Pathophysiology of the periparturient egg rise in sheep: the role of prolactin

The involvement of prolactin in the periparturient rise in the faecal nematode egg count in sheep was investigated. Ostertagia circumcincta larvae (5000 third stage larvae three times weekly) were administered to adult immune ewes from three weeks before parturition to three weeks afterwards. Ten ewes were injected twice daily with 2-bromo-alpha-ergocryptine (bromocriptine), an antagonist of prolactin secretion, for two weeks starting two days after lambing while 10 ewes remained untreated. Bromocriptine treatment was initiated approximately two weeks pre partum in three other ewes. Plasma pepsinogen concentrations rose significantly by one week after the start of O circumcincta larval challenge in all the ewes but faecal egg counts remained negative until approximately one week post partum. Plasma prolactin concentration was reduced to a very low level in all bromocriptine treated ewes but this did not alter the dynamics of the periparturient rise in faecal egg counts. Neither cell-mediated nor humoral immunity of the ewes, as assessed by their sensitivity to BCG inoculation and by antibody titre raised against horse red blood cells, respectively, were impaired during the rise in faecal egg count, nor were these parameters altered by manipulation of plasma prolactin concentration. Lamb growth rate was not retarded by low plasma prolactin concentration in the bromocriptine treated ewes. These results are not consistent with the generally held hypothesis that elevated plasma prolactin concentration is directly associated with the periparturient rise.     

Use of a long acting injectable formulation of moxidectin to control the periparturient rise in faecal Teladorsagia circumcincta egg output of ewes

A field study was conducted in a sheep flock in the south east of Scotland with a history of ivermectin resistance in Teladorsagia circumcincta. The objective of the study was to compare the effects of single anthelmintic treatments in ewes before turn-out onto pasture that was contaminated with a moderate level of overwintered, ivermectin resistant, T. circumcincta infective larvae. The ewes were treated according to label directions with either a long acting injectable formulation of moxidectin (1mg/kg; affording up to 14weeks persistent action against macrocyclic lactone (ML)-susceptible T. circumcincta) or an oral formulation of moxidectin (0.2mg/kg; affording up to 5weeks persistent action against ML-susceptible T. circumcincta). The lambs were enrolled in the normal management of the farm, and received a total of three oral ivermectin treatments during the 16week study. The efficacy of both treatment strategies in controlling the periparturient rise in faecal nematode worm egg counts and subsequent pasture contamination was assessed from the faecal worm egg counts of the ewes and their lambs between lambing and weaning. Ewes that were treated with the oral formulation of moxidectin shed approximately 3.5 times more T. circumcincta eggs between lambing and weaning than ewes that were treated with the long acting formulation of moxidectin. This difference was reflected in the faecal worm egg counts of the lambs that were grazed alongside the different treatment groups of ewes. The results of the current study demonstrate persistent efficacy of the long acting formulation of moxidectin against an ivermectin resistant T. circumcincta population. The decreased pasture contamination after treatment could lead to improved lamb growth and a need for fewer anthelmintic treatments, thus potentially reducing one possible selection pressure for anthelmintic resistance. However, treatment with the long acting formulation of moxidectin would give rise to fewer susceptible nematodes being present in refugia, which could increase another possible selection pressure for anthelmintic resistance, depending on the subsequent grazing management of that pasture. The rationale for use of a persistent anthelmintic drug to control the periparturient rise in faecal ML-resistant T. circumcincta egg output of the ewes is discussed and potential differences in selection for macrocyclic lactone anthelmintic resistance using the different formulations of moxidectin are acknowledged.     

Haemonchus contortus and other trichostrongylid infections in parturient, lactating and dry ewes

From autumn (April) to spring (November), groups of pregnant and dry Merino ewes grazed pasture contaminated with infective larvae of Haemonchus contortus, Ostertagia circumcincta and Trichostrongylus colubriformis. A periparturient rise in faecal egg counts occurred in the pregnant ewes, following the maturation in early spring of arrested fourth-stage larvae of H. contortus and failure of pregnant ewes to expel the resulting adult worms. Peak egg counts were seen in September, just before lambing. In dry ewes, egg counts were slightly elevated at the time of the rise in the pregnant ewes, but fell to very low levels as adult worms developing from previously arrested larvae were expelled. Lactating ewes acquired greater burdens of O. circumcincta and T. colubriformis than did dry ewes, but were equally refractory to new infections with H. contortus. It is therefore suggested that the impairment of immunity to helminth infection seen in reproductive ewes may be more specific than was previously envisaged.     

Effects of protein supply and reproductive status on local and systemic immune responses to Teladorsagia circumcincta in sheep

The effects of protein supply and reproductive status on circulating antibody responses and local inflammatory cell counts were investigated in parasitized sheep, with local immune responses assessed through a recently refined abomasal cannulation methodology. We hypothesized that if breakdown of immunity has a nutritional basis, then protein scarcity would result in a breakdown of immunity to Teladorsagia circumcincta in both periparturient and non-reproducing (barren) ewes. Twin-bearing and barren, abomasally cannulated ewes were fed at either 0.8 or 1.3 times protein requirements from 3 weeks before until 6 weeks after parturition (n = 6). All sheep were trickle infected at a rate of 10,000 infective larvae (L3) per day, for 3 days per week throughout the experiment. Faecal egg counts remained virtually zero in all barren ewes, whilst protein supplementation reduced faecal egg counts in the periparturient ewes during most of the periparturient period. Final worm burdens, taken at 6 weeks into lactation, were lower for the barren ewes than for the lactating ewes, whilst protein supplementation reduced worm burdens in the latter. Protein supply did not affect mucosal mast cell counts, which were consistently higher for the barren ewes than the periparturient ewes, but were temporarily decreased around parturition. Barren ewes and protein supplemented lactating ewes had higher globule leukocyte counts than the unsupplemented lactating ewes. Protein supplementation increased eosinophil counts in the lactating ewes though only during the later part of the lactation period. Plasma IgA anti-L3 antibody was similar for all ewes, but IgE anti-L3 antibody was higher for the protein supplemented periparturient ewes compared to the unsupplemented periparturient ewes and all barren ewes. It is likely that the combination of low protein requirements and large body protein reserves did not result in breakdown of immunity to T. circumcincta for the barren ewes. These results suggest that changes in mucosal mast cell and eosinophil counts are not necessarily associated with changes in host resistance to T. circumcincta. However, the data support the view that increased globule leukocyte counts and plasma IgE anti-L3 antibody may be associated with nutritionally improved expression of immunity in periparturient ewes.     

Evaluation of Haemonchus contortus infections in sexually intact and ovariectomized ewes

Three experiments were conducted to investigate experimentally the occurrence of periparturient nematode egg rise in ewes and the hormonal modulation of Haemonchus contortus infections. In the first experiment, fall-bred and winter-bred pregnant (n = 4 and 14, respectively) and nonpregnant (n = 5 and 29, respectively) ewes were treated with anthelmintic and were pastured together on fields that were contaminated with H contortus. Three weeks before lambing, all ewes were placed in concrete pens; fecal egg counts for the winter-bred group were obtained on alternate days. Pregnant and lactating ewes had significantly larger numbers (P less than 0.01) of H contortus eggs than did the nonpregnant controls 1 week before and after lambing. Lactating, fall-bred ewes had significantly (P less than 0.01) more adult worms in their abomasum through natural acquisition than the nonpregnant controls. In the second experiment, fall-bred and winter-bred, helminth-free, pregnant (n = 4 and 8, respectively) and nonpregnant (n = 3 and 15, respectively) ewes were inoculated on 5 alternate days, beginning 70 days after breeding with 20,000 infective H contortus larvae. The ewes were maintained on concrete pens throughout pregnancy. Fecal egg counts were significantly (P less than 0.05) greater in pregnant ewes, beginning 1 week before lambing until 1 week after lambing. Abomasums of lactating ewes from both lambing seasons yielded significantly (P less than 0.01) more adult worms at necropsy than nonpregnant ewes. In the third experiment, ewes were ovariectomized (n = 15) or sham-operated (n = 9); half of the control ewes were bred.(ABSTRACT TRUNCATED AT 250 WORDS)     

Teladorsagia circumcincta egg output at the onset of natural and induced lactation in ewes

Nineteen ewes maintained under 'worm-free' conditions received anthelmintic and 1 week later were dosed on 5 days each week with 500 Teladorsagia circumcincta larvae for between 10 and 12.5 weeks. Faecal egg counts were determined twice weekly. There were three groups: pregnant (P) ewes (n = 6) which each reared twin lambs, hormone-treated (H) ewes (n = 7) which were not pregnant and were given exogenous hormones (dexamethasone, oestradiol-17 beta, progesterone) for 37 days to induce udder development and milk production, and untreated barren (B) ewes (n = 6). The plasma concentrations of oestradiol-17 beta, prolactin and progesterone were measured. Starting 10-15 days before birth in Group P, there was a biphasic periparturient rise in egg count and the onset of the first peak in egg count preceded the parturient peak in plasma prolactin concentration by several days. Most Group H (5/7) and all Group B ewes remained highly resistant to Teladorsagia infection throughout, despite an induced rise in prolactin concentrations in Group H ewes. It was concluded that factors other than prolactin initiate the periparturient rise but that prolactin is one factor which maintains the response once lactation commences.     

The effect of weather on the occurrence and magnitude of periparturient rise in trichostronglyid nematode egg output in Dorper ewes in a semi-arid area of Kajiado District of Kenya

An investigation was carried out to assess the effects of weather on the occurrence and magnitude of the periparturient rise (PPR) in trichostrongylid nematode egg output in breeding Dorper ewes. The study was conducted over three breeding seasons on a ranch in the semi-arid area of Kajiado District in Kenya between June 1999 and December 2001. During each breeding season 20 ewes randomly selected from the breeding stock and 20 others selected from the unmated yearlings were monitored for faecal strongyle egg counts every 3 weeks. The lambing seasons were timed to coincide with the onset of the short rains (October–November 1999), the mid-short rains (November–December 2000) and the end of the dry season (September–October 2001). In each season higher egg outputs were recorded in the peri-parturient ewes compared to the unmated yearlings. The highest PPR occurred in September 2001, when lambing coincided with the end of the dry season, possibly as a result of maturation of hypobiotic larvae. The lowest PPR occurred in November 2000 when the onset of lambing coincided with the mid-short rains, possibly owing to low pasture infectivity associated with a long dry spell between January and October of the same year. The results of this study indicate that PPR occurred when lambing coincided with both the wet and the dry seasons. However, the magnitude was greatly influenced by the season when lambing occurred. It was also influenced by resumption of development of hypobiotic larvae and the nutritional status of the ewes. Not only should control of gastrointestinal nematode parasites in this area aim at preventing the occurrence of PPR by treating ewes 2–3 weeks before they are to lamb and during lactation, but the anthelmintic used must also eliminate hypobiotic larvae. In addition, the animals must be given supplementary feeding during this period.     

Periparturient and seasonal rise in the trichostrongylid egg output of infected ewes during the dry season in Northern Nigeria

The faecal parasite egg output of lambing and non-lambing ewes in two sheep flocks in the dry season of 1975/1976 and 1976/1977 was determined. Ewes wormed at the end of the rainy season showed a very low parasite egg output during the dry season of 1975/ 1976. No periparturient rise of parasite eggs was detectable in the ewes which lambed in December or January. Towards the end of the dry season the parasite egg counts in the faeces increased and the highest counts were observed in lactating ewes.The parasite egg output at the beginning of the dry season was apparently due to recently acquired infections, mainly Trychostronglus sp. The increase in egg output at the end of the dry season was related to the development of inhibited larvae mainly of Haemonchus sp. This development was detectable from the end of February onwards.The differences in parasite egg output during the two seasons was associated with different rainfall patterns during 1975 and 1976.     

Periparturient rise in faecal nematode egg counts in west African dwarf sheep in southern Ghana in the absence of arrested strongyle larvae

The nematode egg output of two groups of ewes, a pregnant test group (2-3 years of age) mated between March and April and a non-pregnant control group (5-6 years of age) was followed. The worm burdens acquired by worm-free 'tracer' lambs were also followed on the same pasture from March 1988 to February 1989. There was a significant difference (P less than 0.05) in the magnitude of the egg output in the test group compared with the control group. In the test ewes, an initial rise in egg production occurred 2 weeks after lambing and was maintained for five consecutive months. Necropsy worm counts from the 'tracer' lambs revealed that pasture larval levels were directly related to the levels of rainfall. Adult nematodes (Trichostrongylus colubriformis, Trichostrongylus axei, Haemonchus contortus, Oesophagostomum venulosum and Trichuris ovis), and some Moniezia expansa were present in almost all the lambs throughout the year, with the exception of January and February 1989. The absence of immature larvae in these lambs indicates that arrested development is not a feature of the life cycle of these species in Southern Ghana. The occurrence of a periparturient rise of nematode eggs in West African Dwarf ewes, a year-round breeder, means that susceptible lambs could be open to infection throughout the year. Control should, therefore, be conducted by treating ewes after lambing and restricting the breeding season to particular periods of the year.     

Failure of moxidectin to control benzimidazole-, levamisole- and ivermectin-resistant Teladorsagia circumcinda in a sheep flock

Control of a benzimidazole-, levamisole- and ivermectin-resistant population of Teladorsagia circumcincta was attempted using moxidectin administered orally at the manufacturer's recommended dose rate of 200 microg/kg bodyweight. Ewes were dosed after lambing with the aim of controlling their periparturient rise in faecal egg output and lambs were dosed at six-week intervals throughout the summer. This regimen failed to suppress the establishment of significant numbers of infective helminth larvae on the pasture, resulting in unsatisfactory lamb production. Oral dosing with moxidectin was effective in removing adult female burdens of ivermectin-resistant T. circumcincta, but the effect of the drug did not persist against the resistant helminth population.     

Influence of protein supplementation during late pregnancy and lactation on the resistance of Santa Ines and Ile de France ewes to Haemonchus contortus

This experiment aimed to evaluate the effect of periparturient metabolizable protein (MP) nutrition on resistance to Haemonchus contortus in single rearing Ile de France and Santa Ines ewes. The restriction-fed iso-energetic diet was calculated to provide either 0.8 (low MP diet) or 1.3 (high MP diet) times MP, from three weeks before parturition until eight weeks into lactation. The ewes were experimentally infected with 1000 H. contortus infective larvae (L3) three times a week (Mondays, Wednesdays and Fridays), starting five weeks before the predicted date for parturition until a total of 15,000 L3 had been administered. While both breeds showed elevated fecal egg counts (FEC), these values were significantly lower for Santa Ines ewes than Ile de France ewes, but were independent of level of MP feeding. The latter also did not affect lamb weight gain and ewe body weight variation in each breed. Packed cell volume and total plasma protein for Santa Ines in all periods were significantly higher than those for Ile de France ewes (P<0.01) but were not affected by nutrition. In contrast, levels of serum IgG and IgA antibodies against somatic H. contortus infective larvae and adult antigens were similar between breeds but higher in animals that received high MP diets (P<0.05). The reduced body score of ewes at the beginning of the experiment probably influenced their high susceptibility to incoming larvae. Since, unexpectedly, MP scarcity was not achieved in this experiment, our data support the view that Santa Ines ewes are more resistant to H. contortus than Ile de France ewes.     

Can an increased intake of metabolizable protein affect the periparturient relaxation in immunity against teladorsagia circumcincta in sheep?

Within the overall hypothesis that the periparturient relaxation of immunity (PPRI) to gastrointestinal parasites has a nutritional basis, we studied whether an increased intake of metabolizable protein (MP) could affect the expression of immunity to Teladorsagia (Ostertagia) circumcincta of twin-bearing and -rearing Greyface ewes. Two sets of iso-energetic diets were formulated to provide either 85% (L-) or 130% (H-) of the MP requirements during the last 6 weeks of pregnancy (-P) and the first 6 weeks of lactation (-L). Thirty ewes were allocated to LP or HP. Six ewes of each group were slaughtered after lambing (S(0)); the remaining 24 ewes of each group were allocated to either LL or HL, thus creating four treatment combinations during lactation (LP/LL, LP/HL, HP/LL and HP/HL; n=12). Six ewes of each treatment combination were slaughtered at Week 3 (S(3)) or at Week 6 (S(6)) in lactation.The HP ewes were heavier at lambing, produced heavier lambs, had lower faecal egg counts (FECs), a lower concentration of plasma pepsinogen, a higher concentration of plasma albumin during late pregnancy and more abomasal globule leukocytes (GLs) but similar worm burdens at S(0) compared to the LP ewes. The LP/HL and HP/HL ewes had a higher calculated milk production than the LP/LL and HP/LL ewes. FECs increased for each treatment combination during the first 3 weeks of lactation. This increase concurred with fewer abomasal GLs at S(3) compared to S(0). The LP/HL and HP/HL ewes had a higher proportion of immature worms and more abomasal GLs than the LP/LL and HP/LL ewes at S(6). The FECs continued to increase during the second 3 weeks of lactation for the LP/LL ewes only, which concurred with an elevated plasma pepsinogen concentration and an absence of abomasal GLs at S(6). The plasma pepsinogen concentration decreased rapidly for the LP/HL ewes during lactation. We conclude that an increased intake of MP could enhance the expression of immunity to T. circumcincta of twin-bearing and -rearing Greyface ewes.     

The peri-parturient rise in sheep. Faecal worm egg counts in normal and late lambing ewes

The trichostrongylid and strongylid egg output in a group of ewes lambing in March, the middle of the usual lambing time in the Netherlands, was compared with counts in groups of older ewes and yearling ewes lambing in May-June and with counts in a group of barren ewes. The groups lambing in March showed only a rise related to parturition date as normally seen in the Netherlands. A few of the late lambing ewes showed a slight rise in spring, but all of them showed a peri-parturient rise related to parturition date. In the late lambing ewes and in the barren ewes egg counts due to reinfection were seen in summer. It is concluded that under the present circumstances a true spring rise, the calendar phenomenon, occurs very irregularly, while a peri-parturient rise related to the parturition date is a common phenomenon. The occurence of high egg counts in summer, excluding consideration on the barren ewes, in the late lambing ewes only, suggests that changing the traditional husbandry system may influence the epidemiological pattern of the trichostrongylid infections.     

The influence of reproductive physiology and nutrient supply on the periparturient relaxation of immunity to the gastrointestinal nematode Trichostrongylus colubriformis in Merino ewes

A pen experiment was conducted to investigate the interaction of early-weaning and nutrient supply on the periparturient relaxation of immunity to the gastrointestinal nematode (GIN) Trichostrongylus colubriformis in Merino ewes. Mixed-age pregnant and non-pregnant (dry) ewes were infected with 8,000 T. colubriformis L(3)/week, and fed either a high or low quality diet. Following parturition, lambs were either removed from their mothers at 2 days of age or allowed to continue suckling. Systemic immunity began to wane during late pregnancy with circulating eosinophils and plasma total antibody (Ab) levels declining from day -37 (relative to the midpoint of lambing) and day -24, respectively. Pregnant ewes fed the low quality diet exhibited an increasing faecal worm egg count (WEC) from day -24 and had higher intestinal worm burdens on day 13, whereas ewes fed the high quality diet had a delayed transient rise in WEC of lower magnitude. Dry and early-weaned ewes remained highly resistant to T. colubriformis at all times. In the post-lambing/lactation period, ewes fed the high quality diet had higher levels of local total Ab and numbers of goblet cells (GC) in the small intestine on days 13 and 41. Lactating/suckled ewes had a lower anti-parasite local immune response as indicated by reduced titres of total Ab, IgG(1), IgM and IgA and lower numbers of mucosal mast cells (MMC), globule leukocytes (GL) and GC in small intestinal tissue compared to their dry and early-weaned counterparts. Early-weaning resulted in rapid recovery of blood eosinophils and total Ab. On day 13 post-lambing, titres of total Ab, IgG(1), IgM, IgA and IgE, and numbers of MMC and GL were greater than those measured in dry and suckled ewes. When fed the high quality diet, ewes had a higher dry matter (DM) intake, maternal weight, fat score, greater fat depth and eye muscle depth, birthed heavier lambs that had higher growth rates, and produced more milk. The physiological status of pregnancy resulted in a higher DM intake but lower measures of fat depth and eye muscle depth, and suckling led to an increase in DM intake but a reduction in body weight and fat score through mobilisation of fat and muscle reserves. Despite the marked effect of diet quality on production traits, some inconsistencies were observed between body composition and apparent parasite resistance, measured by WEC and worm counts, suggesting that the nutritional influence was not necessarily always mediated through changes in body composition. Although reproductive status affected blood leptin levels, diet had no effect within suckled ewes and therefore it was concluded that leptin has no causative role in maintaining the periparturient relaxation of immunity to T. colubriformis.     

Epidemiological approach to the control of horse strongyles

An investigation of the spring rise in strongyle egg output of grazing horses on two commercial horse farms in northern USA in 1981 and 1982 revealed two distinct spring and summer rises in faecal egg counts, with peaks in May and August/September. There was a marked rise in the concentration of infective larvae on pasture two to four weeks after the peaks in egg output, so that grazing horses were at serious risk from June onwards and pasture larval counts on one farm did not fall to low levels until June of the following year. The spring and summer rises in faecal egg counts appeared to be seasonal in nature, to be derived largely from worms developing from previously ingested larvae, rather than from newly ingested larvae, and to be unrelated to the date of foaling. An epidemiological approach to strongyle control based on prophylactic treatments in the spring successfully eliminated the spring rise in egg output but was inadequate to control the summer rise or subsequent escalation of pasture infectivity in September. It was, nevertheless, superior to a conventional treatment programme at eight week intervals, using the same drug, pyrantel pamoate. Prophylactic spring/summer treatments proved to be much more effective. Both pyrantel pamoate at four week intervals and ivermectin at eight week intervals kept faecal egg counts at low levels during spring and summer. As few as two ivermectin treatments (11 May, 6 July) resulted in a sixfold reduction in pasture larval counts on 9 November and 3 January for the treated group (8872, 8416 stage three larvae [L3]/kg) compared to the control group (52,824, 50,984 L3/kg).(ABSTRACT TRUNCATED AT 250 WORDS)     

THE EFFECT OF ANTHELMINTIC TREATMENT OF MAIDEN EWES IN THE PERIPARTURIENT PERIOD ON PASTURE CONTAMINATION AND PRODUCTION OF PRIME LAMBS

SUMMARY A replicated experiment using first cross Border Leicester/Merino maiden ewes set stocked at the rate of 9.9 per hectare was conducted to examine the effects of treatment of the ewes in the periparturient period (PPP) and treatment of lambs at about 12 weeks of age on the production of prime lambs. All treatments given were thiabendazole. Four groups each of 100 ewes (4 replicates × 25) were treated either pre-lambing, 2 weeks before lambing commenced and again post-lambing, 8 weeks later when lambing was almost complete (DD), pre-lambing only (DO), post-lambing only (OD) or not treated during the lambing period (OO). Lambing commenced 24 July 1972 with the bulk of lambing (98%) occuring within the first 4 weeks of a 6 weeks lambing period. The effectiveness of the drenching regimes was judged by observing bodyweight gains per day during four overlapping intervals between birth and 18 weeks of age. The bodyweight gain of lambs up to 18 weeks and the number of lambs exceeding a target weight of 33 kg at 18 weeks of age were considered to be critical measurements of production efficiency. Significant bodyweight differences were observed within 30 days from birth, favouring DD and DO over OD and OO lambs (p < 0.01). These differences continued until 12 weeks of age at a reduced level of significance (p < 0.05). By 18 weeks of age the results favoured DD over DO (p <0.05) over both OD and OO lambs (p < 0.05). Further differences were created by the treatment of lambs at 12 weeks of age (p < 0.01). At no time during the entire experiment were any significant differences observed between OD and OO lambs. The periparturient rise (PPR) in faecal nematode egg output was monitored throughout the experiment. Anthelmintic treatment of the ewes in the PPP did not eliminate the PPR but did affect the onset, magnitude and duration of the PPR which was either delayed by the prelambing treatment or truncated by the post-lambing treatment; treatments both before and after lambing delayed and depressed the PPR and reduced total pasture contamination by 29%. An effect on bodyweight of ewes was also observed with ewes being treated pre-lambing (Groups DD, DO) being significantly heavier at 18 weeks post-lambing than OD and OO ewes (p < 0.05). The observed effects on lamb production have been attributed in the main to an apparent, though not specifically measured, effect on lactation in those ewes being treated pre-lambing. The effect of delayed (Groups DD, DO) and/or reduced (Group DD) pasture contamination was considered to be additive to the earlier lactation response. Treatment of lambs at 12 weeks did not obscure these responses.     

The epidemiology of abomasal nematodes of sheep in Sweden, with particular reference to over-winter survival strategies

In May 2002, studies on the seasonal patterns of nematode infection of sheep were undertaken on four commercial sheep farms in southern Sweden, which had previously reported problems with nematode parasitism, especially due to Haemonchus contortus. One farm was used for intensive investigation. This entailed the establishment of two replicate groups of sheep, each consisting of 20 ewes and their lambs, on adjacent pasture paddocks. The seasonal patterns of nematode infection were followed by regular (approximately monthly) sampling of both ewes and lambs for nematode faecal egg counts and larval differentiation, and the sequential use of replicate groups of tracer lambs. H. contortus and Teladorsagia circumcincta were the most abundant nematode species, with the former most prevalent in the post-parturient faecal egg counts of ewes. Tracer worm counts showed almost 100% arrested development in the early fourth larval stage for H. contortus as early as mid-summer and the numbers of parasites progressively increased during the season. T. circumcincta also showed high levels of arrested development, but not as early, or as absolute, as for H. contortus. Tracers allocated to the paddocks at the time of turn-out following winter in May 2003, showed virtually a total absence of H. contortus in contrast to exceedingly high infections with T. circumcincta. Results of the three additional monitoring flocks supported these findings. It can be concluded that under Swedish sheep farming conditions, H. contortus has evolved to survive the long, cold winters entirely within the host as the arrested larval stage, relying on the lambing ewe to complete its life cycle. The peri-parturient relaxation of resistance in the ewes triggers the resumption of development to the adult egg-laying parasites to result in pasture contamination and the completion of just one parasite generation/year. In contrast, T. circumcincta can survive well over-winter, both on pasture and within the host.     

Serum and abomasal antibody response of sheep to infections with Haemonchus contortus

Serum and abomasal IgA, IgG and IgM antibody response against adult worm, L3 and egg antigens of Haemonchus contortus was monitored by the ELISA technique after one or two infections with this nematode. Following the first infection, antibody levels in serum did not change materially. After administration of a challenge dose of infective larvae, antibodies of the three immunoglobulin classes in infected animals rose slightly, but this rise appeared later than the fall in the faecal egg counts. In contrast, in abomasal mucosa, IgA anti-larval antibody levels, which did not increase materially after the primary infection, rose rapidly after a transient inhibition when sheep were challenged. A close temporal relationship was observed between the rise in local anti-worm IgA antibodies and the self-cure reaction, but antibody levels fell rapidly after worm diminution. The local antibody response was thus considered to be related to immunity of sheep to H. contortus.     

The immune response of sheep to larval challenge with Ostertagia circumcincta and O. ostertagi.

One of three groups of sheep was challenged twice-weekly with infective-stage larvae (L3) of the sheep parasite O. circumcincta, another with the cattle parasite O. ostertagi while the third received no larval challenge. Positive faecal egg counts (FEC) and a rise in plasma pepsinogen levels were observed only in those animals given O. circumcincta. Anti-O. circumcincta L3 IgG titres were rapidly elevated during parasite challenge with either O. circumcincta or O. ostertagi. Throughout the experiment, no rise in anti-adult IgG titres or eosinophil numbers was observed in peripheral blood in any group. On evidence of self-cure of the trickle-infection, determined by a reduction in FEC, all groups were drenched and challenged with 15,000 O. circumcincta L3. No effect of previous challenge on parasite establishment or FEC was observed, although egg viability was significantly reduced in both groups given prior challenge. Significant differences in adult female worm length were observed between groups. Those recovered from animals previously challenged with O. circumcincta were shorter than from those given O. ostertagi which were in turn shorter than those from previously unchallenged animals. In utero egg counts were significantly lower in worms from animals previously challenged with O. circumcincta than in those from unchallenged control animals. The results indicate that a level of immunity to O. circumcincta can be conferred by exposure to O. ostertagi.     

Effects of trichostrongyle larval challenge on the reproductive performance of immune ewes

Ostertagia circumcincta larvae were administered to adult immune ewes either during the breeding period or around parturition to investigate whether reproductive performance was affected. Two groups of 25 ewes were given water or 12,000 ostertagia larvae three times weekly for five weeks before and five weeks after breeding to synchronised oestrus. Plasma pepsinogen levels were raised in the larval-challenged ewes during the treatment period but no significant effect of treatment was observed on ovulation rate or plasma progesterone concentration. Neither ewe live-weight nor body condition score differed between groups throughout gestation or for four weeks post partum and there was no effect from larval challenge during the breeding period on lambing rate, lamb birthweight or liveweight at four weeks old. A second experiment involved 20 twin-bearing immune ewes which were administered 17,000 ostertagia larvae three times weekly for three weeks before and three weeks after parturition. Plasma pepsinogen concentration was elevated during the larval challenge period and nematode eggs were detected in the faeces after parturition but there was no effect from treatment either on ewe liveweight and body condition score or on lamb birthweight and lamb liveweight at four weeks old. These results indicate that larval challenge with ostertagia larvae at dose rates comparable to larval intake from heavily contaminated pasture did not affect the general well being and reproductive performance of immune Greyface ewes.