Periparturient Relaxation Research Articles

Field testing of recombinant subunit vaccines against Teladorsagia circumcincta in lambing ewes demonstrates a lack of efficacy in the face of a multi-species parasite challenge

IntroductionWe previously demonstrated efficacy of an 8-antigen recombinant subunit vaccine against a single species homologous Teladorsagia circumcincta challenge in lambs and in lambing ewes in pen trials. We subsequently demonstrated efficacy of a simplified, 2-antigen, version of this vaccine in lambs in pen trials. Here, we test both vaccines in lambing ewes in a field setting.MethodsIn the work presented here, 12 adjacent plots were seeded with a mixed infection of several common species of parasitic nematodes of sheep in temperate regions, including T. circumcincta. Ewes (n = 144), in groups of 12, grazed for 2 years on these plots and, in the first year, six of these groups of ewes were vaccinated with a 2-antigen prototype vaccine against T. circumcincta prior to mating and then again prior to lambing. In the following year these ewes were immunised again, this time with the 8-antigen prototype vaccine against T. circumcincta prior to mating and then prior to lambing. Throughout both seasons antigen-specific serum antibody levels in ewes and faecal worm egg counts (FEC) in ewes and their lambs were monitored, along with nematode species diversity at lambing.ResultsImmunised ewes produced elevated serum antibody levels to each of the vaccine antigens following immunisation but their FEC levels were not statistically significantly impacted by vaccination with either vaccine. FEC levels were also not impacted in lambs co-grazing the pastures with these immunised ewes. Nematode species diversity was not significantly impacted by vaccination in either year.DiscussionThe immunosuppressive effects of co-infecting gastrointestinal nematodes, the absence of vaccine cross-protection against co-infecting species and the influence of the periparturient relaxation in immunity probably all contributed to the inability of either vaccine to protect against T. circumcincta infection in field trials in the work presented here.

Dietary protein supplementation results in molecular and cellular changes related to T helper type 2 immunity in the lung and small intestine in lactating rats re-infected with Nippostrongylus brasiliensis.

Acquired immunity to gastrointestinal nematodes reduces during late pregnancy and lactation which is known as periparturient relaxation of immunity (PPRI). Protein supplementation reduces the degree of PPRI in a rat model re-infected with Nippostrongylus brasiliensis, but the underlying molecular mechanisms have yet to be elucidated. Here, we hypothesized that protein supplementation will enhance T helper type 2 immunity (Th2) in the lung and small intestine. Nulliparous Sprague-Dawley rats were given a primary infection of N. brasiliensis prior to mating and restrictedly fed diets with either low protein (LP) or high protein (HP) during pregnancy and lactation. Dams were secondary infected with N. brasiliensis on day 2 post-parturition, and histology and gene expression were analysed for tissue samples collected at days 5, 8 and 11. Genes related to Th2 immunity in the lung, Retnla, Il13 and Mmp12, and in the intestine, Retnlb, were upregulated in HP dams compared to LP dams, which indicates the effect of dietary protein on Th2 immunity. HP dams also had increased splenic CD68+ macrophage populations compared to LP dams following secondary infection, suggesting enhanced immunity at a cellular level. Our data assist to define strategic utilization of nutrient supply in mammals undergoing reproductive and lactational efforts.

Galectin Secretion and Modulation in Sheep Blood

Galectins (GAL) are β-galactoside binding proteins that can modulate both pregnancy and the response to pathogens. Increased susceptibility to infection by pathogens is linked to periparturient immune suppression and a periparturient rise (PPR) in parasite eggs on pasture. The possible role of GAL in periparturient immune relaxation and PRR needs definition.  The objective of this study was to evaluate galectin secretion and its relationship to measures of the immune relaxation and PRR in periparturient sheep. Samples were collected from pregnant St. Croix sheep (n=6) weekly at days -21 to + 21 relative to lambing. Fecal samples were collected and evaluated for strongyle and coccidia parasite eggs. The concentration of IgA and IgE coproantibodies, total microbial DNA, Bifidobacteria and Lactobacillus levels in fecal samples were used as indicators of gut health.  Blood samples were collected by jugular venipuncture and assessed for Packed Cell Volume (PCV), total and differential white blood cell counts. Total protein concentrations and protein profile were evaluated in serum. Secretion of GAL 1, 3, 9 and 14 were evaluated using ELISAs. Data were analyzed by one-way ANOVA and statistical significance was declared at P <0.05. Galectins tested were secreted in sheep blood. Differential modulation of GAL secretion and correlation with periparturient immune suppression and parasite infection was observed. Galectin secretion was modulated by the periparturient period, type and status of parasite infection. This first insight into a possible role of secreted galectins in periparturient immune relaxation and PRR improves the understanding of the immune response, informs development of management programs and therapeutics and presents Galectin profiles as biomarkers with diagnostic potential.

A monitoring study to explore gastrointestinal helminth burdens of ewes of different fecundities in the periparturient period and effects on lamb growth rates

As a result of the periparturient relaxation of immunity(PPRI) in ewes and subsequent rise in parasite burdens it has become common for sheep farmers to worm all ewes prior to lambing. However, with increased prevalence of anthelmintic resistance it is necessary to find more sustainable worming strategies. In this study, the faecal egg counts (FECs) of untreated ewes in different fecundity groups (single-, twin- and triplet-bearing ewes) in the periparturient period, together with lamb FECs and lamb growth rates were monitored on a commercial organic lowland sheep farm in Oxfordshire, UK. Triplet-bearing ewes were supplemented with a high-protein concentrate ration and molasses prior to lambing. All ewe groups showed a relaxation in immunity against gastro-intestinal helminths from the week lambing began until eight weeks later. Twin-bearing ewes had a significantly higher overall worm burden as compared to single- and triplet-bearing ewes. Single lambs had the highest daily live weight gains (DLWGs) during the period from tagging to shearing of their mothers. Faecal egg counts of lambs were performed from six weeks after lambing. A general trend towards increased Trichostrongyloidea FEC with litter size was seen in the lambs. As a result, recommendations for an effective, practical and sustainable nematode control programme for the periparturient period in ewes of different fecundity for UK sheep farmers have been devised. The core recommendations are to not treat single-bearing ewes unless a high parasite burden is detected by FEC, to treat a proportion of twin- and triplet-bearing ewes, and to supplement triplet-bearing ewes with protein prior to lambing and wet foster the third lamb, if practical in indoor-lambing units.

Protection of ewes against Teladorsagia circumcincta infection in the periparturient period by vaccination with recombinant antigens

Teladorsagiosis is a major production-limiting disease in ruminants in temperate regions throughout the world and one of the key interventions in the management of the disease is the prevention of pasture contamination with Teladorsagia circumcincta eggs by ewes during the periparturient relaxation in immunity which occurs in the period around lambing. Here, we describe the immunisation of twin-bearing ewes with a T. circumcincta recombinant subunit vaccine and the impact that vaccination has on their immune responses and shedding of parasite eggs during a continuous T. circumcincta challenge period spanning late gestation and lactation. In ewes which displayed a clear periparturient relaxation in immunity, vaccination resulted in a 45% reduction in mean cumulative faecal egg count (cFEC, p=0.027) compared to control (immunised with adjuvant only) ewes. Recombinant antigen-specific IgG and IgA, which bound each of the vaccine antigens, were detected in the serum of vaccinated ewes following each immunisation and in colostrum taken from vaccinated ewes post-partum whereas low levels of antigen-specific IgG were detected in serum and colostrum from control ewes. Antigen-specific IgG and IgA levels in blood collected within 48h of birth from lambs largely reflected those in the colostrum of their ewes.

Interactions between parasitic infections and reproductive efficiency in sheep

This review article summarises the many reports in the literature, confirming that, in sheep, parasitic infections can adversely affect reproductive efficiency; examples, which refer to all parts of the reproductive cycle of sheep, are as follows: trichostrongylosis in ewe-lambs (which can lead to delayed attainment of puberty), myiosis of the prepuce (which can cause impediment of mating), chorioptic mange or trypanosomosis in rams (which can lead to testicular degeneration or azoospermia, respectively), trypanosomosis or sarcoptic mange in pre-conceptual ewes (which can lead to poor conception rates or reduced number of ovulations, respectively), toxoplasmosis or neosporosis in pregnant ewes (which are causes of abortion), trichostrongylosis or trematode infections in lactating ewes (which can cause reduction of milk yield and can be a risk factor for mastitis, respectively), cryptosporidiosis in newborn lambs (which can be a cause of deaths), coccidiosis in growing pre-weaned lambs (which can cause suboptimal growth rate). In other cases, the reproductive status of the animal can influence the parasitic infection; examples are as follows: the increase in faecal parasitic output during the peri-parturient period (as a consequence of the peri-parturient relaxation of immunity), the heavier trichostrongylid infections of twin lambs compared to lambs from single parities (as a consequence of developmental origin issues in twin lambs). All the above examples support the idea of presence of interactions between parasitic infections and reproductive efficiency in sheep.

Amelioration of the periparturient relaxation of immunity to parasites through a reduction in mammalian reproductive effort

The degree of periparturient relaxation of immunity to gastrointestinal parasites has a nutritional basis, as overcoming protein scarcity through increased protein supply improves lactational performance, enhances local immune responses and reduces worm burdens. Herein lactating rats, re-infected with Nippostrongylus brasiliensis, are used to test the hypothesis that a similar and rapid improvement of immunity can be achieved through reducing nutrient demand at times of dietary protein scarcity. Reducing litter size from 12 to three pups during lactation resulted, as expected, in cessation of maternal body weight loss and increased pup body weight gain compared with dams which continued to nurse 12 pups. This increase in performance concurred with a rapid decrease in parasitism; within 3days post nutrient reduction, a 87% reduction in the number of worm eggs found in the colon and 83% reduction in worm burdens was observed, which concurred with increased local immune responses, i.e. 70% more mast cells and 44% more eosinophils in the small intestinal mucosa, to levels similar to those in dams nursing three pups throughout. However, there were no concurrent changes in goblet cell hyperplasia, serum anti-N. brasiliensis-specific antibody levels or mRNA expression of IL-4, IL-10 or IL-13 in the mesenteric lymph nodes. To our knowledge the current study is the first to employ a litter reduction strategy to assess the rate of immune improvement upon overcoming nutrient scarcity in a non-ruminant host. These data support the hypothesis that periparturient relaxation of immunity to gastrointestinal nematodes can be reduced by restoring nutrient adequacy and, importantly, that this improvement can occur very rapidly.

Sensitivity of periparturient breakdown of immunity to parasites to dietary protein source1

Effects of increased MP supply on the degree of periparturient relaxation of immunity (PPRI) in sheep may be dependent on quality of supplied MP. Here we tested the hypothesis that additional MP supply from rations based on xylose-treated soybean meal would be more effective than from rations based on faba beans in reducing the degree of PPRI, as indicated by nematode egg excretion. Twenty-four multiple-bearing ewes were trickle infected with Teladorsagia circumcincta larvae from d -56 to d 31 relative to start of lactation (d 0). From d -26 onwards, ewes were fed at either 0.8 (LP) or at 1.2 times their respective calculated MP requirements using either xylose-treated soybean (HPS) or faba beans (HPB). Litter size was adjusted to 2 lambs at parturition. Feeding treatments did not affect nematode egg excretion, ewe BW or BCS during late pregnancy (P > 0.10), but HPS and HPB ewes had reduced plasma pepsinogen concentrations (P = 0.003). During lactation, HPS and HPB feeding increased ewe BW gain (P < 0.001) and BCS (P = 0.017), and reduced plasma pepsinogen concentrations (P = 0.008) to the same extent, compared with LP feeding. However, only HPS feeding increased litter weight gain (P = 0.017) and reduced nematode egg excretion (P = 0.015), which were both similar between HPB and LP (P > 0.10). The results support the view that extra MP supply from xylose-treated soybean based rations is more effective in reducing parasitism than MP from faba bean-based rations, suggesting that protein source and/or quality are important factors to consider for the nutritional control of parasitism.

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 L3/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, IgG1, 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, IgG1, 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.

Dietary protein and energy supplies differentially affect resistance to parasites in lactating mammals

Periparturient relaxation of immunity (PPRI) to parasites in mammals results in higher worm burden and worm egg excretion and may have a nutritional basis. Nippostrongylus brasiliensis re-infected lactating rats fed low-crude protein (CP) diets show an augmented degree of PPRI compared with their high CP-fed counterparts. However, such effects of CP scarcity have been confounded by metabolisable energy (ME) scarcity due to increased intake of the high-CP foods. Here, we independently assessed the effects of dietary CP and ME scarcity on the degree of PPRI. Second, parity rats were infected with N. brasiliensis larvae before mating. Upon parturition, dams were allocated to one of six feeding treatments (1-6), consisting of two levels of dietary ME supply, each with three levels of CP supply. On day 2 of lactation, dams were either re-infected with 1600 N. brasiliensis larvae or sham-infected with PBS, while litter size was standardised at ten pups. Dams and litters were weighed daily until either day 8 or 11 of lactation, when worm burdens were assessed as a proxy for PPRI. Increased CP and ME supply independently improved lactational performance. While ME supply did not affect parasitism, increasing CP supply reduced worm burden and the percentage of female worms in the small intestine; the latter was especially pronounced at the lower level of ME supply. The present results support the view that PPRI to parasites may be sensitive to CP scarcity, but not to moderate ME scarcity.

Dissecting the impact of protein versus energy host nutrition on the expression of immunity to gastrointestinal parasites during lactation

Many mammals exhibit a periparturient relaxation of previously established immune responses (PPRI) to gastrointestinal nematodes culminating in increased worm burdens. It has been suggested that the extent of PPRI may have a nutritional basis as it is considerably augmented when protein supply is scarce. Subsequent studies have shown that increased dietary protein intake can ameliorate this phenomenon. However, this effect is often confounded with increased food intake and thus increased energy levels. Herein, we aimed to dissect the effects of protein and energy nutrition on the immune status and resistance to re-infection with gastrointestinal nematodes in the periparturient host. The lactating, Nippostrongylus brasiliensis re-infected rat was utilised as an established model for mammalian PPRI. Experimental animals were assigned to restricted feeding regimens designed to achieve four pre-determined levels of crude protein (CP) at one of two levels of metabolisable energy (ME) and parasitological and immunological measurements taken at either day 6 or day 9 post re-infection. We clearly show that increased supply of dietary CP, but not increased dietary ME, significantly reduced worm burdens. The increased magnitude of worm expulsion with increased dietary CP supply strongly correlated with mucosal mast cell accumulation in the small intestine. In addition, increased CP and not ME supply increased mucosal eosinophil numbers. Furthermore, increased CP led to higher levels of total IgG at high ME only and there were interactive effects of CP and ME on serum levels of IgG1 and IgG2a. Perhaps surprisingly, CP nutrition did not affect expression of either Th1 (IFN-γ) or Th2 (IL-4, IL-13) cytokines in the mesenteric lymph nodes. These data emphasise the role of immunonutrition, and particularly dietary protein, in combating infectious disease such as gastrointestinal parasitism.

Nutritional sensitivity of periparturient resistance to nematode parasites in two breeds of sheep with different nutrient demands

The periparturient relaxation of immunity (PPRI) to parasites in mammals is sensitive to both metabolisable protein (MP) supply and animal genotype (different reproductive outputs). We tested the hypothesis that the sensitivity of PPRI to MP scarcity would not differ between different levels of reproductive output when nutrient intake is adjusted for associated differences in MP demand; this hypothesis assumes that PPRI has a nutritional basis only. Scottish Blackface (BF) and the more productive Mule (MU) ewes were infected with the abomasal parasite Teladorsagia circumcincta, and from day -21 to day 32 (day 0 is parturition), they were fed restrictedly at either 0.8 (low protein (LP)) or 1.3 (high protein (HP)) times their breed-specific estimated MP requirement (n 18 for each breed-feeding treatment combination). During late pregnancy, LP feeding reduced ewe body weight gain in both breeds, tended to increase faecal egg count (FEC), but it did not affect plasma pepsinogen. During lactation, LP feeding reduced litter growth rate and ewe plasma urea and plasma albumin concentrations compared with HP feeding in both breeds. However, breed and feeding treatment interacted for ewe FEC, worm egg excretion and plasma pepsinogen, which were higher for the LP-MU ewes compared with the HP-MU and BF ewes. The lower degree of PPRI of the BF ewes during lactation compared with the MU ewes at a similar degree of MP scarcity suggests that the effect of reproductive output on nutritional sensitivity of PPRI cannot be explained by associated differences in nutrient demand only.

The periparturient relaxation of immunity in Merino ewes infected with Trichostrongylus colubriformis: Endocrine and body compositional responses

The temporal association between the periparturient rise in worm egg count of grazing Merino ewes to infection with Trichostrongylus colubriformis and the underlying causal mechanisms was investigated in an experiment that incorporated two levels of pregnancy (pregnant or unmated), two levels of infection (infected with 6000 T. colubriformis L 3/week or uninfected) and, following lambing, three levels of lactation (non-lactating/“dry”, early-weaned 2 days after parturition, or suckled). In addition to parasitological and host immune responses reported in a previous paper ( Beasley et al., 2010), a range of endocrine and body compositional changes were monitored from day −50 to day 42, relative to the midpoint of lambing (day 0). By day −19, pregnant ewes had begun to mobilise fat and eye muscle, and after 42 days of suckling had lost 31% and 8%, respectively, of their existing depots. In comparison to non-pregnant (dry) ewes, the endocrine profiles of late-pregnant ewes were characterised by low levels of cortisol and prolactin and high levels of progesterone and oestradiol. Lactating ewes had lower levels of cortisol and leptin and higher levels of prolactin compared to both dry and early-weaned ewes. The mobilisation of fat and protein reserves throughout lactation in suckled ewes was closely associated with leptin and cortisol profiles, and provided strong evidence of an underlying nutritional basis for the periparturient relaxation of immunity. Both leptin and cortisol concentrations were also associated with both parasite burden and the immune status of the ewe. It is suggested that lower blood cortisol levels in suckled ewes contribute to a Th1 biased immune response that leads to an increased susceptibility to gastrointestinal nematodes. The results provide a detailed characterisation of the physiology underlying the periparturient relaxation of immunity to T. colubriformis, from which further investigations will aim to expose potential causal factors.

The periparturient relaxation of immunity in Merino ewes infected with Trichostrongylus colubriformis: Parasitological and immunological responses

The temporal association between the periparturient rise (PPR) in worm egg count (WEC) of grazing Merino ewes to infection with Trichostrongylus colubriformis and the underlying causal mechanisms was investigated in an experiment that incorporated two levels of pregnancy (pregnant or unmated), two levels of infection (infected with 6000 T. colubriformis L 3/week or uninfected) and, following lambing, three levels of lactation (non-lactating/“dry”, early-weaned 2 days after parturition, or suckled). The 128 ewes in the experiment were monitored for a range of parasitological and host systemic and local immune responses from days −50 to 42, relative to the midpoint of lambing (day 0). Unmated/non-lactating (dry) ewes remained largely resistant to T. colubriformis infection with WEC remaining below 200 eggs per gram (epg) throughout the experiment. For infected ewes, the PPR in WEC commenced during late pregnancy and peaked at 1552 epg on day 42. There was evidence of a relaxation of immunity that preceded the rise, as demonstrated by a reduction in circulating eosinophils and plasma total antibody (Tab) and lower titres of Tab and IgG 1 in the small intestine in pregnant ewes. Circulating eosinophils and plasma Tab remained low in suckled ewes, and at the local level, lactating ewes also had lower titres of Tab, IgG 1, IgM, IgA and IgE as well as fewer mast cells (MC), globule leucocytes (GL) and goblet cells (GC) in intestinal tissue. These effects were associated with higher worm burdens. Weaning at 2 days after lambing resulted in a rapid amelioration of immunological responsiveness and a reduction in WEC, worm burden and parasite establishment rate. A rapid increase in circulating eosinophils and Tab titre followed by an improvement in components of the local immune response was also observed. Over the 6-week period following lambing, weaned ewes exhibited higher titres of Tab, IgG 1, IgM, IgA and IgE as well as higher numbers of MC, GL and GC present in the small intestinal tissue, compared to lactating ewes. The results provide a detailed characterisation of the periparturient relaxation of immunity to T. colubriformis, and highlight possible causative factors that deserve further investigation.

Consequences of infection pressure and protein nutrition on periparturient resistance to Teladorsagia circumcincta and performance in ewes

The consequences of protein nutrition on the degree of periparturient relaxation of immunity to nematode parasites in sheep may be more pronounced at higher levels of infection pressure. Here, we investigated interactive effects of metabolizable protein (MP) nutrition and infection pressure on resistance and lactational performance of ewes. Twin-rearing ewes were trickle infected with either 1000, 5000 or 10,000 infective Teladorsagia circumcincta larvae and fed either at 0.8 (low protein, LP) or 1.3 (high protein, HP) times their estimated MP requirement. Expected interactions between feeding treatment and infection pressure were not observed. Periparturient relaxation of immunity, as indicated by variation in faecal egg counts, was higher in LP ewes than in HP ewes and FEC showed an inverse relationship with infection pressure indicating possible density dependency effects on worm fecundity. Plasma pepsinogen concentration linearly increased with infection pressure. Daily total nematode egg excretion, assessed at week three of lactation, was not significantly affected by infection pressure but was reduced by 65% in HP ewes compared to LP ewes. MP supplementation improved lamb performance but had little effect on ewe body weight and plasma protein concentrations, whilst lactational performance, as judged from lamb performance, tended to be reduced with increased infection pressure. The results suggest periparturient MP supplementation to ewes reduces nematode egg excretion independent of infection pressure and improves lactational performance of parasitized ewes even in the presence of moderate MP scarcity.

Immunomodulatory effects of dietary protein duringNippostrongylus brasiliensisre-infection in lactating rats

Periparturient relaxation of immunity (PPRI) to secondary infection with nematodes is believed to have a nutritional basis due to differential partitioning of scarce nutrient resources, particularly protein, to reproductive rather than immune functions. At times of protein scarcity, an increase in protein supply has been reported to assuage this phenomenon. The Nippostrongylus brasiliensis reinfected lactating rat model is now being utilized to investigate the immune reactions underlying the modifying role of dietary protein on PPRI. Herein, we demonstrate that lactating rats reinfected with N. brasiliensis under high protein (HP) dietary conditions exhibit decreased worm burdens and reduced colon egg counts compared to their low protein (LP) counterparts. These reductions correlated with increased mastocytosis and greater goblet cell hyperplasia. Additionally, the local antibody profile revealed that HP reinfected lactating rats developed a stronger antigen specific IgG2b response earlier in infection in comparison with their LP counterparts. Our study provides evidence that increased dietary protein content reduces the PPRI to N. brasiliensis re-infection in the lactating rat through improved mucosal immune responses.

Are Scottish Blackface ewes more resistant to nematode parasite infection than Greyface ewes?

The extent of periparturient relaxation of immunity (PPRI) to gastrointestinal nematode parasites is sensitive to metabolisable protein (MP) scarcity but also varies between breeds of sheep (Houdijk, 2008). For example, under ad libitum feeding, Scottish Blackface ewes had a lower extent of PPRI than Greyface ewes (Zaralis et al., 2008). Such between-breed variation in PPRI may not necessarily be associated with genetic resistance per se but could arise from a higher nutrient demand of the more productive Greyface ewes (Houdijk, 2008). This experiment was designed to test the hypothesis that if the extent of PPRI has only a nutritional basis, then the sensitivity of PPRI to MP scarcity will not differ between the breeds when MP feeding is adjusted for between-breed differences in MP demand.

Consequences of protein supplementation for anorexia, expression of immunity and plasma leptin concentrations in parasitized ewes of two breeds

The periparturient relaxation of immunity (PPRI) against parasites in ewes has a nutritional basis. We investigated whether ewes experience a reduction in food intake (anorexia) during PPRI and if the magnitude of anorexia is affected by host production potential and dietary protein supplementation. We also investigated whether nematode infection is linked to plasma leptin concentrations in periparturient ewes. The experiment was a 2 x 2 x 2 factorial design. Two breeds of twin-bearing/lactating ewes (Greyface cross, G (n 32) and Scottish Blackface, B (n 32)) were used. Half of the ewes were trickle infected with 30,000 larvae of the abomasal parasite Teladorsagia circumcincta per week and the other half were not. During the experiment, all ewes had ad libitum access to a low-protein diet that provided less protein than the recommended allowance. In addition, half of the ewes received a protein supplement that resulted in protein intakes that exceeded recommendations. Nematode infection resulted in a breakdown of immunity to parasites and a reduction in food intake in both breeds. The breeds differed in the extent of PPRI (G ewes having higher faecal egg counts than B ewes), but not in the magnitude of anorexia. Protein supplementation resulted in a reduction in faecal egg counts, but had no effect on the magnitude of anorexia. Plasma leptin concentrations changed significantly over time, but were not affected by protein supplementation or infection. It is concluded that infection with T. circumcincta in periparturient ewes results in anorexia that is not alleviated by protein supplementation and seems unrelated to plasma leptin concentrations.

Influence of periparturient nutritional demand on resistance to parasites in livestock

Periparturient relaxation of immunity (PPRI) to gastrointestinal nematode parasites may have a nutritional basis, as the increasing nutrient demand at times of nutrient scarcity during late pregnancy and subsequent lactation may result in a penalty on expression of acquired immunity to parasites. This nutritional basis implies that lowering nutritional demand, at constant nutrient supply, should reduce the degree of PPRI. Evidence to support the latter is reviewed through exploration of the effects of reproductive effort on periparturient resistance to parasites. A large body of evidence shows that a lower nutritional demand arising from rearing single rather than multiple lambs and/or kids consistently reduces the degree of PPRI in small ruminants, as manifested by reduced worm burdens and/or nematode egg excretions. Such variation in reproductive effort may also account, at least to some extent, for the often observed between-breed differences in PPRI, which may arise from differences in production potential, and thus nutritional demand. A reduction in nutrient demand can reduce the degree of PPRI in a matter of days. However, host immune responses associated with this nutritional sensitivity of host resistance remain to be elucidated, which may be achieved through a recently established periparturient rodent model.

Effects of protein supplementation on anorexia and expression of immunity in parasitized periparturient ewes of two different breeds

The breakdown of acquired immunity during the periparturient period in the ewe can be reduced by metabolizable protein (MP) supplementation (Houdijk et al 2001). Recent evidence suggests that the development of immune response results in reduced food intake (anorexia) in many disease models (Materase et al, 2005). However, it is not known whether an immune response following the periparturient relaxation of immunity is associated with a reduction in food intake and whether this is affected by protein supplementation in parasitized ewes. In addition, differences in nutrient partitioning between sheep breeds that differ in production potential may affect the ability of the hosts to express immunity and this may be reflected by differences in their magnitude and/or duration of anorexia. The aim of the present study was to test the hypotheses that: a) nematode infection during the periparturient period results in anorexia and protein supplementation can effect the degree of anorexia in ewes, and b) ewes of a high production potential breed show higher breakdown of acquired immunity than ewes of a low production potential breed, and exhibit higher degree of anorexia.