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Toxoplasma gondii is a parasite that has been extensively studied due to its medical and veterinary importance in terminating pregnancies. Consequently, a satisfactory vaccine is required to control its adverse effects on pregnant animals. The microneme protein, MIC3, is a major adhesion protein that binds to the surface of host cells and parasites, and is therefore a potential vaccine against T. gondii. The viability of MIC3 as a vaccine is investigated in this study. Sheep were injected twice, intramuscularly, with plasmids containing DNA encoding for the mature form of MIC3 protein formulated into liposomes. Control sheep were injected with an empty vector or received no injections. The injection of sheep with DNA plasmids encoding for MIC3 elicited an immune response after the first and second injections as indicated by antibody responses and the production of IFN-gamma. The immune response, as measured by the IgG2 and IgG1 serum levels, was boosted after the injection of the MIC3 DNA vaccine together with high anti-MIC3 antibodies. The results demonstrate that the intramuscular injection of sheep with a plasmid containing DNA coding for MIC3 protein induces a significant and effective immune response against T. gondii.

CD154 is a cell surface molecule expressed by activated T cells. CD40 and CD154 interaction is critically important in regulating humoral and cell-mediated immune responses. In this study we have investigated whether a DNA vaccine encoding rhoptry protein 1 (ROP1) of Toxoplasma gondii, and encoding ovine CD154 induces an enhanced ROP1-specific immune response in sheep. Two groups of twelve animals received two intramuscular injections, of a DNA plasmid encoding T. gondii ROP1 antigen (group 1) or an ROP1 antigen fused to ovine CD154 (group 2). There were two control groups of sheep. One was injected with an empty vector (group 3) and the other received no injections at all (group 4). The injection of the plasmid containing ROP1 (group 1) at weeks 0 and 4 induced a significant IgG2 response at week 2 which was amplified at week 4 after the booster injection and persisted to week 8 compared to the control animals in groups 3 and 4. For IgG1, significant differences from the control animals were only observed from week 5 onwards. The fusion of CD154 and ROP1 elicited significant IgG1 and IgG2 responses from week 1 which were amplified from weeks 5 to 8 compared to the control animals in groups 3 and 4. The IgG1 response was significantly higher in group 2 animals receiving pROP1-CD154 compared to group 1 receiving pROP1 only. There was no significant difference in IgG2 responses between groups 1 and 2. Significant differences in IFN-γ levels were only observed in treatment group 1 at week 2 and treatment group 2 at weeks 1 and 2 compared to the control animals. The results demonstrated that an intramuscular injection of pROP1-CD154 gene to sheep significantly enhanced their immune response and induced a mixed Th1/Th2 response while the intramuscular injection of pROP1 only induced a Th1-specific immune response., (Copyright © 2011 Elsevier B.V. All rights reserved.)

The dense granule proteins of Toxoplasma gondii are investigated as possible vaccine candidates against the parasite. The aim of this research was to evaluate the immune responses of sheep injected twice, intramuscularly, with DNA plasmids encoding T. gondii dense granule antigens GRA1, GRA4, GRA6 and GRA7 formulated into liposomes. Control sheep were injected with an empty vector or received no injections. The injection of sheep with DNA plasmids encoding for GRA1, GRA4, GRA6 or GRA7 elicited an immune response after the first and the second injections as indicated by the moderate to high antibody responses. The injection of pGRA7 induced a significant level of anti-GRA7 IgG2 antibody and IFN-γ responses indicating a Th1-like immune response whereas injection with pGRA1, pGRA4 and pGRA6 stimulated a IgG1 type antibody response with a limited, if any, IFN-γ response. The results demonstrate that the intramuscular injection of sheep with a DNA liposome formulated plasmid coding for GRA proteins is an effective system that induces a significant immune response against T. gondii., (Copyright © 2010 Elsevier B.V. All rights reserved.)

The aim of this study is to compare the immune responses of sheep stimulated by the intramuscular injection of a liposome formulated-DNA plasmid encoding the Toxoplasma gondii MAG1 antigen only or co-expressed with ovine IL-6. Forty-five, 2-year-old sheep were divided into four groups. Group 1 received an empty pVAXIg plasmid, group 2 no treatment, group 3 liposome formulated plasmid pVAXIgMAG1 and group 4, pVAXIgMAG1 plus pVAXovIL-6 plasmids. All the animals were inoculated at weeks 0 and 4. The injection of sheep with a plasmid encoding for MAG1 only or a MAG1 plasmid co-expressed with a plasmid encoding for ovine IL-6 produced humoral immune responses. The plasmids containing MAG1 elevated significantly serum IgG1 and IgG2 levels 2 weeks and onwards after the first injection of the plasmids. Co-expression of IL-6 with MAG1 had no effect on IG1 or IG2 levels illustrating that IL-6 in the formulation used had no modulating effect on any measured immune response., (Copyright 2010 Elsevier B.V. All rights reserved.)

The hypothesis that increases in the concentration of the anorectic peptide leptin may be responsible for the immune-mediated reduction in feed intake (FI) during gastrointestinal parasitism in sheep was investigated. In a 2 x 2 x 2 factorial design, the first factor was age at the start of infection (5 months old v. 17 months old). The second factor was parasite infection (no infection v. administration of eighty L3 infective Trichostrongylus colubriformis larvae/kg live weight (LW) per d three times per week for 77 d). The third factor was immunosuppressive therapy with a corticosteroid (no therapy or weekly intramuscular injection of 40 mg methylprednisolone acetate/30 kg LW). Relative to their uninfected counterparts, a 20 % reduction in FI per unit LW (FI/LW; g DM/kg LW) was observed in infected non-suppressed 5-month-old lambs from 21 to 63 d post-infection (P < 0.001) but not in comparable17-month-old ewes or in corticosteroid-treated lambs or ewes (P>0.05 for all), allowing the suggestion that the anorexia was a consequence of the developing immune response. The reduction in FI/LW in 5-month-old lambs was not associated with an increase in plasma leptin concentration. Furthermore, plasma leptin concentrations were greater in corticosteroid-treated animals (P < 0.001) and in 17-month-old animals (P < 0.001), none of which displayed an infection-induced reduction in FI/LW. Plasma leptin was positively correlated with carcass fat percentage in both 5-month-old (P = 0.016) and 17-month-old (P < 0.001) animals and did not appear to provide a direct feedback mechanism that restricted energy intake. The results do not support the hypothesis that an increase in circulating leptin is directly responsible for the immune-mediated anorexia in lambs during T. colubriformis infection.

The nutritional cost of, and the sequential cellular changes associated with the developing immune response to the abomasal parasite Teladorsagia circumcincta were investigated using corticosteroid-induced immune-suppression. Six-month-old lambs with minimal nematode experience were either infected with 4000 L3 T. circumcincta per day (group IF), similarly infected and concurrently immune-suppressed with methylprednisolone acetate (group ISIF), immune-suppressed only (group IS) or remained as controls (group C). Food intake, faecal egg count (FEC) and antibody titres in plasma were recorded weekly, worm burden at necropsy on day 63 p.i. and body composition by X-ray computed tomography on days -2 and 62 p.i. Furthermore, sequential immunological changes at the site of parasite infestation in the abomasal mucosa were measured from serial biopsy tissue samples taken from additional animals that were fitted with an abomasal cannula and either infected with the same regime as IF animals above (group CnIF) or concurrently infected and immune-suppressed as above (group CnISIF). Corticosteroid treatment resulted in greater FECs (P<0.01) and worm burdens (P<0.01) in both ISIF and CnISIF compared with IF and CnIF sheep, respectively. Infection reduced feed intake by 17% between 14 and 28 days p.i. (P<0.05) and efficiency of energy utilisation by 20% (P=0.07) in IF animals but not in ISIF animals. Mast cells, globule leukocytes and IgA in tissue biopsy samples were elevated in CnIF from 42 days p.i., all of which were abrogated by corticosteroid treatment. The ability to regulate the worm population appeared to be associated with a rise in tissue IgA concentration and numbers of globule leucocytes (GL). The results support the hypothesis that a majority of the production losses that occur during infection of lambs with T. circumcincta in lambs are a consequence of the host immune response. These findings may have implications for regimes that promote the development of a strong host immune reaction to gastrointestinal parasites in lambs.

This study tested the hypothesis that milk has a direct effect on the establishment of infection with Teladorsagia circumcincta, and provides information on the effects of suckling on resilience to infection in young lambs. Of 46 six-week-old twin-born lambs, one from each twin was allowed to continue suckling (S-) and its twin-weaned (W-) while both were concurrently infected with an average of either 0 (-0; n=7/group), 250 (-250; n=8/group) or 1000 (-1000; n=8/group) third stage infective larvae (L3) of T. circumcincta per day, providing six treatment groups. All groups grazed minimally contaminated pasture, and after 42 days were slaughtered for necropsy. Low pasture larval contamination was confirmed in W0 and S0 lambs by faecal egg counts (FEC) of less than 30 eggs per gram (EPG) and burdens of less than 140 worms. There was no difference in FEC between weaned and suckled lambs. Within infection regime, total worm burdens were 55-60% greater in the weaned compared with their suckled counterparts (P=0.05), and represented 27 and 17%, respectively, net establishment of larvae. The greater worm burdens of both groups of weaned animals, which compared with their suckled counterparts, and of those infected with 1000 compared with 250 larvae per day, were associated with shorter female adults that had fewer eggs in utero, perhaps indicating intra-worm population regulation, but highlighting the limitation of FEC in assessing nematode burdens of such young lambs. There was no effect of infection on live weight gain of either weaned or suckled groups and the possibility was raised that, in such young lambs, immune unresponsiveness may be responsible. The major benefit of continued milk consumption appears to lie more in providing nutrients for enhanced growth rather than in improving resilience of the lambs to infection.

A technique to take sequential tissue biopsy samples in multiparous, periparturient ewes from the abomasal mucosa is described, developed in parallel in Scotland and New Zealand. Samples were extracted via abomasal cannulae inserted into the wall of the abomasum and exteriorised through dorso-ventral laparotomy. Animals recovered quickly post-surgery, and tolerated the cannula and sampling without any adverse signs of pain or discomfort. The technique was deployed in two pilot studies to investigate the sequential mucosal inflammatory cell responses in well-defined parasitological models, during the periparturient relaxation of immunity in ewes infected with gastrointestinal nematodes and subjected to different feeding treatments. One experiment (Moredun Research Institute, Scotland) involved the infection of twin-bearing ewes with Teladorsagia circumcincta L3 either before, or after lambing. By feeding ewes with different levels of protein supplementation, preliminary data on the impact of nutrition on the eosinophil, mucosal mast cell and globule leucocyte responses during this period were investigated. A similar study was also performed at Lincoln University, New Zealand, to investigate these cell responses in sheep fed relatively high or low protein diets during pregnancy, and infected with a combined immunisation regime of T. circumcincta and Trichostrongylus colubriformis L3. These studies confirmed the phenomenon termed the periparturient relaxation in immunity (PPRI) where a transitory increase in faecal egg counts is observed during late pregnancy and lactation, and this effect was exacerbated during protein undernutrition. Although the number of animals was low in each experiment and the cell responses variable, the results together suggest a reduction in the number of mucosal mast cells and globule leucocyte during the PPRI when protein supply was restricted. The present paper thus describes a successful technique to monitor ovine mucosal cell populations during local immune responses in normal and pregnant sheep. It is envisaged that this technique will be a powerful adjunct to investigations into mucosal immune mechanisms and disease pathogenesis, and will be employed to confirm the influence of dietary protein on the local inflammatory cell responses during the PPRI.

Haemonchus contortus is known as a prolific parasite, producing high numbers of eggs. It could therefore be used as a cheap source of larvae for immunisation of lambs. The value of immunisation would be improved if the immunity produced gave protection against not only homologous but also heterologous infections. Because antibody cross-reactivity between Haemonchus and Ostertagia has been well established, we wanted to know whether drug-abbreviated infections of H. contortus would stimulate production of antibodies that would cross-react with Trichostrongylus colubriformis. The results obtained from these trials indicated that H. contortus drug-abbreviated infections produced significant immunity against not only Haemonchus but also O. circumcincta. Immunisation increased the level of immunoglobulin E (IgE) and IgE-specific antibodies against T. colubriformis, but the differences between experimental and control animals were not statistically significant. Significantly higher levels of IgG-specific antibodies against T. colubriformis were observed.

Proteinases are known to be capable of prolonging the survival of endoparasites in a host. We were therefore interested in knowing whether immunization of lambs against a proteasome (multisubunit proteinases) preparation obtained from Trichostrongylus colubriformis infective third-stage larvae (L3) would have any effect on the immune response to a single challenge infection with the same organism. A total of 21 penned lambs aged 8 months were divided into 3 equal groups. Group 1 was immunized on three occasions with increasing amounts of a proteasome-enriched fraction obtained from infective L3. Group 2 was given a similar amount of protein from the initial supernatant of homogenized larvae. Group 3 (controls) received adjuvant plus saline solution only. All groups were challenged with 60,000 infective T. colubriformis larvae at 28 days after the last immunization. Significant protection was obtained only when the initial supernatant extract was used to immunize lambs. The proteasome preparation seemed to have immunosuppressive effects through the stimulation of nonspecific IgE production. Significantly lower levels of specific IgE were observed in lambs immunized with the proteasome-enriched fraction, and levels of specific IgG antibodies were increased. We suggest that proteasome fractions of T. colubriformis may serve as useful preparations for the study of mechanisms of IgE production in parasitized sheep.

A very high level of protection was achieved against homologous (up to 97%) and heterologous (up to 87%) infections in 12-month-old Romney sheep immunised with oxfendazole-abbreviated infections of Ostertagia circumcincta and Trichostrongylus colubriformis. No significant protection occurred following ivermectin-abbreviated infections. None of the immunised sheep showed an increase in antibody level against excretory-secretory antigen of T. colubriformis infective larvae. The immunisation procedures did not cause a decrease in wool production, or liveweight gains compared with non-immunised controls.

A protective immune response without liveweight loss can be induced in sheep against T. colubriformis but results depend on the anthelmintic used and duration of immunizing infections. More than 90% protection was achieved in sheep immunized by three 15- or 7-day oxfendazole abbreviated infections or three 21-day nonabbreviated infections. Only 41% protection was induced by 3-day oxfendazole abbreviated infections. Significantly higher worm burden and faecal egg counts were present after challenge in sheep immunized by 7-day levamizole abbreviated infections compared to 7-day oxfendazole abbreviated infection. Liveweight gains of sheep immunized by 15- and 7-day abbreviated infections were not significantly different than non infected controls. Liveweight loss seemed to be associated with high activity of mucus peroxidase and high numbers of eosinophils in the intestinal lumen. High parasite numbers seemed to be associated with low activity of alkaline phosphatase in mucus. Mucus peroxidase, arylsulphatase, larval migration inhibition of mucus, mucus or serum antibody against L3 excretory/secretory antigen or somatic L3, L4 and adult antigen were not associated with protection.

Long-term selection of sheep for resistance to parasite infections may be jeopardized if animals do not retain their normal ability to respond to non-parasite antigens. Therefore the antibody responses to ovalbumin (OVA) and human red blood cells (HRBC), and kinetics of peripheral blood lymphocyte phenotypes were examined in mature grazing sheep, genetically resistant or susceptible to gastrointestinal nematodes. In both lines the HRBC antibody response peaked 2 weeks after the primary injection, 1 week after the second injection and 3 weeks after the second OVA injection. The antibody titres of the resistant line sheep decreased sooner after both primary and secondary injections. The resistant line sheep had higher percentages of CD5+ and CD4+ cells than the susceptible sheep. Two injections of OVA and HRBC did not result in significant alterations in percentages of CD5+, CD4+, CD8+ and CD45R+ lymphocytes in either line. In both lines, the control groups showed a steady increase of 0.29% per week in percentages of T19+ (gamma delta) T cells which was significantly higher than in the antigen injected sheep.

Blastogenic activity, eosinophil and total white blood cell counts (TWBC) were examined over a period of 14 weeks in Romney lambs, genetically resistant or susceptible to gastrointestinal nematodes. The lambs were infected with 5000 infective Trichostrongylus colubriformis larvae twice weekly. Compared to preinfection levels, the blastogenic activity of unstimulated lymphocytes in lambs of both lines peaked at week 3, and was significantly higher in resistant than in susceptible lambs. These changes may have been due to in vivo polyclonal activation. Lymphocytes from susceptible sheep responded more strongly to Con A, PHA and PWM than cells from resistant sheep. Counts per minute (c.p.m) for Con A- and PHA-stimulated lymphocytes increased in both lines of sheep from week 2 to week 7 and then returned to initial levels. An increase in c.p.m. in PWM-stimulated cell cultures was observed from weeks 3 to 5 in both groups. The blastogenic activity for LPS-stimulated cultures was significantly higher for resistant than susceptible sheep at weeks 3 and 4. No significant correlations between the decline in faecal egg counts (FEC) and the blastogenic activity was observed. Eosinophil counts in peripheral blood began to increase one week earlier in resistant than in susceptible sheep. No significant correlation between FEC and eosinophil counts was observed in resistant lambs, whereas in susceptible lambs a significant correlation was found between FEC and eosinophil counts at some sampling times. TWBC in resistant lambs steadily increased with infections whereas susceptible lambs showed a decrease until week 5 and then steadily increased. There was no significant correlation between the decline in FEC and TWBC.(ABSTRACT TRUNCATED AT 250 WORDS)

In sheep that had been given three immunizing infections with Trichostrongylus colubriformis and Ostertagia circumcincta infective (L3) larvae, drenched after the last infection and challenged with larvae of the same species, there was a significant increase in numbers of small intestine mucosal tissue globule leukocytes (TGLs) and lumenal globule leukocytes (LuGLs) compared with sheep that had only been drenched and challenged. There was a positive correlation between the numbers of LuGLs and TGLs in the small intestine but the ratio of these two cell types was lower in non-immunized than immunized sheep. In immunized sheep positive correlations were observed between LuGLs and levels of arylsulphatase and peroxidase in the intestinal mucus and between arylsulphatase and larval migration inhibition (LMI) activity in mucus. Lumen eosinophils correlated with blood eosinophils, serum antibody against T. colubriformis correlated with peroxidase in the mucus and blood eosinophils correlated with nematode specific IgM levels in the intestinal mucus. In the abomasum, TGLs were present but not LuGLs. Sheep repeatedly infected with T. axei also had significantly more LuGLs in the small intestine than control animals. Two sheep that had a surgically prepared isolated small intestinal loop, after oral infection with T. colubriformis had TGLs and LuGLs in the intact intestine, but not in the isolated loop. Significantly more LuGLs were produced in sheep by allowing repeated T. colubriformis L3 infections to develop to adult stages compared to sheep treated with the same number of larvae, but where the infections were terminated by drenching at various intervals.

Romney sheep, 1-2 years old, immunized by at least three anthelmintic abbreviated infections of 80-100,000 Trichostrongylus colubriformis larvae usually produced high numbers of intestinal mucosal mast cells/globule leukocytes (MMC/GLs). In isolating these cells, the importance of maintaining the intestine at 37 degrees C, removal of mucus with dithiothreitol, enzymatic dispersion and careful in vitro handling procedures for maximising cell viability are emphasised. The MMC/GLs were separated from most contaminant cells by using a Percoll discontinuous gradient. MMC/GLs collected at the 60/100% Percoll interface were passed through a complement coated nylon wood column to remove the contaminating eosinophils. Viable MMC/GLs were able to grow in vitro in the presence of Concanavalin A and survive in culture for up to 30 days. The MMC/GLs were readily identified by ultraviolet light microscopy after staining with auramine O.

The presence of globule leukocytes in the lumen (LuGLs) of the small intestine was studied in Romney sheep reared parasite free and then experimentally infected or immunized with Trichostrongylus colubriformis infective larvae, or naturally infected when gazed on pasture. It was discovered that high numbers of LuGLs were associated with both parasitic infection and the sheep protective immune response. A high positive correlation was observed between LuGLs and tissue globule leukocytes. The highest correlation (r = 0.92) observed was between larval migration inhibition (LMI) and numbers of LuGLs. This was the only correlation between LMI and other parameters studied. Significant positive correlations were also observed between LuGLs and IgG1 and eosinophils. There was a negative correlation between LuGLs and the number of parasites in the intestine and parasite egg production. Progeny of genetically resistant and susceptible sires had significantly different abilities to produce LuGLs.

Infection with the intracellular protozoan parasite Toxoplasma gondii ( T. gondii) causes health problems to both humans and livestock and has a large economic impact worldwide. The immune response in sheep following infection with T. gondii was evaluated using six different combinations of plasmid DNA, recombinant antigen and adjuvant. Sheep were generally vaccinated twice by intramuscular injection with plasmid DNA containing gene sequences for either the surface antigen (SAG1) or the rhoptry protein (ROP1) of T. gondii. Two of the groups injected with plasmid DNA SAG1 were boosted with recombinant protein (SAG1). We investigated the efficacy of including oligodeoxynucleotides (ODN) that contain CG motifs (CpG) and the gene coding for ovine granulocyte-macrophage colony stimulating factor (GM-CSF) as potential adjuvants. Administration of the plasmid encoding the ROP1 gene significantly enhanced both IFN-γ production from peripheral blood cells when cultured in vitro with Toxoplasma antigen, and ROP1-specific IgG1 and IgG2 antibody levels present in serum. However, injection with SAG1 did not stimulate IFN-γ production. These results indicate the potential of ROP1, given as plasmid DNA, as a potential vaccine candidate to protect sheep against T. gondii infection. [ABSTRACT FROM AUTHOR]

The nutritional cost of both the acquisition and maintenance of immunity to gastro-intestinal nematodes was investigated using immunologically naïve 5-month-old lambs and immunologically competent 17-month-old ewes. Within each age cohort, animals were either infected with the equivalent of 80 L3 Trichostrongylus colubriformis larvae per kg live weight (LW) per day (IF), similarly infected and concurrently immuno-suppressed with weekly injections of 1·3 mg/kg LW of the glucocorticoid methylprednisolone acetate (ISIF), immuno-suppressed only (IS) or remained as controls (C). Body composition of all animals was estimated using X-ray computer tomography on days -14 and 76 relative to the start of infection. Body weight and faecal nematode egg counts (FEC; eggs per gram of fresh faeces (e.p.g.)) were taken weekly and blood samples for serum proteins and antibodies were obtained every 2 weeks. FEC in IF lambs peaked at 1250 e.p.g. before a typical decline as immunity developed to less than 100 e.p.g. by day 75. FEC of less than 100 e.p.g. in IF ewes indicated immunity was maintained. Successful immuno-suppression in ISIF lambs and ewes was indicated by FEC of 4000 e.p.g. on day 75 and was confirmed by comparative worm burdens and serum antibody titres. The typical reduction in voluntary food intake (VFI)as a consequence of infection was observed in IF lambs (proportionately 0·30, P < 0·001) but not in IF ewes, ISIF lambs or ISIF ewes. Gross efficiency of use of metabolizable energy for net energy deposition was reduced by proportionately 0·20 in lambs during acquisition of immunity and by 0·16 in ewes maintaining an established immunity. Infection in immuno-suppressed animals reduced efficiency by 0·05 and 0·15 for lambs and ewes, respectively. These findings allowed the hypothesis that the reduction in VFI and loss in performance in young parasitised sheep is caused by physiological signalling associated with... [ABSTRACT FROM AUTHOR]

Abstract: The long-term benefit of suckling for the resistance and resilience of lambs to Teladorsagia circumcincta infection was investigated in a 20-week after-lambing trial period; during which time the immune response was anticipated to be strengthening. In a two by two factorial arrangement, one factor was ‘suckling’ with one twin lamb from each of 34 pairs being either weaned (W−) at 39 days of age or allowed to continue suckling (S−) until necropsy at mean ages of either 84, 112 or 140 days. The second factor was ‘parasite infection’ with lambs in each ‘suckling’ treatment group either artificially challenged with an equivalent of 1000 L3 T. circumcincta larvae d−1 (−I) from day 42 of age until necropsy or not artificially challenged (−N). All lambs and their dams grazed ryegrass pastures throughout the trial; these were intended to be relatively parasite safe, although worm burdens of SN and WN groups suggested there was a considerable parasite challenge from pasture from day 84 onwards. Despite this, suckled lambs consistently exhibited lower faecal egg counts (FECs; P <0.05) and worm burdens at slaughter on days 84 and 112 (P <0.05 for both). Worms from suckled lambs appeared to have reduced egg production indicating either a direct effect of milk proteins on parasite fecundity or an indirect benefit of suckling through the provision of nutrients for enhance immune capacity. Lamb growth rate was reduced by weaning but not by artificial challenge (P >0.05). It was concluded that while the enhanced protein supply from suckling will facilitate greater growth rates in young lambs, weaning may not necessarily be associated with reduced resilience to T. circumcincta infection. Furthermore, suckling appears to assist in reducing worm burdens and parasite fecundity either through a direct anti-parasitic effect or through hastening the acquisition of immunity when lambs are exposed to larvae early in life. [Copyright &y& Elsevier]