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.

Implications of nutrition for the ability of ruminants to withstand gastrointestinal nematode infections.

Resistance and resilience of the ruminant host to gastrointestinal (GI) parasitic nematode infections are influenced by many factors, including nutrition. This review examines the effects of host nutrition on the ability of ruminants to withstand GI nematode infections. Firstly the effects of infection on host metabolism are summarised briefly. An important factor in the pathogenesis is a reduction in feed intake by the host. Gut nematodes also increase endogenous protein losses, which result in net loss of amino acids to the parasitised host, though energy and mineral metabolism are also perturbed. The indications are that the major nutritional change is in protein metabolism. Resilience (the ability of an animal to withstand the effects of infection) can be enhanced markedly by increasing metabolisable protein supply and to a lesser extent metabolisable energy supply. Resistance to GI nematodes (ability of host to prevent establishment and/or development of infection) is also influenced by diet, particularly metabolisable protein supply. While there do not appear to be any effects of host nutrition on establishment of infective larvae, the rate of rejection of adult worms can be enhanced by improved nutrition. The exact nutritional requirements or the mechanisms involved are not known. It appears that the effects of improving nutritional status on host resilience are more clearly defined than effects on host resistance. The implication of changes in host resistance with nutritional state for host productivity need to be better described. Understanding the role of nutrition in improving both resistance and resilience of the host to GI parasites will be important if producers are to make better use of host acquired immunity and reduce dependence on pesticides for prophylaxis.

The effect of age and dietary protein on immunity and resistance in lambs vaccinated with Trichostrongylus colubriformis.

Development of immunity and resistance to Trichostrongylus colubriformis was measured in sheep, 8-26 or 33-51 weeks of age, that were offered two levels of dietary protein (11 and 20%). Resistance to challenge infection was ascertained by measuring worm burden, faecal egg count and eggs in utero in the nematode. Development of immunity was measured by an in vitro lymphocyte blastogenesis test to T and B cell mitogens and to somatic antigen from infective (L3) larvae. Young lambs offered the low protein diet showed a significantly lower resistance to parasites than older animals. The young animals on the high protein diet developed better resistance. Age and dietary protein influenced in vitro T lymphocyte responses to parasite antigen and mitogens, which were particularly well developed in vaccinated lambs on the high protein diet. Lymphocyte responsiveness to L3 larval antigen was negatively correlated with parasite status in vaccinated older lambs, and may be important in the development of acquired resistance to T. colubriformis.

The sequential analysis of local inflammatory cells during abomasal nematode infection in periparturient sheep.

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.

Studies using Teladorsagia circumcincta in an in vitro direct challenge method using abomasal tissue explants.

An in vitro direct challenge (IVDC) method in which abomasal tissue explants maintained at 37 degrees C in Hanks/Hepes solution (pH 7.6) in a high oxygen concentration (incubator gassed with pure oxygen and sealed prior to use) were-challenged with exsheathed third stage larvae of Teladorsagia circumcincta has been used to investigate the tissue association phase of larval establishment. Studies using naïve and recently challenged immune sheep and goat abomasal tissues have shown significant reductions (P < 0.05) in the numbers of tissue associated larvae in material from immune animals. The mechanisms that affect the tissue association process appear to be labile since more larvae were recovered from the tissue digests of previously infected sheep that had not been recently exposed to larval challenge in comparison to those that had been recently challenged (P < 0.05). The method has also been used to demonstrate the influence of protein nutrition on the establishment of larvae in abomasal tissues and region specific differences in the efficacy of exclusion mechanisms. The technique appears to have potential as a means of investigating the crucial first phase of the process of establishment.

Enhancing the profitability of pasture-based dairy production in the humid tropics through improved nutrition.

Dairy production in the tropics represents a major challenge, especially when reliant on grazed forages, because of the conflicting factors of a high nutritional demand to sustain lactation on the one hand, and the relatively low quality of tropical grasses and a stressful environment, on the other. This paper focuses primarily on those tropical situations where grazed pasture is the cheapest feed resource. Effective management of the pasture as well as the animals that graze it is required in order to maximize economic viability of the farming enterprise. Feed planning helps to ensure that pasture use is maximized, either directly by grazing or, where profitable, indirectly by cutting for conservation. High rates of pasture utilization at each grazing minimize pasture senescence and decay and ensure that pasture remains in a vegetative state of the highest possible nutritional quality. Total annual feed requirements of the grazing herd must therefore be matched as closely as possible to the total annual pasture production on the farm. Stocking rate is critical in this regard, and is the single most important determinant of productivity in pastoral farming. Periods of pasture surplus or deficit will inevitably arise, but can be minimized by matching the monthly feed requirements of the herd as closely as possible to the monthly feed production on the farm. Herd requirements can be influenced, for example, through altering calving patterns and drying off dates. Even with good pasture management practices, it is highly likely that tropical pasture quality will be low for part of the year. Utilization of low-quality forages can be improved through the appropriate use of feed supplements, and the key principles are discussed. Firstly, conditions for optimum fermentative digestion in the rumen must be promoted through adequate provision of fermentable energy and nitrogen as well as essential minerals, such as phosphorus and sulfur. Further supplementation with metabolizable energy or protein, depending on what limits animal production, may be profitable. Various practical supplementation strategies for tropical forages are discussed. Finally, some suggestions for future research and development for enhancing the profitability of pasture-based dairy production in the humid tropics are made.

The effects of a concurrent infection of Trichostrongylus colubriformis and Ostertagia circumcincta on calcium, phosphorus and magnesium transactions along the digestive tract of lambs.

Twelve lambs, paired on the basis of live weight, were cannulated in the abomasum, proximal jejunum and terminal ileum. Six were infected with 3000 T. colubriformis and 3000 O. circumcincta larvae per day for 18 weeks. The other six lambs were pair-fed to the infected lambs. All animals were offered fresh ryegrass and white clover pasture, cut daily. Dry matter intake, live weight and plasma Ca, P and Mg concentrations were measured throughout the experiment. During weeks 7 and 17 post-infection, digesta flow along the gastrointestinal tract of infected and control animals was measured. Parasitism depressed dry matter intake by 60 per cent and was associated with a reduction in retention of Ca, P and Mg in both infected and control animals. During weeks 7 and 17, parasitism increased the flow of Ca past the proximal jejunum and, during week 17, the flow of P past the terminal ileum. Increased endogenous Ca and P losses, together with a net reduction in absorption of Ca and P, were associated with depressed plasma Ca and P concentrations. These findings, together with evidence of reduced addition of P to anterior regions of the tract, indicate induced Ca and P deficiency. There was evidence for compensatory absorption of Ca and P in the large intestine and distal small intestine, respectively. Magnesium metabolism was apparently not affected by parasitism.

Skeletal and blood biochemical characteristics of sheep during growth and breeding: a comparison of flocks with and without broken mouth.

Two-hundred female Scottish Blackface sheep ranging in age from 3 months to 5 1/2 years were used to investigate the relationships between blood biochemical characteristics, skeletal metabolism and dental health on 5 farms, 3 with broken mouth and 2 with no incidence of the disease. Skeletal growth to 6 months of age was faster in lambs from control flocks than in lambs from affected farms although growth thereafter was slower in the control group. A generally lower protein intake on control farms may account for this reduced growth rate. In general, adult dimensions of bones were attained between 14 and 30 months of age but age of maturation varied between bones; tibial endochondral growth ended after 14 to 21 months while the rib still showed evidence of endochondral growth at 70 months of age. This growth was episodic, being maximal during lactation and possible causes are discussed. There was no significant differences in calcium or phosphorus status between control and affected populations, an inadequate phosphorus intake during pregnancy was common to both groups. Lower serum magnesium concentration in affected than in control sheep was a consistent finding and is discussed.

The rate of recovery of intestinal morphology following anthelmintic treatment of parasitised sheep.

Sixteen sheep which had previously been infected with Ostertagia circumcincta were dosed with 4000 Trichostrongylus colubriformis larvae/day for four weeks, then given anthelmintic and killed at intervals thereafter. Histological examination of the small intestine showed that two to three weeks were required for restoration of a normal mucosal architecture.

Chronic infection with Trichostrongylus vitrinus in sheep. Pathological changes in the small intestine.

Eight four-month-old sheep were infected daily for 90 days with 2500 Trichostrongylus vitrinus larvae. Infections were patent by week 3; mean faecal egg counts reached a peak of 645 epg at week 5, then subsequently declined. Mean worm burden at slaughter was 6100 (range 60--25,080). Two sheep had visible lesions in the small intestine at slaughter. In both sheep, numerous shallow red depressed areas extended for 5 to 7 metres distal to the pylorus while, in one, more extensive irregular depressed areas occupied the first 1.5 metres. Microtopographical examination and scanning electron microscopy showed that these affected areas were devoid of villi and contained numerous worms embedded in the mucosa. Histological examination revealed many similarities with the lesions caused by T colubriformis. Subtotal villous atrophy and crypt elongation were present in the intestines from five of the remaining six sheep, and in these the mucosa contained numerous mononuclear cells and intraepithelial globule leucocytes.

Chronic infection with Trichostrongylus vitrinus in sheep. Some effects on food utilisation, skeletal growth and certain serum constituents.

The effect on food utilisation and skeletal growth of daily infection for 90 days with 2500 Trichostrongylus vitrinus larvae was investigated. Controls were killed initially or fed ad libitum. Food intake, digestibility of dry matter and nitrogen and content of body fat, protein, water and minerals were determined, and selected bones examined chemically and histologically. Serum constituents were determined weekly. Digestibility of food dry matter and nitrogen were reduced during weeks 3 and 5 after initial infection but subsequently recovered. Deposition of fat, protein, calcium and phosphorus in the body was reduced by 36, 58, 67 and 78 per cent respectively. Gross efficiency of utilisation of metabolisable energy was reduced by 36 per cent. Bone changes were associated with a reduction in osteoblastic activity. Hypoalbuminaemia, hyperglobulinaemia and hypophosphataemia were present in infected sheep. There were no significant changes in serum calcium or urea concentrations. It is concluded that the effects of continuous intakes of T vitrinus and T colubriformis larvae on host metabolism are essentially the same.

Chronic subclinical ovine fascioliasis: plasma glutamate dehydrogenase, gamma-glutamyl transpeptidase and aspartate aminotransferase activities and their significance as diagnostic aids.

The effect of low grade chronic Fasciola hepatica infection on the concentration of plasma glutamate dehydrogenase (GD), gamma-glutamyl transpeptidase (gamma-GT) and aspartate aminotransferase (AST) was investigated in sheep dosed daily with three (AL3), eight (AL8) or 14 (AL14) metacercariae over 22 weeks or given a single dose of 200 metacercariae. Significant increases in plasma GD activity first occurred after nine, 12 and 23 weeks and in gamma-GT activity after 12, 24 and 32 weeks for groups AL14, AL8 and AL3 respectively. Changes in AST activity were not as clearly related to dose level. In sheep with single infection, both GD and gamma-GT were capable of detecting liver damage resulting from the migration of 10 or more flukes. Plasma GD and gamma-GT activities are more sensitive indicators of liver cell damage in chronic subclinical fascioliasis than AST activity and gamma-GT may be more suitable as a diagnostic aid on account of its greater stability.

Effect of ostertagiasis on copper status in sheep: a study involving use of copper oxide wire particles.

Lambs infected with Ostertagia circumcincta larvae and uninfected controls were either doses with 5 g copper oxide wire particles (COWP) or remained undosed. The change in abomasal pH was monitored from duodenal digesta and that in liver copper concentration from initial liver biopsy samples and liver obtained at necropsy after 22 days. Infection increased the pH of digesta from 2.5 to 4.5. The change in liver copper content in sheep not treated with COWP was +6.1 mg (12.6 per cent) and -6.8 mg (13.8 per cent) in control and infected sheep, respectively. Significantly greater amounts of COWP were recovered from the abomasa of infected than from control animals (3.6 +/- 0.23 and 1.6 +/- 0.55 g, respectively) and hepatic uptake of copper from COWP was 0.7 and 1.8 per cent of the dose, respectively. There were significant relationships between the pH of duodenal contents and COWP retained, soluble copper concentration in duodenal digesta and hepatic uptake of copper. It was concluded that, through causing an increase in pH in abomasal and duodenal digesta, gastrointestinal nematodes interfere with copper metabolism.

Subclinical trichostrongylosis in growing lambs produced by continuous larval dosing. The effect on performance and certain plasma constituents.

Three groups of eight-four-month-old worm-free lambs were used to investigate the effect of Trichostrongylus colubriformis infection on growth rate, food intake, and concentration of plasma constituents. Groups ALC and ALI were fed ad libitum, the latter being dosed daily with 2500 infective larvae for 14 weeks. Lambs in group PF were paired to individuals in group ALI and offered the same quantity of food as the infected pair consumed. Hypophosphataemia and hypoalbuminaemia developed in ALI at week 3 and hyperglobulinaemia was present from week 6. There was no significant difference in plasma calcium, protein, urea or glucose concentration between the three groups at any stage. No clinical signs of parasitism were seen in seven of eight ALI lambs and egg counts were generally less than 2000 epg. Worm burdens of 1500-82,900 were found at slaughter. Group weight gain was only 50 per cent of PF lambs which received the same digestible energy intake. Group ALI consumed 15-5 kg dry matter per kg gain in bodyweight compared with 8-6 kg/kg and 9-8 kg/kg for ALC and PF respectively. The infections resulted in extensive villous atrophy and flattening of the intestinal mucosa. Bone growth was arrested and osteoporosis present in five group ALI lambs. The possible use of plasma constituents, particularly phosphorus, as aids for the diagnosis of the early stages of trichostrongylosis in the field is assessed.

Effect of copper oxide wire particle treatment on establishment of major gastrointestinal nematodes in lambs.

Eighty-four 10-week-old lambs were used to investigate the effect of copper oxide wire particle treatment on the establishment of major gastrointestinal nematodes. They were maintained on pasture previously treated to minimise larval contamination. Five grams of uniform sized copper oxide wire particles were given orally five days before infection with either 20,000 Trichostrongylus colubriformis larvae, 20,000 Ostertagia circumcincta larvae or 3000 Haemonchus contortus larvae given as three doses at three-day intervals. The animals were slaughtered 21 or 22 days after the last infective dose. Parasite burdens in the lambs treated with copper oxide wire particles were reduced by 96 per cent in the case of H contortus and by 56 per cent in the case of O circumcincta compared to burdens in controls. There was no significant effect of copper particles on the establishment of T colubriformis.

An evaluation of plasma homocysteine in the assessment of vitamin B12 status of pasture-fed sheep.

AIM: To assess the diagnostic potential of concentrations of homocysteine (Hcy) in plasma in relation to those of methylmalonic acid (MMA) and vitamin B12, as predictors of responsiveness of young sheep to supplementation with vitamin B12. METHODS: Eighty-two ewes grazing ryegrass-white clover pasture were used, 39 of which had been supplemented with a Co bullet and 43 unsupplemented. Thirty days after commencement of parturition their lambs (n=53 and 59, respectively) were randomly allocated into one of two treatments, in a 2 x 2 factorial design. Half of the lambs from each group of ewes received an injection of vitamin B12, while the remainder were controls. The trial commenced 31 October 2001 (Day 0), and continued until 01 May 2002 (Day 182). All lambs were weighed, and blood samples taken from 16 identified animals from each treatment group, at approximately monthly intervals. Changes in concentrations of Hcy, vitamin B12 and MMA in plasma, and liveweight gain (LWG) of the treatment groups were evaluated during the suckling (Days 0-89) and post-weaning (Days 90-182) periods. RESULTS: Mean LWG was 40% greater in supplemented than unsupplemented lambs. The concentrations of vitamin B12 and MMA in plasma in the unsupplemented lambs were in the deficient reference ranges of <170 pmol/L and >16 mumol/L, respectively. Mean monthly concentrations of Hcy in plasma ranged from 1.5 to 4.5 mumol/L but showed no pattern of response to vitamin B12 deficiency or supplementation. CONCLUSIONS: Measurement of the concentration of Hcy in plasma as a metabolic indicator of reduced methylation capability of sheep on typical pastures in New Zealand appeared to have little value in detection of vitamin B12 responsiveness, and was less sensitive than the concentration of the vitamin itself or the indicator of adenosyl-cobalamin deficiency, MMA, in plasma. The possibility that concentrations of Hcy in plasma remain low due to re-methylation of Hcy to methionine via the alternative betaine-choline rather than the vitamin B12-dependent methyl-tetrahydrofolate metabolic pathway is rejected, but the possibility is raised that high rates of trans-sulphuration of Hcy to cysteine in the gastrointestinal tract of grazing sheep could be responsible. CLINICAL RELEVANCE: The propionate-succinate pathway appears to be the first rate-limiting pathway in vitamin B12 deficiency, and the product of disruption of this pathway, increased MMA, is the most reliable indicator of metabolic abnormality in predicting responsiveness to supplementation.