The establishment and development of Haemonchus contortus in goats

Twelve goats were inoculated with 40,000 third-stage Haemonchus contortus larvae and two were killed on each of Days 4, 7, 11, 14, 18 and 21 after inoculation (DAI). The number of worms that established, and the site of development were recorded. More worms established in the fundic, than in the middle or pyloric thirds of the abomasum. Early development occurred within the mucosa; emergence into the lumen started between 7 and 11 days after infection. By 4 DAI, all worms had completed the third moult to the L4 stage. At 11 DAI the majority of the worms were adults. A mean of 13.2% of the female worms had eggs in their uteri at 18 DAI; by 21 DAI more than half of the female worms had eggs in their uteri. The development of H. contortus was essentially similar to that described in sheep.     

The establishment and development of Trichostrongylus colubriformis in goats

Twelve goats were inoculated with 20,000 infective larvae of Trichostrongylus colubriformis and two were killed on each of Days 4, 7, 11, 14, 18 and 21 after inoculation (DAI). The number of worms that established, and the site of development were recorded. Most of the worms established within the first 3 m of the small intestine. There was little relocation or loss of nematodes after establishment. The worms started to migrate from the mucosa to the lumen 11 days after infection. At 4 DAI, 63% of the worms were still at the late L3 stage; the remainder of the worm population had completed the third moult to the L4 stage. The population at 11 DAI comprised largely young adults. When 21 DAI was reached, about 57% of the female worms had eggs in their uteri.     

Effect of Ostertagia ostertagi on lamb performance and cross resistance to O circumcincta

Fifteen worm-free lambs (two-and-a-half to three months or four to four-and-a-half months old) were infected with 3500 or 4000 Ostertagia ostertagi larvae on five days each week for six weeks, and their performance compared to that of controls. Eleven lambs were killed after eight weeks and four were challenged with O circumcincta to determine whether any cross resistance had developed. A feature of the O ostertagi infection was the considerable variation in response. Overall liveweight gain was lowered by 24 per cent in the two-and-a-half to three-month-old infected lambs, mainly due to reductions of 27 to 40 per cent in four of the seven lambs. There was no consistent effect in the older lambs. The worm populations consisted mainly of early fourth stage larvae and developing worms, but a small percentage reached sexual maturity and these produced a low faecal egg count (1 to 63 eggs per gram). Numerous intraluminal refractive crystals were present in the gut of both adult worms and developing stages, possibly reflecting degenerative changes. Hypertrophy of the abomasal mucosa with patchy loss of differentiation was a feature of the infection, and in four lambs serum pepsinogen concentrations were elevated. Exposure to O ostertagi did confer some protection against challenge with O circumcincta in that worm counts were reduced to about 60 per cent of those in controls, although no increase was observed in the numbers of arrested larvae. The successful passage of O ostertagi through young lambs could be important in mixed or alternate grazing systems by providing a reservoir of infection for the alternate host.     

Development of immunity to incoming radiolabelled larvae in lambs continuously infected with Ostertagia circumcincta

Infective larvae of Ostertagia circumcincta were radiolabelled with 75selenium by a method which did not affect their viability. Three groups of five-month-old lambs were infected daily with 1000 unlabelled infective larvae for four, eight and 12 weeks, respectively. After each period one of these groups and a group of worm-free controls were challenged with three consecutive daily doses of 1000 radiolabelled third stage larvae. The lambs were killed 13 days after the first dose of challenge larvae and their worm burdens examined. The first indication of immunity was retardation of developing worms observed at four weeks. Resistance to the establishment of incoming worms developed between four and eight weeks and a brief period of population turnover probably took place at this time. Simultaneously a greater inhibition of worm development occurred resulting in an increase in the number of parasites recovered as early fourth stage larvae. By 12 weeks the animals were almost completely immune to incoming worms. The development of resistance to incoming worms correlated with a rise in serum antibody titre and an increase in the number of intraepithelial globule leucocytes in the gastric mucosa.     

Morphogenesis of Trichostrongylus rugatus and distribution during development in sheep

Morphogenesis of Trichostrongylus rugatus was examined in 16 sheep experimentally-infected with 120 000 third-stage larvae and killed 2, 4, 6, 8, 10, 12, 16 and 20 days later (DAI). Third stage larvae moulted between Days 4 and 6, and fourth stage larvae moulted on Day 10 after infection. Four sheep first passed eggs in the faeces between Days 16 and 18 after infection. Rate of growth of larvae was constant between 2 and 10 DAI followed by a period of rapid growth from 10 to 16 DAI. Major features of larval development are described. Nematodes were largely restricted to the first 6 m of gut with 71% of worms occurring in the first 3 m.     

The effects on merino lambs of chronic infection with Trichostrongylus rugatus

Four groups of six 5-month-old merino lambs were exposed to infection with 12,000 larvae of Trichostrongylus rugatus per week and one group was killed at each of 3, 6, 9 and 12 weeks. Numbers of adult worms increased in an approximately linear manner throughout the experiment; the percentage of larvae developing to adults remained between 35 and 25% throughout the course of the experiment. Sub-total villus atrophy was evident in the anterior small intestine of each group of infected sheep and significant reductions occurred in the plasma concentrations of albumin and phosphate. Serum globulin and calcium concentrations rose in infected animals. No clinical signs were observed in infected sheep, no alteration in bone histology was detected and no decrease in growth rate occurred.     

Interactions between Teladorsagia circumcincta and Trichostrongylus vitrinus infections in young lambs

Groups containing six lambs that previously had been infected five times each week for four, eight or 12 weeks, either with 1000 Trichostrongylus vitrinus third stage larvae (L3) or 1000 T vitrinus L3 and 2500 L3 Teladorsagia (previously Ostertagia) circumcincta were challenged, along with six worm-free control lambs, with 3000 75Se-labelled T vitrinus L3. The lambs were killed 10 days after challenge. Those given a dual infection for four and eight weeks had significantly lower total T vitrinus numbers than monospecifically infected lambs. Significantly fewer radiolabelled larvae were recovered from the dual infected group killed after four weeks exposure to infection than from lambs infected monospecifically. The negative interaction between these two nematode species appears to be physiologically mediated and may be an important factor in accounting for differences in their seasonal prevalence.     

The therapeutic efficacy and prophylactic activity of doramectin against Dictyocaulus viviparus in cattle

SUMMARY Three groups of 8, 4-month-old male Jersey or Jersey-cross calves were infected with 2400 Dictyocaulus viviparus L₃ larvae and either left untreated or injected subcutaneously with 200 μg/kg doramectin 5 or 25 days after infection (DAI). Lungworms were found in all untreated cattle (geometric mean = 49) at necropsy 39 or 40 DAI. None was found in any of the treated cattle. In a second experiment, groups of 6, 8-month-old calves were untreated or injected with 200 μg/kg doramectin 28, 21 or 14 days before each calf was challenged with 2700 D viviparus larvae. Lungworms were recovered at necropsy 32 to 34 DAI. The geometric mean worm burden in the untreated cattle was 550. This was reduced by 100%, 99.5% and 94.1% in calves treated with doramectin 14, 21 or 28 days, respectively, before infection. It was concluded that doramectin is a highly effective anthelmintic against D viviparus adult or L₄ infections of cattle, and that reinfection of treated cattle will be significantly reduced for at least 28 days after treatment.     

Development of immunity to incoming radiolabelled larvae in lambs continuously infected with Trichostrongylus vitrinus

Infective third stage larvae (L3) of Trichostrongylus vitrinus were radiolabelled with 75 selenium by a method which did not affect their viability. Three groups of six-month-old lambs were infected daily with 1000 L3 for four, eight and 12 weeks, respectively. After each period, one of those groups (n = 5) and a group (n = 4) of worm-free controls were challenged with three consecutive daily doses of 1000 radiolabelled L3, killed 10 days after the first dose, and their worm burdens examined. After four weeks of continuous infection partial immunity to the establishment of challenge L3 was apparent, and by eight and 12 weeks, with the exception of one sheep, there was almost total resistance to incoming worms. Immunity was also expressed as an inhibition of the development of established worms.     

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.     

Studies on Ostertagia spp. from Greenlandic sheep: arrested development and worm length

Studies were conducted to investigate earlier observations that lambs slaughtered in Greenland in late September/early October have relatively few arrested larvae in the abomasal mucosa and that the adult Ostertagia from such lambs are abnormally large. Post mortem examination of naturally infected ewes in mid-winter demonstrated that a significant proportion of their Ostertagia population was hypobiotic at this time of year, while an experimental study showed that considerable numbers of larvae of Greenlandic origin became arrested following cold conditioning for 10 weeks. A comparison of adult Ostertagia of Greenlandic and Britain origins from lambs experimentally infected with larvae cultured under identical conditions demonstrated that the "giant" size of the arctic worms was attributed mainly to environmental rather than genetic influences.     

Infection of goats with Haemonchus contortus and Trichostrongylus colubriformis: histopathology and pH changes.

Two groups of goats were experimentally infected with Haemonchus contortus and Trichostrongylus colubriformis, and killed at various days after infection (DAI). The percentage of worms that established in the abomasum and the small intestine was low. At necropsy, abomasums from infected goats had thickened walls and oedematous folds. At 7 DAI there was an initial infiltration of eosinophils and some neutrophils which tend to increase with age of infection. The mean pH of the abomasum in goats infected with H. contortus was 5.43 (range 5.3-5.7), while that of the control goats was 3.30 (range 2.8-3.7).     

Observations on the pathogenic effects of Nematodirus helvetianus in dairy calves.

Six Holstein-Friesian calves 3 to 5 months old, were used in a small-scale experiment to determine the pathogenic effects of the thread-necked strongyle Nematodirus helvetianus. Two calves were kept as controls and 4 were infected, 2 with single doses of 25,000, and 2 with 50,000 N helvetianus larvae. Observations made during the 8 weeks of the experiment found that N helvetianus is capable of producing severe pathologic changes in the intestine of previously healthy calves. At 38 days after infection, the calves infected with 25,000 and 50,000 larvae gained 32.9% and 72.4% less weight respectively, than their controls. An increase in total lymphocytes was observed with no significant change in hemoglobin concentration. Adult worms were found to penetrate the intestinal mucosa with considerable destruction, including tunneling. Rectal temperatures were elevated in infected animals. The prepatent period for N helvetianus in single infection was 18 to 19 days. A spontaneous expulsion of the adult parasites was observed in 1 of the calves exposed to 50,000 larvae. Worms recovered in this animal numbered 565, whereas the number of worms recovered in another calf infected with 50,000 larvae was 21,860.     

Seasonal Fluctuation an Inhited Development of Populations of Dictyocaulus filaria in Ewes and Lambs

The fluctuation of populations of Dictyocaulus filaria in sheep was studied under field conditions in which animals are housed during the winter and grazed from late spring to autumn. A comparison was made between residual pasture contamination with overwintered larvae, the fecal larval deposition by ewes from June and both of these factors combined as sources of infection for spring born lambs. Ewes and lambs were killed serially over a year and worms were recovered from the lungs and counted. It was found that during the stabling period most of the ewes were carrying moderate numbers of D. filaria. However, while the vast majority of lungworm populations in the winter was inhibited in development at the early fifth larval stage, virtually all worms in the spring were adults. Any one source of infection studied contributed to the acquisition of important burdens of D. filaria by lambs as well as ewes. Worm counts reached peak in all lambs by November and this pointed to only one important Dictyocaulus generation per grazing season. It would also appear that larvae picked up by ewes and lambs as the grazing season advanced had become inhibited in development with the inhibition rate being most marked in autumn.     

Effects of dietary molybdenum on nematode and host during Trichostrongylus vitrinus infection in lambs.

The addition of molybdenum (0.05 mmol kg-1 dry matter) to the diet of lambs exposed for four weeks to a trickle (2500 third stage larvae per day) infection with Trichostrongylus vitrinus reduced the number and length of adult worms retrieved from the small intestine 11 days later: both effects were particularly marked in female worms from female lambs (P less than 0.01). Worms from lambs given molybdenum contained less proteinase enzyme activity and secreted less proteinases in culture irrespective of the sex of the host. Pathogenicity was not attenuated by molybdenum. Damage to the intestinal mucosa was severe in both dietary groups but infected females given molybdenum developed lower plasma albumin concentrations and lighter dressed carcases than those not given molybdenum. Neither the effects on the parasite nor those on the host could be attributed simply to molybdenum-induced copper depletion, using conventional measures of copper status. Molybdenum may be toxic to T vitrinus but may also facilitate or enhance the inflammatory process limiting larval establishment or increasing parasite rejection.     

The effect of dietary protein on the pathogenesis of acute ovine haemonchosis

Two experiments were conducted to examine the ifluence of dietary proteins on parasite establishment and pathogenesis in Finn Dorset/Dorset Horn lambs infected with Haemonchus contortus. The lambs were introduced to high (169 g) or low (88 g crude protein (CP) kg⁻¹ dry matter (DM)) protein diets at 3 months of age and infected 1 month later with 350 larvae kg⁻¹ body weight (BW). Blood and faecal smaples were collected for analysis and body weights recorded weekly. In the first experiments some of the infected lambs were killed 6 weeks after infection and the remainder 5 weeks later. In the second experimental all the infected lambs were killed 4 weeks after infection.

The results showed that lambs on a low protein diet were less able to withstand the pathogenic effects of infection with 350 H. contortus larvae kg⁻¹ BW than lambs given the higher protein diet. Thus mortality was greater in the low protein group and adverse clinical signs, such as inappetance, weight loss and oedema were osbserved more frequently. This group also had a more severe anaemia, hypoproteinaemia and hypoalbuminnaemia than the high protein group. In contrast, faecal egg counts, total daily faecal egg output and worm burdens were similar in all groups of infected lambs, indicating that the diets did not influence parasite establishment.     

Effect of experimental Ostertagia circumcincta infection on the performance of grazing lambs

Groups of 10 three-and-a-half-month-old Suffolkcross lambs were infected daily, five times a week, for 12 weeks with either zero, 500, 1500, 3000 or 5000 larvae of Ostertagia circumcincta and grazed together on a series of 'clean' paddocks. All the lambs were moved at seven to 10 day intervals to a fresh paddock to prevent autoinfection and were killed after 14 weeks. Another group was killed at the outset as a control for carcase analysis. Mean O circumcincta populations were 34 (grazing control) and 1224, 473, 1092 and 4331 for the groups receiving 500, 1500, 3000 and 5000 larvae per day respectively. The percentage of fourth stage larvae increased with increasing larval intake. Extensive abomasal damage with mucosal hypertrophy and depletion of parietal cells occurred in the lambs receiving 3000 and 5000 O circumcincta larvae per day. Intakes of 1500 larvae per day and above depressed growth rate by 24 to 37 per cent, reduced the deposition of protein in the carcase by 34 to 52 per cent and lowered calcium/phosphorus deposition by 23 to 48 per cent. Plasma pepsinogen concentrations were consistently elevated in lambs receiving 3000 or 5000 larvae per day and may be a useful aid to recognising abomasal damage sufficient to affect productivity.     

Influence of dietary protein on parasite establishment and pathogenesis in Finn Dorset and Scottish Blackface lambs given a single moderate infection of Haemonchus contortus

An experiment was conducted to examine the influence of dietary protein on parasite establishment and pathogenesis of Haemonchus contortus in lambs of two breeds. Finn Dorset and Scottish Blackface. The lambs were introduced to high protein (170 g crude protein (CP) kg-1 dry matter (DM) or low protein (88 g CP kg-1 DM) diets at three months old and infected one month later with 125 larvae (kg bodyweight)-1. Blood and faecal samples were collected for analysis and bodyweights recorded weekly. The lambs were killed 20 weeks after infection. A second experiment was conducted in Blackface lambs solely to determine the influence on parasite establishment of the high and low protein diets following infection with 125 larvae kg-1. These lambs were killed five or 14 weeks after infection. Results showed that in Scottish Blackface lambs dietary protein did not significantly influence the establishment of H contortus. However, in Finn Dorset lambs dietary protein may have influenced parasite establishment since lambs on the low protein diet had a higher faecal egg output four weeks after infection and more severe clinical signs than infected lambs of the same breed on a high protein diet.     

Strongyloides papillosus: prenatal and transmammary infection in ewes

The absence of larvae in lambs of six infected and controlled ewes immediately killed at birth without suckling may indicate the absence of prenatal infection. The ewes had larvae in their milk immediately prior to lambing and several hours afterwards. The persistence of infection, even after intestinal infection was removed with thiabendazole at 50 mg/kg body weigh, may be due to the pool of inhibited larvae. All the suckling lambs had patent infections within 4 to 5 days. Infection of suckling lambs with Strongyloides papillosus via milk from their ewes demonstrates the transmammary route of infection of this nematode.     

Does suckling offer protection to the lamb against Teladorsagia circumcincta infection?

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.