Rumen function and digestion of Merino sheep divergently selected for genetic difference in resistance to Haemonchus contortus

The aim of this experiment was to investigate the effect of Haemonchus contortus infection on rumen function and digestion of Merino sheep from lines divergently selected for genetic resistance to H. contortus. Rumen function and whole-tract digestibility were determined in 29 Merino weaner wethers from the CSIRO Haemonchus selection flock, increased resistance to Haemonchus (IRH), decreased resistance to Haemonchus (DRH) and random bred control (C) selection lines. Wethers were fed a restricted low quality roughage diet (5.9 MJ ME/kg DM, 29 g MP/kg DM) to maintain weight and were either uninfected (NIL) or infected with H. contortus (INF). Measurements were taken at week 7 of infection, while animals were housed in metabolic crates. Animals were then euthanased to determine abomasal worm counts. IRH had significantly lower worm egg count than DRH and C lines however, adult worm and larval counts, though lower in IRH animals, did not differ significantly. DM and OM digestibility and in sacco degradability of IRH and DRH selection lines were greater than C animals. Rumen function of animals from the IRH line was altered in response to H. contortus infection with an increase in fluid outflow and turnover rate and a decrease in propionic acid concentration. These changes may be a component of a greater host resistance and begin to explain the failure of IRH animals to translate lower worm egg count into greater bodyweight gain.     

Voluntary feed intake and diet selection of Merino sheep divergently selected for genetic difference in resistance to Haemonchus contortus

This research was designed to determine if divergent selection for resistance to Haemonchus contortus had produced correlated changes in voluntary feed intake and diet selection. Voluntary feed intake, diet selection and production were determined in 54 Merino weaner rams from the CSIRO Haemonchus selection flock, increased resistance to Haemonchus (IRH), decreased resistance to Haemonchus (DRH) and random bred control (C) selection lines. Weaner rams were fed ad libitum either a high (9.2 MJ ME/kg DM, 90 g MP/kg DM) or moderate (6.3 MJ ME/kg DM, 30 g MP/kg DM) quality diet and given the choice between the two diets, when uninfected (NIL) or infected with H. contortus (INF). Symmetrical response to divergent selection for worm egg count (WEC) was not matched by a symmetrical change in feed intake and there was no difference in diet selection between selection lines. Feed intake, growth and wool production of DRH animals remained the same as that of IRH, yet DRH animals had five times greater WEC than IRH. This study begins to explain the mechanisms that allow resistant animals to effectively prevent establishment and/or development of H. contortus, by maintaining a greater immune response to infection through higher circulating eosinophils, plasma globulin and IgG(1) antibody titres. Susceptible animals have displayed resilience by improving feed conversion efficiency and increasing protein synthesis.     

The response of Awassi and Merino sheep to primary infection with Haemonchus contortus

The possible existence of breed differences in the response of sheep to primary infection with Haemonchus contortus was examined by comparing worm establishment and pathogenic effects of the parasite in Awassi and Merino sheep of haemoglobin type B infected with 500 third stage H. contortus larvae per kg body weight. The results showed that the Merino sheep had lower faecal egg counts and worm burdens and suffered less severe clinical disturbances than sheep of the Awassi breed. This suggests that genetic resistance operates primarily at the level of worm establishment.     

The problem of escalating resistance of Haemonchus contortus to the modern anthelmintics in South Africa

During the past decade in South Africa there has been a continual increase in sheep of strains of gastrointestinal helminths resistant to the modern anthelmintics. Five strains of Haemonchus contortus are described in this paper. Despite the fact that 2 of the 5 strains were tested for susceptibility only to ivermectin, a total of 10 instances of resistance were found. Four of the 5 strains were resistant to ivermectin, 2 to closantel, 2 to rafoxanide and 2 to the benzimidazoles. One of these strains was concurrently resistant to 3 different anthelmintic groups, namely, the ivermectins, the benzimidazoles and the salicylanilides. Resistance to ivermectin developed in 2 strains of H. contortus after a history of only 3 treatments with this compound in one instance and 11 treatments in the other. In the latter case drenching with ivermectin was well interspersed with that of other anthelmintics. This rapid development of resistance suggests that there may be cross-resistance between ivermectin and another anthelmintic group. Two of the ivermectin resistant strains were recovered from separate properties in the south-western Cape Province, where Ostertagia circumcincta, which is usually the dominant parasite in this region, was virtually eliminated by the anthelmintic treatment. On each of these properties it was apparently replaced by a resistant strain of H. contortus. A serious threat to control is the dissemination of worm strains with multiple resistance to anthelmintics. The strain of H. contortus resistant to 3 anthelmintic groups has already been widely dispersed, as the farmer concerned suddenly decided to give up farming with sheep and to sell his flock.     

Selection for resistance to macrocyclic lactones by Haemonchus contortus in sheep.

An anthelmintic-sensitive Haemonchus contortus strain was selected for moxidectin and ivermectin resistance concurrently for 22 generations. Treatment with 0.002 mg moxidectin/kg BW or 0.02 mg ivermectin/kg BW produced >99% efficacy against the susceptible parent strain passaged for 22 generations without any anthelmintic exposure. However, to obtain similar efficacy the moxidectin-selected and the ivermectin-selected strains of H. contortus required 0.05 mg moxidectin/kg BW or 0.4 mg ivermectin/kg BW. These results indicate that development of resistance to one macrocyclic lactone, simultaneously results in resistance to another macrocyclic lactone. However, rates of resistance development differ between compounds and occurs more slowly with moxidectin than with ivermectin.     

Heterosis analysis of Haemonchus contortus resistance and production traits in Rhoen sheep, Merino Land sheep and crossbred lambs

A crossbreeding program was conducted to evaluate the resistance status to Haemonchus contortus in reciprocal crossbred (F(1)) lambs and their Merino Land sheep (Ml) and Rhoen sheep (Rh) purebred counterparts. A total of 406 lambs were included in the study. Faecal egg counts (FEC) and haematocrit values (Hc) of all the lambs were collected 4 and 8 weeks after an artificial infection with H. contortus. FAMACHA scores were given at the same time. Worm counts of all lambs were obtained after slaughtering at 21 weeks of age. The Ml group consistently showed the highest body weights, daily weight gains and best carcass performances, whereas the Rh group showed the lowest results. Most of the body weight parameters in the F(1) group fell intermediately between the Ml and Rh groups, though the heterosis analysis of body weight and carcass parameters showed a tendency to favour the crossbreeding group Rh x Ml. The Rh group showed, on average, the highest FEC and the lowest Hc values compared to the Ml. However, the nematode counts were not significantly different between the pure breeds. In comparison, the worm counts of the Rh x Ml crosses were significantly different from those in the Ml x Rh crosses. A heterosis analysis showed that FEC, Hc and the worm count in the F(1) group favoured the crossbreeding group Rh x Ml. These results suggested that crossbreeding Rh to Ml sheep may be a suitable way of producing lambs with improved resistance to H. contortus infection without any negative effects on production traits.     

Haemonchus contortus resistance to ivermectin and netobimin in Brazilian sheep.

Suffolk, Texel, Hampshire Down and Ile de France sheep from the municipalities of Porto Amazonas, Piraquara and Araucaria in the State of Paraná, and Bagé in the State of Rio Grande do Sul were brought to Sobral, State of Ceará, to be used in a cross-breeding project. On arrival they had clinical signs of nematode parasitosis, and one Suffolk female died. The animals were treated orally with ivermectin (0.2 mg kg-1) and fifteen days later with netobimin (20.0 mg kg-1). Neither drug reduced the egg counts (measured in eggs per gram, EPG) significantly, and this suggested that the nematodes in the sheep were resistant to the anthelmintics used. Haemonchus contortus was the species involved. The egg counts were reduced after oral treatment with trichlorfon (100.0 mg kg-1). Haemonchus contortus larvae obtained from these animals before trichlorfon treatment and passaged through two nematode-free sheep were used in a further experiment. Twenty 6- to 9-month-old nematode-free lambs were infected with the H. contortus larvae (10,000 per animal) and after the infection was confirmed, were randomly divided into four groups of five animals. Group I was orally treated with ivermectin at 0.2 mg kg-1, Group II with oral netobimin at 20.0 mg kg-1, Group III with oral trichlorfon at 100.0 mg kg-1 and Group IV was a non-treated control. Egg counts and faecal cultures were taken before dosing on the day of treatment and seven days later when all animals were necropsied and the nematodes were collected from the abomasa and counted.(ABSTRACT TRUNCATED AT 250 WORDS)     

Functional investigation of a QTL affecting resistance to Haemonchus contortus in sheep

This study reports a functional characterization of a limited segment (QTL) of sheep chromosome 12 associated with resistance to the abomasal nematode Haemonchus contortus. The first objective was to validate the identified QTL through the comparison of genetically susceptible (N) and resistant (R) sheep produced from Martinik × Romane back-cross sheep. The R and N genotype groups were then experimentally infected with 10 000 H. contortus larvae and measured for FEC (every three days from 18 to 30 days post-challenge), haematocrit, worm burden and fertility. Significant differences in FEC and haematocrit drop were found between R and N sheep. In addition, the female worms recovered from R sheep were less fecund. The second step of the characterization was to investigate functional mechanisms associated with the QTL, thanks to a gene expression analysis performed on the abomasal mucosa and the abomasal lymph node. The gene expression level of a candidate gene lying within the QTL region (PAPP-A2) was measured. In addition, putative interactions between the chromosome segment under study and the top ten differentially expressed genes between resistant MBB and susceptible RMN sheep highlighted in a previous microarray experiment were investigated. We found an induction of Th-2 related cytokine genes expression in the abomasal mucosa of R sheep. Down-regulation of the PAPP-A2 gene expression was observed between naïve and challenged sheep although no differential expression was recorded between challenged R and N sheep. The genotyping of this limited region should contribute to the ability to predict the intrinsic resistance level of sheep.     

Genetic parameters for indicators of host resistance to parasites from weaning to hogget age in Merino sheep

Fecal egg count (FEC) has been widely used as an indicator of host resistance to gastrointestinal parasites in sheep and has been shown to be a heritable trait. Two other possible indicators of parasites, dag score (DS; accumulated fecal material) and fecal consistency score (FCS), were investigated in this study, along with BW. All four traits were studied to see how heritability and genetic correlations varied with age from weaning (4 mo) to hogget age (approximately 400 d). More than 1,100 lambs, the offspring of 37 rams, were recorded eight times between weaning (3 to 5 mo of age) and hogget age (13 to 18 mo of age) on two farms. Sire models were fitted to the data from each trait at each recording and in a repeatability model involving the whole data set. Overall, the heritabilities were 0.28+/-0.072 (FEC), 0.11+/-0.036 (DS), 0.12+/-0.036 (FCS), and 0.23+/-0.070 (BW). By fitting random regression models to the time-series data, it was possible to see how these heritability values varied as the lambs aged, from weaning to hogget age. The heritability of FEC rose from 0.2 at weaning to 0.65 at 400 d. Dag score had a higher heritability (0.25) in the middle of the age range and a low value at weaning (<0.1) and hogget age (0.16). The heritability of FCS was low, with a value of 0.2 at weaning reducing to 0.05 as the animals aged. Body weight had zero heritability at weaning, which rose to greater than 0.6 at hogget age. Most traits had low genetic correlations between them, the only exception being that between FCS and DS (0.63). Most genetic correlations varied little over the age range with the exception of FEC and BW, which fell from 0 at weaning to -0.63 at hogget age. Whereas FCS and DS may be good indicators of scouring, they are very different from FEC as an indicator of host resistance to gastrointestinal parasites.     

T-cell receptor and immunoglobulin gene polymorphisms and resistance to Haemonchus contortus in sheep

SUMMARY: DNA extracted from animals originating from three large half-sib sheep families derived from a single, highly resistant ram were probed with ovine T-cell receptor α and β chain and immunoglobulin lambda light chain variable and constant region cDNA probes. When genomic DNA was digested with TaqI, 19 polymorphic bands were detected with the lambda light chain immunoglobulin variable region probe and 6,4 and 5 polymorphic bands were detected with the T-cell receptor α and β probes and the lambda immunoglobulin constant region probes, respectively. All animals had been phenotypically assessed for Haemonchus contortus resistance by faecal egg counts and using best linear unbiased statistical methods, no statistically significant associations were found between RFLP banding patterns detected at these loci and faecal egg counts. The results show that germline polymorphism at these loci does not account for a significant proportion of the variation in resistance to Haemonchus contortus in this flock. ZUSAMMENFASSUNG: T-Zellrezeptor und Immunoglobulin-Genpolymorphismus und Resistenz von Schafen gegen Haemonchus contortus DNS aus Tieren von drei gro?en Halbgeschwister-Schaffamilien aus einem einzelnen hochgradig resistenten Schafbock wurden mit cDNS-Sonden aus ovinen T-Zellrezeptor α- und β-Ketten und Immunoglobulin λ leichten variablen Ketten und konstanten Regionen untersucht. Nach Verdau der genomischen DNS mit TaqI 19 wurden polymorphe Banden mit der λ leichte Ketten Immunoglobulin variablen Region-Sonde entdeckt, 6, 4 und 5 polymorphe Bande wurden mit T-Zellrezeptor α- und β-Sonden und mit der Sonde der λ Immunoglobulin konstanten Region gefunden. Alle Tiere wurden ph?notypisch in Hinblick auf Haemonchus contortus-Resistenz mittels f?kaler Eiproben untersucht. Statistische Analyse mittels linearer unverzerrter Pr?diktion ergab keine statistisch signifikanten Zusammenh?nge zwischen RFLP Bandmustern und f?kalen Eizahlen. Genetischer Polymorphismus an diesen Loci scheint nicht für einen signifikanten Teil der Variabilit?t in der Resistenz gegen Haemonchus contortus verantwortlich zu sein.     

Factors affecting the prevalence and severity of footrot in a Merino flock selected for resistance to Haemonchus contortus

SUMMARY An outbreak of virulent footrot occurred in three lines of sheep selected for different levels of resistance to Haemonchus contortus , providing an opportunity to examine the factors affecting footrot severity. The selection lines did not differ significantly in average footrot score, but various other factors were found to have an effect. In breeding ewes, the average score was 10.2 (maximum possible score 20) and the prevalence of footrot was 96%. There was a significant age effect, with average score increasing from 9.1 ± 0.8 in 2-year-old ewes, to 13.4 ± 0.9 in 7-year-old ewes. Ewes that lost their lambs tended to have higher scores than those that either failed to lamb or successfully reared their litter. Significant differences in footrot score were caused by the lambing paddock in which the ewes had been, during the period seven weeks to one week before infection. Among lambs, aged about 20 weeks, the average footrot score was lower (5.0), as was the prevalence (75%). The score increased by an average of 0.09 ± 0.03 per day of age and the prevalence increased by an average of 0.6%± 0.3% per day of age. Lambs born and reared as twins had significantly lower scores than singles, and progeny of maiden dams had lower scores than progeny of older ewes. There was no significant difference because of the sex of the lamb. The paternal half-sib estimate of heritability of footrot score was 0.20 ± 0.14 and of the genetic correlation between footrot score and worm egg count was essentially zero (0.02 ± 0.20), but these estimates were probably biased by the confounding of sires with lambing paddocks. Nevertheless, the results strongly suggest that there are no adverse genetic associations between resistance to footrot and resistance to internal parasites.     

Performance of hair sheep composite breeds: resistance of lambs to Haemonchus contortus

This study was designed to evaluate breed differences in resistance to Haemonchus contortus in lambs. A total of 181 ewe lambs representing crossbred Dorsets (DO) and Dorpers (DP; out of 1/2-Dorset, 1/4-Rambouillet, 1/4-Finnsheep ewes) and straight-bred Katahdins (KT) were evaluated over 3 yr. An additional 144 DO, DP, KT, and Barbados Blackbelly x St. Croix (HH) wethers were evaluated over 2 yr. Lambs were weaned at 60 to 90 d of age. After deworming at about 4 mo of age, ewe lambs received approximately 10,000 infective larvae and were evaluated for parasite resistance in drylot, whereas wethers were evaluated on pasture under conditions of natural infection. Each sex was analyzed separately. Egg counts per gram of feces (FEC), log-transformed FEC (LFEC), packed cell volumes (%), and body weights (kg) measured at 3, 4, 5, and 6 wk after deworming and reinfection were analyzed. Breed influenced all traits (P < 0.05) except BW in ewe lambs and PCV in wethers. Year and week influenced (P < 0.05) all traits. At most times, DP had the highest FEC, DO had the lowest PCV, and KT and HH had lower FEC and higher PCV than either DO or DP. Clearly, Dorper sheep were not more resistant to parasites than DO, but they were able to cope with infection better by maintaining similar or higher (P < 0.05) PCV and similar BW. Katahdin and HH were more resistant, with lower FEC (P < 0.05) than DO or DP. Breed differences were more apparent when infection levels were higher; DO and DP were less affected when infection levels were low and when animals were on a better plane of nutrition. Caribbean hair sheep originated in hot, humid regions of West Africa. They and the derivative Katahdin breed seem relatively resistant to parasitism. The Dorper, in contrast, was developed in more arid regions and exhibited little parasite resistance.     

Comparison of three in vitro techniques to estimate benzimidazole resistance in Haemonchus contortus of sheep

Three in vitro assays to detect benzimidazole resistance, namely, the egg-hatch assay, tubulin-binding assay, and a larval-development assay, were evaluated by estimating the level of benzimidazole resistance in three field isolates of Haemonchus contortus compared with a susceptible reference strain. Comparisons were also made with estimates of benzimidazole resistance of the three field strains obtained from an in vivo controlled anthelmintic efficacy test. All three in vitro tests showed similar, consistent results which also suggested greater sensitivity than the in vivo assay. These results indicate that selection of an in vitro technique to determine benzimidazole resistance should therefore be based on considerations other than precision, such as technical expertise, availability of equipment, cost and speed in which diagnosis is required.     

High genetic correlation between resistance to Haemonchus contortus and to Trichostrongylus colubriformis in INRA 401 sheep

To compare the protection acquired by infection with Haemonchus contortus or Trichostrongylus colubriformis, successive cross infections with the two species were conducted in two genetically similar groups of 150 ewe lambs from 30 sires of INRA 401 breed. At 6 and 7.5 months of age, each group received two doses of 10000 L3 separated by a treatment of one of the species, followed by a similar infection with the second species at 9 and 11.5 months. Faecal egg counts (FECs) were done 28 and 35 days after each dose. There were very high genetic correlation within and between species suggesting that the environmental conditions were very similar. The heritability of FEC of H. contortus ranged from 0.39 to 0.48 and the genetic correlations between FEC after the first and second infections with the same or different species were all near 1. Selection against this species could be done on the response to only one dose. For T. colubriformis, only the response to challenge infection had a similar heritability (0.47); the genetic correlations within and among species were also near one. These results will assist the selection of sheep resistant to nematode parasites.     

Rapid selection for ivermectin resistance in Haemonchus contortus

Lambs infected with two isolates, one British and one American, of Haemonchus contortus were treated with increasing doses of ivermectin. Eggs from the highest dose that had not eliminated the infection were cultured and larvae used to infect another lamb. After three generations the H. contortus was resistant to 0.2 mg/kg ivermectin. The results stress the ease with which ivermectin resistance can be selected if high selection pressure is applied.     

Hypergastrinaemia of sheep infected with Haemonchus contortus

Radioimmunoassay of plasma gastrin has shown that a hypergastrinaemia occurs in yearling sheep, after infection with 10(6) infective third stage larvae of Haemonchus contortus. Plasma gastrin concentrations began to rise two to four days after infection, that is, at about the same time as, or slightly before, the abomasal pH was observed to increase.     

Trichostrongylus colubriformis and Haemonchus contortus infections in light bodyweight Merino lambs

OBJECTIVE; To show that low bodyweight is a predisposing cause of high Trichostrongylus colubriformis and Haemonchus contortus burdens and egg counts in Merino lambs. DESIGN: A comparison was made, among lambs of different bodyweights, on the effect on immunity of a primary or secondary viable infection with T colubriformis or H contortus larvae. PROCEDURE: Sixty-one Merino lambs, 1 or 6 months of age, were penned indoors and given primary, or both primary and secondary, infection of T colubriformis or H contortus. Faecal egg counts, worm counts and parasite-specific immunoglobulin concentrations were examined for their relationships with bodyweight. RESULTS: Bodyweight at the start of a primary infection was correlated with worm burden, worm fecundity and jejunal IgA antibody concentration. Merino lambs weighing less than 23 kg at the time of first exposure to T colubriformis or H contortus had impaired ability to develop protective mucosal immunity and to resist homologous challenge. CONCLUSION: If the goal is to ensure that lambs develop immunity before weaning, then every endeavour should be made to achieve the combination of critical bodyweight and exposure to moderate levels of nematode infection as soon as possible.