Exploring the assumptions underlying genetic variation in host nematode resistance (Open Access publication)

The wide range of genetic parameter estimates for production traits and nematode resistance in sheep obtained from field studies gives rise to much speculation. Using a mathematical model describing host – parasite interactions in a genetically heterogeneous lamb population, we investigated the consequence of: (i) genetic relationships between underlying growth and immunological traits on estimated genetic parameters for performance and nematode resistance, and (ii) alterations in resource allocation on these parameter estimates. Altering genetic correlations between underlying growth and immunological traits had large impacts on estimated genetic parameters for production and resistance traits. Extreme parameter values observed from field studies could only be reproduced by assuming genetic relationships between the underlying input traits. Altering preferences in the resource allocation had less pronounced effects on the genetic parameters for the same traits. Effects were stronger when allocation shifted towards growth, in which case worm burden and faecal egg counts increased and genetic correlations between these resistance traits and body weight became stronger. Our study has implications for the biological interpretation of field data, and for the prediction of selection response from breeding for nematode resistance. It demonstrates the profound impact that moderate levels of pleiotropy and linkage may have on observed genetic parameters, and hence on outcomes of selection for nematode resistance.     

Accuracy of genomic prediction within and across populations for nematode resistance and body weight traits in sheep

Genomic prediction utilizes single nucleotide polymorphism (SNP) chip data to predict animal genetic merit. It has the advantage of potentially capturing the effects of the majority of loci that contribute to genetic variation in a trait, even when the effects of the individual loci are very small. To implement genomic prediction, marker effects are estimated with a training set, including individuals with marker genotypes and trait phenotypes; subsequently, genomic estimated breeding values (GEBV) for any genotyped individual in the population can be calculated using the estimated marker effects. In this study, we aimed to: (i) evaluate the potential of genomic prediction to predict GEBV for nematode resistance traits and BW in sheep, within and across populations; (ii) evaluate the accuracy of these predictions through within-population cross-validation; and (iii) explore the impact of population structure on the accuracy of prediction. Four data sets comprising 752 lambs from a Scottish Blackface population, 2371 from a Sarda×Lacaune backcross population, 1000 from a Martinik Black-Belly×Romane backcross population and 64 from a British Texel population were used in this study. Traits available for the analysis were faecal egg count for Nematodirus and Strongyles and BW at different ages or as average effect, depending on the population. Moreover, immunoglobulin A was also available for the Scottish Blackface population. Results show that GEBV had moderate to good within-population predictive accuracy, whereas across-population predictions had accuracies close to zero. This can be explained by our finding that in most cases the accuracy estimates were mostly because of additive genetic relatedness between animals, rather than linkage disequilibrium between SNP and quantitative trait loci. Therefore, our results suggest that genomic prediction for nematode resistance and BW may be of value in closely related animals, but that with the current SNP chip genomic predictions are unlikely to work across breeds.     

Genetic parameters for resistance to trichostrongylid infection in dairy sheep

In sheep, the traditional chemical control of gastrointestinal nematode (GIN) parasites with anthelmintics has led to the widespread development of anthelmintic resistance. The selection of sheep with enhanced resistance to GIN parasites has been suggested as an alternative strategy to develop sustainable control of parasite infections. Most of the estimations of the genetic parameters for sheep resistance to GIN parasites have been obtained from young animals belonging to meat- and/or wool-specialised breeds. We present here the estimated genetic parameters for four parasite resistance traits studied in a commercial population of adult Spanish Churra dairy ewes. These involved two faecal egg counts (FECs) (LFEC0 and LFEC1) and two serum indicator traits, the anti-Teladorsagia circumcincta fourth stage larvae IgA (IgA) and the pepsinogen (Peps) levels. In addition, this study has allowed us to identify the environmental factors influencing parasite resistance in naturally infected Spanish Churra sheep and to quantify the genetic component of this complex phenotype. The heritabilities estimated for the two FECs analysed (0.12 for LFEC0 and 0.09 for LFEC1) were lower than those obtained for the examined serum indicators (0.19 for IgA and 0.21 for Peps). The genetic correlations between the traits ranged from 0.43 (Peps-IgA) to 0.82 (LFEC0-LFEC1) and were higher than their phenotypic counterparts, which ranged between 0.07 and 0.10. The heritabilities estimated for the studied traits were lower than previously reported in lambs. This may be due to the differences in the immune mechanisms controlling the infection in young (antibody reactions) and adult (hypersensitivity reactions) animals/sheep. In summary, this study demonstrates the presence of heritable variation in parasite resistance indicator traits in the Churra population studied, which suggests that genetic improvement is feasible for this complex trait in this population. However, further studies in which the experimental variables are controlled as much as possible are needed to identify the best trait that could be measured routinely in adult sheep as an indicator of parasite resistance.     

Confirmation of two quantitative trait loci regions for nematode resistance in commercial British terminal sire breeds

Sheep internal parasites (nematodes) remain a major health challenge and are costly for pasture-based production systems. Most current breeding programmes for nematode resistance are based on indicator traits such as faecal egg counts (FEC), which are costly and laborious to collect. Hence, genetic markers for resistance would be advantageous. However, although some quantitative trait loci (QTL) have been identified, these QTL are often not consistent across breeds and few breeding strategies for nematode resistance in sheep are currently using molecular information. In this study, QTL for nematode resistance on ovine chromosomes (OAR) 3 and 14, previously identified in the Blackface breed, were explored using commercial Suffolk (n = 336) and Texel lambs (n = 879) sampled from terminal sire breeder flocks in the United Kingdom. FEC were used as the indicator trait for nematode resistance, and these were counted separately for Nematodirus and Strongyles genera. Microsatellite markers were used to map the QTL and the data were analysed using interval mapping regression techniques and variance component analysis. QTL for Nematodirus and Strongyles FEC were found to be segregating on OAR3 at 5% chromosome region-wide significance threshold in both Suffolk and Texel sheep, and Nematodirus FEC QTL were segregating on OAR14 in both breeds. In addition, QTL for growth traits were also found to be segregating at 5% chromosome region-wide on OAR3 and OAR14. The confirmation that FEC QTL segregate in the same position in three widely used breeds widens their potential applicability to purebred Blackface, Suffolk and Texel sheep, with benefits likely to be observed in their commercial crossbred progeny.     

Associations of PrP genotype with lamb production traits in three commercial breeds of British hill sheep

The National Scrapie Plan (NSP) was launched in Great Britain in 2001, with the aim of eventually eradicating scrapie, a small ruminant transmissible spongiform encephalopathy, from the national sheep flock. Specifically, a selective breeding programme, the Ram Genotyping Scheme, was devised enabling pedigree ram breeders to reduce the number of scrapie-susceptible genotypes from their flocks. The effect of large-scale manipulation of PrP genotypes on commercially important traits within the sheep industry is, however, unknown. We have therefore examined production traits in a total of 43 968 lambs from 32 pedigree breeders across three British hill breeds, comprising 8163 North Country Cheviot (Hill), 21 366 Scottish Blackface and 14 439 Welsh Mountain lambs. Traits examined included: weights at birth, 8 and 20 weeks; ultrasonic fat and muscle depth, and average daily weight gain from 8 to 20 weeks. Linear mixed models were fitted for each trait, including animal (direct) genetic effects and up to three maternal effects. Potential associations with the PrP gene were assessed by fitting either PrP genotype or number of copies of individual alleles as fixed effects. A number of breed-specific significant associations between production traits and the PrP gene were found, but no consistent significant effects were detected across the three breeds. Breed-specific effects were as follows: (i) 0.37 kg higher birth weights (BWTs) in AHQ homozygous North Country Cheviot (Hill) lambs (P < 0.01); (ii) 0.16 kg higher BWTs in ARR homozygous Scottish Blackface lambs (P < 0.05); (iii) 0.5 kg higher 8-week weights in VRQ heterozygous Scottish Blackface lambs (P < 0.01); (iv) a 0.72 kg decrease in scan weight associated with homozygous ARR Welsh Mountain lambs (P < 0.01); (v) 0.51 mm higher ultrasonic muscle depths in AHQ homozygous Welsh Mountain lambs (P < 0.01); (vi) 0.48 mm lower ultrasonic muscle depths in Welsh Mountain lambs carrying one or more copies of the ARR allele (P < 0.05) and (vii) 0.2 mm higher ultrasonic fat depths in heterozygous VRQ Welsh Mountain lambs (P < 0.05). The use of a Bonferroni correction to define appropriate significance thresholds across the three datasets, which account for the large number of independent comparisons made, resulted in breed-specific comparisons, with P < 0.01 becoming significant at P0.05, and the remaining breed-specific comparisons no longer being significant. The absence of a common effect across the three breeds suggests that any true association found may be due to breed-specific alleles of neighbouring genes in linkage disequilibrium with the PrP locus.     

Effects of genotype and plane of nutrition on growth and carcass characteristics of lambs from hill sheep systems

The objective of this study was to determine the effect of lamb genotype and plane of nutrition on post-weaning lamb growth, and carcass characteristics in hill sheep systems. The lamb genotypes compared were as follows: purebred Scottish Blackface, Swaledale × Scottish Blackface, Cheviot × Scottish Blackface, Lleyn × Scottish Blackface and Texel × Scottish Blackface lambs. During the finishing period, lambs were allocated to a (i) low plane of nutrition (LP), based on a fixed allowance of dried grass pellets representing a grass-based system; (ii) high plane of nutrition (HP-cereal), with a diet consisting proportionately of 0.20 dried grass pellets : 0.80 cereal-based concentrate offered ad libitum; (iii) high plane of nutrition (HP-cereal/oil), with a diet consisting proportionately of 0.20 dried grass pellets : 0.80 cereal-based concentrate containing oilseed rape offered ad libitum. There were significant genotype × diet interactions for dry matter (DM) intake (P < 0.001), live weight gain (P < 0.05) and carcass gain (P < 0.05). On the LP diet, carcass gains were higher for Texel cross compared to Blackface and Swaledale cross lambs (P < 0.05). On both HP diets, carcass gains were lower in Blackface and Swaledale crosses compared with the Cheviot, Lleyn and Texel cross lambs (P < 0.01). Feed conversion efficiency, expressed in terms of carcass gain per unit of DM, was highest for Texel cross and Cheviot cross, lowest in Swaledale cross and Blackface and intermediate for Lleyn cross lambs (P < 0.001). Conformation classification was lowest for Swaledale cross (P < 0.05) and highest for Texel cross lambs (P < 0.001), whilst those of Blackface, Cheviot and Lleyn cross lambs were similar. Carcasses from Blackface and Lleyn cross lambs had higher fat classifications than Texel and Cheviot cross lambs (P < 0.01). Feed conversion efficiency was lowest with the LP diet compared with the HP diets, resulting in a 5-week longer finishing period and lower carcass weights, fat scores, dressing proportions and subcutaneous fat depth measurements. These results indicate that crossbreeding can have a major effect on lamb performance in the hill sheep sector. Feeding a HP increased lamb growth rates, particularly for high growth-potential genotypes, improved feed efficiency, shortened the finishing period but increased carcass fatness. Including oilseed rape in concentrate-based diets did not affect performance.     

Genetic parameters for resistance to the Salmonella abortusovis vaccinal strain Rv6 in sheep

An experimental population (1216 lambs from 30 sires) of the Inra401 sheep was created in an Inra flock to allow QTL detection for susceptibility to Salmonella infection, wool and carcass traits. The Inra401 is a sheep composite line developed from two breeds: Berrichon du Cher and Romanov. At 113 days of age on average, the lambs were inoculated intravenously with 10⁸ Salmonella abortusovis Rv6 (vaccinal strain). They were slaughtered 10 days after the inoculation. Several traits were measured at inoculation and/or slaughtering to estimate the genetic resistance of the lambs to Salmonella infection: specific IgM and IgG1 antibody titres, body weight loss, spleen and pre-scapular node weights and counts of viable Salmonella persisting in these organs. This paper presents a quantitative analysis of the genetic variability of the traits related to salmonellosis susceptibility. The heritabilities of the traits varied between 0.10 and 0.64 (significantly different from zero). Thus, in sheep as well as in other species, the determinism of resistance to Salmonella infection is under genetic control. Moreover, the correlations between the traits are in agreement with the known immune mechanisms. The genetic variability observed should help QTL detection.     

Associations of PrP genotype with lamb production traits in three commercial breeds of British lowland sheep

The Ram Genotyping Scheme was launched in Great Britain in 2001 as part of the National Scrapie Plan and was devised to reduce and eventually eradicate classical scrapie susceptible genotypes from the national pedigree flock. Anecdotal claims from breeders suggest that sheep with more resistant PrP genotypes may have inferior phenotypes. In this study, we test this possibility for lamb production traits in three breeds of lowland sheep: Charollais (22 752 lambs), Poll Dorset (22 589 lambs) and Texel (23 492 lambs). Data were received from 50 breeders and comprised weights at birth, 8 weeks and scanning (from which average daily weight gain was derived), and ultrasonic muscle and fat depths. Animal (direct) genetic effects and up to three maternal effects were fitted in linear mixed models for each trait. Fitting either PrP genotype or number of copies of individual alleles carried as fixed effects allowed potential associations with the PrP gene to be assessed. There were no significant associations seen in the Poll Dorset breed; however, significant associations were found with the number of allele copies carried in the other two breeds included in this study. Charollais lambs carrying one copy of the VRQ allele had significantly (P < 0.01) greater ultrasonic muscle depth (0.58 mm) and fat depth (0.2 mm) than non-carriers. In the Texel breed, lambs with one ARR allele were significantly heavier than those with two or zero ARR alleles; heterozygous ARR lambs were 0.07 kg heavier at birth (P < 0.05), 0.42 kg heavier at 8 weeks (P < 0.01) and 0.17 kg heavier at scan weight (P < 0.01), than non-carriers. After Bonferroni corrections to adjust significance thresholds to account for the large number of independent comparisons made, all significant results remained so at P < 0.05 or greater, except for the ARR allele effect on birth weight in the Texel breed, which was no longer significant. These results compare favourably with others from studies on many continental breeds of sheep, published in recent years, and add credence to the conclusion that selection on PrP genotype is unlikely to have any noticeable impact on the measured growth and carcass traits in sheep.     

Quantitative genetic parameters of agronomic and quality traits in a global germplasm collection reveal excellent breeding perspectives for Jatropha curcas L.

Jatropha curcas L. (jatropha) is an undomesticated plant, which has received great attention in recent years for its potential in biofuel production and in greening and rehabilitation of wastelands. Yet the absence of improved cultivars and the lack of agronomic knowledge are limiting factors for successful jatropha cultivation. The objectives of the present study were to investigate the perspectives of a worldwide jatropha breeding program and specifically to (i) estimate variance components and heritabilities for agronomic and quality traits in the early phase of cultivation; (ii) assess phenotypic and genetic correlations among those traits; and (iii) discuss strategies for breeding high yielding jatropha cultivars. Data on various traits was collected from 375 jatropha genotypes, which were tested at seven locations during the first 3 years of growth. The accumulated seed yields and the seed yields per harvest year differed significantly among the testing locations. The estimates of genetic and genotype‐by‐environment interaction variances were significant and estimates of heritabilities were high for all yield parameters. The estimates of genetic correlations indicated a strong association among yield parameters. Oil yield was strongly correlated with seed yield and only weakly with oil content in seeds. The perspectives of a jatropha breeding program are excellent. Improved cultivars, definition of favorable environmental factors and refinement of agronomic management practices are needed to secure sustainable jatropha cultivation.     

Genetic correlations between reproduction of crossbred ewes and the growth and carcass performance of their progeny

Genetic correlations were estimated between first cross ewe reproduction traits and growth and carcass traits of their second cross lamb progeny. The 2460 crossbred ewes were progeny of 74 maternal breed sires and mainly Merino dams. The ewes had 3 joinings to terminal sire rams with 6824 joining records resulting in 9002 lambs born and 7176 lambs slaughtered. The ewe reproduction traits included: fertility, litter size, ewe rearing ability or lamb survival, number of lambs born (NLBj) and weaned (NLWj) per ewe joined, with traits reflecting ewe productivity being total litter weight weaned (TWWj) per ewe joined and the component trait average lamb weaning weight in the litter (AWW). The lamb growth traits included weight of the lambs at birth (BWT), weaning (WWT) and post weaning (PWWT) as well as growth rate pre and post weaning. The lamb carcass traits included hot carcass weight (HCWT), dressing yield (DRESS%), fat depth (FatGR, FatC), eye muscle depth (EMD) and area (EMA) and meat quality traits (colour and pH). The genetic correlations were estimated by bivariate mixed models using ASReml. The genetic correlations between the composite reproduction traits of the ewes and the post-weaning growth rate of their lambs were high (0.67 for NLBj and 0.65 for NLWj). There were moderate positive correlations between NLWj and WWT (0.36), PWWT (0.49) and pre weaning growth (0.36) and NLBj with PWWT (0.31). BWT was negatively correlated with litter size (−0.34) and positively with ewe rearing ability (0.38). Most of the other genetic correlations were smaller than their standard errors which generally ranged from 0.2 to 0.3. The genetic correlations for HCWT with all the ewe reproduction traits were positive and moderate (0.29–0.53) and high with the ewe productivity traits that included weight of lambs (TWWj 0.98 and AWW 0.96). The genetic correlations among the other traits were variable and had high standard errors, generally ranging from 0.1 to 0.5. However there were generally negative and unfavourable genetic correlations between the reproduction traits and DRESS% and meat colour L*, whereas those with carcass fat (FatC and FatGR) were generally negative and favourable.     

Analysis of prion protein genotypes in relation to reproduction traits in local and cosmopolitan German sheep breeds

Due to the genetic determination of susceptibility to scrapie and other forms of transmissible spongiform encephalopathy (TSE) in sheep breeding to the less susceptible prion protein (PrP) genotype ARR/ARR was advanced within EU. In 4961 ewes of nine German sheep breeds (Coburg Fox sheep, Gray Horned Heath sheep, Merinoland sheep, Rhoen sheep, German Blackheaded Mutton sheep, Shropshire, Suffolk, Texel and White East Friesian Milk sheep) representing local and cosmopolitan breeds the reproductive traits number of lambs born, dead (including abortion at the end of pregnancy, stillbirth and death during the first 56 days post natum), weaned and rearing rate at each lambing were recorded and in 1641 of these ewes the PrP genotype was determined. A linear model was used to evaluate associations between PrP genotype and reproduction traits including the effects of PrP genotype (four classes: ewes with two, one and no copy of the ARR allele and with unknown PrP genotype), breed, interaction of PrP genotype and breed, number of lambing, lambing season and stock. Significant associations were only observed between the PrP genotype and the number of dead lambs at each lambing in Shropshire and Merinoland sheep and the rearing rate at each lambing in Shropshire. These significant associations were mainly caused by differences between animals with unknown PrP genotype and animals of the other PrP classes. In conclusion, breeding for TSE resistant sheep will not lead to a reduction in economically important reproduction traits.     

Tolerance to deer herbivory and resistance to insect herbivores in the common evening primrose (Oenothera biennis)

The evolution of plant defence in response to herbivory will depend on the fitness effects of damage, availability of genetic variation and potential ecological and genetic constraints on defence. Here, we examine the potential for evolution of tolerance to deer herbivory in Oenothera biennis while simultaneously considering resistance to natural insect herbivores. We examined (i) the effects of deer damage on fitness, (ii) the presence of genetic variation in tolerance and resistance, (iii) selection on tolerance, (iv) genetic correlations with resistance that could constrain evolution of tolerance and (v) plant traits that might predict defence. In a field experiment, we simulated deer damage occurring early and late in the season, recorded arthropod abundances, flowering phenology and measured growth rate and lifetime reproduction. Our study showed that deer herbivory has a negative effect on fitness, with effects being more pronounced for late‐season damage. Selection acted to increase tolerance to deer damage, yet there was low and nonsignificant genetic variation in this trait. In contrast, there was substantial genetic variation in resistance to insect herbivores. Resistance was genetically uncorrelated with tolerance, whereas positive genetic correlations in resistance to insect herbivores suggest there exists diffuse selection on resistance traits. In addition, growth rate and flowering time did not predict variation in tolerance, but flowering phenology was genetically correlated with resistance. Our results suggest that deer damage has the potential to exert selection because browsing reduces plant fitness, but limited standing genetic variation in tolerance is expected to constrain adaptive evolution in O. biennis.     

Haemonchus contortus and Trichostrongylus colubriformis did not adapt to long-term exposure to sheep that were genetically resistant or susceptible to nematode infections

We tested the hypothesis that Haemonchus contortus and Trichostrongylus colubriformis would adapt to long-term exposure to sheep that were either genetically resistant or susceptible to H. contortus. Sheep genotypes were from lines with 10 years prior selection for low (resistant, R) or high (susceptible, S) faecal worm egg count (WEC) following H. contortus infection. Long-term exposure of H. contortus and T.colubriformis to R or S genotypes was achieved using serial passage for up to 30 nematode generations. Thus, we generated four nematode strains; one strain of each species solely exposed to R sheep and one strain of each species solely exposed to S sheep. Considerable host genotype differences in mean WEC during serial passage confirmed adequate nematode selection pressure for both H. contortus (R 4900 eggs per gram (epg), S 19,900 epg) and T. colubriformis (R 5300 epg, S 13,500 epg). Adaptation of nematode strain to host genotype was tested using seven cross-classified tests for H. contortus, and two cross-classified and one outbred genotype test for T. colubriformis. In the cross-classified design, where each strain infects groups of R, S or randomly bred control sheep, parasite adaptation would be indicated by a significant host genotype by nematode strain interaction for traits indicating parasite reproductive success; specifically WEC and, for H. contortus strains, packed cell volume. We found no significant evidence of parasite adaptation to host genotype (P > 0.05) for either the H. contortus or T. colubriformis strains. Therefore, we argue that nematodes will not adapt quickly to sheep bred for nematode resistance, where selection is based on low WEC, although selecting sheep using a subset of immune functions may increase adaptation risk. Our results support the hypothesis that nematode resistance is determined by many genes each with relatively small effect. In conclusion, selection of sheep for nematode resistance using WEC should be sustainable in the medium to long-term.     

Heritability of ram mating success in multi-sire breeding situations

Multi-sire mating of a mob of ewes is commonly used in commercial sheep production systems. However, ram mating success (defined as the number of lambs sired by an individual) can vary between rams in the mating group. If this trait was repeatable and heritable, selection of rams capable of siring larger numbers of lambs could reduce the number of rams required for mating and ultimately lead to increased genetic gain. However, genetic correlations with other productive traits, such as growth and female fertility, could influence the potential for ram mating success to be used as a selection trait. In order to investigate this trait, parentage records (including accuracy of sire assignment) from 15 commercial ram breeding flocks of various breeds were utilised to examine the repeatability and heritability of ram mating success in multi-sire mating groups. In addition, genetic and phenotypic correlations with growth and female fertility traits were estimated using ASReml. The final model used for the ram mating success traits included age of the ram and mating group as fixed effects. Older rams (3+years old) had 15% to 20% greater mating success than younger rams (1 or 2 years of age). Increasing the stringency of the criteria for inclusion of both an individual lamb, based on accuracy of sire assignment, or a whole mating group, based on how many lambs had an assigned sire, increased repeatability and heritability estimates of the ram mating success traits examined. With the most stringent criteria employed, where assignment of sire accuracy was >0.95 and the total number of lambs in the progeny group that failed to have a sire assigned was<0.05, repeatability and heritability for log_e(number of lambs) was 0.40±0.09 and 0.26±0.12, respectively. For proportion of lambs sired, repeatability and heritability were both 0.30±0.09. The two ram mating traits (log_e(nlamb) and proportion) were highly correlated, both phenotypically and genetically (0.88±0.01 and 0.94±0.06, respectively). Both phenotypic and genetic correlations between ram mating success and growth and other female fertility traits were low and non-significant. In conclusion, there is scope to select rams capable of producing high numbers of progeny and thus increase selection pressure on rams to increase genetic gain.     

Genetic parameters for growth and faecal worm egg count following Haemonchus contortus experimental infestations using pedigree and molecular information

Background

Haemonchosis is a parasitic disease that causes severe economic losses in sheep industry. In recent years, the increasing resistance of the parasite to anthelmintics has raised the need for alternative control strategies. Genetic selection is a promising alternative but its efficacy depends on the availability of genetic variation and on the occurrence of favourable genetic correlations between the traits included in the breeding goal. The objective of this study was twofold. First, to estimate both the heritability of and the genetic correlations between growth traits and parasite resistance traits, using bivariate linear mixed animal models, from the phenotypes and genotypes of 1004 backcross lambs (considered as a single population), which underwent two subsequent experimental infestations protocols with Haemonchus contortus. Second, to compare the precision of the estimates when using two different relationship matrices: including pedigree information only or including also SNP (single nucleotide polymorphism) information.

Results

Heritabilities were low for average daily gain before infestation (0.10 to 0.15) and average daily gain during the first infestation (0.11 to 0.16), moderate for faecal egg counts during the first infestation (0.21 to 0.38) and faecal egg counts during the second infestation (0.48 to 0.55). Genetic correlations between both growth traits and faecal egg count during the naïve infestation were equal to zero but the genetic correlation between faecal egg count during the second infestation and growth was positive in a Haemonchus contortus free environment and negative in a contaminated environment. The standard errors of the estimates obtained by including SNP information were smaller than those obtained by including pedigree information only.

Conclusions

The genetic parameters estimates suggest that growth performance can be selected for independently of selection on resistance to naïve infestation. Selection for increased growth in a non-contaminated environment could lead to more susceptible animals with long-term exposure to the infestation but it could be possible to select for increased growth in a contaminated environment while also increasing resistance to the long-term exposure to the parasite. The use of molecular information increases the precision of the estimates.     

A review of the effects of the Booroola gene (FecB) on sheep production

Booroola Merino (B^oM) ewes have a high ovulation rate and litter size which in 1980 was postulated to be due to the effects of a major gene (FecB). This was confirmed in breeding experiments and FecB was subsequently shown to be due to a mutation (BMPR-1B) on chromosome 6. The B^oM originated from an Australian commercial fine wool Merino flock (Booroola) and has been used in crossing experiments and for introgression of FecB into many breeds around the world to improve fecundity. The mutation has recently been found in native sheep breeds in India, China and Indonesia and it is likely that FecB in the Australian B^oM was derived from importations of Garole sheep from India in 1792 and 1793.The effects on production traits of the FecB mutation in a range of genetic comparisons, environments and production systems are reviewed. Comparisons involving B^oM crosses with various other breeds and contrasts of FecB homozygous (BB), heterozygous (B+) and non-carrier (++) genotypes in comparable background genotypes, including non-B^oM, have been summarised from 45 reports. The weighted mean effect for ewes carrying one copy of FecB (B+) was +1.3 (range +0.8 to +2.0) for ovulation rate and +0.7 (range +0.4 to +1.3) for litter size. The effect of a second copy (BB) was generally additive for ovulation rate, with little or no increase in litter size for BB ewes among B^oM crosses. However there was generally a further increase in litter size for BB ewes of about half the effect of one copy (B+) in the Indian and Chinese breeds. Poor lamb survival and lamb growth reduced the number of lambs weaned and total weight of lamb weaned by B+ ewes. Most studies still showed a small advantage for B+ ewes, although several reported negative effects. While embryo survival declines at higher ovulation rates, the effects of FecB per se are equivocal. There is some evidence of a higher non-pregnancy rate among homozygous BB ewes. Most studies reported lower birth weight and growth rate from B^oM cross lambs and lambs from crossbred ewes introgressed with FecB. However it is difficult to separate the effects of low background genetic merit for growth of the B^oM and the lower birth weight and growth rate of lambs from larger litters from the genetic effect of carrying FecB. There was little or no difference in growth rate between BB, B+ and ++ genotype lambs. For other traits including, seasonal oestrous activity, carcass and meat quality and wool production, there was no evidence of major effects of FecB. The opportunities for management and nutritional modification of FecB expression and implications for industry adoption are briefly discussed.     

The Norwegian sheep breeding scheme: description, genetic and phenotypic change

The developments in Norwegian sheep breeding since the early 1990s are reviewed. For the largest breeding population, the Norwegian White Sheep, results are presented for both genetic and phenotypic changes. Of the nine traits that make up the aggregate genotype, the largest gain per year, in per cent of the corresponding phenotypic average, was found for carcass grade (1.66%) and carcass weight (0.99%), number of lambs born at 1, 2 and 3 years of age (0.32% to 0.60%) and the maternal effect on weaning weight (0.26%). For fat grade, a genetic deterioration was estimated. This may be due to the too small weighting of this trait in the aggregate genotype and the true genetic parameters being somewhat different from the estimates in the prediction of breeding values. For lamb as well as ewe fleece weight, genetic change was close to zero - interpreted as mainly a correlated response to other traits in the aggregate genotype. Data for the two traits of fleece weight were, respectively, selected and few. Thus, phenotypic change was calculated for all traits except for fleece weight, and in addition for number of lambs at weaning, being indirectly selected for through number of lambs born. For all traits, with the exception of fat grade, advantageous phenotypic change was estimated. For weaning and carcass weight, the phenotypic change was less than the genetic change, while the opposite was observed for carcass and fat grade and number of lambs born. The latter traits can be more easily controlled by environmental actions, and the results thus exemplify the interdependency between environmental and genetic change.     

Recent findings on the genetics of gastro-intestinal nematode resistance in ruminants

The control of helminthiases in ruminants raised in open pasture has been mainly undertaken by using prophylactic measures in the environment, but these are often inadequate due to incorrect application. With the appearance of anthelmintics, the strategy for controlling these parasitoses, passed to pharmacological treatments which became effective in reducing their impact. However, the frequent and incorrect utilisation of these molecules resulted in resistance to anthelmintics and the presence of chemical residues in animal products for human consumption. Anthelmintic resistance is widespread throughout the world, heterogeneous and probably underestimated. This has encouraged the introduction of homeopathic agents and products derived from plants whose effectiveness has not been scientifically assessed. It is well known that it is possible to detect differences in resistance to the most important parasites between breeds. In Europe, it has been reported that some ovine autochthonous breeds, Scottish Blackface and Lacaune, showed higher resistance. The implementation of breeding strategies aimed at obtaining animals with naturally low susceptibility to nematode infestations could therefore play an increasingly important role. Standard animal breeding techniques have been largely successful in improving the performance of domestic animals in the last century. Standard quantitative selection requires field data on: i) individual phenotype performance; ii) expected covariance among animals due to blood relationship between them. The whole process of predicting the breeding value of animals in order to select subsequently the genetically superior parents of the next generation is entirely based on sophisticated computations (BLUP-animal model). In sheep, the main objective is always selecting for milk yield and sometimes, in addition, milk composition. However, due to the evolution of the EU agricultural policy and consumer demand in terms of healthy and organic food, more attention is now being given to traits related to health (resistance to EST, mastitis or parasitic diseases). Some studies conducted in New Zealand and Australia showed that nematode resistance is genetically controlled with high heritabilities and quite low genetic correlations with production traits. In this sense, some studies showed that it is possible to decrease the number of parasites in the framework of a traditional breeding programme. However, in most situations, this trait is not extensively recorded due to the high cost of individual recording. Therefore, it would be useful to implement breeding strategies based on the knowledge of the genes involved in this trait expression. Traditionally, two approaches are available to locate a gene: i) genome scan; ii) candidate gene approach. The candidate gene approach attempts to link general resistance to some particular genes. To date, genetic resistance against parasites is considered to be linked with the MHC and IgE genes. Furthermore, several gene detection studies based on the genome scan approach for this trait are currently being carried out on both crossed experimental populations (fat x lean Blackface lines and Sarda x Lacaune) and pure breeds (Churra). The preliminary results seem promising as to the use of marker assisted or genotype assisted selection for this trait, which is difficult and expensive to measure on a population scale.     

湖羊、东湖杂种羊POMC基因外显子3单核苷酸多态性及其与生长性状的关联分析

阿黑皮素原(Pro-opiomelanocortin, POMC)在动物采食和能量平衡调控中发挥重要作用, 文章对绵羊POMC基因外显子3进行扩增和测序, 筛选多态性位点, 并分析多态位点与湖羊和东弗里生×湖羊杂种羊生长性状的相关性。测序后发现湖羊POMC基因外显子3有2个单碱基突变(g.273 T/C和g.456 G/A), 根据273位点处发生的T/C突变, 建立PCR-RFLP分析方法, 并对162只湖羊和130只东湖杂种羊进行检测分析。结果发现, 在湖羊群体中检测到TT(0.469)、TC(0.438)和CC(0.093)3种基因型, 而在东湖杂种羊群体中仅检测到TT(0.754)和TC(0.246)两种基因型。POMC基因外显子3的273位点多态性与生长性状的相关性研究结果显示：湖羊群体中CC基因型个体的2月龄断奶重、4月龄尻高及TC基因型个体4月龄体长和管围均显著高于TT型个体(P<0.05); CC基因型个体的4月龄重、6月龄重极显著高于TT和TC基因型个体(P<0.01); CC基因型个体的4月龄体高和体长极显著高于TT型个体(P<0.01), 且显著高于TC基因型个体(P<0.05)。此外, CC型个体的管围极显著高于TT基因型个体(P<0.01)。东湖杂种羊群体中TC基因型个体的2月龄断奶重、4月龄重及4月龄体高、体长、胸深和管围都显著高于TT型个体(P<0.05), TC型个体的6月龄重极显著高于TT型个体(P<0.01)。研究结果表明, POMC基因外显子3与绵羊生长性状相关, C等位基因对体重及体尺性状的增加更有利。该结果为进一步探讨POMC基因作为绵羊生长性状的辅助选育标记奠定了基础。     

Genetic parameters for feed efficiency in Romane rams and responses to single-generation selection

Feeding costs represent one of the highest expenditures in animal production systems. Breeding efficient animals that express their growth potential while eating less is therefore a major objective for breeders. We estimated the genetic parameters for feed intake, feed efficiency traits (residual feed intake (RFI) and feed conversion ratio (FCR)), growth and body composition traits in the Romane meat sheep breed. In these traits, selection responses to single-generation divergent selection on RFI were evaluated. From 2009 to 2016, a total of 951 male lambs were tested for 8 weeks starting from 3 months of age. They were weighed at the beginning and at the end of the testing period. Backfat thickness and muscle depth were recorded at the end of the testing period through ultrasound measurements. Feed intake was continuously recorded over the testing period using the automatic concentrate feeders. The heritability of RFI was estimated at 0.45 ± 0.08, which was higher than the heritability of FCR (0.30 ± 0.08). No significant genetic correlations were observed between RFI and growth traits. A favourable low negative genetic correlation was estimated between RFI and muscle depth (−0.30 ± 0.15), though additional data are needed to confirm these results. The selection of low RFI sires based on their breeding values led to the production of lambs eating significantly less concentrate (3% decrease in the average daily feed intake), but with the same growth as lambs from sires selected based on high RFI breeding values. We concluded that in meat sheep, RFI is a heritable trait that is genetically independent of post-weaning growth and body composition traits. A one-generation divergent selection based on RFI breeding values highlighted that substantial gains in feeding costs can be expected in selection schemes for meat sheep breeds.