The effect of feeding sulla (Hedysarum coronarium) or lucerne (Medicago sativa) on lamb parasite burdens and development of immunity to gastrointestinal nematodes

Two experiments investigated the efficacy of the legume Hedysarum coronarium (sulla), which contains condensed tannins (CTs), for reducing gastrointestinal nematode infections relative to lucerne. Experiment 1 was aimed to show whether the lower faecal egg count (FEC) and larval establishment previously reported in lambs grazing sulla were due to direct effects of the forage on Ostertagia circumcincta and Trichostrongylus colubriformis or were mediated through an enhanced immune response. Experiment 2 evaluated the impact of feeding sulla relative to feeding lucerne (Medicago sativa), before, at, or after larval challenge on subsequent FECs and nematode burdens. In experiment 1, 64 Romney lambs were fed either freshly cut lucerne or sulla (32 lambs per herbage) for the duration of the trial. Within each herbage there were four treatment groups (n=8 per group). Initial levels of immunity were assessed in uninfected (UN) lambs which were maintained parasite-free until challenged with 15,000 O. circumcincta and 15,000 T. colubriformis larvae on day 63, and slaughtered on day 81. The other three treatment groups were trickle-infected with each of 5000 O. circumcincta and 5000 T. colubriformis larvae three times per week from day 1 to 35. Non-steroid infected (CONTROL) and steroid-treated (STER) groups were treated with anthelmintic on day 49 and challenged with 15,000 O. circumcincta and 15,000 T. colubriformis on day 63 and slaughtered on day 81. The STER lambs were given dexamethasone trimethylacetate from day 49 to 81 to determine effects of immunity on parasite infection. From day 35 an establishment group (EST) on each herbage was fed a common pelleted lucerne diet and slaughtered on day 56 to determine nematode establishment during trickle-infection. Diet did not affect FECs but feeding lucerne increased (P<0.05) numbers of T. colubriformis in CONTROL lambs compared to those fed sulla. O. circumcincta numbers were lower (P<0.05) in UN lambs fed sulla than lucerne. The sulla diet was associated with higher (P<0.05) antibody titres against secretory-excretory antigens to adult O. circumcincta and to adult and larval T. colubriformis, so there appeared to be some immunogenic response to the sulla diet but effects upon T. colubriformis numbers were not significant. The second experiment involved 48 Romney lambs grazing conventional pasture which were infected with 10,000 each of O. circumcincta and T. colubriformis larvae either 7 days before, 7 days after, or at the time they commenced grazing either sulla or lucerne. Lambs which grazed sulla had lower (P<0.05) FEC and lower (P<0.05) O. circumcincta burdens than lambs which grazed lucerne but timing of infection had no effect on FEC or worm burdens. T. colubriformis numbers were not affected by treatment or herbage. In conclusion, the sulla diet resulted in lower O. circumcincta numbers compared to lucerne outdoors and some evidence of an immunogenic response was obtained indoors. However, neither the herbage nor the immunogenic response reduced T. colubriformis numbers in either experiment.     

Effect of forage legumes and anthelmintic treatment on the performance, nutritional status and nematode parasites of grazing lambs

Recent studies in New Zealand and the UK have shown that certain forages reduce parasitic infection in sheep. The aim of this experiment was to investigate the effects of legume forages compared to ryegrass on interactions between production, nutritional status and nematodes in grazing lambs. Twenty-four male lambs per forage treatment, half of which were treated with anthelmintics on Day 0, grazed monocultures of lucerne (Medicago sativa), red clover (Trifolium pratense) and white clover (Trifolium repens) and were compared with lambs grazing perennial ryegrass (Lolium perenne). Individual faecal egg counts (FEC) and liveweight were determined every 7 days for 56 days, after which half the lambs were slaughtered to determine total nematode intensities (TNI). Results showed that lambs grazed on red or white clover, but not lucerne, had lower pooled mean FEC and improved liveweight performance compared to lambs grazing ryegrass. Lambs treated with anthelmintics had higher TNI compared to lambs not treated, due to a trend for more adult nematodes in lambs grazing lucerne and treated with anthelmintics than all other lambs, except those grazing red clover and also given anthelmintics. Lambs grazing white clover tended to have fewer adult nematodes than lambs grazing other forages. Examination of the nematode species showed a change in female T. circumcincta occurred in all lambs following anthelmintic treatment and that the forage species grazed by lambs affected individual parasite species. Lambs grazing white clover had fewer male and adult T. circumcincta compared to lambs grazing other forages, and lambs grazing lucerne had fewer adult T. circumcincta compared to lambs grazing ryegrass or red clover. Data on small intestine TNI showed that lambs grazing lucerne and given anthelmintics had more male adult nematodes than other lambs, except those grazing red clover and treated with anthelmintics. Results indicate that lucerne and red clover both increase the re-infection of grazing lambs with Trichostrongylus species compared to ryegrass following anthelmintic treatment. In conclusion, legume forages have the potential to contribute to the control of abomasal but not small intestine nematode parasites in finishing lamb systems.     

Experimental infection of Black Belly and INRA 401 straight and crossbred sheep with trichostrongyle nematode parasites

Compared to INRA 401 lambs reared in France, Black Belly (BB) lambs reared in Guadeloupe (F.W.I.) were highly resistant to both primary and secondary experimental infection with Haemonchus contortus. To investigate this huge inter-breed difference, a nucleus flock of BB was constituted, and experiments were conducted to: (i) confirm this difference in lambs born in France, (ii) check whether it was similar for Trichostrongylus colubriformis and Teladorsagia circumcincta, and (iii) find out whether this difference was age-related. Forty BB lambs, 84 F1 lambs (BB siresxINRA 401 ewes) and 88 INRA 401 lambs born in two cohorts were used in an experimental design involving three host breeds, both genders and two age-groups (3.5- and 7-month-old when first infected). The limited availability of BB lambs made the study incomplete. Infection consisted of the administration of two doses of 10,000 infective larvae of one of the nematode species, separated by an anthelmintic treatment and an interval of 1 week before the second dose was administered. Fecal egg counts (FECs) were done on Days 28 and 35 after each infection; ewe lambs of the INRA 401 and F1 breeds were necropsied, the worm burden was established, the length of the female worms measured and the eggs in utero counted. For H. contortus and T. colubriformis, the FEC was lower in the BB than in the INRA 401 lambs, and the FEC found for the F1 lambs was intermediate. For T. circumcincta, only data for the F1 and INRA 401 lambs were available, and the FECs for these breeds were the same. In all three breeds, the FEC determined after the second dose was significantly lower than that found after the first dose in the ewe lambs, but not in the ram lambs. Infecting the lambs when they were 7- instead of 3.5-month-old significantly lowered egg excretion of both parasites in the F1 (P<0.0001), but not the INRA 401 lambs. Worm numbers in the F1 and in the INRA 401 ewe lambs confirmed the FEC data, many F1 lambs being free of T. colubriformis. The H. contortus female worms were shorter and had fewer eggs in utero in the F1 than in the INRA 401 lambs. A higher proportion of T. circumcincta was at the fourth larval stage in the F1 lambs. In conclusion, the BB breed is much more resistant than INRA 401 to H. contortus, with the F1 lambs being more like the BBs after the second dose. This difference was also found for T. colubriformis and, to a lesser extent, for T. circumcincta.     

Reduced Ostertagia circumcincta burdens in milk-fed lambs

AIMS: To compare the susceptibility to parasitism by Ostertagia circumcincta of lambs fed entirely with bovine milk or weaned on to solid feed at 3 weeks of age. In addition, the effect of a single daily feed of milk on worm burdens was assessed. METHODS: Eight lambs were assigned to each of the 3 diets: milk (M), milk plus solid feed (cereal-based pellets and lucerne chaff) (MS), or solid feed only (S). Those to be fed solid feed were converted from complete milk feeding to the designated diet during their third week of life. From 3 weeks of age, all lambs were infected with 1000 O. circumcincta larvae twice weekly for 6 weeks; 4 lambs from each diet group were given normal sheathed L3 and another 4 were infected with exsheathed larvae. Faecal egg counts (FEC) and serum gastrin and pepsinogen concentrations were monitored from Day 17 after first infection, and worm burdens and abomasal pH and morphology were determined at necropsy. RESULTS: Total worm burdens and FEC were significantly lower in the M than MS and S groups, whereas there was no significant difference between those receiving sheathed and exsheathed larvae. The milk-fed lambs had a smaller reticulo-rumen and omasum and a more acidic abomasal pH. Serum gastrin and pepsinogen were increased in all groups, irrespective of diet or type of larvae used for infection. CONCLUSIONS: The cause of the lower worm establishment in lambs fed only milk was probably not failure to exsheath in the immature gastro-intestinal tract, as there were similar worm burdens in lambs whether sheathed or exsheathed larvae were administered. The lower pH of the abomasal contents of the preruminant lambs may have been a factor, as the parasites have previously been shown to die more rapidly in vitro at low pH. Alternatively, the milk itself had adverse effects on the parasites, but was ineffective when combined with solid feed. There was no benefit from feeding a milk plus solid diet over a solid diet.     

Regulation of Teladorsagia circumcincta and Trichostrongylus colubriformis worm populations by grazing sheep with differing resistance status

In an experiment lasting 4 years, changes in the Teladorsagia circumcincta and Trichostrongylus colubriformis populations were compared in lambs and adult sheep with differing resistance statuses. Two flocks of 30 rams (resistant R and susceptible S) grazed separate pastures and 8 rams were slaughtered in the middle and at the end of each grazing season. Five groups of tracer lambs were added each year to estimate the pasture infectivity and were killed for worm counts. The availability of animals with differing resistance statuses (rams and tracer lambs) and differing levels of infection made it possible to investigate the number, size and fecundity of worms of these two species. The inflammatory response was measured in the rams by counting the globule leukocytes, mast cells and eosinophils in the fundic, pyloric and intestinal mucosa. In the tracer lambs, the daily egg production by the female worms of both species was negatively correlated with the worm burden. Worm length accounted for 60 and 70% of the variation in the number of eggs in utero for T. circumcincta and T. colubriformis respectively. Worm length was closely associated with the resistance status of the host; there were greater differences between lambs, and S and R rams for T. colubriformis. T. circumcincta worm lengths were not affected by the worm number. Globule leukocyte counts were related to the worm burdens, and mast cell counts to worm length in the R and S rams. The number, size and fecundity of the worms may well be regulated by similar mechanisms in both species, but T. colubriformis seemed to be more intensively regulated than T. circumcincta. This finding could be useful in devising more effective methods of parasite control.     

Brave or gullible: testing the concept that leaving susceptible parasites in refugia will slow the development of anthelmintic resistance

AIM: To test the theory that creating a reservoir of unselected worms by leaving a proportion of lambs in a flock untreated with anthelmintic, i.e. in refugia, will slow the development of anthelmintic resistance in nematode parasite populations. METHODS: Newly weaned Romney lambs (n=180) were infected with two nematode parasite species, Teladorsagia (= Ostertagia) circumcincta and Trichostrongylus colubriformis. For each species, the challenge doses contained a mixture of infective larvae from benzimidazole-resistant and -susceptible isolates calculated to yield, from the combined population, a 95% reduction in faecal nematode egg counts (FEC) following treatment with albendazole. Once the infections were patent, the lambs were divided into nine groups of 20 animals, and each group was allocated to one of three treatments. In Treatments 1, 2 and 3, 100%, 90% and 80% of animals were treated with an anthelmintic, respectively. For treatments 2 and 3, the heaviest animals remained untreated. Following treatment, each group was moved to its own previously prepared low-contamination pasture. Lambs grazed this pasture for 7 weeks before again being treated and moved to new low-contamination pastures (Shift 1 and Shift 2). The parasite populations on pasture resulting from the different treatments were subsequently sampled using tracer lambs, and worm eggs derived from these were used in both egg-hatch assays (EHA) and larval development assays (LDA), to measure albendazole-resistance status. RESULTS: Treating all animals each time the groups were moved to new low-contamination pastures resulted in higher levels of albendazole resistance (p<0.05), measured using EHA and LDA, in subsequent parasite generations than when either 10 or 20% of animals were left untreated. However, higher FEC in the tracer lambs grazed on pastures in Treatments 2 and 3, compared with Treatment 1, indicated an increased level of pasture contamination as a result of leaving some animals untreated. CONCLUSIONS: The results demonstrate that creating a reservoir of unselected parasites slows the development of anthelmintic resistance, and emphasises the risk of treating all animals prior to a shift on to low-contamination pasture. However, higher levels of pasture contamination, resulting from untreated animals, indicate the difficulty in managing both worm control and resistance.     

Moxidectin: persistence and efficacy against drug-resistant Ostertagia circumcincta

In order to determine whether the efficacy of moxidectin against Ostertagia circumcincta is enhanced by its persistency, therapeutic efficacy was compared at intervals after treatment and with that of ivermectin, a closely related but more transient endectocide. Groups of 7-month-old New Zealand Romney lambs were infected with a strain of O. circumcincta known to be resistant to moxidectin. At patency of the infections, groups of lambs were treated with either moxidectin or ivermectin at the manufacturer's recommended dosages, or left untreated. At 3, 6 and 10 days post-treatment, faecal egg count was measured and groups of lambs were slaughtered for estimation of adult worm burden. Drug-resistant worm burdens were significantly reduced in those animals treated with moxidectin but not in those treated with ivermectin. No effect of time of slaughter on worm burden was observed with either drug, demonstrating that the higher therapeutic efficacy of moxidectin against this parasite was not due to an increased period of drug exposure. Faecal egg counts in the moxidectin treated animals increased with time after treatment indicating a temporary suppression of egg output by surviving worms. The implications of these findings on selection for anthelmintic resistance are discussed.     

Multiple species of nematodes resistant to ivermectin and a benzimidazole-levamisole combination on a sheep farm in New Zealand

AIM: To confirm the presence of multiple anthelmintic resistance on a sheep farm in New Zealand. METHODS: Three groups of 10 weaned Romney-cross lambs were treated either with an oral dose of ivermectin (0.2 mg/kg), or a benzimidazole/levamisole (BZ/LEV) combination (4.75 albendazole and 7.5 mg/kg levamisole), or were left untreated. Ten days later, animals were necropsied, and adult worms recovered and identified from the abomasa and small intestines. Pre- and post-treatment faecal nematode egg counts (FEC) were recorded, and larval cultures were performed. RESULTS: In a faecal egg count reduction test (FECRT), adjusted to reflect pre- and post-treatment larval culture results, ivermectin resistance was detected in Teladorsagia (Ostertagia), Trichostrongylus and Haemonchus spp, while BZ/LEV combination- resistant Teladorsagia and Trichostrongylus spp were also present. Adult worm counts confirmed these results, and identified the species involved as Teladorsagia circumcincta, Trichostrongylus colubriformis and H. contortus. CONCLUSION: Multiple, multi-generic anthelmintic resistance was confirmed on a sheep property in New Zealand. This included the first confirmed case of ivermectin resistance in T. colubriformis from sheep in New Zealand.     

The effect of diet fed to lambs on subsequent development of Trichostrongylus colubriformis larvae in vitro and on pasture

Contrasting herbage diets were fed to lambs to evaluate their effect on subsequent development of Trichostrongylus colubriformis larvae in faeces and on pasture. The diets had either no condensed tannin (CT), lucerne (Medicago sativa cv. Otaio), white clover (Trifolium repens cv. Tahora), or had moderate to high concentrations of CT, sulla (Hedysarum coronarium cv. Grassland Aokau), Lotus corniculatus (cv. Grasslands Goldie), L. pedunculatus (cv. Grassland Maku), Dorycnium pentophyllum, and Dorycnium rectum. Trials were carried out in summer (warm) and in autumn (cool and moist). In summer, egg viability was evaluated in vitro with egg hatch and larval development assays. In both seasons faeces were placed on pasture to compare recovery of eggs and larvae from faeces and larvae from herbage on the high and low fertility farmlets on the AgResearch Ballantrae Hill Country Research Station. D. rectum and D. pentophyllum diets decreased (P<0.01) egg hatching and larval development in laboratory assays relative to other diets. In summer, the number of larvae recovered from faeces placed on pasture was far greater (P<0.001) if the lambs had been fed lucerne than any other diet, whereas recovery was always lowest from faeces of sheep fed D. rectum and D. pentophyllum. Although dietary differences were lower in autumn than in summer, larval recoveries were lower (P<0.05) from faeces of lambs fed D. rectum and L. corniculatus than from white clover, lucerne and sulla diets. This study indicates that the diet of the host can have a significant impact on egg hatching and the subsequent development of T. colubriformis larvae in the laboratory and in the field. In particular, D. rectum consistently reduced T. colubriformis development. Effects measured in vitro generally under-estimated effects measured under field conditions.     

The effect of birdsfoot trefoil (Lotus corniculatus) and chicory (Cichorium intybus) on parasite intensities and performance of lambs naturally infected with helminth parasites

Conventionally, farmers rely upon the routine use of anthelmintics to control helminth parasites and their use has proved highly cost-effective. However, several factors, including the emergence of helminths resistant to pharmaceutical anthelmintics, are forcing farmers to seek alternative approaches to parasite control. Studies in New Zealand have shown that some alternative forages may reduce parasitic infestation in sheep. In the current study, it was found that under UK environmental conditions lambs with naturally acquired helminth infections grazing chicory (Cichorium intybus) and birdsfoot trefoil (Lotus corniculatus) had fewer helminth parasites than sheep grazing ryegrass/white clover (Lolium perenne/Trifolium repens). Twelve pure-bred Lleyn male lambs grazed replicated 0.5ha plots of birdsfoot trefoil, chicory or ryegrass/white clover for 5 weeks. Liveweight and faecal egg counts (FECs) were determined weekly and eight lambs per forage were slaughtered at the end of the trial to determine total helminth intensities. Lambs grazing birdsfoot trefoil had a lower FEC on day 7 (P<0.05) and fewer total adult helminths than those grazing the other forages on day 35 (P<0.01). Lambs grazing chicory did not have significantly lower FEC than lambs grazing other forages but these lambs were found to have fewer total adult abomasal helminths than lambs grazing ryegrass/white clover (P<0.001). As the performance of grazing lambs is inversely correlated with the intensity of helminth parasites, these alternative forages could be used to improve the liveweight gain of lambs produced in the UK. Overall, the results support the contention that alternative forages could have a positive role in the control of helminth parasites in sheep, subject to successful agronomic development and integration of these forages into whole farm systems.     

The effect of continuous drug exposure on the immune response of lambs challenged with drug-susceptible or drug-resistant nematode larvae

Groups of lambs either with or without controlled-release albendazole (ABZ) capsules (CRCs) were challenged twice weekly for 6 weeks with either drug-susceptible or drug-resistant Ostertagia circumcincta and Trichostrongylus colubriformis. Groups with and without CRCs remained unchallenged as controls. There was minimal establishment of drug-susceptible parasites of either species in those lambs with CRCs. However, drug-resistant parasites of both species established adult worm burdens in the presence of the capsules. The humoral immune response, as measured by the serum anti-larval (L₃) antibody (Ab) titre, was pooled for weeks 4–6 and compared for each group. With the exception of anti-T. colubriformis Ab in group 2, anti-L₃ Ab titres were significantly higher in all the parasite-challenged groups as compared to the control animals. Also, with the exception of anti-resistant O. circumcincta Ab levels in the CRC-treated animals (group 5), no significant difference was observed in Ab titres between the four groups challenged with either resistant or susceptible larvae. The results demonstrate the inability of CRCs to prevent establishment of drug-resistant parasites and that immune stimulation is not inhibited by the capsules.     

Evolution of nematode community in grazing sheep selected for resistance and susceptibility to Teladorsagia circumcincta and Trichostrongylus colubriformis: a 4-year experiment

Field trials were undertaken to compare nematode population dynamics in two experimental flocks of rams selected for high and for low faecal egg counts (FECs) following two doses with 20000 infective larvae, 12000 Teladorsagia circumcincta (Tcirc) and 8000 Trichostrongylus colubriformis (Tcol) separated by a treatment. The selection was done by dosing 200 ram lambs (INRA 401 breed) and FECs proceeded with egg identification on days 28 and 35 post-each-infection. The 30 lambs with the lowest FECs constituted the group of permanent resistant rams (R), the 30 with the highest values the group of susceptible ones (S). Each group grazed separated pastures from April to November, this during 4 consecutive years. The contamination (number of eggs deposited) was estimated every 3 weeks on each pasture. The larval populations was measured by successive groups of tracer lambs. The infection of the permanent rams was done by slaughtering eight R and S rams on mid and end of each grazing season. Each autumn, new R and S ram lambs were selected to complete the R and S groups before turning out in spring. On third year, a third species (Haemonchus contortus, Hcon) was added to compare its behaviour with the two others.The regulation of parasite populations were studied by comparing what happens in R rams and pasture to S ones. The selected R ram lambs had a FEC of Tcirc 50% lower than S ones. This induced a similar lowered pasture contamination, a 25% decrease of infective larval population in years 2 and 3, and the worm burdens in R rams were the half of that in S rams after 2 years. The FEC of Tcol was 75% lower at the selection and induced similar difference in pasture contamination, but 65 and 96% reduction in L3 population, respectively, after 1 and 2 years. The worm burden was reduced by 99%. Adding Hcon species during the third year, in year 4, the L3 population was half in R pasture compared with S one, and R rams had 60% less worms of this species than S ones.This selection without any drench prevent outbreaks observed in young tracer lambs on S pasture. The comparison of selected resistant rams to selected susceptible ones and not to unselected controls gave more rapid information about the tendency of the regulation of the communities of nematode parasites.     

Influence of selenium status in merino weaners on resistance to trichostrongylid infection

Weaned merino lambs, grazing pastures low in selenium, were used to investigate the effect of selenium status on immunity to trichostrongylids. Six weeks following selenium supplementation to 14 of the 27 sheep using intraruminal selenium pellets, 5000 Ostertagia circumcincta and 5000 Trichostrongylus colubriformis larvae were administered orally to all sheep. At four weeks after infection, the mean total worm burden in the selenium supplemented sheep (5537 +/- 343, n = 14) was not significantly different (P greater than 0.05) from that in the unsupplemented sheep (5614 +/- 374, n = 12) and faecal worm egg concentrations were also similar in the two treatment groups. At this time, mean red cell glutathione peroxidase activities in the supplemented and unsupplemented groups were 430 and 11 U g-1 haemoglobin, respectively, and clinical white muscle disease had been observed in the latter group. These results suggest that increasing selenium status of selenium deficient sheep by the use of intraruminal selenium supplementation, has a negligible effect on resistance to an artificial challenge infection of O circumcincta and T colubriformis.     

Isolation of benzimidazole resistant strains of Ostertagia circumcincta from British sheep

Two strains of Ostertagia circumcincta were isolated from sheep in Great Britain; one (CVL strain) from a breeding flock maintained at the Central Veterinary Laboratory, the other (H2 strain) from a commercial flock in southern England. Their resistance to benzimidazole anthelmintics was assessed by means of in vitro egg hatch assays and slaughter trials. In vitro egg hatch assays gave calculated ED50 estimates of 0.799 micrograms thiabendazole/ml for the CVL strain and 0.794 micrograms thiabendazole/ml for the H2 strain, compared with ED50 estimates of 0.038 micrograms thiabendazole/m and 0.036 micrograms thiabendazole/ml for two known susceptible strains of O circumcincta. There was a 40.7, 28.4 and 66.9 per cent reduction in the group mean worm burdens of lambs infected with the CVL strain following treatment with thiabendazole, fenbendazole and oxfendazole, respectively, and 23.8, 0.0, 79.6, 52.7, 99.9 and 100 per cent reduction in the group mean worm burdens of lambs infected with the H2 strain following treatment with thiabendazole, fenbendazole, oxfendazole, albendazole, levamisole and ivermectin, respectively. Detailed field histories for both strains are given.     

Comparative susceptibility of Romanov and Lacaune sheep to gastrointestinal nematodes and small lungworms

The susceptibility of Romanov and Lacaune sheep to nematode parasites was studied in a permanent flock grazing an extensive pasture. The output of strongyle eggs and protostrongyle larvae was higher in Romanov ewes. Faecal samples of the Romanov ewes more often harboured Nematodirus eggs while the larvae recovered from cultures of these samples contained a higher percentage of Teladorsagia. Romanov tracer lambs had a significantly heavier adult Teladorsagia burden. This higher natural susceptibility of the Romanov breed was confirmed by experimental infections with T. circumcincta and N. spathiger but not with Trichostrongylus colubriformis. In infections with Muellerius capillaris and Protostrongylus rufescens, the worm burdens were similar but parasite fertility (larvae per adult) seemed to be higher in Romanov tracer lambs.     

Production and immunological responses associated with controlled-release-capsule vs 5-drench preventive anthelmintic programmes for parasite control in lambs

AIMS: To determine whether: a) using a controlled-release anthelmintic capsule (CRC) instead of a programme of 5 oral drenches administered at 3-4 week intervals, would delay the development of anti-parasite immunity in lambs; b) the use of ivermectin instead of albendazole, administered either as a CRC or as a programme of 5 oral drenches, would delay the development of anti-parasite immunity in lambs; c) lambs treated with CRCs would have higher liveweight gains than lambs drenched orally 5 times at 3-4 week intervals, and; d) delayed onset of anti-parasite immunity is associated with reduced liveweight gains in the period following anthelmintic treatment. METHODS: Three field trials were conducted, 1 on a research farm and 2 on commercial sheep farms, in which groups of 30 lambs were treated with either a CRC containing albendazole, a CRC containing ivermectin, 5 oral drenches with albendazole, or 5 oral drenches with ivermectin, administered at 3-4 week intervals. Liveweights and faecal nematode egg counts (FECs) were recorded in all trials. Immunoglobulin-G (IgG) antibody levels to Ostertagia circumcincta and Trichostrongylus colubriformis adult and larval antigens were measured in Trials 1 and 3, and fleece weights and resistance of animals to nematode challenge infection were measured in Trial 1. RESULTS: CRC-treated lambs had higher levels of antibodies to O. circumcincta infective-stage larvae (L3) than orally drenched lambs in Trial 3, but no other immunological differences due to mechanism of delivery were detected. Antibody levels were lower in lambs treated with ivermectin than albendazole, as a CRC or oral drench in Trial 1, but this was not associated with any measurable effects on FEC or productivity. No significant differences (p>0.05) were detected between drench types (albendazole vs ivermectin) or delivery mechanisms (CRC vs oral drenching) in any of the production parameters measured, in any of the trials. Albendazole-CRCs failed to control FECs in all 3 trials. CONCLUSIONS: Although some differences between treatments in antibody levels were detected these were not associated with measurable differences in level of parasitism or productivity of lambs. CRC use did not appear to offer substantial gains in productivity over a structured programme of 5 oral drenches administered at 3-4 week intervals.     

Managing anthelmintic resistance--use of a combination anthelmintic and leaving some lambs untreated to slow the development of resistance to ivermectin

A field study was conducted to test the hypotheses that use of a combination anthelmintic and/or increasing the pool of unselected worms 'in refugia' by leaving a proportion of lambs untreated would slow the development of resistance to ivermectin. Twelve suites of four paddocks (farmlets) were seeded with a mixture of resistant and susceptible isolates of both Trichostrongylus colubriformis and Teladorsagia (Ostertagia) circumcincta calculated to yield a 95% reduction in faecal nematode egg count (FEC) after treatment with ivermectin. Each year for three years the farmlets were stocked in the spring with mobs of lambs which were treated five times at 28-day intervals with either ivermectin or an ivermectin+levamisole combination. In addition, in half the mobs the heaviest 10% of lambs remained untreated at each treatment occasion, resulting in a 2 × 2 factorial treatment structure (i.e. two drench types × two percentage treated) with three complete replicates. The development of resistance to ivermectin, and at the end to levamisole, was measured by larval development assays (LDA) and worm counts from treated and untreated tracer lambs. For T. colubriformis the development of resistance to ivermectin, as measured by tracer lamb worm burdens, was delayed by treatment with the combination and by leaving 10% of lambs untreated. In addition, the interaction between these factors approached significance (p=0.052). Similarly, results of the LDAs indicated a slower development of resistance when lambs were treated with the combination and when 10% of lambs were left untreated. For T. circumcincta, results were compromised by the rapid development of resistance, which appears to be the result of low viability in the field of the susceptible isolate used to contaminate the pastures. Although a small delay in the development of resistance to ivermectin was indicated, this was off-set by an increase in the level of resistance to levamisole. A post-study modelling experiment simulating the conditions of the field study and the starting efficacies for the two nematode species produced equivalent outputs to those measured in the field. Overall, results support the conclusions that use of combination anthelmintics and deliberately increasing 'refugia' of unselected genotypes will slow the development of anthelmintic resistance. However, as indicated in modelling studies, once resistance to all the constituent actives is well developed (efficacy<70%) the value of combinations for slowing the development of resistance is largely lost.     

In silico exploration of the impact of pasture larvae contamination and anthelmintic treatment on genetic parameter estimates for parasite resistance in grazing sheep

A mathematical model was developed to investigate the impact of level of Teladorsagia circumcincta larval pasture contamination and anthelmintic treatment on genetic parameter estimates for performance and resistance to parasites in sheep. Currently great variability is seen for published correlations between performance and resistance, with estimates appearing to vary with production environment. The model accounted for host genotype and parasitism in a population of lambs, incorporating heritable between-lamb variation in host-parasite interactions, with genetic independence of input growth and immunological variables. An epidemiological module was linked to the host-parasite interaction module via food intake (FI) to create a grazing scenario. The model was run for a population of lambs growing from 2 mo of age, grazing on pasture initially contaminated with 0, 1,000, 3,000, or 5,000 larvae/kg DM, and given either no anthelmintic treatment or drenched at 30-d intervals. The mean population values for FI and empty BW (EBW) decreased with increasing levels of initial larval contamination (IL(0)), with non-drenched lambs having a greater reduction than drenched ones. For non-drenched lambs the maximum mean population values for worm burden (WB) and fecal egg count (FEC) increased and occurred earlier for increasing IL(0), with values being similar for all IL(0) at the end of the simulation. Drenching was predicted to suppress WB and FEC, and cause reduced pasture contamination. The heritability of EBW for non-drenched lambs was predicted to be initially high (0.55) and decreased over time with increasing IL(0), whereas drenched lambs remained high throughout. The heritability of WB and FEC for all lambs was initially low (～0.05) and increased with time to ～0.25, with increasing IL(0) leading to this value being reached at faster rates. The genetic correlation between EBW and FEC was initially ～-0.3. As time progressed the correlation tended towards 0, before becoming negative by the end of the simulation for non-drenched lambs, with increasing IL(0) leading to increasingly negative correlations. For drenched lambs, the correlation remained close to 0. This study highlights the impact of IL(0) and anthelmintic treatment on genetic parameters for resistance. Along with factors affecting performance penalties due to parasitism and time of reporting, the results give plausible causes for variation in genetic parameter estimates previously reported.     

Efficacy of the nematode-trapping fungus Duddingtonia flagrans against three species of gastro-intestinal nematodes in laboratory faecal cultures from sheep and goats

The ability of the nematode-killing fungus Duddingtonia flagrans to reduce number of infective larvae of three species of gastro-intestinal parasitic nematodes developing in dung was investigated in both goats and sheep. Groups of lambs and kids (12-20 weeks old) were given mono-specific infections of Haemonchus contortus, Ostertagia (Teladorsagia) circumcincta or Trichostrongylus colubriformis. Following patency of the infections (t1) faecal samples were collected for determination of faecal nematode egg count (FEC) and culture of parasite larvae. Groups of animals were then dosed on 2 consecutive days with one of the two dose rates of the fungus (250,000 or 500,000 spores/kg liveweight). One (t2) and 5 (t3) days after the second dose of fungus samples were again collected for FEC and culture. The number of larvae recovered from the faecal cultures at t1 and t3 were used as controls to assess the efficacy of the experimental treatment at t2. Average efficacy was 78% with group means ranging from 40 to 93%. Dose rate of fungus appeared to influence efficacy against O. circumcincta but not against H. contortus or T. colubriformis. Overall, there were no differences in the efficacy of the fungus against any of the parasite species or in either host animal. The results of this trial indicate the potential use of this fungus as a broad spectrum anti-parasite agent for use in both goats and sheep.     

Stocking rate effects on the worm burdens of Angora goats and Merino sheep

Worm egg counts of yearling Angora or Merino wethers grazing in mixed flocks at either 4/ha or 8/ha were observed over a 5 month period. Based on faecal egg counts infection levels in the sheep increased for the first 3 months and then decreased as the animals developed resistance. Egg counts in the goats did not decrease during the experiment. Post mortem worm counts indicated that there was a significant host species--stocking rate interaction on worm burdens of Haemonchus contortus adults and Ostertagia circumcincta adults and larvae. Goats had more H. contortus and O. circumcincta adults than sheep at the 8/ha stocking rate. At 4/ha goats had fewer O. circumcincta fourth stage larvae than sheep. There was no stocking rate effect on the numbers of Trichostrongylus axei or T. colubriformis. Sheep had more T. axei and fewer T. colubriformis than goats.