaminated or aggregated contaminated patches. Patch contamination comprised of either faeces from sheep infected with O. circumcincta larvae, faeces from uninfected sheep, or O. circumcincta larvae only. Infected sheep discriminated against faeces from parasite-infected animals and faeces from uninfected animals equally. Sheep did not discriminate against patches contaminated with parasite larvae only. In experiment 2, sheep infected with O. circumcincta and uninfected sheep grazed experimental plots with differing spatial patterns of faecal-contaminated patches, allowing animals the opportunity to forage in contaminated or uncontaminated patches of herbage. Plots were also grazed by infected and uninfected animals that were fistulated at the oesophagus to enable the collection of ingested herbage. Sheep spent a greater proportion of their time foraging in uncontaminated patches than in contaminated patches. Where patches were highly aggregated, infected animals spent a greater proportion of total grazing time in uncontaminated patches than did uninfected animals, and grazed uncontaminated patches for longer on each sampling occasion. On grazing plots where all patches were contaminated, the difference between the numbers of larvae isolated from pasture herbage and ingested herbage was greatest for infected animals. In this situation, infected animals avoided parasites most. On grazing plots consisting of both contaminated and uncontaminated patches, the difference between the numbers of larvae isolated from pasture herbage and ingested herbage was greatest for uninfected animals. In this situation, uninfected animals were most effective at parasite avoidance as they consumed fewer parasite larvae relative to what was available on pasture.     

Population dynamics of the parasitic stages of Ostertagia spp. in sheep

Callinan A. P. L. and Arundel J. H. 1982. Population dynamics of the parasitic stages of Ostertagia spp. in sheep. International Journal for Parasitology12: 531–535. The development and survival of continuing infections of Ostertagia spp. in weaner sheep were studied in order to develop a general model of the parasitic stages of the life cycle of these sheep nematodes. After 10 days, 13.8% of infective larvae (L₃) given at the rate of 1 dose of 1000 L₃ twice per week (group 1) and 20.8% of L₃ given at the rate of 10,000 L₃ twice per week (group 2) were recovered in the first of the serial nematode counts. In the final counts at 137 days, 7.7 and 0.7% were recovered in these groups. The build up and maintenance of nematode populations was regulated and related to the level of infection. A model in which the death rate of the parasitic stages was a function of the time of exposure to infection and rate of infection was used to describe the serial total nematode counts. During the experiment there was no noticeable trend in numbers of fourth stage larvae (L₄) in nematode counts, the size of adult nematodes, nematode egg counts (EPG) and egg output per female nematode (EPF). After 112 days, liveweight gains were significantly reduced in group 2 only, but increases in wool lengths were significantly reduced in both groups.     

The influence of weather and egg contamination on the development of third-stage larvae of Cooperia oncophora on pasture.

The influence of weather and egg contamination on the dynamics of herbage contamination with infective larave of Cooperia oncophora was investigated on artificially contaminated grass plots and in a grazing experiment with 24 first-year grazing calves from May to October 1987 in Lower Saxony, Germany. On the experimental plots the larval translation was highest at the beginning of July and in the second part of September, following high mean weekly temperatures. Between July and September peak recovery of larvae from herbage occurred 4 weeks after contamination. A seasonal pattern of larval translation similar to that on the experimental plots could be demonstrated on the grazed pastures when the number of larvae per m2 of pasture had been adjusted to the previous egg output by means of a contamination index. The resulting 'relative larval density' is regarded as a good indicator for larval development on pasture. From July to September the larval population on pasture resulted mainly from the egg contamination 2-3 weeks earlier. The short persistence of the infective larvae on herbage was probably due to the frequent and heavy rainfall throughout the season, causing a passive washout of larvae into the soil. On single pastures the larval density started to increase within 1 week after the calves had first contact with these fields. The impact of the calves on the distribution of larvae is discussed.     

A preliminary study of the effect of microclimate on third-stage larvae of Haemonchus contortus and Haemonchus placei on irrigated pasture.

Assessments were made on the influence of several microclimatic variables on the availability of third-stage larvae of Haemonchus contortus and Haemonchus placei on four strata of irrigated Kikuyu pasture. Three replicates of these pasture samples were collected on 18 sample days over 12 months and the log10 mean counts of the larvae recovered were analysed by a step-wise regression model. Predictors for the log counts of the four strata for the two nematode species included relative humidity, illumination, air temperature and windspeed. The effect of air temperature on larvae of both Haemonchus species was similar; as air temperature increased, the number of larvae on pasture increased. The inverse was true for windspeed; as windspeed increased larval counts decreased. For H. contortus, relative humidity increased as the number of larvae increased on all strata except upper herbage. The R2 values ranged from 0.11 to 0.21 for H. contortus and from 0.04 to 0.12 for H. placei. Under the conditions of this study, only 21% of the effect on H. contortus and 12% on H. placei third-stage larvae on pasture can be explained by microclimatic conditions.     

The statistical distribution of trichostrongylid nematodes in grazing lambs

Barger I. A. 1985. The statistical distribution of trichostrongylid nematodes in grazing lambs. International Journal for Parasitology15: 645–649. Worm counts of the genera Haemonchus, Ostertagia, Trichostrongylus and Nematodirus from 104 naturally-infected lambs were found to follow negative binomial distributions. Although mean counts covered a 10-fold range, k values did not differ significantly among the four genera. It was concluded that all genera were equally overdispersed. Some implications of this overdispersion for flock productivity and for parasite control were examined     

The response of sheep to parenteral vaccination and immunizing infections against the abomasal nematode, Haemonchus contortus

Adams D. B., Beh K. J. and Davies H. I. 1982. The response of sheep to parenteral vaccination and immunizing infections against the abomasal nematode, Haemonchus contortus. International Journal for Parasitology12: 445–449. Subcutaneous injection of relatively large amounts of unfractionated homogenates of adults plus infective larvae of Haemonchus contortus emulsified in complete Freund's adjuvant produced a degree of protective immunity when challenge infection was given eight weeks after the first or only dose of vaccine. In an attempt to improve acquired immunity, parenteral vaccination was either followed or preceded by a short immunizing infection with H. contortus, which was terminated by anthelmintic before patency. This treatment aimed at stimulating general responsiveness to worm antigens and invoking mucosal immunity in the abomasum. Disparate results were obtained; immunizing infections either increased immunity or made sheep more susceptible to challenge infection. In this latter situation, the unresponsiveness associated with primary infection with H. contortus may have been increased.     

A Study on the Transmission of Oesophagostomum dentatum and Hyostrongylus rubidus among Outdoor Reared Pigs in Denmark

This study was carried out to obtain basic information on the transmission of Oesophagostomum dentatum and Hyostrongylus rubidus in outdoor reared pigs in Denmark. Eighteen 10 weeks old worm-free pigs were allocated into 3 groups of 6 pigs each. In May, all pigs were turned out on the same parasitologi-cally naive pasture, and after 2 weeks the pigs in groups 2 and 3 were experimentally infected with 10,800 O. dentatum and 8,700 H. rubidus infective larvae, respectively. Pigs in group 1 served as non-infected controls. All pigs were reared together on the experimental pasture for further 134 days until slaughter in October. Strongyle egg counts, differentiation of infective larvae at species level, serum pepsinogen, and herbage larval infectivity were monitored at regular intervals throughout. Both strongyle species established in the originally parasite-free pigs (group 1) and cross infections were established in group 2 and 3. The pigs were exposed to steadily increasing herbage infectivity of both species of strongyles. At the end of the experiment, geometric mean worm burdens of O. dentatum in groups 1,2 and 3 were 1202, 6136 and 1431 respectively, the burden in group 2 being significantly higher (p<0.05) than that of the 2 other groups. The geometric mean worm burdens of H. rubidus in groups 1, 2 and 3 were 4907, 3679 and 5246 respectively, showing no significant differences between groups.     

Effects of reproduction,genotype and anthelmintic treatment of ewes on Ostertagia spp. populations

Donald A. D., Morley F. H. W., Waller P. J., Axelsen A., Dobson R. J. and Donnelly J. R. 1982. Effects of reproduction, genotype and anthelmintic treatment of ewes on Ostertagia spp. populations. International Journal for Parasitology12: 403–411. Merino and Border Leicester × Merino (BL × M) ewes, nearly all of the same age and reared at the same site, were either unmated or mated to Border Leicester rams. Ewes of each genotype and reproductive status were untreated or were given a single pre-lambing drench with thiabendazole at 50 or 100 mg/kg a week before the start of lambing in spring on pastures at Canberra which had been contaminated during autumn and winter by adult sheep. The two genotypes grazed together within each combination of reproductive status and anthelmintic treatment which grazed separately. Thiabendazole was highly effective in removing both fourth stage larvae and adults of Ostertagia spp., the most abundant genus. Eight weeks after the pre-lambing drench lactating ewes carried larger Ostertagia spp. populations than did unmated ewes of both genotypes, but as a result of reinfection after treatment, differences between drenched and undrenched ewes were not significant. At this time lactating as well as unmated ewes harboured large populations of arrested early fourth stage larvae of Ostertagia spp. acquired during the last 8 weeks, showing that arrest of development is not prevented by lactation. There was strong evidence that some ingested larvae which became arrested in lactating ewes were rejected by unmated ewes. At all stages of the reproductive cycle studied, BL × M ewes were substantially more resistant to Ostertagia spp. infection than Merinos. No persistent benefits in parasite control or in animal production were detected from the pre-lambing drench.