Impact of strong selection for the PrP major gene on genetic variability of four French sheep breeds (Open Access publication)

Effective selection on the PrP gene has been implemented since October 2001 in all French sheep breeds. After four years, the ARR "resistant" allele frequency increased by about 35% in young males. The aim of this study was to evaluate the impact of this strong selection on genetic variability. It is focussed on four French sheep breeds and based on the comparison of two groups of 94 animals within each breed: the first group of animals was born before the selection began, and the second, 3–4 years later. Genetic variability was assessed using genealogical and molecular data (29 microsatellite markers). The expected loss of genetic variability on the PrP gene was confirmed. Moreover, among the five markers located in the PrP region, only the three closest ones were affected. The evolution of the number of alleles, heterozygote deficiency within population, expected heterozygosity and the Reynolds distances agreed with the criteria from pedigree and pointed out that neutral genetic variability was not much affected. This trend depended on breed, i.e. on their initial states (population size, PrP frequencies) and on the selection strategies for improving scrapie resistance while carrying out selection for production traits.     

Prion‐like Doppel gene polymorphisms and scrapie susceptibility in portuguese sheep breeds

The establishment of an association between prion protein gene (PRNP) polymorphisms and scrapie susceptibility in sheep has enabled the development of breeding programmes to increase scrapie resistance in the European Union. Intense selection for PRNP genotype may lead to correlated selection for genes linked to PRNP. We intended to investigate if any association exists between genetic variation in prion‐like protein Doppel gene (PRND) and scrapie susceptibility, determined through PRNP genotyping. Sampling included 460 sheep from eight Portuguese breeds and the PRND gene coding region was analysed by multiple restriction fragment‐single strand conformation polymorphism (MRF‐SSCP), whereas PRNP genotyping was carried out by primer extension. A synonymous substitution (c.78G>A) was detected in codon 26 of the PRND gene, in all breeds except Churra Mondegueira. Linkage disequilibrium was found between the PRND and PRNP loci (P = 0.000). Specifically, PRND was monomorphic in the 45 animals with the more resistant ARR/ARR PRNP genotype (P = 0.003), whereas a higher frequency of PRND heterozygotes (GA) was associated with ARQ/AHQ (P = 0.029). These results constitute preliminary evidence of an association between a polymorphism in the PRND gene and scrapie susceptibility, and indicate that the possibility of undesirable consequences from widespread selection for PRNP genotype on genetic diversity and reproduction traits needs to be further investigated.     

p.Asn176Lys and p.Met137Thr dimorphisms of the PRNP gene significantly decrease the susceptibility to classical scrapie in ARQ/ARQ sheep

In this study, we investigated the susceptibility to scrapie of Sarda breed sheep carrying the genotype ARQ/ARQ with additional polymorphisms at the PRNP gene. To do this, we examined 256 scrapie-affected sheep and 320 flock-mate negative controls from 24 flocks. Logistic regression analysis demonstrated that sheep carrying the ARQ/ARQ genotype with additional dimorphisms had lower risk of becoming scrapie affected when compared with those with ARQ/ARQ_wildtype genotype. ARQ/ARQ genotypes that were detected with heterozygous or homozygous p.Asn176Lys and p.Met137Thr dimorphisms were associated with the lowest susceptibility to the disease. A significant lower risk was also associated with the p.Arg154His dimorphism, while p.Leu141Phe had a protective effect that was not statistically significant.     

Genetic variability of six French meat sheep breeds in relation to their genetic management

Some demographic parameters, the genetic structure and the evolution of the genetic variability of six French meat sheep breeds were analysed in relation with their management. Four of these breeds are submitted to more or less intense selection: the Berrichon du Cher (BCH), Blanc du Massif Central (BMC), Charollais (CHA) and Limousin (LIM); the other two breeds are under conservation: the Roussin de La Hague (RLH) and Solognot (SOL). Genealogical data of the recorded animals born from 1970 to 2000 and of their known ancestors were used. The most balanced contributions of the different flocks to the sire-daughter path was found in the SOL. In the BCH, a single flock provided 43% of the sire-AI sire path, whereas the contributions of the flocks were more balanced in the BMC and LIM (the only other breeds where AI is used to a substantial amount). The distribution of the expected genetic contribution of the founder animals was found to be unbalanced, especially in the BCH and LIM. The effective numbers of ancestors (founders or not) for the ewes born from 1996 to 2000 were equal to 35 (BCH), 144 (BMC), 112 (CHA), 69 (LIM), 40 (RLH) and 49 (SOL). Inbreeding was not analysed in the BMC, due to incomplete pedigree information. From 1980 on, the rates of inbreeding, in percentage points per year, were +0.112 (BCH), +0.045 (CHA), +0.036 (LIM), +0.098 (RLH) and +0.062 (SOL). The implications of the observed trends on genetic variability are discussed in relation to the genetic management of each breed. The need for a larger selection basis in the BCH, the efficiency of the rules applied in the SOL to preserve the genetic variability and the need for a more collective organisation in the CHA and RLH are outlined.     

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.     

Community-based sheep breeding programs generated substantial genetic gains and socioeconomic benefits

Community-based breeding programs (CBBPs) for small ruminants have been suggested as alternatives to centralised, government-controlled breeding schemes which have been implemented in many developing countries. An innovative methodological framework on how to design, implement and sustain CBBPs was tested in three sites in Ethiopia: Bonga, Horro and Menz. In these CBBPs, the main selection trait identified through participatory approaches was 6-month weight in all three sites. In Horro and Bonga, where resources such as feed and water permitted larger litter sizes, twinning rate was included. Ten-year (2009 to 2018) performance data from the breeding programs were analysed using Average Information Restricted Maximum Likelihood method (AI-REML). Additionally, the socioeconomic impact of CBBPs was assessed. Results indicated that 6-month weight increased over the years in all breeds. In Bonga, the average increase was 0.21 ± 0.018 kg/year, followed by 0.18 ± 0.007 and 0.11 ± 0.003 kg/year in Horro and Menz, respectively. This was quite substantial in an on-farm situation. The birth weight of lambs did not improve over the years in Bonga and Horro sheep but significant increases occurred in Menz. Considering that there was no direct selection on birth weight in the community flock, the increased weights observed in Menz could be due to correlated responses, but this was not the case in Bonga and Horro. The genetic trend for prolificacy over the years in both Bonga and Horro flocks was positive and significant (P < 0.01). This increase in litter size, combined with the increased 6-month body weight, increased income by 20% and farm-level meat consumption from slaughter of one sheep per year to three. The results show that CBBPs are technically feasible, result in measurable genetic gains in performance traits and impact the livelihoods of farmers.     

A 12 000-rad whole-genome radiation hybrid panel in sheep: application to the study of the ovine chromosome 18 region containing a QTL for scrapie susceptibility

Whole-genome radiation hybrid (RH) panels have been constructed for several species, including cattle. RH panels have proven to be an extremely powerful tool to construct high-density maps, which is an essential step in the identification of genes controlling important traits, and they can be used to establish high-resolution comparative maps. Although bovine RH panels can be used with ovine markers to construct sheep RH maps based on bovine genome organization, only some (c. 50%) of the markers available in sheep can be successfully mapped in the bovine genome. So, with the development of genomics and genome sequencing projects, there is a need for a high-resolution RH panel in sheep to map ovine markers. Consequently, we have constructed a 12 000-rad ovine whole-genome RH panel. Two hundred and eight hybrid clones were produced, of which 90 were selected based on their retention frequency. The final panel had an average marker retention frequency of 31.8%. The resolution of this 12 000-rad panel (SheepRH) was estimated by constructing an RH framework map for a 23-Mb region of sheep chromosome 18 (OAR18) that contains a QTL for scrapie susceptibility.     

Multiplicative genetic effects in scrapie disease susceptibility

Despite experimental evidence that scrapie is an infectious disease of sheep, variations of the occurrence of the natural disease suggest an influence of host genetic factors. It has been established that the genetic polymorphism of the prion protein (PrP) gene is correlated to the incidence of scrapie and to the survival time: five polymorphisms have been described by variations at amino-acid codons 136, 154 and 171. In this paper we study the effect on scrapie susceptibility of the pairing of the five allelic variants known to exist: we show that scrapie susceptibility is given by the produce of the elementary allelic factors. This first well-documented evidence of a multiplicative property of genetic risk factors could give hints on the underlying mechanisms of prion-induced neurodegenerative diseases.     

Frequencies of PrP genotypes and their implication for breeding against scrapie susceptibility in nine Pakistani sheep breeds

Prion protein (PrP) gene of 308 sheep was genotyped to investigate polymorphisms at scrapie-associated codons 136, 154 and 171 to assess the resistance of nine different Pakistani sheep breeds to natural/typical scrapie. As a result six genotypes were established on the basis of polymorphic codons 154 and 171. The most scrapie-susceptible codon 136 (A/V) was monomorphic (A) in all breeds. Wild-type genotype ARQ/ARQ was detected with maximum prevalence ranging from 63.2% in crossbred Pak-karakul to 100% in native Buchi, Kachi and Thalli breeds. The most frequent of typical scrapie-associated genotypes was ARQ/ARR as indicated by five of nine breeds. The coding region of PrP gene of 49 animals from the total sampled was also sequenced to ascertain additional polymorphisms. Polymorphism was found in 13 animals of the six breeds in codons 101(Q/R), 112(M/T), 146(N/S) and 189(Q/L) and ten genotypes were established on the basis of these polymorphic codons. Only Hissardale possessed five of the ten genotypes. The most frequent genotype was M¹¹²ARQ/T¹¹²ARQ detected in Hissardale, Pak-karakul and Awassi, whereas genotypes ARQr²³¹/ARQr²³¹ and ARQR²³¹/ARQr²³¹ (established on the basis of silent polymorphism agg/cgg-R/R) were detected in all breeds. Some animals consisted of three polymorphisms at different PrP codons that are not common in European breeds. An infrequent double heterozygosity (c/c a/g g/t) for codon 171 resulting in a genotype R/H was also detected in three animals each one from Kajli, Hissardale and Pak-karakul. This study concludes that all native sheep breeds are poor in scrapie-resistant PrP genotypes and could contract scrapie if exposed to prions.     

Genetic variation within the Merino sheep breed: analysis of closely related populations using microsatellites

Genetic relationships among six populations of Merino sheep were investigated using microsatellites. The history of the six populations is relatively well documented, with all being derived from the Spanish Merino breed within the last 400 years. Genetic variation was highest amongst the Spanish and Portuguese populations, although the preservation of genetic diversity within the other populations was high. By a variety of different statistical tests the French Mutton, German Mutton and New Zealand Merino populations could be differentiated from each other and the Iberian Merinos, indicating that microsatellites are able to track relatively recent changes in the population structure of sheep breeds. The dendrograms constructed on the basis of microsatellite allelic frequencies showed that populations that have shared selection criteria (meat vs. wool) tend to cluster together.     

Tracing genetic differentiation of Chinese Mongolian sheep using microsatellites

The genetic consequences of population differentiation and isolation have been the subject of conservation biology. In this study, we analysed the genetic diversity and structure of Mongolian sheep in China. These animals belong to a traditional local breed with high production, extensive adaption, early maturity and roughage resistance. For this purpose, 26 microsatellites were genotyped for five Mongolian sheep populations. The Bayesian clustering indicated five clusters as the most probable genetic structure of the populations investigated. In addition, a clear genetic structure was revealed in three populations distributed at large geographical scales, while the other cluster encompassed UQ and HLBR sheep that displayed no clear differentiation, probably due to their close and small geographical distributions. Overall, our results are helpful in understanding the interplay of population dynamics in these close genetic lineages of Mongolian sheep.     

Defining the breeding goal for a sheep breed including production and functional traits using market data

In this study, the economic values for production and functional traits of dairy sheep are estimated through the application of a profit function model using farm-level technical and economic data. The traits incorporated in the model were milk production, prolificacy, fertility, milking speed, longevity and mastitis occurrence. The economic values for these traits were derived as the approximate partial derivative of the specified profit function. A sensitivity analysis was also conducted in order to examine how potential changes in input and output prices would affect the breeding goal. The estimated economic values of the traits revealed their economic impact on the definition of the breeding goal for the specified production system. Milk production and fertility had the highest economic values (€40.30 and €20.28 per standard genetic deviation (SDa)), while, mastitis only had a low negative value of −0.57 €/SDa. Therefore, breeding for clinical mastitis will have a minor impact on farm profitability because it affects a small proportion of the flock and has low additive variance. The production traits, which include milk production, prolificacy and milking speed, contributed most to the breeding goal (70.0%), but functional traits still had a considerable share (30.0%). The results of this study highlight the importance of the knowledge of economic values of traits in the design of a breeding program. It is also suggested that the production and functional traits under consideration can be categorized as those which can be efficiently treated through genetic improvement (e.g. milk production and fertility) while others would be better dealt with through managerial interventions (e.g. mastitis occurrence). Also, sub-clinical mastitis that affects a higher proportion of flocks could have a higher contribution to breeding goals.     

The growth of sheep scab lesions in relation to sheep breed and time of the year

Numerically equal groups of between five and ten young Merino and Dorper sheep were artificially infested with sheep scab mites, Psoroptes ovis. Separate groups of sheep were infested during the early winter of two successive years as well as during the early summer of the second year. Lesion growth was measured at two-weekly intervals for a period of six weeks during the first winter and for eight weeks during the second winter and in the summer. The rate of lesion growth and mean lesion size on Merino sheep was significantly greater than that on Dorper sheep at each two-weekly recording during the winter of both years. The rate of growth of the lesions did not differ significantly between the two sheep breeds during summer, although, with the exception of eight weeks post-infestation, the mean lesion size on Merino sheep was significantly larger than that on Dorper sheep at each assessment. This revised version was published online in July 2006 with corrections to the Cover Date.     

Profitability of a dairy sheep genetic improvement program using artificial insemination

This simulation study investigated the farm-level economic benefits of a genetic improvement scheme using artificial insemination (AI) with fresh ram semen in dairy sheep of the Chios breed in Greece. Data were collected from 67 farms associated with the Chios Sheep Breeders' Cooperative 'Macedonia', describing the percentage of ewes that would be artificially inseminated in the flock, pregnancy rate, annual ram costs that could be saved using AI rather than natural mating, expected improvement in milk production, annual costs of semen and feed, milk price and number of years of AI usage. The study considered 77 760 possible scenarios in a 3 × 4 × 4 × 3 × 3 × 3 × 4 × 15 factorial arrangement. Analysis of variance was used to investigate the effect of each factor on farm profitability. All factors considered were statistically significant (P < 0.001), but their effect varied. The number of years using AI had the greatest effect on profitability and farmers should become aware that using AI is a long-term investment. Semen price, pregnancy rate and improvement in milk production also had substantial effects. The price of milk and feed had a considerably lower effect on profitability, as did the annual cost of maintaining rams that would be replaced by AI. A positive annual and cumulative return was achieved in the model within the first 6 years. The cost of semen was estimated at 8€ to 10€ per dose for the first 5 years. Where the annual improvement in milk production was 1% of annual phenotypic mean (e.g. 3.0 kg) profitability of the scheme was improved greatly.     

The effects on genetic variability following a recent colonization event: the Australian sheep blowfly, Lucilia cuprina arrives in New Zealand

The Australian sheep blowfly, Lucilia cuprina, was first identified in New Zealand in 1988 and is now found to have spread throughout many sheep-farming regions. L. cuprina is estimated to have been present in New Zealand < 20 years, while in Australia L. cuprina has been estimated present > 100 years. The aim of this study was to determine the genetic effects of colonization of L. cuprina and to compare populations of L. cuprina from these two countries in terms of genetic variability and differentiation. Allozyme electrophoresis was used which revealed variability at six loci. 1680 blowflies were examined from 56 sites throughout L. cuprina's range in both countries. Genetic variability at each locus in terms of allele composition was found to be high and genetic differentiation varied considerably in New Zealand in comparison to Australia. Temporal sampling in New Zealand suggests seasonal fluctuations of population size in the recently colonized region of the South Island.     

The population genetic effects of ancestry and admixture in a subdivided cattle breed

The genetic structure of the Dexter, a minority cattle breed with complex demographic history, was investigated using microsatellite markers and a range of statistical approaches designed to detect both admixture and genetic drift. Modern representatives of two putative ancestral populations, the Devon and Kerry, together with the different populations of the Dexter, which have experienced different demographic histories, were analysed. Breed units showed comparatively high levels of genetic variability ( H _E = 0.63–0.68); however, distinct genetic subgroups were detected within the Dexter, which could be attributed to known demographic events. Much lower diversity was identified in three small, isolated Dexter populations ( H _E = 0.52–0.55) and higher differentiation ( F _ST > 0.13) was found. For one of these populations, where strong selection has taken place, we also found evidence of a demographic bottleneck. Three methods for quantifying breed admixture were applied and substantial method-based variation in estimates for the genetic contribution of the two proposed ancestral populations for each subdivision of the Dexter was found. Results were consistent only in the case of a group consisting of selected Traditional Dexter animals, where the ancestor of the modern Kerry breed was also determined as the greater parental contributor to the Dexter. The inconsistency of estimation of admixture proportions between the methods highlights the potentially confounding role of genetic drift in shaping small population structure, and the consequences of accurately describing population histories from contemporary genetic data.     

母体遗传效应对青海细毛羊生产性能遗传参数估计的影响

为了研究母体遗传效应对青海细毛羊生长性状、产毛性状的影响,文章采用平均信息最大约束似然法应用不同混合动物模型估计青海细毛羊生产性状的遗传参数,并采用似然比检验对不同模型进行比较分析。各模型中均包括固定效应、个体直接加性遗传效应、残差效应;随机效应为:个体永久环境效应、母体遗传效应、母体永久环境效应。不同模型对随机效应作了不同考虑:模型1不考虑个体永久环境效应、母体遗传效应、母体永久环境效应;模型2考虑母体永久环境效应;模型3考虑母体遗传效应;模型4考虑母体遗传效应和母体永久环境效应;模型5考虑个体永久环境效应和母体遗传效应;模型6考虑个体永久环境效应、母体遗传效应、母体永久环境效应。各模型估计的初生重遗传力为:0.1896~0.3781;断奶重遗传力为:0.2537~0.2890;周岁重遗传力范围:0.2244~0.3225;成年羊体重遗传力范围:0.2205~0.3983;产毛量遗传力为:0.1218~0.1490;羊毛细度遗传力为:0.0983~0.4802;羊毛长度遗传力为:0.1170~0.1311。与模型1相比,模型3对于初生重、断奶重差异显著(P<0.01),对于周岁重、成年羊体重各模型与模型1的似然比检验差异不显著(P>0.05);与模型6相比,模型4、5对于羊毛细度差异显著(P<0.01),模型4对羊毛长度差异显著(P<0.05),对于产毛量各模型与模型6似然比检验差异不显著(P>0.05)。生长性状中初生重、断奶重受母体遗传效应影响显著,周岁重、成年羊体重受母体遗传效应影响不显著;产毛性状中羊毛细度、长度受母体遗传效应影响显著,产毛量受母体遗传效应影响较弱。     

母体遗传效应对青海细毛羊生产性能遗传参数估计的影响

为了研究母体遗传效应对青海细毛羊生长性状、产毛性状的影响, 文章采用平均信息最大约束似然法应用不同混合动物模型估计青海细毛羊生产性状的遗传参数, 并采用似然比检验对不同模型进行比较分析。各模型中均包括固定效应、个体直接加性遗传效应、残差效应; 随机效应为：个体永久环境效应、母体遗传效应、母体永久环境效应。不同模型对随机效应作了不同考虑：模型1不考虑个体永久环境效应、母体遗传效应、母体永久环境效应; 模型2考虑母体永久环境效应; 模型3考虑母体遗传效应; 模型4考虑母体遗传效应和母体永久环境效应; 模型5考虑个体永久环境效应和母体遗传效应; 模型6考虑个体永久环境效应、母体遗传效应、母体永久环境效应。各模型估计的初生重遗传力为：0.1896~0.3781; 断奶重遗传力为：0.2537~0.2890; 周岁重遗传力范围：0.2244~0.3225; 成年羊体重遗传力范围：0.2205~0.3983; 产毛量遗传力为：0.1218~0.1490; 羊毛细度遗传力为：0.0983~0.4802; 羊毛长度遗传力为：0.1170~0.1311。与模型1相比, 模型3对于初生重、断奶重差异显著(P<0.01), 对于周岁重、成年羊体重各模型与模型1的似然比检验差异不显著(P>0.05); 与模型6相比, 模型4、5对于羊毛细度差异显著(P<0.01), 模型4对羊毛长度差异显著(P<0.05), 对于产毛量各模型与模型6似然比检验差异不显著(P>0.05)。生长性状中初生重、断奶重受母体遗传效应影响显著, 周岁重、成年羊体重受母体遗传效应影响不显著; 产毛性状中羊毛细度、长度受母体遗传效应影响显著, 产毛量受母体遗传效应影响较弱。     

湖羊结构基因座遗传共适应性分析

根据多座位电泳法检测的结构基因座上等位基因频率分析湖羊群体的遗传共适应性。结果发现, 在显隐性－显隐性模式中未发现座位间的遗传共适应现象, 显隐性－共显性模式中X-p-Cat组合座位以及共显性－共显性模式中Po-CA、Po-Cat组合座位均存在遗传共适应, 表明在湖羊群体中性结构基因座间存在遗传共适应, 而且起主要的作用, 维持着座位间的遗传平衡或者使座位间处于遗传不平衡状态。