Rémy Arquet

Genetic variability in resistance to gastro-intestinal strongyles during early lactation in Creole goats

The study was undertaken in a Creole goat flock at INRA-Gardel in Guadeloupe, to evaluate the opportunity to use artificial selection as a means of controlling gastro-intestinal infection during early lactation. The flock grazed all year on Digitaria decumbens pastures. Faecal and blood samples were taken from kids at 11 months of age and from does at kidding before drenching (week 0) and at weeks 4 and 6 after kidding. Faecal egg counts (FEC) were estimated using a modified McMaster method. Blood samples were used to determine packed cell volume (PCV) and eosinophil concentrations (EOS) values. Haemonchus contortus , Trichostrongylus colubriformis and Oesophagostomum columbianum were the main strongyle species identified in faecal cultures. The data came from 1092 litters obtained from 688 does sired by 142 bucks and 413 dams. Variance and covariance components for genetic and residual effects were estimated with multivariate animal models using the restricted maximum likelihood VCE package. Repeatability and overall heritability for FEC during the post-partum period were 0·17 and 0·10±0·02. The genetic correlations between FEC and PCV were −0·56±0·11 at 4 weeks after kidding and −0·79±0·13 at 6 weeks after kidding. The genetic correlations between FEC and EOS were 0·37±0·15 at 4 weeks after kidding and 0·68±0·17 at 6 weeks after kidding. Hence, does that contributed least to pasture contamination during the postpartum period also had low EOS and high PCV breeding values. The genetic correlations between FEC measured at 11 months of age and FEC during periparturient period ranged from 0·57±0·12 to 0·76±0·16. Therefore, breeding goats for increased resistance during the post-weaning period will lead to a less marked and less persistent rise in doe FEC during early lactation. The epidemiological implications of this selection have to be quantified in terms of lower pasture contamination, lower kid parasitism, and higher milk production of does.

Serum antibody responses in Creole kids experimentally infected with Haemonchus contortus.

The objective of this study was to evaluate the relationship of parasite-specific serum antibodies with the resistance status of Creole kids. The average breeding values on egg output predicted in a context of natural infection at 11 months of age were distant of 1.07 genetic standard deviation between resistant and susceptible animals. After drenching the animals were maintained worm-free during 1 month until experimental infection with 10,000 Haemonchus contortus infective larvae (L3). Enzyme-linked immunosorbent assay was carried out in serum samples to determine the level of IgG, IgA and IgE anti-H. contortus L3 crude extracts and adult excretion/secretion products (ESP). Parasitological and blood immunological parameters were measured on the 2 extreme groups. Despite the absence of any typical signs of haemonchosis, susceptible kids had more than 11 times higher faecal egg counts (FEC) at 35 days post-infection (d.p.i.) than resistant kids had. Levels of immunoglobulin against H. contortus L3 and ESP increased significantly after infection in both groups. However, no difference in the host immune response mediated by immunoglobulin against H. contortus was evidenced between groups. This finding suggests that, in goats previously infected by H. contortus, a degree of protection occurred and the phenotypic and genetic segregation in resistant and susceptible animals were not related to the humoral immune response. The correlation coefficients between FEC and IgE anti-ESP (r=0.593; P<0.05 was significant in both resistant and susceptible animals. Such correlation suggesting a hypersensitivity reaction dependent on worm prolificacy has never been described. This result needs further studies to understand the mechanisms underlying this observation.

Effect of dietary supplementation on resistance to experimental infection with Haemonchus contortus in Creole kids

The aim of the present study was to test the effect of dietary supplementation on resistance to experimental infection with Haemonchus contortus in Creole kids. One trial with three replicates involved a total of 154 female kids that were chosen from three successive cohorts of the Creole flock of INRA-Gardel in 2007. The kids were placed into four treatments according to the amount of concentrate they received: G0 (no concentrate and a quality Dichantium spp. hay ad libitum, HAY), G1 (HAY+100g commercial concentrate d(-1)), G2 (HAY+200 g commercial concentrate d(-1)), G3 (HAY+300 g commercial concentrate d(-1)). The G0-G3 groups were infected with a single dose of 10,000 H. contortus third stage larvae (L(3)) at Day 0 (D0). Each infected group was comprised of one half resistant and one half susceptible genetically indexed kids. The average breeding values on egg excretion at 11 months of age were distant of 0.70, 0.65, 0.61 and 0.61 genetic standard deviations in G0, G1, G2 and G3, respectively. The faecal egg count (FEC), packed cell volume (PCV), eosinophilia (EOSI) and dry matter intake (DMI) indices were monitored weekly until 42 days post-infection. Enzyme-linked immunosorbent assay was carried out on serum samples to determine the level of IgA anti-H. contortus L(3) crude extracts and adult excretion/secretion products (ESP). The 10,000 L(3) dose received by the kids induced a severe infection: 8000 eggs per gram at the FEC peak, a PCV less than 15% and mortality. Interestingly, the supplemented animals in G3 showed a higher level of EOSI but a lower level of IgA anti-L3 and IgA anti-ESP than non-supplemented animals (G0). Resistant and susceptible kids had significantly different FEC variations within the groups. Susceptible kids had a 1.6 times higher egg output than resistant kids in G0. This difference was not found in the supplemented groups. The results of this study showed that supplementary feeding improved resistance of Creole kids to H. contortus experimental infection.

Intake and digestibility of naive kids differing in genetic resistance and experimentally parasitized (indoors) with Haemonchus contortus in two successive challenges.

We evaluated the effects of infection with Haemonchus contortus on feed intake, digestibility, fecal egg count, circulating eosinophils, and packed cell volume in Creole kids differing in genetic resistance (susceptible, S; resistant, R) to gastrointestinal parasitism and maintained on a similar level of nutrition. The experiment was carried out during 2 periods of 6 wk each differing in immunity development stage. In the first period (acquisition of immunity; period I), 22 naïve male kids (23.4 +/- 0.65 kg of BW) were housed in individual boxes and fed a hay-based diet, and a primary infection was induced. In the second period (expression of immunity; period II), 15 of the initial 22 kids (28.4 +/- 0.77 kg of BW) were submitted to a secondary infection. Housing and management were uniform throughout the experiment. For each period, measurements of intake and digestibility were made at 0, 2, and 4 wk postinfection (WPI) with a single dose of 10,000 infective larvae (L(3)). The DMI and total-tract DM, OM, CP, NDF, and ADF digestibilities were determined using the total feces collection and ad libitum forage supply method. Fecal and blood samples were collected weekly to measure fecal egg count, circulating eosinophils, and packed cell volume. Infection with Haemonchus contortus decreased feed intake during period I. The absence of anorexia in period II was probably due to the acquired immunity of kids. The DMI was affected (P = 0.05) by genetic predisposition to resistance (626 vs. 583 +/- 26 g/d, for R vs. S) and WPI, being greatest in the second WPI (693 vs. 614 and 657 g/d, for WPI-2 vs. WPI-0 and WPI-4, respectively). The latter was related to worm establishment phase and was linked to the lower total tract digestibilities at this point. Digestibilities were least at WPI-2. The fecal egg counts were greater (P < 0.001) in period I than II, and differences between S and R were evident after the fifth WPI in period II. Circulating eosinophils were greater (P < 0.001) in S vs. R. The results suggest that effects of these parasites on intake and digestibility are influenced by the individual genetic resistance and the immunological stage, and the strongest impact occurs between the second and the third WPI, a period during which the immune response is more pronounced, probably due to parasite maturation.

Genetic analysis of the potential role of IgA and IgE responses against Haemonchus contortus in parasite resistance of Creole goats.

Small ruminants are affected by gastrointestinal nematode (GIN) infection. A promising alternative strategy for control of GIN infection is to increase the level of resistance in the population by taking advantage of the hosts immune response. Immunoglobulin A (IgA) and E (IgE) are known to be involved in immune response to GIN. The aim of this study was thus to investigate genetic parameters of IgA and IgE responses against Haemonchus contortus in Creole kids naturally challenged at pasture and to determine the relationship with other resistance criteria such as faecal egg counts, packed-cell volume, eosinophil counts and bodyweight. Variance and covariance components for genetic and residuals effects for each trait were estimated on 3862 males at 11 months of age. Heritability estimates for IgA and IgE ranged between 0.15 and 0.57. Strong positive genetic correlations were observed between either IgE or IgA responses against L3 and adult excretory/secretory products (ESP) antigens of H. contortus, suggesting that the humoral immune response is not specific to the life cycle stage of the parasite suggesting that there is substantial cross recognition between the different parasite antigens. Heritability estimates for faecal egg count (FEC), packed-cell volume (PCV) and bodyweight (BW) were in accordance with previous results in Creole kids. Blood eosinophil counts were found moderately heritable and negatively correlated with FEC, suggesting that this cell population plays a role in resistance to nematode parasite infection in Creole goats. IgA response was positively correlated to FEC, in contrast with the negative correlation between IgE against L3 of H. contortus and FEC. In Creole goats, IgA response against L3 or ESP of H. contortus would rather be associated with the worm burden than an immune protective response. The immune response involving activity of IgE against L3 of H. contortus may be one important pathway for development of resistance to gastrointestinal nematode infections in Creole goats.

Economic values of body weight, reproduction and parasite resistance traits for a Creole goat breeding goal

A specific breeding goal definition was developed for Creole goats in Guadeloupe. This local breed is used for meat production. To ensure a balanced selection outcome, the breeding objective included two production traits, live weight (BW11) and dressing percentage (DP) at 11 months (the mating or selling age), one reproduction trait, fertility (FER), and two traits to assess animal response to parasite infection: packed cell volume (PCV), a resilience trait, and faecal worm eggs count (FEC), a resistance trait. A deterministic bio-economic model was developed to calculate the economic values based on the description of the profit of a Guadeloupean goat farm. The farm income came from the sale of animals for meat or as reproducers. The main costs were feeding and treatments against gastro-intestinal parasites. The economic values were 7.69€ per kg for BW11, 1.38€ per % for FER, 3.53€ per % for DP and 3 × 10(-4)€ per % for PCV. The economic value for FEC was derived by comparing the expected profit and average FEC in a normal situation and in an extreme situation where parasites had developed resistance to anthelmintics. This method yielded a maximum weighting for FEC, which was -18.85€ per log(eggs per gram). Alternative scenarios were tested to assess the robustness of the economic values to variations in the economic and environmental context. The economic values of PCV and DP were the most stable. Issues involved in paving the way for selective breeding on resistance or resilience to parasites are discussed.

Genetic parameters for body weight, reproduction, and parasite resistance traits in the Creole goat

We estimated the genetic parameters for BW, reproduction, and parasite resistance traits to implement a breeding program for the Creole goat. The traits were preweaning BW at 70 d of age (BW70d), BW at 11 mo of age (BW11), fecal egg count at 11 mo of age (FEC11) for all animals, packed cell volumes of lactating does (PCV), and their fertility (FER) and litter size (LS). We analyzed about 30 yr of data, which included 18,450 records on 11,970 animals from the INRA experimental flock in Guadeloupe (French West Indies). Heritability estimates were low for reproduction traits (0.11 ± 0.02 for LS and FER) to moderate for production traits (0.32 ± 0.03 for BW11; 0.20 ± 0.03 and 0.08 ± 0.02 for the direct and maternal heritability estimates of BW70d, respectively). Heritability estimates for gastrointestinal nematode resistance traits were situated in an intermediate range (0.13 ± 0.05 for PCV and 0.18 ± 0.04 for FEC11). Genetic correlations between FER, PCV, BW11, and the maternal effect of BW70d were altogether positive, whereas LS and FEC11 were almost uncorrelated phenotypically and genetically. These correlations are very favorable for setting up a breeding program, making it possible to improve BW, reproduction, and parasite resistance traits simultaneously.

Production systems of Creole goat and their implications for a breeding programme.

The Creole goat is a local meat breed well adapted to the tropical environment of Guadeloupe, a French island in the Caribbean. A survey of 47 goat farmers was conducted in May 2008 to describe the Guadeloupean goat farming systems. It was the preliminary step for the implementation of a breeding programme for Creole goats. Farmers had 31 does on average. A small number (4%) kept only Creole goats. Most of them (62%) had a mixed herd of Creole and crossbreds. One-third of them (34%) reared only crossbred goats. Farmers appreciate the rusticity and resistance of the Creole goat but consider its growth as too slow. The most desired traits for goat selection were conformation and growth for males (77% of the answers). These traits were also important for females (30% of the answers). Maternal qualities were also frequently cited (maternal behaviour 23%, reproduction 20% and milk production 17%). Disease resistance was not seen as an important trait (10% and 7% of the answers for bucks and does, respectively). A typology constituted of five groups of farmers was also created. Farmers of three groups were retained to participate at a selection programme. They kept Creole goats and have expressed a strong willingness to join a selection programme. The results of the survey suggest that a breeding programme should mostly focus on the Creole goat as a maternal breed. Real consideration should be given to disease resistance. The Creole goat has indeed a key role to play in the sustainability of local farming systems.

Detection of quantitative trait loci for resistance to gastrointestinal nematode infections in Creole goats

This study aimed to identify regions of the genome affecting resistance to gastrointestinal nematodes in a Creole goat population naturally exposed to a mixed nematode infection (Haemonchus contortus, Trichostrongylus colubriformis and Oesophagostomum columbianum) by grazing on irrigated pasture. A genome-wide quantitative trait loci (QTL) scan was performed on 383 offspring from 12 half-sib families. A total of 101 microsatellite markers were genotyped. Traits analysed were faecal egg count (FEC), packed cell volume (PCV), eosinophil count and bodyweight (BW) at 7 and 11 months of age. Levels of activity of immunoglobulin A (IgA) and activity of immunoglobulin E (IgE) anti-Haemonchus contortus L3 crude extracts and adult excretion/secretion products (ESPs) were also analysed. Using interval mapping, this study identified 13 QTL for parasite resistance. Two QTL linked with FEC were found on chromosomes 22 and 26. Three QTL were detected on chromosomes 7, 8 and 14 for eosinophil counts. Three QTL linked with PCV were identified on chromosomes 5, 9 and 21. A QTL for BW at 7 months of age was found on chromosome 6. Lastly, two QTL detected on chromosomes 3 and 10 were associated with IgE anti-L3, and IgE anti-ESP was linked with two QTL on chromosomes 1 and 26. This study is the first to have identified regions of the genome linked with nematode resistance in a goat population using a genome scan. These results provide useful tools for the understanding of parasite resistance in small ruminants.

A long term experiment of integrated control of nematode parasitism in Creole goats

Infection with gastro-intestinal nematodes (GIN) is one of the most prevalent diseases affecting small ruminants in the tropics. Mixed grazing between sheep and cattle reduces infection of the small ruminants and improves growth performances (Mahieu et al., 1997). Also, it is possible to select Creole goats on resistance to GIN during the post weaning growth and the periparturient period (Mandonnet et al., 2006). This study presents the potential benefits of combining these two control methods.